ro Late Ordovician Benthic Marine Communities in North-Central New York by Peter Bretsky BULLETIN 414 NEW YORK STATE MUSEUM AND SCIENCE SERVICE The University of the State of New York/ The State Education Department /Albany, 1970 AUG 16 197' Late Ordovician Benthic Marine Communities in North-Central New York by Peter Bretsky BULLETIN 414 NEW YORK STATE MUSEUM AND SCIENCE SERVICE The University of the State of New York/ The State Education Department / Albany, 1970 THE UNIVERSITY OF THE STATE OF NEW YORK Regents of the University (with years when terms expire) 1984 Joseph W. McGovern, A.B., LL.B., L.H.D., LL.D., D.C.L., Chancellor . New York 1985 Everett J. Penny, B.C.S., D.C.S., Vice Chancellor . White Plains 1978 Alexander J. Allan, Jr., LL.D., Litt.D. . . . Troy 1973 Charles W. Millard, Jr., A.B., LL.D., L.H.D. Buffalo 1972 Carl H. Pforzheimer, Jr., A.B., M.B.A., D.C.S., H.H.D . Purchase 1975 Edward M. M. Warburg, B.S., L.H.D. . . . New York 1977 Joseph T. King, LL.B . Queens 1974 Joseph C. Indelicato, M.D . . Brooklyn 1976 Mrs. Helen B. Power, A.B., Litt.D., L.H.D. . Rochester 1979 Francis W. McGinley, B.S., LL.B., LL.D. . . Glens Falls 1980 Max J. Rubin, LL.B., L.H.D . New York 1971 Kenneth B. Clark, A.B., M.S., Ph.D., Litt.D. Hastings on Hudson 1982 Stephen K. Bailey, A.B., B.A., M.A., Ph.D., LL.D . Syracuse 1983 Harold E. Newcomb, B.A . Owego 1981 Theodore M. Black, A.B . Sands Point President of the University and Commissioner of Education Ewald B. Nyquist Executive Deputy Commissioner of Education Gordon M. Ambach Associate Commissioner for Cultural Education Hugh M. Flick Assistant Commissioner for State Museum and Science Service John G. Broughton State Paleontologist for State Museum and Science Service Donald W. Fisher Contents Abstract 1 Introduction 2 Acknowledgments 4 Stratigraphic Setting and Environmental Framework 5 Faunas 7 Benthic Invertebrate Communities 9 Ambonychia-Modiolopsis Community 10 Nuculites-Colpomya Community 19 Dalmanella-Sowerbyella Community 20 Environmental Interpretations 23 Summary and Conclusions 27 References 3 1 Locality Register 33 Figure Page 1. Upper Ordovician (Lorraine) localities — Tug Hill Plateau 3 2. Diagrammatic reconstruction of Late Ordovician depositional regimes in north-central New York 6 3. Inferred distribution of Lorraine shelf sediments 6 4. Diagrammatic composite columnar section — Upper Lorraine Group 11 5. Detailed section of Upper Pulaski at Taberg, New York 12 6. Reconstruction of the Ambonychia-Modiolopsis Community 13 7. Faunas of the Ambonychia-Modiolopsis Community 14 8. Interference ripple marks 15 9. Rock exposure at locality 5, southeast of Pulaski 15 10. Biogenic reworking- — “ turkey tracks ” 15 11. Reconstruction of the Nuculites-Colpomya Community 16 12. Faunas of the Dalmanella-Sowerbyella and Nuculites-Colpomya Com¬ munities 18 13. Reconstruction of the Dalmanella-Sowerbyella Community 21 14. Late Ordovician communities— -north-central New York 23 15. Inferred distribution north-central New York shelf faunas 24 16. Late Ordovician communities central Appalachians — New York 29 Table 1. Abundant genera and/or species from Late Ordovician rocks, north- central New York 8 2. North-central New York Late Ordovician communities 9 3. Comparison of Late Ordovician faunal communities 17 Late Ordovician Benthic Marine Communities in North-Central New York' by Peter Bretsky2 ABSTRACT Invertebrate faunas in the shales and sandstones of the Lorraine Group (Upper Ordovician) in north-central New York provide an excellent example of well- defined onshore-to-offshore benthic marine communities. The distributional trends of the New York faunal associations and their environmental setting can, further¬ more, be traced into the central Appalachians. About 35 species dominate the Lorraine fauna, mainly trepostomatous bryo- zoans, brachiopods, gastropods, bivalve molluscs, crinoids, and trilobites. Three benthic marine communities are recognized on the basis of species which con¬ sistently occur together. These communities are: (1) Dalmanella-Sowerbyella Community: widespread, lived on muddy silts of the outer infralittoral zone; abundant dalmanellid (Dalmanella), strophomenid (Sowerbyella, Rafinesquina) , and atrypid (Catazyga) brachiopods, crinoids, trepo¬ stomatous bryozoans (Hallopora), and trilobites (Flexicalymene, Cryptolithus) with locally abundant pleurotomariid (Lophospira and Ruedemannia) gastropods. (2) Nuculites-Colpomya Community: widespread, lived on or buried in silts and fine sands of the outer and inner infralittoral zone protected environments; abundant nuculoid (Nuculites, Ctenodonta?), modiomorphid (Coipomya) and actinodontoid (Lyrodesma) bivalve molluscs, crinoids, and orthid brachiopods (Glyptorthis). (3) Ambonychia-Modiolopsis Community: patchy distribution, lived on or partially buried in medium to coarse sands of the shallow infralittoral and inter¬ tidal exposed environments; abundant ambonychiid (Ambonychia), modiomorphid (Modiolopsis), and cyrtodontid (Cyrtodonta, Ischyrodonta) bivalve molluscs, and monoplacophorans (Cyrtolites) with locally abundant pleurotomariid (Clathrospira) and belle rophontid (Sinuites) gastropods, crinoids, trepostomatous bryozoans (De- kayia), and orthoconic cephalopods. The pattern of offshore brachiopod-dominated communities transitional shore¬ ward into molluscan-dominated communities shown by these benthic marine faunas from New York also is characteristic of Late Ordovician deposits in the central Appalachians. The zoogeographic distribution of major community types is correlated with the position of a major deltaic complex in central Pennsylvania. ^Manuscript submitted for publication January 29, 1970. ^Department of Geological Sciences, Northwestern University, Evanston, Illinois. Introduction Fossiliferous shales, siltstones and sandstones of the Lorraine Group (Upper Ordovician) crop out along the fringes of the Tug Hill Plateau (=Uplift) in north- central New York (fig. 1, insert). Located between the Black River Valley on the east and Lake Ontario on the west, the Tug Hill Plateau has been an area of classic paleontological investigations dating from the reports of Conrad (1839) and Hall (1847). These studies provided the basis for the more expanded in¬ vestigations of Foerste (1914, 1916) and Ruedemann (1925a, b; 1926a, b). Since the 1920’s, however, little work has been done on the Upper Ordovician rocks of north-central New York except for some re¬ gional reconnaissance prior to publication of the 1961 New York State geological map (Fisher and others, 1961). The present investigation concentrates on the zoo¬ geographic distribution and ecology of the Late Ordo¬ vician benthic marine invertebrates in north-central New York. In addition ecological comparisons are made with Late Ordovician invertebrate communities in the central Appalachians (Bretsky, 1969) . The first and second generation studies in New York were con¬ cerned, of course, mainly with the description and subsequent , identification of Late Ordovician species and the main stress was placed on using these newly compiled species lists for the interpretation of bio- stratigraphic relationships between north-central New York and the Cincinnatian type area. This is not to say that other aspects of the fauna were totally ne¬ glected; in fact, Foerste (1924) and Ruedemann (1925a, 1926a) briefly outlined New York Ordo¬ vician paleogeography and paleoecology. Furthermore, Foerste (see especially 1924, p. 32) discussed pos¬ sible environmental controls on the distribution of Late Ordovician New York species. The fossiliferous units studied in this report are referred to the Lorraine Group of Emmons (1842; emended by Ruedemann, 1925a), which is composed of two main intergrading lithologic units; an upper medium-bedded fine-grained sandstone, siltstone, and shale — the Pulaski Shale (Vanuxem, 1840) and an underlying thin-bedded shaly siltstone and shale — the Whetstone Gulf Formation (Ruedemann, 1925a). No major break in bedding or lithology is evident throughout the Lorraine Group, though there is a noticeable increase upward in the amount of arenace¬ ous material, as well as in the thickness and irregu¬ larity of bedding. In fact the overlying Oswego Sand¬ stone can be distinguished from the upper Lorraine strata only because the progressive increase in arena¬ ceous content is at the expense of shale and siltstone intercalations. The Oswego has been characterized as being unfossiliferous, and unfossiliferous Oswego-like sandstones also are common in the upper parts of the Pulaski Shale. These lithologic gradations, as well as the lack of continuous exposures, have caused the definition of these lithologic units to lack precision. The reader is referred to Ruedemann (1925a) and Foerste (1916) for reviews of the earlier strati¬ graphic nomenclatural difficulties. The present study of the distribution and ecology of benthic marine invertebrates is restricted strati- graphically to the Pulaski Shale and some portions of the upper Whetstone Gulf Formation. The total thickness of the fossiliferous Upper Ordovician strata investigated amounts to about 400 feet (122 meters), whereas the overall thickness of the Lorraine may be as much as 2000 feet (609 meters) . The most com¬ plete Tug Hill exposures are normally less than 20 to 30 feet (6 to 9 meters) thick and not closely spaced; because of this and the monotonous and gradational lithologic sequence, precise tracing of individual beds is unlikely. Thus the estimate of 400 feet was pieced together from about 50 localities (fig. 1) . Study of the Lorraine fauna was begun along the southwestern fringe of Tug Hill (near Pulaski and Bennett Bridge, localities 4, 5, 6, 6A, and 7) where the fossiliferous strata crop out beneath the massively bedded, unfossiliferous, orthoquartzitic Oswego Sand¬ stone. The faunal assemblage was traced northward and eastward around the plateau, terminating at ex¬ posures along the southeastern fringe of Tug Hill near Taberg, Lee Center, and Starr Hill (localities 33, 36, and 37). About 35 abundant species dominate the Pulaski and upper Whetstone Gulf faunas, which are com¬ posed mainly of trepostomatous bryozoans, crinoids, brachiopods, and bivalve molluscs, associated with lesser numbers of monoplacophorans, gastropods, and trilobites. The dalmanellid and strophomenid brachio- pc^s, modiomorphid bivalve molluscs, and trilobites of the Lorraine Group have been correlated by Ruede¬ mann (1925a), Foerste (1914), and Ulrich (1911, 2 Figure 1. Locality map. Numerals represent localities where Lorraine Group strata are exposed and in place (see locality register, p. 33) . Exposures of glacial drift containing Lorraine Group fossiliferous boulders are not indicated on this figure. 1913) with strata of Eden and lower Maysville age in the Cincinnati region. Furthermore, Ruedemann (1925a, p. 122-138) has divided the Lorraine Group fauna into six assemblage [?] zones, some of which are found only at the classic Lorraine Gulf sections (my localities 12 to 18, 20; fig. 1); other zones ap¬ parently are correlative with Eden and Maysville beds near Cincinnati. I have interpreted the Pulaski and upper Whetstone Gulf fauna as comprising three benthic marine invertebrate communities which can be traced south of central New York and are ecologically compatible with other Late Ordovician faunas from central Pennsylvania and northern Virginia (Bretsky, 1969). The three New York communities are defined 3 on the basis of the recurrence of abundant species throughout the Tug Hill region, and are interpreted as having occupied relative onshore to offshore posi¬ tions; the bathymetric analysis is based primarily on stratigraphic position, zoogeographic distribution and autecology of the fauna as well as primary sedi¬ mentary and biogenic structures. It is likely that the age of the Pulaski Shale is not everywhere the same and, in fact, the entire Lorraine Group may become progressively younger to the west (Fisher, 1962). Although I believe that unequivocal paleontological evidence for this age differentiation is lacking, the hypothesis seems probable in view of the Late Ordovician stratigraphic-sedimentological setting which emphasizes a westward progradation of the Queenston deltaic complex. The Pulaski and upper Whetstone Gulf thus are interpreted as westwardly time-transgressive, nearshore marine phases within this deltaic setting. ACKNOWLEDGMENTS I am indebted to my colleague Laurence L. Sloss at Northwestern University and Donald W. Fisher, New York State Museum for reading the manuscript and offering helpful suggestions. I also appreciate the care spent in drafting the text figures by Miss Pauline Mohr and the typing of numerous drafts of this manuscript by Miss Linnea Danchi. Special thanks are due Mr. Richard L. Snyder, Adams, New York, whose knowledge of local geography helped the author in locating several rock exposures, and Mr. Robert Iredale of the Niagara-Mohawk Power Company, Pulaski, for permission to examine an exposure in the channel at the Red Brick Power Station, Bennett Bridge. The project was supported in part by research grant No. 443-07 from Northwestern University. Stratigraphic Setting and Environmental Framework Lorraine Group rocks studied in this report crop out around the western, northern, and eastern fringes of the Tug Hill Plateau, which is capped by the Oswego Sandstone and presumably some isolated out¬ liers of the Queenston Formation (fig. 1) . A thick sequence of Trenton Group limestones and shales underlie the Lorraine (Fisher, 1962 for stratigraphic details) . Fossils were collected from only the upper part of the Lorraine Group, specifically the Pulaski Shale and the uppermost portions of the Whetstone Gulf Formation. The total thickness of the Upper Ordovician section examined is estimated to be about 400 feet (122 meters) . The exact position of the Pul- aski-Whetstone Gulf boundary is difficult to place because of a gradual lithologic transition from the dominantly thin-bedded siltstones and shales of the Whetstone Gulf into the overlying medium-bedded sandstones, siltstones, and shales of the Pulaski. This lithologic gradation is accompanied by a gradual change in the faunal content of these rocks; the fossils closely resemble Eden and lower Maysville species in the Cincinnati region. The strata of the Lorraine Group constitute part of the major Late Ordovician Queenston deltaic com¬ plex which had its origin in a progressive uplift that took place along the present eastern seaboard of the United States from New Hampshire to Virginia (Kay and Colbert, 1964, figs. 9 and 21). The Late Ordo¬ vician regression is well documented from sedimento- logical studies in the central Appalachians (McBride, 1962; Yeakel, 1962) and eastern Canada (Mason, 1967). These Late Ordovician sediments, in all cases, were transported in a generally westward direction, giving rise in central New York to the Queenston clastic wedge (King, 1959, p. 61, fig. 34; for an in¬ formative and concise review of the Queenston clastic wedge see Clark and Steam, 1968, p. 286). The Late Ordovician regression in New York is documented by an upward coarsening of sediments and progressive increase in bedding thickness; primary sedimentary structures emphasize a progressive shallowing trend that culminated in the deposition of the red shales and sandstones of the Queenston Formation. These red beds represent migrating deltaic deposits laid down on a broad alluvial plain. This apron of predominantly terrestrial red shales and sandstones graded westward into the terrestrial and shallow marine (supratidal?) virtually abiotic Oswego sandstones which, in turn, passed into the shallow marine faunally-rich Pulaski sandstones, siltstones, and shales. The shales and silt¬ stones of the underlying Whetstone Gulf Formation represent progressively deeper marine deposits. Thus the Late Ordovician environments in central New York represent a series of transitional marine-ter¬ restrial depositional regimes that exhibit an overall east-to-west regressive pattern (see Fisher, 1962, five left-hand columns, Niagara Falls through Rome Quad¬ rangles for east-west interpretation cross-section) ; dur¬ ing any given instant of time the eastern subaerial and western subaqueous environments would have existed simultaneously (fig. 2; see Grabau, 1913; Broughton and others, 1962, p. 20). Mason (1967) reached similar conclusions from a sedimentological study of Upper Ordovician rocks along the Chambly- Fortierville Syncline in Quebec. The sediments of the Pulaski Shale were deposited in nearshore marine regimes, most likely inner in¬ fralittoral, perhaps in part intertidal, whereas the sedi¬ ments of the Whetstone Gulf Formation were laid down in deeper water, outer infralittoral regions (fig. 3) . Changes in the faunal content of these sediments take place very gradually and are accompanied by numerous repetitions of faunal associations, thereby emphasizing the periodic cycling of benthic marine con¬ ditions that favored one faunal assemblage then another on a gently sloping submarine shelf. Bayer (1967) has documented somewhat similar Late Ordovician bottom conditions in southeastern Minnesota, where repetitious changes in sediment influx and bottom chemistry led to periodic changes in the benthic fauna. 5 DIAGRAMMATIC RECONSTRUCTION Figure 2. Diagrammatic reconstruction of Late Ordovician depositional regimes in north-central New York; interpretation adapted from Broughton et al. 1962. LATE ORDOVICIAN REGRESSIVE PATTERN Figure 3. Inferred distribution of Lorraine shelf sediments from stratigraphic and sedimentological evidence. The sinuosity of the Oswego-Pulaski and Pulaski- Whetstone Gulf boundaries is a diagrammatic representation of the very grada¬ tional and recurrent nature of these lithologies. It is important to note that the diagram portrays lithologic distributions at any one time during the Late Ordovician in north-central New York. Present-day reference points are included specifically for scale and do not imply present geographic relationships to the distribution of Ordovician rocks. 6 Faunas Numerically dominant taxonomic groups in the cen¬ tral New York Upper Ordovician fauna are treposto- matous bryozoans, brachiopods, bivalve molluscs, and crinoids, with lesser numbers of trilobites, gastropods, and monoplacophorans; the latter three taxa are nor¬ mally more restricted geographically, and presumably ecologically, than the dominant taxa. The fossils are distributed throughout the Lorraine Group, but are most often concentrated in lenses and layers. In fact, crossbedded crinoid-bryozoan coquinites are present in the upper parts of the Pulaski Shale. Table 1 is a list of the abundant genera and / or species in the Pulaski and upper parts of the Whetstone Gulf Formation. Although only about 35 taxa are listed, it is prob¬ able that there are more than 100 species of macro¬ fauna preserved in these Late Ordovician sediments. I would expect that a percentage of abundant to total macrofaunal species, if precisely documented, would fall between 10 and 25 percent. A scale of relative taxonomic diversity was employed, however, in this introductory study and applied at each local¬ ity where areal exposure and preservation deemed it applicable. Taxa were thus identified in the field, relative abundances being recorded at the locality along with stratigraphic relationships and associated lithology. These preliminary results were refined in the laboratory and then again re-examined in the field. Although vertical and horizontal trends in species com¬ position were evident, the observed density of some species commonly varied from one to another of closely spaced localities. I suspect that this variation not only reflects actual differences in species diversity, but also is, to some extent, an artifact of areal extent of the exposures. What is important, however, is the consistent stratigraphic and zoogeographic occurrence of the same species. Utilizing this systematic occur¬ rence these Late Ordovician faunal communities can be outlined. 7 Table 1. Abundant genera and/or species from Late Ordovician rocks, north-central New York BRYOZOA: Dekayia Hallopora ?Heterotrypa BIVALVIA: Nuculites “scitulus” Lyrodesma poststriatum Ctenodonta? cf. pulchella Colpomya faba Ambonychia radiata Ambonychia praecursa Modiolopsis modiolaris Ischyrodonta unionoides BRACHIOPODA: Lingulapholis? Dalmanella multisecta Sowerbyella (Sowerbyella) sericea Rafinesquina “altemata” Rafinesquina mucronota? Glyp tort his crispata Catazyga erratica Zygospira modesta Cyrtodonta Cyrtodontula MONOPLACOPHORA: Archinacella Cyrtolites omatus GASTROPODA; Sinuites? Clathrospira subconica Eotomaria? Ruedemannia? lirata Loxoplocus (Lophospira) abbreviata Loxoplocus (Lophospira) ventricosta Loxoplocus (Lophospira) bowdeni Hormotoma gracilis CRINOIDEA: columnals TRILOBITA; Llexicalymene Cryptolithus Isotelus CEPHALOPODA: orthocones CONICAL SHELLS OF UNKNOWN AFFINITY; Comulites Benthic Invertebrate Communities The consistent and recurrent associations of the abundant invertebrate species (i.e. bryozoans, brachio- pods, bivalve molluscs, gastropods, monoplacophorans, crinoids, and trilobites) has led to the recognition of three faunal associations, termed communities. These are outlined in table 2. Each of these Late Ordo¬ vician communities is named for its most conspicuous and co-dominant species. The communities are as follows: 1 . Ambonychia-Modiolopsis Community 2. Nuculites-Colpomya Community 3. Dalmanella-Sowerbyella Community TABLE These three communities are composed of groups of species all of which show a high degree of affinity and pronounced tendency to occur together through¬ out the Pulaski Shale and upper Whetstone Gulf Formation. Some species are completely restricted to one community, but normally those species listed as members of a particular community (table 2) are numerically significant in that community alone. Furthermore, this designation of communities does not imply that community boundaries, either faunally or geographically portrayed, are rigid, and that no mixing of species among communities takes place. 2. AMBONYCHIA - MODIOLOPS/S COMMUNITY DALMA NEL L A -SOWERB YELL A COMMUNITY NUCUUTES - COL POM YA COMMUNITY HALLOPORA DEKAYIA'f. DEKAY/A DALMA NELLA MULTISECT A GLYPTORTH/S CR/SPATA SOWERBYELLA (SOWE RBY E LLA) SERICEA Lt N G ULA PHOLIS ? RAFINESOUINA "ALTER NAT A" R. MUC RONOTA ? CATAZYGA ERRATIC A ZYGOSPIRA MODESTA CLATHROSP! RA SUBCONICA HORMOTOMA GRACILIS SINUITES ? LOXO PLOCUS (LOPHOSP! RA) ABBREVIATA LOXO PLOCUS (LOPHOSPIRA) BOWDEN! LOXOPLOCUS (LOPHOSPIRA) VENT RICOSTA RUEDEMA NNIA'f DR AT A AMBONYCHIA PRAECURSA AMBONYCHIA RADIATA MODIOLOPSIS MODIOLARIS CYRTODONTA ISCHYRODONTA UNIONOIDES NUCULITES "SC IT ULUS" LYRODESMA POSTSTRIATU M CTENODONTA? cf. PULCHELLA COL PO MY A FA BA FLEX I C ALY ME NE C RY PTOLITHUS ISOTELUS CYRTOLITES ORNATUS ARCH! NACELLA ORTHOCONIC CEPHALOPODS COR NULITES CR I NOI D COLUMNALS Table 2. North-central New York Late Ordovician communities 9 The genus Ambonychia Hall and the crinoids are excellent examples of significant ubiquity among com¬ munities. The stratigraphic distribution of the faunas compris¬ ing these three communities shows considerable over¬ lap but a definite gradational trend from one into another (fig. 4) . The obvious effects of post-mortem transport accounts for some of this overlap, especially in the upper parts of the Pulaski, where comminuted fragments of crinoids, bryozoans, and some brachio- pods were reworked into crossbedded coquinas. Aside from the obvious instances of post-mortem transport, evidence of abrasion is not significant and measure¬ ments of disarticulated valve ratios and of shell sizes of abundant species on bedding planes shows little evidence of significant winnowing and highly sorted shell concentrations. I think it is reasonable to state that most of the benthic fauna was preserved in the area in which it was living. Because the communities are defined on the replication of numbers of abundant species, the chance accumulation of fossil assemblages as mere artifacts of physical energy and mixing is minimized. This consistent association of certain species was the basis for Petersen’s (1914) concept of a marine bottom community. A more rigorous statistical delineation of these communities has been carried out at a few of the sample localities (Bretsky, unpub. data) . At the present time, it adds little background to this preliminary study, but this work is continu¬ ing on larger numbers of samples from the more continuously exposed Upper Ordovician Nicolet River section in the St. Lawrence Lowlands of Quebec. Each of the three benthic communities extends across the 2000 square mile Tug Hill region. Some zoogeographic differences, however, exist, but at pres¬ ent can be incorporated within the community struc¬ tures as defined. Thus a local abundance of Rafines- quina mucronota? Foerste along the northern fringes, or of Loxoplocus (Lophospira) Whitfield and Ruede- mannia Foerste in the southeast, are more simply in¬ corporated into the Dalmanella-Sowerbyella Com¬ munity and do not merit a separate designation. These and other examples are discussed in the following sections. Ambonychia-Modiolopsis Community (fig. 6) The fauna of the Ambonychia-Modiolopsis Com¬ munity (table 2) is exposed only along the south¬ eastern (fig. 1, localities 33, 34, and 35) and south¬ western (localities 5, 6-A, and 7) portions of Tug Hill Plateau, but the Ambonychia-Modiolopsis faunas are found in numerous glacial erratics to the north of the present exposures. Thus I suspect that this community was more widespread than would be in¬ ferred from present exposures. The community is dom¬ inated by large epifaunal suspension-feeding bivalve molluscs, associated with somewhat lesser numbers of detritus-feeding gastropods and monoplacophorans, (fig. 7, no. 1-4) . Suspension feeders consist of the mussel-like species Modiolopsis modiolaris (Conrad), Ambonychia praecursa (Ulrich), Cyrtodonta Billings, and Ischyrodonta unionoides (Meek) , whereas the most abundant detritus feeders are the coiled monoplaco- phoran Cyrtolites omatus Conrad and pleurotomariid gastropod Clathrospira subconica (Hall) . Orthoconic cephalopods are found in large numbers only at lo¬ cality 33 (East Branch of Fish Creek) where several bedding planes of south-eastward oriented orthocones are exposed immediately above the stream bed. While specimens of Ambonychia, Modiolopsis, Ischyrodonta, and Cyrtolites are most abundant in the west, Modio¬ lopsis and Clathrospira are more common in the south¬ eastern exposures. Cyrtolites appears much more common in weathered glacial erratics, and thus may have been dominant to the north. Most of the previously described and photographed species that are now included in the Ambonychia- Modiolopsis Community came from weathered glacial erratics uncovered in the vicinity of Trenton Falls, where bedrock are limestones of the Middle Ordo¬ vician Black River or Trenton Groups. Actually, these species were found to be much more abundant in glacial boulders from the vicinity of Pulaski and, in general, from the more southwestern portions of Tug Hill. Furthermore, fine specimens can be obtained from partially weathered exposures of Lorraine bed¬ rock. The fossils of the Ambonychia-Modiolopsis Com¬ munity are common in thin- to medium-bedded, fine- to medium-grained sandstones, which are interbedded with irregularly crossbedded coarse orthoquartzites and dark grey-black silty shales. The coarse tabular and planar crossbedded sandstone lenses commonly thicken from 5 inches to 3 feet over a lateral distance of less than 6 feet (fig. 5) . Thinner, but similarly crossbedded or laminated, crinoid-bryozoan, sand-pebble conglom¬ erates are generally enclosed within or occur at the base of orthoquartzites. The orthoquartzites alternate and intertongue with dark grey-black silty shales that are commonly mottled, less commonly fissile and con¬ tain a patchy fauna composed of small-sized taxodont, modiomorphid, and desmodont bivalve molluscs, grap- tolite fragments and more rarely long, oriented crinoid stems. This fauna is not considered part of the Am- 10 HETSTONE ^ GULF I PULASKI . OSWEGO DIAGRAMMATIC COMPOSITE COLUMNAR SECTION UPPER LORRAINE GROUP DESCR IPTION Massive unfossllif erous orthoquartzites, few thin black shales. Shale pebble conglomerate, Irregularly bedded sandstones, siltstones and shales. Crossbedded sandstones (tabular, planar) with intercalated bryozoan-crinoid coquinlte. Cut and fill. "U"-shaped tubes and "turkey Interference ripple marks. Small scale interference ripple marks. rhin brachiopod coqulnites very common. Meandering surficial biogenic reworking. Trilobites more common; occasionally only a sparse trilobite (Triarthus)- graptolite fauna. Figure 4. Diagrammatic composite columnar section Upper Lorraine Group in central New York showing the vertical distribution of the three benthic marine communities and a short description of the associated lithologies, primary sedi¬ mentary structures and biogenic structures. 11 DETAILED SECTION (LOCALITY 33) OF UPPER PULASKI AT TABERG, N.Y. Figure 5. Detailed section of Upper Pulaski at Taberg, New York (locality 33) . Of particular interest are not only the faunal associations but also the extreme irregularity in bedding and lithology. Coarse crossbedded sandstones thicken and thin by several orders of magnitude over very short distances. 12 AMBONYCHIA-MODIOLOPSIS COMMUNITY Figure 6. A reconstruction of the Ambonychia-Modiolopsis Community. The community is dominated by large epifaunal suspension-feeding bivalve molluscs, AmbonycMa praecursa (A) , Modiolopsis modiolaris (M) and Ischyrodonta union- ides (I) which are associated with detritus-feeding gastropods, Clathrospira sub- conica (C) and Hormotoma gracilis (H) , and the coiled monoplacophoran Cyrto- lites omatus (Cy) . Excluded from this reconstruction are the common burrows, grazing and/or resting tracks (see fig. 5 and 10). bonychia-Modiolopsis Community, but actually consti¬ tutes the partial remains of another faunally distinct community. The oriented crinoid stems are especially common at the base of the silty shales overlying the massive crossbedded orthoquartzites. Figure 5 diagrams in partial detail a stream section from Taberg (local¬ ity 33) that shows the stratigraphic relationships of these various lithologies. In the midst of the grey- black silty shales occur fine- to medium-grained sand¬ stones that contain elements of the Ambonychia-Modio¬ lopsis Community. These fossiliferous sandstones and silty shales are cut by shallow channels 2 to 3 inches wide and normally less than 1 inch deep, that are filled with crinoid-bryozoan debris. Additional current reworking of the bottom sediments is evident in the large-scale flattened interference ripples (fig. 8) and the disarticulated crinoid stems and worn frag¬ ments of trepostomatous bryozoans in the sand-pebble coquinites. Thus the overall stratigraphic sequence is characterized by extreme lithologic irregularities and little gradation of one rock unit into another (fig. 9) . Lithologic variation is rather abrupt and includes lens¬ like crossbedded orthoquartzites and numerous shallow cut-and-fill structures. The entire sequence grades upward into the unfossiliferous orthoquartzites and shale pebble conglomerates of the Oswego Sandstone. The extreme irregularity in bedding is accompanied by numerous burrows, grazing and/or resting tracks (also fig. 5) . Evidences of biogenic reworking in the upper part of the Pulaski can be grouped into three main morphological types that occur in a partichlar stratigraphic order and normally within specific litho- 13 Figure 7. Faunas of the Ambonychia-Modiolopsis Community. 1. — Modiolopsis modiolaris, composite mold natural XI (locality 33). 2. — Ambonychia praecursa, latex impression of left valve XI (locality 41). 3. — Cyrtolites omatus, latex impression X2 (locality 29) . 4. — Clathrospira subconica, latex impression X2 (glacial drift) . logics. The most numerous and widespread biogenic structures are burrows or grazing tracks which form a pattern of 3 or 4 broad, blunt, shallow finger-like projections on bedding planes (fig. 10); I have seen this type of biogenic structure, called “ turkey tracks ” by workers in the Cincinnati region, in the Corry- ville Member of the McMillan Formation at the Stone- lick Creek section about 20 miles east of Cincinnati (also see Hofmann, 1964 for locality data). “ Turkey tracks ” are most common in New York on the bed¬ ding planes of coarse sandstones that alternate with flattened interference-rippled sandstones. Shinn (1968, pi. 110, fig. 2) and Schafer (1962, fig. 52) photo¬ graphed present-day crustacean burrows and bivalve mollusc grazing patterns that appear somewhat ana¬ logous morphologically to these Ordovician “ turkey tracks.” A second, less common biogenic structure consists of longitudinally striated surface burrows that inter¬ sect bedding planes at low angles and which are 4 to 5 inches long and about V2 inch wide; consid¬ erably deeper than the “ turkey tracks ” and in some places filled with crinoid-bryozoan debris (fig. 12, no. 12). These striated burrows are very similar to some of the Cruziana facies burrows of Seilacher (1964 see esp. fig. 8, showing basal parts of Diplocraterion) , and occur in mottled shaly sandstones with patches of abundant Ambonychia and Cyrtolites; therefore, they may represent a soft-bodied member of the Am¬ bonychia-Modiolopsis Community. The third type of biogenic structure is least com¬ mon and consists of “ U ’’-shaped tubes that are found only in the massive-bedded orthoquartzites. The tubes, which are filled with the dark grey silty shales, 14 Figure 9. Rock exposure at locality 5, southeast of Pulaski Seilacher’s Cruziana facies (1964, p. 309-310) and Schafer’s U-Bau von Arenicola (1962, Abh. 174, 175) are morphologically very similar. Stratigraphically the “ U ’’-shaped tubes are the highest of all three bio¬ genic structures and presumably may cross the tran¬ sitional Pulaski-Oswego boundary. As outlined in figures 4 and 5, the faunas of the Am- boychia-Modiolopsis Community are interbedded and in some places mixed with other species believed to be more diagnostic of other communities. The other species are small taxodont, modimorphid, and des- modont bivalve molluscs which are of patchy abund¬ ance throughout the grey black silty shales. These small bivalve mollusc species constitute a major part of another faunal assemblage, the Nuculites-Col- pomya Community, which appears to have occupied part of the same geographic area as the fauna of the Ambonychia-Modiolopsis Community. A third faunal assemblage is composed of fragmented remains of the Figure 8. Interference ripple marks Figure 10. Biogenic reworking “ turkey tracks ” outline a dumbbell shape on the bedding plane (the dumbbells are normally IVi to 4 inches apart; see Ruedemann, 1926b, p. 81 and Foerste, 1924, p. 78); they were called Arthraria Billings by these early in¬ vestigators. The tubes have been found only in the coarse unfossiliferous Oswego-like sandstones that are interbedded with the fossiliferous shaly sandstones, siltstones, and shales of the Pulaski. Corophioides in 15 brachiopods Dalmanella and Sowerbyella, crinoids, and bryozoans, and forms the thin coquinites. Some of these species, especially the crinoids and bryozoans, are occasionally observed mixed with the faunas of the Ambonychia-Modiolopsis Community but normally form distinct layers and lenses of shell debris. The local abundance, however, is evidence of the reason¬ able proximity of a third distinct faunal assemblage — the Dalmanella-Sowerbyella Community; but because the species occur abundantly only in the coquinite and show signs of abrasion and size sorting, it is probable that no significant portions of the Dal¬ manella-Sowerbyella Community lived in this particu¬ lar area in which they are now preserved. Bretsky (1969) outlined a similar Late Ordovician faunal association in the central Appalachians termed the Orthorhynchula-Ambonychia Community, which is common at the top of the Reedsville Formation from south-central Pennsylvania into central Virginia. The similarities that exist between the New York and central Appalachian communities are evident not only in the number of dominant taxa common to both (table 3) , but also in quite analogous strati- graphic-sedimentologic situations. The faunas of both communities are abundantly represented in fine- to medium-grained sandstones interbedded with crossbed- ded orthoquartzites and grey-black laminated shales, a succession which represents a shallow marine de- positional regime within the Late Ordovician eastern North America regressional sequence; differences are evident in that the shales are less abundant and un- fossiliferous in the central Appalachians and the cri- noid-bryozoan coquinite is common only in New York. Species of the large epifaunal suspension-feeding bi¬ valves Ambonychia, Modiolopsis, and Ischyrodonta are abundant in both communities but faunal differences are evident beyond this comparison. The most appar¬ ent difference is that the Ambonychia-Modiolopsis Com¬ munity of New York contains no extensive brachio- pod fauna such as the extremely abundant Lingula? NUCULITES-COLPOMYA COMMUNITY Figure 11. A reconstruction of the A^wcw/Zto-Co/pomja Community. The com¬ munity is composed of predominantly small infaunal detritus feeding bivalve molluscs, Nuculites “ scitulus ” (N) and Ctenodonta? cf. pulchella (C), and an infaunal suspension (?) feeding bivalve, Lyrodesma poststriatum (Ly) . There are two numerically important epifaunal suspension feeders, the small modio- morphid bivalve mollusc, Colpomya faba (Co) and the orthid brachiopod, Glypt- orthis crispata (G) . The detritus feeding, epifaunal cap-shell monoplacophoran, Archinacella (A), is common at some locales. 16 APPALACHIAN COMMUNITIES ABUNDANT SPECIES NEW YORK COMMUNITIES 17 Figure 12 18 Figure 12. — Faunas of the Dalmanella-Sowerbyella and Nuculites=Colpomya Communities. 1. — Ruedemannial lirata, latex impression X2 (locality 37). 2. — Sowerbyella (Sowerbyella) sericea, pedicle valve internal mold natural X2 (glacial drift) . 3. — Dalmanella multisecta, latex impression of pedicle valve external X2 (locality 16). 4. — Catazyga erratica, brachial valve external X2 (glacial drift) . 5. — Loxoplocus (Lophospira) bowdeni, latex im¬ pression X2 (glacial drift) . 6. — Dalmanella multisecta, latex impression brachial valve internal X2 (locality 39) . 7. — Nuculites ” scitulus, ” composite mold natural X2 (locality 33). 8. — Lyrodesma poststriatum, internal mold natural X2 (glacial drift). 9. — Glyptorthis crispata, latex impression pedicle valve external X2 (locality 3). 10. — Nuculites “scitulus,” internal mold natural X2 (locality 41). 11. — Colpomya faba, latex impression X2 (local¬ ity 14). 12. — Longitudinally striated “worm” burrow XI (locality 7) . Brugiere and Orthorhynchula linneyi (James) in the central Appalachians. Bellerophontid gastropods (Plec- tonotus? Clarke and some Bucania Hall) common in the central Appalachians appear to be absent in New York and perhaps were replaced in part by the large coiled monoplacophoran Cyrtolites and the pleu- rotomariid gastropod Clathrospira. The protobranch bi¬ valve mollusc Tancrediopsis cuneata (Hall) is also entirely missing from the New York community but is ubiquitous throughout the central Appalachians. Thus, the most significant differences are the absence of the Tancrediopsis-Lingula?-Plectonotus and Orth¬ orhynchula linneyi fauna in New York (Bretsky, 1969, figs. 5 and 6), and conversely the abundance of Cyrtolites only in New York. Despite these faunal differences, however, the two communities are com¬ parable in their overall dominance by large epifaunal, suspension feeding bivalve molluscs, and, of course, their environmental setting. Nuculites-Colpomya Community (fig. 11) The fauna of the Nuculites-Colpomya Community (table 2) is exposed around the western, northern, and eastern fringes of the Tug Hill Plateau (local¬ ities 7, 14, 17, 20, 33, and 41). The community is composed of predominantly small infaunal detritus and suspension (?) feeding bivalve molluscs (fig. 12, no. 7-11). The detritus feeders consist of the nuculoid bivalves Nuculites “scitulus ” Emmons and Ctenodonta? cf. pulchella (Hall) with lesser numbers of Praenucula Pfab and Palaeoconcha Miller. Infaunal suspension (?) feeders are less numerous and include the actinodontid bivalve Lyrodesma poststriatum (Emmons) and assorted desmodonts-Rhytimya Ul¬ rich, Cuneamya Hall and Whitfield, Cymatonota Ulrich, and Psiloconcha Ulrich. In addition, there are two numerically important epifaunal suspension feed¬ ers, the small modiomorphid bivalve Colpomya faba (Emmons) and the orthid brachiopod Glyptorthis crispata (Emmons) . Some of these species are not widely distributed; Lyrodesma poststriatum is abund¬ ant only in the west, Glyptorthis crispata is common only in the north, whereas the desmodont bivalve molluscs are more commonly found in the southeast. The fossils of the Nuculites-Colpomya Community are common in silty shales and siltstones, which are high in the Pulaski Shale section and are sporadically interbedded with irregularly, crossbedded orthoquartz¬ ites (see especially localities 7 and 33). Other silt- stones and shales, however, contain the Nuculites- Colpomya fauna and are found lower in the strati¬ graphic section, where the bedding is thinner and more 19 regular, the shales are fissile, and the fine sandstones sporadically exhibit only small symmetrical ripple marks (see especially localities 14, 17, 20, and 41). The faunas of the Nuculites-Colpomya Community thus extend over a broader stratigraphic range within the Pulaski Shale than do the faunas of the Am- bonychia-Modiolopsis Community (fig. 4) . Differences in the faunal composition of the Nuculites-Colpomya Community do exist stratigraphically, as well as zoo- geographically, although an overall consistency is main¬ tained; Nuculites and Colpomya are abundant through¬ out, but there is a trend toward increased numbers of smaller-sized (3 to 10 mm) Nuculites lower in the section whereas larger (1.5 to 2.0 cm) ones exist higher up (fig. 12, no. 7, 10). Lyrodesma post¬ striatum and the desmodont bivalve molluscs are more abundant in the upper Pulaski, whereas Glyptorthis crispata is more abundant lower in the section. The fossils of the Nuculites-Colpomya Community are, of course, associated with other species more rep¬ resentative of the other two communities. High in the Pulaski section, remnants of the Ambonychia- Modiolopsis Community, especially Ischyrodonta, Am- bonychia, and a few Modiolopsis are mixed with the dominant Nuculites-Colpomya fauna, but the faunas of the two communities are normally interbedded and mixing would be expected since the two communities presumably occupied part of the same geographic area. Occasionally, long-stemmed crinoids without calyces and graptolite fragments are found with the abundant species of the Nuculites-Colpomya Com¬ munity, but crinoid columns and graptolites (uniden¬ tified) are few in number and are believed to be derived from outside the immediate area. Lower in the Pulaski section the Nuculites-Colpomya fauna is interbedded with the Dalmanella-Sowerbyella fauna, and as might be expected, some mixing of these faunas is common. Crinoids become more commonly mixed with the Nuculites-Colpomya fauna as well as scarce trilobites, dalmanellid brachiopods, and a few trepo- stomatous bryozoans. Species which are numerically significant in the Nuculites-Colpomya Community are either not present or occur in much lesser numbers in the central Ap¬ palachians. Small patches of nuculoid bivalves and Lyrodesma poststriatum, however, occur in north- central Pennsylvania. These forms were, because of their scarcity, incorporated into the central Ap¬ palachian Sowerbyella-Onniella Community (Bretsky, 1969). Specimens of Nuculites are insignificant, and Colpomya was not found, in the north-central Appa¬ lachians, but the nuculoids Ctenodonta? and Prae- nucula, as well as Lyrodesma poststriatum, are com¬ mon in some places in a regularly bedded siltstone associated primarily with the dominant dalmanellid- strophomenid-crinoid and trilobite fauna of the central Appalachian Sowerbyella-Onniella Community. Because of the increased development of the protobranch fauna to the north, the protobranch-rich assemblage is no longer considered part of the strophomenid brachio- pod community as in the north-central Appalachians but now stands as a separate major faunal assemblage in north-central New York. At the present time there is no evidence for the development of such a com¬ munity south of north-central Pennsylvania. The pro¬ tobranch bivalve and Lyrodesma fauna is strictly a north-central Appalachian and New York Late Ordo¬ vician development. Dalmanella-Sowerbyella Community (fig. 13) The fauna of the Dalmanella-Sowerbyella Com¬ munity (table 2) encircles the Tug Hill Plateau and is very common in weathered glacial boulders. Ex¬ cellent exposures yielding the faunas of the Dalma¬ nella-Sowerbyella Community are seen at localities 3, 4, 11, 14, 16, 17, 19, 20, and 41. The approximate stratigraphic position of these exposures is indicated on figure 4. The community is dominated by large numbers of epifaunal suspension feeding brachiopods and bryozoans (fig. 12, nos. 1-6). Orthid [Dalma- nella multisecta (Meek)] , and strophomenid {Sow- erbyella (Sowerbyella) sericea (Sowerby)] brachio¬ pods are most abundant, in addition to trepostomatous bryozoans Hallopora Bassler and Dekayia Milne-Ed- wards and Haime, and crinoids. Less common over¬ all, but locally highly concentrated in patches, are strophomenid, [Rafinesquina “ altemata ” (Hall) and R. mucronota? Foerste] and atrypid {Catazyga er- ratica (Hall) and Zygospira modesta (Hall)] brachio¬ pods, and species of Loxoplocus (Lophospira) Whit¬ field and Ruedemannia Foerste. Specimens of Ca¬ tazyga and Rafinesquina are found in layers and lenses along the northern portions of Tug Hill, espe¬ cially in the Lorraine Gulf section, and also appear much more frequently in drift blocks than would be expected from their present limited areal exposure (also see Ruedemann, 1925a, p. 122 and 134). Am- bonychia radiata? Hall and the trilobites Flexicaly- mene Shirley and Cryptolithus Green are ubiquitous but less numerous than the brachiopods and crinoids. The fossils of the Dalmanella-Sowerbyella Com¬ munity are common in thin-bedded calcareous silt- stones and shales. Bedding is regular throughout, but there is a definite tendency for thicker and more 20 DALMANELLA-SOWERBYELLA COMMUNITY Figure 13. — A reconstruction of the Dalmanella-Sowerbyella Community. The community is dominated by large numbers of epifaunal suspension feeding brachiopods, Dalmanella multisecta (D), Sowerbyella (Sowerbyella) sericea (S), Rafinesquina “ altemata ” (R), Catazyga erratica (C), and trepostomatous bryozoans, Hallopora (H) . In addition there are less common crinoids (Cr), three species of Loxoplocus (Lophospira) (L) and trilobites, Flexicalymene (F) . numerous calcareous beds to occur higher in the sec¬ tion. Fragmented specimens, especially crinoids and bryozoans, are common high in the Pulaski section, interbedded with the massive Oswego-like ortho¬ quartzites and the siltstones and shales bearing faunas of the Ambonychia-Modiolopsis and Nuculites-Col- pomya Communities. These reworked crinoid-bryo- zoan lenses continue downward in the Pulaski sec¬ tion for a short distance, but rapidly give way to fossiliferous siltstones that show little evidence of the higher physical energy present in the overlying beds. The regularly thin-bedded siltstones are interbedded with gray-green silty and non-silty shales; within this regular alteration there is a gradient toward increased shale in the middle and lower portions of the Pulaski Shale as well as the upper parts of the Whetstone Gulf Formation. The fossiliferous silts finally grade downward into fissile gray-black organically rich shales and a few thin-bedded silty shales that are dominated by a sparse trilobite (Triarthus Green) -graptolite fauna. The extremely regular, thin-bedded siltstones are marked in places by small symmetrical interference ripples (localities 14, 16). The wave length and height of these ripples are about 1/10 the size of those found in the coarser sandstones of the upper parts of the Pulaski Shale. Biogenic reworking also varies signifi¬ cantly. Instead of the large burrowing and grazing pat¬ terns associated with the faunas of the Ambonychia- Modiolopsis Community, fine meandering patterns cover the bedding plane (see Seilacher, 1964, p. 308, for fine meandering grazing patterns in the Nereites facies) ; very rarely do these burrows penetrate from one bed into another. 21 The faunas of the Dalmanella-Sowerbyella Com¬ munity are closely associated with those of the other two communities. Crinoids and bryozoans in the upper Pulaski Shale are interbedded and in places mixed with the faunas of the Ambonychia-Modiolopsis Com¬ munity apparently constituting a part of an adventi¬ tious fossiliferous sandstone or coquinite. This brachio- pod- and crinoid-dominated community is also inter¬ bedded and mixed with the bivalve fauna of the Nu- culites-Colpomya Community, again especially through¬ out the upper and middle parts of the Pulaski Shale. The north-central New York Dalmanella-Sowerby¬ ella Community is very similar to the Sowerbyella- Onniella (=Dalmanella) Community of the central Appalachians and Shochary Ridge in eastern Penn¬ sylvania (Bretsky, 1969; Bretsky and others, 1969), and the Late Ordovician Thaerodonta-Onniella Com¬ munity in southeastern Minnesota (Bayer, 1967). Differences that exist among the faunal components in these three communities are much less significant than their similarities, especially in the domination of the dalmanellid and strophomenid brachiopods, trilo- bites, and numerous crinoids (table 3) . A very close relationship exists between the New York and central Appalachian communities, although dalmanellids are more abundant and strophomenids less common in New York. The pleurotomariid genera Loxoplocus (Lophospira) and Ruedemannia are common in the north-central Appalachians (Pennsylvania and north- central Virginia) and New York and, in both cases, they occur along^ the eastern- edges of the Upper Or¬ dovician outcrop belt. As indicated above, the Sower- byella-Onniella Community in the central Appala¬ chians included some species that belong to the New York Nuculites-Colpomya Community. Environmental Interpretations The definition and recognition of these three New York Late Ordovician communities permits a recon¬ struction of the inferred onshore to offshore com¬ munity distribution (fig. 14) , based on the stratigraphic relationships (fig. 4) , present geographic distribution of the faunas and autecological interpretations of the species. The reconstruction is drawn on the Late Or¬ dovician sedimentological fremework of a prograding depositional regime (fig. 3). Figure 15 is a profile taken across this reconstructed Late Ordovician shelf showing the inferred distribution of the abundant faunal elements from their present vertical distribu- LATE ORDOVICIAN COMMUNITIES CENTRAL NEW YORK Figure 14. — Environmental reconstruction Late Ordovician communities north-central New York based on stratigraphic, sedimentologic and faunal evi¬ dence. The zoogeographic pattern of the communities is portrayed as they may have been distributed at any given instant in time during the deposition of the Late Ordovician Lorraine sediments. As in figure 3 the present day reference points are for scale and do not imply present geographic distribution of the Late Ordovician communities. 23 INFERRED DISTRIBUTION NORTH-CENTRAL NEWYORK SHELF FAUNAS Figure 15. — Inferred distribution north-central New York shelf faunas. Refer to figure 14 for areal view of the environmental setting. Species ranges are drawn from their present vertical distribution in the Pulaski Shale and upper parts of the Whetstone Gulf. No estimate of water depth are inferred although the deepest portions of these Late Ordovician shelf environments was probably not greater than 50 to 75 meters. tion in the Pulaski Shale and upper parts of the Whetstone Gulf Formation. This figure emphasizes the pronounced nearshore-to-offshore change in the abundant faunas, although there is considerable mix¬ ing and overlap of the individual species. The fauna of the Dalmanella-Sowerbyella Com¬ munity was probably outer infralittoral and was abun¬ dant away from nearshore turbid environments and regions of relatively high sediment influx. The numer¬ ous crinoids and dalmanellid, atrypid, and stropho- menid brachiopods are most likely indicative of waters of normal marine salinity. The most favorable sub¬ stratum was a calcareous silt or muddy silt, although some individuals lived in coarser sands; the community as a whole is more common in silts. Local patches of slightly mobile, calcareous silt and sand may have given rise to locally higher concentrations of treposto- matous bryozoans, some branches of which show growth around crinoid columnals. The Dalmanella- Sowerbyella Community intergrades at some localities with the Nuculites-Colpomya Community which was composed primarily of infaunal detritus feeders liv¬ ing in silty muds. Conversely, the Dalmanella-Sow¬ erbyella Community was dominated by epifaunal sus¬ pension feeders presumably on a firmer silt or fine sand. The Dalmanella-Sowerbyella Community graded 24 shoreward (east) into sands and silts containing a fauna dominated by large epifaunal suspension feed¬ ing bivalves comprising the Ambonychia-Modiolopsis Community, and graded seaward (west) into fissile gray-black muds and silts, containing only a sparse trilobite-graptolite fauna. The silty bottom was oc¬ casionally marked by low, closely spaced interference ripples, and trails and tracks exhibit a fine meander¬ ing pattern that is correlated with the deeper or at least offshore Nereites facies of Seilacher (1964). The Nuculites-Colpomya Community occupied both outer and inner infralittoral environments and was characterized by the major expansion of infaunal de¬ tritus feeding bivalves that were only sparsely dis¬ tributed in the offshore waters of the brachiopod-dom- inated Sowerbyella-Onniella Community in north-cen¬ tral Pennsylvania. The fauna appears to have been normal marine, although some individuals in nearer shore environments may have experienced periods of variable salinity. The substratum presumably was a silty mud, but the community shows some noticeable onshore to offshore changes in faunal composition. Intergrading with both offshore Dalmanella-Sowerby- ella and nearshore Ambonychia-Modiolopsis Com¬ munities, the deeper-water faunas of the Nuculites- Colpomya Community contained greater numbers of the orthid brachiopod Glyptorthis crispata, the modio- morphid bivalve Colpomya faba, and the monopla- cophoran Archinacella. In contrast, the nearer shore faunas, those interbedded with the Ambonychia-Mo¬ diolopsis Community, exhibit a greater diversity of burrowing, infaunal desmodont bivalves Rhytimya, Cy- matonota, Cuneamya, and Psiloconcha? . Lesser num¬ bers of Colpomya faba are found in the nearshore regions but do occur with some patches of graptolites; it is suspected that these small mussels may have been attached to other planktonic forms and floated in like the graptolites. Nuculites also undergoes considerable size changes from offshore deposits where it is only 4 mm in length, to nearshore deposits, where it mea¬ sures 1.7 cm (measurements medians of slightly less than 100 specimens) . The size differential, if sig¬ nificant, may be the result of this protobranch having to maintain a certain distance beneath the sediment water interface; therefore, small size would compen¬ sate for decreased firmness of the substratum with increased water content of the muds. The dominant nearshore assemblage was the Am¬ bonychia-Modiolopsis Community, composed primar¬ ily of an epifaunal, suspension-feeding, bysally-attach¬ ed bivalve fauna. This fauna was seemingly well adapted to regions of irregular sedimentation and in areas of mobile bars and barriers; thus most of the species presumably existed on a firm, but slightly mobile sandy substratum. In some areas cut-and-fill channels, bars, crossbedded coquinites, large scale in¬ terference ripples and oriented orthoconic cephalopods testify to a reasonably high energy nearshore environ¬ ment. These fossiliferous marine environments graded landward (east) into transitional nearshore marine and alluvial Oswego orthoquartzites and seaward (west) into the finer silts and shales dominated by the extensive epifaunal brachiopods of the Dalmanella- Sowerbyella Community. The lower Oswego sand¬ stones enclose only a very few disarticulated valves of Modiolopsis modiolaris and it is thought that much of the Oswego may have been supratidal or alluvial in origin. Biogenic reworking also emphasizes, by anal¬ ogy with Recent and ancient deposits (cf. Cruziana facies of Seilacher, 1964), the intertidal and shallow sublittoral pattern of the upper Pulaski deposits. U- shaped tubes and lobate grazing patterns (“ turkey tracks ”) appear to represent possible parts of inter¬ tidal and shallow sublittoral tunnels, burrows or graz¬ ing patterns (cf. Shinn, 1968 and Schafer, 1962). The total Ambonychia-Modiolopsis fauna was presumably broadly tolerant and able to withstand the rigors of salinity fluctuations, occasional harsh physical regimes and possibly desiccation, having been attached and partially buried in a slightly mobile substratum. 25 Summary and Conclusions The major objectives of this paleoecologic study were to outline the benthic marine invertebrate com¬ munities that were present in north-central New York during the Late Ordovician, and to compare these community structures with those to the south in the central Appalachians (see Bretsky, 1969). Precise tracing of the faunas between the two areas is not possible because of the Late Paleozoic overlap in south-central New York and northern Pennsylvania, but it appears that a reasonably integrated Late Ordo¬ vician benthic community pattern can be constructed for eastern North America despite this gap in the record. The Late Ordovician benthic communities occu¬ pied a westward sloping shelf that was receiving sedi¬ ment from uplifts to the east, prograding the shore¬ line westward throughout the Late Ordovician. This shoreline and associated shelf environments have now been traced from north-central Tennessee into cen¬ tral New York; and it has been shown that the area of central Pennsylvania was the region of greatest sediment influx (see especially Yeakel, 1962; Horo¬ witz, 1966). A clastic wedge is readily apparent on isopach and sediment distribution maps of the Late Ordovician (Sloss et al, 1960, p. 8; Kay and Colbert, 1965, p. 170). Previous workers have interpreted this region as representing the development of a major deltaic complex. Although subsidiary clastic wedges developed immediately to the north (Queenston) and south, the same intensity in the amount of sediment contributed to the basin is not exhibited. Thus the development of this major deltaic complex in central Pennsylvania gave rise to a well-delineated onshore- to-olfshore sediment gradient, and it is thought that longshore currents reworked some of this deltaic ma¬ terial and transported it southward along the shore¬ line where it was deposited in the form of bars and barriers (Bretsky, 1969) . Thus to the south of central Pennsylvania and extending into central Virginia are developed a number of Late Ordovician nearshore barrier-bar-lagoonal environments that are quite distinct from those delta-front environments in central Penn¬ sylvania (also Horowitz, 1966 and Thompson, 1967). Somewhat similar nearshore environments are found in north-central New York, but they are not as ex¬ tensively developed, and are physically cut off from those to the south of the deltaic complex. In south¬ western Virginia and northern Tennessee, the rate of sediment influx and deltaic progradation is markedly lower so that much more significant amounts of car¬ bonate mud and shells were deposited. In the south the accumulation of substratum depended more on shell debris and less on sediment influx into the area. Thus the onshore-to-offshore sediment gradient becomes very slight in the south-central Appalachians when compared to those in the north-central Appalachians and central New York. Into this general sedimentological scheme are placed four major benthic community types, each one closely associated with the progradation of the main deltaic complex in central Pennsylvania. The four community types are arranged on p. 28 on a very general bathy¬ metric scale (also see table 3 and fig. 16). As depicted in figure 16 the distribution of these community types is correlated with the position of the major deltaic complex. Environmental implications may be determined from the amount and type of sed¬ iment being delivered to a particular region of the shelf, which in turn would have a direct bearing on the attachment sites or burrowing and feeding habits. De¬ tails of the distribution patterns within this overall structure emphasize further environmental controls, such as the selection for broadly tolerant species in some nearer shore environments where the effects of vari¬ able salinity and desiccation are added to the sedi¬ mentological effect on feeding habits and substratum type. The degree to which the distribution of the Late Ordovician communities is affected by the location of source areas is observed in the abrupt change that takes place in the nearshore environments on either side of, and immediately off of, the deltaic complex; whereas offshore communities (Dalmanella-Sowerbyella and Sowerbyella-Onniella) on the apron or fringes of the delta front show little change in faunal composi¬ tion. (table 3). Although both Orthorhynchula-Ambonychia and Am- bonychia-Modiolopsis Communities are dominated by 27 Offshore: f Sowerbyella-Onniella Community (north-central Appalachians) \ Dalmanella-Sowerbyella Community (New York) dominant fauna: strophomenid and dalmanellid brachiopods crinoids trilobites Offshore and Onshore: jj Zygospira-Hebertella Community (south-central Appalachians) ■ dominant fauna: trepostomatous bryozoans atrypid and orthid brachiopods IJI J Nuculites-Colpomya Community (New York) dominant fauna: nuculoid, actinodontoid, and desmodont bivalves Onshore: , Orthorhynchula-Ambonychia Community (north-central Appalachians) dominant fauna: rhynchonellid and linguloid brachiopods rV ■< ambonychiid and modiomorphid bivalves bellerophontid gastropods ^ Ambonychia-Modiolopsis Community (New York) dominant fauna: ambonychiid and modiomorphid bivalves monoplacophorans large, byssate, epifaunal suspension-feeding bivalves, there are significant differences in faunal composi¬ tion from the community in south-central Pennsylvania and the one in north-central New York, and, in fact, neither community occurs off of the delta front in cen¬ tral Pennsylvania. The most significant faunal differ¬ ences are seen in the abundance of the rhynchonellid brachiopod (Orthorhynchula linneyi), the linguloid brachiopod (Lingula! ), the bellerophontid gastropod ( Fleet onotus?) and the nuculoid bivalve mollusc (Tan- crediopsis) in the north-central Appalachians. Seem¬ ingly related to this barrier-bar-lagoonal faunal suite (i.e. the bellerophontid, linguloid, and nuculoid) in New York is a much more widespread community, the Nuculites-Colpomya Community, that shows neither the bellerophontid or lingulid, but rather an abun¬ dance of nuculoid (Nuculites, Ctenodonta?, Palaeo- concha, Praenucula) and desmodont bivalve molluscs that were common in some offshore delta-front en¬ vironments in central Pennsylvania, but have expanded in numbers, diversity, and areal extent in New York. It appears that the offshore nuculoids and desmodonts of central Pennsylvania came to dominate the infaunal nearshore niche in New York, where the detritus feed¬ ing nuculoid Tancrediopsis is completely absent. This much broader development of the more southerly offshore nuculoid fauna into the nearshore regimes may be a result of larval dispersal problems, Tan¬ crediopsis, simply being unable to migrate around the environmental-sedimentological barrier. Possibly cur¬ rent or temperature controls were operative so that the more southerly deeper-water nuculoid came to occupy a nearshore environment in central New York. The acceptance of a possible north-south bathymetric temperature control of the distribution of the nuculoid bivalve molluscs appears to be in conflict with prev¬ ious paleomagnetic and faunal interpretations of Or¬ dovician paleogeography. Allowing for individual in- terpretational differences, Irving (1964), Opdyke (1962), Spjeldnaes (1960), Whittington (1966) and most recently Fell (1968) have emphasized that east¬ ern North America during Ordovician time would have been a tropical or sub-tropical environment. In fact, in one instance (Fell, 1968, fig. 2), the latitudes parallel the reconstructed central Appalachian and New York Late Ordovician shoreline. The large epifaunal, suspension-feeding bivalve molluscs continued to dominate the nearshore regime 28 LATE ORDOVICIAN COMMUNITIES CENTRAL APPALACH IANS - NEW YORK Figure 16. — Late Ordovician communities central Appalachians — New York. Note that the Dalmaneila-Sowerbyella (New York) and Sowerbyella-Onniella (central Appalachian) Communities have been combined into a single pattern (community type I) , whereas for clarity in graphical presentation the Ambonychia- Modiolopsis (New York) and Orthorhynchula-Ambonychia (central Appalachian) Communities have remained as separate patterns although forming a distinct onshore community type (IV) . (See text) . with, however, the addition of the large coiled mono- placophoran Cyrtolites, which was not present in the central Appalachians. This represents a possible niche replacement for the browsing detrital-feeding bellero- phontid gastropod that was so abundant to the south. Only a few of the nearshore species occur in any numbers in offshore environments; these are the large modiomorphid and ambonychiid bivalve molluscs, both of which are found in small numbers in the Dal- manella-Sowerbyella and Sowerbyella-Onniella Com¬ munities and, incidentally, immediately off the delta front in central Pennsylvania. These are the only species that are common in the nearshore environments both to the north and to the south of the deltaic complex and it is possible that they are the only broadly tol¬ erant forms able to exist, albeit in small populations, around the perimeter of the delta. Here they could take advantage of the organically rich nearshore re¬ gimes in both north and south. Single specimens of Orthorhynchula, Lingula?, Plectonotus?, and Tancredi- opsis are rarely if ever found in the off-delta front environments in central Pennsylvania. Detailed ex¬ planations of the probable migration patterns must remain until further studies are completed in the Mid- Continent and Canada, but it is evident that this major deltaic complex in central Pennsylvania was a major environmental factor in the distribution of Late Ordovician benthic marine invertebrate communities. 29 References Bayer, T. N., 1967, Repetitive benthonic community in the Maquoketa Formation (Ordovician) of Minnesota: Jour. Paleontology, v. 41, p. 417- 422, pi. 51, 2 text figs. Bretsky, P. W., 1969, Central Appalachian Late Or¬ dovician communities : Geol. Soc. American Bull., v. 80. p. 193-212, 11 figs., 2 pis. - , Flessa, K. W. & Bretsky, S. S., 1969, Brachiopod ecology in the Ordovician of east¬ ern Pennsylvania: Jour. Paleontology, v. 43, p. 312-321, 4 text figs. Broughton, J. G., Fisher, D. W., Isachsen, Y. W. & Richard, L. V., 1962, The Geology of New York State: New York State Museum and Sci¬ ence Service, Map and Chart Series No. 5 (Text) , 42 p., 24 figs. Clark, T. H., & Steam, C. W., 1968, Geological evolution of North America: New York, The Ronald Press, 570 p. Conrad, T. A., 1839, Second annual report on the paleontological department of the Survey (of New York): State of New York Assembly Document (?) No. 275, 1839, Commun. transmitting re¬ ports Geological Survey, Annual Report 3, p. 57- 66. Emmons, E., 1842, Geology of New York; Part II comprising the survey of the second geological district: Albany, (Carroll and Cook), Natural History of New York, 437 p. Fell, H. B., 1968, The biogeography and paleoecology of Ordovician seas, in Drake, E. G., ed.. Evo¬ lution and environment: p. 139-162, 2 figs.. New Haven, Conn., Yale Univ. Press. Fisher, D. W., 1962, Correlation of the Ordovician rocks in New York State: New York State Mu¬ seum and Science Service, Geological Survey, Map and Chart Series No. 3. - , and others, 1961, Geologic map of New York: New York State Museum and Science Service, Map and Chart Series 5. Foerste, A. F., 1914, Notes on the Lorraine faunas of New York and the Province of Quebec: Bull. Science Lab. Denison Univ., v. 17, p. 247-340, 5 pis. - , 1916, Upper Ordovician formations in On¬ tario and Quebec: Canada Geol. Survey, Memoir 83 (No. 70 Geol. Series), 279 p. - , 1924, Upper Ordovician faunas of Ontario and Quebec: Canada Geol. Survey, Memoir 138 (No. 121 Geol. Series), 255 p., 46 pis. Grabau, A. W., 1913, Early Paleozoic delta deposits of North America: Geol. Soc. America Bull., v. 24, p. 399-528, 1 pi., 12 figs. Hall, J., 1847, Descriptions of the organic remains of the lower division of the New York system: Palaeontology of New York, v. 1, 338 p., 98 pis. Hofmann, H. J., 1964, Mud cracks in the Qrdovician Maysville Group: Jour. Geology, v. 72, p. 638- 641, 1 pi. Horowitz, D. H., 1966, Evidence for deltaic origin of an Upper Qrdovician sequence in the central Appalachians, in Shirley, M. L. and Ragsdale, J. A., eds.. Deltas in their geologic framework: Houston, Texas, Houston Geol. Society, p. 159- 169, 2 pis., 5 figs. Irving, E., 1964, Paleomagnetism and its application to geological and geophysical problems: New York, J. Wiley, 339 p., 259 figs. Kay, M. & Colbert, E. H., 1964, Stratigraphy and life history; New York, J. Wiley, 736 p. King, P. B., 1959, The Evolution of North America: Princeton, N. J., Princeton Univ. Press, 189 p., 96 figs. McBride, E. F., 1962, Flysch and associated beds of the Martinsburg Formation (Qrdovician), cen¬ tral Appalachians; Jour. Sed. Petrology, v. 32, p. 39-91, 34 figs. Mason, G. D., 1967, Depositional environments of the Lorraine and Richmond Groups in the St. Lawrence Lowlands of Quebec: McGill Univer¬ sity, M. S. Thesis, 69 p., 30 pis. Opdyke, N. D., 1962, Paleoclimatology and conti¬ nental drift in Runcorn, S. K., ed.. Continental drift: Chap. 2, p. 41-65, 14 figs.. New York, Academic Press. 31 Petersen, C. G., 1914, On the distribution of the animal communities of the sea bottom: Report Danish Biol. Station, v. 22, 7 p. Ruedemann, R., 1925a, The Utica and Lorraine for¬ mations of New York: New York State Museum Bull. 258, Part 1. Stratigraphy, 175 p., 7 pis. - , 1925b, The Utica and Lorraine formations of New York: New York State Museum Bull. 262, Part 2. Systematic Paleontology No. 1, plants, sponges, corals, graptolites, crinoids, worms, bryo- zoans, brachiopods, 171 p., 13 pis. - , 1926a, Faunal facies differences of the Utica and Lorraine shales: New York State Museum Bull. 267, p. 61-77. - , 1926b, The Utica and Lorraine formations of New York: New York State Museum Bull. 272, Part 2. Systematic Paleontology No. 2, mollusks, crustaceans and eurypterids, 227 p., 28 pis. Schafer, W., 1962, Aktuo-Palaontologie nach Studien in der Nordsee: Frankfurt am Main, Kramer, 666 p., 277 abb., 78 figs. Seilacher, A., 1964, Biogenic sedimentary structures, in Imbrie, J. and Newell, N. D., eds., Approaches to paleoecology: New York, J. Wiley, p. 296- 316, 8 figs. Shinn, E. A., 1968, Burrowing in Recent lime sedi¬ ments of Florida and the Bahamas: Jour. Paleon¬ tology, V. 42, p. 879-894, pis. 109-112, 17 text- figs. Sloss, L. L., Dapples, E. C. & W. C. Krumbein, 1960, Lithofacies maps: New York, J. Wiley, 108 p., 153 maps. Spjeldnaes, N., 1960, Ordovician climatic zones: Norsk. Geol. Tidsskr., v. 41, p. 45-77, 7 figs. Thompson, A. M., 1967, Facies succession in a ma¬ rine-continental transition in the Upper Ordo¬ vician of central Pennsylvania (abs.) : Geol. Soc. America, Northeast Section Meeting (1967), p. 61. Ulrich, E. O., 1911, Revision of the Paleozoic sys¬ tems: Geol. Soc. America Bull., v. 22, p. 281- 680, 5 pis., 19 figs. - 1913, The Ordovician-Silurian boundary: Etude faite a la XII e session du Congres geo- logique international, reproduite du Compte Rendu, p. 593-667. Vanuxem, L., 1840, Fourth annual report of the geo¬ logical survey of the third district: New York Geological Survey, Annual Report 4, p. 355-383. Whittington, H. B., 1966, Phylogeny and distribution of Ordovician trilobites: Jour. Paleontology, v. 40, p. 696-737, 16 text-figs. Yeakel, L. S., 1962, Tuscarora, Juniata and Bald Eagle paleocurrents and paleogeography in the central Appalachians: Geol. Soc. America Bull., V. 73, p. 1515-1540, 17 figs. 32 Locality Register (not including exposures of glacial drift containing Lorraine Group fossils; all quadrangles 1:24,000 unless otherwise noted) Locality Description 1 Richland Quadrangle — Road cut at intersection of Oswego County Route 22 (Lacona-Orwell Road) and Lester-Jerry Look Road. 2 Sandy Creek Quadrangle — Road cut 2.3 miles east of Lacona on Oswego County Route 15 (Lacona-Smartville Road); exposure immedi¬ ately west of cutoff to Boylston Center. 3 Boylston Center Quadrangle — Stream cut 0.3 mile southwest of Winona where stream crosses the Mannsville-Winona Road. 4 Pulaski Quadrangle — Stream cut (Salmon River) in Pulaski Village; exposure along river bank below River Street. 5 Richland Quadrangle — Stream cut (Salmon River) beneath overpass of Rome, Watertown, and Oswego Railroad bridge; 1.3 miles south¬ east of Pulaski Village where tracks cross Pulaski-Centerville Road. 6-A Orwell Quadrangle — Stream cut, Salmon River Falls, 1.1 miles east on dirt cutoff from Or- well-Bennett Bridge Road. 6 Orwell Quadrangle — Stream channel cut in front of Red Brick Power Station (Niagara- Mohawk Power Company) Bennett Bridge; 0.1 mile north of Bennett Bridge along Or- well-Bennett Bridge Road. 7 Richland Quadrangle — Road cut along U.S. Interstate 81 immediately north of intersec¬ tion with New York Route 13. 8 Barnes Corners Quadrangle — Road cut 0.2 mile south of East Rodman along Whitesville Road. 9 Barnes Corners Quadrangle — Road cut 0. 1 mile east of Jefferson-Lewis County line along Whitesville Road. 1 1 Barnes Corners Quadrangle — Stream cui (Shingle Gulf) at a point 0.5 mile south of junction of Whitesville and Cronk Corners Roads. 12 Barnes Corners Quadrangle — Stream cut (South Sandy Creek) immediately south of Seven-by- Nine Corners and extending west downstream for 1.1 miles. 13 Adams Quadrangle — Stream cut (South Sandy Creek) 0.2 mile south of Allendale where New York Route 178 crosses South Sandy Creek. 14 Rodman Quadrangle — Stream cuts (Fox Creek? and Big Brook) southeastern part of Lorraine Village. 15 Rodman Quadrangle — Stream cut (Fox Creek south branch) 1.2 miles south of Lorraine along Lorraine-Pierrepont Manor Road. 16 Boylston Center Quadrangle — Stream cut (Fox Creek) 1 . 1 miles south of Lorraine along Lor¬ raine-East Boylston Road; 0.4 mile north of Watertown Road cutoff. 17 Rodman Quadrangle — Stream cut (Abijah Creek) at intersection of creek and New York Route 178; 1.0 mile west of Bullock Corners. 18 Rodman Quadrangle — Stream cut (South Sandy Creek) 0.1 mile north of Worth on New York Route 178. 19 Barnes Corners Quadrangle — Road cut imme¬ diately east of Babbits Corners (junction of New York Routes 177 and 178) on New York Route 177. 20 Rodman Quadrangle — Stream cut (South Sandy Creek; Lorraine Gulf) 0.3 mile north of Bul¬ lock Corners along Bullocks Corners-Tremaines Corners Road. Barnes Corners Quadrangle — Road cut 0.2 mile east of locality 9 along Whitesville Road. 33 21 22 23 24 25 26 27 28 29 30 31 Rodman Quadrangle — Stream cut (Fish Creek) 1.0 mile south of Rodman on Rodman-Ross Corners Road. Adams Quadrangle — Stream cut (Towle Gulf) 0.9 mile east of Allendale in Washington County Park off of Washington Park Road. Barnes Corners Quadrangle — Stream cut (Gulf Stream; Inman Gulf) 0.1 mile north of inter¬ section of New York Route 177 and Whites- ville Road at Barnes Corners. New Boston Quadrangle — Stream cut 0.1 mile east of Bellwood immediately off of New York Route 177. West Lowville Quadrangle — Road cut 0.6 to 0.9 mile west of West Martinsburg along West Martinsburg-Rector Road. Sears Pond Quadrangle — Road cut and stream cut 0.2 to 0.4 mile northeast of Sears Pond along Sears Pond-Rector Road. New Boston Quadrangle — Stream cut (Deer River) immediately west of New Boston along New York Route 177. New Boston Quadrangle — Stream cut near sec¬ ondary road 0.5 mile north of junction with New York Route 177; junction 0.4 mile west of Bellwood. Port Leyden Quadrangle (1:62,500) — Stream cut (Whetstone Gulf) exposures extending 1.8 miles upstream from park entrance. Port Leyden Quadrangle (1:62,500) — Road cuts 1.0 to 1.5 miles southwest of Martins¬ burg along Corrigan Hill Road to Tabolt Cor¬ ners. Page Quadrangle — Stream cut (Whetstone Creek) where Corrigan Hill Road crosses Whetstone Gulf. 32 Port Leyden Quadrangle (1:62,500) — Stream cut (Mill Creek) 1.5 to 1.7 miles west of Turin along Turin-Welch Hill Road. 33 Lee Center Quadrangle — Stream cut (East Branch Fish Creek) immediately west of inter¬ section of stream and old New York Route 69 at village of Taberg. 34 Point Rock Quadrangle — Stream cut (East Branch Fish Creek) 0.8 mile northwest of Point Rock along Yorkland Road. 35 Lee Center Quadrangle — Stream cut (Florence Creek) 3.1 miles north of Taberg along Coal Hill Road. 36 Lee Center Quadrangle — Stream cut (?Wood Creek or Canada Creek) in Lee Center where stream crossed by Center Road. 37 North Western Quadrangle — Quarry at Fink Hollow immediately north of the intersection of the Steuben-Boonville Road and Big Brook. 38 Boonville Quadrangle — Road cut 0.9 mile west of North Steuben Church along Buck Hill Road. 39 West Leyden Quadrangle — Stream cut 0.7 mile east of Ava along East Ava Road and in Mo¬ hawk River. 40 Point Rock Quadrangle — Stream cut (East Branch Fish Creek) 0.3 mile east of Swan- cott Mill along Swancott Mill Road. 41 West Leyden Quadrangle — Road cut 0.3 mile east of West Leyden along New York Route 294. 34 BULLETIN NUMBER 415 ( Al'" it New York State Museum and Science Service^^^'-’j The University of the State of New York The State Education Department JANUARY 197T The Behavioral Patterns of the Eastern Bluebird ( St aha stalls) David C. Krieg BULLETIN NUMBER 415 New York State Museum and Science Service The University of the State of New York The State Education Department JANUARY 1971 THE UNIVERSITY OF THE STATE OF NEW YORK Regents of the University {with years when terms expire) 1984 Joseph W. McGovern, A.B., LL.B., L.H.D., LL.D., D.C.L., Chancellor . New York 1985 Everett J. Penny, B.C.S., D.C.S., Vice Chancellor - - - - White Plains 1978 Alexander J. Allan, Jr., LL.D., Litt.D. . Troy 1973 Charles W. Millard, Jr., A.B., LL.D., L.H.D. . Buffalo 1972 Carl H. Pforzheimer, Jr., A.B., M.B.A., D.C.S., H.H.D. - Purchase 1975 Edward M. M. Warburg, B.S., L.H.D. . New York 1977 Joseph T. King, LL.B. . Queens 1974 Joseph C. Indelicato, M.D. . Brooklyn 1976 Mrs. Helen B. Power, A.B., Litt.D., L.H.D. . Rochester 1979 Francis W. McGinley, B.S., LL.B., LL.D. . Glens Falls 1980 Max J. Rubin, LL.B., L.H.D. . New York 1971 Kenneth B. Clark, A.B., M.S., Ph.D., Litt.D. . Hastings on Hudson 1982 Stephen K. Bailey, A.B., B.A., M.A., Ph.D., LL.D. - - - - Syracuse 1983 Harold E. Newcomb, B.A. . Owego 1981 Theodore M. Black, A.B. . Sands Point President of the University and Commissioner of Education Ewald B. Nyquist Executive Deputy Commissioner of Education Gordon M. Ambach Associate Commissioner for Cultural Education John G. Broughton Principal Scientist, Biological Survey Donald L. Collins State Zoologist Ralph S. Palmer Dedicated to the Memory of ELON HOWARD EATON author of The Birds of New York 1910 and 1914 NEW YORK STATE BIRD On May 18, 1970, Governor Nelson A. Rockefeller signed chapter 824, section 78 of the State Laws of 1970 and made the Eastern Blue¬ bird the State Bird of New York. ^^*'’ T© fti3 r»^ ^.r «’ Hum )* ^ ijttil • . . >■ .. ;.. «MS^.ir3J,';. -1 hi #-.,'h. )^£i: ". T . .ffiH lytk. li^ fU^eS'i": iKt a* ^ fi^ittiai, Ui.t'* I K •* M. - i;tei» f»i|(^ '«tll| THM*^ i^|ix<4 . ... , ^ btui » i * f,y rjrJj . . fwhon^ J V, V*"^ " * ” ■*'* * ' -•■»■ •V'.;^:; *1. V », ;. . /; pihtaiH -jAfv^f/^ ^ijsiflogA . ’'^* '** ■*•'*' ‘ ■ ' i: yy'JiiaMw •‘f-j < • «'(A|<^'ftKI bwii eaiuJso*! J/»t(fr . . • ^^'■‘bBSff^aoV ' i i , , ; . .,; , -lorverfsH tKn&Xh ^ ‘ • J * ' ' • iJUtrnasdqaJa '* ";■ '■ ''^'.h ' • < •■• -I f H. larVfidifl 4n;)f/j‘>iS89qqA ’ • - *>.^t i Vj.r. . ,...,, . lofvofsff sbbl^ T. ' * '-?•■ ' . , r viojmaT ^ ’ * ‘ '■’ ’-'f ■ • • • ‘ * • ►« *X7 ^oS ^ V , ^ . ,v , 4,y , . ,, fi^tslgaid IjchsA . ,-- V.., . , -vv »_y ffinsinoK , Fmoiht»T‘ . ;»>">KbfinT5^ 3o «f®Ojj3(U/^ ' ^ *?;-“• i^A noiiwmo^ lUsfj : ' 1 * /*!***'*’' -*’-* 5 V '^r Sv/tiunrfioO ^ * : ;*^^’i ■' ' ^ ‘ JJ‘>bj/i300^ feared? • • .r'l^ v-*^ v't^* <; r‘.v^ . , . . , 9^j .Jo nbiifivtJoW . ■ ’ e^«rq£ia ' .1 The Behavioral Patterns of the Eastern Bluebird f Si a Ha sialis)^ By David C. Krieg^ INTRODUCTION This study describes the species-typical behavioral patterns of the eastern bluebird (Sialia sialis) and interprets this behavior from an ethological standpoint. The author intends to make a comparative ethological study of the genus Sialm, but prior to any such study, an attempt must be made to study intensively one species to obtain a com¬ parative standard for the group. The available literature on bluebird behavior is scattered and con¬ cerned primarily with breeding biology (Butler, 1907; Harper, 1926; Low, 1933, 1934; Musselman, 1935, 1939; Pettingill, 1936; Smith, 1937; Brodrick, 1938; Hamilton, 1943; Bent, 1949) with major work being done by Laskey (1939, 1940, 1943) Thomas (1946) and Hartshorne (1962). Reported information on agonistic and courtship behavior is cursory because no bluebird has yet been studied ethologi- cally. Therefore, the conclusions presented here are to be considered first approximations. Bluebird refers to Sialia sialis in this publication. Sialia sialis was studied in the field through five breeding seasons, 1963 to 1967. Most observations on behavior of the bluebird were made within a IS mile radius of St. Bonaventure University, the same 1 Manuscript submitted for publication May 21, 1970. 2 Holder of Student Honorarium, New York State Museum and Science Service, 1966. Present address : Department of Biology, State University of New York, New Paltz. [1] area covered by the annual Christmas Bird Census {Audubon Field Notes, April, 1951, pg. 67). Approximately 75 percent of this area is wooded, with the broad Allegheny River valley and tributary valleys open and used to some extent for agricultural purposes (Eaton, 1953), although few farmers gain a livelihood from the soil. The river valley has an elevation of 1400-1500 feet and the tributary valleys extend to 2000 feet. These open tributary valleys provide ideal nesting habi¬ tats, at this time, for bluebirds. Although field studies were made in all the tributary valleys within this area. Wing Hollow, in the township of Allegany, Cattaraugus County, New York, was selected for intensive behavior work. A sheep farm is located in the middle of the hollow and a larger dairy farm is at its mouth. These two operations provide approximately 300 acres of permanent pasture and many rows of fence lines. In addition to these, abandoned orchards, a centrally located brook with a few tributary streams, and electric utility lines, provide good habitat for bluebirds. Wintering behavior of N. sialis was studied from January 10 to January 24 and February 21 to February 28, 1967, in the Apalachicola National Forest, 15 miles south of Tallahassee, Leon and Wakulla Counties, Florida. Observations were also made at Tall Timbers Research Station, 20 miles north of Tallahassee and near the city of Vero Beach, Indian River County, Florida. Population Buildup The lack of a sizable breeding population of bluebirds to facilitate an ethological study was overcome by placing nest boxes in the study area. The nest box used was basically the one described by Pettingill and Hoyt (1963) and measured 7 inches deep with a floor space of 5x5 inches, a li-inch opening, and a hinged top. More than 100 such boxes were placed in the study area and table 1 shows the number of bluebird pairs using these boxes during the 5 years of this study. Trapping, Banding, and Marking Birds were banded and marked for individual recognition. Males, due to their strong territorial attachment and Nest Demonstration Display, could be caught almost anytime during the breeding cycle without abandonment of territory. The most effective trap was a nest box with a manually operated shutter similar to that described by Fischer (1944). A rectangular piece of cardboard (2" x 5"), with [2] TABLE 1 Number of Bluebird Pairs Using Nest Boxes Over a 5-year Period AREA YEAR NUMBER OF BOXES NUMBER OF NESTING PAIRS Wing Hollow 1963 3 1 1964 3 2 1965 10 7 1966 40 11 1967 43 10 Morgan Hollow 1963 4 3 1964 3 1 1965 10 2 1966 9 1 McClure Hollow 1963 3 2 1964 3 3 1965 12 3 1966 9 2 Smith Hollow 1963 3 2 1964 4 2 1965 6 3 1966 11 3 Five Mile Valley 1963 8 1 1964 6 2 1965 13 5 1966 8 1 Four Mile Road 1963 4 1 1964 2 2 1965 6 4 1966 6 2 Lippert Hollow 1963 4 2 1964 3 2 1965 5 2 1966 5 2 West Branch 1963 4 0 Four Mile 1964 4 2 1965 9 5 1966 12 7 [3] a long string attached, was nailed to one side of the entrance. After the bird entered the box, the string was pulled from a distance of about 100 feet and the cardboard moved up to cover the entrance. In the later stages of the nesting period, however, this method was not effective because the males, feeding the young at the entrance, seldom enter the box. Females are more sensitive and often will abandon the territory after being handled, and therefore, were usually not trapped until the later stages of incubation. The shutter trap was used in some instances but the female was more effectively captured at night as she roosted with her eggs or young. After banding and returned to the nest, she seldom abandoned the territory. Sometimes a stretch of cold weather early in the breeding season offered an unusual opportunity to capture and band most of the popu¬ lation in the study area. The birds would roost inside the nest boxes and (see roosting, pg. 21) were easily captured and banded at night. Early in the study, mist nets, and a modified bow trap (that re¬ leased a net over the box top and entrance when sprung) were used but proved too time-consuming and less effective than the shutter trap and nightly checking of nest boxes. To facilitate behavior observation, both adults and young were banded. An aluminum U.S. government band, plus one to three colored aluminum bands were placed on the birds’ legs. Different color combinations of aluminum bands identified each individual in the area. The most successful colors were white, yellow, red, and green. To make instant recognition from a distance possible and to gain valuable information on the timing of the molt, the remiges or rectrices were painted with white, red, or yellow airplane dope (see Lanyon, 1957). TABLE 2 Number of Adult and Young Bluebirds Banded During Each Year YEAR ADULTS YOUNG 1963 _ 90 1964 10 180 1965 47 81 1966 28 66 1967 7 13 Total 92 430 [4] Collection of Data Field observations were made almost daily from the arrival of the males in mid-March until the young had fledged. The visits varied from 1 to 7 hours, depending upon the weather, time of the breeding- cycle, and activity of the birds. Occasional shorter visits were made in late summer when the birds were molting, and again in the fall when they were flocking. Observations were facilitated by 7 x 35 binoculars and a 20X spot¬ ting scope. Notes in the field were tape recorded for later transcrip¬ tion. Outline maps, taken from aerial photographs, were used to record the location of activities and to chart the location, size, and shape of territories. The portable blind used initially proved unnec¬ essary, for observations could be made in the open from 20 to 30 yards without disturbing the birds. To obtain data on pair formation, agonistic behavior, and main¬ tenance of territory, both stuffed (with and without recorded vocaliza¬ tions), and live decoys were placed in the territory and the pairs’ reactions recorded. These experiments were highly successful and similar to those used by Lack (1953) and Dilger (1956a). Power (1966) using this type of investigation on Shlia currucoides had little success, because the birds ignored the decoy. The decoy and tape recorder (if vocalizations were used) were placed on a fence post by the nest box or elsewhere in the territory. A 50-foot wire extended from the tape recorder to the observer and by closing the switch the vocalizations could be played. Live decoys were kept in ordinary bird cages (12" x 7" x 12") tied to fence posts inside the territory. No tape recorded vocalizations were used. The live decoys elicited much stronger responses from the territorial pairs. The results of some experiments were recorded on 16 mm color film for later analysis. Over 100 experiments were conducted. Vocalizations were tape recorded at 7^ IPS (inches per second). Sometimes in the field the birds were so close that vocalizations were recorded on the portable tape recorder with the microphone mounted on a 24-inch parabolic reflector. In July 1966, eight young were taken from three nests at the age of 8 days. They were kept in a 10' x 10' x 10' (length x width x height) indoor flight cage connected (by a window that opened) to a smaller outdoor cage, 8' x 6' x 8'. In March 1967, three pairs were removed and placed in separate indoor cages approximately 8' x 6' x 8'; the fourth pair remained in the indoor-outdoor cage. Three of the four pairs mated and laid eggs, but no young were successfully raised. In the fall of 1967, the flock was reduced to two pairs and kept [5] in an indoor cage (8' x 6' x 8') until March 1968, when they were separated for breeding. The first cages, constructed of -^-inch mesh hardware cloth, proved unsatisfactory as rectrices and feathers at the base of the bill were lost when caught in the small mesh. Welded wire of 1 inch by 2 inch rectangular mesh was then used satisfactorily to cover the front and top. Each side consisted of two 4' x 8' sheets of ^-inch plywood nailed together. The wooden sides with the wire front and top rested against the back wall of the room. Each cage contained perches, nest box, infrared light, and a continual supply of food and water. In one cage, the birds were observed through a “ one-way glass ” window on the side of the cage. Data on the ontogeny of maintenance activities came from three young, hand-reared from 5 days of age. The birds were released when 30 days old. Some observations were also made on two pairs of hybrid bluebirds (Sialia sialis x Sialia currocoides) obtained from Dr. Robert Nero, Manitoba Museum of Man and Nature, Winnipeg, Canada (see Lane, 1968). The birds were housed in a 7' x 5' x 8' cage from September 1967 to March 1968 and then placed in separate 7' x 5' x 8' cages for breeding. Behavior of the captive birds was recorded in notebooks and on color film, using a Bell and Howell Model 432-A Super 8 camera with a 5 to 1 zoom lens. These movies were later analyzed ; 35 mm photo¬ graphs were made in the field and laboratory with 125 mm and 300 mm lenses. The laboratory diet of captive birds consisted mainly of Ken-L- Ration Burger for dogs, dried currants, and sumac (Rhus typhina), mealworm (Tenebrio sp.) larvae, insects, and various fresh fruits were given as supplements. Commercial vitamins (e.g. Vi-Sol) were added to the drinking water or mixed in the food. [6] ACKNOWLEDGEMENTS I am especially indebted to Stephen W. Eaton, Professor of Biology at St. Bonaventure University, St. Bonaventure, for his advice and guidance during this study. He was very helpful in obtaining nest boxes, tape recorders, photographic supplies, aviaries, and other equip¬ ment needed during field and indoor work and generous in allowing unlimited use of his personal library. Thanks are due to Wilson Baker for his hospitality at Tall Timbers Research Station, Tallahassee, Fla. ; Anthony Moriello for preparing the drawings of the bluebird postures and displays ; Jerry Noles for assistance in photographing wild bluebirds ; Vernon Rounds for tech¬ nical advice ; and Marvin Wing for permission to use his land as a study area. Field work was supported by a Student Honorarium grant from the New York State Museum and Science Service and one from the Chapman Fund of the American Museum of Natural History. The cover photograph was taken by Ralph S. Palmer, State Zoolo¬ gist, who has also helped in many other ways. [7] [8] MAINTENANCE ACTIVITIES Marler (1956) defines maintenance activities as “those activities which are concerned with locomotion and the general health and effi¬ ciency of the body, mostly occurring throughout the year.” As Ficken and Ficken ( 1966) point out, maintenance activities are not only important in the animal’s survival but also in understanding the evolution of bird displays. They may even be useful as taxonomic characters. Because maintenance activities often provide the raw material from which agonistic and sexual displays have evolved, understanding these activities is essential to understanding the specialized displays derived from them. CARE OF THE BODY SURFACE Bill-wiping The body is lowered and directed along the perch. The bill is wiped from base to tip, on alternate sides of the perch several times in suc¬ cession, often with short pauses between. Incomplete bill-wiping movements were first seen in young 12 days old. Wing-fiapping Young in the nest rise up on their legs, stretch out both wings and flap them vigorously. The bird then settles back in the nest and may preen the wings. Wing-flapping was first seen in young 11 days old and either disappears or is rare in adult birds (some flapping may be used in drying movements). Wing-flapping helps arrange the flight feathers and with preening may aid in breaking the sheaths enclosing the feathers. [9] Head-shaking Head-shaking is uncommon in nestling bluebirds although Dilger (1956b) commonly found it in fledgling and young thrushes (Catharns sp. and Hylocichla sp.), but not in adult birds. I observed it only once in nestling bluebirds. A 5-day old stretched its neck in the air then shook the head and neck laterally. This is quite common in adults, but the neck is not extended. Head-shaking can occur with bill-wiping, bathing, drying, and often is associated with feather settling movements, especially ruffling and shaking the plumage. Head-shaking is also associated with food rejection. Both young and adult birds will open the mouth wide then shake the head violently until the unacceptable object is removed. Pellets (compacted indi¬ gestible food remains) can be cast without violently shaking the head. As the bill is opened there are throat movements, and the pellet is voided. Nodding Nodding is a deliberate downward head movement with the bill pointing directly at the perch. The nod, held momentarily as the bird Fig. 3 Eastern Bluebird scratching the throat indirectly [10] “ inspects ” the feet or surrounding area, may be successively repeated, then completed by p)ecking the tarsometatarsus, leg band, or toes, as the bird cleans its feet. After inspecting the feet, the bird frequently preens the belly. If on a flat surface (top of a nesting box), the bird may hop backwards between nods. Nodding frequently precedes bill-wiping, head-rubbing, and cleaning the feet, and probably is an intention movement to do so. Yawning Yawning was first seen in young 9 days old. The bill is opened widely then closed in a deliberate manner, often while the bird is rest¬ ing. Hartshorne (1962) reports that incubating female bluebirds preen, scratch, and yawn. Head-scratching Bluebirds scratch the head indirectly, with the foot moving up and over the dropped wing (Ficken and Ficken, 1958; pers. obs.). The bill and eyes may be open. The crown, sides of the head, and throat are scratched. Extended-scratching (Simmons, 1961) is used to oil the head feathers. After bathing, the bird obtains oil from the uropygial gland with its bill, scratches first the bill, then the head, moving the head in an irregular fashion underneath the foot. Head-scratching was first seen in young 12 days old. The first movements appear awkward with many incomplete attempts as the bird loses its balance. One young scratched under the wing (direct) with the foot, scratching the breast instead of the head. Stretching The bluebird uses the same three main types of stretching as the other North American thrushes {Hylocichla sp. and Catharus sp.) studied by Dilger (1956b). 1. One wing and leg stretch. The tail is spread, then the wing is lowered and extended as the leg, on the same side, is lifted from the perch and stretched down and outward with the wing. The tail is spread toward the side of the wing and leg stretch. 2. Both wings up stretch. Both wings are stretched up over the back, partly unfolded, then returned to the side. 3. Both legs stretch. Both legs are stretched together lifting the bird from the perch with a corresponding lowering of the head, lifting of the rump, and depression of the tail, giving the bird an arched appearance. This is the rarer type of stretching. An [11] [12] Eastern Bluebird stretching. Left — one wing and one leg stretch ; right — both wings upstretched adult female in captivity gave both wings up stretch and both legs stretch simultaneously. Any of these types may occur alone or in a sequence with the others. The most common sequence is : one wing and leg stretch, both wings up stretch. This can be continued to include the other wing and leg or both legs stretch. A typical full stretching sequence in the thrushes studied by Dilger (op. cit.) is to first stretch both wings over the back, then stretch a wing and leg on one side, stretch the wing and leg on the other side, then stretch both legs. This kind of sequence, one type (wing and leg) followed by the same type (wing and leg), did not exist in the bluebirds I studied in the field and aviary. Stretching was first seen at the age of 9 days. In addition to the three types described above, young bluebirds employ the both wings down stretch. This movement has been reported for other species (Andrew, 1956; Dunham, 1964; Ficken, 1962b; Nice, 1943) and becomes rare or nonexistent in adults. The both wings down stretch is usually given with the both legs up stretch in young bluebirds. The bird pushes up on the legs lifting the rear end and at the same time stretches both wings out and down. The head is tilted down and the back arched up. The wings down stretch can be given without the both legs stretch but they usually occur together. Both wings down never was observed in adult birds and Ficken (op. cit.) considered it a transitional movement in the redstart (Seto- phaga ruticilla) giving rise to the wing and leg sideways stretch. This is not a transitional movement in the bluebird. Both types (both wings down; one wing and leg) are well developed by the 10th day and both are used until the young bird leaves the nest (17th day, Hartshorne, 1962; pers. obs.). Possibly, the disappearance of the both wings down stretch in adults is due to body mechanics prevent¬ ing the fledged bird from performing this stretch without losing bal¬ ance. (A captive 16-day-old bird spent most of the day out of the nest and performed the both wings down only once, but performed it fre¬ quently in the nest at night.) Preening Preening was not studied in detail. The bird can reach all areas of the body with the bill except the head and neck. Remiges and rectrices are cleaned by being drawn through the bill. The eyes are usually closed. Young will preen the remiges and rectrices before the feathers have broken through the sheaths and by the age of 9 days all major feather [13] Fig. 6 Fig. 5 Eastern Bluebird preening the breast Eastern Bluebird preening under the wing after bathing [14] tracts are worked by the bill. Nibbling on a feather or sheath is interspersed with preening. Bluebirds were not seen to eat dislodged feathers but Dilger (A. Weisbrod, 1965, Unpublished Masters Thesis; Cornell Univer¬ sity) has observed juvenal and adult thrushes {Catharus sp. and Hylocichla sp.) eating feathers and feather sheaths. The tarsus and toes are pecked (sometimes scales are nibbled) to clean them. Both wild and captive birds show considerable irritation after being marked with aluminum bands. They peck at and even grasp the band with the bill, pulling at it, causing a loss of balance and sometimes toppling from the perch. Aluminum bands were never removed from the legs by the birds themselves as Dunham (1964) reported for the rose-breasted grosbeak {Phencticus ludovicianns) . The feathers near the base of the bill are cleaned by rubbing this area on the side of the perch. The bill is generally open and the rubbing is a slow and deliberate movement progressing from either the front of the head toward the back or from the back toward the front. Bill-wiping is often combined with head-rubbing. Preening, like many other maintenance activities, acts as a social stimulus with both members of a pair preening together unless one is engaged in another activity. Members of a flock frequently preen in concert. A winter flock (wild) of 18 birds landed in the top a leafless oak tree (Quercus sp.) and all preened for 5 minutes before leaving. Feather-ruffling This is a feather-settling movement where the body feathers are ruffled simultaneously, in various intensities, then lowered. Ruffling can occur by itself or in association with other feather-settling move¬ ments. A typical full sequence would be : the feathers are ruffled momentarily, the bird leans forward, the body is shaken from side to side with a corresponding shuffling of the wings and tail (the tail may also be fanned), the feathers are then lowered as the head is shaken and the bill wiped. Bluebirds ruffle the body feathers during preening, sunning, drying, and bathing. It is a common displacement activity in many conflict situations and resembles (except the plumage of the head is not lat¬ erally compressed nor is the tail spread) the spread display of the wood thrush {Hylocichla mustelina) (Dilger, 1956a), but in the blue¬ bird feather-ruffling has not yet become ritualized to serve as a display. [15] Bathing Bluebirds have tv^^o methods of bathing, dipping and complete- bathing. Either can be preceded by drinking or dipping the bill in the water with a sideways flick of the head, splashing water upon the bird. Both methods consist of two alternating phases : 1. The head and breast are lowered into the water, the bill is shaken from side to side, and both wings are beaten. Some¬ times the wing movement may be omitted. 2. The head and breast are raised, the rump is lowered into the water, the tail is spread, and the wings are fluttered. In dipping, the bird is in the water briefly and may only perform phase ( 1 ) before leaving. If both phases are performed they are done quickly and without ruffling the feathers. Bluebirds rarely take one dip but usually return two to three times in succession without thoroughly wetting the plumage. In complete-bathing (almost always preceded by dipping) the bird remains in the water and alternates between both phases, while ruf¬ fling the feathers until the plumage is thoroughly wet. Sometimes the bird will rest between the two phases with the body partly im- merged, feathers ruffled, and tail spread on the water, giving the bird a floating appearance. When leaving the water, flight is labored. A bird taking a complete bath is reluctant to leave the water when chal¬ lenged by a conspecific, whereas a dipping bird will leave immediately. In captivity bluebirds bathe daily. It is generally a social affair, for when one bathes, the others usually follow. Low ranking birds in the social hierarchy, when thoroughly wet from complete-bathing, were often attacked by dominant birds. Wintering bluebirds bathed in ditches along highways, swampy areas, and puddles in fields. As many as seven were seen bathing at once. (Also see Thomas 1946, pg. 171.) The alternate wing movements reported by Nice (1943), Coutlee (1963), and Andrew (1956) were not observed in the bluebird. Drying Upon reaching a perch after bathing, bill-wiping, head-scratching, head-rubbing, and preening are common. The body is ruffled and shaken repeatedly and excess water is removed from the plumage by a variety of wing and tail movements. The wing is extended sideways and forwards, carpometacarpus and carpals raised. The head is bent beneath the exposed wing and the ventral surface of the remiges are preened from base to tip, the bird [16] giving a slight flick of the wing as the tip of the feather is pulled through the bill. The bird preens the opposite wing, then rapidly alternates between both wings. Interspersed with underwing preening is a rapid opening and closing of the wings and tail as the wings are repeatedly flicked outward and the tail is fanned. In synchrony with underwing preening and lateral flicking of the wings and tail, is an intense whirling of wings that almost lifts the bird from the perch. The bird may interrupt drying movements and fly rapidly around the cage. As the plumage dries, the bird perches in a fluffed position and continually shuffles the wings and performs other cleaning move¬ ments such as preening, head-scratching, rubbing, and bill-wiping. Sunning Bluebirds show two responses to the sun (infrared lamp in cap¬ tivity). In the first the crest is raised, the plumage is ruffled, and the body is directed at right angles to and tilted away from the light source. The bill and eyes may be open. This is essentially the level III of Hauser (1957) except that the wings are not drooped, the tail is not spread, and the feathers are ruffled. Fig. 7 Eastern Bluebird Sunbathing (Level IV) [17] In the more exaggerated (level IV of Hauser, op. cit.) form, both wings are completely spread (with each primary being exposed) and somewhat drooped, the tail is spread, crest raised, and body plumage ruffled. The bird lies in a horizontal position on the perch with the head cocked to one side. The bill and eyes may be open. Any part of the body can be directed toward the light source. There is some variation in the degree of spreading the wings. If only one is spread it is the one toward the light source and may be spread in a series of hitches as it moves toward the light until fully spread. Spread¬ ing of both wings is more common. Hauser (op. cit.) mentions the social quality in sunbathing. In cap¬ tivity as many as four bluebirds would sun together. In the wild, a pair on territory would often be observed sunning together. While sunning, the bird may perform other maintenance activities without leaving the original perch. A 14-minute recorded sequence for a captive female is : bird lands 1 5 inches from infrared lamp — scratches side of head — rubs head across perch — sunbathes (level HI)- — -nods, scratches bill — sunbathes (level HI) — pivots — sun¬ bathes (level IV) — preens — scratches crown — scratches side of head — preens breast — sunbathes (level HI) — scratches side of head — sunbathes (level IV) — scratches chin — nibbles wrist — sunbathes (level IV) — scratches crown — sunbathes (level HI) — preens primaries — scratches chin — nibbles wrist • — stretches both legs — ruffles feathers and shakes — stretches wing and leg — bird leaves. Anting Although captive birds were fed ants (Camponotns sp.), they were never observed to ant, i.e. place ants among their feathers. Whitaker (1957), in her resume of anting, listed all the other North American thrushes, except the bluebird, as anting. FEEDING AND FORAGING Drinking Bluebirds normally do not enter the water to drink. They remain at the edge, the bill is dipped into the water and opened, there are throat movements. The head is tilted back and with the bill slightly open there are swallowing movements. Usually three to seven sips are taken at each visit. Head flicks can be given between sips to re¬ move water from the bill. [18] As many as six wintering bluebirds have been seen drinking at once from ditches and swampy areas along the highways. Caged birds pre¬ fer to drink from running, rather than, standing water. Power (1966) when studying 6". currucoides never observed the birds drinking and felt this need was satisfied by animal juices and green plant food, but Criddle (1927) reported that to induce 6'. currucoides to nest one must provide a nesting box and a watering place for the birds to drink and wash. Feeding The bluebird feeds on both animal and vegetable matter ; Beal (1915), Bent (1949), Forbes (1880), Forbush (1929), Hamilton (1943), and Hartshorne (1962) list the foods consumed. The bluebird takes its food mostly from the ground but captures are also made in the air and from the outside foliage of trees and shrubs. Exposed lookout posts afford a clear view of the surrounding terrain (see Preston and McCormick’s (1948) report on the eyesight of the bluebird) . Once prey is detected there are a few wing beats, the bird glides to the ground, landing with wings raised, seizes the prey, and flies back to the same or adjacent perch. Sometimes they hover just above or skim the top of the ground to make captures. Bent (1949), Power (1966), and Criddle (1927) report that the mountain bluebird (S’, currucoides) hovers in the air like a sparrow hawk (Falco sparverius) before descending to make a capture. I have never seen this type of hovering in S', sicilis. The prey is eaten immediately on the ground or carried in the bill to a perch where it is either “ scissored ” (a rapid sideways movement of the prey back and forth between maxilla and mandible) or “ ham¬ mered ” (beating the prey over and over against the perch). Some¬ times the bird will combine both methods. In hammering a large insect (e.g. June Bug) the appendages are often broken off, the elytra removed, and the body swallowed whole. A controlled fall occurs when the bird is “ working ” the prey on a high perch (telephone pole) and it is dropped or escapes. The bird falls vertically with fluttering wings, sometimes recapturing the prey before it reaches the ground. Power (op. cit.) reported that S', currucoides always eats the prey on the spot of capture unless feeding young. Perches are usually utility lines, fence posts, bare tree branches, small bushes, and even “ weed ” stalks. When feeding in pine for¬ ests, (Sprunt, 1954; pers. obs.) bluebirds will use the lower branches for perches but seem to prefer the bare trunks of the trees as they perch with the body parallel to the ground. When using a utility [19] line the bird either perches in the typical round-shouldered position (Peterson, 1961), pivoting frequently, or directs the body along the wire in a semicrouch as it peers down. The head may be cocked to one side in either position. Two methods of fly catching are used in foraging. One is to snap at an insect passing a fixed position ; the other is flying from the perch to make the capture in midair. I have never seen 5. sialis make more than one capture in the air and Marshall (1957), observing both sialis and mexicana, reported only mexicana making several captures in one flight. Bluebirds sometimes feed on the ground in a manner similar to the robin (Tttrdns migratorius) , (Bent, 1949; pers. obs.). When thus engaged I have never seen them use their bill or feet to turn over foliage in search of food as reported by Baird et al. (1905). When feeding in winter flocks along the highway berm the birds are spread out, mainly along utility lines. There is a constant drifting movement of subgroups and individual birds within the flock as they move back into cutover pine forests to feed on sumac (Rhus sp.) and mistletoe (Pharadendron fiavescens) , before returning to the utility lines. Drifting continues throughout the day as the birds alternate between feeding, digesting, and maintenance activities. Stoddard (1963) lists the bluebird as being especially attracted to recently burned-over ground. While feeding on sumac, the bird usually perches on top of the spike and bends the head down, pecking the fruit around the feet. The bird may, however, perch on the stem and peck forward or upward at the fruit. Sumac and adult stoneflies such as Taeniopteryx nivalis, are an important food source during spring snow storms (Krieg, 1962 ; Eaton, pers. comm.). Defecation For the first few days of nestling life the parents will eat the fecal sacs (Hartshorne, 1962; pers. obs.). A fecal sac is not presented at each feeding. At 5 days the young back to the nest rim, lift the rear, and spread both wings as the sac is voided. This is similar to behavior described by Dilger (1956b) for a young olive-backed thrush (Catharns nstulatus) except ustulatus does not spread the wings. Adult birds spread the belly feathers, lower the rear, and lift the tail slightly when defecating. Defecation in fear-producing situations (e.g. holding the bird in the hand while banding) rarely occurred. [20] LOCOMOTION Nestlings crawl using their wings as legs. At 14 days the bird can right itself and make short shuffling movements forward and back¬ ward. Hopping is well developed by the 16th day. Movement is straight ahead and there are no conspicuous corresponding head and tail movements. Hopping can be rapid or slow. Bluebirds move along a perch by hopping sideways, sidling (a lateral movement with short alternate steps), and pivot-hopping. In pivot-hopping the bird hops and simultaneously turns its body through a 90 to 180° arc, repeating this, as it moves first one way then the other along the perch. This is similar to “ pirouetting ” (Ficken, 1962b) in the American redstart except the bluebird does not use it for foraging. When fully drenched from bathing, bluebirds will climb (with cor¬ responding wing beats) up the cage wire. During molting they will run along a semivertical branch to reach a higher perch. Pivoting, without lateral movement, is common while perching. The young leave the nest at approximately 18 days (Hartshorne, 1962; pers. obs.) and can fly short distances (sometimes up to 100 yards). Thomas (1946) reported nestlings, out only a day or two before they can fly, are able to get into a tree by climbing the trunk. ROOSTING Resting In captivity bluebirds spend long periods of time resting in con¬ cert. The highest perch is used and an individual distance of 2 feet is normal. The body feathers are fluffed, the head is drawn in and may be turned toward another bird, the bill may be a slight degree upward. Sometimes the legs are fully flexed so the bird rests with the abdomen on the perch. Either foot may be tucked up into the ventral feathers. The eyes may be closed for short periods of time and yawning occasionally occurs. Rehearsed song (Lanyon, 1960) is given repeatedly. Sleeping Young nestlings sleep with the head drooped down over the abdo¬ men. At 5 days the head is held limply in front or curled to the side. [21] Soon they begin to sleep with the head and neck either retracted or extended, and the head held a little above the horizontal. These two positions persist until the 12th day. Laying the head on the scapulars was first seen at the age of 9 days and by the 13th day the young all slept in the typical adult manner described by Dilger (1956b) and Nice (1943). Roosting During the breeding season pairs do not normally roost inside the nest box. The female will, however, spend the night inside the box, throughout the incubation and fledging periods, but rarely will be joined by the male (Hartshorne, 1962; pers. obs.). In cold weather the pair sometimes roost inside the box or even join other bluebirds and roost in one box. As Thomas (1946) and my observations show, pairs that fight over a territorial boundary may roost together at night during cold weather. Wintering bluebirds, however, will roost communally and a few observations on roosting behavior were made in the Apalachicola Na¬ tional Forest, 15 miles south of Tallahassee, Florida. Two roosting sites were used. One was the top branches of pine trees {Finns sp.) located in a cutover area 100 yards from the high¬ way. The flock fed in this area during the day and went to roost at sundown in a drifting manner with small groups of two to three or single birds leaving the utility lines and flying to the pines. More than one bird would try to roost in the same branch causing consider¬ able unrest, squealing, aggressive calling, and pushing as birds moved from one tree to another. Sometimes two birds would topple off a branch, grasping one another, and fighting part way to the ground. By darkness the birds were settled and the whole flock (25 ± ) roosted in six or seven medium-sized pine trees growing close together. By shaking the trees at dark, an estimate could be made of the num¬ ber of birds roosting on one branch. Seven birds came out of one treetop but the usual number was one to three. The other roosting site was an electric power substation located next to the highway approximately 2 miles from the one previously described. When first discovered the weather had turned cold (Feb. 23, 36°; Feb. 24, 33°; Feb. 25, 26°; Feb. 26, 21° (low for month and winter) ; Feb. 27, 26°) and it is not known if the birds roosted at this site during warm weather. At sundown small flocks (four to 10 birds) gathered on utility lines along the highway and silently, or with occasional location notes, flew into the power station area to roost. Some roosted on the labyrinth of wires, condensers, and beams [22] inside the station fence but the majority preferred roosting on the top of a steel I-beam that held three large transformers. As many as seven at once flew silently up under the same transformer. Due to the difficulty of observation, the number, posture, and arrangement of the birds could not be determined. The transformers gave off a continual “ hum ” and probably some heat suggesting that birds roosting under them in cold weather would benefit from this heat and shelter. Frazier and Nolan (1959) reviewed and described communal roost¬ ing in the bluebird. Their report was based on observing five to 14 bluebirds roosting together in a nest box during cold weather. I ob¬ served this same behavior (five birds involved) at Wing Hollow, Allegany, New York on March 27, 1966. The temperature was 22° F. and there was 3 inches of snow. On checking the nest boxes at night, box 1 1 S contained five bluebirds, two pairs that had ad j oining territories and a migrating male. All five were on the floor of the box with heads facing the south-east corner, bills straight ahead and somewhat raised ; the birds were crowded together with those in back snuggled in between with their bills on the side of the heads of those in front. This arrangement is different from that reported by Frazier and Nolan (op. cit.) where their birds slept with heads together and bodies pointed downward, forming an inverted cone. In captivity, preferred roosting spots were unpredictable. Birds would seldom roost side by side but at times as many as four roosted together. As the birds began to roost much unrest and fighting took place. The fighting seemed due more to violation of individual dis¬ tance than over a “ select ” roost. Two birds would fight violently on a particular perch with the winner (after some settling movements) leaving, only to try to supplant another bird on a different perch. Movement, therefore, was aggressively from perch to perch. A bird appeared to have no preferred roost or partner. However, in the winter of the second year in captivity, the flock was reduced from eight to four birds and these four (two pair that had bred in captivity) did show a preference for a particular roost. The one female roosted inside a nest box (the only time in 2 years of observation that a bird roosted inside a nest box) with the male on a perch next to the box. The other male and female roosted on the other side of the cage, each with its preferred perch (not side by side). [23] #^|\ii^H '^-'M . . rfe{Vq,ipii!fe »f^i^jr! -jc-iroi *cfcfxidonqi>fli^ " muA .i^«U bftfc ijib- mJ(«^ i^7rt5td J>f»Josn iiflifiow bko jui ggmIjfUJOMrw ;^:K{1?^ i^ k>Ym>¥^ (Oacr } tuihn fMW laiiTj^ apttiy^^o «di tti my ;ov^viJtttSUitufifo*«^ tjo ' >f rr? cr ov MrfSbfc r.3?itis sw/idiftbA r Uigit«ttbw<>3> ftbtjd 35^dJtt4«9W>4rf<^49K-^ »i>itlii#i»i^t atianal om M 10^ 3r;.m ir , nk. ?*5r.2f*‘ i 21"^ now ua j^ooiH) 4r4>^1i}(N')'; fm ^ kfiiiWh' if (Hc VioJ* tvahA- ; • JW\uf1ov\ro '*et;»U flocks i6 1^^) «»f tlflsy itnfcff »)qAh tUK^^wsy sM siktittj^* W. rflif WUiw ltJ»o fhe I’oww Italian- Iir«a mc on tk^ OOn.liHiscrft, tud ' m\ " AGONISTIC BEHAVIOR The word “ agonistic ” comes from the Greek root agon which means “ to struggle ” and refers to behavior concerned with attack and escape. The term agonistic behavior, as used in this study, has been described by Scott and Fredericson (1951) and includes the gen¬ eral group of behavioral patterns that involve aggressive and defensive fighting, escape behavior, and passivity. Such behavior results from variations in the actual and relative strengths of the attack and escape tendencies. It is assumed that these tendencies are usually, if not always, simultaneously present (Moyni- han, 1955b) but either may be so predominant that the animal simply flees or attacks the opponent. However, as these two tendencies are incompatible (Tinbergen, 1940) the animal is usually in some state of conflict resulting in various displays depending upon the actual and relative strengths of the two conflicting tendencies. The agonistic displays of the bluebird will be described and their context given. Where possible, the cause and function of the display will be postulated according to the methods described by Tinbergen (1959). Daanje’s work (1951) forms the foundation for describing the derivation of display components. The following definitions of ethological terms are essential to this investigation : * Display — Those peculiarly standardized and often exaggerated performances, including all vocalizations and many movements and postures, which have become specialized and modified as social signals or releasers. (Moynihan, 1955a) Ritualization — The evolutionary process responsible for the exist¬ ence of inter- and intra-specific signaling movements. (Blest, 1961) _* The names of stereotyped display-activities are ordinary words used hereinafter in a special sense, hence are capitalized. [25] Intention movement — Applies to various incomplete and low in¬ tensity movements, so called by Heinroth because they reveal what the animal is “ intending to do ” to the observer. (Marler, 1956) Tendency — ^Is used in the sense of Hinde (1955-56) as given by Marler (1956) : . .the readiness to show a particular type of behavior as observed under natural conditions.” Feather postures — (Morris, 1956a) : Sleeked — The feathers are fully depressed against the body, giving it a slim appearance. Fluffed — The feathers are erected, but only partially, giving the body a very rounded appearance, with a smooth, unbroken outline. Ruffled — The feathers are fully erected and the body, although rounded, has a ragged appearance, with a broken outline. ATTACK RESPONSES Supplanting Attack The Supplanting Attack of bluebirds is similar to that of most pas¬ serine species. The aggressive bird flies toward the perch occupied by another bird, which vacates its perch. Supplanting Attacks, often accompanied by Bill Snaps, may occur in rapid succession resulting in a prolonged Chase. Supplanting Attack reflects a strong tendency to attack, and as Hinde (1955-56) and Marler (1956) suggest, a more confident ex¬ pression of hostility than any other form of agonistic behavior. This “ confidence ” is shown by both captive and wild bluebirds. In captivity Supplanting Attacks were the commonest expression of aggressiveness as shown by female DG who made 41 Supplanting At¬ tacks upon female W within a 1-hour period. Subordinate birds often left their perch just as the aggressor started to fly. Sometimes this resulted in aggressive Chases with the submissive bird being Sup¬ planted as soon as it alighted on perch after perch. These aggressive Chases were terminated either by the aggressor “ losing ” interest or by the subordinate refusing to fly and showing defensive threat or some other willingness to Fight. Some Chased individuals landed close to a bird dominate to the chaser thereby ending the Chase with the aggressor perching quietly nearby. In the wild, a territorial male will Supplant any bluebird it finds in its territory; females often do the same. The Supplanting Attack [26] is extremely important during pair formation because the pair bond is not formed until, leaving the nest box, the female Supplants the male. As the courtship period progresses this act of dominance by the female becomes more intense. Collisions occur when the bird being Supplanted is highly motivated to stay, i.e. between members of a pair during early pair formation, e.g., females will fly at and knock males off the top of the nest box. Both sexes make Supplanting Attacks on other species that loiter near the nest box. Chases Chases occur during skirmishes between males within and at the boundary of the territory. These appear to be lowly motivated if the birds know one another (i.e. hold adjacent territories). In such cases there may be a series of Supplanting Attacks each followed by a silent, slow, short Chase as the birds move along a fence line or hedge row. If an unknown male (or pair) moves into the area the Chase is more violent and sometimes prolonged over 300 yards. The pursuer may fly at top speed, dive, and catch the pursued in midair, then topple to the ground. Fighting. Most Chases occur between males but females have Chased other females out of their territory. After a Chase the male may return and land by his female or nest box and start to sing. Chases appear to be nonritualized for the pursuer flies at top speed and tries to catch the pursued, often succeeding, resulting in a violent Fight. Low motivated Chases were rarely seen in captivity ; rapid aggres¬ sive Chases were the rule, as a dominant bird Chased a subordinate around and around the cage. Fighting The average Fight took place between males or between females, as stated by Thomas (1946), but when I placed a live or mounted decoy in the territory of an established pair, both members of the pair attacked whether the decoy was male or female. In the aviary both members of the pair attacked intruding birds of either sex. In the wild. Fights between male and female were occasionally observed. The situation here resembled that described by Lack (1953, pg. 57) with the English robin (Erithacus rubecula). During contact, both combatants may strike with their wings, grapple at one another with their feet, and peck viciously at each [27] other’s head. On April 14, 1966, one combat lasted for 15 minutes; the two males, feet locked together, wings outstretched, and bills open were Pecking at one another as they tumbled about in a ditch. Experiments with decoys and mirrors showed bluebirds strike one another with both the beak and feet, as do the English blackbird, Turdus meriila (Brown, 1937) and the American robin (Young, 1951). In this study Fights never resulted in real injury but Pettingill ( 1936) reported a female with the crown featherless and bleeding as a result of a Fight with another female and Blake (1954) reported a Fight between two females in which one died. In captivity where escape was sometimes impossible, wounds were inflicted. On one occasion male DY had invaded pair GY and DR’s cage through a hole in the wire. When the cage was examined, male DY was wedged in the corner, his head protected by an overlapping 2x4, with both male GY and female DR Pecking him viciously in the shoulder region that was completely featherless with a raw wound. Fights between members of a pair are frequent. The male lands on the female’s back, violently Pecking her head several times in suc¬ cession. Some females will Fight back but the normal response is passive ; the female crouches with closed eyes as the male Pecks. These attacks are not strictly agonistic but have a sexual tendency in conflict with the attack and escape tendencies and are discussed under court¬ ship. If perches are lacking during a boundary dispute, birds may Fight on the ground like robins. They face one another with sleeked plum¬ age, one bird flies up about 3 feet with wings and tail spread and lands, then the other flies up and lands. This is repeated several times and ends either in combat as both fly up and meet in midair, or by one or both returning to guard the nest box or female. Sometimes Fight¬ ing is interrupted by a series of zigzag hops by both birds that re¬ sembles “ pushing ” in the American robin (Young, 1951). Thomas (1946) also mentioned this type of Fighting ”... the males were on the ground within a few feet of each other, teasing at grass. They came face to face, and there was a brief encounter, the two jumping like little cocks; then they hopped in opposite directions and pecked at the ground ; . . . ”. Pecking When individual distance is violated, one bird may turn and Peck the other. These Pecks seem weak but usually cause the recipient to flee. [28] During courtship when the males attack tendency is highly moti¬ vated, he lands on the female’s back and delivers a series of Pecks to the top of her head. These are impetuous but apparently never harm the recipient. Repeated Pecking is also seen when two bluebirds are fighting. Both captive and wild bluebirds bite when held in the hand. THREAT POSTURES AND DISPLAYS Facing This is the simplest act of threat in the bluebird and involves turning the head to face the opponent. Facing is usually seen when individual distance (Conder, 1949) is violated among birds in a flock or between members of a pair. In its lowest intensity the bird merely looks at the opponent with no noticeable feather adjustments. However, if the opponent ap¬ proaches, the aggressor will sleek the head and neck feathers and lean somewhat toward the approaching bird. In its highest intensity the threat valence of Facing is reinforced by Gaping, Bill Snap, or Rasp and may even develop into a lunge at the opponent. Gape In this display the bird turns the head toward the opponent and holds the bill open for several seconds. The closer the opponent ad¬ vances the wider the bill is opened. In its highest intensity the head is sleeked and the bird leans toward the opponent with the bill held open and finally, may move toward or lunge at the other bird. Gape reflects a stronger tendency to attack than to flee and is often reinforced with Bill Snap and Rasp giving the display a strong threat valence. In captive birds this is a common display when individual distance is violated and during such social activities as feeding, bathing, and sunning. In wild birds it is most often seen during encounters between members of a pair, especially during pair formation. For example, when courtship feeding commences the female will often Gape as the male approaches with food or when the male advances to copulate. Gape is probably derived from an intention movement to bite. Wings-fiicked In Wing-flicking the bird perches in an oblique position and rapidly flicks both wings out from, and back to the body. The wings are com- [29] [30] Two Bluebirds involved in an agonistic encounter. The bird on the left Facing while the one on the right shows Head Forward (stronger attack tendency) . Both displays are reinforced with Gape. pletely unfolded and are flicked simultaneously to a plane level with the body. Tail-fanning and Chatter usually accompany this display. In the wild, Wing-flicking was employed by both sexes, usually when a live decoy was introduced into a pairs’ territory. The pair became greatly disturbed and one will start Wing-flicking (usually ac¬ companied by Tail-fanning and Chatter) as it perched within 5 feet of the decoy, usually not facing it. Wing-flicking may then give way to attack. It is also employed in interspecific encounters as when trying to dislodge a starling (Sturnns vulgaris) from a nest box. Dilger (1956a), describing a similar display in Hylocichla mustelina and Catharus sp., stated that Wing-flicking was indicative of low in¬ tensity attack and escape drives and was employed in a vertical plane. The bluebird differs in that Wing-flicking appears to reflect a high attack tendency, and Tail-fanning accompanying the Wing-flicking is given in a horizontal plane. In captivity when a bird became extremely aggressive and started to harass a subordinate, Wings-flicked was the predominate display. The aggressor would continually give Chatter, Wing-flicking, Tail- fanning and then Supplant and Chase the subordinate. Wings-flicked is probably derived from a flight-intention movement. Tail-fanning The tail is rapidly fanned in and out in a horizontal plane. Tail- fanning never occurred alone, but was always employed with Wings- flicked. Tail-fanning occurs in the same contexts as does Wings-flicked and is probably a flight-intention movement. Wings Out The bird faces its opponent, with legs fully flexed, body horizontal, plumage sleeked, and the wings held out horizontally to the side. The degree that the wings are lifted horizontally from the supporting feathers varies with the attack motivation of the bird. In bluebirds this display was performed by both sexes (wild and captive) in response to Scream given by a bird held in the hand. The display reflects a strong attack tendency and after adoption a strong aerial attack is given as the bird, uttering Chatter and/or Bill Snaps, dives at the adversary. Wings Out was only observed in response to Scream and was never seen to be directed toward a conspecific. To typify this : July 11, 1967 ; St. Bonaventure Aviary. Six bluebirds are in an outdoor flight cage ; all show different stages of postnuptial molt. [31] Fig. 9 Wings Out threat display of Eastern Bluebird Fig. 10 Eastern Bluebird in Head Forward display [32] ! I Male Y is captured (to examine the progress of the molt) and while being handled, he Screamed. The other five birds immedi¬ ately line up on a wire perch in varying degrees of the Wings Out display. Uttering Chatter, they now start to mob me. As they return to the perch Wings Out is again adopted. This display is similar to the Wings Out Display described by Ficken (1962a) for the redstart (Setophaga ruticilla) except in that species it is directed at conspecifics. The flexed horizontal body with sleeked plumage and extended wings are all flight-intention movements and express a strong attack tendency. Head Forward The bird faces the opponent, body horizontal with head somewhat retracted, legs flexed, and tail slightly lowered. The body feathers usually are sleeked and rarely are fluffed. Gaping usually accom¬ panies this display and Bill Snap and Rasp are common. Head For¬ ward often precedes the Supplanting Attack. In captivity Head Forward often was unsuccessful in causing an opponent to flee and even in its most intense form (orient toward, plumage sleeked, Gape, Bill Snap) the displayer often was Supplanted ; thus suggesting a weak threat valence. Head Forward is a common threat posture in the passerines (Andrew, 1961) and is similar to the Horizontal Stretch of Catharns (Dilger, 1956a) and possibly the Attack Run of Tiirdiis migratorins (Young, 1951). Bill-raising Bluebirds have two postures that involve raising the bill above the horizontal. a. Oblique Bill-up — In this display the body is oblique with the head, bill, and tail in line with the rest of the body. The tail may be slightly lifted. b. Horizontal Bill-up — The body is horizontal, legs extended, neck somewhat stretched, the head and bill are pointed upward. In both displays the tail is not spread and Gaping may occur. Normally the plumage is compressed but this varies with the strength of motivation. When the bird is highly motivated (a strong attack tendency in conflict with a tendency to flee or to feed young) the neck and head are extremely sleeked with the belly and breast feathers remaining fluffed. In this conflict state the bird may alternate between the Oblique and Horizontal Bill-up showing ambivalent movements [33] [34] Fig. 11 Eastern Bluebird Bill-raising display. On left — the Oblique Bill-up; on right — the more aggressive Horizontal Bill-up. toward and away from the opponent. The sleeking of the head and neck reflect a stronger tendency to attack than to flee. When the bill is raised the two black chin stripes (occurring in both sexes) are exposed to the opponents’ view. However, the Bill-up Display has a weak threat valence for the opponent rarely flees. The Horizontal Bill-up is not as common a display as the Oblique Bill-up and was seldom seen. When observed it seemed to reflect a shift in the strengths of the attack and escape tendencies, with attack seeming more dominant. Bill-raising is not always directed toward the opponent for when in this posture, a bird can face the opponent, stand parallel to it, or even face away. The displaying bird can incorporate movement into the Bill-raised Display by sidling, pivoting, or pivot-hopping toward the opponent ; or when in conflict show intention movements to both approach and flee by hopping in a circle. In the wild, Wings-flicked and Tail-fanning were never employed with the Bill-raised Display. In captivity, however, both were some¬ times used with the Oblique Bill-up. This indicates a stronger attack tendency as the bird adopts the Oblique Bill-up followed by Wing- flicking, Tail-fanning, and Chatter. The bird then flies at and Sup¬ plants the opponent. The following extracts from my notes provide typical observations involving Bill-raising. June 30, 1967 ; Wing Hollow ; Pair 3S. A live decoy is placed on a fence post approximately 40 yards from the nest box. A predator had just destroyed the nest. Male 3S flies and lands in front of the cage. He assumes an Oblique Bill-up and keep¬ ing this posture moves in a circle on the fence post then pivots and starts to move away ; assumes another Oblique Bill-up, nods, and touches the tarsus ; assumes a Head Forward and flies closer to the cage. Assumes an Oblique Bill-up with breast fluffed then turns away, down into a horizontal position, back to Oblique Bill-up, nods. Oblique Bill-up, flies over cage, assumes Oblique Bill-up with head and neck sleeked, now assumes a Horizontal Bill-up and flies closer to the cage. February 4, 1966; Indoor aviary; St. Bonaventure University. Female DG assumes Oblique Bill-up then rapidly gives Wings- flicked, Tail-fanning, and Chatter. Now DG Supplants female W. After Supplanting W, DG assumes Oblique Bill-up, Wings- flicked, Tail-fanning then repeats the Supplanting Attack. January 16, 1967 ; Indian Springs Camp ; Leon County, Florida. 3 :20 p.m. Bluebirds are spread out on utility lines along Rt. 369. Suddenly a male flies in and Supplants another male who then moves a short distance. The aggressor now assumes a Horizontal [35] Bill-up, gives a Location Call then flies at and Supplants the same male again. 3 :25 p.m. Two females are close together on the utility line and one assumes an Oblique Bill-up and Gapes. The other female leaves. 4:10 p.m. A female on a utility line assumes an Oblique Bill-up and sidles laterally toward another female who then leaves. May 18, 1967; Wing Hollow; Pair ION. 5:15 p.m. Boundary dispute with pair IN. Two males are on utility line and male IN adopts an Oblique Bill-up as he faces male ION. Male IN assumes another Oblique Bill-up then a Head Forward as he goes and Supplants ION. After attack IN assumes Oblique Bill-up but is now lateral to ION ; IN pivots while still in the Bill-up. Female IN comes to utility line and male IN moves back and guards female. Female ION comes and two pair move closer together and male IN adopts Oblique Bill-up then three more in succession (now head and neck are sleeked with body fluffed). Male ION sits Fluffed by his female. Now both pairs leave. The Oblique Bill-up is similar to the Upward of Catharus studied by Dilger (1956a) except that Catharus do not sleek the head and fluff the body feathers nor do they incorporate movement into the display. Hartley (1949), however, describes a similar posture in the mourning chat (Oenanthe lugens) where the belly and lower breast feathers are fluffed. Turdiis merula and T. migratorius (Andrew^ 1961) spread the tail in the Bill-up Display while Erithacus ruhecula (Lack, 1953) in a similar display raises the tail. Bill-raising is probably derived from an upward flight-intention movement. VOCALIZATIONS Bill Snap Bill Snap is given (by both sexes) in interspecific and intraspecific encounters. It usually accompanies the Supplanting Attack but can be given with stationary displays such as Facing, Gaping, or Head Forward. Bill Snapping reflects a strong attack tendency and is presumably, a ritualized intention movement to bite. It occurs in the thrushes of the genera Catharus and Hylocichla (Dilger, 1956a) and in the American robin (pers. obs.). [36] Screech Screech, a single harsh sound given by captive birds, was rarely heard. It is usually uttered when individual distance is violated and may accompany Facing or Gaping. Rasp Rasp is a loud nasal sound which can be repeated. It is given in the same situations as Screech and is often uttered by birds engaged in combat. Rasp can accompany a Supplanting Attack but this is rare. Rasp indicates an attack tendency and seems to function as a threat since it often stops the advance of the other bird when individual dis¬ tance is violated. Squawk In captivity. Squawk is given by a bird being Chased. It is a single vocalization which may be repeated as the Chase continues. Birds caught by the pursuer and sometimes bitten, would utter Squawks. Similar vocalizations are given in much the same context in such diverse species as Stellers jay (Cyanocitta stelleri) (Brown, 1964) and rose-breasted grosbeak (Dunham, 1964). Chip Chip, a low pitched soft sound, may be given by females when the male approaches during early courtship. Normally the female is in a Fluffed posture and Faces the male as Chips are uttered. At times Chips plus Facing quell the approach of the male thus indicating a weak threat valence. “ Turring This is a single high pitched “ turr ” which can be repeated to make a series of two to four such notes. Long pauses can occur between “ turrs ” as the bird remains motionless. “ Turring ” is given by both sexes in a variety of situations. Some¬ times the acts performed by the displaying bird following this vocaliza¬ tion seem to indicate a relatively strong attack tendency in conflict with a weaker escape tendency. The following excerpts from my notes typify this. May 14, 1967, Aviary, St. Bonaventure University. Male YW “turring,” he gives a high pitched (barely audible) “ turr-turr- [37] Fig. 12 The Crouch posture of the Eastern Bluebird Fig. 13 The Alert posture of the Eastern Bluebird, adopted during mild fright [38] turr ” call for 6 minutes from a frozen oblique posture then makes a violent attack upon female W. April 20, 1967 ; Wing Hollow. A live decoy is placed two posts away from the nest box of Pair 2 IS. The female is nest build¬ ing. Male 21 S sits in a tree and preens then flies to a fence post by the decoy and assumes an Oblique Bill-up, pivots, assumes a Head Forward, pivots, then makes ambivalent movements toward and away from the decoy. The male now flies to the top of the cage (decoy inside) and becomes quiet, looks around then gives the high pitched “ turr-turr-turr ” over and over, then attacks the decoy. “ Turning ” is also given during territorial disputes and sometimes, when engaged in a Song Duel, one male will cease Warbling and start “ turning.” In the wild, “ turning ” may be given in such a muted form that the observer can only detect it by seeing the pulsating throat feathers every few seconds as the bird remains motionless. “ Turning ” is frequently heard during early pair formation. “ Turning ” is also given in situations that suggest a strong escape tendency thwarted by physical factors, such as a cage or being held in the hand. For example, female DR had laid an egg on the floor of the cage and as I opened the cage to remove it she flew out. After much effort the female was caught and returned to the cage. She then assumed a frozen oblique posture and kept giving the single high pitched soft “turr” over and over (13 “ turns ’’/minute). Some¬ times birds that have just been caught and are being hand held will give this soft “ turning.” In this situation there are no pauses between each “ turr.” It may be possible to subdivide “ turning ” on the basis of form and function into different vocalizations when more observations are available. Chatter Chatter is a loud continual “ chit-chit-chit ” uttered in rapid suc¬ cession but can, during mild alarm, be given as a single harsh “ chit.” Chatter has a low alarm threshold and may be given in response to any general disturbance, e.g., ground predators (cats, dogs, Eluphe ohsoleta, Peromyscus sp., mounted saw-whet owl) ; novel stimuli introduced into the cage (aquarium, sumac) ; disturbances around the nest boxes (humans, starlings, cows). Chatter given in these contexts seems to indicate a stronger escape tendency in conflict with a tendency to approach (novel stimuli) or to attack (predators). The birds will Chatter as they hover over the form of stimulus or will fly rapidly to the top of the cage, clinging to the wire and Chattering. [39] Chatter is also given by both sexes in intraspecific encounters. In this context Chatter usually accompanies Wings-flicked and Tail- fanning Display but may, depending on the motivation, be associated with Bill-raised Display, Supplanting Attack, or Chasing. In these situations Chatter seems to indicate a stronger attack tendency in conflict with a weaker escape tendency. This was especially evident in captivity where dominant birds would harass subordinates by repeated Supplanting Attacks preceded by Wings-flicked, Tail-fanning, and Chatter. Territorial males, involved in disputes with neighboring males or a new female will Chatter in flight as they fly to a nest box, song perch, or territorial boundary. Chatter in these instances can precede and become continuous with Warble. Chatter was observed to be associated with panicking only once. A live Cooper’s hawk (Accipter coo peril) was brought into the indoor aviary and all eight captive bluebirds immediately panicked. There was an outburst of wild activity as the birds flew around and around the cage, often striking the sides. Chatter was given continuously. Scream Scream was only heard to be given by a captured bird. While being held in the hand for banding or examination the bird uttered a loud harsh Scream. Scream may be associated with an attack tendency for while Screaming, the struggling bird will often bite the captor’s finger. Also, Wings Out Display, associated with a high attack tendency, was the only display seen in response to Scream. ALARM BEHAVIOR Crouch Young bluebirds inside the nest box will Crouch as a response to the parent’s Chatter, jarring of the nest box, and opening of the top. They flatten themselves against the bottom of the nest, close their eyes, and remain motionless. If removed for banding, they immedi¬ ately assume this posture when placed back in the nest. Captive adults may Crouch while giving the Flying-predator Alarm Call, when frightened, or when unable to locate a source of disturb¬ ance. The body is lowered as the legs are flexed, neck extended below the body level, throat fluffed, plumage relaxed to sleeked. Birds in indoor flight cages often adopt this posture as they peer out windows or try to locate unusual noises. [40] Crouching has also been observed in the wild. Females when incu¬ bating or brooding will Crouch as the top of the box is opened and both males and females, after being trapped inside the box, will flatten themselves against the bottom of the box before being removed for banding. Birds on utility lines will sometimes Crouch as they appear to scan the horizon. During early pair formation, when the males are extremely aggressive toward the females, the female sometimes will, upon seeing the male approaching. Crouch with sleeked plumage as the male lands on her back and violently Pecks her head. During interspecific encounters bluebirds sometimes combine Crouch with elements of attack when showing defensive threat (see page 29 for explanation of defensive threat). When being mobbed by tree swal¬ lows (Iridoprocne bicolor) bluebirds will Crouch on the top of the nest box, Gape, and Bill Snap (both elements of attack) as the swallows dive at them. In captivity, where escape is impossible, they sometimes show a strong but thwarted escape tendency. The bird will Crouch and flatten itself on the perch, head extended and slightly raised, eyes bulged, plumage extremely sleeked with the lateral breast feathers ruffled over the tightly retracted wings. The bird remains motionless peering at the source of stimulation. If startled or approached, the bird will fly explosively, hitting the sides of the cage hard. Crouching is similar to the Freezing posture described by Dunham ( 1964) for the rose-breasted grosbeak {Pheiicticus ludovicaniis) and by Ficken (1962a) for the American redstart (Setophaga riiticilld). An alarmed bluebird may also Freeze. Usually the body is held in an oblique position, plumage compressed, legs flexed, bill up some¬ what, and eyes bulged (see Eaton, 1914, pg. 538 for a picture of young Freezing). However, many times there is no characteristic Freezing posture as the bird remains motionless for varying lengths of time. Sleeking of the plumage and Crouching are both flight intention movements and exophthalmos may increase the field of view (Marler, 1956). Alert Alert is an expression of mild fright and appears to indicate a con¬ flict between the tendency to flee and the tendency to remain. The alarmed bird stands very erect with body feathers sleeked, pronounced extension of the neck, wings somewhat raised from the supporting feathers. The bird is silent, makes ambivalent movements, and may [41] alternate between a strict vertical and a horizontal position as it peers in the direction of the disturbance. This posture is assumed when the bird is confronted with a novel situation and upon habituation, gives way to a more relaxed position. Dunham (1964) pointed out the biological significance of this dis¬ play. He felt that it minimizes deleterious consequences and maxi¬ mizes possible advantages of novel situations. Wing-jiashing Wing-flashing (sensu Hailman, 1960; Horwich, 1965; Selander and Hunter, 1960) observed in captive birds, is similar to Wing¬ flashing in the mockingbird {Mimiis polyglottos) except, in the blue¬ bird, the wings are not “ flashed ” in a series of hitches. Monroe (1964) observed Wing-flashing in the foraging behavior of the red- backed scrub-robin (Erythropygia zambesiana) . This was the first report of Wing-flashing in the Turdidae that was identical to that in the Mimidae. Dilger (1956a) reported Wing-flashing in the Catharus sp. he studied but this appears to be quite different from Wing-flashing in the Mimidae both in form and context. The bluebird’s body is held in a normal position, head slightly extended, wings somewhat drooped. Both wings are brought up in a deliberate manner and fully extended horizontally or vertically, level with, or just above the body, held in this position for a second or two, then returned quickly to the sides. The tail may be slightly lifted and spread. Wing-flashing was first observed in captive birds, 4 months old when sumac (Rhus typhina) was introduced into the flight cage. The birds became frightened and began to Chatter and mob the sumac. They approached the sumac in an Alert posture with a hesitant man¬ ner, Wing-flashed, and finally Pecked at and ate the sumac. In captivity, this behavior was observed in many different situa¬ tions. Birds Wing-flashed to a variety of familiar and unfamiliar objects. The following, taken from my notes, will serve as examples. December 8, 1966; St. Bonaventure Aviary ; 12 :00 noon. Nailed a branch of sumac to the feeding table. The birds were in the out¬ side aviary and came in immediately. All line up on the wire perch and give Chatter. Females W, R, DG, Wing-flash. Female DG Wing-flashes as she lands on a window ledge, water tray, stove pipe, and perch. Others Wing-flash as they land to feed. 12 :30 p.m. Birds appear habituated to sumac. February 2, 1967 ; St. Bonaventure Aviary ; 12 :00 noon. A funnel trap with a song sparrow (Melospiza melodia) inside is put on the floor inside the aviary. Female DG hovers over the cage, lands next to it and Wing-flashes four times in a row, then bends over [42] and picks something off the floor, hops, Wing-flashes, hops. Wing- flashes, lands on top of the trap. Wing-flashes, flies to a perch and ruffles and shakes the body feathers. February 21, 1967; St. Bonaventure Aviary; 10:15 a.m. Put a live black rat snake (Elaphe ohsoleta) insides the flight cage. Female DG lands on wire above snake and Wing-flashes. Female DR lands on top of nest box and Wing-flashes, flies to perch and Wing-flashes. 10 :30 a.m. The snake is removed and male YW goes to perch where the snake had been and Wing-flashes twice. Birds keep hovering and looking in box 3 where snake was last seen. Although I looked for this behavior in wild bluebirds, it was seen only once. On January 16, 1967, in Wakulla County, Florida, a female bluebird, feeding along the berm of the highway, Wing-flashed then seized an insect in the grass. The function and biological significance of this display, or whether, in fact, it even exists in the behavioral repertoire of noncaptive blue¬ birds, remains obscure. In captive birds. Wing-flashing is sometimes given as an alarm reaction when the birds are confronted with strange objects or unfamiliar situations. It has also become individually con¬ ditioned and is given irrelevantly without being associated with alarm or fear situations. Fig. 14 Captive Eastern Bluebird Wing-flashing to sumac (Rhus typhina) [43] Mobbing Captive bluebirds less than a year old were observed to show Mobbing behavior in response to the following objects: a. High intensity Mobbing response shown to : Live black rat snake (Elaphe obsoleta) Mounted saw-whet owl {Aegolins acadica) Dead deer mouse (Peromysciis sp.) Sumac (Rhus typhina) b. Low intensity Mobbing response shown to : Mounted tree swallow (Iridoprocne bicolor) Mounted eastern bluebird, male (Sialia sialis) Mounted great horned owl (Bubo virginianus) Mounted red-winged blackbird (Agelaius phoeniceus) The first reaction during a high intensity response is immediate Chatter as the birds fly rapidly around the cage. As the escape tend¬ ency becomes less motivated the birds come to rest on a perch above the object. The plumage is sleeked and the birds may make constant restless movements as they pivot and fly to another perch or they may remain motionless and peer down at the object. Wing-flicking and Tail-fanning can be given as the birds start to hover above the object, uttering Chatter, then flying to a different perch. Diving attacks may be made. Sometimes subordinate birds would not participate in Mobbing but would assume a Fluffed posture and watch from a high perch in the cage. Females were more responsive than males, as males appeared more fearful of the objects and sometimes completely avoided them. When a bird finally did approach the object, it was slow and from the Alert posture ; there would be incomplete intention movements to fly and finally the object would be Pecked. Displacement feather-ruffling was common and Wing-flashing sometimes occurred. Chatter in low intensity response, if given, was weak and intermit¬ tent. A few birds would perch above the object, hover, then lose inter¬ est. Often one would approach in an alert posture. Peck the object, then avoid it. I never observed Mobbing (involving more than one pair of birds) in the wild, yet Sindelar (1967) reported that several eastern blue¬ birds dive-bombed a meadow vole until it took shelter. However, when I stood by a nest box after the young had hatched, the adults often delivered an aerial attack. This took the form of a Pendulous Display as the bird would hover in midair Chattering, then dive and [44] give Bill Snaps, return to a point in midair on the other side of me, and repeat the attack. To show this diagrammatically : Hovering/Chatter Hovering /Chatter Thomas (1946) mentions that bluebirds respond to the distress of other species (“. . . He will join a robin, Turdus migratorhis, in attacking a blue jay near the robins’ fledgling.”) but I have never observed this. DISPLACEMENT BEHAVIOR Tinbergen (1952) defines a displacement activity as “. . . an activity belonging to the executive motor pattern of an instinct other than the instinct (s) activated.” The problem of displacement is very complex and several hypotheses (Tinbergen, op. cit. ; van lersal and Bol, 1958; Rowell, 1961 ) have been presented concerning its cause. Several times throughout this study, bluebirds in agonistic encoun¬ ters performed “ activities ” that were irrelevant and out of context, especially during experiments with live decoys conducted on territorial birds. It is difficult to know when one is observing displacement activities (Dilger, 1956a) so these observations are tentatively considered as such because they were performed during a conflict situation and were typically incomplete and out of context. [45] Displacement Bill-wiping Commonly seen in agonistic situations, the movement is often incomplete with the bill not touching the perch. Displacement Preening Any part of the plumage may be preened in a nervous, jerky manner. Usually the feathers are not rufifled (as in preening) or drawn through the bill. The bird may make quick “ swipes ” at the feathers. Often the head remains sleeked and the bird may give Muted Warble. Displacement Feather-ruffling This is similar to the Spread Display of Hylocichla mustelina (Dilger, op. cit.) but is not ritualized to serve as a display (see pg. 15) . Displacement Nodding This is a downward movement of the head varying from a deliberate nod to an obvious lowering of the head with the bill touching the perch. The nod may be held momentarily as the bird wipes the bill, pecks at the feet, or touches the lower breast feathers. Sometimes the bird will deliver several Pecks at the perch. It is quite possible that nodding is not a displacement activity but has some appeasement value by concealing the bill from direct view of the opponent (it is the opposite of the Oblique Bill-up). However, the nod is seldom directed toward the opponent and often occurs before or after agonistic encounters, when the opponent is no longer present. Displacement Yawning Dilger (1956a) mentioned yawning as a possible displacement activity in Catharus juscescens. In the bluebird, yawning was not observed to occur in conflict situations. Displacement Bathing Bathing, as a possible displacement activity, was seen twice. On both occasions a male, engaged in a boundary dispute, interrupted fighting and bathed in a creek that flowed along his territorial boundary. The following extract from my field notes will serve as characteristic examples of displacement activities during conflict situations. Dis¬ placement activities are underlined. [46] June 30, 1967, Wing Hollow ; 11 ;00 a.m. Territory of pair 3S ; Nest destroyed by predator. A live decoy (male GY) is placed on a fence post 30 yards from the nest box. Male 3S flies to telephone pole then nods, assumes Oblique Bill-up, pivots, preens nervously around neck, breast, and wing (feathers not ruffled, head is sleeked). Male now flies close to the cage (decoy inside), assumes Oblique Bill-up, keeps his posture and makes a circle on the fence post, nods from Oblique Bill-up and touches tarsus ; assumes Head Forward and flies closer to the cage. Feather- ruffles and shakes the plumage, turns away, assumes Oblique Bill-up, nods, nods (both from Oblique Bill-up), nervously preens back and breast . etc. 11:15 a.m. Decoy is removed. Pair 3S come to where cage was. Both are excited and male starts “ turring.” Now both start nodding with the bill touching the perch as they look all around. Male preens very erratically then looks all around, male gives Rapid Warble, Wing-lifting Display, nods from a fluff, and touches bill to post. 1 1 :26 a.m. Male comes back to post where decoy was and hovers, lands, nods. Feather-ruffles, looks around, nods over and over, leaves. 11:30 a.m. Male comes back to where the decoy was, looks around, bill-wipes, then leaves. Displacement feeding, pecking, and/or collecting nest tnaterial This is usually seen during agonistic encounters between pairs at a territorial boundary. When the birds meet, the encounter is often interrupted as one or both glide to the ground to feed, peck at grass, or make intention movements to do either one. Often intense Fighting is broken off as the birds leave the fence line, glide to the ground, look around, and return to the fence line. This can be repeated several times. Thomas (1946) mentioned this behavior in bluebirds and made the following statement . . the pecking at the ground and tossing of leaves was apparently substitute behavior for fighting at a boundary line.” My observations agree with this. May 16, 1967 ; Wing Hollow ; Boundary dispute between Pair 19N and ION. Male ION advances from bush to bush, he keeps going to the ground as if to feed or pick up nest material ; male 19N does the same except he keeps his back to ION. Now 19N goes to the ground and pecks (pulls?) at grass over and over then flies up to a tree. Male 19N now has a small piece of nest material in bill as he goes from tree to tree. Both birds keep [47] Fig. 15 Two captive Eastern Bluebirds showing Turning-away (ap¬ peasement behavior). The bird on right is more submissive. Fig. 16 The Fluffed posture of the Eastern Bluebird [48] going to ground, sometimes feeding, other times not. Now 19N makes a Supplanting Attack upon ION. APPEASEMENT BEHAVIOR These are displays “ designed ” to prevent attack by directly reduc¬ ing the actual and relative strength of an opponents attack tendency, without provoking escape by the opponent or any general reaction by neighbors and companions (Moynihan, 1955b). Smith (1966) stressed their importance as a major feature in loosening the phe¬ nomenon of individual distance during pair formation. In the bluebird two postures were observed that appeared to have an appeasement function. T urning-away Turning-away is usually seen during agonistic encounters between members of a pair. It may be observed during boundary disputes between males. It usually consists of fluffing the body feathers and then turning the head away from the opponent. Turning-away may, however, involve the whole body as the bird turns its back on the opponent. When engaged in Song Duels at territorial boundaries the males sometimes perch with their backs to one another as they sing. In the early stages of pair formation Turning-away is frequently seen when members of a pair are in close contact. This is especially evident during the advent of courtship feeding; if the male does not immediately leave after he feeds the female, either one or both birds will Turn-away as they perch together. This probably signifies a weak escape tendency, giving the bird a nonaggressive appearance, therefore quelling any agonistic encoun¬ ters. Fluked Posture The bird perches in a normal position, head retracted, and plumage fluffed. The eyes may be closed and the bill pointed slightly upward. This is similar to the resting posture but differs in context. The Fluffed posture shows an absence of aggressive tendencies and a lack of readiness to act and, in some instances, was observed to reduce the likelihood of attack by another bird. The fluffing of the plumage is associated with the escape tendency, can be considered a submissive display, and in some instances may serve an appeasement function. [49] This posture is assumed by subordinate birds, by females during pair formation, and, at times, by one or both members of a pair when close together. Often this served as appeasement but several instances were observed where the aggressor attacked even though the attacked bird remained in the Fluffed posture. It appears, therefore, that in the bluebird the Fluffed posture is not a ritualized appeasement display. It may also occur when the escape tendency is thwarted by an incompatible tendency (feeding, sexual) or by the inability to flee due to confinement. December 23, 1966 ; Indoor flight cage ; St. Bonaventure Uni¬ versity Aviary. Male Y is in a submissive Fluffed posture with the bill slightly raised. He looks at male GY (despot), who is 2 feet away, and closes his eyes. GY Faces Y, gives Rasp twice, then leaves. Now male DY lands on one side of Y and female DG on the other side (both dominant to Y). Y remains in Fluffed posture and looks toward DY then toward DG. Y re¬ mained in Fluffed posture for 6 minutes and was not attacked. January 16, 1967 ; Wakulla County, Florida. 3 :30 p.m. A male assumes an Oblique Bill-up then flies at and Supplants another male who moved only 2 feet and immediately assumed the Fluffed posture with bill slightly raised. The dominate male again as¬ sumes an Oblique Bill-up and Supplants the Fluffed male who again moved 2 feet and assumed the Fluffed posture. The male while in a Fluffed posture was Supplanted 20 times in a row by the dominate male. After each attack the Fluffed posture was assumed. These 20 attacks occurred within 3 minutes as the birds moved along a fence line. May 4, 1967 ; Wing Hollow. Pair formation between male and female 5N. 4:30 p.m. After being violently Chased by the male, the female goes to a hedgerow and assumes the Fluffed posture (female looks like a round ball with the head drawn in). Male gives Chatter then flies at the female, knocking her from the tree and Chasing her through a hedgerow. The male goes to box 5N : the female to a tree and assumes the Fluffed posture. Bluebirds also combine the Fluffed posture with elements of attack when showing defensive threat. The body is fluffed with the head retracted but the bird Faces and Gapes at the opponent (both elements of attack). December 20, 1966; Indoor flight cage; St. Bonaventure Aviary. Female R bathes and on leaving the water is Chased four times by female DR. Now male YW Chases her, hits her in midair and both DR and YW drive her to a corner of the cage where she assumes a Fluffed posture with head retracted but Faces and Gapes at DR who finally leaves. R stayed in this position for 3 minutes. [50] Dilger (1960) noted that redpolls (Acanthis fiammea) in captivity, when subjected to persistent attacks by dominate birds, and escape was impossible, showed defensive threat. He pointed out the biologi¬ cal advantages gained by defensive threat. The bird indicates fear but also a willingness to attack if further molested, and if this infor¬ mation is communicated to the aggressor, further attack may be avoided. REDIRECTION ACTIVITIES If two or more incompatible drives are strongly and simultaneously activated by the same stimulus (or complex of inseparable stimuli, such as those provided by the mere appearance of an animal or object), the conflict of drives may be resolved by the animal “ venting ” one of these drives upon some third animal or object (Bastock, Morris, and Moynihan, 1953). Here the motor patterns appropriate to one of the conflicting tend- cies are shown, but are directed toward an object other than that which elicited the conflict. This behavior was seen in captive and wild blue¬ birds during the reproductive season. Four pairs of bluebirds were housed in separate (8' x 6' x 8') cages, in the same room, so the pairs could see and hear one another. Redi¬ rection was observed when males in adjoining cages engaged in a Song Duel which intensified gradually until the male attacked his own mate instead of the other unavailable male. After such an attack the male would resume the Song Duel and again Warble would intensify to a certain threshold whereupon he attacked the female. One par¬ ticular instance is worth mentioning : April 13, 1967 ; St. Bonaventure Aviary. Pair YW and W. Male YW’s Song Duel with the male GY reaches a peak and he starts Wing-lifting and looking around for female W (who is inside the nest box). Finally the Song Duel reaches such a high intensity that male YW flies to the nest box and starts pecking rapidly at the hole. He returns to the wire and starts answering GY again. This reaches a peak and as female W comes out of the box the male attacks, driving her to the floor, Pecking her head over and over. In this situation it is difficult to imagine that the male’s escape tendency is also strongly activated and in conflict with the attack tend¬ ency. However, Bastock et al. (1953) mention that, although most redirection activities seem to have resulted from conflicting drives [51] within an animal, such activities may also occur when a single drive (tendency) is strongly activated and then thwarted. This appears to be the case in the preceding example. Male YW’s attack tendency was strongly activated (due to the Song Duel with GY) and then thwarted (due to captivity), hence the bird attacked its mate. Of the four pairs of bluebirds in captivity, two showed this type of redirection during early pair formation. Kortlandt (1940) men¬ tioned that when an intervening fence prevented two angry animals from attacking one another, one or both would often turn and attack other animals. In the wild, during spirited boundary disputes, females often showed possible redirection activities. This usually involved a female direct¬ ing her attack upon her mate. June 24, 1966; Wing Hollow; Boundary dispute between pair 21S and 20S. “. . . males are now close together on the fence line with their females behind. Now both females go to the ground Fighting. The males remain on the fence posts and wave both wings. The females stop Fighting and return to the fence line and then the males go to the ground Fighting. As the males return to the line there are a lot of Supplanting Attacks. Female 21 S keeps Supplanting her mate (male 21S) as they move down the fence line. Male 20S rufFes the feathers, then female 20S does the same, nods, false preens, then her male comes and lands next to her. Rapid Warble is given by both pairs. Female 21S again starts Supplanting her male driving him back up toward the other pair. Female 21 S keeps up these Supplanting Attacks on her mate. . .” This behavior was also observed when both members of a pair were attacking a decoy. As the female was pecking the decoy she would turn and attack her mate. Indeed some females became so aggressive toward the decoy, the mate would be attacked whenever he approached. It is difficult to label this female behavior as redirection because the attacked male was previously showing threat behavior and may have provoked the female attack. This would also hold true for the female but a male rarely attacks his mate during a boundary dispute or during an attack upon a decoy. It may be that this female display of dominance in conflict situations may strengthen the pair bond. FLOCK BEHAVIOR Bluebirds in the nest box form social bonds to one another and will, if separated, crawl together again. Heinroth (1924) points out that nestlings of some species (i.e., chats wheatears, thrushes) lose this [52] first bond to one another when they first leave the nest and seek sep¬ arate perchs. This is not the case with fledged bluebirds for the young remain together in a close family unit, sometimes loafing and preening side by side and even roosting together the first night out of the box. In captivity, young of the same brood would sometimes roost side by side up to 155 days of age. Low (1934) mentioned the persistence of family groups and reported several instances of trapping young of the same brood a month or two after leaving the nest. Sometimes the social bond between the parents and young remained through a second nesting with the juveniles helping to feed the second brood (Hamilton, 1943; Laskey, 1939; Wetherbee, 1933; pers. obs.). In this study juveniles from different family units joined together to form small mobile flocks in late summer. Agonistic behavior then became fairly common, usually consisting of Supplanting Attacks and Chases that occasionally developed into actual combat. Incipient reproductive behavior was also common with females pecking at and mandibulating nest material or males attempting copulation. Juve¬ niles of both sexes eagerly participated in nest inspection. Table 3 shows the size and composition of these small mobile flocks in the Wing Hollow area. TABLE 3 SIZE OF DATE FLOCK COMPOSITION 1966 8/29 4 adult male and female; two young (migrants) 5 female 7N ; female ION ; female 20S ; two young ION 9/12 6 ? 3 male 7N ; female 17S, female ? 5 ? 5 male, female 23 N ; male and two females banded but not identified 11 ? 9/13 5 female, young ION ; male, young ION ; other young ? BEHAVIOR maintenance actvity on utility line one young attempts to copulate with other young- feeding along hedgerow nest inspection feeding maintenance activity on utility line seen by S. W. Eaton on utility line nest inspection, much chasing [53] TABLE 3 (cont.) DATE SIZE OF FLOCK COMPOSITION BEHAVIOR 3 male young lOS ; fe¬ male young 2 1 S ; female young ION nest inspection 4 young 1 N ; other young ? nest inspection 10 ? on utility lines and left right away 5 male and female 40S ; male and two females (migrants) nest inspection 9/16 3 young 7N ; (one male, two females) nest inspection 6 p feeding on Primus vir- giniana 9/24 4 adults ; one female banded maintenance activity, then left 4 male 40S ; male and two females (migrants) nest inspection 9/25 5 two males ; three females nest inspection 9/28 2 male young ? ; female young ION nest inspection 4 male 17S ; female 40S ; male and female ? (mi¬ grants ) nest inspection 2 female 23N ; male young 7N nest inspection 10/8 4 male and female young ION ; male and female young lOS nest inspection 4 two male young 7N ; fe¬ male young 7N ; young ? nest inspection 10/9 2 male 17S ; female ? (mi¬ grant) feeding in upper pasture 1 male 12S feeding from utility line 4 two females, young ION ; two males, young ? feeding along fence line [54] The size of migrant flocks of bluebirds is quite variable. Bent (1949) reports some flocks with only six birds while other contained “ some hundreds ” of birds. In southwestern New York, migrant flocks of one to 30 have been seen (Eaton, pers. comm.; pers. obs.). Buck and Bates (1962) report four separate flocks of 50 birds each in the area just north of St. Bonaventure University. In the winter of 1967, wintering flocks were studied in the vicinity of Tallahassee, Florida. As many as 100 bluebirds (both sexes about equal) could be counted along a 3-mile stretch of highway. I inter¬ preted this to represent several subgroups acting somewhat indepen¬ dently of the main group. These subgroups contained two to 40 birds with the number continually changing as birds moved independently to feeding areas. Thomas (1946), studying a nonmigrant population, stated that the size of a winter flock was determined by the number of nest sites (boxes) in a locality and the bluebird population of the surrounding country. I did not find this to be the case in a migrant population. To see if wintering birds in the Tallahassee area would show any interest in nest boxes, two were placed on telephone poles where the flock normally fed. Although the boxes were present for 10 days and the birds were always feeding in close proximity to them, no bluebird ever showed an interest (i.e., nest inspection) in them or, in any natural nesting holes. The noteworthy behavior of these wintering flocks was the relative paucity of agonistic encounters. When feeding, bathing, preening, or loafing, bluebirds showed little aggressiveness toward conspecifics (even when in such close association as feeding upon sumac and mistletoe). Most agonistic encounters recorded involved direct attack on the opponent rather than attack when individual distance was violated. I had determined that a relatively strong social bond remained among fledgling nest mates so an investigation was made to determine what type of social structure existed in a group composed of birds from different broods. These observations were made on juvenile birds, hand-raised from 8 days of age, and may not be representative of wild bluebirds. Subgroup 1 consisted of two males (DY, Y) and two females, ( W, R) all from the same brood ; subgroup 2 contained two males (YW, GY) and two females (DG, DR) from the same brood except male YW. The two subgroups were housed in separate (3' x 3' x 6') cages until approximately 60 days old. On September 21, 1966, both sub- [55] groups were placed in a 10-cubic-foot flight cage. Recorded observa¬ tions were begun a week later and the social hierarchy was determined by examining conflicts involving dominance between caged individuals. A bird was considered the winner of an encounter if its opponent re¬ treated. Between October 1, 1966 and April 1, 1967, 1,480 agonistic encounters were recorded in approximately 76 hours of observations distributed over 82 days. Table 4 shows the social structure of each subgroup over a 6-month period. Both subgroups were organized in a peck-dominance hier¬ archy where the relationship between any two birds is determined by which bird wins the majority of encounters. In subgroup 1 the males dominated the females whereas in subgroup 2 the females dominated the males. TABLE 4 Social Hierarchies of Subgroups 1 and 2 for a 6-Month Period SUBGROUP 1 SUBGROUP 2 DY^ Y5 W$ R2 DG$ DR 5 GY^ YW5 DY^ 6 23 6 DG$ 22 31 26 YS 10 3 2 DR$ 6 16 19 W5 7 3 GY 5 8 14 72 R2 1 1 1 YW^ 11 3 1 October 1, 1966 to April 1, 1967 TABLE 5 Social Hierarchy of the Flock (Both Subgroups) for a 6-Month Period DG5 DR 2 GY^ YW5 DY5 Y^ W2 R2 DG2 22 31 26 14 17 167 31 pecks : 6 DR 5 6 16 19 9 9 52 80 5 GY S 8 14 72 271 42 85 37 5 YWS 11 3 1 121 19 47 33 4 DY$ 74 55 1 15 6 23 6 4 Ys 2 4 10 3 2 3 W2 7 3 1 R9 3 1 1 1 0 October 1, 1966 to April 1, 1967 [56] Read from left to right in the horizontal rows. The numbers indicate the number of wins by the bird in question. For example in the first hierachy, DY won six observed agonistic encounters with Y. Based on the number of encounters, members of subgroup 1 were not as aggressive toward one another as were the members of sub¬ group 2. Table 5 shows the social structure of the flock (now considering both subgroups as a single flock) over the same 6 months. The flock was generally organized in a peck-dominance hierarchy but the two subgroups never became integrated. Thomas (1946) reported that winter flocks rarely mingled in the area of their nest sites ; in bitter winter weather small flocks might join together but with return of mild weather the newcomers were driven off by the resident birds. Subgroup 2 completely dominated subgroup 1, especially during social activities such as nest inspection. Nest inspection was a social affair with all members of subgroup 2 participating. The birds derived much excitement from this activity and at times appeared playful even though agonistic encounters were frequent. Members of subgroup 1 were never allowed to participate and if they tried, were immediately Chased. The aviary contained five nest boxes and usually when sub¬ group 2 was participating in nest inspection at one box, a single mem¬ ber of subgroup 1 would inspect a different box. At times this would be tolerated, but nest inspection never became a social affair for sub¬ group 1. The following notes typify nest inspection. Members of subgroup 1 (subordinate) are underlined. November 5, 1966; St. Bonaventure Aviary. Members of sub¬ group 2 are at box 3 participating in nest inspection. Male DY looks in and is Supplanted by male YW. Male GY rocks, goes in, YW is on top. Female DG goes in with GY, now YW goes in. All three inside the box when GY comes out, lands on a post, YW follows and tries to copulate with GY but GY Gapes and YW leaves. February 2, 1967 ; St. Bonaventure Aviary. Members of sub¬ group 2 at box 5. YW inside box, DG, DR both at hole, DR and YW keep Pecking at each other. YW comes out and pushes past DR who then rocks, DG goes to top of box. Now all leave and go to box 4. DR rocks, goes in, DY lands on top but is Sup¬ planted by GY. Now YW rocks, goes in, DG comes, rocks, goes in, GY comes, looks in (other three inside). YW pushes past GY, W comes to box top and is Chased by GY. DR comes out, GY comes, looks in, rocks, DR Supplants GY at the hole, W goes to box 2, looks in, then leaves right away. R goes to box 5, looks in, leaves. DG still inside box 4, GY comes, rocks, DR Supplants GY at hole. DG comes out and now birds go and feed. [57] Subgroup 2 also expressed its dominance over subgroup 1 in other social activities, such as preroosting behavior. To typify this : November 1, 1%6; St. Bonaventure Aviary. DG, YW, GY are in the corner of the cage pushing and shoving under and over one another. DR comes in and lands on the back of YW and pushes in between the two birds. Now all either shove under or over one another as they change positions. W and DY now go to opposite corner of cage and start to do the same thing. YW comes and Supplants both of them. Members of subgroup 1 never roosted next to members of subgroup 2. Birds of subgroup 2 would occasionally roost side by side, but those of subgroup 1 roosted alone throughout the cage. Group dominance was not expressed toward objects inside the aviary except nest boxes. The members of dominant subgroup 2 de¬ fended no special perches, roosting sites, or feeding areas. Marler (1955) and Dilger (1960) studied the effects of starvation on the social order of chaffinches (Fringilla coelebs) and redpolls (Acanthis flammea). They found that when starved, less dominant birds became more “ reckless ” about violating the individual distance of more dominant birds. The same may be said for bluebirds although I did not design specific experiments to show the effects of starvation. Bluebirds are extremely fond of mealworms {Tenehrio sp.) and were fed them only once every 2 weeks. At this time (starved in the sense of mealworms) subgroup 1 would usually feed first, followed by subgroup 2. It was not unusual to have both subgroups feeding together, side by side, all uttering a soft, hardly audible “ pump handle-like ” call with little or no agonistic encounters. Although both subgroups were organized in a peck-dominance hierarchy, because they were not integrated, a complex peck order, resulted, maintained by both peck-dominance and peck-right. Mem¬ bers of the dominate subgroup 2 held peck-right relations over all or some of the birds in subgroup 1 ; subgroup 1 lacked peck-right rela¬ tionships. Table 5 shows this but the following arrangement gives a better indication of peck-right and peck-dominance relationships. Peck-Right GY $ *dominates YW5,DY5,Y^,W2,R5 DR 5 Y^ , W5, R2 DG2 W$, R5 YW5 W5, R5 [58] Peck-Dominance DG 2 dominates DR 2 , GY $ , YW ^ , Y ^ DY^ " DG2, DR2, W~2, R2 YW 5 " DY 5 , Y 5 DR 2 " GY g , YW ^ Y 6- " DY $ W 2 " R 2 ^ Underlined birds are members of dominate subgroup 2. Note that male GY held peck-right relationships with all members of the flock except his two nest mates, females DR and DG, who both dominated him. Male YW, although a member of subgroup 2 (but never a nest mate for he was from a different brood) was never com¬ pletely accepted into the group for male GY held a peck-right rela¬ tionship with him. Table 6 shows the flock’s social structure for each month of the investigation. Because both complete and partial dominance relation¬ ships were formed, the flock showed both triangles and dominance reversals. Two of the most notable triangles involved male DY. He was dominated by males GW and YW ; yet he dominated both females (DG and DR) that dominated these two males (see below). Early in the investigation females DG and DR both domiinated male DY but he reversed this dominance and by the end of the study completely dominated both females. No evidence indicated that this reversal was due to redirection (sensu, Bastock, Morris, and Moyni- han, 1953) by male DY because he was completely harassed by males GY and YW. [59] Figure 17 shows the aggressiveness of each individual over a 6- month period. The relative aggressiveness of each bird was measured by the number of encounters it won per hour of observation each month. Aggressiveness reached its peak during the winter months and declined with the advent of the breeding season (Figure 18). There was no shift (i.e. change in agonistic relations of the sexes as they came into reproductive condition) in dominance such as reported by Hinde (1955-56), Dilger (1960), and Coutlee (1967). This is natural for wild bluebird pairs have an incomplete dominance relation¬ ship. [60] Fig. 18 Seasonal variation in total aggressiveness for the captive flock of Bluebirds in 1966-67. HRS of OCT NOV DEC. 'AN. FEB. MAR. OBSRV 13 10 10 7 15 19 Fig. 19 Seasonal variation in the total aggressiveness of eight caged male and female Bluebirds in 1966-67. [61] Five members of the flock showed a cyclic aggressiveness phenom¬ enon while the three totally subordinate birds showed none. In general the position in the hierarchy corresponds to the relative aggres¬ siveness of each individual. Figure 19 shows that males were generally more aggressive than females. It can be concluded that the social structure in the juvenile captive bluebirds studied, consisted of both peck-right and peck-dominance relationships. Indeed, Marler (1955) in quoting Ritchey (1951), stated “ that it is possible to obtain an apparent “ peck-right ” relation¬ ship in a society that is actually organized in a “ peck-dominance.” In the bluebird this complex social structure resulted from subgroup I’s inability to integrate with subgroup 2 suggesting that members of a brood (in captivity) retain a social bond to one another and that complete dominance is attained only over members from different broods. TABLE 6 Social Hierarchy of the Flock (Both Subgroups) for Each Month of the Investigation Period October 1966 DG? GY^ DR 2 YW^ DY$ Y5 W2 R2 DG9 1 2 5 3 8 4 pecks : 6 GY^ 2 13 16 4 24 17 6 DR 2 2 5 2 7 9 5 YW5 6 2 5 11 2 DYS 2 6 2 2 4 2 Y$ 1 3 1 1 1 1 W2 2 2 1 R2 3 1 0 [62] TABLE 6 (cont.) November 1966 DG5 GY^ YWS DY^ DR 2 Y5 W2 R2 DG$ 1 5 6 1 1 6 3 pecks : 7 GY^ 6 4 1 5 6 6 5 YW^ 1 1 16 2 8 16 14 4 DY^ 1 1 2 1 7 3 5 1 4 DR 2 1 2 2 5 6 2 Y5 1 1 1 2 W2 1 1 1 R2 0 December 1966 GY^ YW5 DY^ DR 2 DG2 Y5 W2 R2 GY 5 16 21 3 4 14 21 6 pecks : 7 YW5 73 5 8 6 3 5 DY^ 3 5 1 3 1 5 DR 2 2 3 2 2 3 7 41 4 DG2 1 1 3 2 5 25 5 3 Y5 0 W2 1 0 R2 0 January 1967 DR 2 DG2 GY 5 YW SDYS Y^ R2 W2 DR 2 4 6 5 2 18 17 pecks : 6 DG2 2 6 5 4 14 13 5 GY^ 3 2 12 44 12 7 7 5 YW^ 11 1 3 6 4 DY^ 12 17 1 2 4 Y^ 9 1 2 R? 1 1 W2 0 [63] TABLE 6 (cont.) February 1967 DG2 GY^ DYs DR 2 YW5 R2 Y^ W2 DG2 10 5 6 5 4 32 pecks : 6 GY $ 2 101 6 9 1 7 9 6 DYs 29 19 3 4 4 DR 2 3 4 6 2 5 4 YW5 5 7 1 2 6 3 R2 1 1 Y5 1 0 W2 0 March 1967 (R and Y removed) DG2 DR 2 GY^ DY^ YW5 W2 DG2 12 12 7 83 pecks : 4 DR 2 4 3 11 3 GY 5 85 16 18 3 DY5 20 14 3 7 3 YW5 8 8 2 W2 3 0 [64] TERRITORY I follow Noble (1939) in defining territory as “ any defended area.” The territory of the Eastern bluebird can be classified as Hinde’s (1956) Type A. This is a large breeding area in which nesting, court¬ ship, mating, and most food-seeking occur. Powers (1966) found the territory of the mountain bluebird to be Type A. PHYSICAL ASPECTS OF THE TERRITORY The nest box or natural cavity is the most important element in the selection of the territory. After selecting a nest hole the male estab¬ lishes a rather large territory around it. Other thrushes (ex. Ameri¬ can robin; Young, 1951) occupy a territory first and then choose a suitable nesting place. Von Haartman (1956) stated that a suitable hole seems to be the key-stimulus for inducing hole-nesters to choose a territory and that the terrain in which the hole is situated is of slight importance. Again, the bluebird differs, for the terrain in which the hole is located is extremely critical. The territory must contain open fields or aban¬ doned orchards with many perches that can be used for singing, feeding, and display. Grazed pastures are preferred to planted fields. Streams and stands of sumac (Rhus sp.) are also important for they assure a food supply (e.g., adult stoneflies and sumac fruit) during early spring snow storms. Bluebirds cannot be induced to nest just by putting up nest boxes and disregarding the terrain. The number of nest boxes a territory contains has some influence on the maintenance of the territory and hence, maintenance of the pair bond. Once males have established territory, the attachment is stead¬ fast and abandonment is rare. In early pairing-behavior, the male’s attachment to the territory (nest box) remains stronger than his [65] Fig. 20 Territorial changes, Wing Hollow study-area, 1967. [66] bond to the female. If, for some reason, the female abandons the nest site early in the nesting cycle the male will remain and start advertising again. Some males have mated with as many as four females before a successful nesting was accomplished. Therefore the number of nest boxes during early prenesting is of little importance in, main¬ taining the pair bond. However, once the bond is more firmly estab¬ lished and the pair are midway through the nesting cycle (i.e., incubat¬ ing or feeding young) other nest boxes within the territory take on added importance in its maintenance. If this nesting is prematurely terminated, the pair usually will remain and renest if other nest boxes are available, maintaining the territory and hence the pair bond. Pair 17S (1966) had five empty boxes in their territory, lost their young on June 7, and maintained their territory for the next 46 days without renesting. Maintenance was carried out mainly by the male leading the female daily from box to box and going through the Nest Demon¬ stration Display while the female watched nearby. If the territory lacks other nest boxes the male will either abandon the territory or the female, for females rarely will renest in the same box. Although quantitative data are lacking, the male’s attachment to the female at this stage appears to be stronger than his attachment to the territory; hence the pair desert the territory and the area completely. Nest boxes are also instrumental in determining the size, shape, and boundary changes of territories. When first establishing a ter¬ ritory bluebirds do not just select and defend one “ box.” Depending on the population density they inspect all boxes in the vicinity, using these for Nest Demonstration Display. The general procedure is for early arrivals to frequent a much larger area than they will later defend. Once the female starts to build, the pair confines its activities to one box, but will still defend other boxes in its territory. As the population density increases, nesting bluebirds will surrender certain boxes in the marginal areas of their territory to new arrivals. This results in many boundary changes throughout the breeding season. Figure 20 shows some territorial changes that took place in Wing Hollow during 1967. Pair ION were the first arrivals and frequented a large area. By mid-April, males 20S, 14S, and IN had established territories along the margin of lON’s original area, thus reducing its size considerably. By mid-May, males IIS, 17N, and 3S had carved out territories in the center of the breeding population thus reducing the size, shape, and boundaries of lON’s, MS’s, and IN’s territory. [67] This figure illustrates several important factors about bluebird terri¬ tories : 1. Bluebirds frequent a much larger area than they will later defend. 2. Bluebird territories are compressible but their compressibility has definite limits. 3. There appears to be a minimum territorial size and any encroach¬ ment is strongly resisted. 4. Unlike most hole-nesters that defend only a nest hole (Von Haartman, 1957), bluebirds defend a relatively large area sur¬ rounding their nest cavity. Von Haartman (op. cit.) also stated that one is able to increase enormously the number of most hole-nesting birds by putting up nest boxes. He increased his population of pied flycatchers {Muscicapa hypoleiica) from 10 to 70 pairs in the same study area. Once again the bluebird differs. Figures 21, 22, and 23 show the increase in blue¬ bird population at Wing Hollow from 1965 to 1967. In 1965, 10 boxes were available with five pairs nesting, in 1966 the number of boxes was increased to 35 with nine pairs nesting, and in 1967, 43 boxes were available with 10 pairs nesting. The most apparent reason for the moderate increase in population was not the increased number of boxes, but rather the selective place¬ ment of boxes throughout the hollow, thereby covering all available habitat. The five territories maintained in 1965 remained approxi¬ mately the same in 1966 and 1967 with some having as many as four empty boxes. New territories were usually established in favorable terrain that lacked boxes in 1965. This same trend prevailed in the other tributary valleys studied. Bluebird populations, in a given area, cannot be prodigiously increased by just increasing the number of nest boxes. Bluebirds have certain behavioral mechanisms that prevent crowding; namely, the vehement defense of a relatively large area around the nest box rather than just the nest box itself. Once established, territorial boundaries not containing nest boxes remained fairly rigid. These were formed where there were satisfac¬ tory perches for carrying out territorial defense patterns. Early in the season, stretches of bad weather weaken territorial behavior and pairs tend to wander in search of food with hostile encounters when pairs meet. [68] [69] [70] [71] However, boundaries containing empty nest boxes remained rather fluid (sensu, Howard, 1920) with ownership going to the neighboring male or pair in the prenesting phase of the breeding cycle. To show this diagrammatically : When the pairs were nesting, they ignored this fluid area and main¬ tained a rigid territorial boundary as indicated by the solid lines. If, however, a male lost his mate or a pair had an unsuccessful nesting, they would extend their territory to include the fluid area around the empty nest box, both defending it and using it for display. This situation existed through most of the nesting season in 1967 among pairs ION, 17N, and IIS. The area around box 16N remained fluid, with ownership continually changing. Territories were mapped after becoming fairly stable as determined when the female started to carry nest material, concentrating most activities around one box. Field sketches were made of each pair’s activities and plotted on an aerial map with size determined by the square grid method (Stains, 1962). The average size was 15.1 acres in 1966 and 12.1 acres in 1967 (table 7). Figures 22 and 23 show the distribution of territories. In 1966 and 1967 the territories in the center of the study area became smaller and more compact while those at both ends remained approxi¬ mately the same. This was because the males spread out themselves in early April as seen in figure 20. As the season progressed, late arriv¬ ing, unpaired males established territory in the center of the popula- [72] tion rather than at its ends ; apparently attracted by the frenzy of con¬ tinual courtship display. This was especially noticeable in 1967 when males 3S, 17N, and IIS, all late, single male arrivals carved out territories in the middle of the population, rather than at its ends where plenty of empty nest boxes were available. In fact, the area between territories of pair 5N and 3N (see figure 7) contained five empty nest boxes and remained vacant all season. Late arriving pairs, however, never attempted to “ carve out ” ter¬ ritories in the center of the population. They always established territories at the ends or extreme margins of the population, slipping in unnoticed with little advertisement. TABLE 7 Approximate Size of Bluebird Territories, Wing Hollow, 1966 and 1967 Territory 1966 Size Territory 1967 Size Pair lOS 8.9 acres Pair 14S 5.1 acres Pair 23N 9.9 acres Pair IN 5.3 acres Pair 17S 12.8 acres Pair ns 5.5 acres Pair 12S 15.0 acres Pair 17N 6.2 acres Pair ION 15.7 acres Pair ION 8.4 acres Pair 21 S 15.9 acres Pair 3S 9.9 acres Pair 3S 18.7 acres Pair 5N 14.6 acres Pair 7N 19.4 acres Pair 21S 16.9 acres Pair 40S 19.8 acres Pair 3N 20.5 acres Pair 20S 28.7 acres Mean 15.1 acres Mean 12.1 acres ARRIVAL AND ESTABLISHMENT OF TERRITORY In the North American thrush genera Hylocichla, Cathariis and Tiirdns, the males of the species arrive on the breeding grounds be¬ fore the females and set up and defend territories against birds of their own species (Dilger, 1956a; Young, 1951). Despite observations to the contrary (Bent, 1949; Todd, 1940) I found that arrival of blue¬ birds in the Wing Hollow area differs from this thrush pattern (see table 8). Males do not always arrive first, followed by the females. The general pattern was a staggered arrival of male and female birds from mid-March to the beginning of June. As table 8 shows, some birds arrived already paired, especially in 1967 ; others unpaired. Cold [73] weather late in March held up the advancing birds farther south, resulting perhaps in pair formation south of their breeding territories. Most bluebirds that arrived in Wing Hollow stayed throughout the breeding season. The reappearance of both young and adults banded in previous years indicated a tendency to return to the same general area. Kibler (in lett.) however, informed me of a male bluebird he captured for 2 consecutive years in the Jamestown, New York area that was banded as an adult in the Ithaca, New York area some 200 miles east. Thomas (1946) asserted that at the start of the season unmated males do not take up territory ; in Wing Hollow this was not the case. After arrival, the male selects a suitable nest box and confines his activities to its immediate area. At first he is quite secretive, mostly foraging and silently inspecting the nest box. Warble is weak and seldom given although Location Calls are frequent. After a day or two he becomes more conspicuous and starts to advertise the territory in earnest. He becomes very aggressive, continually challenging neighboring pairs. At this time the male enlarges his territory almost daily, taking over as many nest boxes and suitable terrain as the TABLE 8 Arrival Dates and Pairing Status of Bluebirds, Wing Hollow, 1966-67 Bird Arrival Date Status When Previous First Seen History Male lOS March 19, 1966 Paired Banded as adult. Female lOS a March 19, 1966 Paired — left 1965 Female lOS b p Unpaired * Male 40S April 14, 1966 Unpaired Female lOS c ? Unpaired Male ION March 21, 1966 Paired ? Female ION March 21, 1966 Paired ? Male 12S March 22, 1966 Unpaired Female 12S March 23, 1966 Unpaired Banded as adult. Female 12S a June 14, 1966 Unpaired 1965 Male IN March 23, 1966 Unpaired Female IN March 30, 1966 Unpaired Male 3S March 23, 1966 Unpaired Female 3S April 14, 1966 Unpaired Female 3S a June 17, 1966 Unpaired Male 38S March 27, 1966 Unpaired — left [74] TABLE 8 (cont.) Male 7N Female 7N Female 7N a April 11, 1966 April 15, 1966 May 24, 1966 Unpaired Unpaired — left Unpaired Male 17S Female 17S April 26, 1966 April 29, 1966 Unpaired Unpaired Male 23N Female 23N May 6, 1966 May 6, 1966 Paired Paired Banded as adult, 1965 Male 2 IS Female 21S June 10, 1966 June 10, 1966 Paired Paired Male 3N Female 3N Female 3N a March 29, 1967 March 29, 1967 June 17, 1967 Paired Paired — left Unpaired Banded as adult, 1966 Male ION Female ION March 29, 1967 March 29, 1967 Paired Paired Banded as adult, 1966 Male IN Female IN April 2, 1967 April 2, 1967 Paired Paired Banded as adult, 1966 Male 22S Female 22 S April 2, 1967 April 2, 1967 Paired Paired Banded as young, 1966 Male 20S Female 20S April 8, 1967 April 8, 1967 Paired Paired Male 14S Female 14S April 9, 1967 April 27, 1967 Unpaired Unpaired Male 5N Female 5N April 28, 1967 May 4, 1967 Unpaired Unpaired Banded as adult, 1966 Male 19N Female 19N May 13, 1967 May 18, 1%7 Unpaired Unpaired Male 17N Female 17N May 20, 1967 May 21, 1967 Unpaired Unpaired Banded as adult, 1966 [75] TABLE 8 (cont.) Male ns Female IIS Female IIS a Female IIS b Male 3S Female 3S May 21, 1967 May 25, 1967 June 5, 1967 June 26, 1967 May 26, 1967 May 31, 1967 Unpaired Unpaired — left Unpaired — left Unpaired Unpaired Unpaired * Bond formed between two birds. neighboring birds permit. Once territory is established the male re¬ stricts all his activity to this area. However, in early spring during adverse weather conditions, there are hours and even days when some males cannot be located. Whether they leave the territory in search of a mate or food is not known. After the male has established the territory he begins to continually advertise his presence in it. Advertisement is carried out by a variety of displays that function both as epigamic and gamosematic displays. Birds that arrive already paired, select and establish the territory together. ADVERTISEMENT OF THE TERRITORY The displays will be treated here only as they pertain to self- advertisement by the male. Once the female has accepted the ter¬ ritory most of these displays still persist but take on a different func¬ tion. Therefore, the relationship between the male and female will be treated in detail under Pair Formation. SONG Warble does not always begin when the first migrants arrive. Weather conditions play an important role because cold spells inhibit song. The highest perches in the territory (tops of dead trees, utility poles and lines) seem to be preferred but Warble is also given in the box vicinity and along fence lines. I have observed bluebirds singing from any posture, even during extensive preening bouts. However, Warble usually is given from an upright posture with the plumage somewhat fluffed (especially the throat feathers). When singing the bird may pivot and then sing in the opposite direction. Two movements can be performed by the [76] singing bird. The tail can be spread with each burst of song or the wing may be lifted rapidly in a vertical plane. Both of these indicate a sexual tendency and consequently the presence of a female in the vicinity. Saunders (In Bent, 1949) described the advertising song of the bluebird as three to eight notes grouped in phrases of one to three notes, with short pauses between them. In the Wing Hollow area. Warble consisted of two different song types ; the “ tury, cherwee, cheye-ley ” and the “ turr, heur, lee, em.” The latter can be intro¬ duce with either a “ turr ” or “ pew ” note. Sometimes “ pew ” is repeated over and over. Both song types can be alternated. Each Warble lasts about 1 to 2 seconds then is repeated after a short pause. In a random sample of 11 wild, unpaired males, the average number of Warbles per minute was 17 (table 9). Warble in this context func¬ tions to attract unpaired females. TABLE 9 Rate of Song Delivery of 11 Wild Unpaired Bluebirds Rate of Warble per minute 15 16 17 18 19 20 Number of songs 4 2 3 8 6 3 Male bluebirds use other vocalizations to advertise their presence in the territory. The most notable is Rapid Warble, which sounds like Warble (“tury, cherwee, cheye-ley”) without pauses between the phrases. Chatter is also frequently given, usually during Rapid Flight. Unpaired males frequently give Whisper Song near the nest box, usually in the form of Rapid Warble, but Warble and “ turring ” are also common. Muted vocalizations appear to have the same form as regular songs but are delivered at much lower volume, scarcely audible. Whisper Song may be detected (at least to human observers) by the pulsating throat of the apparently silent bird. I do not know if Whisper Song in this context functions as advertisement. AERIAL DISPLAYS Semicircling The male flies out from his perch, in a Semicircle and returns farther down the fence or utility line. Circling is usually at the same height as the perch but may be performed by dipping almost to the ground as he flies down the utility line. Rarely is song associated [77] with Semicircling. This is not the Circling described by Dilger (1956a) in pair formation but is typical of bluebirds as they adver¬ tise their presence in the territory. Females were also observed to Semicircle. Fig. 24 June 11, 1967. Male 3S giving Warble and Rapid Flight song while advertising his territory after losing 5 3S [78] Gliding Flight In this display the tail can be spread and the wings are fully extended but not moved as the bird loses altitude. Glides are normally brief (3 to 40 yards) but long flights (over 100 yards) are not rare. In long Glides a few dips or banking movements to either side may be made. Rapid Warble can accompany this display. When landing the wings are held up and out for a brief second then folded to the sides. Gliding is usually used as the bird goes from perch to perch or when approaching the nest box. It is also employed in foraging. Rapid Flight Song This display, given by males, is a rapid, straight flight between two perches ; one often being the nest box. Normally the flight is well over 30 yards and is accompanied by either Rapid Warble or Chatter, both of which can be given in a continuous or interrupted series. Rarely are the two alternated during one flight. Although given throughout the courtship, it is especially evident during establishment of territory. After losing a female this is a frequent display as the male again advertises his territory (see Fig. 24) . Of all the thrush species studied by Dilger (1956a) apparently only Catharus minimus bicknelli has a well-developed flight song. Impeded Flight Male bluebirds have three impeded flight displays. Hovering accents the presence of the male and nest box in the territory. The bird Hovers in front of the box, body somewhat vertical, wings rapidly beating. The tail is spread and the bird remains stationary in mid¬ air. A flight toward the nest box may end in a Hover for several seconds before landing at the hole. During Butterfly Flight the wings are fully extended and the tail is spread. The wings move in large amplitude but slow frequency so that actual flight speed is considerably slower than normal. Usually the flight is straight, between 5 and 20 feet above the ground, but wide circles and zigzags are not rare. Both Warble and Rapid Warble can accompany this display. Butterfly Flight is the most common Impeded Flight Display and usually ends by landing at the nest box. In Lopsided Flight the bird flies slowly, tail spread and both wings out of synchrony as he appears to “ flop ” along. This seems to be a form of Butterfly Flight with a strong sexual tendency present for [79] the bird appears to Wing-lift as he flies, resulting in the unsymmetrical flight. Sometimes bluebirds combine different forms of aerial display giving a rather bizarre aerial performance appearing to function as advertise¬ ment. The following two extracts from my notes are characteristic observations involving “ bizarre ” aerial displays. June 9, 1967; Wing Hollow; Pair 20’S ; 11 ;00 A.M. Pair 17S’s nest was destroyed by a predator and male 17S has begun to sing again. Most of the morning he has been intermittently Nest Demonstrating at box 18N and singing from different perches m the territory. Male 20S gives an aerial display along the border between his and 17S’s territory. He flies up 300 feet in the air and while singing uses Butterfly Flight, Glides, then gives Lop¬ sided Flight as if Wing-lifting, all the time moving along the ter¬ ritory boundary. He then suddenly “ tumbles ” down through the branches of a large oak ( Q uerciis sp. ) tree and lands next to his female. May 31, 1967; Wing Hollow; Male 3S. Male 3S is not yet mated. A female was seen in the north pasture but has not yet entered 3S’s territory. Male 3S gives Rapid Flight Song to the north fence line. Wing-lifts, then leaves and Glides for 100 yards with tail spread (few dips in Glide), lands in tree, and Warbles. Male repeats this again but female still does not follow. Now male uses Butterfly Flight all the way up to the north fence line, turns in midair, and Glides back with many “ banks,” first one way then the other, then starts Rapid Flight Song, lands at box ION, and goes in. Female ( ?) comes to box right away. To show this diagrammatically : NONAERIAL DISPLAYS Wing-lifting Unpaired males rarely Wing-lift unless a female is in the vicinity. Once the male becomes aware of a female’s presence he immediately begins a Wing-lifting Display, a particularly sensitive “ barometer ” in this situation. The wing is lifted away and up from the body in a vertical plane. Depending on the intensity, the wing can remain folded or be unfolded as it is rapidly lifted above the back then back to the side. When highly motivated the display appears almost as a “blur”; low intensity Wing-lifting may be just a quiver with the wing hardly leaving the supporting feathers or a frequent jerk of the wing up to the level of the back. Wing-lifting can involve one or both wings. This display indicates an activated sexual tendency and also makes the male’s presence more conspicuous. Wing-lifting can be given in conjunction with other displays such as Warble, and Nest Demonstration. It persists throughout the breeding season, often acting as a greeting between members of a pair. Wing-waving This display is also given when a female is in the vicinity. Some¬ times Wing-lifting can develop into Wing- waving, especially if the male is highly motivated. Wing-waving may represent Wing-lifting in its highest intensity. Both wings are held out to the sides, fully extended and then moved up and down. The body is in an oblique to horizontal position, the tail is somewhat raised but not spread. Sometimes in its highest intensity the wings are completely unfolded and held above the back for several seconds. Nest Demonstration The nest box (hole) is the most important element in the bluebird territory and a ritualized Nest Demonstration Display is used to advertise its presence. Nest Demonstration is the most important advertising display here as in pair formation and courtship. The display will be discussed here only as it pertains to territory adver¬ tisement by unmated males. The relationship of the female to Nest Demonstration will be treated in detail on page 114 (pair formation). Several authors (Burroughs, 1871; Bent, 1949); Thomas, 1946; etc. ) liave commented on the “ male’s persistent attempts to persuade a female to accept a nest-hole.” Von Haartman (1957), anthropo- morphically expressed the display’s function in hole-nesting birds [81] quite well when he wrote: “ This behavior evidently functions as an advertisment : good-looking bachelor with own apartment wants a mate.” In the bluebird, the complete display has several elements which can be performed in varying sequences. During high-intensity display (i.e., a female is present in the vicinity) the male flies to the nest box using any one of the aerial displays already mentioned, although But¬ terfly Flight is the most common. He perches at the hole with tail spread and wings partly open, showing the blue color of the back conspicuously. Wing-lifting or Waving may be given, especially if a female is nearby. While at the hole he may look around and then either peer into the entrance or put his head in and out of the hole in quick succession. The male then starts to rock back and forth, putting his head and shoulders inside with every forward rock, some¬ times looking around with the backward rock. Finally the male enters the box, and as he inspects, may peck at the sides and floor. Before leaving the box, the male shows his face at the hole for several sec¬ onds often ducking back inside only to face-show again. Face-showing probably has some signal value and appears to be somewhat ritualized. It is most frequently given when a male is highly motivated to entice the female to approach the nest box. For example, within 10 min¬ utes male 17N flew to box 18N and went in and out 11 times, face-showing in the following sequence when inside the box : 4_3-3-9-5-3-^1-4-2-1-0. The male’s exact manner of leaving the nest box varies. He may hop from the entrance, turn in midair. Hover, then land at the hole, and repeat the display. Sometimes he flies to a distant or nearby perch and sings, or flies to the top of the box and sings before repeat¬ ing the display. Often he flies from the hole to another nest box in his territory and performs Nest Demonstration there. At lower intensities one or more elements of the display drop out and at its lowest intensity the male merely flies to the nest box, look in, and leaves. The male bluebird usually incorporates song and nest material into Nest Demonstration Display. Both Warble and Rapid Warble can be given during any phase of the display and “ turring ” is frequently heard. While at and inside the box. Warble and Rapid Warble can be given in a muted form. Sometimes the male stays inside the box for over 3 minutes continuously giving Whisper Warble as he alter¬ nates between remaining in the box and face-showing. When face¬ showing, the pulsating throat is clearly evident. It is difficult to assess the function of Whisper Warble from within the box; espe- [82] cially during advertisement when the female is not present in the ter¬ ritory. However, during pair formation and courtship, Whisper Warble is a frequent part of Nest Demonstration and probably has some adaptive value in courtship by minimizing the pair’s presence to other bluebirds and possible predators. It may be that the unpaired male performs the complete Nest Demonstration Display during adver¬ tisement with Whisper Warble taking on meaning only during reac¬ tions between the pair during courtship. The male may carry nest material in the bill during some or all phases of the display. The material is usually obtained from inside the box and the male holds a strand, often mandibulating it, while face-showing. He can, however, obtain the material outside and take it to the box, mandibulating it, throughout the whole display. The male may also advertise himself and the nest box by dancing on the box top. Dance Display can be continuous with Nest Dem¬ onstration or can occur by itself. It is usually given on the box top but may be given on the ground or on a fence post. The male assumes the Oblique-sleek Display with tail spread and wings slightly drooped. He then moves all over the box top dragging the spread tail. He hops in a circle, stops to look around, then hops again. While displaying he may sing or carry nest material in the bill. Wing-lifting can ac¬ company this display. TERRITORIAL DEFENSE Territorial studies (Nice, 1941 ; Tinbergen, 1939) usually conclude that the established bird is nearly invincible in his territory. Indeed, Thomas (1946) stated . . . “ I have found that bluebirds are invincible in their territories only in the course of a nesting, not after their young are fledged.” I found that bluebirds in the Wing Hollow area were not always invincible in their territory. Paired males (or females) will some¬ times invade and usurp territory holders. The history of one follows : Wing Hollow, 1966 ; Pair lOS March 19 — Male lOS, a return from the 1965 season, arrives already paired with female lOS. March 25 — Female lOS has left. Male lOS showing weak ter¬ ritorial advertisement. April ? — Female lOSa arrives and forms a bond with male lOS. [83] [84] At hole with nestmaterial ; note dropped ^ Looking around wings and spread tail Fig. 25 Nest Demonstration display of male Eastern Bluebird showing several different elements. [85] Peers into entrance d Begins rocking [86] Rocking, continued f Pace showing, with nest material April 12 — Female lOSa is now starting to carry nest material to box 40S. Male lOS singing from different perches within the territory. April 13 — Female lOSa building in earnest now. Male lOS stays close to the nest box as female builds. April 14 — 7:30 a.m. Female carrying nest material to box. Female leaves box and male lOS Chases her across field. 7 :45 a.m. A new male (unbanded and unpaired) comes to the nest box, Warbles, goes in box, comes out, and Warbles from top. Male lOS comes immediately and a violent fight takes place in a ditch in front of the box. Female lOSa comes and Hovers over Fighting males then leaves giving Location Calls. 8 :01 a.m. Female lOSa comes back to the nest box. The males stop Fighting and one Chases the other across the field with the female following. 8:05 a.m. New male and female lOSa going through Nest Dem¬ onstration at box 41 S. April 15 — New male has taken over territory of male lOS and has formed a bond with female lOSa. Female again carrying nest material. April 20 — Male lOS has established a new territory in the south¬ ern part of the study area. Pettingill (1936) reported the case of an unpaired female bluebird invading a pair’s territory and successfully driving out the resident female. However, bluebirds were usually successful in defending their ter¬ ritories. Most disputes were settled by display and when actual combat occurred it was usually over a female rather than territory (nest box). The agonistic behavioral patterns used by bluebirds in territorial defense have been described under Agonistic Behavior. These depend on the male’s position in reference to his territory, his mate, and the behavior of the other bird. At territorial boundaries those displays showing conflicting attack and escape tendencies were prevalent, result¬ ing sometimes, in prolonged encounters. In the vicinity of the nest box, the attack tendency was strong and little display was seen as the owner simply attacked the intruder. Of all the agonistic displays. Warble is the most important in main¬ taining the territory. It, like that of most song birds (Marler, 1956), has the dual function of attracting females and repelling males. Loud Warble is usually the only display given during intraspecific encoun¬ ters in the vicinity of the nest box and probably reflects a high attack tendency as shown by the following observations. When decoys are placed near the nest box Warble is given immedi¬ ately as the resident male (or pair) attacks the decoy. Males will even attack a microphone emitting Warble in the absence of a decoy. [87] In captivity, even though visual contact was reduced to a minimum by attaching either 4' x 8' sheets of plywood or large pieces of cloth to the wires separating the cages, my bluebirds were never successful in fledging young. I believe the reason for the nesting failure was that the birds were in constant auditory contact with one another. Regard¬ less of the phase of the nesting cycle, on hearing Warble, a pair would try with great persistence to attack other pairs through small cracks in the cages. This caused a constant turmoil among the caged pairs with incubation and feeding of young ignored or suppressed by the stimulation of the stronger attack tendency. Mrs. Laskey (personal communication) says that in her experience bluebirds would only nest close together if a natural barrier (such as a hedgerow or barn) separated the nest boxes thereby reducing both visual and auditory contact between nesting pairs. Hartshorne (1962) successfully bred bluebirds in captivity; how¬ ever, his birds were kept in sound isolation chambers making auditory (and visual) contact between pairs impossible. Song Duels between neighboring males were a common display whenever territorial boundaries were in dispute. Normally this in¬ volved two males perched at the boundary, rendering Warble, but yet Song Duels often involved more than two birds. If one was unpaired and establishing territory the Duel would develop into a noisy “ melee,” reminiscent of the communal display of the house sparrow. Passer domesticus, especially if the females of the other participating birds were present. The unpaired male would Warble with the greatest volume and frequency and as the Duel intensified it would develop either into a communal Chase with two or three males Chasing a female or actual Fighting between males or pairs. Young (1951) never observed “ Song Duels ” between male robins {Tiirdiis migra- torius) but Siivonen ( 1939) describes a Song Duel between two song thrushes (Turdns ericetoriiim) . Power (1966) said of the mountain bluebird (Sialia currucoides) . . . “ two males would frequently be seen answering each other’s song along the common borders of their territories.” When females are present during boundary disputes the males guard them by staying between the female and the other male. If the female flies to the ground the male will follow and Hover over her, returning with her to the perch. Boundary disputes are frequently interrupted by Nest Demonstration. After vigorous encounters between the pairs, one or both will interrupt Fighting, fly back to their nest box, and give Nest Demonstration Display. This was also common during experiments with decoys ; the pair would attack the decoy, leave [88] and give Nest Demonstration Display, only to return and attack the decoy again. The nest box does not have to be the one used for nesting; any box in the territory can be used. Von Haartman (1956) mentioned this same type of behavior in the pied flycatcher {Muscicapa hypolenca) and interpreted it as ambivalent behavior, the male react¬ ing momentarily as if the intruder were a female. Hinde (1952) also mentioned that in the great tit (Pams major), hole inspection sometimes occurred during a lull in reproductive fighting. In the bluebird a possible explanation may be that the presence of the intruder (or decoy) was sufficient to stimulate an attack response. After pro¬ longed Fighting (or a lull in Fighting) the attack tendency waned and the presence of both the female and nest box released the next type of behavior for which both internal and external stimulation had reached threshold value, in this case, Nest Demonstration. Territorial defense does not decline when the female starts to incu¬ bate but remains intense throughout the nesting cycle (see table 10). This is important in a species that has special territorial requirements (nest cavity) in short supply. Usurpation is rare and hence the pair are able to raise more than one brood without wasting energy and time trying to locate a new nest cavity. Bluebirds usually tolerate other birds in their territory except in the immediate vicinity of the nest box. They show a marked antipathy, however, toward other hole-nesters, especially tree swallows (Irido- procne bicolor), house sparrows (Passer domesticiis) , and starlings TABLE 10 Results of Decoy Experiments Stage Reaction of Male or Pair ABC Male unpaired . 3* 3 Prenest building . . . . 2 5 Nest building . . 1 1 8 Laying . . ..... 5 1 2 Incubating . . 9 4 15 Feeding young . ..... 2 1 17 Young have fledged . . . . 8 — 9 Totals . . . * Number of experiments . 25 12 59 A = No reaction, birds ignore decoy B = Moderate reactions, birds show threat toward decoy C = Strong reaction, birds attack decoy [89] (Sturnus vulgaris). Sometimes bluebirds use a defensive display when defending the nest box from tree swallows. The bird will enter the box and face-show, while Gaping and Bill Snapping at the diving tree swallows. Although brown-headed cowbirds {Molothrus ater) have been reported to parasitize bluebird nest boxes (Friedman, 1929; Thomas, 1946), this never occurred in the Wing Hollow area. Cowbirds were Supplanted and Chased whenever they came near a nest box. Hamil¬ ton (1943) reported a case where two cowbird eggs were deposited in a bluebird’s nest and the female promptly covered both with a new lining. Bluebirds completely ignored the sparrow hawks {Falco spar- veriiis) nesting in area. Drinkwater (1953) reported a female spar¬ row hawk taking young bluebirds from a nest box but he felt this was because the box had a perch in front of the hole. Bluebirds were seen to Chase the following species from the vicinity of the nest box; robin (Tnrdus migratorius) , eastern phoebe (Sayor- nis phoebe), song sparrow {Melospiza melodia), and field sparrow (Spizella piisilla) . FUNCTIONS OF TERRITORY IN THE BLUEBIRD Tinbergen (1957) stated that territory is the result of two tenden¬ cies in the owner : 1. Attachment to a site. 2. Hostility towards a certain category of other animals, usually members of the same species and the same sex. The territorial behavior of the bluebird clearly exhibits both these tendencies. Because of its extreme hostility to conspecifics and its tenacious attachment to a nesting site (box) the following functions of territory are suggested. The male’s vehement defense of the nest site has selective advantages for the nest site plays an important role in pair formation. Bluebird nest sites are scarce ; therefore it is advantageous for females to pair with males who already possess one. The male’s advertising within his territory makes him more con¬ spicuous to roaming females, thus helping to bring the sexes together and form sexual bonds. The pair’s aggressiveness toward conspecifics and other hole-nesters secures a nest site (box) for the pair and the attachment to this area helps maintain the pair bond. Strong terri¬ torial defense also reduces the likelihood of interference with coition. [90] Males from adjacent territories are attracted to the intense courtship activities of the pair and sometimes are successful in interfering with copulation. By defending the territory per se instead of just the nest cavity the population density can be somewhat limited thereby securing for the pair a certain food supply and allowing them the opportunity to raise more than one brood. [91] WW ttAiHht^iil^m i^nt$mmm i5/S»y W^0^'tB^ •hi- ,)iai^Mfa4^tig- «vl «,[} STi.r{tln«; sUt ^hoifc iiiii bJBd^Sf^ »f(i ^jmUioteb iw-y^.ihik nfA’frpimr94 )■. % Wltfe ? fwlbo^«fiia SjiffijM!t«r iAi pfj(»Hp4 wbwtftvwtliojr V#i»iuiw^-fiiw Hiw >itt{i.(i94iy » tm «t^jioi»b«il ] fM»i $tvi ii\.. lo ttip tlurii'ciiiifjr-fj <4.ik iMflW i iwicf »t*fr*»wf{yik,9pki Ii^k>4k)4 ^firfila pmiUa), ^'l|(H|MMrt||' MM9H ^glwHMW^iKpSwi ‘-'^ -si!'^ • .f ™ j FUNCnONS QP rnfcRITQI^y ftlUCeiRD' *'f - Tt>c |4n^0rjAt ti»3l,H,Mtt,f ttf Itp bM ' jK^kSi^nfikfi/ UtifMAe »4 l4# ^ ' , . fttLvKjwPijii bJ. t iit* 1 '■ ' ■» * ’ , JX^vi fMV«.,vvhtu*>rftt td thf ftfjtftJb ^ j • W *lt» )tbyr| tm roi# {ttliilf ftn Wkmv . ^ I . ; .^m rJtffixQ^K it «o’ ynU wj#'* »4trfU4^ wbHM i4)4«irni^ ^twunhiir tlw pi4y« ^ onu^lxroetu M #«ii\|i«l^ i4i t-ftftd ‘'>5^1^ ^rrt f/wial dtttfnAtf- ftlMk Mim: jtm ■■'■* J.miV PAIR FORMATION Lack (1940b) in his review of the literature stated that in many species, pair formation occurs in the territory, while in others, it occurs in the flocks. Pair formation in the bluebird apparently occurs in both situations for some birds returned to Wing Hollow already paired. However, it will only be discussed as it occurred within the territory of unpaired males in the Wing Hollow area. In all, 10 natural pair formations were observed and these were supplemented by introducing live caged females into the territory of four unpaired males. THE MECHANISM A typical sequence when an advertising male detects a female in his territory follows. On seeing a female in his territory, an advertising male immediately begins Wing-lifting Display and Rapid Warbling. He may then fly toward either the female or the nest box, using Impeded Flight and uttering Rapid Warble. He ends up at the nest box and assumes the Oblique Sleek Display (see pg. 101) with continual Rapid Warble or performs Nest Demonstration Display. He lands at the hole with tail spread and wings partly spread, starts Wing-lifting and Waving, rocks, then enters. While inside he continues to Warble, face-shows several times with nest material, comes out to the top or nearby perch, and mandibulates the nest material as he Wing-lifts and Rapid Warbles. If the female does not approach the nest, the male may : 1. Repeat Nest Demonstration Display, 2. Perch on or by the nest box (often in Oblique Sleek), and give a variety of vocalizations such as Warble, Rapid Warble, or “ turring ” as he continually Wing-lifts, [93] 3. Fly with Impeded Flight and Rapid Warble to another nest box in the territory and give Nest Demonstration Display, 4. Fly toward the female and repeat the aerial display back to the box, 5. Chase the female. , The continued aerial and Nest Demonstration performances with inter- ] mittent Chases accentuate the presence of the male and the nest box \ and stimulate the female to approach the nest hole. Depending on the female’s motivation this may take minutes or several hours, but the ! female eventually approaches, either by following the male or coming while he is displaying at the box. Whatever the approach, the male immediately goes inside the box and begins Rapid Warbling (usually in muted form) and face-showing with nest material. The female may remain passively perched by the box in a Fluffed posture and appear to ignore the displaying male. If she does fly to the hole she is very tense as witnessed by her withdrawn posture with her body held way back and turned slightly sideways to the hole, the head sleeked and drawn in between the shoulders. One foot holds to the hole, the other near the base of the box and she keeps looking away as if she will flee at any moment. This is a critical moment for the female must enter the nest box while the male is inside to insure the establishment of the pairing bond. However, she seldom enters on her first approach to the hole. Usually she flies rapidly from the box but does not leave the territory. She remains perched (frequently in thick foliage such as a hedgerow or tree) in the Fluffed posture and may perform maintenance activities. The male continues his aerial and Nest Demonstration Displays and the female occasionally warbles and moves closer to the nest box. At this time the male commonly shows Wings-raised Sleek Display which develops into either Nest Demonstration or a violent Chase of the female ending with the male using Butterfly Flight and Rapid Warble back to the box; the female remains Fluffed on some perch within the territory. If, in the interim, the male is not present at one of the boxes, the female may suddenly fly to the box, go inside, remain a few seconds, then leave. Eventually the female approaches the nest again and remains Fluffed by the box as the male is inside giving Rapid Warble and face-showing. She may give a weak Wing-lift, Rapid Warble, fly to the hole and look in, engage in Mutual Warbling with the male, then return to her perch and again assume the Fluffed posture. When the male comes out he goes to the opposite side of the box and shows the Oblique Sleek Display (see pg. 101) with his back to the female. The female may then go to the hole and look inside. This causes the male to become greatly excited and while still in the Oblique Sleek he begins [94] to quiver and lift the wings as he shows intention movements to both leave and approach the box while uttering Rapid Warble and Squeals. This interplay between the two continues as the female begins to look in the nest box more often and starts to show signs of sexual excitement herself. Her tendency to flee lessens and the male’s Wings-raised Sleek Display now develops into actual attack as he lands on her back and Pecks her head. The crucial time comes when the female finally enters the nest box while the male is inside. After she enters, the male always comes out first and lands near the box usually in an Oblique Sleek with his back to the nest. The female follows and directly Supplants the male. Now the female takes an active part in Nest Demonstration as she follows the male to the different boxes in his territory. She shows reduced fear of the male and increases her Supplanting Attacks upon him. As the pair continue Nest Demonstration Display, one gets the impression (as did Meyerriecks in his study of the green heron, Butorides virescens, 1960) that the male is now the intruder ! The female can Supplant the male in any of the interactions between them while in the immediate vicinity of the nest box but it usually occurs as just described. I feel this act of dominance by the female is of prime importance in establishing the pair bond. Some evidence exists that the female dominates the male as soon as she enters the nest. When observing pair formation in captive birds the female would start Pecking the male as soon as she entered the box to the extent that the male would sometimes get confused and try desperately to leave through the side observation glass in the box. To show the importance of the Supplanting Attack by the female, the following section from my field notes is given ; May 4, 1967, Wing Hollow. Pair formation between male 5N and migrant female. Female has not yet come to box. 3:15 p.m. Male gives Butterfly Flight with Rapid Warble to box 9N, looks in, goes in, face-shows with nest material, comes out, turns in midair, lands at the hole with tail spread, looks in, goes to top with nest material. Wing-lifts, Hovers, looks in, goes in, comes out with nest material to the top and moves all over touching nest material to the top. Hovers, looks in, goes in, comes out with nest material. Wing-lifts . . . keeps up the same pattern and goes in the box at least 12 times. Now a female flies across the road and the male goes very fast, cuts her off, and he lands at box 8N, female goes to utility line. 3 :50 p.m. Female still has not come to the box. Female on utility line and male assumes Wings-raised Sleek, Chases female. [95] turns, and uses Butterfly Flight with Rapid Warble to box 8N. Males goes in .... 4 :35 p.m. Female nervously preens in hedgerow, male gives Rapid Warble, flies right at the female and Chases her among the trees, then goes to box 8N and Nest Demonstrates. Female remains in tree in a Fluffed posture .... 5 :40 p.m. Male inside box, female Fluffed nearby and Wing- lifts a few times, goes to hole with leg down and head back, looks in, back to wire. Male face-shows with nest material, comes out. Hovers, lands at hole, back in, face-shows with nest material (female looks towards box) three times, out. Hover, right back in. Female still in Fluffed posture. Male face-shows four more times, goes to wire on opposite side of box, and assumes Oblique Sleek. Female goes to hole, looks in, male Wing-lifts and female leaves the hole and Supplants the male. Male leaves and female looks in, goes to wire. Male Butterflies to the hole, goes in, much Whisper Warble heard. Female looks in, goes in (both in), male face-shows and out immediately with nest material and as¬ sumes Oblique Sleek on the wire. Female comes out and Sup¬ plants the male, male goes back in, female looks in, goes to wire. Male out with nest material and in Oblique Sleek, female in Fluff and Wing-lifts, male back in, out to opposite side of the box. Female goes and Supplants male, male doesn’t leave so female Gapes and bites him .... 5 ;50 p.m. Female comes toward box, male assumes Oblique Sleek and goes in, face-shows, female on wire, male out to the opposite side and female goes and Supplants him. Female now looks in. Wing-lifts, male comes to post and female Supplants him then keeps Supplanting him in rapid succession .... This partial dominance at the nest box is extremely important. The box is the most important element in the male’s territory and his de¬ fense of it is most vehement. Thus if the female is going to be ac¬ cepted, she must show some form of dominance in its vicinity to reduce the likelihood of attack by the male. Therefore the extent of the male’s aggressiveness partly depends on the degree of dominance exerted by the female at the nest box. DOMINANCE RELATIONS From the preceding discussion one would expect that a reversal of dominance in the sexes occurs during pair formation and that the female establishes a microterritory around the nest box. Yet this is not the case, for when the dominance relations between the pair [96] are considered for the entire courtship it becomes evident that neither sex completely dominates the other; a partial or sometimes incom¬ plete dominance exists. This is shown in table 11 where, of 255 recorded encounters, the males were dominant only 54 percent of the time. TABLE 11 Agonistic Encounters in Wild Pairs of Sialia sialis Wing Hollow, 1966-67 Pair Number of Encounters Male Wins Eemale Wins 1966 3S 16 14 2 7N 12 7 5 ION 2 2 0 lOS 1 1 0 12S 15 10 5 17S 18 7 11 23 N 1 1 0 40S 25 14 11 1967 3N 10 1 9 3S 11 9 2 5N 21 6 15 IN 15 5 10 ION 21 8 13 IIS 37 33 4 14S 3 1 2 19N 2 2 0 20S 18 9 9 22S 4 0 4 17N 23 6 17 Totals 255 136 119 Note the variation in the dominance relations between different pairs. In some relationships the males were extremely aggressive (35S, US), in others, the females (5N, 17N) were. Those birds that arrived already paired showed little aggression. Table 12 shows the types of agonistic displays used in the inter¬ actions between the pairs during courtship. [97] TABLE 12 Types of Agonistic Displays Used in Interactions Between Pairs Male Display Stage Number of Encounters Chase A 43 B 19 C 14 Attack A 9 B 5 C 34 Supplanting A 4 B 4 C 2 Diving at A _ B — C 2 Number of Female Display Stage Encounters Supplanting A 18 Attack ; B 48 at nest box C 12 Supplanting A 2 Attack; away B 3 from nest box C 2 Supplanting A 8 Attack; during B 5 boundary dispute C 2 Face-Gape-Peck A 3 B 8 C 8 A = Pair formation C = Nest building B = Prenest building Males mainly intimate females by Chasing but as courtship con¬ tinues the female’s tendency to flee becomes inhibited and actual attacks become predominant. Attacks were always expressed by the male Pecking the female’s head and occurred mainly during nest building. The female reveals her dominance by Supplanting Attacks which usually (65 percent) occur at the nest box. These are most frequent [98] during pair formation and prenest building then are reduced during nest building. Therefore the female does not establish a microterritory around the nest box that excludes the male. Throughout courtship the male is ever present at the box and he rarely shows fear when approaching it. SEXUAL RECOGNITION Bluebirds are sexually dichromatic and when a territorial male meets a male intruder he reacts aggressively, but when a female intrudes his first reaction is overtly sexual. Instead of Warbling, flying directly at the female, and delivering a series of Supplanting Attacks (as he would to another male), he immediately starts Wing¬ lifting, shows Impeded Flight with Rapid Warbling to the nest box where he gives Nest Demonstration Display. This indicates the male is able to recognize the female as distinct from the male and suggests that morphological rather than behavioral characteristics are used in sexual recognition. This is also supported by the fact that in four experiments in which a caged live female was introduced into the ter¬ ritory of an unpaired male, the male’s first reaction was always sexual (as above) rather than aggressive. This was also true when mounted female decoys were used. For example : May 27, 1966 — Wing Hollow. Male 7N is unpaired. I put a mounted male and female bluebird on a fence line 3 feet from male 7N’s nest box. Warble is played through the tape recorder. .... Male 7N Warbles and flies close to the decoys. Male Warbles then goes to box and gives Rapid Warble, goes in box, face-shows, back in, stays inside 3 minutes and gives Whisper Rapid Warble, face-shows, looks toward decoys, comes out, lands by box, and assumes Oblique Sleek Display. Now I play tape recording. .... Male flies up and over the male decoy and lands on other side. He assumes Head Forward, Oblique Bill-up, then attacks the male decoy violently and knocks it to the ground. Male regards the female decoy, gives Whisper Rapid Warble, goes to the ground, and pulls feather ofl the male decoy. I remove male decoy. .... Male moves closer to female then goes to ground where male decoy was and pecks ground. Male back by female decoy, regards it. Warbles, goes to box, looks in, goes in, out to fence line and Warbles, preens nervously. Wing-lifts. Male now Warbles from Fluffed posture then starts to preen. [99] I play recorded Warble and male moves away and gives Whisper Warble then feather-ruffles, goes to ground, back to fence line. Now male goes to ground where male decoy was, looks around, then back to fence line. Thus pair formation in bluebirds is quite different from the Hylo- cichla and Catharus thrushes studied by Dilger (1956a). He found males initially react aggressively toward females entering their ter¬ ritory, much as they would to intruding males. Thus in these thrushes there are no special displays associated with pair formation like those in the bluebird. MOTIVATION OF THE SEXES When the male first detects the presence of the female, the sexual tendency is clearly predominant. He shows Wing-lifting, Rapid Warbling, Impeded Flight, and Nest Demonstration; all sexual dis¬ plays. The attack tendency is not shown at first, but in later encoun¬ ters becomes clearly evident by the appearance of the Wings-raised Sleek Display, Sexual Chasing, and attacks upon the female. The escape tendency, though weakly activated, is betrayed by the Oblique Sleek Display given when the female is at the box. The female at first shows no aggressive tendency and usually remains in the Fluffed posture. With increasing visits to the nest box her sexual tendency rises as she shows Wing-lifting, Rapid Warbling, and an interest in the nest box. Finally, she takes an active part in Nest Demonstration and becomes increasingly aggressive as shown by her frequent Supplanting Attacks upon the male. Since pair formation seems to increase the relative strength of the attack tendency in both sexes, a principal function of bluebird court¬ ship is to suppress this attack tendency in the sexual partners. This is achieved by much mutual sexual displaying between the pair during the courtship period with Nest Demonstration and courtship feeding being especially important. [100] COURTSHIP I follow Morris (1956b) in defining courtship as “the hetero¬ sexual reproductive communication system leading up to the con- summatory sexual act.” This includes the activities from pair forma¬ tion through copulation. It has now been established that in both sexes there is a three-point conflict between the incompatible tenden¬ cies to flee, attack, and mate (Hinde, 1952, 1953; Marler, 1956; Morris, 1954, 1956b; Tinbergen, 1952). Courtship, therefore, does not involve a single sexual activity, but also the tendencies to attack and to flee from the sexual partner. In the sexual behavior of the bluebird all three tendencies are expressed but the bird is principally under the influence of conflicting tendencies to attack and behave sexually towards the mate. MALE DISPLAYS Oblique Sleek Display This is a variable display with many possible combinations of com¬ ponents. It is similar to the Oblique Bill-up Threat Display and may be derived from it. In its lowest intensity the body is held in an oblique position with the plumage sleeked. This is especially notice¬ able when the male is advertising his territory. However, most pair formation and early nesting behavior involve the full (more intense) Oblique Sleek Display, usually in the vicinity of the nest box. As the female approaches the box (or comes out of it) the male turns his back to her and assumes the Oblique Sleek with wings drooped, bill slightly raised, and tail maximally spread. The male thus presents a “ mass of blue ” to the female as she either Supplants him or he moves away. As courtship continues and the motivation levels of the different tendencies shift, the male incorporates more components [101] into the display. He may remain stationary (and tense) and lift or quiver the wings instead of drooping them. Vocalizations indi¬ cating a sexual tendency can be given ; Rapid Warble, “ turring,” and Squeal. Orientation may change as the male no longer turns away from but either faces or glances over his shoulder at the female. Some even pivot or turn in a complete circle showing alternating movements to approach and flee from the female. A number of facts suggest this display is due primarily to con¬ flicting escape and sexual tendencies. The oblique posture, sleeking of the feathers, and flexing of the legs are all flight intention move¬ ments and coupled with the male turning his back to and showing movement away from the female suggest a strongly activated escape tendency. Turning the back is a “ reversed movement ” (e.g., Marler, 1956), that is, the opposite in form from the posture of a bird which is ready to attack. Sometimes, when displaying, the male will even fly before the female makes the Supplanting Attack. The spread tail, movements of the wings, vocalizations, and the occasional attempts at rape all represent a sexual tendency. Also the male will sometimes mandibulate nest material while displaying. Yet the attack tendency is also weakly activated as evidenced by the sometimes open bill (Gape), change in orientation, and rare attacks on the female. Vocali¬ zations indicating an attack tendency such as Warble and Chatter can be given. In captivity where escape was impossible this display sometimes developed into a forced rape of the female. For example : May 14, 1967. St. Bonaventure Aviary. Pair YW and Y. Female is nest building. Male is in the corner of the cage in a frozen Oblique Sleek. He keeps “ turring ” for 6 minutes then makes a violent rape of the female. For the next 45 minutes he assumes the Oblique Sleek Display (full intensity), keeps his back to the female most of the time, and repeatedly gives Rapid Warble as he flies around the cage rapidly as if trying to get out. Within this period though he attacked and raped the female seven times, each time trying desperately to copulate. Isenberg (1953) bred the loochoo robin, Icoturns komadori koma- dori (Erithacus komadori komadori, Peters, vol. 10, pg. 39, 1964) in captivity and reported a similar display. He wrote “. . . he struts, not unlike a Turkey, with lowered wings, tail fanned out and held up over his back the while a torrent of lovely song pouring forth.” Sometimes during periods of intense courtship neighboring males will perform a very exaggerated form of the Oblique Sleek at the ter¬ ritorial boundary. The display appears to be directed toward the neighboring female and includes ambivalent movements, postures. [102] and displacement activities. The duration of the display is much longer (giving the impression of a dance) and the body can alter¬ nate betw^een an oblique and horizontal position. These w^ere observed eight times in the wild and they always occurred either at the boundary or within the neighbors’ territory. A typical example is taken from my field notes : June 17, 1966. Wing Hollow. Male IIS (not paired) is invad¬ ing (establishing territory?) the territory of pair 12S during the courtship period. He appears very nervous and flies to the tele¬ phone crossbar and assumes a horizontal sleek and walks along dragging the depressed and fully spread tail. Now he flies to a post and Warbles .... male flies back to crossbar and assumes an Oblique Sleek, tail spread and wings drooped, then walks along dragging both tail and wings, bows, leaves .... flies back to the crossbar, assumes Oblique Sleek, tail spread, droops wings then walks fast with head down one way, pivots, walks the other way (head still down), bill wipes, giving Rapid Warble con¬ stantly, feather ruffles, back into Oblique Sleek, tail spread, wings drooped, and walks along crossbar. Warbling. Thus, in this exaggerated Oblique Sleek, the bird has a strong tendency to behave sexually and also a strong tendency to flee, the conflict being revealed by the walking movements, displacement activities, and the obvious ambivalence. The walking is probably an ambivalent movement that rapidly alternates between intention movements to approach and flee. As far as I know this is the only situation in the bluebird’s life where it walks (see locomotion, p. 2). Two displacement activities are seen in the display, bill-wiping and feather-ruffling. I am uncertain of the Oblique Sleek posture with the head down. It might be an ambivalent posture with the Oblique Sleek being a flight intention movement and the head down an inten¬ tion movement to peck. Wings-raised Sleek In this display the male faces the female. In its most intense form the body is horizontal, plumage sleeked, and the wings raised to or above the shoulders with a slight downward rotation relative to their articulation with the body. The tail is maximally spread and arched downward, legs flexed, and the bill open and slightly raised. The variable components are the wings which can be stationary when raised or quivered, lifted, or waved. The bill is usually closed and directed forward and nest material can be carried in it. Whisper song often accompanies this display and Squeals may be given. The dura¬ tion of the display is very short, rarely lasting more than 5 seconds. [103] Fig. 26 The Oblique Sleek display of the Eastern Bluebird accom¬ panied by Rapid Warble. Dotted line indicates pulsating throat Fig. 27 The Wings Raised Sleek display of the male Eastern Bluebird [104] Wings-raised Sleek was observed 37 times in the wild, often near the nest box. Table 13 shows the behavior of the male immediately following the display. TABLE 13 Wings-raised Sleek Display : Behavior of the Male Immediately Following the Wings-raised Sleek Display Stage Nest demon- None stration Chase Chase- attack Attack Copula¬ tion attempt Pre-pair Formation (female in vicinity) Pair formation 4 4 1 6 2 2(1) 1 Prenesting building 1 4(1) Nest building 1 2(1) (6) (1) Numbers — the number of observations. ( ) — number of observations when display was preceded by Song Duel or Whisper Rapid Warble. When neighboring or migrating females are in the immediate vicinity of an unpaired male’s territory he may show a low intensity Wings- raised Sleek and then give Nest Demonstration Display. During early pair formation the same holds true but once the pair bond is formed this display leads to increasing attacks on the female, especially in the nest building period. On five occasions it appeared to be initiated by intense Song Duels with neighboring males, perhaps as a result of redirection. On five other occasions it was preceded by Whisper Rapid Warble which would increase in frequency until the male would suddenly assume the Wings-raised Sleek and violently attack the female. Table 14 shows the context of the display in the wild for one male during different stages of courtship. [105] TABLE 14 Context of Wings-raised Sleek During Different Stages of Male IIS Courtship Period Copula- Nest demon- Chase- tion Stage None stration Chase attack Attack attempt Pair For. June 5 5 1-2-3-4* 6-10 8 7-9 345 minutes** Nest build June 6 1-2-3-4— 5 240 minutes Female leaves June 7 Female No. 2 Pair For. June 20 1-3 2 315 minutes Female leaves June 21 Female No. 3 Pair For. June 25 — no observations Nest build June 26 1-2-3 300 minutes Nest build June 27 1 180 minutes Nest build June 28 1 30 minutes * Note ; The numbers indicate the sequential pattern of change in the males' behavior following each display and therefore refer to one observation, not to the number of observations. ** Total period of observation. In the beginning of pair formation before the female has entered the box, the male does not attack the female from the Wings-raised Sleek but gives Nest Demonstration. Once the female has entered (and accepted?) the nest box, the male becomes more aggressive and this behavior leads to Chases or Attacks on the female. When the female starts to carry nest material she no longer flees in response to Wings- raised Sleek but remains and sometimes give a low intensity Solicit ; yet the male still attacks. The evidence (tables 13 and 14) indicates that the Wings-raised Sleek primarily involves a strong tendency to attack the mate. How¬ ever, when observing this display in captivity it becomes evident that a strong sexual tendency is also activated. Table 15 shows the ontog- [106] eny of this display in one pair of captive birds. During early courtship the display leads to attack but as courtship continues Wing-raised Sleek becomes a precopulatory display. To cite typical examples : April 12, 1967. St. Bonaventure Aviary. Pair DY and DG. Male assumes Wings-raised Sleek and female Solicits. Male mounts and copulates, hops off. Faces the female and gives Wings- raised Sleek, mounts female and copulates, then leaves. April 14, 1967. 1 :10 p.m. Male is Fluffed on the stove pipe. Female flies up to the male and Solicits. The male shows Wings- raised Sleek, mounts female and copulates, then leaves. 1 :15 p.m. Male assumes Wings-raised Sleek, faces female who Solicits and Squeals, male mounts and copulates. Male does this twice then leaves. 1 :19 p.m. Male copulates with female without showing Wings- raised Sleek. However, in the wild, Wings-raised Sleek was never observed to func¬ tion as a precopulatory display. Note also that in the 11 displays that led to attack, seven were preceded by Song Duels between the captive birds. TABLE 15 Wings-raised Sleek Display by Male DY in Captivity, 1967 Stage Prenest Building Building April 7 8 Behavior of male following display None 9 10 12 1 13 14 17 18 Attack 1* 2 4 3 1 (1)** Copulation (4) (1) (1) attempt Copulation with¬ out display 1 3 Starts 4 1 * number of observations **display was preceded by Song Duel [107] I display Fig. 29 The Wing-lifting Tail-up display of the fe¬ male Eastern Bluebird [108] Not only does this display often lead to actual attacks or Chases (59 percent in the wild) but it resembles both the Head Forward and Wings Out Threat Displays. The elements of horizontal body, sleeked plumage, raised wings, and frontal orientation all are intention movements of attack. Wing quivering, lifting, and waving, all impor¬ tant elements in the display, occur only in the breeding season. The depressed spread tail is a common element during copulation as is Squeal. Hence these elements, plus the wing movements, probably all denote a sexual tendency. It appears therefore that the Wings-raised Sleek involves a strongly activated attack tendency, a fairly strong sexual tendency, and a rela¬ tively weak tendency to flee. Fluffed Posture This is similar to the Fluffed posture used in agonistic situations (see pg. 49), but usually is more marked. In its more intense form during courtship the body feathers are fluffed, neck withdrawn, wings slightly drooped below the tail, and the crest and rump are ruffled. This posture is frequently used, along with tail spreading, when ren¬ dering Warble. It is frequently seen in situations when the escape tendency would be expected to be relatively high, for example after pair formation when the female becomes more aggressive toward the male around the nest box. At this time the male usually sits by the box, in a Fluff, watching the female build. The Fluffed posture often involves an activated sexual tendency as the male gives Wing-lifting (or Waving) and Rapid Warble. In this posture the tendency to attack the mate is relatively low. However, when accompanied by Whisper Rapid Warble the male sometimes attacks the female. To cite one example : June 28, 1967. Wing Hollow. Male IIS is in an oblique Fluffed posture. He Wing-lifts and gives Whisper Rapid Warble faster and faster. He now pivots on the perch and the Wing-lift changes to Wave, the Whisper Rapid Warble becomes more in¬ tense, and then the male flies 50 yards, lands on the female’s back, and Pecks her head violently. It appears that Whisper Rapid Warble and Wing-lifting raise the relative strengths of the sexual and attack tendencies, and on close contact with the female, the attack tendency is expressed. The Fluffed posture is also frequently shown between paired birds just prior to copulation. The birds remain immobile and quite pas¬ sive for long periods of time. [109] The Fluffed posture is opposite in form to the two sleeked dis¬ plays just described and probably functions as appeasement by signal¬ ing the submissiveness of the bird. Wing-lifting and Waving These are both very common displays during courtship and are associated with an activated sexual tendency. Both are discussed on pg. 81. Symbolic Nest Building This is the manipulation of nesting material by a member of a pair which does not ordinarily help construct the nest (Nice, 1943). It is a very common display in the bluebird, especially during Nest Demon¬ stration. The male will hold a piece of nest material in the bill, some¬ times mandibulating it, as he proceeds through the different phases of Nest Demonstration. Usually he gets the material from inside the nest box but will occasionally carry it to the box. It almost always occurs during pair formation and continues well into the nest building period. On two occasions, captive males were observed to go through nest¬ shaping movements while inside the nest box. This same behavior was also noticed by Hartshorne (1962). Power (1966) observed symbolic nest building in Sialia currucoides where the male went through all the motions of picking up nest material but did not actually pick up anything. Curio (1959), in describing the behavior of the male semicollared flycatcher {Ficedula semitorquata) at the nesting hole, stated that the male brings out nest material (left by starlings) but he called this cleaning. Although the female constructs the nest (Hartshorne, 1962; Thomas, 1946; pers. obs.), there are reports of males sometimes par¬ ticipating in this activity (Bent, 1949; Hamilton, 1943; Hartshorne, 1962; Knight, 1908). Manipulation of nest material may be a vestige of behavior of a former time when the male may have had a more important role in nest building. This behavior may stimulate the female to approach the nest and/or to build. When the nesting cycle is interrupted (predators, etc.) the male will almost always resume Nest Demonstration Display while carrying nest material in his bill. One such male gave Nest Demon¬ stration Display 38 times in 1 hour ; most of the time with nest material in the bill. Incomplete activities often have great importance because of their signal significance or stimulatory effect upon the mate (Tin¬ bergen, 1951). [110] Flight Display Male bluebirds have several flight displays to advertise their ter¬ ritory (see pg. 77 for description). These persist after the female has entered the territory and becomes a definite part of courtship. Table 16 shows the occurrence of such displays throughout the court¬ ship period. The displays are performed during advertisement (24 percent), reach a peak during pair formation (53 percent), and then occur sporadically through nest building. TABLE 16 Occurrence and Number of Observations of Flight Displays by Male Bluebirds During the 1967 Breeding Season, Wing Hollow Flight Display Stage Not Paired Pair Formation Prenest Nest building building Butterfly Flight 19** 38 6 4 directed toward box 12 36 5 1 perch 7 2 1 3 Gliding Flight 7 24 9 8 directed toward box 5 19 9 6 perch 2 5 - 2 Lopsided Flight 5 7 2 - directed toward box 3 6 - - perch 2 1 2 - Butterfly/Glide 1 2 - - Total of all flight displays 32 71 17 12 ** Number of observations. Of the 132 flights that I observed (1967), 104 (78 percent) were directed toward the nest box, almost always ending in Nest Demon¬ stration Display. These flights show (to the female) the presence of the male and the location of the nest box. They are primarily gamosematic displays (see Lack, 1940b, pg. 281) by assisting the members of a pair to find one another and probably influence the female in selecting a mate at pair formation. [Ill] FEMALE DISPLAYS Sleeked Display Females were never observed to give Oblique Sleek or Wings-raised Sleek. During pair formation, however, they sometimes sleek the head while at the nest box. Fluked Posture This is similar to that given by the male but not as marked. It is usually associated with a thwarted tendency to flee and probably functions as appeasement. It is especially evident during pair forma¬ tion. Wing-lifting and Waving Similar to the male. Symbolic Nesting During Nest Demonstration the female may carry nest material in the bill. However, in the female it may result from low motivation for nest building. Solicit This is a stationary display and at low intensity the female crouches with plumage compressed, head drawn in, and bill slightly raised. The wings are drooped with their tips below the base of the tail ; the tail is slightly raised (also see Hartshorne, 1962). Low intensity Solic¬ iting can be given in early courtship in response to approach by the male. This rarely leads to successful copulation but rather results in threat or even attack ; the female maintains the Solicit as the male lands on her back and repeatedly Pecks her head. Occasionally the attack tendency is expressed in the female as she Faces, Gapes, Rasps, and sometimes Pecks the male. At higher intensity (usually when the nest is completed) the female crouches in a horizontal position, plumage compressed, with the feath¬ ers of the crown and back especially sleeked. The bill is slightly raised and the tail is raised markedly. The wings are drooped and can be quivered. Usually Squeal and peeps are given. The Soliciting posture is the preliminary to actual copulation and clearly involves a strong tendency to behave sexually (Hinde, 1954). In the bluebird the attack tendency is also weakly activated as shown by the occasional threat toward the male. Rarely does the female flee. [112] Wing-lifting Tail-up Display The body is held in a horizontal position with the plumage either flufifed or compressed. The legs are slightly flexed and the tail is raised markedly but not spread. The head is raised but can (rarely) be lowered in a bow. The wings are continually lifted or even waved and the bird may move by pivoting or hopping. A variety of vocaliza¬ tions can be given- — Rapid Warble, Squeal, “ whines,” or Peeps. In 1967 this display was seen 14 times in the wild and in two of these situations it was repeated 10 and 12 times in succession. In 10 cases it was the female who displayed ; three cases, the male ; once, both displayed. The display was observed when the bird was greatly ex¬ cited and usually in a conflict situation as shown by the following figures : 1. Boundary disputes 6 cases 2. Courtship feeding 4 cases 3. Nest Demonstration 3 cases 4. Soliciting 1 case In addition to these it was observed in captivity. The Wing-lifting Tail-up Display resembles the female precopula- tory display of other passerines, e.g. the chaffinch, Fringilla coelebs (see Andrew, 1961, pg. 345). In the bluebird it may be the highest intensity of the Solicit posture, yet it differs from the Solicit in many ways : 1. it (usually) is not a stationary display 2. Wing-lifting and Waving occur 3. it never resulted in copulation (or even an attempt) 4. it is usually given in situations that suggest conflict Five typical Wing-lifting Tail-up Display situations from my field notes are described below : 1. Pair 5N are excited around their nest box. The female flies to a tree and gives Wing-lifting Tail-up Display. Male follows and lands by female who then Supplants him. Female remains in Wing-lifting Tail-up Display and flies from branch to branch, male gets excited and female Supplants him. 2. Male 12S and IIS fight at the territorial boundary. Female 12S gets excited and gives Wing-lifting Tail-up Display on wire above Fighting males. Female pivots on wire as she displays. . . . .after more disputes the male is guarding the female when she flies to ground, returns, and lands close to the male. The male then gives Wing-lifting Tail-up Display toward his female. Male IIS comes and both males fall to ground Fighting. [113] 3. I put a decoy eight feet from the nest box (young 3-days old) of pair 2S. Male comes immediately and attacks decoy so I re¬ move it and just play song. Male and female get excited and go to the nest box. On top, the female assumes the Wing-lifting Tail-up Display while turning and Squealing. Male approaches, hovers over the female but does not mount. As I play song the female does this 12 times in a row each time the male hovering but never mounting. 4. Male IN is banging a large caterpillar on the fence post. Fe¬ male IN assumes Wing-lifting Tail-up Display, flies to male, and gives Wing-lifting Tail-up Display in front of him. The male keeps moving away and 10 consecutive times the female flies to him and gives Wing-lifting Tail-up Display. Male Anally leaves and female goes back to nest box. 5. Male IIS forms a bond with female 14S who has a broken lower mandible. Female apparently cannot carry nest material. Female is on the fence line and assumes Wing-lifting Tail-up Display and male comes and feeds her. When observing this display one gets the impression that the female is highly motivated and in some situations invites copulation, in others, feeding. Yet in all observations she achieved neither (except number 5 above). Morris (1954) stated that in some species the ferqale’s invitation-to-copulate display is identical with the juvenile food¬ begging display. In the bluebird, the Wing-lifting Tail-up Display definitely resembles both the female’s Solicit and food-begging posture, yet differs from each. In Soliciting the wings are not lifted or waved and in food-begging (courtship feeding) the tail is not raised. Since sufficient observations are lacking the function and signifi¬ cance of this display remain obscure. Both a highly activated sexual and feeding tendency are involved and when the display occurs during boundary disputes it may possibly be a displacement activity. The occurrence of this display in the male (four cases) may be due to a thwarted sexual tendency and hence a pseudo-female display (see Morris, 1955). MUTUAL DISPLAYS Nest Demonstration This is the display that the male uses to advertise himself and the nest box to the female (see pg. 81) for complete description. During this display the pair bond is formed. The female takes an active part in the display and can perform all its components. After pair forma- [114] tion and throughout courtship the pair still perform the display. In this context it probably functions to sexually stimulate both members of the pair and to strengthen the pair bond by suppressing the agon¬ istic tendencies of each. This is also evidenced when a nesting is terminated; Nest Demonstration again becomes predominate. This appears to be a critical test, for if the female will not participate in the display and therefore not accept the nest box, either the female or the pair will leave the territory. Usually, the female does participate (especially if other nest boxes are in the territory) and the pair re¬ nest. However, if she does not participate and leaves, the male’s at¬ tachment to the territory is usually stronger than the pair bond and he remains and starts to advertise again. John Burroughs (1871) recognized this when he wrote ; “. . .He (Eastern Bluebird) thought the box just the thing, and that there was no need of alarm, and spent days in trying to persuade the female back. Seeing he could not be a stepfather to a family, he was quite willing to assume a nearer relation. He hovered about the box, he went in and out, he called, he warbled, he entreated ; the female would respond occasionally and come and alight near, and even peep into the nest, but would not enter it, and quickly flew away again. Her mate would reluctantly follow, but he was soon back, uttering the most confident and cheering calls. If she did not come he would perch above the nest and sound his loudest notes over and over again, looking in the direction of his mate and beckoning with every motion. But she responded less and less frequently. Some days I would see him only, . . . . ” Sexual Chasing Sexual Chases in the bluebird involve a rapid pursuit of the female by the male. Most occur during the pair formation and prenesting period. The motivation of the male is variable. Sometimes Chases were preceded by some form of aggressive display. For instance, on 11 occasions the male assumed the Wings-raised Sleek (strong attack tendency) and moved toward the female who responded by flying off rapidly with the male in close pursuit. Chases could also be preceded by signs of sexual excitement. The pair could be actively involved in Nest Demonstration when a Chase would suddenly develop or the male could give Rapid Warble with Wing-lifting Display then chase the female. Some Chases appeared to develop “ spontaneously ” ; the pair inactive when a rapid Chase would suddenly take place. The Chase is a rapid, sometimes twisting flight, yet in only one Chase did the male actually catch the female. A variety of vocaliza¬ tions can be given, especially Chatter (attack tendency). Rapid [115] Warble, and Squealing (sexual tendencies). Chases always occur within the territory and the female may only leave its boundaries dur¬ ing pair formation. On two occasions the male overtook the female and both times he landed at the nest box and performed Nest Demon¬ stration. The pair rarely end the Chase close together ; the male usually lands several feet from the female. The female usually assumes the Fluffed posture, the male the same or the Oblique Sleek. Table 17 shows that 68 percent of the recorded Chases ended with no interaction between the pair. However, in 21 cases, after the female landed, the male went directly to the nest box and gave Nest Demonstration. In the bluebird, it appears that Sexual Chases can be an expression of either an aggressive or sexual tendency in the male. At their termination though these tendencies usually become inhibited by the escape tendency. While the male shows either an activated attack or sexual tendency at the beginning these are rarely expressed at the end by actual attack (1 case), or copulation attempts (2 cases); usually he lands several feet from the female (escape tendency being predominant). However, the male may still show sexual excitement at the end by giving Nest Demonstration Display. In the 11 Chases preceded by the Wings-raised Sleek, five ended in Nest Demonstra¬ tion; the male’s apparent motivation changing from aggressive to sexual. It appears therefore that during the Chase either the male’s (depend¬ ing on the motivation at the start) attack or sexual tendency is low¬ ered. This is advantageous to a species that has a strong aggressive element in its courtship. Sexual Chasing then may provide a pro- TABLE 17 Occurrence of Sexual Chasing and Resultant Pair Behavior Stage observa¬ tions attack Chase ends in copula- nest demon- tion stration* breaks off termi¬ nates in dense foliage Pair formation 39 1 0 11 19 8 Prenest building 22 0 0 9 9 4 Nest building 15 0 2 1 8 4 Totals 76 1 2 21 36 16 *only the male participates [116] visional outlet (as suggested by Marler for the chaffinch, 1956) for high copulatory or attack motivation thus allowing the mating activi¬ ties of the male and female to become better synchronized. The effects of Chasing on the female are not clear. The escape tend¬ ency is certainly activated yet she seldom leaves the territory. She certainly benefits from those Chases preceded by aggressive display by avoiding an actual attack. Courtship Feeding Fully developed courtship feeding actively involves both mates ; the male gives the prefeeding call and brings the food to the begging female. There is, however, much variation in the pattern, depending on the level of motivation of each sex and on the conflicting agonistic tendencies associated with the sexual and feeding tendencies. In its early stages it may be incomplete and sometimes results in agonistic encounters between the pair. The male usually initiates courtship feeding with the female generally responding in one of four ways ; As the male approaches with food the female may 1. flee (rarely) 2. threaten him 3. remain passive and accept the food, or 4. adopt a begging posture similar to that used by young bluebirds. When actively begging, the body is in a horizontal to oblique position, plumage somewhat fluffed, one or both wings lifted in rapid succes¬ sion, and begging calls are given. In some cases (especially in captivity) courtship feeding first in¬ volved fleeing or threat, then passivity, and finally active begging by the female. However, in the wild it was extremely variable with some females actively begging all the time. Later in the courtship period the female may start to beg even be¬ fore the male captures the prey or else fly to him and beg as he mouths the prey in his bill. During incubation, apparently without stimulus from the female, the male feeds the female inside the box. Evidence exists that the male’s feeding tendency is waxing (espe¬ cially in late arrivals) before and during pair formation. An unpaired male was observed giving the prefeeding call as he mandibulated an insect before swallowing it, and during symbolic nest building males will sometimes work nest material as if it were an insect. In one unusual case male 5N, who had recently (approximately 4 hours) formed a bond with female 5N, was observed to perform a possible overflow activity. Overflow activities (vacuum activities of Lorenz, 1937) occur when an animal has one drive strongly activated and the [117] releasing stimuli, indispensable for the performance of the executive motor patterns of the drive, are inadequate. They are apparently nothing but reactions to suboptimal stimuli (Bastock, Morris, and Moynihan, 1953). Male 5N was on a fencepost 10 yards from the nest box. He bent over in a horizontal position and moved the bill as if “ working ” an insect. Yet the bill was empty ! He touched the bill to the post and then banged it one way then the other as he worked the invisible insect. Feeding the female by regurgitation was only observed in captivity. This may occur in the wild for one pair was observed to engage in billing where both mates faced each other in Fluffed postures as they touched bills. The male left, returned immediately, and repeated the same performance. This is the only time billing was observed in the wild but captive young birds will often touch bills and then gently Peck one another. In Myadestes imicolor (grey solitaire) Isenberg (1948) reported that the male feeds the female by regurgitation. In the Wing Hollow population, courtship feeding is a common and necessary part of courtship and appears to serve two major functions : 1. to maintain and strengthen the pair bond, and 2. to reduce the agonistic tendencies between the pair. Evidence for the former is that courtship feeding occurs in all phases of the reproductive cycle (table 18). The male also frequently feeds the female 1. during lulls in boundary disputes, 2. after attacks upon her, 3. during the interval before renesting after an aborted attempt. Regarding the latter function, the bluebird in the Wing Hollow area has a strong aggressive element in its courtship. Throughout the courtship period the male makes repeated attacks upon the female whereas the female makes Supplanting Attacks upon the male in the vicinity of the nest box. It is suggested therefore that courtship feed¬ ing may suppress the agonistic tendencies in the pair by permitting each bird to become habituated to the proximity of its mate. For instance, a male aggressive to its mate at one moment will, in the next moment, approach and feed the female with no sign of aggressiveness. This is best illustrated by the following field observation : Wing Hollow, June 6, 1967 ; Pair IIS. 11 :25 a.m. Female comes to fence line by the box, male turns toward her, assumes Oblique Sleek, shivers the wings, female goes in box. Female comes out and lands in front of box. Male (with no display) comes at and hits her with his feet, then Pecks [118] her head, hops up in the air and repeats this 8 to 10 times finally knocking female upside down on the fence line. Male leaves and female goes back to nest building. 1 1 ;28 a.m. Male comes with insect and sidles up to female and feeds her ; both face away, male leaves. Also when the male approaches the female with food his feeding tend¬ ency dominates both the attack and escape tendencies. In well over 100 observations I have never seen a male display strong aggression during courtship feeding. Likewise, while studying the relationships between agonistic behavior and the social hierarchy in captive blue¬ birds, out of more than a thousand encounters, very few occurred over food. Dominant and subordinate birds often would feed together. Therefore, despite threat or even fleeing by the female the male is persistent in his attempt to feed her. TABLE 18 Occurrence of Courtship Feeding During the Different Phases of the Nesting Cycle Stage Number of observations Pair Formation 11 Prenest Building 24 Nest Building 66 Incubation frequent Lack (1940a), in his review of the literature, stated that the eastern bluebird’s habit of courtship feeding is rare and many individuals apparently do not show it. Thomas (1946), in her 14-year study of an apparent nonmigrant population in Arkansas, found that court¬ ship feeding did not occur at all, nor were females ever observed to beg during the courtship period. Yet both Hartshorne (1962) and I (working with migrant populations) found courtship feeding common. It is possible that courtship feeding occurs prominently in social interactions between the pair in a migrant population but is apparently rare or nonexistent in some nonmigrant populations. Thomas (ibid.) did not mention any aggressive element during the courtship period. She stated that most pairs were formed between November and the last of January, nesting started approximately March 7, and in old mates there appeared to be a real bond the year around. Nonmigrant populations may have no need for a mechanism such as courtship feeding to reduce the agonistic tendencies between the pair during courtship. Apparently this is resolved before nesting begins, possibly [119] [120] Fig. 30 Courtship Feeding in the Eastern Bluebird. Male is on right. by Nest Demonstration Display which, according to Thomas, occurs throughout the nonbreeding season. The Wing Hollow population not only incorporated courtship feed¬ ing into their reproductive cycle, but also had “ helpers at the nest ” (i.e., young of first brood help feed young of second brood). Thomas (op. cit.), never observed feeding or helpers at the nest. Copulation Successful copulation is sometimes initiated by the male; he flies toward the female who adopts the Soliciting posture as the male lands on her back. Usually though the female initiates copulation by giving a marked Soliciting posture. The male flies either directly to her back or else lands beside her, then up and on. During copulation the male waves his wings as the tail is brought down and cloacal contact is made. Two vocalizations can be uttered, Squeal by either partner and Peeping by the female. No post-copulatory displays were noted. Successive mountings were seen only in captive birds. Once the male mounted two times, and twice, three times before leaving. Copulation attempts may begin before nest building and terminate part way through the incubation period. They can occur on any convenient perch within the territory. In most copulation attempts observed (both in the wild and cap¬ tivity) the male was influenced by conflicting tendencies to attack and behave sexually with the female. The attack tendency was almost always expressed by Pecking the female’s head while standing on her back. It was frequently difficult to determine if the male was actually attempting copulation or just attacking the female. Only if the male depressed the spread tail was it considered a copulation attempt. In the wild, 19 such attempts were recorded and in 14 the male Pecked the female’s head. Only once did the female Peck the male and this occur ed after a successful copulation. In captivity, attacks on the female during copulation were common. For example : Nest box has two eggs. Male goes in the box and female Squeals. Male comes out with nest material in bill. Wing-lifts and gives Rapid Warble, now he approaches female who is Soliciting. Male lands on her back and Pecks her head over and over with tail spread and waving both wings. Male now stops Pecking and depresses his tail as if to copulate. Male leaves and then returns and feeds female. Hartshorne (1962) saw one of his laboratory males Peck the head of the female during copulation but did not see this in other bluebirds. [121] Fig. 31 Female Soliciting and male attacking behav¬ ior. In upper figure the female (on left) is giving a low intensity Solicit as the male is about to attack. The lower figure shows the male now violently attacking the female by pecking her head. [122] On four occasions neighboring males interrupted copulation attempts by attacking the copulating pair. I never observed reverse mounting but Pearse (in Bent, 1949) reported this in the western bluebird (Sialia mexicana) ; the male mounted the female and then the female mounted the male. SYNOPSIS OF THE COURTSHIP PERIOD The length of the bluebird’s courtship period varies. In birds that pair early in the season (late March-early April), four weeks may elapse before successful copulations take place whereas in late pair formations (early June) only 24 hours may elapse. Weather is an important factor, for during a period of low temperatures, courtship activity ceases and the pairs become inactive and difficult to locate. Before actual nest construction begins the male still Warbles and patrols the territory. The pair forage and move about the territory together keeping in contact by calling “ tu-a-wee,” yet individual dis¬ tance remains strict. Boundary disputes are common and both mem¬ bers actively defend tbe territory. Most sexual behavior is centered around the nest box. The male takes the initiative and displays at the box showing Wing-lifting, symbolic nest building. Rapid Warble, and Nest Demonstration. Shortly, the female takes an active role and the frequency of her Supplanting Attacks on the male increase. Nest Demonstration now becomes the principal display as the pair inspect (with either sex leading) all the boxes in the territory. While at the boxes the female may show searching behavior (Hinde, 1954, pg. 229) by hopping along the ground and going through the motions of looking for nest material but not actually picking any up. Later she may start to carry pieces of grass into and sometimes out of the nest boxes. The male still Chases the female but she never leaves the territory. Later still the pair center their activities around one box and engage in much mutual sexual display. Finally the female starts to build and this causes a rise in the male’s sexual and attack tendencies. He remains close to the box and may inspect the nest in her absence, often bringing nest material out, or he may fly inside and face-show as she approaches. Some males follow the female back and forth to the box or Cbase her as she comes out. Low intensity Solicits are frequent and courtship feeding is common. At this time the female’s Supplanting Attacks on the male decrease and the male becomes more aggressive and increases his attacks on [123] the female. As the nest nears completion these attacks become more associated with copulation attempts as the male rapes the female. When the nest is completed the pair spend little time near the box. They remain together and retire to the shrubs and trees in the ter¬ ritory, spending considerable time in maintenance activities. Both show the Fluffed posture, the male feeds the female quite regularly now and there are bursts of sexual excitement as copulation takes place with no aggressiveness between the pair. During the parental phase some of the courtship displays can be given in certain situations. Wing-lifting remains prevalent and is a mutual display given much like a “ greeting ” when the pair meet after a sustained absence. For example, if one bird is perched and the other, after an absence, lands close by, both start Wing-lifting as they perch close together. This may be a form of reassurance (sensu. Smith, 1966) in which each bird communicates that it will not likely attack the other. [124] SUMMARY The general behavior of the eastern bluebird, Sialia sialis, was studied for 5 years, the last two primarily devoted to agonistic and courtship behavior of both captive and wild color-banded birds in western New York State. Eight nestlings were reared by hand and observed in captivity for 2 years. Observations on wintering bluebirds were made in the vicinity of Tallahassee, Florida. Maintenance activities and the behavioral patterns associated with agonistic and courtship behavior are described in detail. Interpreta¬ tions are offered for the functions, motivations, derivation, and bio¬ logical significance of some of these behavioral patterns. Comparisons and homologies are made with behavioral patterns of other species, especially other chat thrushes and true thrushes (Ripley, 1952). Previous studies such as those of Laskey (1939, 1940, 1943), Thomas (1946), and Hartshorne (1962) have stressed different as¬ pects of bluebird behavior. The information summarized here is either new or reinterpreted in the light of recent ethological thought (see Dilger, 1962). Maintenance Activities Maintenance activities of the bluebird were studied in the field and in captivity and resemble those of most passerines that have been studied. They have been described and their ontogeny traced whenever possible. Care of the body includes 12 activities. Head-shaking is common in adults and is associated with other toilet (for use of this term see Andrew, 1956) behavior. The head is scratched indirectly. The three stretching movements are similar to other North American Hylocichla and Catharns species but the sequence is different. Both wings down occurs only in nestlings and is not a transition movement. [125] Two methods of bathing are used and alternate wing movements . (motion 3 of Nice, 1943) do not occur. Preening, bathing, and sun- j ning all have social qualities. 'i Typically the species forages by scanning the ground from elevated j perches. Midair catches are made and food may also be obtained by '< hopping along the ground. Preparation of prey before swallowing j is by “ scissoring ” and/or “ hammering.” Both animal and vegetable matter are consumed, the latter mainly in the winter. The ontogeny of locomotion is given. Movement along a perch is by hopping, sidling, or pivot-hopping. Pivoting is a common move¬ ment while perching. The behavior and location of communal roosting, during adverse weather, is described. Agonistic Behavior Agonistic behavior in captive and wild bluebirds generally resembles that of other North American thrushes studied but differs in detail. Four attack responses are described : Supplanting, Chasing, and Fight¬ ing are associated mainly with territorial defense, and Pecking when individual distance is violated. The threat code consists of seven dis¬ plays which showing varying intensities of the attack and escape tend¬ encies. In S', salis, Wings-flicked reflects a high attack tendency and is accompanied by Tail-fanning in a horizontal plane rather than Tail- flicking in a vertical plane as in Hylocichla and Catharns. Wings-out was only given in response to Scream and was not directed toward conspecifics. Bill-raising has two intensities and incorporates ambivalent movements and feather adjustments. Eight vocalizations accompanying agonistic displays are described. Crouch and Alert are adaptive responses to unfamiliar stimuli. Wing-flashing, which occurs frequently in captive birds in response to novel stimuli, was only observed once in the wild. Feather-ruffling is a common displacement activity and is similar to the Spread Display of Hylocichla miistelina but differs in not being- ritualized to serve as a display. The Fluffed posture seen in pair interactions is not a ritualized appeasement display. In captivity redirected attack occurred as a result of intensive song duels between males in adjoining cages. On leaving the nest, young bluebirds remain together in a close family unit and may help to feed the young of the second brood. In late summer, juveniles from different family units join together to form small mobile flocks with the individuals showing much agonistic and incipient sexual behavior. The size of migrant flocks is variable and [126] on the wintering grounds a flock of 100 or more bluebirds represents several subgroups. Nest inspection was not part of the behavior of wintering flocks but was common in captive birds during the nonreproductive season. In captivity young from two different broods (subgroups) were kept together from October to March in a large 10-cubic-foot aviary. Both subgroups were organized in a peck-dominance hierarchy; the males in one were dominant over the females but in the other sub¬ group the females were dominant. The subgroups never became integrated, as one subgroup completely dominated the other. This resulted in a complex peck order maintained by both peck-dominance and peck-right. It is suggested that (in captivity at least) the mem¬ bers of a brood retain a social bond to one another based on incom¬ plete dominance. Complete dominance is only attained over members from different broods. T erritory Bluebirds defend large territories in which courtship, mating, nest¬ ing, and most food-seeking occurs. Possession of a nest cavity in open terrain is a prerequisite before the territory can be established. Nest cavities are in short supply and both interspecific and intraspecific competition for ownership is intense. As a result, aggressive behavior is employed by males in acquiring nest cavities, which provide the opportunity to mate and begin nesting. It is important to both the maintenance of the territory and the pair bond that the male select and defend more than one nest box. Nest boxes are also instrumental in determining the size and shape of the territory. As the population increases some territorial boundaries are frequently changed. The changes are caused by early arriving males (or pairs) taking posses¬ sion of a very large territory, and then being forced to yield parts of it to new arriving males. Boundaries that contain empty nest boxes remain rather fluid with possession determined by variations in the breeding activities of the neighboring pairs. Territories in the Wing Hollow area average 13.6 acres in size, but they vary all the way from less than 5.3 to 28.7 acres. The population could not be increased significantly by putting up more nest boxes. The males do not always arrive first. Arrival is a staggered affair with male and females arriving from mid-March to early June. Some birds arrive already paired. The territory is advertised mainly by Warble, six Aerial Displays, Wing-lifting and Waving, and Nest Demonstration Display. Warble is the principal display in main- [127] taining the territory and Song Duels between neighboring males are common. At the territorial boundaries, displays showing a conflict of attack and escape tendencies are prevalent, especially the Oblique Bill-up Display. When conspecific intruders come within the ter¬ ritory and near the nest box the resident males usually react with Supplanting Attacks or displays showing a strong attack tendency. Bluebirds in Wing Hollow usually maintain the territory throughout the breeding season. Experiments with decoys showed that the ter¬ ritory, per se, and not just the nest box is defended and defense con¬ tinues throughout the nesting cycle. This allows more than one brood to be raised and has certain selective advantages. The birds do not have to waste time and energy locating new nest cavities (which are in short supply) and reestablishing territory. The nature of territory in the bluebird is applicable to Tinbergen’s ( 1957) concept that territory results from two tendencies in the owner, site attachment and hostility. Natural selection would favor these two in the bluebird which has special territorial requirements (nest cavity) in short supply and no hole-forming capabilities, as with the titmice, to compensate for the lack of nest cavities. The most important functions of territory in the bluebird are the facilitation of pair formation and maintenance of the pair bond. Pair Formation and Courtship The mechanism of pair formation as it occurs within the territory of unpaired males is described. In the developing pair bond, the female must take an active part in Nest Demonstration Display and enter the nest box while the male is inside. To insure the establishment of the bond, she must express her dominance by making Supplanting Attacks on the male in the vicinity of the nest box. After the pair bond is formed, its structure is based on incomplete dominance between male and female. Bluebirds are somewhat sexually dichromatic, so the male can recognize the female as distinct from other males and reacts aggressively to a male and sexually to a female intruder. Pair formation increases the relative strength of the attack tendency in both sexes and a main function of courtship is to suppress this attack tend¬ ency. This is achieved by much mutual sexual displaying in which Nest Demonstration Display and Courtship Feeding are especially important. Six male displays are described. The Oblique Sleek Display, seen primarily in early courtship, includes many variable elements. It reflects a strong sexual tendency in conflict with the escape tendency and when given at territorial boundaries may develop into a dance. [128] The Wings-raised Sleek Display involves a strong attack tendency and during the nest-building period develops into attacks on the female. In captivity this display developed into a precopulatory display result¬ ing in successful copulation without aggressive tendencies. Symbolic Nest Building probably has some stimulatory effect on the female. Butterfly, Gliding, and Lop-sided Flight are primarily gamosematic displays. The primary female display is the Solicit which has two intensities. Wing-lifting Tail-up Display resembles both the female’s Solicit and food-begging posture yet is different from each. The function and significance of this display is not known. Sexual Chasing is an expres¬ sion of either an aggressive or sexual tendency in the male and func¬ tions by lowering such tendencies. Nest Demonstration is a variable display and when mutually performed functions to stimulate the pair sexually and helps to strengthen the pair bond by suppressing their agonistic tendencies. Courtship Feeding occurs throughout the court¬ ship period and serves to maintain and strengthen the pair bond and to reduce aggressiveness by allowing each bird to become habituated to the proximity of its mate. The ontogeny of courtship feeding is described. Copulation can be initiated by either sex and may occur without preliminary display by the male. Most copulation attempts in the wild involved an aggressive element as the male pecked the female’s head. A synopsis of the courtship period is given. [129] [130] Fig. 32 Wing Hollow, looking east, April 29, 1970 [131] LITERATURE CITED Andrew, R. J. 1956. Normal and irrelevant toilet behavior in Emheriza spp. Brit. J. Anim. Behav., 4:85-91. - . 1961. The displays given by passerines in courtship and reproductive fighting: a review. Ibis, 103a :3l5-348, 549-579. Baird, S. F., T. M. Brewer, and R. Ridgway. 1905. A History of North American Birds. Little, Brown and Co. Bastock, M., D. Morris, and M. Moynihan. 1953. Some com¬ ments on conflict and thwarting in animals. Behavior, 6:66-84. Beal, F. E. 1915. Food of the robins and bluebirds of the United States. U.S. Dept. Agr. Dept. Bull. 171. Bent, A. C. 1949. Life histories of North American thrushes, kinglets, and their allies. U.S. Natl. Mus. Bull. 196. Blake, C. H. 1954. Death of a Bluebird. Bird-Banding, 25 :59. Blest, A. D. 1961. The concept of ritualisation. In W. H. Thorpe and O. L. Zangwill (ed.). Current problems in animal behavior. Cambridge Univ. Press. Brodrick, H. J. 1938. Nesting and re-mating of a pair of blue¬ birds. Auk, 55 :538-539. Brown, G. 1937. Aggressive display of birds before a looking- glass. British Birds, 31 :137-138. Brown, J. L. 1964. The integration of agonistic behavior in the Steller’s Jay (Cyanocitta stelleri). Univ. of Cal. Pub. in Zoology, 60:223-328. Buck, S. E., and N. Bates. 1962. Bluebirds. Kingbird, 13 :25. Burroughs, John. 1871. The Bluebird. The Writings of John Burroughs, 1 :205-216. Butler, A. G. 1907. The Bluebird in captivity. Avicultural Magazine, 6:50-53. [133] Conder, P. J. 1949. Individual Distance. Ibis, 91 ;649-655. Coutlee, E, L. 1963. Maintenance behavior of the American Goldfinch. Wilson Bulletin, 75 :342-357. - . 1967. Agonistic behavior in the American Goldfinch. Wilson Bulletin, 79:89-109. Griddle, N. 1927. Habits of the mountain bluebird in Manitoba. Can. Field Nat., 41 :40-44. Curio, E. 1959. Beobachtungen am Halbringschnapper, Ficedula semitorquata, in Mazedonischen Brutgebiet. J. Orn., 100:176-209. Daanje, A. 1950. On locomotary movements in birds and the intention movements derived from them. Behavior, 3 :49-98. Dilger, W. C. 1956a. Hostile behavior and reproductive isolat¬ ing mechanisms in the avian genera Catharus and Hylocichla. Auk, 73 :3 13-353. - . 1956b. Hylocichlid thrushes in captivity with notes on their behavior. Avic. Mag., 62:183-189. - . 1960. Agonistic and social behavior of captive Redpolls. Wilson Bulletin, 72:114-132. - . 1962. Methods and objectives of ethology. Living Bird, 1 :83-92. Drinkwater, H. 1953. Young Bluebird taken from nest box by Sparrow Hawk. Auk, 70:215. Dunham, D. W. 1964. Behavior of the Rose-breasted Grosbeak, Pheucticus ludovicianus (Linnaeus). Ph.D. thesis, Cornell Uni¬ versity. Eaton, E. H. 1914. Birds of New York. New York State Mus. Memoir 12, Part 2. Eaton, S. W. 1953. Birds of Olean and Salamanca Quadrangles. Science Studies, 25:1-27. Ficken, M. S. 1962a. Agonistic behavior and territory in the American Redstart. Auk, 79:607-632. - . 1962b. Maintenance activities of the American Redstart. Wilson Bulletin, 74:153-165. Ficken, R. W., and M. S. Ficken. 1958. Head-scratching in Seinrus (Parulidae) and other passerines. Ibis, 100:277-278. - . 1966. A review of some aspects of avian field ethology. Auk, 83:637-661. Fischer, R. B. 1944. Suggestions for capturing hole-nesting birds. Bird-Banding, 15 :15 1-156. [134] Forbes, S. A. 1880. The food habits of thrushes. Amer. Ent., new sen, 1 : 12-13. Forbush, E. H. 1929. Birds of Massachusetts and other New England States, Vol. 3. Frazier, A., and V. Nolan, Jr. 1959. Communal roosting by the Eastern Bluebird in winter. Bird-Banding, 30:219-226. Friedmann, H. 1929. The cowbirds, a study in the biology of social parasitism. Charles C. Thomas, Springfield, Ill., 421 pp. Haartman, L. von. 1956. Territory in the Pied Flycatcher, Miiscicapa hypoleiica. Ibis, 98:460-475. - . 1957. Adaptation in hole-nesting birds. Evolution, 11:339-347. Hailman, J. P. 1960. A field study of the Mockingbirds’ wing- flashing behavior and its association with foraging. Wilson Bul¬ letin, 72:34^357. Hamilton, W., Jr. 1943. Nesting of the Eastern Bluebird. Auk, 60:91-94. Harley, P. H. T. 1949. The biology of the Mourning Chat in winter quarters. Ibis, 91 :393-413. Harper, W. T. . 1926. A Bluebird’s nest. Bird-Lore, 28 : 187-190. Hartshorne, J. M. 1962. Behavior of the Eastern Bluebird at the nest. Living Bird, 1 : 13 1-149. Hauser, D. 1957. Some observations on sun-bathing in birds. Wilson Bulletin, 69:78-90. Heinroth, O. 1924. Lautavsserungen der Vogel. Jour. f. Ornith., 72 :223-244. Hinde, R. A. 1952. The behavior of the Great Tit (Pants major) and some other related species. Behavior, Supplement No. 2, 1-201. - . 1953. The conflict between drives in the courtship and copulation of the Chaffinch. Behavior, 5 :1-31. - . 1954. The courtship and copulation of the Greenfinch (Chloris chloris). Behavior, 7 :207-232. - . 1955-56. A comparative study of the courtship of certain finches (Fringillidae). Ibis, 97:706-745; 98:1-23. - . 1956. The biological significance of the territories of birds. Ibis, 98 :340-369. Horwich, R. H. 1965. An ontogeny of wing-flashing in the Mockingbird with reference to other behaviors. Wilson Bulletin, 77:264-281. [135] Howard, H. E. 1920. Territoiy in Bird Life. Murray, London, 308 pp. lersel, J. J. A. van, and A. C. A, Bol. 1958. Preening of two tern species. A study on displacement activities. Behavior, 13:1-88. Isenberg, A. H. 1948. Breeding of ‘ Clarin-Jilgaro ’. Avicult. Mag., 54:160-166. - . 1953. Breeding of Riv Kiv Island Robin. Avicult. Mag., 59 :208-210. Knight, O. W. 1908. The birds of Maine. Bangor, Maine. Kortlandt, A. 1940. Wechselwirkung zwischen Instinklen. Arch. Neerl. Zook, 4:442-520. Krieg, D. C. 1962. Bluebird nest box project. Kingbird, 14:2^27. Lack, D. 1940a. Courtship feeding in birds. Auk, 57 :169-178. - . 1940b. Pair-formation in birds. Condor, 42:269-286. - . 1953. The Life of the Robin. London : Penguin Books, 240 pp. Lane, J. 1968. A hybrid Eastern Bluebird X Mountain Bluebird. Auk, 85-684. Lanyon, W. E. 1957. The comparative biology of the meadow¬ larks (Sturnella) in Wisconsin. Publ. Nuttall Orn. Club, 1 :l-67. - . 1960. The ontogeny of vocalizations in birds. Pp. 321- 347 in Animal sounds and communication ( W. Lanyon and W. Ta- volga, eds.). A. I. B. S. Publ. No. 7. Washington, D.C. Laskey, A. R, 1939. A study of nesting Bluebirds. Bird- Banding, 10:23-32. - . 1940. The 1939 nesting season of Bluebirds at Nashville, Tennessee. Wilson Bulletin, 52:183-190. - . 1943. The nesting of Bluebirds banded as nestlings. Bird-Banding, 14:39-43. Lorenz, K. 1937. Veber die Bildung des Instinktbegriffs. Naturwiss, 25 :289-331. Low, S. H. 1933. Notes on the nesting of Bluebirds. Bird- Banding, 4:109-111. - . 1934. Bluebird studies at Cape Cod. Bird-Banding, 5 :39^1. Marler, P. 1955. Studies of fighting in Chaffinches (1) Behavior in relation to the social hierarchy. Brit. Jour. Anim. Behav., 3:111-117. [136] - . 1956. Behavior of the Chaffinch, Fringilla coelebs. Be¬ havior, Supplement V. E. J. Brill, Leiden, 184 pp. Marshall, J. T., Jr. 1957. Birds of Pine-Oak Woodland in south¬ ern Arizona and adjacent Mexico. Pacific Coast Avifauna, 32 :l-25. Meyerriecks, A. J. 1960. Comparative breeding behavior of four species of North American herons. Publ. Nuttall Orn. Club, 2:1-158. Monroe, B. L. 1964. Wing-flashing in the Red-backed Scrub- robin, Erythropygia zambesiayia. Auk, 81 :91-92. Morris, D. 1954. The reproductive behavior of the Zebra Finch (Paephila guttata), with special reference to pseudofemale behavior and displacement activities. Behavior, 6:271-322. - . 1955. The causation of pseudofemale and pseudomale behavior : a further comment. Behavior, 8 :46-56. - . 1956a. The feather postures of birds and the problem of the origin of social signals. Behavior, 9:75-113. - . 1956b. The function and causation of courtship cere¬ monies. Pp. 261-286 in L’instinct dons le comportement des ani- maux et de I’homme. Paris, Masson et Cie. Moynihan, M. 1955a. Remarks on the original sources of dis¬ plays. Auk, 72:240-246. - . 1955b. Types of hostile display. Auk, 72 :247-259. Musselman, T. E. 1935. Three years of Eastern Bluebird band¬ ing and study. Bird-Banding, 6:117-125. - . 1939. The effect of cold snaps upon the nesting of the Eastern Bluebird. Bird-Banding, 10:33-35. Nice, M. M. 1941. The role of territory in bird life. Amer. Midi. Nat., 26 :441-487. - 1943. Studies in the life history of the Song Spar¬ row II. Trans. Linn. Soc. New York, 6:1-329. Noble, G. K. 1939. The role of dominance in the life of birds. Auk, 56:263-273. Peterson, R. T. 1961. A field guide to western birds. 2d ed. Houghton Mifflin, Boston. Pettingill, O. S, 1936. Breeding behavior of Bluebirds. Auk, 53 :86-87. - , and S. Hoyt. 1963. Enjoying birds in Upstate New York. Laboratory of Ornithology, Ithaca, New York. Power, H. W., Ill, 1966. Biology of the Mountain Bluebird in Montana. Condor, 68:351-371. [137] Preston, F. W., and J. M. McCormick. 1948. The eyesight of the bluebird. Wilson Bulletin, 60:120. Ripley, S D. 1952. The thrushes. Postilla, No. 13, 48 pp. Ritchey, F. 1951. Dominance-subordination, and territorial relations in the common pigeon. Physiol. Zool., 24:167-176. Rowell, C. H. F. 1961. Displacement grooming in the Chaffinch. Animal Behavior, 9:38-63. Scott, J. P., and E. Fredericson. 1951. The causes of fighting in rats and mice. Physiol. Zool., 24 :273-309. Selander, R. K., and D. K. Hunter. 1960. On the functions of wing-flashing in Mockingbirds. Wilson Bulletin, 72 :341-345. Siivonen, L. 1939. Zur Oekologie und Verbreitung der Sing- drossel {Turdiis ericetorum philomelos Brehm). Ann. Zool. Soc. Bott. Fenn. Vanamo, 7:1-289. Simmons, K. E. L. 1961. Problems of head-scratching in birds. Ibis, 103a:37-49. Sindelar, C. 1967. Eastern Bluebird versus Meadow Vole. Pas¬ senger Pigeon, 29:115. Smith, W. J. 1966. Communication and relationships in the Genus Tyrannus. Publ. Nuttall Orn. Club, 6:1-250. Smith, W. P. 1937. Some Bluebird observations. Bird-Banding, 8:25-30. Sprunt, A. 1954. Florida Bird Life. Coward-McCann, New York, 527 pp. Stains, H. S. 1962. Game Biology and Game Management. A Laboratory Manual. Burgess Publishing Company, 143 pp. Stoddard, H. L. 1963. Bird habitat and fire. Proceeding Sec¬ ond Annual Tall Timbers Fire Ecology Conference, 163-175. Thomas, R. H. 1946. A study of Eastern Bluebirds in Arkansas. Wilson Bulletin, 58:143-183. Tinbergen, N. 1939. The behavior of the Snow Bunting in spring. Trans. Linn. Soc. N.Y., 5 :l-94. - . 1940. Die Ubersprungbe wegung. Zeits f. Tierpsychol., 4:1-40. - . 1951. The Study of Instinct. Clarendon Press, Oxford. - . 1952. “ Derived ” activities ; their causation, biological significance, origin and emancipation during evolution. The Quart. Rev. Biol., 27:1-32. - . 1957. The functions of territory. Bird Study, 4:14-27. [138] - . 1959. Comparative studies of the behavior of gulls (Laridae) : a progress report. Behavior, 15 :l-70. Todd, W. E. 1940. Birds of Western Pennsylvania. Univer¬ sity of Pittsburgh Press, 710 pp. Wetherbee, K. B. 1933. Eastern Bluebirds in juvenal plumage feed young of second brood. Bird-Banding, 4:199-200. Whitaker, L. M. 1957. A resume of anting, with particular reference to a captive Orchard Oriole. Wilson Bulletin, 69 : 195-262. Young, H. 1951. Territorial behavior in the Eastern Robin. Proc. Linn. Soc. N.Y., Nos. 58-62:1-37. [139] ' >he *w^tsin*| jt ; (a^ ii4iimol(« knw*MhiJw^^ />-'V 7 A .J osifjslixiv attjiig37 J fmSo^-iriiwliw3lii4«^|;aife !i4 , am/o’ ^ej^dof. li. k:: ici’D, v.tn|5s^4e lll‘Md<^ i # 71 si(fOT^«^u wV Hto ^^i’fjrti^ jsjtesw^ 2y<»1 tC. Hfi; JU. T*f ^bftms ^1 li^4^jk'hij(!QHli^£|r)n^ hfr^t. ^ . ii»u, ^ ' ; ^.■' ^ ‘ ; C I'a^iiVvft elu<^Tr$^. BN Uk Covv*raWftfcvvVN<^' ■ ■ ; vp^5j7j,p, ^ -■ ■ ^ - ,. ^ ■; %iAim, }L S.i Came iJiblbop., %jwrCi»w jV i^uY^jt. W Vri;.;gt 3Lr, iVriS. jttftlniild and iU;*^. -’l(*l^#)caCi\ff 1^ IpioeJbKf Cwto'rb^ ^)u)nm«i W -A it«1nbw«BR, N. tm Tht tH -^yii l^.. Will jn>^"UU4pui^ 2<^v471[«f«)(c^ itMn^ ' ;■' .. 5a "-Vji'.iik h>5 1 . 8ftt^ of-fflci^. Qisn4!!^ iT^xtOrd, ^ Ty»ir^«iM4kirMli^ii ■liiJ. ' IJ. j. jL.,..! ^ ™ , ljte2.' i)'eri<*e Other Information This bulletin is based primarily on the small mam¬ mal collecting program, but T have also attempted to assemble information on the occurrence of the other land mammals and the marine mammals. For this region of the State, there is a rather extensive and scattered natural history literature going back many years; mammal speci¬ mens and records are located in various museums and private collections. Long Island has been well endowed with many resident and visiting naturalists, collectors, and students. A more intensive search of these sources would reveal additional information and records of inter¬ est. However, it seems worthwhile to summarize the pub¬ lished and unpublished information now available to me on the status of the various mammals, as well as our own findings in the field. The most important published accounts of Long Island mammals in general are by Helme (1902) and Hamilton (1949). Publications dealing exclusively with the bats of this area are by Murphy and Nichols (1913) and Nichols and Nichols (1934). A recent publication of the Nature Conservancy (Anon., 1968a) has interesting firsthand accounts of the larger land mammals. Other Long Island naturalists who have considerable knowledge of the subject and who have made collections of mammals include William T. Helmuth of East Hampton (deceased) , Roy Latham of Orient, and Gilbert S. Raynor of Manor- ville. The accounts of species in the present paper deal mainly with the native and naturalized wild mammals which now occur on Long Island and in its adjacent waters, or for which there are at least occasional recent records. Available records of interest concerning mam¬ mals on outlying islands such as Gardiners Island, Fishers Island, and Staten Island, have been worked into the species accounts. My information on the status of most species in these peripheral areas is less complete than for Long Island proper. Additional species, such as some which have been extirpated on the island, introduced forms of temporary or local occurrence, and certain small mammals of the nearby mainland which have not been found on Long Island, are discussed briefly under “Other Mammals” following the species accounts. Extinct species, such as the mastodon, are not discussed. Maps of eastern United States and North America in several books which show ranges of species of mammals by shading are deceptive as far as Long Island is con¬ cerned and are puzzling to many people. Such maps may be large enough to show Long Island as a discrete area, yet the island may be incorrectly shaded in or left blank for certain species. This probably happens from lack of local information, through oversight, or because the area seems insignificant in size. Most of the mammals living on Long Island at the present time probably were on hand to greet the first white settlers from Europe in the 1600’s. Denton (1670), for example, lists muskrats, foxes, raccoons, skunks, ot¬ ters; and deer among the “wilde beasts” of Long Island. At several Indian archaeological sites on Long Island, with radiocarbon dates ranging between 1043 B.C. and 763 B.C., bones of woodchuck, gray fox, raccoon, mink, white-tailed deer, and various small rodents have been found, and at later prehistoric sites, muskrat, raccoon, whale, and deer have been reported (Ritchie, 1969). Roy Latham (personal communication) has records from about 100 excavated Indian camps and village sites from Montauk and Orient to Riverhead. At these sites he found the same species mentioned by Ritchie, and also discovered the remains of cottontail {Sylvilagm sp.), Eastern chipmunk, gray squirrel, flying squirrel (River- head only), red fox, weasel (Mustela sp.), striped skunk, and harbor seal. This material was identified by Roy Latham and authorities at the United States National Museum, Washington, D.C., and other museums. Also small bones of mice, moles, and bats were collected but no attempt was made to name them. Evidently only a very few of the established species, as mentioned elsewhere, have arrived since the time of the first Europeans settlers. Marine Mammals The accounts of marine mammals are based mainly on a brief survey of the literature and museum collec¬ tions, and correspondence with a few observers. But it is hoped that all species known from Long Island and its offshore waters have been included. Cetaceans travel wide¬ ly, and additional species may eventually be recorded from the area. Stranding records include species which regularly frequent coastal waters and those which nor¬ mally live farther out. Some strand alive for one reason or another, but certain species seem more careful in avoiding this; some float in after death or disablement, 3 but many species are likely to sink when dead. All strand- ings are considered local records, but I have indicated, when known, if the animals were actually alive when found on the shore. There are no separate species accounts for those marine mammals which have been reported only north or south of Long Island, but some which approach the area quite closely are mentioned under accounts of related species. A few species for which I have no firm Long Island records, but which have been recorded nearby to the north and south of the island, or well offshore, have been included as complete accounts; for these, the Long Island coast is considered to be at least on the edge of their range or occasionally visited. Marine mammals in general; i.e., all the large whales, dolphins, porpoises, and the seals, were very numerous originally, but have been greatly decimated in this area and occur at only a very small fraction of the population present before the heavy settlement of the coast and the era of whaling. Several of the small, little-known whales, however, have never been common. As far as I know, there has not been a general report on the cetaceans of Long Island since the brief statements on a few species by Miller (1899) and Helme (1902). Checklist The species accounts, as listed in the Contents, may be considered a checklist of known present-day native and naturalized mammals of Long Island, with certain qualifications in the case of the marine species. This list totals 59 species. Of these, 35 are land mammals, col¬ lected on this survey or examined as museum specimens except for gray fox and short-tailed weasel (published records exist for these). Marine species total 24, and at least 14 of these are represented in museums by Long Island specimens (strandings or killed near shore). I was unable to locate specimens of the other 10, as follows — killer whale, little piked whale, blue whale, and hump¬ back whale, for which there are published records of occurrence; white whale, with one probable sight rec¬ ord, not previously published; gray grampus, sei whale, hooded seal, harp seal, and gray seal, species which have been found both north and south of this area, and also within a very few miles of our coast or directly offshore (details given in text). The author would be very pleased to receive additional records or information concerning land and marine mammals in the Long Island area. Little attention was given to the question of sub¬ species, and most of the subspecies names used are merely those of the forms which would, on geographical grounds, be expected to occur; in some especially doubtful cases no subspecific name is given. Accounts of various species have been handled some¬ what differently; for example, information on the small mammals we collected is arranged under subheadings for convenience, and the marine mammals have very brief, general descriptive infonnation for the benefit of readers unfamiliar with these animals. DESCRIPTION OF REGION Location Long Island, approximately 118 miles long and 12 to 20 miles wide for most of its length, is the largest island adjacent to the eastern coast of the United States. A coastal-plain extension of New York State south of Connecticut, Long Island is surrounded by salt water, primarily the Atlantic Ocean and Long Island Sound. At its western end it is narrowly separated from other por¬ tions of New York: from the Bronx mainland and Man¬ hattan by the East River, a tidal channel or tidal strait partly occupying a passage cut in rock, and no more than one-half mile wide in places, and from Staten Island by The Narrows, about 1 mile wide. New Jersey is about 2 miles away across the waters of Upper New York Bay. The principal political units of Long Island are the coun¬ ties of Nassau and Suffolk, and the New York City boroughs of Brooklyn and Queens. Suffolk County, the largest unit, also includes various islands off the east end of Long Island; the most remote is Fishers Island, which is closer to Rhode Island and Connecticut than to Long Island. Staten Island (borough of Richmond), the least developed part of New York City, together with Long Island and its small outlying islands, make up the coastal- plain portion of New York State. Topography and Geology Long Island is mostly flat and elevations axe low. High Hill, south of Huntington, about 420 feet above sea level, is the highest point. Aside from a few small out¬ crops of ancient bedrock near the East River in Queens, the island is composed of unconsolidated materials. The basal layers are Cretaceous in origin, but these are cov¬ ered nearly everywhere by glacial deposits of gravel, sand, clay, boulders, and till. Long Island was built primarily by the glacier; that is, without the extensive deposits left by the ice, the area above sea level today would be much smaller. Long Island and Staten Island mark the southern limit on the coast of the Wisconsin stage, the most recent advance of the Pleistocene ice. Two distinct terminal 5 moraines (accumulations of debris dropped by the ice) were formed at the melting edges of the halted ice mar¬ gins during two different substages of the Wisconsin. These moraines, existing as two ranges of low hills, are prominent topographic features of Lon^ Island. The older moraine runs through the center of the island and then east along the south fork to Montauk Point and islands beyond. Later, another moraine was developed. On western Long Island this moraine overlaps the earlier one, but eastward the ice did not reach as far south and the moraine forms most of the hilly north shore and ex¬ tends on to Fishers Island and Cape Cod. Much of Long Island’s surface consists of outwash sediments which form extensive, flat, and often sandy plains south of the moraines. Lesser, but striking, re¬ minders of the glacier are the occasional large boulders, or erratics, some as big as a house, which were carried down from farther north, and the numerous kettle holes (now ponds or hollows) which formed where detached blocks of ice, partly or completely buried, finally melted away. The sea level fluctuated during the Pleistocene. Ma¬ rine waters covered most of the Long Island area prior to the Wisconsin advance. Later, during the Wisconsin glacial stage of low sea level (within the last 25,000 years), much of the present continental shelf was evi¬ dently a broad emerged coastal plain; teeth of mastodons {Mammut americaiium) , mammoths {Mammuthus sp.), and remains of other large Pleistocene mammals which lived on the plain have been dredged up by fishermen south of Long Island (off New Jersey), and east of Long Island (on the Georges Banks) (Whitmore et al., 1967). With the melting of the ice, the sea level rose again. The Long Island area was probably freed of ice by about 15,000 years ago. The gradual post-glacial submergence of the coastline, which followed, formed Long Island Sound, New York Bay, and Raritan Bay, and also flooded the narrow valley of the East River (Schuberth, 1968). The approximate present outline of Long Island thus was established. Besides land erosion by streams and wind in post¬ glacial times, the shape of the coastline has been greatly modified by the actions of ocean waves, shore drifting, and tidal currents; this has caused, for example, much land loss at Montauk, and the buildup of miles of bar¬ rier beach islands along the south shore. The construction of these low, sandy beaches and the growth of salt marshes behind them have added many square miles to the island in Recent times, although probably more than counterbalanced by land lost to erosion and subsidence (Fuller, 1914). Where higher ground meets the coast, as on the north shore, the shores between the forks, and in the Montauk region, bluffs and rocky beaches face the water. Plant life covered Long Island following the melt¬ ing of the ice, the species composition changing as the climate ameliorated (temperate conditions may have ar¬ rived about 9,000 years ago) . Mammals could have reached the island in various ways, such as by swimming, flying (bats), being transported on floating debris or by man, and walking across seasonally frozen bodies of water and early land connections. A broad expanse of the coastal plain now under water probably remained exposed for a while after the ice front receded, and this would have permitted rapid invasion of the Long Island area. But the rising sea level eventually flooded the shelf and finally severed all land connections between the island and the mainland. Surface soils of the island, which are well to exces¬ sively drained, and tend to be acid, vary from gravels to sands to silt loams. Although the climate (temperature, precipitation) is generally quite favorable, about one- half of the soils are so sandy and porous and of such low water-holding capacity that they are nonagricultural (Cline, 1955; Free et al, 1957). The extensive pine-oak barrens and abandoned fields of central and southern Suffolk County are on such soils. The more fertile areas are on the western end (now under intensive urban pressure) and along the north shore. The production of potatoes is an important industry on soils with good water-holding capacity in northeastern Suffolk County, and locally elsewhere. There are no large streams on Long Island. Much of the rain water sinks through the deep, porous ground materials to form ground water. The Peconic River, flow¬ ing east into the depression between the north and south forks, is the largest stream. Ponds and swamps are rather numerous, however. The coast is indented with many tidal inlets, bays, and estuaries. Climate Much of the following is from Taylor (1927). Long Island has a milder climate than the remainder of New York State, and has a long growing season which aver¬ ages between 180 and 210 days in length. The climate in general resembles that of the nearby mainland, but the ocean has a moderating influence here, reducing to some extent the extremes of summer heat and winter cold. The climate differs somewhat at the two ends of the island. Spring is later on the eastern end, where the in¬ fluence of the cold ocean water retards the growth of vegetation (leafing of tret's, etc.), and this difference, 6 which may be as much as 2 weeks, is very noticeable if one travels from Brooklyn to Montauk at this season. There is, however, a long frost-free season on the east end, since killing frosts rarely occur late in spring, and the moderating effect of the ocean also tends to delay fall frosts. Central Long Island occasionally experiences very hot periods in summer; the heat may be especially op¬ pressive in the pine barrens where the open canopy lets down much more sunlight than other forest types. The average annual precipitation is 40-50 inches over most of the island, and somewhat less than 40 inches in the Napeague-Montauk region. Snowfall averages about 20-40 inches, less than mainland New York, and a snow cover rarely lasts because of the relatively mild climate (winter of 1960-61 unusually severe). There is a good deal of rain during the winter. Summer months are the driest, the lack of rain producing occasional droughts destructive to plants on the highly porous soils, especially when combined with high temperatures and winds. Long Island is a windy place, and where there are no obstructions, as at Montauk, Shinnecock Hills, and along the outer barrier beaches, wind has a pronounced local effect on the vegetation. In such areas during winter the wind makes it feel much colder than the thermometer indicates. Weather Bureau records show that Montauk Point is one of the windiest spots along the Atlantic Coast. In summer, regular, cooling “sea breezes” make eastern Long Island and the outer beaches pleasant. Severe coastal storms such as hurricanes (usually in late summer) and northeasters (usually in winter), occa¬ sionally batter the coast. At times they break through the outer beach strip to create new tidal inlets, which may increase the salinity and affect the life of the bays. The inlets may become sealed off again naturally or shift from east to west; man has worked to keep some of them open for boating by building jetties and dredging. Some of our field work was conducted in the vicinity of Shin¬ necock Inlet, which was created during the exceptionally destructive hurricane of September 21, 1938, and Mor¬ iches Inlet, which was opened during a storm in March 1931. Helmuth (1954) wrote a fascinating account, based on his personal field observations, of the great hurricane of 1938 on eastern Long Island and its effect on the bird and small mammal populations of the area. Western Atlantic coastal waters between far northern and tropical seas may be considered to belong to three main zones or faunal provinces, significant in the distri¬ bution of many marine invertebrates and vertebrates. There is a cold-water zone north of Cape Cod and a warm- water zone south of Cape Hatteras; Long Island is in the northern part of the temperate-water zone between the two capes. Vegetation The vegetation of Long Island has been profoundly affected by the activities of modem man, especially on the urbanized western end, where very little remains of the original flora. Indians had been living on Long Island for many centuries prior to the arrival of the white men. Undoubtedly the former had some effect on the vegeta¬ tion through their deliberate burning of the woods, help¬ ful in hunting game, and their clearing of the ground to plant corn and other crops. But the big change began in the 1600’s when the whites replaced the aboriginal in¬ habitants. The first Dutch farm was established in what is now Brooklyn in 1625, and English settlers, by way of Connecticut, arrived on eastern Long Island in 1640. The English spread west rather rapidly along the coasts of Long Island, while the Dutch for a time controlled the western portion. By 1670 Denton was able to write: “Long Island ... is inhabited from one end to the other. On the west end is four or five Dutch towns, the rest being all Eng¬ lish to the number of twelve, besides villages and farm houses.” He also wrote of meadows and hayfields which supported a plentiful supply of cattle and other livestock. As for the Indians, Denton stated that in his own time they were reduced from six towns to two small villages, although the survivors still went hunting and fishing and raised crops of com. Svenson (1936) discussed the early vegetation of Long Island, quoting extensively from Denton (1670) and other writers of long ago. In early Colonial days there were tall and impressive stands of timber, espe¬ cially on the western end and along the north shore; tall red oak, white oak, hickory, black walnut, American chestnut, and beech abounded. Much of the valuable timber was removed very early. Long Island was praised by the settlers for its wild fruits, an abundance of straw¬ berries, blueberries, huckleberries, cranberries, grapes, and beach plums; the wild strawberries were large and eagerly sought. The vast pine and oak barrens, typical of central and southern Long Island today, were mentioned by early travelers. Apparently much of this forest became de¬ graded, however, by heavy cutting and ever more fre¬ quent fires. Swamps of southern white cedar formerly were much more widespread than 'at present, and typi¬ cally bordered the heads of tidal streams all along the south shore. The Hempstead Plains, unusual in being considered a true natural prairie in the East, originally comprised an area 16 miles long, covered with tall grass. Denton (1670) stated that it was being mowed for hay, and used to graze sheep and other livestock and for rac- ing horses. A sizable portion of the Plains remained until a few decades ago, but now only a small remnant has survived the spread of suburban housing. According to Taylor (1923), early Colonial documents described the hilly coastal grasslands still existing in the windy Mon- tauk area, known as the Montauk Downs. The grassy, rolling Shinnecock Hills apparently are more stabilized than formerly; older writings refer to a lack of vegeta¬ tion and drifting sand in this area. The woods of Gardi¬ ners Island reportedly grew the largest trees in all of the Long Island area up until the destructive hurricanes of 1938 and 1944. According to Taylor (1923), woods in a relatively sheltered area near Montauk Point had trees as large as those of Gardiners Island before a severe storm in 1815. The barrier beach and salt marsh habitats, once relatively inaccessible, are endangered by steadily increasing recreational use of the entire south shore. The climate and elevation are fairly uniform throughout Long Island and small differences are of minor importance in the distribution of plants. Under¬ lying geological materials, soil types, and moisture are significant. The pine barrens, for example, grow largely on the sandy, gravelly outwash plains, while a mixed deciduous forest covers richer, moister sites on the north shore moraine. Brodo (1968) points out that there is a strong correlation between soil types and vascular vege¬ tation on the island. Fires play an important role in the barrens, favoring fire-resistant pitch pine and scrub (bear) oak; where fires are especially frequent the pines and oaks are low and shrubby. Brodo (1968) described the vegetation types of Long Island, and referred to the many botanical papers deal¬ ing with local areas and plant associations. Peters (1949) gave an account of the dominant types of vegetation. A brief and sketchy classification, based on these and other references and my own field work is given below. All of the 14 major categories listed were searched for small mammals except for the nearly vanished Hempstead Plains; some of the study areas were in miscellaneous or edge habitats not resembling any of the major types. Habitats where the various mammals were found are mentioned in the species accounts, and some of the col¬ lecting areas are shown in the photographs (figures 1 to 26). Mixed deciduous forests. The forest of the richer soils of western Long Island (now surviving only in parks and other small areas) and also on the north shore (in estates and woodlots). Large trees, growing especially tall in the western areas — black oak (Quercus velutina) , white oak {Q. alba), red oak {Q. rubra), beech (Fagus grandifolia) , tuliptree (Liriodendron tulipifera) , black birch {Betula lenta) , hickory [Cary a spp.) , and others. 7 Oak forests. These occupy extensive areas on drier soils in central Long Island and on the south fork (both west and east of the pine barrens, and elsewhere). Scar¬ let oak (Q. coccinea), black oak, white oak, and other oaks. Pine barrens {or Pine-oak barrens). The typical pine barrens on Long Island consist of nearly pure stands of pitch pine {Pinus rigida) , with an extensive undergrowth of scrub oaks, especially the species known as bear or barren oak {Quercus illicifolia). Also blueberries, huckle¬ berries, and the low ground cover, bearberry {Arctosta- phylos uva-ursi) may be found. This habitat is especially characteristic of dry, very well-drained soils, and high incidence of fire. Areas where all the vegetation is low and scrubby are sometimes called “pine plains” or “oak brush plains.” The pine region extends from eastern Nassau County east to Riverhead and Hampton Bays. Pine-oak forest. Similar to the pine barrens, but more mature, with older trees and a large proportion of tree¬ sized oaks, such as white oak and scarlet oak. Pitch pine, the dominant conifer of Long Island, and the various scrub and tree oaks, the dominant deciduous trees, also are found growing in all sorts of complex combinations. The predominantly oak woods in turn merge with the mixed deciduous forests. Other forests. Of interest are certain woodlands near the coast of eastern Long Island and on the outer barrier beach, which are relatively small in extent but apparently long-established and unlike woods elsewhere. Most publicized is the Sunken Forest, where holly {Ilex opaca), tupelo {Nyssa sylvatica), sassafras {Sassafras al- bidum), shadbush {Amelanchier) , and other species form a deeply shaded woodland sheltered behind the inner dunes on Fire Island. In Hither Hills there is a wood¬ land (“Hither Woods”) of old oak trees somewhat dwarfed by the wind and formerly festooned with lichens. A stretch of woods near Montauk Point (“Point Woods”) contains a beautiful stand of beech, oaks, holly, red maple, etc. Other rich woods with large beech and oaks on east¬ ern Long Island are on Shelter Island and Gardiners Island. On the survey, we collected in the woods of Hith¬ er Hills, and to a lesser extent in the woods of Montauk and Shelter Island, and only visited the Sunken Forest and Gardiners Island. Southern white cedar swamps. Coast or southern white cedar {Chamaecyparis thyoides) , growing closely together, is the dominant tree. Ground cover consists of sphagnum moss. Now of very limited distribution on Long Island, cedar swamps are found mainly near ponds and streams in the eastern part of the pine barren region. Red maple swamps. This is the most frequent type of swampy woods; common along streams throughout the island. Red maple (Acer rubrum) is numerous, and tupelo is often present. Sphagnum hogs. Small areas of leatherleaf (Cha- maedaphne calyculata) , sedges, sphagnum, and other bog plants are found in wet areas of the pine barrens and eastern Long Island. Chamaecyparis may be present as scattered trees or along the edge. Abandoned cranberry bogs also are present locally. Fresh marshes. Fresh-water marshes, with grasses, sedges, cattails, and the like, are found mainly along south shore streams, around ponds, and in kettle holes. Hempstead plains. This natural prairie land, formerly extending across much of central Nassau County, is (or at least was) dominated by a species of beardgrass (An- dropogon scoparius) . Downs (grassy coastal uplands). The Montauk Downs, described in detail by Taylor (1923), are grassy, rolling hills with occasional islands of shrubs and a very few widely scattered trees. Here the wind seems to be principally responsible for preventing or greatly slowing down the development of trees. The Shinnccock Hills are somewhat similar and are exposed to winds blowing across the ocean and Great Peconic Bay. The vegetation con¬ sists of Andropogon spp. and other grasses, with clumps of bayberry (Myrica pensylvanica) , beach plum (Prunus maritima), etc., and some scattered trees (pitch pine, red cedar) . Dunes (grass and thickets). Sand dunes are the backbone of the outer beach strip, and also occur in some other areas. Probably the largest dunes on Long Island are the spectacular moving or “walking” dunes on the east side of Napeague Harbor. Desert-like conditions prevail on the dry sandy dunes; beachgrass (Ammophila breviligulata) is the most vigorous dune plant in exposed situations, while certain shrubs (bayberry, beach plum) are frequent. Along the ocean, primary or outer dunes lie just behind the beach, while a bit farther back are the more stable secondary or inner dunes. In the inter¬ dune valleys, and in depressions and hollows, may be found moist, protected situations where the vegetation is relatively dense (and small mammals numerous). Salt marshes. Located primarily in the bays along the south shore, these marshes are still fairly widespread. They are covered periodically by salt water during very high tides, and contain certain plants, such as salt-mead¬ ow grass (Spartina patens) and marsh-elder (Iva fru- tescens) , not found in other habitats. The extensive Napeague Meadows, situated between Napeague Harbor and the ocean dunes, does not seem to fit any of the above categories exactly. Quite a few small mammals were collected in this flat, open, mari¬ time area containing both moist meadow (fresh to salt) and dry sandy ground. Abandoned fields. Dry, sandy, abandoned farm fields with grasses (especially Andropogon virginicus) and various weeds, are frequent on the poor soils of central Long Island and were covered in this survey. ACCOUNTS OF SPECIES Opossum Didelphis marsupialis virginiana Kerr The opossum is generally common on Long Island, but apparently it has been a resident here only during the past 100 years or so. It became established late in the 1800’s, probably as the result of repeated accidental introductions and releases. Seemingly contrary to the prevailing evidence is a statement by Denton (1670) naming the opossum as a food of Long Island Indians; however, Denton did not include this marsupial in a list of Long Island mammals presented elsewhere in his pub¬ lication. Audubon and Bachman (1851) predicted that the opossum would in time become established on Long Island, and in other areas east of the Hudson River, be¬ cause the living animals were constantly being carried there. Helme (1902) mentioned accumulating reports of opossums in the early 1880’s and stated that in a few years they became common with a wide distribution the full length of the island. Opossums were quite common at Montauk in 1893, but had become common there only a few years earlier (Dutcher and Dutcher, 1893). By the late 1880’s several towns were paying bounties for the destruction of opossums, but this seemed to have no ap¬ preciable effect on their abundance, even though thou¬ sands were reported killed in some years. Nowadays, this is one of the mammals frequently found dead along many of the highways on Long Island, indicating a high population present. The opossum seems to thrive in the more thickly settled areas, and is even found within the limits of New York City. In our field work, this species was frequently noted along the south shore, throughout much of the south fork, in the central pine-oak region, and in some north shore areas. We regu¬ larly saw opossums in sandy and marsh-edge habitats on the outer barrier beach between Moriches Inlet and Shin- necock Inlet. However, in comparing today’s abundance with that suggested in the early reports, and judging from the rec¬ ords of experienced observers extending over many years, there are fewer opossums in some sections of the island at present than during their first decades of explosive in¬ crease. Roy Latham (personal communication) writes that opossums were common in Orient and throughout the north fork from 1900 to 1930, but have become scarce within the past 10 years, while red foxes and rac¬ coons have increased dramatically since 1930. Our field work in 1963, and conversations with local residents, in¬ dicated opossums to be quite rare on Shelter Island; opossums were formerly common there, according to Latham, and he has records for Shelter Island up to 1943. The opossum seems to have had a checkered career on Staten Island, including a marked increase late in the past century. Masked Shrew Sorex cinereus cinereus Kerr Distribution and habitat. The masked or common shrew is rarely seen and is not as familiar to Long Island residents as the eastern mole, short- tailed shrew (also often called “mole”), and the various kinds of mice. Nevertheless, it is quite possible that this is the most nu¬ merous mammal on Long Island. Trapping results indi¬ cated this to be the case at least for the less developed areas of Suffolk County. Sorex was outnumbered in our total Long Island catch by Blarina, Peromyscus, and Mi- crotus, but these last three are more easily trapped than this tiny Sorex, which, when it does encounter a trap may often escape untouched or fail to spring it. Furthermore, on Long Island this species is found in a great variety of environments, as discussed below. This is the smallest mammal found here; in New York State only the pigmy shrew (Microsorex) is smaller, and it is not known to oc¬ cur on Long Island. Trapping results indicated the presence of this species in just about every habitat on Long Island with sufficient ground cover, including both damp and dry situations, and areas with or without woody growth. Grasslands of every description, except where the grass was too sparse, were especially productive: over half of the specimens were taken in grass. Such habitats included abandoned fields, the Shinnecock Hills, fresh marshes (along streams, etc.), salt and brackish marshes, moist and dry grassland at Montauk Point, and dry sandy places near the shore (such as depressions among the dunes, meadows at Na- peague, etc.). In these habitats various shrubs, sedges, and miscellaneous herbs were often present, but grass of MAP 3. LONG ISLAND, SHOWING MOST OF THE 12 one kind or another was the dominant feature. This shrew and Microtus were frequently found together, the former often traveling the runways of the latter. This little shrew occurs all along the outer barrier beach of the south shore, such as Jones Beach, Fire Is¬ land, and between Moriches and Shinnecock inlets. Indi¬ viduals have been found under old logs and other drift¬ wood on the upper beaches of both the north and south shores and at Montauk Point. Sorex cinereus was also common in the woods. Oak woods, mixed deciduous woods, pitch pine barrens, swampy woods of red maple and other trees, southern white cedar swamps, and both dry and damp coastal woods near Montauk Point, all yielded their quotas of this shrew. Probably it occurs in the Sunken Forest on Fire Island, although we did not trap that part of Fire Island. In the woods the shrews were trapped under the leaf litter and fallen branches, about stumps and logs, in mole tunnels, and in mossy cavities in maple swamps and at the base of cedar trees (figure 8). Hamilton (1949) wrote of trapping many in woods at Roslyn (north shore of Nassau County), and they are still found in wooded parks in eastern Queens. Sphagnum bogs were also good for finding Sorex (figure 9). Here, leatherleaf, other shrubs, and sedges grow above the thick sphagnum carpeting the ground; the shrews were found in runways and cavities on and under the moss. In several of the cool cedar swamps and sphagnum bogs Sorex cinereus was the only species of smalb mammal collected. These habitats are very reminis¬ cent of areas in the New Jersey pine barrens, many of the same plants being found in both regions. But the New Jersey region has a richer mammal fauna in that the cedar swamps and sphagnum bogs there are inhabited also by tbe red squirrel (Tamiascurus) , red-backed mouse (Clethrionomys) , bog lemming (Synaptomys) , and meadow jumping mouse (Zapus) which do not occur on Long Island, exc^t for the jumping mouse, which is scarce. Sorex cinereus is present on Staten Island and Shel¬ ter Island. I do not know of any records of this shrew on Gardiners Island, Plum Island, or Fishers Island, but would not be surprised if it was found on these islands off the east end of Long Island. Reproduction. Masked shrews were found to pro¬ duce young on Long Island for at least half the year, from April through October. On April 27, one female examined was carrying seven embryos, while five other females in April were already nursing litters — April 18 (1), April 26 (3), and April 27 (1). Reproduction continued through the summer and in October as well, although at a reduced rate since there was evidence of fewer pregnan¬ cies after June. On October 14, a female carried six quite small (about 2.5 mm.) embryos, and another one on October 20 was lactating. Litter size based on eight em¬ bryo counts averaged 5.9 (range four to seven) ; embryo size ranged from about 1 mm. (diameter of swellings) to 11 mm. (crown to rump length). Adult males in evident breeding condition (reproductive structures markedly en¬ larged) were collected from March 18 to October 14. The first juveniles out of the nest were trapped June 7 and 8. Measurements. Forty adult males, collected from March to October at various localities in Suffolk County, and all in apparent breeding condition, average: weight, 4.7 grams (3.4-6.4) ; total length, 99.3 mm. (92-105); tail, 38.7 mm. (35-42) ; hind foot, 12.0 mm. (11-12.5). Females are closely similar in size and 30 adult females (eight with embryos) from March to November (same localities) average: 4.8 grams (3.2-7.9) ; total length, 99.2 mm. (92-105); tail, 38.2 mm. (33-41); hind foot, 11.9 mm. (10-12.5). If the eight pregnant females are exclud¬ ed, the 22 remaining adult females weigh slightly less, averaging 4.5 grams (3. 2-6.0). Individuals taken. 216 Short-tailed Shrew Blarina brevicauda talpoides (Gapper) Distribution and habitat. On Long Island, Blarina is distributed from Brooklyn to Montauk Point and on some of the offshore islands. This large shrew is generally abundant in many different habitats throughout its range, which blankets much of eastern North America. On the island we found Blarina most numerous in woods with deciduous trees (133 taken), such as oak or mixed de¬ ciduous woods, oaks with pines, and red maple swamps; and in inland fields (91 taken). Only a relatively small number (26), were taken in all other habitats, that is, seven or less were taken in each of the following: pine barrens, southern white cedar swamps, sphagnum bogs, fresh marshes, salt marshes, and coastal dunes and grasslands. Several extensive traplines in these habitats did not yield a single specimen, and also we failed to find any at our outer barrier beach localities. This was quite different from our experience with Sorex cinereus. Especially in sandy grasslands near the shore (including the outer beaches), in salt marshes, and in sphagnum bogs, Sorex was much more numerous than Blarina. The following are some examples of this among trap returns in the period 1961-63: Napeague Meadows 13 on the south fork (traps mostly in dry sandy grassland) — 40 Sorex, no Blarina; Hither Hills (in grassy depres¬ sions near Napeague Harbor — 28 Sorex, two Blarina; various sphagnum bogs — 17 Sorex, four Blarina. Many of the traps were set in Microtus runways in these areas. On the other hand, in woods and also in many fields away from the immediate vicinity of the coast Blarina seemed to be the more numerous shrew. Blarina also was common on wooded bluffs of the north shore, over¬ looking Long Island Sound. In woods and fields, short-tailed shrews were fre¬ quently taken in close association with pine mice, in the same burrows in the humus or sandy soil. Also, mouse traps set in tunnels of the eastern mole took many of these shrews. Nineteen were caught in star-nosed mole tunnels in red maple swamps at Belmont Lake State Park, Baby¬ lon. Of course, being so numerous in general Blarina was also found living together with the other small mammal species in one area or another. Blarina was common during this survey in 1963 on Shelter Island, in both woods and fields. Richard Van Gelder (verbal communication) has trapped this species on Gardiners Island. The short-tailed shrew occurs at Orient Point and also on Fishers Island, but I do not know of any records for Plum Island. This island is a quarantined animal disease laboratory and few naturalists visit. Blarina is common on Staten Island, where the least shrew {Cryptotis parva) also occurs. Reproduction. Only five pregnant females were taken, the earliest on March 22. Embryo counts ranged from five to eight (average 6.7). Males in apparent breeding condition were collected from January 16 (1962) and March 9 through the first half of October. Measurements. Fifty adult males from various months average: weight, 17.9 grams (13.4-24.1) ; total length, 119.4 mm. (109-129); tail, 24.9 mm. (21-30); hind foot, 14.2 mm. (13-16). Twenty nonpregnant adult fe¬ males (various months) average: weight, 16.1 grams (12.9-20.9); total length, 119.5 mm. (112-128); tail, 25.4 mm. (23-28) ; hind foot, 14.3 mm. (13-15). Individuals taken. 250 Eastern Mole Scalopus aquaticus aquaticus (Linnaeus) Distribution and habitat. This is the common species of mole on Long Island, and it is also present on Staten Island. The hairy-tailed mole ( Parascalops) which is the common mole in woods and drier ground in most of “upstate” or mainland New York, has not been found in this region. The eastern mole occupies most of Long Island where the ground is not too wet. Probably it is most numerous in some of the mixed deciduous woods along the north side of the island where richer soils accommo¬ date high populations of invertebrates. However, the mole also occurs in the sandy ground throughout the pine and oak areas, and in abandoned fields, of the central and south shore portions of Suffolk County. The conspicuous ridges, marking the course of the shallow subsurface tunnels, may be seen even in exceedingly barren situa¬ tions, such as under scrubby waist-high “pine plains” vegetation south of Riverhead. The mole also invades suburban yards, where its tunnels and mounds disfigure lawns to some extent. East and west, its range extends nearly the full length of the island — from parks within New York City limits to well out on both the northern and southern peninsulas, or forks, of eastern Long Island. On the north fork a mole was collected within 3 miles of Orient Point. On the south fork we did not find the eastern mole in the immediate vicinity of Montauk Point, but it was common in the Hither Hills region a few miles from the point. In their mammal survey of the Montauk re¬ gion before the turn of the century. Butcher and Butcher (1893) wrote: “No moles of any kind were observed east of Napeague Beach, although careful search was made, nor did we hear of any mole ridges ever being seen on Montauk. Napeague seems to them to be an im¬ penetrable barrier.” In the present survey (early 1960’s) we found mole tunnels numerous at Hither Hills, which is east of the narrow neck at Napeague. Perhaps Scalopus is extending its range eastward in this sector. Scalopus is present on Shelter Island, according to Roy Latham (personal communication). Moles have not been collected or reported on Fire Island, Gardiners Island, Plum Island, or Fishers Island, as far as I know. It seems likely that moles are absent from some if not all of these islands. This mole frequently digs through pure sand in the vicinity of tidal water. The tunnel ridges may be seen in sandy ground along the very edge of salt marshes or meandering across dunes near the ocean. Such areas seem insufficiently provided with food, and are probably not permanently occupied by the moles, but the tunnels in these situations are conspicuous. Near Moriches Bay (in April) I noted a mole tunnel in a rather high, sandy bank bordering a salt marsh stream. Long, raised mole tunnels across the surface of the smooth sand are fre- 14 quent in the moving dunes area at Hither Hills State Park. At nearby Napeague Harbor in June, mole tunnels were seen which traversed the rolling surface of beach dunes and wandered over the level upper beach. Near East Hampton I trapped a mole during December in a sand tunnel on inner dunes about 800 feet from the ocean; cover consisted of bare sand alternating with patches of woody hudsonia {Hudsonia tomentosa) and lichens. A remarkable spot to see a mole tunnel was on a sandbar about 1,000 feet long separating Peconic Bay from a pond and salt marsh. Here (in March) a mole had tunneled for hundreds of feet through low sand dunes, with beach-plum (Prunus maritima) and beach- grass (Ammophila) for scattered cover. The mole finally stopped upon encountering an outlet stream which crossed the beach. Some of the tunnels, such as those at Napeague Harbor, were in locations which would occa¬ sionally be covered by salt water during storms or exceptionally high tides. But 1 did not see Scalopus tun¬ nels in the true intertidal zone of beaches, such as have been reported by McCully (1967) for the California mole {Scapanus latimanus) . Fresh swamps and meadows on the island are mostly without moles, since the semi-aquatic Condylura is extremely local here, and the present species generally avoids water. Occasionally though, tunnels of Scalopus penetrate a short distance into the damp soil of creek banks and wet woods. In the shrubby grassland of Shinnecock Hills, moles seemed scarce; tunnels were noted only in a few depressions among the hills. On the grassy, hilly Montauk Downs I did not find any evidence of mole activity. Moles seemed to be uncommon in some cultivated areas, such as farmlands on the north fork. In this sec¬ tion moles were found primarily in the lighter soils of hilly, wooded areas, such as Browns Hills, where they are common. Roy Latham (personal communication) ex¬ plains further: ''Scalopus is restricted to the hills in Orient and dry soil on the north fork. I never saw it in the heavy farm land that is cultivated. It is uncommon in Greenport and Orient. Common in most sections of eastern Long Island.” In Browns Hills, in June 1963, in a woodland of hickory, hackberry, and other deciduous trees, with an undergrowth of rank weeds, the loose, fertile soil teemed with earthworms, insects, and other invertebrates, and was riddled with mole tunnels (one Scalopus was collected). Specimens were trapped in the different habitats men¬ tioned above, such as various types of deciduous woods, pine barrens, fields, suburban yards, and sand dunes. Most specimens were caught with ordinary mouse snap traps placed crossways in the tunnels, level with the lower surface, and the opening above covered over with card¬ board, although they are harder to catch this way than Parascalops or Condylura. Some were collected in each of the four seasons; in cold, midwinter weather, trapping was possible in tunnels where the ground was not frozen, such as those under deep pine-needle litter. Moles re¬ mained active in winter in tunnels near the surface as long as the ground was not deeply frozen. Also at this season (December, January) I occa¬ sionally noted in yards and gardens, large, conspicuous “mole hills,” larger and more numerous than usually made by this species. These mounds probably consisted of earth brought up from deep burrows undergoing ex¬ cavation. The ground often had a lumpy appearance in such areas, apparently the result of irregular up-and- down tunneling or shifting of earth, instead of the usual horizontal digging. In the warmer months a surge of increased activity near the surface was common when it rained following a dry period, especially in sandy areas. Short-tailed shrews and pine mice were found to travel through the mole tunnels frequently and to make holes opening up on the surface. Other species such as masked shrews, meadow mice, and white-footed mice also used the tunnels in some localities. Food habits. Eighteen stomachs were examined (five spring, four summer, six fall, three winter). In¬ sects are foremost in the diet in much of the area judg¬ ing from this small sample, being found in all the stomachs checked and comprising 68.8 percent of the total volume. Earthworms have been reported as the pri¬ mary food of this mole in many regions, but I found them in only two stomachs (6.7 percent of volume). A mole from a north shore locality where earthworms were numerous,' however, had been feeding almost entirely on these animals (June). In most other Suffolk County localities where moles were collected, such as Riverhead, Hampton Bays, etc., earthworms are not so common, and insects were first in the diet. Most important in this category were “white grubs,” the large larvae of June beetles (Scarabaeidae: Phyllophaga) , 26.1 percent of total volume and present in 10 stomachs; and ants (Formicidae) , 21.2 percent of total volume and in 11 stomachs. Over 100 ants were counted in one stomach (March). Other insects eaten included other Coleoptera (adults and larvae) and larvae of Lepidoptera and Diptera. Other invertebrates, which were present in only small numbers were slugs, sow-bugs (Isopoda), spiders, and centipedes. The only vertebrates detected as food were two red- backed salamanders, [Plethodon cinereus, lead-back color phase), in the stomach of a mole collected at Flanders 15 on March 9. The eastern mole feeds to a limited extent on plant material. Vegetation was found in eight stomachs (8.9 percent of total volume); this was chiefly pieces of an unidentified slender root. Reproduction. The eastern mole breeds early in spring, producing one litter a year. Two males collected in the first week in March were in apparent reproductive condition, with much enlarged testes and accessory struc¬ tures. These males showed pronounced orange-brown glandular stains on the chin, snout, wrists, and along the ventral midline of the body. Males from May to January all appeared to be nonbreeding. A female on May 3 showed four distinct placental scars and evidence of having very recently given birth. Another female on June 6 also showed four placental scars and was lactating. Hamilton (1943) found a nest containing four young in mid-April on Long Island. Remarks. Two adults which showed distinct evi¬ dence of molting were taken on June 6 and 13, 1963. These individuals had extensive areas of new, shorter fur coming in dorsally and ventrally, contrasting with patches of old faded fur. These animals were probably molting from winter to summer pelage (spring molt). A pure white albino mole of this species was plowed out at Calverton on July 4, 1952 (Roy Latham, personal communication ) . Measurements. Ten adult males from Suffolk County average: weight, 64.3 grams (56.5-74.8); total length, 163.3 mm. (155-172); tail, 28.6 mm. (24-32); hind foot, 19.7 mm. (19-21). Nine adult females from Suf¬ folk County average: weight, 49.9 grams (38.4-58.4); total length, 154.6 mm. (140-159); tail, 26.9 mm. (23- 30); hind foot, 18.9 mm. (18-19.5). The smallest mole collected was an adult female in mid-January which weighed only 38.4 grams, and measured 140 mm. in total length. Individuals taken. 22 Star- nosed Mole Condylura cristata cristata (Linnaeus) Distribution and habitat. The star-nosed mole is very scarce and local on Long Island, where it seems to have escaped detection by many of the active field natur¬ alists and collectors. Helme (1902) searched various parts of the island without finding any evidence of its presence. The only record he had was one he found lying dead in the street at Miller Place on the north shore, about 4 miles east of Port Jefferson; Helme thought perhaps the ani¬ mal had been dropped by a hawk which had carried it over from the Connecticut mainland. Jackson (1915) listed other New York, but no Long Island localities for this species, and he did not include Long Island in his map showing the geographic range of Condylura. Three specimens, all males, collected on western Long Island after Helme’s publication are in the American Mu¬ seum of Natural History. Two are from Baldwin in Nas¬ sau County — September 1, 1907 (A.H.H.), No. 35155, and September 28, 1908 (D. Franklin), No. 73626; the third specimen was taken north of Jamaica in Queens, May 17, 1913 (H. Boyle), No. 37271. Apparently these specimens did not come to the attention of Jackson and others. Suitable habitat probably no longer exists in those localities. There are also some published references attest¬ ing to the presence of this species on Long Island. Audu¬ bon and Bachman (1851) stated they had received speci¬ mens from a collector on Long Island. Nichols (1907) on April 18 received a star-nosed mole (caught by a cat), from Great Neck, north side of Nassau County. Turrell (1939) termed the star-nosed mole as “fairly abundant” in the Smithtown region but gave no addi¬ tional details. Probably he found the species present in this area without realizing its general scarcity on the island. The star-nosed mole is present on Staten Island, where some have been found in barred owl {Strix varia) nests and pellets (Chapin, 1908; Davis, 1908). On our survey, a very few reports were received of animals seen (or found dead) which were recognized as this species by the observer, or consisted of a convincing description of it. Reports represented both north and south shores of western Long Island — Nassau County and into Suffolk County as far east as the Connetquot River. Because Condylura is so distinctive in appearance, such reports are considered reliable. Gilbert S. Raynor and Richard Van Gelder (verbal communications) also have received a few sight reports of this species on Long Island. In and near Belmont Lake State Park, at Babylon, I collected six star-nosed moles and found numerous signs of this species. Whenever visiting meadows, wooded swamps, or streamside areas on the island which looked suitable for Condylura I examined the ground closely, but never found definite signs of its presence other than in this one locality. Favorable-looking but apparently un¬ inhabited spots were examined at Carmans River, East- port, East Hampton, East Quogue, Flanders, Forge River, Hauppauge, Middle Island, Nissequogue River, North Hills, Peconic River, and Westhampton Beach. On Great Hog Neck, on the north fork of eastern Long Island, mole tunnels were present in wet, black, swamp soil where there were many earthworms, tunnels which may have 16 been the work of this species. This seems somewhat questionable, however, since no specimens were taken and tunnelings of the eastern mole were abundant in dry soil nearby. Thus it appears that the star-nosed mole is, or was, most common towards the western end of the island; however, in this section probably most of its habitat has been destroyed by the expanding metropolitan area. Evi¬ dently its range extends eastward in a spotty manner along both the north and south shores of the island. If it occurs in eastern Suffolk County, it must be very local, since we did much field work there without finding the species. Possibly the widespread boggy, acid conditions in the wet areas, with an accompanying scarcity of earth¬ worms or other food, is unfavorable. However, Christo¬ pher McKeever has shown me an unpublished list of Long Island mammals by William T. Helmuth (consisting of marginal notes written in a copy of Hamilton, 1943) which records East Hampton as a locality for Condylura. This is far to the east of other specimens and reports, although I have not learned of any star-nosed moles ac¬ tually collected at East Hampton. The Babylon habitat was situated along a small stream, tributary to Carlls River (figures 13, 14). The moles were found inhabiting the ground at least one-fifth of a mile along the stream, in the vicinity of the west boundary of Belmont Lake State Park (on both sides of the boundary line). Most of the mole activity was found within a few yards of the creek, which was shallow, sluggish, and about 2 to 3 feet wide. The ground was wet, mucky, and black and contained many earthworms. The site was rather heavily wooded with red maple and tupelo, and many shrubs and vines such as highbush blueberry, arrow-wood, common elderberry, Smilax, etc. Skunk cabbage and cinnamon fern also were plentiful. No evidence of mole activity was detected in this area at first glance. The colony was located by probing the ground by hand until some rather deep tunnels were located. No surface ridges of pushed up soil were pres¬ ent. Eventually, small, inconspicuous “mole hills” were noticed — mounds of mucky soil pushed up from tunnels below and partly covered by fallen leaves. The larger mounds were about 2 inches high and 6 inches in dia¬ meter. These mounds apparently were made in the fall; by spring they had almost entirely disappeared, although the moles were still numerous in the area. Also, shallow tunnels of Condylura were noticed when setting traps — these were typically along the sides of logs and were merely furrows covered by leaves and other debris. From some of the tunnels vertical shafts led down to the water level a short distance below. Two individuals were trapped on December 1, 1961, and four were caught on April 24, 1963. All were taken using standard wood pedal mouse traps set down in the tunnels in spaces excavated with a knife. Other small mammals present were Blarina, Peromyscus, and Micro- tusi Blarina and Microtus frequently used the Condylura tunnels. Other wildlife along the creek included wood¬ cock, raccoon, opossum, cottontail, and gray squirrel. I have since learned that this creek area has been sched¬ uled for drainage and other “improvements,” which probably means the end of the colony of star-nosed moles. Food habits. Earthworms ranked first in the stomach contents of the six moles examined, comprising 55 percent of the total volume and occurring in five stomachs. In¬ sect larvae, including numerous craneflies (Tipulidae), were second at 26 percent of volume, present in five stomachs. Other foods included beetles, spiders, and plant material. Reproduction. All three males taken on April 24 were in apparent breeding condition, with much enlarged reproductive structures (testes 20-21 mm.). Measurements. Three April adult males average: weight, 49.4 grams (48.8-50.0) ; total length, 185.7 mm. (185-186); tail, 68.0 mm. (64-71) ; hind foot, 26.8 mm. (26-27.5). An adult female, in April, without embryos, measured: weight, 58.5 grams; total length, 185 mm.; tail, 67 mm.; hind foot, 27 mm. Individuals taken. 6 Little Brown Myotis My Otis lucifugus lucifugus (Le Conte) Distribution and habitat. At the present time the little brown appears to be the most numerous summer bat over most of Long Island. Evidently it has increased considerably in numbers since the early part of the cen¬ tury. Helme (1902) does not mention this species, except lor stating, under a brief account of the big brown bat, that a smaller unidentified brown species is of occasional but rare occurrence on Long Island. Murphy and Nichols (1913) stated that Myotis lucifugus was scarce at that time, that it rarely frequented houses or towns on the island, and that the species was not collected here until 1900, when three were shot from a flock of a dozen or more at Cold Spring Harbor. The little brown myotis was known to be common in many mainland regions of New York State before 1900 — such as the Adirondacks, Catskills, and Oneida 17 Lake region, and to form large colonies in buildings, at least locally, as summarized by Miller (1899). DeKay (1842) in his account of the little brown bat, which probably refers to Myotis in general, but this species pri¬ marily, stated that it was “very numerous ... in the southern counties” (of New York). However, neither author mentions Long Island in his account of this species. Today, summer colonies are well established at scat¬ tered localities over most of Long Island. During the survey, small bats observed hunting over ponds or clear¬ ings in most sections of the island usually appeared to be this species. Some accounts for the 1930’s indicate it was common then: Nichols and Nichols (1934) found this species the most plentiful bat in summer evenings near Mastic on the south shore in 1934; Turrell (1939) had little to say about bats but remarked that this species was familiar in the Smithtown region near the north shore of the island. Roy Latham, in a personal communication, writes that the little brown species is common on Shelter Island. Reports and complaints of colonies of brownish bats in houses (especially attics), churches, and other build¬ ings during the summer were frequent. Such cases are generally referable to either this species or the big brown bat, but I did not visit very many of the sites. Most ob¬ serving and collecting of this species was done at a readily accessible summer maternity colony discovered in the attic of an old abandoned house near Manorville, in central Suffolk County. However, one other summer colony was visited, situated in a large, old, inhabited house north of Southampton. Also, reports of colonies which were probably Myotis lucifugus, or were identified as this species by the observers, were received from lo¬ calities representing the central, north shore, and south shore sections of Suffolk County. I did not learn of any really large summer colonies, consisting of many hun¬ dreds or even thousands of individuals, such as have been reported for some regions. In evening observations we never discovered large concentrations of hunting little brown myotis such as are frequently encountered in many areas of mainland New York State. Observations, mainly at certain ponds and clearings in eastern and central Suffolk County, only occasionally produced as many as three or four individ¬ uals in a group. At some localities none at all were seen, at others only a single one, even when weather and insect conditions seemed very favorable. This may indi¬ cate a relatively small population, in spite of the known colonies present, or the better hunting grounds may have been overlooked. More little browns were seen in August and September than in June and July, probably refiect- ing increase in the population and dispersal after raising of young. Habitats where bats identified as this species were seen hunting included large ponds in both wooded and residential areas, small pond hidden in dense maple swamp, clearing in pine barrens, field near pond and house, vicinity of a dump, and once or twice at a street light. Myotis lucifugus is also present on Staten Island; in fact all the species of bats which occur on Long Island have also been collected or reported on Staten Island, except perhaps Myotis keenii, for which I have been un¬ able to locate any Staten Island records. The little brown myotis hibernates chiefiy in caves and mines, situations lacking on Long Island. Judging from what is known of seasonal movements of Myotis lucifugus in New England and elsewhere, most of the Long Island population probably migrates north or west to mainland cave regions for the winter. There is very little evidence of hibernation in situations other than caves and mines in the Northeast, although this possibility has not been fully investigated. Of interest in this con¬ nection is a personal communication from Roy Latham; he reports finding little brown bats [Myotis]^ thought to be this species (or possibly M. keenii?) , in winter in the cellar of an old house near a swamp on Shelter Island, and some flying activity on mild days in February. Reproduction {maternity colony). Occasional ob¬ servations were made in 1962 and 1963 at a maternity colony, where, as is the custom of these bats, females gather in summer to give birth and raise their young. The colony was discovered May 23, 1962, in a low attic of an old abandoned farmhouse (figure 10) near Manor¬ ville. About two dozen bats of different ages were col¬ lected here and in outlying buildings during the two seasons. The attic had a broken window at each end, and was about 6 feet high at the center; the bats com¬ monly clustered just under the peak of the roof on top of the ridgeboard. They were readily visible through a nar¬ row space on either side, between the roof and ridge- board. The colony was not large; usually 30 to 50 little browns were visible. A maximum number of 51 adult females was counted on June 26; I believe this was most of them, and allowing for a few probably missed, the colony of adult females probably numbered about 60 in both years. Later in the season, the number was increased somewhat by the addition of young. Numbers remained high into the first week of August in 1962 but by August 9 there was a definite decline in the popula¬ tion as the bats dispersed. Some bats were present at least until September 25, when six remained. Visits in 18 October (9lh and 18th) revealed no little browns, indi¬ cating the site had been deserted for the season. In 1963 no bats were found in mid-April (visits on the 12th and 18th), but the next visit, on May 15, revealed some of the bats had arrived, with about a dozen or more on hand, suggesting that the colony had begun to form at the end of April or in early May. It appeared that most young probably were born in June. Nine females, each bearing a single embryo, were examined between May 15 and June 26. Embryos ranged from 5 to 20 mm. in size; the largest (20 mm.) were examined on June 2 and 26. The first small young, two which were probably a few days old, were seen June 26. These were nursing, were partly furred, and could crawl but not fly (weight of one: 3.7 grams, total length, 62 mm.). One large young close to early flight stage (weight 6.4 grams, total length 78 mm.) and probably 2-3 weeks old, was collected June 29; it could flap its wings strongly but was still nursing and had not yet begun to feed on insects. By August 1, active, approximately fully grown young bats at least a month old were noted in the colony. A few bats occupied a large shed 50 yards away from the house. Most of the time only one or two males plus an occasional Eptesicus were to be seen. But on June 2 an adult female carrying an embryo was found tucked away in a crack, well removed from the main colony. Also from late August to late September some females and many of the active young were present, and had presumably moved over from the main building. In the fall of 1962 the chimney of the house col¬ lapsed, leaving a gaping hole in the roof, which admitted much more light and air into the attic. This change in the environment did not seem to adversely affect the bats, and their production of young the following year, al¬ though observations were terminated July 2, 1963; on this date over 40 adult females and several crawling young were seen. On the evening of June 8, 1962, (weather warm and calm) I watched from outside for bats to emerge, but saw only 20 fly out of the house, fewer than were actually present. Some may have been missed, since the bats came out of several openings beside one of the windows. After 9:15, when it was completely dark, I went inside and found several flying about within the house and at least nine still inactive in the roost. The bats were carrying em¬ bryos at this time, and may have been less active for that reason. On August 29, 1962, after two nights with much wind and rain associated with a tropical storm (“Alma”), three bats collected had virtually empty digestive tracts. Measurements. Fifteen adult females in summer (May 23 to September 25) average: weight, 10.0 grams (7.7-12.0) ; total length, 94.2 mm. (90-101) ; tail, 40.2 mm. (37-44) ; hind foot, 9.5 mm. (9.0-10.0) ; ear from notch, 15.1 mm. (14-16) ; tragus, 6.9 mm. (6.5-7.0) ; forearm, 38.8 mm. (36.5-40) ; wingspread, 264.1 mm. (250-275). Eight of the May and June females carried embryos; some of the August and September bats were quite fat — the heaviest individual (12.0 grams) was one of those collected on September 12. An adult male on June 26, 1962: weight, 8.5 grams; total length, 87 mm.; wingspread, 252 mm.; forearm, 37.5 mm. Individuals taken. 34 Keen's Myotis Myotis keenii septentrionalis (Trouessart) Distribution and habitat. Very little is known about this species on Long Island, and few specimens have been collected here previously as far as I know. The first published occurrence was of an individual found in a house at Mastic, Suffolk County, on August 18, 1933 (Nichols and Nichols, 1934). Two other specimens from Long Island, collected in August 1886 (R. Waldo) and at Mt. Sinai, July 1940 (E. A. Williams, Jr.) are in the American Museum of Natural History. Roy Latham col¬ lected one at Mattituck, September 19, 1931 (skin and skull in his collection). The mapped range for Myotis keenii in Hall and Kelson (1959) seems misleading in that it does not include Long Island; the island is small on the range maps, which are designed to show much of the continent, but the other bats occurring here are correctly indicated. This long-eared, northern myotis is not considered a rare species but it never seems to be abundant either, and it has been studied only casually. Most Keen’s bats have been encountered in caves and mines in winter and during the late summer swarming period; relatively few have been observed on the summer range away from such retreats. Thus a large concentration of this species discovered on a Long Island feeding ground seems note¬ worthy and is described here in some detail. All of our specimens were collected at Fresh Pond, Hither Hills State Park, on the south fork near the east end of the island (figure 22). Here Keen’s myotis was found hunting in the evenings throughout the summer and early fall season and appeared to be the most numer¬ ous bat. Using Japanese mist nets, 11 (three males, eight females) were taken on June 26, 1963; two (one male, one female) on August 22, 1963; and nine (four males, 5 females) between October 2 and 18, 1962. During these operations about 15 other myotis were netted and re- 19 leased, and all of them were Keen’s; no other species of myotis was taken. Besides those netted, additional scores of small bats were seen flying about in a similar manner; probably many or all of these also were M. keenii. Such information suggests that Keen’s may be the commonest summer bat in sections of extreme eastern Long Island, replacing the little brown and big brown bats of farther west. Recent reports received of small brown bats in sum¬ mer on Gardiners Island just a few miles north of Hither Hills may possibly indicate the presence of this species there. Fresh Pond, about one-half mile long and two-tenths of a mile wide, is completely surrounded by woods, chief¬ ly oaks (especially black oak, white oak) with a fairly dense shrub layer. The area is rather hilly. There are some fields and meadows within half a mile of the pond, where an occasional small bat (Keen’s?) was seen. The peninsula is a little more than a mile wide here, with the bat collecting site situated about three-tenths of a mile from its north coast. Keen’s bats are known to live in summer in such retreats as cottages, barns, and beneath the bark of trees. No buildings were noted in the im¬ mediate vicinity of the pond, and I did not find any bats in a hurried check of a few park buildings in the gen¬ eral region, although small colonies could easily be over¬ looked. The Keen’s bats at the pond may represent a gath¬ ering from small colonies scattered in trees in the sur¬ rounding woods, or they may spend the day in buildings farther away, outside the park. This species was first encountered in October 1962, when mist-netting was tried out at Fresh Pond after some red bats were seen. Much of the pond was bordered by impenetrable bushes and small trees, but netting was possible at a tiny shallow cove with a sandy beach. The nets (about 40 feet long, % inch mesh) were set at right angles, approximately, to the shoreline. Usually one pole was placed on shore a few feet back from the water’s edge, the other pole out in the .shallow water; the net was attached between the poles, the lower edge of it a few inches above the water, the upper edge about 8 feet high. One to three nets were set up on netting nights; they were up by early twilight and taken down before mid¬ night. Netting was restricted by the weather — often the evenings were windy here, rendering the nets ineffec¬ tive. On most evenings Keen’s myotis did not appear be¬ fore dark. On October 11, for example, none was seen during twilight in more or less continuous observation; yet four were netted after it was thoroughly dark (8:00, 8:30, 10:00, and 10:15 p.m., E.D.T.). After becoming active, these bats often hunted very low over the water close to shore, sometimes flying in tight circles in the cove and frequently touching the water. Also they were seen at times closely following the shoreline, and flying above the bushy foliage at the water’s edge. Individuals were caught above both water and land, from several inches above the surface to 4 feet high. Often these bats were seen to avoid the nets repeatedly, while at other times they flew directly into the nets. On one occasion a Keen’s bat was observed chasing another one, and thus off-guard both plunged headlong into a net. Bats were not sought here after October 18, although conditions remained warm and favorable for another week. Late in the month the weather turned much cold¬ er, with strong winds, light snow, and frosty nights. This species, like Myotis lucifugus, generally hibernates in caves during the winter, and all or most of the Long Island individuals presumably cross to the mainland for this purpose. In the following year Fresh Pond was first visited on June 26, a warm and calm evening with many insects visible in the air. Many Keen’s bats were about, but I was unable to detect any other species. Eleven keenii were collected using two mist nets placed near the wooded shore, in the usual manner, between 9 and 11 p.m., E.D.T. Scores of the bats were seen; many bounced off the nets, others were caught and released. All females collected carried well-developed embryos. The pond was next visited on August 22, and again Keen’s myotis, but no other species, was recorded. No bats at all were seen until late, darkening twilight (8:30 p.m., E.D.T. ), when suddenly two were flying about close by, often to within a foot or two of my head and shoul¬ ders as I stood in shallow water. Soon more appeared. Three were caught in a mist net. The bats on this date hunted about 2 to 6 feet above the water and shore. Also they sometimes entered the woods by flying very low along paths leading away from the shore. It was not de¬ termined how far into the woods they went, but under the canopy of the trees seemed to be part of their “beat,” at least on this evening. The flight of feeding Keen’s myotis showed a fairly steady course (in straightaway or when circling), with occasional sudden veering to one side; it would not be termed a fluttering flight. Regarding voice, these bats usually seemed silent in flight except for an infrequent short high-pitched squeak. When alarmed or excited, as when caught in a net or being handled, they gave a squeaking chatter very reminiscent of Myotis lucifugus. In partial summary of the above observations on iheir foraging habits at Hither Hills, Keen’s myotis are active in the area at least from June to October, appear late in the evening when it is quite dark, are numerous and associate together when feeding, and have a direct 20 manner of flight; both summer and fall they hunt low over the water (often 1 to 6 feet) ; and they also fly low over the shore, close to foliage, and low under the can¬ opy of woodland trees. In this locality keenii seemed to be the commonest bat and was the only Myntis taken. Stomach contents, containing the remains of many in¬ sects have been saved, but not as yet examined closely to determine the identity of the insects. Reproduction. Apparently small nursery (maternity) colonies are formed, but essentially nothing is known of reproduction in this species (Barbour and Davis, 1969). Reproductive females netted at Fresh Pond in late June indicated the presence of such colonial roosts in the area, but we did not locate them. Each of the eight females taken on June 26 contained a single, fairly large embryo, suggesting birth of young mainly in July (embryo crown to rump measurements, in millimeters, were 8, 13, 13, 14, 14, 15, 15, 16). The date seems a little later than with most of our local bats, and agrees with Hamilton’s (1943) information for elsewhere in New York (he mentioned 3 females with single large embryo taken in late June and early July, indicating parturition in July). Measurements. Eight adult females in June with embryos average; weight, 8.3 grams (6.4-9.2) ; total length, 91.6 mm. (86-96); tail, 38.1 mm. (36-40) ; hind foot, 9.6 mm. (9.0-10.5) ; ear from notch, 17.6 mm. (17.0-19.0); tragus, 9.1 mm. (8.5-9.5) ; forearm, 36.0 mm. (34.5-37.5); wingspread, 249.8 mm. (241-253). Three adult males in June average: weight, 6.0 grams (5.5-6.4) ; total length, 85.7 mm. (83-88); tail, 39.0 mm. (38-40) ; hind foot, 9.2 mm. (8.5-9.5) ; ear from notch, 17.3 mm. (16.5-18.0); tragus 9.0 mm. (all about 9.0); forearm, 34.7 mm. (34.0-35.0) ; wingspread, 243.3 mm. (232-237). Weights of nine specimens of both sexes in October average: 7.2 grams (5.9-9.7). All of the above bats had food in their stomachs. The October specimens had accumulated considerable body fat. Individmds taken. 22 Silver-haired Bat Laisonycteris noctiva^ans (Le Conte) DeKay (1842) termed the silver-haired bat as com¬ mon on Long Island (season not stated). Since then, various writers (Helme, 1902; Murphy and Nichols, 1913; Nichols and Nichols, 1934) have shown that this northern hat is rare on Long Island in spring and early summer, is occasionally plentiful in late summer and early autumn (period of southbound migration), and that, singly or in groups, a few remain to hibernate dur¬ ing the winter in hollow trees, in ships, and in buildings of every description. It has been reported from Staten Island, New York Harbor, and Brooklyn east to Orient Point, Montauk Point, and over the ocean. Nichols and Nichols (1934) collected five specimens between June 17 and July 10, 1933, at Mastic, establish¬ ing the first definite information on the presence of La- sionycteris through the summer. The authors, experi¬ enced in collecting bats on Long Island, were surprised to encounter this species in June, outside the period of south¬ ward migration; two of the June ones were females carry¬ ing well-developed embryos. As with the red bat, the highly migratory silver- haired bat has been encountered migrating south along our beaches (both north and south shores) and even out over the ocean. Migrating individuals may linger to hunt over woods, clearings, and bodies of fresh water; on the coast quite a few have been found resting on vessels at anchor. Helme (1902) stated that in some years during migration this species even outnumbers the red bat, but that in other years very few are seen. Large numbers have been seen gathering in late summer in Prospect Park, Brooklyn, and flying over the lake there (Murphy and Nichols, 1913). We did not collect or positively identify this species on the survey; some of the bats seen in the Hither Hills area in October 1962, may have been silver- hairs — if so, they were apparently outnumbered by Keen’s bats and red bats. Roy Latham has in his collection sev¬ eral Lasionycteris secured during the month of September on the north fork of eastern Long Island and on Shelter Island; two specimens from Shelter Island were found drowned in a pail of water on September 3, 1929, by W. W. Worthington. Latham’s latest fall record is Oc¬ tober 10 (1939). Robert Cushman Murphy has ma'de many observa¬ tions on the migration of this species along our coast. Early in the morning of September 6, 1907, while in a boat 4 or 5 miles off Sandy Hook, New Jersey, in blustery weather, he saw several individuals struggling towards the Staten Island shore (Murphy and Nichols, 1913). A speci¬ men in the American Museum of Natural History was col¬ lected by Murphy while 3 miles at sea off Long Beach, Nassau County, on September 7, 1918. Murphy and Nich¬ ols also wrote that silver-haired bats have been seen in September evenings flying above the beaches of Long Island Sound and circling high over Mt. Sinai Harbor and other salt water inlets. A far-offshore record is one col¬ lected on August 19, 1953, after it circled about and landed on the rigging of a vessel in 39°36' N, 71°03^ W, about 95 miles SSE of Montauk Point, the nearest land (Mackiewicz and Backus, 1956). 21 Long Island seems to be near the northern limit of the winter range of this species. Regarding winter habits in our area, Murphy and Nichols (1913) wrote: “Silver- hairs . . . are not infrequently found in hollow trees cut in winter for firewood, and they also have been discov¬ ered hibernating in sky-scrapers, churches, wharf-houses, and the hulls of ships in New York City and Brooklyn, during the months between December and March.” The authors also referred to over 20 “black bats” found in January in Port Jefferson Harbor aboard a yacht, and then thrown overboard by the captain, as probably be¬ longing to this species. Roy Latham (personal communi¬ cation) has one winter record for Orient, a lone individ¬ ual found in a building on January 23, 1946. Another winter record is a female Lasionycteris in the American Museum of Natural History which was collected at Mastic in February 1937, by David G. Nichols. Eastern Pipistrelle Pipistrellus subflavus subflavus (F. Cuvier) The little pipistrelle is apparently uncommon on Long Island. Helme (1902), Murphy and Nichols (1913), and Nichols and Nichols (1934) did not mention this species. But Hamilton (1949) wrote that one summer evening he nearly succeeded in capturing one with an insect net. Roy Latham collected a male pipistrelle at Montauk on August 25, 1931; preserved as skin and skull in his collection, this is the only Long Island speci¬ men I have examined. During our survey, a bat identified as a pipistrelle was observed on the night of October 10, 1962, at Bel¬ lows Pond near Hampton Bays. At 8:00 p.m., E.D.T., in bright silvery moonlight, I watched a tiny bat with an erratic and moth-like fluttering flight hunting back and forth above the tops of tall bushes near the pond. It appeared smaller and had a different manner of flight than the little brown. Keen’s and other bats that were ob¬ served on the island. Known hibernation sites of the pipistrelle are chief¬ ly caves and mines, and the deficiency of such sites on Long Island may account for the scarcity of this species. Not much is known about distances traveled by this small bat to and from its winter quarters. On Staten Island, pipistrelles have been found in barns and other buildings during the summer. Several which have been collected there are in the American Museum of Natural History. Big Brown Bat Eptesicus juscus fuscus (Palisot de Beauvois) Distribution and habitat. This large bat may be seen anywhere on the island, and since it is hardy and often hibernates in buildings it may also be encountered during any month. Eptesicus fuscus is a rather sedentary species, as stated by Barbour and Davis (1969), and it seems like¬ ly that most of the population remains on Long Island throughout the year. Maternity colonies as well as hiber¬ nating sites appear to be most frequent in the western part of the island, and this bat is not uncommon in the urban areas of Brooklyn and Queens. Murphy and Nichols (1913) wrote that Eptesicus frequents large parks and cemeteries in Brooklyn, and hibernates abundantly in city buildings of the area. Several old publications in¬ dicate that the big brown was also common on Long Island during the last century, and DeKay (1842) ob¬ tained this species in Kings County (Brooklyn). Eptesi¬ cus also raises young (at least locally) and hibernates in Suffolk County, but becomes less common towards the east end of the island. Complaints about bats in buildings frequently came from western Suffolk and eastern Nassau Counties. On investigating a few of these reports, some big brown bats were found. In a small village in eastern Nassau County, a maternity colony was located high in a house (where 20-30 bats were shot in the previous year in an unsuccessful attempt to eliminate them). The bats were not readily accessible, but big browns were seen peer¬ ing out of holes near the roof on July 27. On the previous day, which was very hot, the bats had been reported hanging on the outside surface of the house with their young. Also, bats were heard squeaking during the day in woods near the house, possibly in hollow trees, al¬ though we were unable to locate them. At another house nearby in the same village, during late May, 25 bats (Eptesicus?) were reported shot while flying out of the attic at dusk, after which no bats were seen there. In a nearby town up to 60 bats had been reported emerging at dusk in May from the attic of a large building. On entering the attic June 2, I found the floor littered with many droppings which seemed to be of this species, but the site appeared to be almost de¬ serted, even though these bats had not been shot or mo¬ lested as far as I know; only a solitary male Eptesicus was present. Big browns occasionally leave a nursery roost at least temporarily; one reason, as mentioned by Barbour and Davis (1969), is that this species is not as tolerant of high temperatures as Myotis lucifugus, and during a hot spell may move to a cooler part of the building or 22 abandon it altogether. Or possibly this was a recently- abandoned winter roost. A maternity colony of big brown bats has occupied a large bam near Yaphank in central Suffolk County for some years (LeRoy Wilcox, verbal communication). At the attie colony of little brown bats near Manor- ville, one or two adult male Eptesicus were often found in small sheds near the main house, from May to October. Although maternity colonies of these two species do not occur together, individual male big brown bats were ap¬ parently attracted to the vicinity of the Myotis colony. In October, after the little browns had abandoned the location for the season, single big browns were occa¬ sionally seen hanging in the attic itself (where the small¬ er species had raised young). The big brown bat was frequently detected in an area by seeing lone individuals, just before dark, flying rather high on a straight, steady course to their feeding grounds. Localities in eastern Suffolk County where Ep¬ tesicus was seen or collected, although maternity colo¬ nies were not found, include Amagansett, Calverton, Flanders and Squiretown. Near Amagansett, well out on the south fork, several were seen in late June hunting around street lights near the ocean. Solitary individuals were noted on two or three occasions flying before dark near buildings on the outer beach strip (between Shin- necock and Moriches inlets). Summer maternity colonies and wintering individuals have been found on Staten Island. Food habits. Only two stomachs were examined, and these were filled almost entirely with beetles (Coleoptera) , including large June beetles of the family Scarabaeidae; some remains of moths (Lepidoptera) were noted in one of the stomachs. Individuals taken. 4 Red Bat Lasiurus borealis borealis (Muller) Distribution and habitat. Helme (1902) designated this species as the most abundant bat on Long Island. Likewise, Murphy and Nichols (1913) stated that “Dur¬ ing most of the summer a bat seen anywhere on Long Island is, nine times out of ten, a red bat. The species is abundant from the busy streets of Brooklyn to Orient and Montauk . . As far as I have been able to deter¬ mine, this no longer seems to be true on most of the island, although the red hat is not rare. In many areas the little brown myotis, which was found to be scarce by Murphy and Nichols, is the bat most likely to be encountered. Locally, other species, namely Keen’s and big brown bats, appear to be first in abundance. Also, in some apparently favorable situations, such as certain ponds and fields on still summer nights when insects were abundant, we failed to detect any bats at all. There does not seem to be much information on the status of this bat during the early summer season (June, July), before the fall migration, except that it appears to be widely distributed but not abundant then. In 1933, four red bats were collected between July 3 and August 7 among more plentiful Myotis lucifugus at Mastic (Nich¬ ols and Nichols, 1934). Lanyon (1961) termed the red bat a regular but uncommon summer resident on the Kalbfleisch Field Research Station in the Dix Hills area of western Suffolk County, and mentioned a specimen collected on July 12, 1961. Red bats have been collected on Long Island in every month from late May to late November; many specimens are in the American Museum of Natural His¬ tory. Murphy and Nichols (1913) stated that the red bat appears on Long Island on the first warm spring days, and that there is a Staten Island record as late as December 5. Kimball and Nichols (1940) have recorded this species flying about at Mastic, Suffolk County, on mild days in December (as late as December 26, 1936, and December 25, 1937). I do not know of any winter records later than December. Most red bats move well to the south of Long Island for the winter; but the late December sight records suggest that a few individuals, hidden and dormant, may remain this far north all win¬ ter, emerging to fly only during unseasonably warm spells. The red bat does not hibernate in caves, and gen¬ erally roosts in trees throughout the year. The red bat is most frequently observed in this region during late summer and early fall. This species, like the silver-haired and hoary bats, is highly migratory, and southbound flights and coneentrations are often no¬ ticed along the coast at this season. Red bats may be seen flying by day over the beach or ocean, or found roosting in unusual locations. Carter (1950) reported on a flight of 200 bats seen around a ship enroute to New York, about 65 miles offshore (40°10' N, 71'’00' W) on September 29, 1949; three individuals collected were red bats. Several reports we received, from south shore lo¬ calities in the fall, of bats hanging in trees and bushes during the day or found dead on the ground, seem to refer mainly to this species. LeRoy Wilcox and Walter Terry have informed me that they usually catch one to several red bajts in mist nets each fall, while banding migratory birds on the outer barrier beach strip near 23 Quogue. These bats probably come across Long Island Sound, too, and during October many have been found hanging asleep on exposed roots under the overhang of the beach cliffs which fringe the north shore (Murphy and Nichols, 1913). The seven specimens I collected were taken between August 21 and October 18; also all of my field observations of this species were made dur¬ ing August, September, and October. Field observations. Red bats were seen in August in the area of Shinnecock Bay and adjacent barrier beach. Individuals were occasionally seen flying over the bay in all directions at twilight, apparently dispersing to feed rather than migrating on these flights. Here, on the outer beach strip near Shinnecock Inlet, I watched one mi¬ grating by day on August 10, 1968, at 4:30 p.m., E.D.T., in bright sunlight. The bat flew over the marshy bay side of the strip, heading in a generally southwesterly direc¬ tion, that is, following the coast. When “on course” it flew high with a steady flight, but also it occasionally in¬ terrupted its progress to dip down low, perhaps to feed. Once it flew low around a small salt marsh pond close by; the red color was striking in contrast to the green of the marsh grass. Numerous birds, including terns, shore birds, starlings, and swallows were present, but they showed no interest in the bat. After a few moments the bat resumed its trip. I followed its course with binoc¬ ulars as it flew, occasionally dipping down again, before it was lost to sight. Also on this date there was a nearly continuous passing of barn swallows (Hirundo rustica) , tree swallows {Iridoprocne hiocolor) , and monarch but¬ terflies (Danaus plexippus) , all migrating in the same direction as the bat. During our survey, these bats were most regularly encountered on the eastern part of the south fork, hunt¬ ing over ponds and clearings and around certain street lights during the early fall season. Here they seemed to gather and linger, and could be seen whenever the weath¬ er was not too bad. Rather intensive searching in these same areas in June, however, failed to indicate the pres¬ ence of red bats. Red bats were observed on mild October evenings at Fresh Pond (figure 22), the large pond at Hither Hills described in the account of Keen’s myotis. Here they were seen flying high over the pond and adjacent woods, and also hunting close to dense vegetation along the shore or skimming over the water. Occasionally one was seen early, while it was still quite light, but normally red bats did not appear until late twilight or after dark, when they were caught in Japanese mist nets or spotted momen¬ tarily in the light of a strong flashlight. The presence of hunting red bats seemed to be determined in part by the occasional large concentrations of moths along the shore of the pond. Bats of both sexes appeared to be common. Migrating hawks of various species were occasion¬ ally noted in this area in the fall, and as many as seven pigeon hawks (Falco colunibarius) were seen in one afternoon flying over or perching near the pond. One of the early flying red bats observed was apparently pur¬ sued by a pigeon hawk. As I watched a red bat circling above the pond late in the afternoon, on October 11, 1962, a pigeon hawk darted from its lookout perch on a tree on the opposite side of the pond and headed directly toward the bat. The bat turned toward the woods, the hawk followed, and both disappeared behind the trees without my learning the outcome of the brief episode. But it seems likely that this swift little falcon, a fairly com¬ mon coastal migrant in early fall, may at times prey on migrating or other day-flying bats, as well as birds. At Fresh Pond, red bats were taken in mist nets placed along the shoreline over the water and beach; net captures were made from just a few inches above the water to over 6 feet high. Sometimes these bats were observed to detect the nets as they hunted low over the water, veering off at the last possible moment time and again. Once one was seen to collide with a net, and then fall into the water; but it managed to fly up into the air after flapping its wings on the water for a few mo¬ ments. These bats occasionally gave rather loud hissing screams when caught or handled. Near some villages on the south fork red bats were seen hunting around bright street lights, often quite near the ocean. Usually they were seen repeatedly at certain lights to the exclusion of other similar lights nearby, probably indicating rather fixed hunting territories of individual bats. At the lights the bats made swift passes at the clustering insects, then disappeared into the dark for a while before returning. Food habits. The stomachs of four red bats col¬ lected at Fresh Pond during October 1962, contained many moths. Many, perhaps most, of the moths belonged to the family Geometridae, and numerous moths of this family were also observed swarming over the water and near shore during warm evenings. Small beetles and other insects were also eaten. Measurements. Four October specimens average: weight, 10.6 grams (9.2-11.8) ; total length, 102.3 mm. (96-106) ; tail, 44.5 mm. (42-48) ; hind foot, 8.6 mm. (8.0-9.0) ; forearm, 39.3 mm. (37.0-40.5) ; wingspread, 301.0 mm. (275-313). Individuals taken. 7 24 Hoary Bat Lasiurus cinereus cinereus (Palisot de Beauvois) On Long Island this large, handsome bat is the rar¬ est of the three highly migratory species, which (like birds) fly south in the fall and north in spring. As with the red and silver-haired, most occurrences are in late summer and autumn. Murphy and Nichols (1913) knew of records of hoary bats being collected on Long Island in August, September, and October, including one dis¬ covered under a driftwood plank on a north shore beach. Kimball and Nichols (1940) reported an individual hang¬ ing on the porch of a cottage facing the sea at Oak Is¬ land, south shore of Suffolk County, on December 2, 1939, apparently the first published record later than October. It was assumed by the authors that this individ¬ ual was still migrating on this late date, not disturbed from hibernation. This heavily furred species may be considered a rare possibility any time during the colder months. There are a few December, January, and February records for the general latitude of Long Island (Connecticut, New Jer¬ sey, and Pennsylvania). DeKay (1842), in his discus¬ sion of this bat in New York State, mentioned seeing two individuals actively flying about shortly before noon on December 12, 1841 ; he did not mention the locality, but I assume it was somewhere in the State, possibly even Queens, Long Island, then DeKay’s place of residence. In the American Museum of Natural History there is a female hoary bat, preserved in alcohol, which was col¬ lected on January 16, 1969, at Far Rockaway, southern Queens, by John Bull; this is the only midwinter record for Long Island, as far as I know. Such records may in¬ dicate that a few individuals hibernate in this region, well to the north of most of their fellows. Hoary bats, like red bats, normally roost among the twigs and foliage of trees and generally do not frequent caves or buildings. Perhaps trees serve as winter retreats also, with the bats leaving them to fly after insects on warm days, although practically nothing is known for certain about the win¬ ter habits of this species. There seems to be no information on spring migra¬ tion of hoary bats in the Long Island area. Nor do I know of any records of these elusive bats during the period from about late May to early July when they give birth to young. They appear to be more common farther north at this season, but a few may occur here since Roy Latham has recorded this species as early as July 14, and specimens have been collected in nearby New Jer¬ sey and Pennsylvania during June. Our only observation on the museum survey was of one flying low, near a street light after dark, at Hamp¬ ton Bays on September 29, 1962. For Orient, Roy Latham (personal communication) has four records, dates rang¬ ing from July 14 (1963), as mentioned above, to Oc¬ tober 29 (1919). Eastern Cottontail Sylvilagus floridanus mallurus (Thomas) Distribution and habitat. Two cottontail rabbits, the New England cottontail {Sylvilagus transitionalis) and the eastern cottontail (S. floridanus mallurus) occur on Long Island, as indicated by Nelson (1909), Hamilton (1949), and others. Both kinds appear to be rather wide¬ ly distributed on the island, based on specimen records. But the exact status of the two species — relative abun¬ dance and patterns of distribution, perhaps in the process of change — awaits clarification. Nelson (1909) and others have shown that transitionalis was probably the only native cottontail in an extensive area of the Northeast, including all of New York State and adjacent states to the north and south, and that floridanus has extended its range into this region rather recently. Possibly floridanus is in the process of replacing transitionalis on Long Is¬ land, as it has in some mainland areas of eastern New York and southern New England. Also, cottontail rabbits of other subspecies of S. floridanus from certain midwestern states have been introduced on Long Island. Very likely some of these arrivals survived to interbreed with the local stock (5. /. mallurus), although presumably the native form tends to prevail in most areas of the island. Western cottontails became very numerous on Fishers Island following intro¬ duction (Joseph Dell, verbal communication). Although fluctuating in abundance from year to year, cottontails are generally common from highly de¬ veloped residential areas and parks in eastern Queens and even parts of Brooklyn to the very tip of Montauk Point, and near both the north and south shores. They are numerous on some outlying islands, including Shelter Island, Gardiners Island, and the full length of the outer barrier beach of the south shore, including Fire Island. Apparently S. floridanus is the common cottontail in most of these areas, judging from the many museum specimens of this species (American Museum of Natural History, U.S. National Museum, N.Y. State Museum, and elsewhere) from more than 30 localities throughout the island. The eastern cottontail is also common on Staten Island. 25 On the field survey, we collected 17 cottontail rab¬ bits in the area south and southwest of Great Peconic Bay, that is, from Riverhead and Flanders south to Mo¬ riches, and outer beach near Quogue, Hampton Bays, and Mecox Bay. These animals all proved to be flori- danus (based primarily on skull characters commonly used in separating this species from transitionalis) . Ad¬ ditional eastern cottontails collected included specimens from Shelter Island and Hither Hills. It seems amazing that these fairly sizable, defense¬ less animals manage to survive in many built-up areas where their habitat consists solely of the yards of sub¬ urban homeowners. Here they thrive in spite of the many dogs, cats, people, and automobiles. Although not often noticed during the day, a visit to such areas at night may reveal rabbits feeding in the open on grass, clover, and weeds, or sometimes on the more valuable plants of yard and garden. In the less developed countryside cottontails are widespread, wherever suitable food is found, and they feed on a vast array of shrubby and herbaceous plants. We found rabbits present in all sorts of woods, from dry to swampy, as well as in fields, bogs, and various grassy and bushy areas. Cottontails are found throughout the pine barrens and dry oak woods, although presumably in fewer numbers than in areas endowed with a greater variety of food plants. Often they were seen in or along the edge of red maple swamps; here there was evidence of their browsing heavily in winter on young red maples and other plants, such as greenbrier {Smilax) . On the highly agricultural north fork of the island, cottontails were very numerous wherever there were trees or bushes, at least in 1963. Cottontails do well in the maritime areas. A walk through the grass and low shrubs among the ocean- front dunes often results in one to several individuals being startled from their forms and hiding places. Their droppings often seem to be “nearly everywhere” on the sand. The rabbits are even found in the beachgrass on the very summits of the outer or primary dunes, one of the few mammals regularly found in this environment. They may be encountered here any time during the year, although they probably withdraw someWhat to¬ wards the better cover about the inner dunes and edge of salt marshes during winter. In many sections, the salt marsh edge, with its rather good variety of food and cover plants is home for numerous cottontails. They also venture out into these marshes to a limited extent. On Fire Island we saw cottontails (and abundant signs of their presence) among the dunes, including the vicin¬ ity of the Sunken Forest. Measurements. Four winter adults average: weight, 1,167.5 grams (984-1,279) ; total length, 407.8 mm. (398- 421) ; tail, 43.5 mm. (40-45) ; hind foot, 93.5 mm. (91- 96); ear from notch, 62.0 mm. (59-67). Individuals taken. 20 New England Cottontail Sylvilagus transitionalis (Bangs) The New England cottontail is a small rabbit which is very similar in external appearance to the eastern cot¬ tontail, the two differing only slightly in pelage and form. But the two cottontails are distinct and separate species, even though closely related, and apparently they rarely hybridize in the wild. The New England cotton¬ tail does not presently appear to be as common on Long Island as the eastern cottontail. Nevertheless, in various museums there are specimens of transitionalis which have been collected at localities in Nassau County and east through Suffolk County (including some north shore and south shore localities) to Shelter Island and Montauk Point. During the 1930’s about 20 were collected at Mastic by David G. Nichols and John G. Nichols (specimens in American Museum of Natural History). This species has also been collected on Staten Island. The various locality records indicate that the two species of cottontail may be found in the same general area on Long Island. But I have no information on habi¬ tat differences, if any, although it has been shown that on the mainland transitionalis tends to prefer wooded and brushy areas, as opposed to more open areas often favored by floridanus. Nelson (1909) observed that specimens of S. flori¬ danus mallurus from New Jersey and southeastern New York, including Long Island, are especially close to S. transitionalis in size, and often in color. Benton and Atkinson (1964) add that identification of the two species is virtually impossible in the field and is difficult in the hand. This situation is frustrating to the active naturalists of the island who observe and study the local fauna. During the survey, we occasionally observed rabbits at very close range, encouraging tentative species identifi¬ cation based on several external characters such as ap¬ parent length and shape of ear (shorter and more rounded in transitionalis). Individuals thought to be transi¬ tionalis were observed at several south shore localities from Babylon to Montauk Point, and also on the north fork of eastern Long Island. However, one such individual was collected and it proved to be floridanus instead, thus 26 our field identifications cannot be relied on, although perhaps some of them were correct. A close examination of key skull characters, which are consistently different in the two species, is the de¬ pendable way to distinguish the cottontails; Hamilton (1943) and Fay and Chandler (1955) are helpful ref¬ erences for Identifying specimens. Recently, Hinderstein (1969) made a comparative study of skull characters of the eastern and New England cottontails in the region of northern New Jersey to Connecticut, including Long Island; she also studied geographical variation between Long Island and mainland specimens of each species. Woodchuck Marmota monax rujescens A. H. Howell Woodchucks are only locally common, being distrib¬ uted in a spotty manner over an extensive portion of central Long Island. I saw very few woodchucks during the survey, these primarily in fields and scrubby woods in the Peconic River region of Suffolk County. However, woodehucks occur westward about as far as central Nas¬ sau County, and are still common enough to cause com¬ plaints from gardeners in some areas of that county. They are also seen regularly along the Northern State Parkway north of Hicksville. Lanyon (1961) terms this species as regular and fairly common on the Kalbfleisch Field Research Station, in the Dix Hills area near Hunt¬ ington, with daily observations (including den sites and juveniles) from mid- April to late November. Several writers have mentioned various habitats frequented hy woodchucks on Long Island, including fields, meadows, wooded fence rows, brush, scrub oak woods, oak-hickory woods, hillsides, and highway and railroad embankments. Woodchucks are found near the north and south shores in places, but today are rare towards the eastern end of Long Island as well as in the built-up areas of the western part of the island. Evidently the species was formerly much more numerous, but has declined in numbers; the reasons for this are not entirely clear. Helme (1902) maintained that woodchucks, although still com¬ mon at the turn of the century, had become greatly re¬ duced in numbers in many localities after several towns began paying bounties for their destruction. Also the rather barren, sandy pine and oak areas with their aban¬ doned fields, although wild enough, may be eapable of supporting only a low population of chucks. According to Roy Latham (personal communication) years ago woodchucks were very abundant and wide¬ spread on the east end of Long Island, ineluding Shelter Island; there were many in the agricultural Orient re¬ gion before 1910, but Latham recorded his last one there in 1915. He reports that they are still present in the Sound Avenue section north of Riverhead and are found eastward from there, in small numbers, as far as Great Peconie Bay. Latham’s earliest seasonal record of this hibernator on the north fork was one caught near Orient on February 1, 1902. Long Island specimens (skins and skulls) are pre¬ served in the American Museum of Natural History, in the U.S. National Museum, and in the collection of Roy Latham. Although uncommon now, woodchucks have been on the island for a long time, and their remains have been found in Indian archaeological sites dating back many centuries before the white men arrived. Eastern Chipmunk Tamias striatus fished A. H. Howell Distdbution and habitat. Chipmunks are moderately common to conspicuously abundant in most wooded areas of Long Island. We saw them in and near parks in Queens and in residential areas on the north shore of Nassau County, eastward to Hither Hills State Park well out on the south fork. Also, chipmunks are reported as still present in Prospect Park, Brooklyn [where they are less common than formerly] and to occur as far east as Montauk (Anon., 1968a). Roy Latham collected one at Montauk in June 1927 (in his collection). Pos¬ sibly this species is relatively new in the Montauk area, since Butcher and Butcher (1893) stated that there were no chipmunks in the woods of Montauk. Tamias is also present on Staten Island. Chipmunk distribution appears to be “spotty” com¬ pared with many mainland regions, however, sinee we were unable to find it in several seemingly suitable areas near both the north and south shores. Also, Tamias shuns the immediate vieinity of the ocean beaches here, and is absent from most if not all of the small outlying is¬ lands off the east end and south shore of Long Island. Tlae majority of the specimens were colleeted in central Suffolk County south of Riverhead and Manor- ville, in pine barrens, and in oak woods and other de¬ ciduous growth. They are quite common in the pine barrens, especially near the ponds and streams found in parts of this region; pitch pine, bear oak, and blueberry comprised the principal woody cover in many of the pine-region localities. Chipmunks were noted as common in oak woodlands, also locally in various scrubby woods and residential areas near bays along the south shore. 27 Chipmunks were numerous throughout the period of this survey on the south fork of eastern Long Island in woods composed primarily of various species of oaks (white, black, scarlet). In contrast, this species seemed to be very rare on the north fork, at least during the summer of 1963 when observations were made in a number of wooded areas, including Browns Hills and Great Hog Neck (none seen, although probable burrow noted on Great Hog Neck) . Gray squirrels, however, were recorded as being very numerous in the same north fork areas. Roy Latham (personal communication) writes that there are no chipmunks in Orient, and he has never seen them in Greenporl, although he reports that a few chipmunks are found between those localities, at East Marion. From this it appears that Tamias striatus is quite scarce and local in this northeast section of the island. Shelter Island appears to lack chipmunks altogether. In a week of active field work in June 1963, no chip¬ munks were found. Roy Latham adds that as far as he knows there are no Shelter Island records. Food habits. Chipmunks feed on many kinds of nuts, seeds, fruit, and other plant material, and also on inver¬ tebrates and small vertebrates. Our observations were few and casual. Chipmunks were noted as consuming acorns and hickory nuts, the former probably an impor¬ tant staple here. In the pine barrens they feed on the seeds of pitch pine, as do gray squirrels and flying squir¬ rels. During August I saw chipmunks climb wild black cherry trees (Prunus serotina) to feed on the fruit. In September some were seen joining robins and catbirds to feed on the fruit of tupelo {Njssa sylvatica). Wild grapes and blueberries also were eaten in season. Remarks. Little information was gathered relating to reproduction. A few females were collected in June and September with three to six placental scars, prob¬ ably indicating spring and summer litters (there are two breeding periods in the eastern chipmunk). Chipmunks sleep underground through the winter, although occasionally individuals are active above ground at this season. We did not see any chipmunks between November 22 and April 5, and most of our records were from May to October. We received two or three reports of all-white chip¬ munks on Long Island. However, albino chipmunks are rare and were not seen by us. One day during June, a red-tailed hawk {Buteo ja- maicensis) was observed to catch a chipmunk in low, scrubby pine plains habitat just inland from Napeague Beach, an area, incidentally, where I had not previously suspected the presence of chipmunks. The hawk watched from the top of a telephone pole for several minutes, then it dove down, disappearing briefly among some bushy pitch pines, before emerging with a chipmunk in its talons. It flew off with its prey, pursued by a pair of noisy sparrow hawks {Falco sparverius) which apparent¬ ly had their headquarters nearby. Measurements. Eight adult males (Suffolk County; April to September) average: weight, 88.8 grams (79.0- 105.3); total length, 237.7 mm. (230-251); tail, 87.3 mm. (85-90) ; hind foot, 34.0 mm. (31.5-36). Eight non¬ pregnant adult females (Suffolk County; June to Sep¬ tember) average: weight, 82.5 grams (75.5-86.6) ; total length, 233.3 mm. (228-238); tail, 86.7 mm. (83-97); hind foot, 34.0 mm. (31-38). Individuals taken. 26 Gray Squirrel Sciurus carolinensis pennsylvanicus Ord Distribution and habitat. This large squirrel is the conspicuous diurnal tree squirrel of Long Island; the red squirrel (Tamiasciurus) does not occur. We observed gray squirrels in all types of woodland: coniferous pitch pine barrens and small stands of southern white cedar as well as deciduous types such as oak, mixed deciduous, and swampy woods of red maple and tupelo. An adapt¬ able species, it is found in city parks and crowded sub¬ urbs as well as in predominately open agricultural land, provided there are scattered trees, groves, or small wood- lots. Occasionally individuals were encountered traveling on the ground at considerable distances from trees in both suburban and farm areas. As one would expect, general observations indicate that gray squirrels are more numerous in the richer and more diversified leafy woods such as occur on the north shore, than in the pine barrens region. The former pro¬ vides a more plentiful and dependable food supply. The species was noted as very numerous — at least in 1963 — on eastern Long Island’s north fork, where it was seen or heard almost every place where there was woody vege¬ tation. For example, in Browns Hills, which extend to within 3 miles of the tip of Orient Point, it was noted as very numerous in June of that year, in scrubby areas as well as among tall trees. There is a great variety of trees here, including much hickory. On the south fork in the same year, gray squirrels seemed much less numerous, although they were observed in various areas east to Montauk Point State Park. These squirrels were also recorded on Shelter Island and on outlying Gardiners Island. In the pine barrens of Suffolk County gray squirrels are fairly common; they are found 28 throughout the region wherever the trees are of appre¬ ciable size and they are often seen near streams or other bodies of water. Occasionally gray squirrels were noted in bushy pine plains habitat, probably foraging for food or passing through rather than residing in such areas. These squirrels are probably hard pressed to find food in the pine barrens during years of failure in pine cone or oak acorn production. Gray squirrels are found near the ocean towards both ends of the island, but they become very rare and local on the barrier beach strip where it is widely sep¬ arated from the mainland by Great South Bay and other bays. Gray squirrels were noted to inhabit cavities in trees and also to build conspicuous outside leaf nests (often of oak leaves), as is the custom of this species. Reports were received of gray squirrels causing rather severe damage inside buildings, by chewing holes through walls in the attics of houses, and in summer camp and gun club buildings in the woods. Food habits. Casual observations indicated gray squirrels feed on such items as nuts of hickory and black walnut, acorns of various oaks (including white oak, black oak), and seeds of pitch pine. Many of the feed¬ ing signs seen under pitch pines — little piles or scat¬ tered litter of scales and stripped cores of cones — were probably the work of gray squirrels. Nichols (1927, 1958) made interesting observations on the food habits of gray squirrels in a suburban area of western Long Island. Color. Gray squirrels in the black color phase are un¬ common here, but are occasionally observed. Several in¬ dividuals essentially black throughout, although often with a noticeable brownish tinge to the pelage, were noted in Queens, Nassau, and Suffolk Counties. Appar¬ ently the black form is especially scarce on eastern Long Island, but I saw one at Hampton Bays in October 1961 ; Haft (1930) reported on a melanistic gray squirrel from East Hampton (in the collection of Roy Latham). No white (albinistic) gray squirrels were seen, but reports were received of partially and completely white squir¬ rels in certain areas, including Shelter Island and central Suffolk County; Roy Latham reports two white ones observed on Shelter Island in the winter of 1951. Measurements. Seven adult males from Suffolk Coun¬ ty average: weight, 609.7 grams (528.5-661.0) ; total length, 491.5 mm. (488-496) ; tail, 215.7 mm. (208-230) ; hind foot, 69.7 mm. (68-74). Individuals taken. 11 Southern Flying Squirrel Glaucomys volans volans (Linnaeus) Distribution and habitat. This nocturnal species is still common in the more heavily wooded sections of central Long Island. There we found flying squirrels in woods with assorted oaks or other deciduous trees, pitch pine, mixed pitch pine and oaks, and southern white cedar. Glaucomys is present also on the north side of the island, near the Sound, and on the south side, near the bays, wherever there are suitable woods. Eastward, however, the flying squirrel apparently extends only as far as Riverhead and Hampton Bays. It seems to be absent from woods on both the north and south forks and on the neighboring islands, such as Shelter Island and Gardiners Island; at least I have not heard of any records for these areas. Roy Latham (per¬ sonal communication) has never encountered this spe¬ cies in years of field work on the north fork. In the present survey, many hours of evening observations and rodent trapping in the extensive oak woods of Hither Hills failed to indicate the presence of flying squirrels, although gray squirrels and chipmunks were common enough; the last two are evidently much more accom¬ plished in crossing treeless ground and necks of land to reach such areas. To the west, Glaucomys is restricted mainly by the expanding metropolitan area. Flying squirrels are pres¬ ent on the Kalbfleisch Field Research Station, in the Dix Hills area of western Suffolk County (Lanyon, 1961). In Nassau County a few are still present locally, and they also occur, or formerly occurred, within the limits of New York City, including Queens and Staten Island. Flying squirrels were observed (a few collected), and seemed to be fairly common, well out in the extensive sandy pine barrens of Suffolk County south of the Pe- conic River (figure 3). They were not noted in the low, shrubby so-called “pine plains,” but frequented areas where the pine trees were of fairly good size. But there was rarely much variety in the flora in these localities — mainly just the rather well-spaced pitch pine {Pinus rigida) , some low understory oak, usually bear or scrub oak (Quercus ilicifolia) , with the prostrate shrub, bear- berry {Arctostaphylos uva-ursi) covering much of the ground. There was evidence of flying squirrels feeding on the seeds of pitch pine, and it may be that in this sparse habitat much of the diet consists of pine seeds, acorns, and perhaps fungi and insects or other animal food. Evening parties of flying squirrels were observed in stands of southern white cedar {Chamaecyparis thyoides) 29 in this region, especially along the edge of a large (for Long Island) cedar swamp near Riverhead (figure 7). Remarks. Flying squirrels live in hollows of trees and stubs and also may build outside nests. One January day in a pitch pine and oak woodland near Hampton Bays I briefly shook a tall pine stub which had a con¬ spicuous hole about 11 feet above the ground. A flying squirrel immediately came out of the hole and scurried to the top of the stub, after which a second one ap¬ peared at the entrance and gazed down at me, without emerging. After I backed off a short distance, they both settled back down in the cavity. Woodcutters reported up to a dozen or so flying squirrels found together in one large hollow tree or stub, and also we heard that they occasionally take up residence in buildings near woods, habits characteristic of this species. At another locality, near Flanders, two females were caught during April at the base of an old pine stub (which had numerous Jioles). This area was of interest in that the stub was on a dry slope just above a small white cedar swamp, and in some of the cedars were outside nests somewhat resembling in general appearance the work of red squirrels (which do not occur on Long Island). The nests, most of which were fairly high up and on branches near the trunk, were globular, compact, and constructed mainly of shredded cedar bark with some cedar twigs and leaves included. These nests may have been constructed by flying squirrels rather than gray squirrels, at least the nests differed considerably from the usual leaf nests built by gray squirrels. Measurements. The following figures are for speci¬ mens taken in Suffolk County during winter and spring. Four adult males average: weight, 72.3 grams (65.8- 76.1) ; total length, 231.5 mm. (221-237) ; tail, 99.5 mm. (96-101) ; hind foot, 31.5 mm. (30.5-34). Three adult females without embryos average; weight, 67.9 grams (63.1-73.3) ; total length, 234.3 mm. (230-238) ; tail, 100.0 mm. (97-105) ; hind foot, 31.7 mm. (30.5-33.5) . Individuals taken. 10 White-footed Mouse Peromyscus leucopus novehoracensis (Fischer) Distribution and habitat. The white-footed mouse probably is the most widespread or generally distributed mammal on Long Island, although perhaps not the most numerous. This species was taken on more traplines than any other small mammal; very often only one or two Peromyscus were caught in a given area, but this was enough to indicate its presence. It seems to be rivaled only by the masked shrew in its adaptability to the vari¬ ous environments here. Woods of every general type trapped were occupied by Peromyscus, including oak woods, mixed deciduous woods, maple swamps, pine barrens, and cedar swamps; dry, damp, mature, and second-growth woods were all inhabited. Peromyscus was numerous in the dry, sandy pine and oak woods of Suffolk County, and was defi¬ nitely the most abundant mammal in the drier barrens, under pitch pine and low bear oak (figure 3). Also this is the common mouse of the cedar swamps, generally avoided by the other mice. Other moist areas inhabited included sphagnum bogs near woods, and bushy zones around ponds and along streams in wooded areas. Open, treeless areas of various sorts were also oc¬ cupied, and accounted for approximately one-fourth of the total catch. These areas were predominately grassy and often included low shrubs. The areas were primarily in two categories — abandoned inland fields and various coastal habitats. In the coastal areas, trees were often far away and of no apparent significance to Peromyscus. Here the ground was usually dry and sandy, rarely marshy (fresh to salt), where these mice were collected. Little valleys and depressions among the sand dunes, such as are found at Hither Hills and at East Hampton Beach (figure 17), were found to harbor many wbite-footed mice. Fewer were found on the harsh outer dunes, a habitat of sand and beachgrass. Also Peromyscus was present in exten¬ sive, level, open areas at Napeague Meadows, on Shelter Island, etc.; beachgrass and other grasses, plus low shrubs (bayberry, beach plum, etc.), formed low cover in these areas. On the outer strip, Peromyscus was found in grassy and grassy-shrubby areas far from any trees; areas in¬ cluded Fire Island and the strip east of Moriches Inlet. Peromyscus was common in various types of vege¬ tation at the very end of Montauk Point, including the vicinity of the old lighthouse. Butcher and Butcher (1893, in their survey at Montauk, found this species present in woods, wet and dry meadows, and even under logs and rocks just above the tidal zone, in fact every¬ where except on the open uplands. Three white-footed mice were taken well out in salt marshes, one on the north shore (Mt. Sinai) and two in the Moriches Inlet area, but probably the mice had wandered out from drier, sandy areas. Salt marsh ap¬ peared to be the least preferred by Peromyscus of all the major habitats on the island. 30 Peromyscus has been collected on various smaller islands in this region besides Fire Island and Shelter Island. This species is present on Plum Island off Orient Point, and one collected by A. H. Howell in 1898 is in the U.S. National Museum. It is common on Fishers Island, and three collected by G. G. Goodwin in 1924 are in the American Museum of Natural History. For Gardiners Island, Richard Van Gelder (verbal commu¬ nication) found Peromyscus abundant in woods there during a brief period of trapping. White-foots are com¬ mon on Staten Island, where I have found individuals living in roofed-over bird nests. Remarks. This species, in common with many other rodents, fluctuates in abundance, being much more nu¬ merous some years than others. This may be why our trapping indicated white-footed mice to be rare or ap¬ parently absent in many seemingly favorable woodland areas. In the winter and spring of 1962, especially, these mice were notably scarce in woods throughout much of eastern Long Island. Roy Latham (personal communica¬ tion) reports a marked low at Orient during 1968. Usu¬ ally, many white-footed mice come into Orient houses during the winter. But they were so scarce in the winter of 1967-68 that not one came into Latham’s house, and he did not see any of these mice during 1968; Peromys¬ cus seemed to be virtually extinct locally. By the spring of 1969, however, Latham found Peromyscus to be on the increase again, becoming common in 1970. White-footed mice were usually trapped on the sur¬ face of the ground, but a few (mainly in winter) were caught underground in tunnels of the eastern mole; also catches were made several feet above ground on the trunks of leaning trees. A few individuals from tunnels were dirty white or grayish beneath, presumably a stain from contact with the soil. This species is found in many houses and other buildings in the more rural areas, but is not as much of a nuisance as the house mouse. An amusing instance of the wide variety of nest sites utilized was noted near Great Peconic Bay in late September 1960. I found an old wooden kitchen table standing on bare ground under some oak trees, evidently the abandoned remnant of a summer campsite. When the drawer was pulled out, four more or less fully grown white-foots looked up from a cozy nest within. One scur¬ ried out of a hole in the back of the drawer and then down to the ground, but the others remained in place in the nest as I shut the drawer. Helme (1902) on two different occasions on Long Island found as many as 16 individuals of this species in one nest. Reproduction. The reproductive season extended at least from March to October (female with embryos on March 29, 1961, lactating individuals in mid-October, 1960). First young juveniles out of the nest were trapped April 24 (1961). Average litter size based on 12 em¬ bryo counts (March 29 to August 23) is 4.3, with a range of three to six. Thirty females showed placental scars; individual counts ranged from three to 10, some of the higher counts apparently representing two litters. Males in evident breeding condition were collected from late March through most of September. Measurements. Seventy adult males from all seasons (fall, 1960 to summer, 1963) average; weight, 22.0 grams (16.8-27.8) ; total length, 175.3 mm. (155-197) ; tail 81.2 mm. (67-93) ; hind foot, 20.8 mm. (19.5-22) ; ear from notch, 16.9 mm. (15.5-18.5). Thirty-eight non¬ pregnant adult females in the same period average: weight, 21.2 grams (16.3-29.5); total length, 175.0 mm. (156-197); tail, 81.0 mm. (71-92) ; hind foot, 20.6 mm. (19-22); ear from notch, 16.8 mm. (15.5-18). Localities range from western Nassau County to Montauk Point. Individuals taken. 219 Meadow Mouse Microtus pennsylvanicus pennsylvanicus (Ord) Distribution and habitat. The meadow mouse, also commonly known as field mouse or meadow vole, is abundant in many grassy areas, from the outer dunes and salt marshes to inland fields and bogs. This is one of the most numerous mammals of Long Island, but it is not found in as many different habitats, and thus prob¬ ably does not equal in total numbers such species as the masked shrew, short-tailed shrew, and white-footed mouse. As far as is known, all meadow mice in the Long Island region today are the same as those on the mainland [M.p. pennsylvanicus) ; the recently exterminated subspecies on tiny Great Gull Island, M.p. nesophilus, is discussed at the end of this account. A surviving insular form just outside this region is M. pennsylvanicus provectus, con¬ fined to Block Island, Rhode Island, about 15 miles east of Montauk Point. Microtus is especially numerous (and far outnum¬ bers other mammals) in the extensive salt marshes which fringe the south shore bays and which also dot the edge of Peconic Bay (figure 6) and the north shore. In these marshes, areas of low grass, such as Spartina patens, are highly favored by the mice. Surface runways, cuttings, and nests of meadow mice are usually the only signs of mammals in the mole-shrew-mouse size range to be seen in the Long Island salt marshes. Occasionally we caught a few masked shrews or individuals of other species in the meadow mouse runways, but often Microtus appeared 31 to be the only small mammal present. As a general indi¬ cation of meadow mouse abundance in this habitat, 25 single-catch live traps which were set in a small area of salt marsh near Moriches Inlet (figure 16) for two nights and the intervening day in August 1962, and checked three times during this period, caught a total of 49 dif¬ ferent individuals. Dry, sandy, coastal areas, with assorted grasses and shrubs, also produce many meadow mice. They are com¬ mon on the tops of the outer dunes which border south shore beaches (figure 17). Here there is little vegetation save beachgrass [Ammophila breviligulata) , but it grows in fairly dense stands where the ground is reasonably level along the summits of the dunes. To judge from the abundant cuttings and other signs, Microtus feeds on the leaves, stems, and seeds of beachgrass; but gen¬ erally no distinct runways are to be found in this habi¬ tat, just the grass cuttings littering the bare sand where meadow mice are present. Signs of Microtus were found even on the summits of the steep moving or “walking” dunes in Hither Hills State Park near Napeague Harbor (figure 19). These dunes are much higher than the ocean beach dunes, and yet the mice were present, at least periodically, wherever there were stands of beach¬ grass growing on the tops of these otherwise bare peaks of sand. Microtus was common in fields island, especially where the grass was thick and luxuriant. It was often present in the same sandy field with Pitymys, but Micro¬ tus generally favored the denser stands of Andropogon or other grasses, while the more fossorial pine mouse was less restricted, and was apt to occur in the more open, sparsely covered field areas, and in adjoining woods as well. Fresh-water habitats included moist places adjacent to ponds and streams, cattail marshes, meadows, and sphagnum bogs; usually sedges or grasses were present. In the bogs, covered with shrubs, sedges, and the like, Microtus was locally abundant (figure 9) ; here it tun¬ nels through and under the sphagnum moss. In central and eastern Long Island the shrubby, sedgy bogs strik¬ ingly resemble those on the New Jersey coastal plain, where Synaptomys and Clethrionomys live, but on the island Microtus seems to be the only microtine present in such areas. This species was rare in wooded areas in general, and apparently nonexistent in the dry oak and pine woods covering much of the region. Only six were collected in areas that were classified as woods — three in a red maple swamp, three in grass under a stand of black locust trees. We collected specimens of Microtus on Fire Island and Shelter Island, and found abundant signs of these mice on Gardiners Island, but we did not work farther offshore. Frank M. Chapman collected this species on Gardiners Island in 1902 (specimens in American Mu¬ seum of Natural History). Eight specimens collected on Plum Island by A. H. Howell in 1898 are in the U.S. National Museum. This form also occurs on Fishers Island. Meadow mice are common on Staten Island, al¬ though landfill operations in salt marshes have destroyed much of their habitat. In Roy Latham’s collection there is an all-white meadow mouse from moist woods on Shelter Island, Food habits. Many kinds of plants are eaten by meadow mice. Food habits were not studied, except that we could hardly avoid noticing signs of feeding, such as the many cuttings of grass or sedges where these mice were present. Frequently noted examples of this included cuttings of cord-grass (Spartina alterniflora, S. patens) and bulrush {Scirpus sp.) in salt marshes, beachgrass (Ammophila breviligulata) on the ocean dunes, and broomsedge (Andropogon virginicus) in sandy fields. In a few salt marsh areas many cuttings were found of the odd succulent plant, glasswort (Salicornia sp.). Also we observed extensive diggings by meadow mice in stands of reed (Phragmites communis) on the barrier beaches, suggesting feeding on the rhizomes of this large plant. Meadow mice of the genus Microtus are noted for storing quantities of food, especially underground parts of plants belonging to the buckwheat, pulse, morning- glory, and composite families. In North America this habit is especially developed in certain northern and western areas. In March 1961, on Long Island, I found rather striking accumulations of the tuberous enlarge¬ ments from rootstocks of the groundnut or wild bean (Apios americana); apparently these had been gathered by meadow mice, rather than pine mice, which also store plant roots. The tubers, about an inch long when dry, resembled tiny sweet potatoes in general shape and color (figure 24). Groundnut grows in moist ground and thickets, often in or near areas frequented by Microtus, although it is not an abundant plant. Most of tbe stores were found east of Hauppauge in an area of ponds, and moist meadows near tbe northeast branch of the Nis- sequogue River. Piles of the tubers were found on the surface of the ground in grass, and associated with Mi¬ crotus runways. Two piles were ^examined closely; one contained over 100, the other about 175 tubers. One of the piles was next to a well-used Microtus runway, the other was at the end of a short spur connecting with a main runway. Remarks. Meadow mice are destroyed by a host of vertebrate predators. We found their remains in pellets 32 of several species of hawks and owls. Short-eared owl pellets were noted as especially numerous in salt marshes and fields along the south shore in the winter and early spring of 1961, and they contained many meadow mice. On one occasion a short-eared owl was observed to catch and eat a meadow mouse during the day on snow-covered dunes at Tiana Beach. There was also much evidence of red foxes preying on meadow mice in different areas of the island. Occasional hurricanes and other severe storms un¬ doubtedly drown many meadow mice in the shore areas, although the reduction in the numbers of these mice is only local and temporary. A prolonged and devastating east-northeast storm which reached its peak on March 6, 1962, washed over or broke through the outer barrier beach in several places between Moriches Inlet and Shin- necock Inlet. A week later along this same strip I noticed several dead meadow mice that had apparently drowned or died of exposure. Also some beachgrass and salt marsh areas were buried under a deep layer of sand probably carried by storm waves from the beach and outer dunes. One depression back of the dunes, formerly grassy and with a population of meadow mice, when revisited after the storm was found to be covered with 2 to 4 feet of new sand resembling a vast snow-drift. However, in the summer and fall of 1962 meadow mice were generally numerous along this strip of barrier beach. The experi¬ enced Long Island observer. Dr. William T. Helmuth (1954) wrote that meadow mice were still plentiful on eastern Long Island near the shore after the destructive 1938 hurricane, even though many storm-killed mice were noted. Reproduction. Average number of young based on 57 spring, summer, and fall embryo counts (Suffolk County) is 4.3, with a range of one to seven; three to five was usual (four the most frequent number). These figures are somewhat smaller than usually reported for this prolific species. Size of embryos ranged from small swellings 3 mm. in diameter to large embryos near birth about 30 mm. in length (crown to rump). A long breeding season was indicated, with some individuals probably breeding in winter. Embryo dates ranged from April 16 to December 15. Also, females which had already given birth (lactating, placental scars) were collected on the south fork in mid-April. Males in breeding condition were trapped primarily in the period March 22 to December 19, but two such males were also collected in central Suffolk County on February 7 and 8, 1962. Measurements. The following refers to specimens taken at various Suffolk County localities during all sea¬ sons of the year. One hundred and twenty-five adult males average: weight, 44.5 grams (28.4-75.7). Eighty adult males average: total length, 171.3 mm. (151-211); tail, 50.9 mm. (38-67); hind foot, 21.7 mm. (20-24). Fifty-eight adult females average: weight, 39.5 grams (28.3-65.4). Seventy-two adult females average: total length, 167.6 mm. (150-197); tail, 48.3 mm. (42-59); hind foot, 21.3 mm. (19-23). Individuals taken. 396 GuU Island Mouse. The Gull Island subspecies {Mi- crotus pennsylvanicus nesophilus V. Bailey) apparently became extinct 9 or 10 years after its discovery. Formerly it was considered a separate species, but it appeared to differ only in minor skull and pelage characters (skull shorter and wider, color of pelage darker) from the common meadow mouse of Long Island and the main¬ land. This form was evidently confined to Great Gull Island. This island, and nearby Little Gull Island are located at the entrance to Long Island Sound between Plum Island and Fishers Island. Great Gull is long and narrow, slightly over one-half mile long, and is about 2 miles east of much larger Plum Island, which in turn is about 11/^ miles off the end of Orient Point across Plum Gut. Plum Island mice, however, resemble the mainland form rather than nesophilus (Miller, 1899). Dutcher (1889) collected the first known specimen of nesophilus on August 6, 1888, and he described the birdlife, habitats, and dimensions of the two Gull Is¬ lands. Great Gull, 14 or 15 acres in extent, was composed of sand, with a shoreline and outlying reef of rocks; it was hilly (probably 25 feet high at highest point) , covered with coarse grass and some clumps of bushes, and had a small fresh-water marsh. Dutcher stated that Great Gull Island was purchased by the Government to serve as a garden for the keepers of Little Gull light¬ house, but it was so overrun with the mice that it was useless for that purpose. Little Gull Island was only about 100 by 50 yards in size, and consisted mainly of gravel and boulders, with a lighthouse. Dutcher also wrote that in the summer of 1888 com¬ mon terns {Sterna hirundo) were nesting in abundance on Great Gull Island, and that song sparrows {Melospiza melodia) were quite common, especially in and near the marsh on Great Gull. Marsh hawks {Circus cyaneus) visited the island and apparently preyed on the mice. Dutcher secured one of these hawks on Great Gull Island on August 12 and the stomach contents included the remains of a mouse (I presume nesophilus) ; he also stated that the marsh hawk was known as “mouse hawk” to the keepers of Little Gull Light. 33 More of these mice were collected on Great Gull Island in the years immediately following Butcher’s 1888 visit. Frank M. Chapman collected seven in July 1889 (skins and skulls in American Museum of Natural His¬ tory) and Basil Hicks Butcher returned to collect more specimens in 1890 (deposited along with type specimen in U.S. National Museum). Bailey (1898, 1900), who examined 15 specimens, published a description of the new mouse. Buring 1897, fortifications were erected on the island and in conjunction with this the entire surface of the island was disturbed, wiping out the original habi¬ tat. Reed (1898) described the rapidly changing environ¬ ment, and its effect on the tern colony, on Great Gull Island during the summer of 1897. On August 8, 1898, Arthur H. Helme and Arthur H. Howell visited Great Gull Island, searched every part of the island, noted the nearly complete destruction of the vegetation, and found no trace of the voles; they concluded that the Gull Island mouse was probably extinct (from Miller, 1899, who quoted notes furnished by Howell). At the present time the introduced house mouse is the only mouse on Great Gull Island as far as I know. Pine Mouse Pitymys pinetorum scalopsoides (Aud. and Bachman) Distribution and habitat. The pine mouse, or pine vole, spends much of its life underground, is highly adapt¬ able, and is one of the common mammals of Long Island. Perhaps it is exceeded in numbers only by the masked shrew, short-tailed shrew, white-footed mouse, and mead¬ ow mouse. The pine mouse has long been known to occur on Long Island, the type locality of scalopsoides; Audu¬ bon and Bachman (1841) obtained many Long Island specimens, on which they based their original description of this form, naming it Arvicola scalopsoides. Pitymys is found in most of New York State, but becomes progressively less common northward, where it is confined mainly to the humus of the forest. In southern mainland New York (such as lower Hudson Valley, Bela- ware River drainage), and on Long Island and Staten Island it is quite common, even abundant in places. On Long Island this mouse is widely distributed in the light sandy soils of fields and woods. Eastward on the island we found it common as far as Hither Hills on the south fork and Great Hog Neck on the north fork, and it is known to occur all the way to Montauk Point, where William Butcher collected this species in 1893 (specimen in American Museum of Natu¬ ral History). I was unable to find the pine mouse in sev¬ eral days spent collecting on Shelter Island. It is possible that it was overlooked on this large island, although Roy Latham, in a personal communication, writes that he has no record of Pitymys on Shelter Island either. He also writes that this species has not been recorded on the north fork east of Southold. Pitymys also appears to be absent from Gardiners Island and Plum Island, judging from the lack of records. Pine mice and meadow mice may occur together in the same field on Long Island (figure 11), but there are so many differences in habitat preference that more fre¬ quently only one is found in an area. The pine mouse is scarce in the immediate vicinity of the ocean front along the south shore. This fossorial species avoids the salt marshes (where Microtus abounds), although it is locally common at the dry edge of this habitat. Also Pity¬ mys was not found in grassy dunes and hollows behind the ocean beaches of eastern Long Island, even though Sorex, Blarina, Scalopus, Peromyscus, Microtus, Mus, and Zapus were found in such areas. We were unable to find Pitymys on the outer barrier beaches, where our collect¬ ing localities included Moriches Inlet, Tiana Beach, and Fire Island opposite Smith Point. Inland in Suffolk County, however, Pitymys finds more territory suitable to it than Microtus does. The pine vole tunnels under the leaf litter of oak and pine-oak woods; also it finds satisfactory the dry sandy ground of pine barrens, and old sandy fields where ground vegeta¬ tion is too sparse for Microtus. The latter often does pre¬ dominate where the grass is thick, and invariably does so where the ground is wet, as along streams and in marshes and bogs. Around some of the bays, pine mice may be found living right up to the edge of tidewater. We found this to be especially true in the vicinity of Great Peconic Bay, which penetrates far inland (figure 6). Here Pitymys was associated with Microtus in a dry sandy zone (often 30-40 feet wide) along the very border of the salt marsh under beachgrass [Ammophila breviligulata) , a tall un¬ identified grass growing in tussocks, groundsel-tree (Bac- charis halimifolia) , bayberry, and the like. The pine mice were found occupying subterranean burrows, and also surface runways in the grass. Windrows of old storm debris from the bay were conspicuous in this zone. This edge zone was a clear-cut area of overlap for the two microtines: Microtus alone occurred out in the wet tidal marsh, while inland, under pines and oaks, only Pitymys was found. This species is sometimes said to be poorly named and rarely found under pines. Probably it would be more accurate to say that Pitymys does not appear to be re¬ stricted to pines in any region, and it is also found in re¬ gions where there are no pines. In the Long Island pitch- 34 pine region Pitymys is actually rather common; in fact, it is only microtine present in this environment (fig¬ ure 3). Here it is found even in rather barren areas, with few green plants besides pitch pine, an undergrowth of bear oak, and for ground cover, bearberry and some scattered sedge. Pine mice were also taken in tunnels un¬ der fallen leaves in deciduous woods. At Hither Hills it was trapped under black oaks and white oaks (figure 21), many stunted and bent by the wind, near the shore of Block Island Sound. In old sandy fields, formerly cultivated and now covered with broomsedge {Andropogon virginicus) , com¬ posites, and other herbs, and sometimes scattered young pitch pine, wild black cherry, etc., pine mice are often abundant, and easily trapped because the holes leading to their tunnels are numerous and readily seen. Occa¬ sionally we found Pitymys in parts of a field where the ground was nearly bare, with only a scanty growth of grass, while the less fossorial Microtus and Sorex were rather closely confined to dense grass cover. The burrows appeared to be smaller than those of meadow mice, and in fields were more apt to be seen in bare, open ground, as mentioned above. In one small field surrounded by woods, where no meadow mice were taken, pine mice fre¬ quented surface runways in dense broomsedge. I had mis¬ taken the runways for those of Microtus before trapping, but they did seem to be narrower than those of Microtus. Blarina appeared to be the most regular mammal associate of Pitymys, both showing a somewhat similar habitat distribution, and often found in the same tunnels. The eastern mole was also quite characteristic of the same areas, pine mice often using the larger mole tunnels. Be cause of a more subterranean existence than Microtus, probably relatively fewer pine mice are destroyed by predators, although various birds, mammals, and snakes eat them. We did see remains of pine mice apparently killed and eatem by red foxes and other wild predators, and at Hampton Bays our pet cat occasionally caught them. Food Habits. This omnivorous and rather voracious vole is of economic importance, although its mainstay may be said to be succulent roots and other underground parts of wild plants, with seasonal variation. At times this species causes considerable damage in orchards and nur¬ series on Long Island and elsewhere by eating the bark and cambium of tree roots, girdling them. Some damage to potatoes and other root crops of farm and garden may occasionally be caused by Pitymys. Roy Latham, an ex¬ perienced potato farmer, writes (personal communica¬ tion) that on Long Island pine mice nibble on potatoes, carrots, rutabagas, and other roots in weedy fields but apparently do no notable damage. Moles, which make more conspicuous tunnels, are frequently blamed for the depredations of underground pine mice. Some green foliage is consumed, although the cut¬ tings are generally not as numerous and conspicuous as those of meadow mice. In a number of abandoned fields visited there was evidence of pine mice feeding on various herbaceous plants. For example, goldenrod (Solidago sp.) was being eaten during May in some fields in central Suffolk County; many tender shoots and small plants a few inches high were cut off and carried into the burrows. Also, daisy fleabane {Erigeron sp.) was similarly used, and cuttings were found in the burrows. In June in the same region it was noted that a cinquefoil {Potentilla sp.) had apparently been added to the diet, with fresh cut¬ tings in burrows frequented by Pitymys. Some individuals, especially in spring, gave off strong odors from feeding on pungent herbs, including wild onion {Allium sp. ). Seventy stomachs were saved but most have not been examined at the time of this writing. A few that were examined contained only plant material — the roots and green parts of grasses and other vege¬ tation. These mice are also inclined to eat animal food; quite a few specimens were destroyed in the traps, many presumably eaten by their own kind. Reproduction. Litters are small in Pitymys, and this was borne out in our Suffolk County information based on 14 embryo and placental scar counts. The average was 2.4, with a range of two to four. The season of re¬ production extended at least from early April (female with embryos on April 17, another nursing on April 18) to September (one recently nursing on September 27, several with placental scars in October) . Although lit¬ ters are small, Raynor (1960) found three litters and a pregnant female in a single nest at Manorville, Long Island, in April; it was assumed that at least one of the nest litters belonged to another female. Color. The only noticeable color variants collected were two adult females near Flanders in October 1960, which tended !o melanism, showing a dark grayish- brown (nearly blackish) hue on the upper parts, rather than the bright chestnut-brown characteristic of this species. Buffy specimens have been found at Mattituck and nearby Cutchogue, as described by Hatt (1930) from a skin in Roy Latham’s collection. Measurements. The following pertains to specimens collected in central and eastern Suffolk County at all seasons of the year. Thirty adult males average: weight, 25.2 grams (21.2-30.7) ; total length, 123.9 mm. (120- 129) ; tail, 23.1 mm. (19.5-26) ; hind foot, 16.5 mm. (15- 35 17.5). Thirty adult females without embryos average: weight, 25.5 grams (20.1-32.0) ; total length, 126.3 mm. (117-131); tail, 23.4 mm. (20-27) ; hind foot, 16.6 mm. (14.5-18). Individuals taken. 85 Muskrat Ondatra zibethicus zibethicus (Linnaeus) Muskrats are common throughout the Long Island area wherever their modest aquatic requirements are met. All the earlier references dealing with the mammals of the island, going back to early Colonial days, mention this val¬ uable rodent, and it was known to the Indians even earlier. Through the years, many thousands have been trapped for their fur. At the time of DeKay (1842) the skins sold for 25(i apiece and were extensively used in the man¬ ufacture of hats. Muskrats are still present in Brooklyn (Jamaica Bay) and Queens, and on Staten Island; east¬ ward they extend to Montauk Point, Orient Point, Shelter Island, and Gardiners Island (we saw several on Gardin¬ ers Island in 1962). Their most extensive habitats, and where they flour¬ ish, are the great salt and brackish tidal marshes which encircle the bays along the south shore (including Fire Island). The similar but more scattered marshes around the Peconic bays and on the north shore also harbor many muskrats. Inland, muskrats live in almost all fresh-water localities, especially in marshy areas along the major streams (figure 5), but also in the many ponds in the pine barrens (figure 4) and elsewhere, and in bushy sphagnum bogs, old cranberry bogs, and wooded maple swamps. The overall muskrat population is much smaller than in the past, because of the many acres of aquatic en¬ vironment which have been drained or otherwise lost. Thus Hamilton (1949) remarked that in the winter of 1919, when the rats brought up to $4.25 a pelt, over a thousand dollars worth of the furs were harvested from the salt marshes and creeks (Flushing Meadows area in Queens) that later became the site of the 1939 World’s Fair. Roy Latham (personal communication) used to be able to trap about 80 muskrats in a season on the agri¬ cultural north fork at Orient, an area where they are now scarce. Hatt (1935) described an unusually dark muskrat (black phase) collected by Roy Latham at Orient on December 28, 1929. Formerly, Latham (personal com¬ munication) saw or obtained about two individuals in the black phase each trapping season at Orient. There seem to be relatively few muskrat houses, or lodges, compared with the many muskrats obviously pres¬ ent in the various areas. Many must make their homes in burrows dug into banks. We did see some houses in marshes, both tidal and inland, also on ponds, and in overgrown bogs. House Mouse Mus musculus Linnaeus Distribution and habitat. This familiar species, intro¬ duced from the Old World about two centuries ago, is numerous in and around buildings in cities and towns and on farms, where it lives in association with man. Since little field work was done in such situations, we did not collect many house mice. This species is sometimes found in fields and less often in woods in this region; also it lives along the seashore and on outlying islands in vari¬ ous parts of its worldwide range, including the Long Island area. In winter, at least in 1960-61, feral or wild-living house mice seemed to be rather numerous and widespread on the outer ocean dunes along the south fork of eastern Long Island. At East Hampton Beach, in December, sev¬ eral were trapped on the outer dunes (figure 17) just back of the beach in nearly pure stands of beachgrass (Ammophila breviligulata) with some scattered herbs, chiefly composites. This was during severe weather, a period of low temperatures followed by unusually heavy falls of snow. I do not know how dependent this hardy colony was on the presence of buildings; the nearest in¬ habited houses were about one-half mile inland, although several summer cottages were somewhat closer. The beach individuals were probably feeding on seeds (and perhaps green parts) of beachgrass; several were trapped on the sand next to little piles of the seed heads. Microtus was the only other small mammal col¬ lected on the outer dunes at East Hampton Beach; the two species appeared to be living closely together on the same ground. Some of these outdoor house mice were breeding during the winter, and a female on December 8 contained five small embryos. House mice also were found living on the barrier beach strip near Moriches Inlet in rather wild, deserted dunes country far removed from any buildings. Here they were considerably outnumbered by Microtus and Pero- myscus. One January day in this area we saw a sparrow hawk [Falco sparverius) feeding on a freshly caught house mouse. House mice are present on remote Great Gull Island (specimens in American Museum of Natural 36 History), the island formerly inhabited by the Gull Island vole (Microtus pennsylvanicus nesophilus) . Most trapping in natural habitats inland, away from buildings, farms, and the like, gave only negative results for this species. One was collected in a sphagnum bog bordering a cold stream, along with a large number of meadow mice. Individuals taken. 14 Norway Rat Rattus norvegicus (Berkenhout) Distribution and habitat. The Norway rat, also known as brown rat, house rat, barn rat, and even water rat in some localities, is probably our most destructive and hated mammal. It is usually associated with man and is often extremely abundant in and around barns and other buildings. Hundreds of rats were seen at a time at certain Suffolk County dumps visited after dark, and rat burrows were noticed on duck farms and even in the better residential areas. Rats were encountered near water, both salt and fresh, often at a considerable distance from human habitations. On Long Island this species is distributed from the Brooklyn waterfront to the rocky tip of Montauk Point. Signs of rats are frequently noted near inlets and at rock jetties on the ocean front. At the jetty which hugs the west side of Shinnecock Inlet (figure 12) rats make their home among the rocks, and I have observed them boldly active on the nearby open beach during daylight hours. When pressed too closely, they take cover among the rocks. Rats appeared to be present throughout the year here, and they were abundant even in January (noted in 1963). Their numerous footprints formed beaten trails in the sand next to the jetty and among nearby sand dunes. On the bay side of the inlet there are fishing stations and assorted buildings which prob¬ ably shelter many other rats of the area, and where the jetty rats may take refuge during storm tides and when¬ ever necessary. Rats are also found at Montauk Point in a some¬ what similar situation. They live among the rocks and boulders at the base of the cliff at the very end of the point, and judging from the many tracks, are present here even in the bitter, windy midwinter season. Turrell (1939) mentions that rats are occasionally seen in broad daylight foraging on the beaches of the Smithtown Bay region on the north shore. A few individuals were trapped in the Great Peconic Bay region in salt and brackish marshes and in adjacent dry grassy areas (figure 6). Norway rats are said to be a problem in some Peconic localities (and probably else¬ where) when exceptionally high tides or storms drive them out of the marshes, at which times they appear in abun¬ dance in and around houses. Rats are also present in fresh-water areas, and individuals were seen foraging along the shores of ponds in the evening, tracks were seen along streams in red maple swamps, tracks were numer¬ ous around waterways in certain parks, and the like. A few even frequent small streams in the dry pine-oak woods. Norway rats inhabit muskrat burrows in some of the marsh and pond areas. Norway rats are present on most of the eastern is¬ lands, including Shelter, Gardiners, Plum, and Fishers, and, reportedly, even Great Gull Island. Food habits. Rats eat all sorts of animal and vege¬ table food, including carrion, garbage, and food stored by man. In the shore and marsh areas of Long Island, rats probably feed extensively on certain readily avail¬ able forms of animal life (both living and dead). Rats living among rocks at the ocean, such as inlet jetties and boulders at Montauk, may subsist mainly on dead birds, fish, and other marine life washed ashore, and probably also eat the remains of bait and fish discarded by fisher¬ men. One January morning I saw a Norway rat feeding on the carcass of a herring gull on the beach beside the Shinnecock Inlet jetty. Audubon and Bachman (1851) told of Norway rats along the East River which were re¬ ported to dig up and feed on a small (%-inch long), thin-shelled species of clam which was then abundant in sandy places below high-water mark. Individuals taken. 8 Black Rat Rattus rattus Linnaeus The black rat (also called ship rat or roof rat), like the Norway rat, lives in human habitations and was ac¬ cidentally introduced from the Old World. However, the black rat arrived much earlier than the Norway, prob¬ ably with the first colonists. The latter species is said to have been introduced to the seaboard states during or about the time of the Revolutionary War; it spread rapidly in the Northeast, replacing the black rat, which seemed to disappear entirely from most areas where it had been numerous. The two species of Rattus cannot be distinguished by color alone, regardless of common names; melanic (black) Norway rats are not rare, and also R. rattus has 37 different color phases or variants, some closely resembling typical Norways in color. Such features as the longer tail, larger ears, and smaller body size of R. rattus are more reliable. The present status of the black rat in the Northeast is not clear; perhaps it may be described as either very rare or very local. This species may have been abundant on Long Island at one time, as it seems to have been in some other areas of New York and neighboring states, but practically nothing is known of its history here. It has certainly long since disappeared from most of the island. Very likely the black rat in this region has con¬ tinued to exist in city waterfront areas (e.g. Brooklyn), where its numbers are maintained by new arrivals brought in by ships. Naturalists rarely visit or collect in such areas. Helme (1902) wrote that “many years ago” some specimens of Rattus rattus were caught in a stable in Brooklyn. The only existing Long Island specimen I know of was collected at Douglaston on the north shore of Queens near the Nassau County line, March 13, 1921 (Dr. Fisher). It is a male in the black color phase, preserved as skin and skull at the American Museum of Natural His¬ tory, No. 42978. Kieran (1959) stated that this species is far behind the house mouse and Norway rat in abun¬ dance in all five boroughs of New York City, but that recent collecting by professional exterminators has re¬ vealed that sizable numbers of Rattus rattus are present. I do not know if any specimens collected in this manner have been preserved. Meadow Jumping Mouse Zapus hudsonius americanus (Barton) Distribution and habitat. The meadow jumping mouse is rather rare and local on Long Island; it seems to be more numerous on the adjacent mainland. This species has been collected from Brooklyn and Queens east to Montauk Point, and is still found thinly spread throughout the full length of Suffolk County, but at the present time appears to be common only near the eastern end of the island. There jumping mice are found in very dry, as well as moist, open areas with grasses and other low vegetation. They thrive in sandy areas near salt water marshes and bays, and in the grassy hollows just back of the ocean dunes. But even though we found these mice in places along the very edge of salt marshes, they rarely venture out into this habitat. Fourteen individuals were collected in dry, sandy areas east of Amagansett, in the region of Napeague Meadows and Hither Hills (figure 18). The vegetation consisted of a variety of grasses, including much beach- grass {Ammophilia breviligulata) , and also many shrubs and vines, especially bayberry (Myrica pensylvanica) , poison ivy {Rhus radicans) , and wild rose {Rosa Caro¬ lina). One was collected on Shelter Island in similar habitat, adjacent to a salt marsh (figure 25). Such areas of excessively drained ground (nearly pure sand), often remote from any standing fresh water, appeared to be the most productive habitat for Zapus on eastern Long Island judging from our trapping results. The remaining four jumping mice were collected in moist, somewhat boggy situations with grasses and sedges. Sorex cinereus and Microtus pennsylvanicus were com¬ mon in all of the Zapus collecting areas, but other small mammals were scarce. In the pine barrens of Suffolk County, jumping mice are found in some of the more luxuriant areas near water, and we saw one or two among shrubs and grass on the edge of a pond near Flanders. Butcher and Butcher (1893) reported Zapus to be the “least numerous” species of mouse at Montauk, but they found it widespread there, present in swamps and wet and dry meadows, and also on the Montauk Downs. Eight meadow jumping mice (and other mammals) col¬ lected by the Butchers at Montauk are in the U.S. Na¬ tional Museum. Although found at Montauk and on Shel¬ ter Island, Zapus has not been recorded from the more outlying eastern islands, such as Gardiners Island and Plum Island, as far as I know; nor do I know of any records for Fire Island, although there appear to be ex¬ tensive areas of suitable habitat there. This species is a resident of Staten Island, where it has been collected in several localities. Reproduction. Several breeding females were col¬ lected in June: one with six tiny embryos on June 8, one lactating and with four placental scars on June 22, and two lactating and with placental scars on June 27. Measurements. The following refers to specimens taken on the southern fork of eastern Long Island and on Shelter Island. Nine adult males without hibernation fat (May and June specimens) average: weight, 15.0 grams (11.6-17.3); total length, 201.3 mm. (194-205); tail, 116.3 mm. (101-128); hind foot, 27.9 mm. (26-29). Five nonpregnant adult females without hibernation fat (June and August) average: weight 17.4 grams (14.7- 19.3); total length, 208.4 mm. (193-227); tail, 122.2 mm. (112-134); hind foot, 27.8 mm. (26-30). The hea¬ viest individual, weighing 24.9 grams, was a nonbreeding female with much fat on October 19. Individuals taken. 18 38 Common Dolphin Delphinus delphis Linnaeus This graceful and beautifully marked dolphin, which reaches a maximum length of about 81/^ feet, frequents New York waters on occasion, and is fairly common well offshore. Schevill and Watkins (1962), who point out that Delphinus delphis is known as the saddleback por¬ poise by fishermen (because of the shape of the dark dorsal area), encountered a group of 20 individuals and made sound recordings in August of unstated year some 60 miles south of Martha’s Vineyard, Massachusetts; in relation to Long Island, this general location is the open sea several score miles southeast of Montauk Point. Ed¬ wards and Livingstone (1960) observed two in mid-winter 1959, feeding on fish escaping from an otter trawl net, depth of water 200 feet, lat. 39°48' N, long. 72°28' W (about 65 miles off the south shore of Long Island). The New York coastal records (sightings and strand- ings) now available are more numerous than when Stoner (1938) documented a remarkable movement of common dolphins far up the Hudson River in October 1936. Roy Latham has numerous records of this species in the waters about the east end of Long Island; he writes (per¬ sonal communication) that it is recorded every year and is more frequent than Tursiops. Some of his locality rec¬ ords are as follows: East Marion, January 24, 1947; Orient, Gardiners Bay, May 26, 1947; Robins Island, Great Peconic Bay, September 11, 1947; Plum Island, July 14, 1951. Latham also has records for Shelter Island and Gardiners Island. Specimens in Latham’s collection include one from Orient, Gardiners Bay, April 1928, and one from Cutchogue, Peconic Bay, 1923. On December 6, 1960, an individual 7 feet 2 inches long stranded alive on the eastern shore of Staten Island and was transported to the New York Aquarium, where it lived 4 days (Ray, 1961). Long Island specimens (skulls and skeletal material) of the common dolphin, presumably strandings, in the American Museum of Natural History include the follow¬ ing: Montauk Point, September 9, 1931; Jones Beach, August 1938 (H. C. Raven) ; Fire Island Beach, Novem¬ ber 1, 1908 (skull found on beach) (H. C. Raven). Also in the collection are the Staten Island individual men¬ tioned above and specimens from the Hudson River, 1911 and 1936. Another record for Montauk (Dead Mans Cove) is a skull in the Cornell University collection dated January 2, 1951 (M. Gordon). True (1889) listed com¬ mon dolphins taken in the last century near both ends of Long Island, including New York Harbor and off Block Island, Rhode Island (specimens in U.S. National Mu¬ seum) . This species, common enough in the North Atlantic, is a particularly conspicuous inhabitant of many warm- temperate regions, such as the Mediterranean Sea, where it is regularly seen playing about ships and leaping high out of the water. This is the dolphin most frequently de¬ picted and written about in the ancient art and literature of the Old World; the ancients of the Mediterranean area also knew the next species, S. coeruleoalbus, judging from surviving portrayals of it on vases and other works of art. Striped Dolphin Stenella coeruleoalbus Meyen This little-known pelagic species, which is about the same size as the common dolphin, but with a different color pattern and heavier beak, has not often been re¬ ported on the east coast of North America. The striped (or Gray’s or euphrosyne) dolphin infrequently comes near shore or strands, but it is not really rare. A group of about 200 was noted passing by in deep water over the lower Hudson Canyon about 140 miles out from New York (south of Long Island) in August of unstated year, when sound recordings were made (Schevill and Watkins, 1962). At least two striped dolphins have been found on Long Island. A male weighing about 85 lbs. was found alive on the beach at the foot of the Throgs Neck Bridge on May 6, 1967, and was transported to the New York Aquarium, but it had a broken neck-vertebra and died on May 9, 1967 (Robert A. Morris, personal communica¬ tion). This locality is on the north shore of Queens, where the East River meets Long Island Sound. The skeleton is preserved at the American Museum of Natural History. Another one (skull at the same museum) was found at Bellport, on the south side of Suffolk County, in May of 1929 (P. White). A previously published locality record which is close to eastern Long Island — some 30 miles northeast of Montauk Point — is Narragansett Town Beach, Rhode Island, where a male 7 feet 9 inches in length stranded on December 5, 1966 (Cronan and Brooks, 1968). The taxonomy of Stenella, ar large genus of dolphins, is poorly known; it is being studied by F. C. Fraser of the British Museum according to Rice and Scheffer (1968). Striped dolphins of northern oceans, including our Atlantic coast, have been called Stenella euphrosyne (=S. Styx), but apparently just one worldwide species of striped dolphin exists. There are two strongly marked 39 races, however; thus local specimens may be designated S. c. euphrosyne. Another member of this genus, the spotted dolphin {Stenella plagiodon, also known as S. pernettyi or S. dubia) is a southern species which has not been recorded in the Long Island area. The island is perhaps too far north to be visited by this species, but it has been re¬ ported as far north as southern New Jersey. Voice re¬ cordings were made from a group of two or three dozen spotted dolphins 60 miles off Cape May, New Jersey, in August of unstated year (Schevill and Watkins, 1962) ; this location is roughly 110 miles south of Long Island. Bottle-posed Dolphin Tursiops truncatus (Montagu) The bottle-nosed dolphin, a large grayish species which may attain a length of 12 feet, is a common coastal species of the Atlantic, and is especially abundant along the shores of our southern and mid-Atlantic states. It is migratory along the coast and visits the Long Island area, including Long Island Sound and various bays, in the warmer months. Roy Latham (personal communication) writes that this species is recorded every year at various stations along the eastern Long Island coast from May 28 (year? ) , Greenport, Long Island Sound, to November 17 (1937), Bay View, Peconic Bay, and also in 1932 a single one was present as late as December 2, at Southold, Peconic Bay. He has records of Tursiops, as well as Delphinus, at Shelter Island and Gardiners Island. Latham (1954) re¬ ported on an adult female Tursiops truncatus 9 feet long, and bearing an unborn young 43 inches in length, which stranded and died on the Sound beach at Orient, June 11, 1953. Specimens (chiefly skulls) in his collection include the above female and young, and also one from Mattituck, Peconic Bay, May 1928, and another from Shelter Island, Gardiners Bay, 1921. Helme (1902) wrote that dolphins in general, either this species or Delphinus, may be seen as early as April and as late as December, and are plentiful in Long Island Sound from June to late October. Some of the reports we received in the 1960’s of “dolphins” or “porpoises” observed in Gardiners Bay, Peconic Bay, and lower New York Bay may refer to the present species. On October 17, 1960, we found a bottle-nosed dolphin, which was about 8 feet long and had been dead for several days, on Tiana Beach, ocean side of Suffolk County. A highly unusual account by Stone (1964) tells how the author, during many July and August days in 1945 and 1946 when she was a girl of 13 and 14, played with a group of about six dolphins in Long Island Sound near a boat 2 or 3 miles off the north shore of Suffolk County. By the end of the second summer the play developed to the point where on several occasions she grasped a fin of a dolphin and was actually towed around on the sur¬ face. Late in the day the boat was sometimes taken into Port Jefferson Harbor, and the dolphins would follow far into the harbor. These animals were apparently small (young) Tursiops truncatus. Two specimens collected near western Long Island are in the American Museum of Natural History: one (skeleton preserved) from Pelham Bay, New York, main¬ land side of extreme western Long Island Sound, August 16, 1906, (W. Dolan), and another (skull only) from Navesink River, New Jersey, May 25, 1960. A group of five to eight Tursiops was observed in the tidal Shrews¬ bury River, New Jersey, in an area 31/^ miles from the mouth of the river at Sandy Hook Bay, from April 17 to June 16, 1965, (Clark, 1965). The Shrewsbury River, its tributary the Navesink, and Sandy Hook Bay are not indicated on the map but are just south of Lower New York Bay on the west side of Sandy Hook. A few have over-wintered in the Navesink River (Ulmer, 1961). Dolphins, especially those of the genus Tursiops, are called “porpoises” in America by many cetologists as well as by most mariners and landsmen. The word “dolphin,” although an old name for small beaked cetaceans, is con¬ fusing in that it is also used for two species of fast, color¬ ful, warm-water ocean fishes (genus Cnryphaena) ; the species C. hippurus grows to about 5 or 6 feet in length and is popular with offshore sport fishermen as far north as our region. Tursiops truncatus is the species of dolphin or porpoise which has become especially familiar in re¬ cent years as a performer in public oceanariums and as a much written-about experimental subject in studies of echolocation, intelligence, etc. Dolphins and porpoises feed largely on fishes, squids and other marine animals. The Atlantic white-sided dolphin {Lagenorhynchus acutus) is a northern species found at times in schools as far south as the ocean waters off Cape Cod. Apparent¬ ly it is unknown in the Long Island area, although it would be possible to find a dead or stranded one on this coast. The nearest record I know of is a dead one found in the Town of Naragansett, Rhode Island, on July 22, 1967, (Cronan and Brooks, 1968). Killer Whale Orcinus orca Linnaeus The killer whale is a large (males up to 30 feet long) relative of the dolphins which is noted for its attacks on 40 other marine mammals. Prey includes seals and porpoises, many of the larger cetaceans ranging in size up to the huge but defenseless whalebone whales, and also school¬ ing fishes, squids, and seabirds. It has a strikingly tall dorsal fin, up to 6 feet high in the large adult males. Grampus, firmer, and killer are among the various com¬ mon names that have been applied to this species. The range of the killer whale is essentially worldwide, but it is most numerous towards the Arctic and Antarctic re¬ gions where large warm-blooded prey is plentiful. Few killer whales have been captured or stranded along the Atlantic coast of the United States, although they are sighted offshore fairly regularly at least as far south as New Jersey, where two stranded specimens have also been found. A small number have been found beached in recent years on the southern coasts of nearby Rhode Island and Massachusetts. For Long Island, Roy Latham (personal communication) reports that a killer 24 feet in length stranded alive on a clam flat in Orient Harbor, Jan¬ uary 11, 1944, and then died. Cook and Wisner (1963), in an exaggerated but profusely illustrated account of the killer whale, reported on one which followed a 28-foot fishing boat one July day in 1958, from 35 miles off Mon- tauk Point towards land to about 15 miles off the point; included in the book are several photographs of this indi¬ vidual, apparently a large male. Orcinus was probably more abundant in earlier days when various other ceta¬ ceans and seals abounded. DeKay (1842) mentioned see¬ ing killer whales off the coast of Long Island on several occasions, but asserted that they were formerly more numerous here, at a time when right whales were also abundant. Gray Grampus Grampus griseus G. cuvier The gray grampus, white-headed grampus, or Risso’s dolphin, is about the size of the bottle-nosed dolphin, but is beakless, with a blunt rounded snout, and a rather high, recurved dorsal fin. This species is widely distributed in the oceans of the world, but is not very well known. It appears to be a deep-diving form which feeds on cepha- lopods (squids and relatives). In the western North At¬ lantic this small whale has been recorded, though rarely, from Massachusetts to the coast of New Jersey. Schevill (1954) made observations suggesting greater abundance than previously suspected off our coast; on August 20, 1952, a group of over 60 grampus was watched from a ship near Lat. 40°00' N, Long. 71°31' W. The animals consorted in small bunches of four to six within the larger grouping, and they were quite playful, often leap¬ ing out of the water. The position indicated is roughly 75 statute miles out to sea in a southeasterly direction from Southampton, Long Island. Blackfish Globicephala melaena Traill Blackfish, or pilot whales, which are related to dol¬ phins and killer whales, are almost entirely black, have a high, bulging forehead, and frequently travel in large migratory schools. Maximum length is about 28 feet. Squids, cuttlefish, and various fishes are eaten by pilot whales. This is one of the more numerous cetaceans, and it is common offshore in this region, especially during the warmer months. Helme (1902) and other observers have mentioned frequent sightings of blackfish off the coast of eastern Long Island. There are also records for western Long Island and Long Island Sound. Strandings of pilot whales are frequently reported, and occasional mass live strandings of many individuals at one time are known to occur, although in this area, at least, usually only single individuals come ashore. On May 13, 1960, a live pilot whale, a female 12 feet 9 inches long, was found stranded alive on Brighton Beach, Coney Island, Brooklyn, about half a mile from the New York Aquarium; it was retrieved and lived at the aquarium for 29 days (Ray, 1961). At Atlantic Beach (west end of Long Beacli strip), Nassau County, a male 13 to 14 feet long came ashore on April 15, 1966, and died later the same day (Robert A. Morris, personal communication). An immature female was collected near Massapequa, south shore of Nassau County, on April 28, 1960, (skeleton preserved in American Museum of Natural History). That blackfish penetrate extreme western Long Island Sound and even the East River is indicated by one which stranded in Flushing Creek, Queens, off the upper East River, June 13, 1944, (Anon., 1956) and others which have grounded on the Sound shore of the Bronx. The name “blackfish” is also used by fishermen for a popular salt-water fish less than 3 feet long, the tautog {Tautoga onitis) . The Short-finned blackfish {Globicephala macrorhyn- cha) may visit; difficult to distinguish from the common blackfish, it ranges north at least as far as New Jersey. Harbor Porpoise Pliocoena phocoena Linnaeus The harbor porpoise is smaller than the dolphins, reaching a maximum size of 6 feet. It seldom leaps clear 41 of the water, and the head is blunt and rounded, lacking the beak characteristic of the dolphins. Phocoena fre¬ quents coastal waters and enters inlets, bays, and even rivers; New York is near its southern limit of abun¬ dance on the coast. Miller (1899) termed this species the commonest cetacean in New York tidal waters. Per¬ haps this is still true today, although for a long time now the porpoise has been gradually declining in numbers in our area. Apparently the harbor porpoise no longer forms large schools here of up to 100 or even “hundreds” of in¬ dividuals such as were occasionally reported late in the last century. There are numerous published reports of harbor por¬ poises in Long Island Sound. Rowley (1902), for exam¬ ple, stated that Phocoena was then very common in sum¬ mer in this body of water, and that schools could be seen on almost any clear day, while Goodwin (1935) also mentioned the presence of this species in the Sound. Tur- rell (1939) wrote that harbor porpoises enter Smith- town Bay on the north shore of Suffolk County in large schools in summer. We received verbal reports for the 1950’s and 1960’s of sightings of porpoises in summer in Long Island Sound and also Great Peconic and Little Peconic Bays, although some of these reports may refer to dolphins (also called “porpoises”) rather than Pho¬ coena. For the Orient region, Roy Latham (personal com¬ munication) writes that Phocoena is recorded every year at all stations, but that the big schools which were re¬ corded yearly up to 1920 are no longer seen. The last sizable group recorded in this area, in Long Island Sound near Orient, was 25 individuals on December 7, 1921. Latham also reports that one came ashore, chilled, in the Sound at Orient on January 12, 1943; two specimens from the Orient region are in Latham’s collection. Gil¬ bert S. Raynor informs me that a harbor porpoise was found dead on the beach near Orient Point (Gardiners Bay side) about 1960. Harbor porpoises may still be seen on occasion in Lower New York Bay and in Raritan Bay (waters bounded by Long Island, Staten Island and New Jersey). I have seen small groups in the summer months (during the 1940’s) in this area. Porpoises also ascend rivers in this general region (New Jersey, Connecticut, and Hud¬ son River), although I have no definite reports of this for the small rivers of Long Island. On the south shore of the island stranded individuals have been collected during various months of the year from Brooklyn to Montauk Point (specimens from Montauk Point in Ameri¬ can Museum of Natural History and N.Y. State Museum). Anthony S. Taormina (personal communication) has many Long Island records, including photographs, of both the harbor porpoise and the harbor seal. Of interest is DeKay’s (1842) account of this spe¬ cies. He related that, although still common in his day, the harbor porpoise was formerly so abundant on the shores of Long Island that it was avidly pursued for its oil (from the blubber), and also for its durable leather (from the hide). He described in detail the procedure used in seining schools of harbor porpoises at the east end of Long Island based on a paper published in 1792 by E. L’Hommedieu. At the time of DeKay’s publication harbor porpoises were evidently being killed in Long Island Sound, on the Connecticut coast at Stratford (op¬ posite Port Jefferson), according to Linsley (1842), who wrote: “Numbers of the common (harbor) porpoises are taken in this town for the sake of the oil, which is usually from three to seven gallons.” White Whale Delphinap terns leucas (Pallas) The white whale, or beluga, a small whale usually 11-14 and rarely up to 18 feet in length, and which (after early life) is completely milk-white in color, inhabits arctic and subarctic seas including the northern North Atlantic. It occurs regularly in the Gulf of St. Lawrence and adjacent bays and rivers. But southwards this spe¬ cies is rare, occurring as a straggler only as far as Cape Cod, Massachusetts, according to Miller and Kellogg (1955) and other recent authorities. There are two cen¬ tury-old records for that area — outer Cape Cod — hut no recent records (Waters and Rivard, 1962). Although never collected on Long Island as far as I know, Delphinapterus has probably visited the area at least once judging from the following sight report. A small whale, white in color, which was believed to be this species was observed in Long Island Sound off the north fork for 4 days in June 1942 (Roy Latham, per¬ sonal communication). Latham first observed the whale off Orient, noted that it was moving west, and estimated the animal to be 10-12 feet long. He immediately alerted a friend in Mattituck who observed a white whale there the following day, presumably the same individual, 25 miles west of Latham’s first sighting; this second ob¬ server reported the whale moving east. In earlier years Roy Latham had heard of fisher¬ men seeing white whales in the Sound, but he was un¬ able to establish a record for one prior to 1942. Latham has considered the possibility of albinos of other species of whales being responsible for at least some of the 42 sightings. Albinism in whales is rare but has been re¬ ported in a number of species ranging in size from the huge sperm whale {Physeter catodon) to the little har¬ bor porpoise {Phocoena phocoena) ; Tomilin (1967) wrote that rare instances of partial to complete albinism (one specimen out of tens of thousands) occur in the harbor porpoise. Also, increasing whiteness comes with age in some species, and old individuals of the bottle¬ nosed whale {Hyperoodon ampullatus) may be encoun¬ tered which are yellowish-white in color. For Latham’s sight record, however, his original field identification, the estimated size of the animal, and his familiarity with the common small cetaceans (dolphins, porpoises), as well as the white coloration, all strongly indicate Del- phinapterus leucas. The bottle-nosed dolphin (Tursiops), although rarely over 10 feet long here, overlaps Del- phinapterus in size, but the latter is shaped differently and lacks the conspicuous dorsal fin of Tursiops. Sperm Whale Physeter catodon Linnaeus This species is the largest of the toothed whales, the males, which are much larger than the females, at¬ taining a length of about 60 feet. The massive squarish head and relatively small lower jaw are distinctive. Food of this deep-diving whale is primarily squids, also sharks, skates, and fishes. This is essentially a whale of tropical and warm temperate seas, but it is migratory, and some of the males go far to the north in summer. According to DeKay (1842) and some other ac¬ counts sperm whales were formerly abundant “along” or “on” our coast. Also, some sperm whales were taken in Long Island waters during the early days of American whaling (Murphy, 1918). But unlike finback whales and right whales, probably the sperm whales only occasion¬ ally came close enough to be sighted from shore, even in the early days of whale abundance. Sperm whales gen¬ erally avoid shallow waters of sandy coasts, although they may approach shore closely where the water is deep, such as near the Azores. Today, sperm whales are very rare near the shores of Long Island, and are uncommon offshore although occasionally reported from ships out at sea. Some in¬ shore records are as follows. On March 13, 1928, a young male 18 feet 31/^ inches long strayed into New York Harbor (Upper New York Bay), where it was captured alive, but it died soon after being towed to Gowanus Canal, Brooklyn (Raven and Gregory, 1933). A female 39 feet long became stranded on Great South Beach (Fire Island) opposite Bellport on February 28, 1918 (Murphy, 1918) ; this individual came ashore alive during the night but died before daylight as the tide receded. On December 9, 1894, a sperm whale 40 feet long was captured in Fishers Island Sound, between Fishers Island and the Connecticut coast. No very re¬ cent Long Island records have come to my attention, but according to Cronan and Brooks (1968) the only au¬ thentic record for Rhode Island is a 14 foot 5 inch indi¬ vidual which washed ashore in Charleston, Quonchontaug Beach, February 20, 1967; this locality is in the south¬ western corner of Rhode Island, near eastern Long Island (about 12 miles east of Fishers Island), New England whalers, seeking the sperm whale pri¬ marily, ushered in a famous era of American whaling. They extended the fishery from coastal waters, where the right whale had been pursued, out on to the open sea, eventually leading to lengthy voyages to other oceans. Sag Harbor on Long Island also became one of the leading deep-sea whaling ports during this period, along with New Bedford and Nantucket, Massachusetts, and New London, Connecticut. This enterprise began early in the eighteenth century, and finally declined after the middle of the nineteenth century. This period was for the most part still one of harpoons thrown by hand from open boats which were carried to sea by a mother ship, before modern methods used in pursuit of the fast blue and fin whales were put to use. Sag Harbor was established as a whaling port be¬ fore the Revolutionary War and its peak whaling years came in the late 1830’s and in the 1840’s (biggest year — 1847); a rapid decline followed the peak years, and it is said the last whaling ship set sail from this port in 1871. It was not until 1785 that the first successful trips were made from Sag Harbor by ships fully equipped for long (South Atlantic) whaling voyages (Willey, 1949). Before this time the ships made short voyages lasting no more than a few weeks, often near shore, then returned to Sag Harbor to process the blubber for oil. According to Howell (1941) in this period the ships ventured out only two or three hundred miles from port, usually to the southeast of Montauk. Although I am uncertain which species of whale comprised the bulk of the catch on these early offshore trips, such a voyage in summer would have taken whalers into an area of sperm whale abundance, as described below. Although the sperm whale is almost cosmopolitan, there were favored feeding areas or “grounds” known to the whalers. Townsend (1935) studied whale distribu¬ tion based on whalers’ logbook records dating from 1761 to 1920; the logs indicate positions and dates of whales 43 taken by seagoing whaling vessels from New England and Long Island, enabling Townsend to plot on maps the location of the take by month. An extensive sperm whal¬ ing ground centered roughly southeast of Long Island and northwest of Bermuda (approximately 33°-40°N, 60° — 75°W), where sperm whales may still be observed, was known as the “Southern Ground.” The western edge of this ground coincided approximately with the edge of the continental shelf; and like other North Atlantic grounds it was influenced by the Gulf Stream. This was a summer whaling ground, sperm whales being taken here primarily from May to September (especially May to July at the northern limit of this ground, near the latitude of Long Island). Few were taken in winter here; at that season attention was directed chiefly to waters below 25°N latitude. Pigmy Sperm Whale Kogia breviceps Blainville Although related to the great sperm whale, Kogia is very small (maximum length 13 feet), and in outward appearance somewhat resembles a porpoise. Also the pro¬ truding snout and underslung lower jaw with sharp teeth are suggestive of a shark, for which it is sometimes mis¬ taken by fishermen. The rather sluggish pigmy sperm whale is rarely observed and is little known, although it appears to be rather widely distributed in the tropical and temperate waters of the world. Evidently Kogia was never common in historic times; however, it has stranded fairly frequently along the east coast of the United States, where more than 50 records are now known (Handley, 1966). Since 1914, at least eight specimens of Kogia have become stranded on the New York coast or captured a .short distance offshore, and information is available on several. A large female with an unborn young (male fetus) stranded at Long Beach, on the south shore of Nassau County, Long Island, on February 28, 1914 (Schulte, 1917; Schulte and Smith, 1918; Allen, 1941). A pigmy sperm whale 9 feet long and weighing 700 pounds was captured off South Beach, Staten Island, on March 2, 1920 (Davis, 1920) ; this individual was re¬ ported as a porpoise in the local newspapers. Another one, a female, was captured 10 miles south of Shinnecock Inlet, near Hampton Bays, Long Island, in July 1941 (J. Carter). A specimen has also been collected at West- hampton Beach, Long Island (H. Raven). All of the above specimens (skulls and/or other skeletal material) are in the American Museum of Natural History. In August 1942, Roy Latham collected the skull and some other bones of a Kogia breviceps about 9 feet long. This animal had become caught 10 days earlier in the leader of a fish trap at Major Bank, Orient Harbor, and then washed ashore dead. Latham also recalls another Kogia taken near shore at Hampton Bays about 1930. James W. Romansky, Jr. (personal communication) reported on a pigmy sperm whale he found on the south shore of Captree Island, approximately 200 feet east of Fire Island Inlet Bridge, on the morning of November 24, 1968. Total length of the animal, measured 32 hours after it was found, was 157.5 cm. (less than 6 feet) ; it was an immature male. The cause of the death was an apparent gunshot wound just behind the left pectoral fin. It is believed the wound was inflicted by commercial fish¬ ermen, since the skin had parallel marks as if caused by rubbing against metal trawling rigs or offshore pound- nets. Careful measurements and notes were taken, and an articulated skeleton and internal organs in formalin are being kept at the Bay Shore High School. This indi¬ vidual was identified by Romansky as K. breviceps (iden¬ tification confirmed at Smithsonian Institution and Ameri¬ can Museum of Natural History). It is possible that some of the smaller New York specimens of Kogia are referable to the closely related dwarf sperm whale, K. simus Owens. As shown by Hand- ley (1966), simus is a strongly differentiated species of Kogia, which, like breviceps, commonly strands along the coasts of eastern United States. Considerable taxo¬ nomic confusion has existed regarding members of this genus, and simus has been overlooked and not distin¬ guished from breviceps by most authors and collectors. Long Island may fall within the geographical range of simus as well as of breviceps, but I do not know if the former has been collected here. Kogia simus is even smaller than breviceps, adults under 9 feet in total length, while the latter commonly ranges from 9 feet to 11 feet when adult. Size and skull differences and other dis¬ tinguishing characteristics are tabulated by Handley (1966). I have closely examined only one Long Island specimen of Kogia, the Orient Harbor skull collected by Roy Latham in 1942 (now at the N.Y. State Museum), and identified it as breviceps. Besides this one and Ro- mansky’s breviceps, probably the Long Beach and Staten Island specimens cited above are also breviceps, judging from available measurements and illustrations. 1 44 Dense-beaked Whale Mesoplodon densirnstris Blainville This species is frequently called the Atlantic beaked whale or Blainville beaked whale. Members of the genus Mesoplodon are small, mostly rare whales, the life his¬ tories of which are very little known; food is chiefly squids and fishes. Some species are known only by a small number of stranded specimens. Moore (1966) dis¬ cusses distinguishing skull characters and apparent distributions of species of Mesoplodon which strand in North America. On May 12, 1925, a young female densirostris was found stranded on eastern Long Island at Southampton (Raven, 1942). A few other individuals of this species have come ashore to the north and south of our area, including Massachusetts and New Jersey. Antillean Beaked Whale Mesoplodon europaeus Gervais The Antillean beaked whale is also known in the literature as Gervais’ beaked whale or Gulf Stream beaked whale. This rare whale, of which a total of 14 verifiable records of occurrence are known, is principally southwestern North Atlantic in distribution (Moore, 1966). An adult female 15 feet 4 inches in length be¬ came stranded on the south shore of Long Island at Rockaway Beach, Queens County, on December 22, 1933 (Raven, 1934, 1937). This is the northernmost record for the western North Atlantic, although another one came ashore in 1905 on the nearby northern New Jersey coast at North Long Branch. True's Beaked Whale Mesoplodon mirus True The principal range of True’s beaked whale ap¬ pears to be the North Atlantic, from which about a dozen stranded specimens are known. Raven (1934, 1937) re¬ ported on an adult female mirus about 16 feet long stranded at Edgemere, Rockaway Beach, on January 14, 1934, less than a month following the discovery of the specimen of europaeus, also at Rockaway Beach. Another adult female (15 feet, 6 inches long) stranded alive and died on Mason Island, off Mystic, Connecticut, on No¬ vember 19, 1937 (Thorpe, 1938) ; this locality is near the entrance to Long Island Sound and about 3 miles north of Fishers Island, New York. As mentioned by Moore (1966), the North Sea beaked whale [Mesoplodon bidens) is even rarer on this side of the Atlantic (two strandings known) than the three other members of the genus discussed above. M. bidens is unrecorded on Long Island; the nearest record is a stranded male found on Nantucket, Massa¬ chusetts, in 1867. Cuvier's Beaked Whale Ziphius cavirostris G. Cuvier This whale, also known as the goose-beaked whale, is somewhat larger than the species of Mesoplodon, adults generally ranging from 18 to 28 feet in length. Cuvier’s beaked whale is nearly worldwide in distribution, but is rather scarce and little known. Two, a female 17 feet 6 inches in length, and a recently born young measur¬ ing 8 feet 3 inches, came ashore alive and then died at Long Beach, on the south shore of Nassau County, on August 15, 1914 (Rockwell, 1914; Ulmer, 1941). A third individual reportedly ran ashore briefly, but escaped capture by lifeguards who secured the other two as soon as they beached. Skeletons of the two Long Beach speci¬ mens of Ziphius as well as the New York individuals of Mesoplodon mentioned above, are in the American Mu¬ seum of Natural History. A goose-beaked whale 18 feet 11 inches long beached to the northeast of Long Island at Newport, Rhode Island, on March 14, 1961 (Cronan and Brooks, 1968). The North Atlantic bottle-nosed whale [Hyper oodon ampullatus) , another member of the family of beaked whales, is a far northern species which has not, appar¬ ently, been recorded this far south on the western side of the Atlantic. According to Miller and Kellogg (1955), Hershkovitz (1966), and others, the bottle-nosed whale is unknown south of Rhode Island, where it is an ex¬ tremely rare visitor. However, it is worth noting that this species is often mentioned in the literature as having occurred in the Long Island region, locations cited in¬ cluding Lower New York Bay, the south shore of Long Island, and also the nearby Connecticut coast of Long Island Sound; but, for one reason or another, all such published reports are incorrect as far as I know. Of course, there is always the possibility of a straggler being found here. 45 Little Piked Whale Balaenoptera acutorostrata Lacepede Least rorqual and minke whale are among the many vernacular names for this species. The little piked whale is the smallest of the rorquals or fin whales of the genus Balaenoptera; rarely does it exceed 30 feet in length. A broad white band on the outer side of the front flippers is a distinctive identifying mark. As with many other whales, the minke whale is widely distributed in the oceans of the world, and is most frequently encountered in colder waters. It is generally considered to be rare south of the latitude of Long Island. There are quite a few records for the Cape Cod-Nantucket-Rhode Island area to the east and north. Southward the story is dif¬ ferent. For many years a single record for New Jersey (Long Beach, fall 1866) was the southernmost known occurrence; however, it is now known to occur occasion¬ ally as far south as Florida. The first recorded instance of the little piked whale in the Long Island area appears to be an individual captured in 1822 in the Lower Bay of New York. DeKay (1842), under the name beaked ror¬ qual (Rorqualus rostratus) gives a detailed description of this specimen, indicating a fin whale 18 feet long with “swimming paws white in the middle.” At the other end of the island, Helmuth (1931) examined a specimen about 25 feet long which was killed off Montauk Point and towed to shore on August 16, 1931. Finback Whale Balaenoptera physalus Linnaeus The various species of Balaenoptera are known col¬ lectively as fin, finner, or finback whales or rorquals. They are streamlined and mostly very large whales, with a small dorsal fin in contrast to the right whales which lack this fin. The spout, or blow, of rorquals is single and vertical (if not deflected by the wind) ; sperm whales have a single spout directed forward at an angle, while right whales have a double V-shaped spout. B. phy¬ salus, the commonest rorqual, is huge (commonly 60-70 feet long), and exceptionally fast. This species feeds on planktonic crustaceans, also herring, mackerel, and other fishes. The finback whale appears to be migratory, as are the other rorquals. In the North Atlantic this species moves northward in spring to feed at the higher lati¬ tudes during the summer, and southward in autumn to warmer waters, although its movements are imperfectly known. Finback whales were rarely taken by whalers in the early period when right whales were sought, the former being faster and more difficult to handle, with a lower yield of oil and whalebone. Finbacks were caught in large numbers following tbe development of faster ships and more efficient equipment; there was an active fishery in the New England area from 1850 to 1896, as described by Allen (1916). The finback whale has been greatly reduced in numbers by whaling, but is still generally the most numerous of the large whales off the coast of north¬ eastern United States, including the latitude of Long Island, and also is the one most commonly stranded. Al¬ though probably not as common here as in the vicinity of Cape Cod, which extends farther out to sea, finbacks are occasionally seen near the Long Island shore. They are reported fairly frequently from fishing boats off Mon¬ tauk Point. Finbacks are most frequent in summer, al¬ though they may be encountered in any month. Whales this large only rarely penetrate Long Island Sound be¬ yond its eastern end, however. Finback whales were frequently sighted, but rarely tackled, by eastern Long Island shore whalers who pur¬ sued right whales in small open boats launched from the shore as recently as the early 1900’s. Edwards and Rattray (1932), who stated that finback whales often approach the Long Island shore closely in pursuit of small fish, related an instance of a crew fastening a har¬ poon to a finback off Amagansett; they were reportedly towed 3 miles away from shore in a fast and frightening “sleighride” until the whale broke free of the iron. Murphy (1918) wrote that finback whales regularly feed offshore in the Long Island area in summer, and reported that six finbacks came inside the inlet to Jamaica Bay in July 1916; one of these, 50 feet long, perished after becoming stranded on a bar. Other strandings include one at Huntington Harbor (inside Huntington Bay, western Long Island Sound) on October 22, 1946, and another at Sheepshead Bay, south shore of Brooklyn, on Novem¬ ber 14, 1936, (Anon., 1956). Sei Whale Balaenoptera borealis Lesson This species resembles in general appearance the fin¬ back whale {Balaenoptera physalus), but is somewhat smaller, attaining a maximum length of 50 or 60 feet. The word sei, correctly pronounced “say,” is from the Norwegian, and refers to a kind of fish with which this whale often associates, both feeding together on the same crustaceans. The sei whale is widespread throughout the 46 oceans of the world, and is numerous at times along the Norwegian coast and also off Newfoundland. Generally, though, it is considered to be rare in the western North Atlantic. 1 have no records for the sei whale on the Long Island coast, but probably it is a rare visitant; it has been recorded from Labrador south to Florida and Mexico, and has stranded on the Massachusetts coast at least twice in this century. Also, although borealis is smaller and darker than the more numerous phy solus, it is so similar to the latter that it is difficult to identify at sea. Thus it is possible that undetected schools of sei whales are present at times offshore. This species is mi¬ gratory but is more irregular in its occurrence than the other rorquals. Blue Whale Balaenoptera musculus Linnaeus This species, also known as the sulphur-bottom, is the largest of all mammals. Total length of fully grown adults in the Northern Hemisphere is about 75-95 feet (females somewhat larger than males); blue whales grow even larger in the Antarctic, where specimens over 100 feet long have been recorded. Very large individuals are now rare, however. Food is almost entirely shrimp-like crus¬ taceans less than 3 inches long which swarm in the cooler oceans of the world. The blue whale is pelagic and highly migratory, those in the Northern Hemisphere moving to far northern waters in spring or early summer to feed, and then southward in autumn for the winter breeding season. The breeding grounds have not been pinpointed, and most blues may spend this period far out in the central or southern North Atlantic; apparently only a few occasionally migrate as far as the tropics. In the North Atlantic this species has been so over- exploited that probably only a very few hundred remain, and it is the most endangered of all the large whales. The blue whale is rare everywhere along the east coast of the United States, although it has been found off Newfound¬ land at times, and there is a sparse scattering of records of individuals which have stranded, or beached, as far south as Ocean City, New Jersey. I know of only one record of a blue whale on the Long Island coast. An individual came ashore at Sagaponack several decades ago (date unknown to me). This stranding is mentioned by Edwards and Rattray (1932), who also provide a photograph of the dead whale lying on the ocean beach. Humpback Whale Megaptera novaeangliae Borowski This species is closely related to the rorquals but has a stout body and long pectoral fins; adults are about 40 to 50 feet in length. It occurs in all oceans and is highly migratory along often rather well-defined routes, spending the winter in tropical and subtropical waters and migrating to high latitudes for the summer. In spring, northbound humpbacks pass our region, and large herds have been seen in April moving north some 200 miles east of the North American coast. Kellogg (1929) provides a distribution map with probable migra¬ tion routes. Humpback whales may be seen in spring near the coasts of the northeastern states, and a few may also be encountered in summer and fall, but the fall mi¬ gration south appears to be farther offshore. Also, the larger schools are usually fairly well offshore at all times according to Allen (1916). Formerly common, Megaptera has been badly depleted and is now considered to be rare in the North Atlantic, although increasing slightly in recent years and more numerous on the western side (Kenyon, et ah, 1965; Simon, 1966). In the Long Island area, probably a few humpbacks were taken by early shore whalers pursuing right whales, with more attention being directed to this species from boats cruising offshore following the decline of the right whale. According to Allen (1916) a favorite whaling ground for humpbacks was on the Nantucket Shoals, which are about 100 miles due east of Montauk Point. They were killed here, and also on the Georges Banks farther east, during the eighteenth and nineteenth centuries. Everett J. Edwards, who participated in 15 late period shore whal¬ ing expeditions for right whales off the Long Island beaches, and whose father. Captain Josh Edwards, led the Amagansett whaleboats for nearly 50 years, until 1915, recalled only one humpback sighted locally. According to the younger Edwards (in Edwards and Rattray, 1932) about 1913 his father headed a boat which fastened a harpoon to a humpback which had been sighted from the Amagansett beach. But the bomb failed to go off, and the whale towed the boat at high speed until the line parted. Humpback whales rarely become stranded and I do not have any such records for Long Island, although un¬ doubtedly this has happened. One stranded at nearby Matunuck, Rhode Island, June 1957 (Cronan and Brooks, 1968). 47 Black Right Whale Balaena glacialis Miiller Right whales, like the rorquals, are huge whalebone whales which lack teeth but have horny plates of whale¬ bone or baleen, which are frayed on the inner side and serve to strain small food organisms from the water. Right whales lack both a dorsal fin and grooves on the throat characteristic of the fin whales, but have a rela¬ tively larger head and chunkier appearance; adults may be 45 to 55 feet long. This species was the main object of pursuit by the early North Atlantic shore whale fish¬ ery, in the day of small boats and hand-held harpoons, because of its slowness, buoyancy when dead, and high yield of oil and whalebone. The coastal area from Massachusetts to Long Island, where this whale was numerous in season, was one of several important centers of the right whale fishery in the North Atlantic. One after the other these various areas of abundance became depleted, until the right whale verged on extinction. The species survived, is now pro¬ tected by international law, and appears to be slowly increasing; but in all the North Atlantic perhaps only a few hundred right whales exist (Anon., 1968b). Shore whaling, using open boats launched from the shore, became established on Long Island during the 1640’s and 1650’s, and was particularly active along the southern coast of the east end of the island, although right whales were taken as far west as Brooklyn. Also larger boats were used to hunt right whales a short dis¬ tance from shore. In the early decades of whale abun¬ dance scores were caught in some years, and also the many “drift whales,” those which had been cast up on shore, were utilized; probably many of these had been disabled by harpoons, others may have been driven ashore by storms or illness. Native Long Island Indians were skillful whalers willing to work for low wages, and thus were frequently employed in manning the whale¬ boats. After the early part of the eighteenth century, intensive shore whaling rapidly declined here (1707 peak year for amount of oil sold). A few right whales continued to appear along this coast, and sporadic shore whaling continued through the eighteenth and nineteenth centuries, even though the major whaling effort was then far at sea in larger vessels. Especially in the region of Amagansett, East Hampton, and Southampton, where Long Island shore whaling be¬ gan, whales were still pursued whenever sighted from shore. At times this amounted to only one or a few taken at intervals of several years. DeKay (1842) wrote; “The right whale was formerly captured in great numbers from sloops and whaleboats, along our whole coast, chiefly from February to May, although they appeared occasionally at all seasons of the year. Along the southern coast of Long Island, whaleboats are still kept in readi¬ ness; and upon the appearance of a whale, the people in the vicinity quickly assemble, and soon are in pursuit of the animal.” Four which were harpooned on January 30, 1885, near Southampton attracted collectors attempting to sal¬ vage skeletal material, including Frederick W. True from the U.S. National Museum (True, 1885). The skeletons of a few other right whales killed off eastern Long Is¬ land in the late 1800’s found their ways to various mu¬ seums. Some were still being killed along this same coast early in the twentieth century; one of these, an adult female 54 feet long (from tip of snout to notch of flukes) captured off Amagansett on February 22, 1907, and the skeleton collected for the American Museum of Natural History, was one of the largest American speci¬ mens ever recorded (Andrews, 1908, 1916). A young whale 38 feet long believed to be the calf of the large one, was captured^nearby. Andrews (1909) also studied another small specimen captured by shore whalers off Amagansett on December 10, 1908. A lively firsthand account of this late period (1880’s - 1918) of shore whaling from open whaleboats in the Amagansett region is included in the book by Edwards and Rattray (1932). According to the authors the year 1918 was the last in which a right whale was caught and tried out for oil (30 barrels, but never sold) ; two were sighted off the East Hampton shore early one summer morning and the larger of the two was pursued to Napeague and killed. According to Sleight (1931) right(?) whales were sighted off Southampton and else¬ where in 1923, but the era was definitely over, and no boats were launched. Still to be seen on some old houses in the same region is the “scuttle,” a trapdoor in the roof where a person could look out over the ocean and watch for whales. True (1904) and Allen (1916) compiled reports of right whales captured off the Long Island shore, gleaned from state and local histories and records, and news¬ papers such as the Nantucket Inquirer, which kept its readers well posted on whaling matters. Years covered range from 1669 to 1908, with most data from the 1800’s. Such reports often indicate date of capture, number of whales seen or killed, approximate location along the coast, number of barrels of oil produced or expected, and occasionally length of animal and if calves seen. Dates given indicate that right whales through the years have been most numerous from about late February to late May, with a few appearing earlier (November, De- 48 cember, January). According to Allen (1916) right whales are, or were, more numerous in April than dur¬ ing any other month of the year in the Long Island region. Cows with calves have been noted in March, April, and May. That occasionally a right whale caused at least accidental damage to its pursuers after being wounded also is recorded in the documents; for example, a whale was struck on April 28, 1854, off Southampton, and though mortally wounded it demolished the whaleboat, injuring several men. Edwards and Rattray (1932) tell of right whales off Amagansett overturning whaleboats, even as recently as 1911, and in some instances cutting boats in half with their flukes. Human fatalities were infrequent, however. This species generally avoids shallow waters and rarely strands, but there are old records (late 1800’s) of individuals entering Upper New York Bay and Raritan Bay (near Staten Island), and in 1850 one was captured in Shelter Island Sound near Greenport. According to Edwards and Rattray (1932) right whales seldom came over the shallow sandbar said (by the authors) to lie about a quarter of a mile off the Long Island south shore beaches, whereas the more slender finbacks reportedly came over it frequently in pursuit of small fish. Right whales continue to appear off the southeast coast of Long Island, and I assume mainly at the same time of the year as in the past, but reports are few, in part probably because the whales are no longer watched for at the proper season by many well-trained eyes on shore. In recent years right whales have reappeared in numbers in the Cape Cod area, with up to 40 or 50 present in Cape Cod Bay during May, and occasional occurrences as late as June (reported by Waters and Rivard, 1962, and others). A recent record for Long Island is that of a right whale observed on June 8, 1960, in the ocean off Shinnecock by Stanley E. Poole ( William E. Schevill, personal communication). The East Hampton Town Marine Museum at Amagansett has a right whale skull found on the beach there in the summer of 1965. Although frequenting temperate rather than tropical or arctic seas, in the western Atlantic these whales sum¬ mer well to the north of Long Island, and many of them winter some distance to the south. In fall, most south¬ bound ones pass by well out to sea. As Allen (1916) points out, in spring right whales apparently migrate fairly close to the coast and are turned abruptly east¬ ward by the projecting coasts of Long Island and Cape Cod. Thus they converge on the south and east shores of Long Island and Massachusetts to round Cape Cod. Red Fox Vulpes vulpes fulva (Desmarest) The red fox is surprisingly common at the present time in most parts of Suffolk County, and it is frequent also in the less developed areas of Nassau. Writing well over a century ago, DeKay (1842) mentioned a large male red fox killed in Queens, and evidently this species was common enough then, as it had been at least as far back as the late 1700’s, although DeKay did not comment on the relative abundance of the red fox to the then very numerous gray fox. Later, at the turn of the century, Helme (1902) briefly noted that the red fox was very common throughout Suffolk and portions of Nassau counties. Through the years red foxes have been popular in furnishing sport for hunters, and also many have been killed for their pelts, which were formerly in de¬ mand and brought good prices. The gray fox was more numerous on Long Island in prehistoric times than the red fox, judging from finds at Indian archaeological sites. I do not know of any published information confirming the presence of the latter on Long Island during Indian times. Waters (1967), who discussed archaeological finds of Vulpes and Urocyon on Martha’s Vineyard, knew of no records of Vulpes from archaeological sites on Long Island or farther south. Thus the faet that Vulpes is represented in Roy Latham’s archaeological collection from eastern Long Island is of interest. He writes (personal communication) that although more gray foxes were found in his excava¬ tions, both species were recorded from sites at Three Mile Harbor, Southold, and Shelter Island. European red foxes, representing a different geo¬ graphical race of V, vulpes, are said to have been re¬ leased on Long Island for hunting purposes during the 1700’s. Apparently such introductions, here as elsewhere, did not take hold, or at least had little effect on the natural increase and spread of the native American form, fulva, which extended its range southward into the middle Atlantic and southern states during Colonial times. Assuming the red fox was present on Long Island when the European settlers arrived, the changing en¬ vironment which followed may have favored the increase of this species (or if it disappeared during late Indian or early Colonial times it could have reinvaded the island during the spread southward) . During the survey, we saw red foxes or signs of their presence in a great variety of habitats. Red foxes cover most of the wilder open country on the island in their 49 search for such food as cottontail rabbits, meadow mice, birds, carrion, insects, and other varied items. They are very common on the outer beach strip, such as Fire Island and the section between Moriches and Shinnecock inlets; here they patrol the dunes, thickets, and salt marshes, and even forage along the water’s edge on the ocean beach. Inland, red foxes are present in woods, near streams, and around large fields. Eastward on the island they seem especially common, and are very much at* home at Montauk, Hither Hills, Shinnecock Hills, Napeague meadows and dunes, and on Shelter Island, and elsewhere. Apparently eastern Long Island is one of many regions where this species has shown a general increase in recent decades. According to Roy Latham (personal communication) red foxes were encountered only occasionally in the Orient region during four decades or more prior to 1930, but have become very common since then. During the late 1800’s the wilder Montauk region was noted for its red foxes, and it attracted hunting parties from farther west on the island. Naturally, the fox has not fared so well on western Long Island, but it still thrives in some built-up areas near New York City. It was common in now densely settled eastern Queens at least as late as the 1920’s and still is plentiful as far west on the south shore as Jones Beach (Anon., 1968a). Dens, some actively in use, were found in various situations: on shrubby, grassy hillsides in the Shinnecock Hills (figure 15), in a dike crossing an old, abandoned cranberry bog, in sandy pitch pine barrens, and in the sand of the more sheltered inner dunes on the barrier beaches. In the Long Island sand dunes red foxes are reported to favor the concave face of a wind-carved and overhanging dune as a burrow site (Anon., 1968a). In an area of woods and estates on the north shore of Nas¬ sau County, in June, we were shown where red foxes had apparently raised four or more young in a long drainpipe under a paved drive; remains of cottontail, rabbits and other prey were present when we visited the site. Latham (1954) wrote: “Probably the food of the red fox is more varied than that of any other wild mammal on Long Island, and no other is so clever in the methods of obtaining it.” In the same article he wrote about two different occasions at Montauk when he observed red foxes catch a weasel and a muskrat. Hamilton (1935, 1949), drawing partly on information supplied by Roy Latham and others, mentioned some of the diverse foods of red foxes on the island, such as various kinds of frogs, birds, and mammals, and also discarded marine fish, snapping turtle eggs, blueberries, and other tidbits. Audubon and Bachman (1851) wrote an interesting ac¬ count of the abundance of red foxes, and their food habits, on the New Jersey outer beaches, an environment similar to the south shore of Long Island; rabbits, water- fowl, wading birds, and crabs and fish thrown up by the surf were reportedly eaten. Near the entrances to red fox burrows in the Shin¬ necock Hills, in the spring of 1963, I noted conspicuous remains of various food species, including eastern cotton¬ tail, meadow mouse, pine mouse, muskrat, pheasant, and meadowlark. A nursing red fox, examined after being shot by a hunter in central Suffolk County, was found to be carrying three adult meadow mice in its mouth. In unusually deep snow present in January and February 1961, we found a number of cottontails which had been cached by red foxes, several near small streams in the pine barrens. Scats (droppings) are a good indication of what foxes actually eat; only a small number were ex¬ amined, and they contained cottontail and meadow mouse remains. Of course, the diet of red foxes on Long Island does not differ basically from those living elsewhere in the State, except for their propensity to feed on available marine organisms and sea birds. Gray Fox Urocyon cinereoargenteus cinereoargenteus (Schreber) Originally the common native fox of Long Island, the gray fox became greatly reduced in numbers during the past century and it has remained rare and local since then. DeKay (1842) stated that the gray fox was then very abundant on Long Island, where it was often called the “plain fox” or “grass fox.” Helme (1927) wrote that although formerly common, this species had become extinct on the island, and that few if any grays were to be found later than 1880; he thought that any instances of their occurrence in the present century must be at¬ tributed to introduction or escape from captivity. However, the status of the gray fox remained in doubt, with a scattering of sightings reported since 1927, while perhaps a majority of the interested field natural¬ ists believed the species had disappeared. I do not know of any specimens actually collected and preserved in re¬ cent years, and at least some reports of gray foxes killed have turned out to be red foxes. A few gray foxes have been included in the reported take of foxes by hunters and trappers on Long Island, as were mapped for the year 1940 by Seagears (1944). Even assuming that all of the reported grays were correctly identified, these records indicate that this species comprises only a very small percentage of the total number of foxes killed here. 50 Hamilton (1949) maintained that the gray fox is generally distributed (or at least was then) in the scrub oak and pine of the central portion of Long Island. Also, this fox apparently still exists in the Montauk-Hither Hills area according to Sam Yeaton (Anthony Taormina, per¬ sonal communication). In recent years, gray foxes and their trails have been reported in dense thicket country at Montauk (Anon., 1968a). During our field survey we glimpsed single individuals believed to be this species on two occasions, although both of the sightings were dis¬ appointingly fleeting; localities were wooded areas near Hampton Bays and Montauk Point. From the scanty evidence, it seems likely that the gray fox declined drastically in numbers without entirely disap¬ pearing from the island. It apparently survives today in the wilder sections of central and eastern Long Island, especially in the pine barrens region and the eastern part of the south fork. However, so little seems to be on record regarding this species here that any additional informa¬ tion, if made known, would be of considerable interest. It should be noted that the gray fox tends to be more nocturnal and secretive than the red, and less inclined to frequent the open country and beaches. Raccoon Procyon lotor Intor (Linnaeus) The familiar raccoon, like the red fox, is a sizable member of the Carnivora which is more than holding its own on much of Long Island. It is common in various parts of the heavily settled western portion, including certain areas in eastern Queens, and ranges eastward to Montauk Point, Orient Point, and Shelter Island. While it has fluctuated in abundance through the years, prob¬ ably according to the degree of persecution (many have been killed for sport and for their pelts) and in re¬ sponse to a rapidly changing environment, the raccoon has persisted; evidently it has been well served by its intelligence, adaptability, and highly omnivorous food habits. The raccoon has been more or less widespread on Long Island throughout a long period of man’s existence here. Helme (1902) stated that it was common in most parts of the island. For the Montauk region, Dutcher and Butcher (1893) found raccoons common, especially around the various ponds there. On the north shore, Tur- rell (1939) reported raccoons numerous in the swamps of the Smithtown region, and also along the Nissequogue River and various brooks. DeKay (1842) did not men¬ tion Long Island specifically, but stated that the species was well known and found in every part of the State. Going back three centuries, Denton (1670), in his ac¬ count of Long Island, listed the raccoon as a member of the fauna, and also stated that it was eaten by the In¬ dians. Also, bones have been found at prehistoric Indian sites of different periods. In recent decades raccoons have been on the up¬ swing in the area, becoming very common and spreading into new or reoccupying old neighborhoods. For example, Roy Latham (personal communication) writes that for at least 40 years, from about 1888 to 1930, there were no raccoons in the Orient region, but that they have become very common since 1930. Latham states further that they have become exceptionally numerous and locally trouble¬ some to the farmer in the past 20 years. They eat potatoes by digging into the hills, and destroy corn, fruit, and other crops, and even damage buildings by pulling shingles off roofs to get inside. Other writers and ob¬ servers also tell of a general increase of raccoons else¬ where on Long Island, especially since about 1940. We saw tracks and other signs of “coons,” and sometimes the animals themselves, in a variety of situa¬ tions. These included margins of streams and ponds throughout the island, swampy maple woods, bogs, pine and oak barrens near water, small patches of deciduous woods in suburban areas, woods near brackish and salt marshes, grassland at Montauk and Napeague, and even the bare sand of the “walking dunes” (although near trees and marshes) at Napeague Harbor. Although fre¬ quenting the edge of protected salt water areas, such as Long Island Sound, and the Peconics and other bays, the raccoon does not seem to be as common on the ocean barrier beaches (with their dunes and salt marshes) as the red fox; this may be because of the lack of trees or other denning sites in many of these areas. Raccoons are on Fire Island, however. Inland, raccoons often survive in a patch of woodland trees, especially if a stream or pond is present, after it has been largely transformed into a suburban area or a developed park; their char¬ acteristic footprints are still to be seen long after most of the larger forms of wildlife have disappeared. Long-tailed Weasel Mustela frenata novebpracensis (Emmons) Distribution and habitat. This is the common weasel found more or less throughout Long Island, except in the heavily built-up areas. Helme (1902) recounted his experiences with weasels on the island — family of young found under a pile of wood, one shot from a squirrel’s nest about 20 feet above ground in a cedar tree, etc. In Queens this species was formerly common and probably 51 still occurs locally; Hamilton (1949) once observed a nest with young at Bayside in Queens. Quite a few Long Island specimens are in other museums and private col¬ lections, localities recorded including Flushing in Queens, various north shore and south shore points, and east to Orient and Shelter Island. In eastern Suffolk County we occasionally observed long-tailed weasels, trapped a few, and found others killed on the highway; localities were Flanders, Hampton Bays, Montauk Point, Quogue, Riverhead, Sagaponack, South¬ ampton, Squiretown, Tiana Beach, Wainscott, and Water Mill. Several of our specimens were caught at two trap sites near Flanders, one located at the base of a fallen tree in a densely wooded, bushy area by a fresh stream, the other in the base of a hollow stub in woods bordering a salt marsh. It is not unusual to encounter this species near the ocean shore and on the outer barrier beaches. At Tiana Beach near Shinnecock Inlet a long-tailed weasel was observed in a salt marsb and another was found dead on Dune Road; this area is chiefly salt marsh, sand dunes, and flats. Tracks judged to be of this species were noted in the sand on Fire Island (opposite Smith Point and near the Sunken Forest), and on the east side of Moriches Inlet in salt marsh and sand. One January day in grassy ocean dunes near Wainscott, Christopher Mc- Keever and I found a large adult male weasel which had been killed just a short while previously, apparently by a marsh hawk {Circus cyaneus) . The weasel, decapitated and partially eaten, was found lying on the ground near a summer cottage, after we had startled a marsh hawk into flight from the spot. Next to the weasel were several pellets probably cast by the hawk, three of which con¬ tained the remains of weasel, others the remains of mead¬ ow mice. Tracks and other signs of weasels (probably frenata) were also noted in the following habitats: large field near Manorville with many meadow mice and pine mice pres¬ ent, bog near Speonk with many meadow mice, pine barrens near Flanders, and red maple swamp at East Hampton. Two were found dead on roads next to duck farms — the weasels may very well have been seeking Norway rats often abundant in such areas. Mustela frenata is the species of weasel usually found on Staten Island, too, although it no longer seems to be very common. Some years ago (1941) I saw one hunting far out in a salt marsh there, near New Springville, My impression, in limited trapping for this species, was that it is much less common on Long Island than in many “upstate” (mainland) New York areas. Roy Latham, in a personal communication, writes that weasels were common and widespread up to about 1950, but he believes they have become very scarce, at least on eastern Long Island, during the last two decades. Remarks. All the winter individuals collected or ob¬ served were in brown pelage. Although most long-tailed weasels in northern and central New York turn white in winter, virtually all of them remain brown on Long Island. Helme (1902) and Hamilton (1949), both familiar with this species on Long Island, never recorded any in white winter pelage. During the survey, we received a few sight reports of white weasels in winter in Suffolk County. This could indicate that a small proportion of the population assumes a white winter coat, or that the observations refer to Mustela erminea (which turns white throughout its eastern range). Based on what is already known, the latter seems more likely, although erminea is rare on Long Island. Most of the stomachs examined were empty, but one held the partial remains of a gray squirrel. Some of the prey species on record as having been caught by weasels on Long Island are Sorex cinereus (Nichols and Nichols, 1935); moles, Scalopus Aquaticus (Helme, 1902); cot¬ tontail rabbits; chipmunks; meadow mice; white-footed mice; and Norway rats. A female on April 17, 1961 (Flanders, Suffolk County) contained six embryos, the swellings 8 to 9 mm. in diameter. Measurements. Data for the female with embryos mentioned above are; weight, 109.5 grams; total length, 298 mm.; tail, 90 mm.; hind foot, 34 mm. An adult male on the same date: weight, 205.4 grams; total length, 397 mm.; tail, 138 mm.; hind foot, 45.5 mm. Individuals taken. 9 Short-tailed Weasel Mustela erminea cicognanii Bonaparte The short-tailed weasel, or ermine, a smaller and more northern species than the long-tailed weasel, is very rare in the coastal plain portion of New York State. An individual in white winter pelage has been taken at Babylon, Suffolk County, and was the only Long Island record known to Hamilton (1949), who obtained a photograph of the specimen. Some nineteenth century pub¬ lications refer to possible but not well substantiated early records of this species on Long Island (summarized by Helme, 1902). Mustela erminea is also rare on Staten Island, where it has been collected by Crowe (1939). There are. a Jew sight reports by biologists of erminea on both Long Island and Staten Island. This region is not 52 included in the range of erminea as described by Hall and Kelson (1959) ; for the east coast they state only that this species occurs south to Rhode Island, although they also note an inland Maryland record. We did not obtain any firsthand information regard¬ ing short-tailed weasels on this survey, although reports received from Suffolk County residents of “very small” weasels and white winter weasels may refer to this species. It is said that one or two least weasels {Mustela nivalis) have been collected on Long Island and the skins mounted or otherwise preserved, but I have been unable to locate the whereabouts of any such specimens. Weasels so labeled and actually from Long Island may be Mustela erminea, since this rather small weasel has, in the past, been called “least weasel,” as it was in Crowe’s paper cited above; the range of the true least weasel, nivalis, is not known to extend this far east. Mink Mustela vison mink Peale and Palisot de Beauvois Although the mink is not often seen and is much less common than formerly, it is not really rare and in fact has a rather wide distribution in the region. This species may be encountered near water, mainly in the less popu¬ lated areas, for almost the full length of Long Island and on both the north and south shores. Like the muskrat, the mink is at home in the coastal salt marshes as well as fresh water areas inland. Quite a few mink live on the open stretches of the outer barrier beaches, especially Jones Beach, Fire Island, and the strip between Moriches Inlet and Shinnecock Inlet. In these areas I noted their footprints quite frequently in mud, sand, and snow, and also occasionally found troughs in the snow made by traveling mink when coast¬ ing for short distances. In the salt marshes on the bay side of the outer beaches (figure 16) the mink appear to spend much time following the many creeks and ditches, judging from the tracks. In these marshes are many musk¬ rats, meadow mice, birds, fishes, and aquatic invertebrates, while cottontail rabbits abound in the adjacent areas, all potential food for hunting mink. Regarding eastern Long Island, Roy Latham (per¬ sonal communication) writes that mink are still on Montauk Point, at Three Mile Harbor, and on Shelter Island (12 taken in winter of 1952) and are not uncom¬ mon around the Peconic River. At Orient, mink were formerly common, but Latham’s last record in that area was one collected on November 28, 1910; before 1910 he averaged about two per winter on his traplines. In Hither Hills State Park, 1 saw a mink skirting the shore of Fresh Pond one October evening in 1962 (figure 22). As an indication of former abundance, Butcher and Butcher (1893) remarked that mink were very common on Mon¬ tauk Point, with some trappers securing as many as 50 skins apiece in the winter of 1892-93. The status of this species on Gardiners Island was not determined. Chapman (1908), in a chapter devoted to Gardiners Island, referred to a lack of minks, weasels, and other mammalian carnivores. However, Roy Latham was informed by an earlier caretaker there that mink were present before 1900. On a 1-day visit on July 22, 1961, we found partial remains of what appeared to be a mink on the south point of Gardiners Island, but this material was lost on the return boat trip before being definitely identified. For the New York metropolitan area, Hamilton (1949) mentioned trapping mink as a boy near Bouglas- ton, north side of Queens, as well as nearby in western Nassau County. A few mink survive on Staten Island, although much of their habitat has been destroyed in recent years. Study skins and skulls from Long Island are repre¬ sented in various collections, including the U.S. National Museum, the American Museum of Natural History, Cornell University, and the collection of Roy Latham. Striped Skunk Mephitis mephitis nigra (Peale and Palisot de Beauvois) The skunk, like the woodchuck, is a medium-sized mammal which has decreased markedly on Long Island in this century and is generally much less common here than in the mainland portions of New York State. For¬ merly common the length of the island, apparently the skunk has declined in numbers over a long period. At the time of Helme (1902) it was still numerous, although less so than formerly. Helme suggested that perhaps the Paris green poison used to control the Colorado potato beetle (“potato bug”), which arrived in the 1890’s, was reduc¬ ing its numbers; it was thought that the skunks ate the poisoned beetles and suffered a rather abrupt decline at about this time. Murphy (1964) and some other veteran observers agree, although there is not unanimity of opin¬ ion on this point. Skunks seemed to remain common much longer in certain wilder, nonagricultural sections, such as Montauk and in pine-oak barrens but have grad¬ ually become scarce there too. This animal is a frequent victim of automobile traffic, and the dense network of roads and ever-increasing traffic on the island has pos¬ sibly been one cause of continued reduction in more recent years. 53 Hamilton (1949) mentioned that 20 years earlier skunks were common, even in sections of western Long Island. Roy Latham informs me that when he was a boy, skunks used to make burrows in potato fields at Orient; his last record for that area was in the year 1911, but he noted skunks as still common in some other Long Island areas where potatoes are commonly grown, many years after they were gone from Orient. Several residents of eastern Long Island, from Hampton Bays to Montauk, told us (in the early 1960’s) they believed skunks were much more numerous 15 to 20 years earlier. Skunks were formerly abundant on Shelter Island, and specimens col¬ lected there are at the U.S. National Museum and in the collection of Roy Latham. In the 1920’s Roy Latham oc¬ casionally saw albino or white skunks at Montauk, includ¬ ing two observed together on June 14, 1928, in the day¬ time. On Staten Island, a few skunks were present at least as late as 1940, but evidently they have become extirpated or very rare (I have no recent information). It is possible that the skunk is holding its own or even increasing slightly in a few of the less developed areas of eastern and central Long Island. The only speci¬ men we obtained was one killed by a car, but in good condition when found, near Sag Harbor in October 1961. No other road kills were seen, although some were re¬ ported to us from the Montauk area. I noted signs of skunks including a den, odor, tracks, and droppings, in an area of fields and pine-oak barrens near Manorville in 1962 and 1963. The den, a burrow occupied by skunks in the winter of 1962, was in level, sandy ground under scrubby oak trees, near the edge of a large field. Tracks were seen in the smooth sand of the moving dunes in Hither Hills State Park (figure 20) in 1962. Skunks are reported to be common in the Hither Hills section where they sometimes raid the garbage cans at the beach camp¬ ing grounds (Anon., 1968a). Various observers reported seeing skunks in the following localities in the early 1960’s: Montauk, Calverton, and Yaphank in Suffolk County, and North Hills in Nassau County. Three hundred years ago, Denton (1670) listed the skunk as one of the wild animals of Long Island. He also mentioned this species as an article of food of the local Indians. In his account of the Indians, by then consider¬ ably reduced in numbers, Denton wrote: “The meat they live most upon is Fish, Fowl, and Venison; they eat like¬ wise Polecats, Skunks, Raccoon, Possum, Turtles, and the like.” Roy Latham (personal communication) writes that the skunk was one of the more common mammals dis¬ covered in his Indian archaeological excavations on east¬ ern Long Island, found at most sites, as were muskrat, raccoon, deer, beaver, and wolf. River Otter Lutra canadensis (Schreber) The time is long past since the otter was common on Long Island. Denton (1670) included the otter in his brief listing of Long Island land mammals, but 172 years later DeKay (1842) wrote that this species, although formerly numerous throughout the State, had become ex¬ tirpated on Long Island and Staten Island. Nevertheless, during the past 100 years or so, a few otters have been observed at various localities on Long Island and some have been killed. Possibly the otter was never completely extirpated here, but it has certainly been very rare, at least, for a long time nearly everywhere on the island. Most of the locally active field naturalists have never seen it. According to Helme (1902) at least lour otters were killed, then mounted by taxidermists, between 1875 and 1901, as follows: Yaphank, about 1875; Carmans River, 1898; Peconic River near Calverton, winter of 1900-01, also other individuals seen; Patchogue, 1901. Roy Latham (personal communication) states that one trapped at Three Mile Harbor in 1881 may have been the last one taken at that south fork locality. Another record is an otter shot in Great South Bay near Smith Point in 1902. In Roy Latham’s collection is the skin of an otter trapped on March 5, 1929, at Oyster Pond on Montauk Point; it was a male weighing 20 pounds and had been skinned for the fur market before Latham acquired it. Latham writes that otters were present on Montauk for years before this one was trapped, and he saw them there on several occasions in the 1920’s when this was still wild country. He first saw them in 1925 at Oyster Pond and Great Pond (Lake Montauk or Montauk Harbor on recent maps) before the latter was opened to the sea; he also saw them at Big Reed Pond, a smaller body of water in the same area. In the fall of 1928 he saw two otters at Great Pond and the slide they were using. Otters still persist in this general area; in a recent communica¬ tion Roy Latham writes that in February 1969, an adult female otter was caught in a fishtrap at a Montauk Point locality. Otters were on Shelter Island in the late 1920’s, and one was seen several times by a reliable observer during the winter of 1940-41, according to Latham. Also, Latham writes that otters were sighted in the Peconic River near Calverton in the 1920’s and 1930’s. No otters were en¬ countered during the museum survey, but several reports were received from reliable persons of single otters seen during the 1950’s: Shelter Island (about 1955), locality 54 near north shore of Nassau County (about 1957), Peconic River, and two south shore localities near Great South Bay, Many otter reports, especially certain recent ones and those from western Long Island, can perhaps best be explained as individuals which have come across the Sound from Connecticut, where the species has been in¬ creasing in numbers. On the other hand, Roy Latham has shown that the otter has had a long history at Montauk, and the frequency of reports from certain other areas sug¬ gests additional centers where this species may have be¬ come at least temporarily established from time to time. Harbor Sea! Phoca vitulina Linnaeus This is the only seal which regularly visits the area and, as its name implies, it frequents harbors, bays, and inlets, and also even enters rivers on occasion. Phoca vitulina is a small (up to 6 feet), densely spotted species. Here, in the southern part of its range, where it is a migratory visitor, it is most numerous during winter and early spring, is occasionally observed during the fall, and is rare in summer. This species may be seen swim¬ ming along with its head out of the water, when it appears somewhat dog-like, or intermittently diving beneath the surface; also the harbor seal may be found hauled out on the more deserted beaches, islands, rocks, marshy banks, and points of land. Harbor seals in New York State are most numerous about the east end of Long Island, where even in recent years up to half a dozen or more together on shore have been reported. Roy Latham (personal communication) writes that seals, (usually singles, sometimes three to five in a group) are recorded every winter in Long Island Sound, Gardiners Bay (from Orient west to Cutchogue), and from all the islands of the east end region, including Gardiners Island and Shelter Island. His dates are from November 30 (1929) to May 5 (1932), the latter record at Orient Harbor. Also, 50 seals were counted on a single sailing trip around Plum Island on February 2, 1933. Christopher McKeever informs me that reports he has received indicate fairly large numbers of harbor seals still inhabit the bight on the south side of Plum Island every winter. Butcher and Butcher (1893) reported har¬ bor seals as present every winter on rocks that lie off Montauk Point, usually arriving in the area in late No¬ vember or early Becember and remaining until spring. Robert A. Morris, Curator at the New York Aquar¬ ium, has been keeping records in recent years of reported sightings of harbor seals in the Long Island area. He writes (personal communication) that over the past 2 years — 1968 and 1969 — the first reports of seals have come early in January. These reports continue until the end of April. Only one report was received of a harbor seal in August, one at Montauk Point on August 12, 1968. Harbor seals may be seen just about anywhere along the entire Long Island coast, and small groups are occa¬ sionally noted in western Long Island Sound and along the south shore (ocean and bays). It is more usual to see lone individuals (as mentioned by Latham) and I have seen singles in winter swimming in Shinnecock Bay and in Lower New York Bay. Buring periods in the win¬ ters of 1960-61 and 1962-63 coast guard observers at the Shinnecock Inlet station reported seeing a seal, prob¬ ably this species, swimming every morning in the inlet (figure 12). We also received reports of seals in winter in Moriches Inlet and Great South Bay. The tendency for harbor seals to enter rivers and fresh waters is well known, which explains sightings in the Hudson River. In the collection of the American Museum of Natural History are several specimens (skins, skulls, skeletons) taken between 1928 and 1959, mainly during winter, also one in late October, at both ends of Long Island. Locali¬ ties include New York Harbor and Rockaway Beach on the west and Montauk Point on the east; a few are from Long Island Sound. Richard Van Gelder informs me that harbor seals stll occur regularly in the vicinity of Mon¬ tauk Point, and that quite a few have been shot there in recent years. Many are short for sport. Some fishermen are prejudiced against seals, which do little harm, but eat the common and more available fishes, mollusks, and crustaceans, and also occasionally damage nets. But, as Van Gelder and others have urged, it would be desirable to pass a law protecting all seals on Long Island, while this interesting species is still present, for the benefit of the vast numbers of people who enjoy seeing seals. For many years the harbor seal has been scarce com¬ pared with its former abundance in days before the heavy settlement of the coast, its history paralleling many of the cetaceans in this respect. Thus BeKay (1842) men¬ tions the comparative scarcity of harbor seals in his time compared with their former great abundance, while much earlier Benton (1670) wrote of the “innumerable multi¬ tude of seals” present all winter on the marshes, beaches, and sandbars of the south shore of Long Island. Harbor seals in the western North Atlantic are usual¬ ly designated P.v. concolor. The type locality of this form is Long Island Sound, near Sands Point, Nassau County, New York. However, the seals frequenting our coastline are very similar to those of the eastern North Atlantic and Europe, which are designated a different subspecies, P.v. vitulina. 55 Harp Seal Pagophilus groenlnndicus (Erxleben) There seem to be few records for the Long Island area of seals other than the harbor seal. The harp seal, gray seal, and hooded seal undoubtedly visit these shores on occasion; a few stray individuals of all three have been recorded along the New Jersey coast south of Long Island. These seals have centers of abundance in the region of Newfoundland and the Gulf of St. Lawrence. Harp seals and hooded seals migrate to that region from farther north each winter. Gray seals and the now much depleted hooded seals number a few thousand each, while the harp seals are more numerous but even they are declining. Long Island is situated south of the regular range of these northern seals and only an occasional wanderer can be expected. According to Linsley (1842) the harp seal was very rare on the Connecticut coast and was seen only at Ston- ington a few times on the rocks; he was told of an un¬ successful attempt to take a seal there, which from the description be believed to be this species, during the winter of 1841-42. This area is less than 3 miles from Fishers Island, New York, near the entrance to Long Island Sound. Kieran (1959) briefly mentions that the harp seal has been seen offshore at Coney Island in winter; I do not have additional information on this report. Stragglers have occurred as far south as Virginia. The gray seal and hooded seal are large spotted species (larger than the harbor seal), while the harp, which is about the size of the harbor seal, is pale grayish or yel¬ lowish, usually distinctively marked when adult with a large irregular, dark band on the back. Gray Seal llalichoerus grypus (Fabric! us) An interesting find, confirmed in 1958, was that of a small southern outpost colony of gray seals located in the vicinity of the west end of Nantucket (Andrews and Mott, 1967). The gray seals, or “horseheads” as they are known locally, are present year round there. The Nantucket lo¬ cality is some 80 miles east-northeast of Montauk Point. Southward, a gray seal appeared at Atlantic City, New Jersey, after a severe northeast storm in March 1931 (Goodwin, 1933). I know of no definite reports of this species on Long Island, but it should be looked for around the east end of the island. The relatively deserted coasts and subsidiary islands favored by harbor seals in this area may very likely shelter an occasional wandering gray seal from the Nantucket colony or elsewhere. Young gray seals are known to disperse widely. This species may be distin¬ guished from the harbor seal by its larger size, heavier head, and longer, more pointed snout, and by its slower, more deliberate manner; the color is variable, but usually it is a shade of gray, marked with spots or blotches. Archaeological finds indicate Indians killed gray seals in prehistoric times as far south as Block Island, about 15 miles from Montauk Point, and also on the Connecticut coast (Waters, 1967). Harp seal remains have also been found on Block Island, but as far as I know, the harbor seal (Phoca vitulina) is the only seal which has been reported from material collected at Long Island archae¬ ological sites. Hooded Seal Cystophora cristata (Erxleben) At least two hooded seals have been found near western Long Island. According to DeKay (1824, 1842), an adult male over 7 feet long was killed at Eastchester and then put on exhibition in New York City. DeKay (1824) wrote: “It was taken in a small creek emptying into Long Island Sound at Eastchester, about 14 miles from this city. The animal made considerable resistance, but exhibited no symptoms of fear. The captor suc¬ ceeded in diverting his attention by means of a dog, and was thus enabled to destroy him by repeated discharges of his musket.” This fearless, even pugnacious, behavior in face of attack is characteristic of male Cystophora. Eastchester, which is not shown on the map, is located on the New York mainland side of the Sound, a few miles across from western Nassau County. Also, within just the last few years a hooded seal was found in a tribu¬ tary of New York Harbor, according to unpublished information, but I have been unable to locate specific details on this recent record. White-tailed Deer Odocoileus virginianus Zimmerman The white-tailed or Virginia deer has had a long and varied history on Long Island. Archaeological finds and early Colonial records show that for centuries this was the principal big game species bunted by Long Island Indians and was one of their important sources of food. Van der Donk (1656) in referring to New York 56 in general (Colony of New Netherlands), wrote: “The deer are incredibly numerous in the country. Although the Indians throughout the year and every year (but mostly in the fall) kill many thousands, and the wolves, after the fawns are cast, and while they are still young, also destroy many, still the land abounds with them everywhere, and their numbers appear to remain un¬ diminished.” On Long Island, the number of deer de¬ clined with the spread of white settlers throughout the area; this was followed by enactment of various laws designed to conserve the deer herd. DeKay (1842) claimed that, in New York State, attacks of men and wolves were daily decreasing the number of deer, but then went on to say: “In some insulated districts, as on Long Island, where the wolf has been extirpated and the deer are placed under the pro¬ tection of the laws during the breeding season, although more than a hundred are annually killed by sportsmen, yet it is believed that their number is actually on the increase.” Audubon and Bachman (1851) likewise re¬ marked that on Long Island, where the game laws were strictly enforced, deer seemed to be increasing. The deer population continued to decline later in the 1800’s, however. According to Helme (1902) the deer had by then become restricted to a small area (6 X 4 or 5 miles) near the south shore of Suffolk County in the townships of Islip and Brookhuven. Pro¬ tection afforded by game preserves and private estates enabled the deer to survive in this area. White-tailed deer were also introduced from the mainland. In recent decades deer have increased again, and have spread widely over much of the island, although the expanding metropolitan area restricts their numbers in the western portion. At the present time deer are numerous on the eastern half of Long Island wherever there is suitable cover, in fact they have become a nuisance in some areas. Hamilton (1949) wrote that the Suffolk County deer herd, then numbering between 1,500 and 2,000 animals and increasing, causes considerable damage to vegetables, principally potatoes, and to nursery stock, such as the growing sprouts of young apples and other fruits. Hamilton predicted that in a few years radical measures would have to be considered to control the number of deer. Mild winters, abundance of food, no large predators except dogs, and the general absence of hunting have favored deer abundance in the more thinly settled areas of Long Island. Many are killed by dogs and automobiles, but the deer have continued to increase j and now number several thousand individuals. I In January 1969, a special deer hunt was held on Long Island, the first deer hunt since 1928; with permits carefully regulated by the State Conservation Depart¬ ment, 162 deer were taken by hunters using shotguns during two 5-day periods (Jackson and Miller, 1969). We saw deer or their tracks at most of our trapline localities in Suffolk County. Deer were encountered in all sorts of woods, including the inland pine barrens, where they are numerous, and around fields, ponds, and bogs. Occasionally they were seen in woods on the very edge of large residential areas. Deer are abundant all through the Montauk area. Here they are easily observed and often appear quite tame in the extensive open and shrubby areas such as the Montauk Downs; we noted groups of up to 16 or more here in the evening. Signs of deer were even found on the ocean shore at the very end of Montauk Point. At Hither Hills, deer are common in oak woods and other habitats. In Peconic Bay, deer were met with on Jessup Neck, and I understand they are present on Robins Island. On the north shore they are present locally in deciduous woods overlooking Long Island Sound, east to the vicinity of Orient. Roy Latham, in a recent per¬ sonal communication, writes: “Orient is overrun with white-tailed deer. We counted 28 in a field east of us early in the spring (1970) and I had a herd of nine in my little garden; they have done much damage to plants and small trees.” In western Suffolk County, deer are regular but uncommon in the vicinity of the Kalbfleisch Research Station, Dix Hills near Huntington (Lanyon, 1961) and are said to be numerous in Heckscher State Park on Great South Bay. Deer were seen on visits to Gardiners Island, Shelter Island, and Fire Island (including the Sunken Forest). Controlled marketing is practiced on Gardiners Island, where deer are very abundant (Anon., 1968a). Gardiners Island, incidentally, has had a uniquely long record of one-family ownership, and the deer were carefully pro¬ tected in the days when they had disappeared elsewhere on eastern Long Island. I am not familiar with the his¬ tory of this deer herd, but Dutcher and Dutcher (1893) in their account of the mammals of Montauk wrote, “Deer sometimes swim across from the Gardiners Island game preserve some three miles away, causing tremendous ex¬ citement and great loss of breath among all the native inhabitants of Montauk.” OTHER MAMMALS Vanished Recent Mammals A number of modern-day species of mammals which evidence indicates lived (or probably lived) on Long Island, have become extirpated. Several members of the original mammalian fauna which coexisted with the Indians, including all of the larger carnivores, dis¬ appeared early in the settlement of Long Island by the Dutch and English. Little is known about these formerly important species, which include the gray wolf {Canis lupus), bobcat {Lynx rufus), black bear {Ursus Ameri- canus), and beaver {Castor canadensis). Because this area is coastal and insular, and was heavily settled early in Colonial times, most of the large wild animals were rather quickly exterminated. A few mountain lions, or panthers {Felis concolor), may have been present then, too, but there seems to be no information about them on Long Island; if a few of these large cats roamed the island originally, they probably disappeared very early in the Colonial period, which is apparently what hap¬ pened in various other areas along the East Coast. The white-tailed deer is the one large land mammal which managed to survive to the present, although severely re¬ stricted at one time to one or two small protected areas of Suffolk County, and the present stock is much diluted by animals introduced from the mainland. Denton (1670) mentioned the presence of deer, bears, and wolves on Long Island, besides various small furbearers. A number of early documents and accounts refer to the great abundance of wolves and bobcats, upon which the settlers made war for many years. In the 1600’s and beginning of the 1700’s bounty payments for wolves killed were made by towns throughout Long Island from Brooklyn to East Hampton (Thompson, 1839, and other sources). Often it was required that the heads of the wolves be displayed in public, such as hav¬ ing them nailed to the door of the constabulary. Large pits and guns set overnight were among the methods used in effecting their capture. As for the bobcat, DeKay (1842) related that it was believed to be extirpated on Long Island then, but that it was so numerous 130 years earlier that the General Assembly passed an act [regard¬ ing bounty payments] to encourage the destruction of wildcats in Suffolk County, and that this act was renewed in 1745. Very little is on record concerning bear and beaver here, except that apparently they did occur in early historic times. Thompson (1839) recounts that beavers were reportedly common at a well-known body of water called Beaver Pond near an Indian settlement not far from present-day Jamaica in Queens, during the time of first settlement of the area. Murphy (1964) has been unable to find anything in the literature about beaver trapping on Long Island, but has found the term “beaver dam” in written records. Latham (1940) lists beaver, bear, wolf, and bobcat as recorded for the Town of Southold during the Colonial period, but with dates of latest captures uncertain. Van der Donk (1656) described in much greater detail than Denton the abundance and habits of animals in early Colonial days. Van der Donk, however, wrote not about Long Island or of any one area in particular, but of the entire New Netherlands colony, the area now consisting of southern and eastern New York State and parts of adjacent states. But his words help give some idea of the abundance of certain species in the general region in the early and mid-1600’s, including the follow¬ ing mammals then present on Long Island as well as on the mainland: beavers — -“numerous,” wolves — “numer¬ ous,” and preying on deer, calves, sheep, etc.; and bears — “many.” Mountain lions, however, were evidently very scarce or absent near the settled areas and coast according to Van der Donk, although known to the Dutch settlers from skins brought in by the Indians to sell. Scanty remains of some large game mammals have been found at Indian archaeological sites on the island, species which did not exist on the island in colonial times, but which are still found today on the North American mainland. These include wapiti or elk {Cervus canadensis), moose {dices alces) , and apparently bison {Bison bison). These species, as well as some others which survived to early Colonial days, are in Latham’s archaeological collection from eastern Long Island. He writes (personal communication), in response to a re¬ quest for information on vanished species, as follows: “I have teeth from the black bear from Indian village sites on Shelter Island, Three Mile Harbor, and Southold, I have a section of an antler from a wapiti from a pit on an Indian site in Noyack near Sag Harbor and a sec¬ tion of a moose antler from an Indian site in Acquebogue, These antlers were determined by [Robert T.j Hatt. I have the leg bone of a bison from Noyack. This member was identified at the U.S. National Museum as ‘either a bison 57 58 or a domesticated cow.’ It could not have been a cow as it was in the bottom of a 4-foot deep pit in ground not disturbed since the Indian occupation in the pre¬ historic period. Remains of beaver have been found in all Indian sites from Montauk to Riverhead. None of the Indian sites excavated have been contact sites. We found wolf remains, but we know that they were common in early settler times. We must take into consideration that antlers and even bear teeth could have been brought to Long Island in trade.” In a later letter Latham adds that the Shelter Island bear has been dated at about 1,000 B.C.; also he recorded a bobcat from a Shelter Island site. Latham feels that the wapiti, moose, and bison re¬ mains, being of late prehistoric times (200-300 years before white 'settlement), were probably brought over from the mainland by the Indians. Moose remains have been found on Fishers Island; Goodwin (1935) mentioned a fragment of an antler found in an Indian shell heap there, and Robert Cush¬ man Murphy (personal communication) recalls that the late Harry Ferguson uncovered moose teeth on Fishers Island and had them identified by a qualified authority. Remains of Indian dogs have also been reported from Long Island archaeological sites. The Dutch settlers of New Netherlands in the early 1600’s had knowledge of living wapiti, moose, and bison, but not in the immediate vicinity of their coast. For ex¬ ample, it is well known that Van der Donk (1656) was familiar with bison. At the beginning of a rather long passage on this species, he wrote: “Buffaloes are also tolerably plenty. These animals mostly keep towards the southwest where few people go.” A well documented example of a small mammal (sub¬ species) which became extinct quite recently (late 1800’s) in the Long Island area because of the complete destruc¬ tion of its habitat by man is the Gull Island mouse [Microtus pennsylvanicus nesophilus) , which is discussed in the species account. Other small, inconspicuous mam¬ mals may have disappeared without notice in the years following European settlement of Long Island. Fox squir¬ rels vanished during the last century from the Northeast; the subspecies known as the northern fox squirrel (Sciurus niger vulpinus) once ranged north to Connecti¬ cut and New York. But Bangs (1896) wrote that this shy form could not withstand persecution, and clearing and settlement of the land. For Long Island, there seems to be little information available concerning vulpinus, and also much confusion existed in the past regarding the various races and color forms of fox and gray squirrels. But Audubon and Bachman (1849) apparently had this subspecies primarily in mind when they wrote that the “cat squirrel” [Sciurus cinereus) , which they explain is the fox squirrel of New York, Pennsylvania, and New Jersey, and intermediate in size between the gray squirrel and the fox squirrel of the southern states, “is rather a rare species — and is met with on Long Island and some other portions of the state of New York.” Introduced Mammals The Norway rat and house mouse are familiar asso¬ ciates of man which came over from the Old World. So did the black rat, which is now rare. At least one and probably more native American species also made their way to Long Island by one method or another during the historic period. This is fairly well documented in the ease of the opossum. Other species, such as the eastern cottontail, and possibly the little brown bat, also may have invaded the island and prospered in response to changing ecological conditions since the coming of the white man. All of these well-established mammals are dis¬ cussed in the species accounts. But in addition to these. Long Island has played host to an unusually large number of deliberately im¬ ported and escaped exotic mammals, some of which have become temporarily established locally. The many wealthy landowners and sportsmen, with their private estates and game preserves, and numerous travelers re¬ turning from other lands have contributed to this situa¬ tion. Also certain outlying islands have proved to be havens for some of the newcomers. None of the species has become widely distributed, as far as I know, and lacking details on the history and status of most of them, they will be discussed only briefly. But the possibility of encountering such animals is something to be kept in mind by the student or observer afield in this area. Various hares and rabbits have been introduced, besides western races of Sylvilagus floridanus, and some of these may be found locally in small numbers. These include — European rabbit (Oryctolagus cuniculus) , said to be on Great Gull Island and possibly one or two areas on Long Island proper; black-tailed jackrabbit [Lepus californicus) , reported in some limited areas of western and central Long Island in recent years; European hare [Lepus europeaus) , a few reportedly in Nassau County, possibly also central Suffolk County; and varying hare [Lepus americanus) , introduced in several localities years ago. Some varying hares introduced into Orient in 1915 lasted about 4 years (Roy Latham, personal communica¬ tion). Among the hares [Lepus), authentic recent records (within the past 2-3 years) are known only for the black¬ tailed jack, of which two skins have been examined 59 (Joseph Dell, Anthony Taormina, personal communica¬ tions). As far as I know, the varying hare was not a native resident of Long Island during historic times. The citing of Locust Grove, Nassau County, as a peripheral locality record for this species by Hall and Kelson (1959) ap¬ pears to be an error. After tracing this record back to Nelson (1909), who did not include Long Island in the range of this species, I believe the probable location for this record is the very small village of Locust Grove in Lewis County (northwestern New York), where C. Hart Merriam collected many specimens in the late 1800’s, rather than the much larger town of the same name on Long Island. For deer, there are black-tailed deer {Odocoileus hemionus) on Shelter Island and sika deer {Sika nippon) near the Carmans River. The former, which are well known to the local residents, were introduced in 1910 and still survive in an undeveloped area of Shelter Island; I noted several black-tailed and white-tailed deer asso¬ ciating together and feeding in an open field there in June 1963. The sika deer were introduced at Suffolk Lodge (the Hard property on the Carmans River, now a county park) many years ago; the small herd remains in a wild state, but is largely confined to the park (Anthony Taormina, personal communication). Beavers {Castor canadensis) have been reintroduced and established locally and temporarily, but I have no knowledge of any presently-existing colonies. Other oddities include escaped hedgehogs {Erinaceus euro- paeus) in central Long Island, and thirteen-lined ground squirrels {Citellus tridecemlineatus) on Fishers Island; I have not heard of any recent reports of these species. The hedgehogs are said to have escaped from green¬ houses where they were kept to control insects (as is done in Europe). Even escaped California sea lions {Zalophiis calif or - nianus) have been encountered. In July 1965, about 15 escaped from a marineland at Ocean City, Maryland, and although most of them eventually headed south, two of them were recovered as far north as the Long Island area (Robert A. Morris, personal communication). Missing Land Mammals Long Island lacks several kinds of small mammals presently found on the nearby mainland or in similar terrain along the New Jersey coastal plain. For some of the species this seems surprising, but probably the situa¬ tion is about what one might expect for a large island near the mainland — the island fauna is mostly similar, but has somewhat fewer species. Certain forms apparently did not make the short jump from the mainland at the western end of the island, or if they occurred earlier, conditions may have become unsuitable for them in the limited Long Island area. With the paved metropolis now blocking this approach, Long Island seems more effec¬ tively isolated than previously to natural invasion by four-footed land mammals (although not to accidental or intentional introduction by man). The red squirrel {Tamiasciurus hudsonicus) is miss¬ ing, without any evidence that it ever occurred here as far as I know, although it is found in southern New England and also abundantly southwards on the New Jersey coastal plain in pitch pine — oak woods resembling those growing on Long Island. The red squirrel formerly occurred as close as Bronx and Manhattan on the other side of the East River. The red-backed mouse {Clethrion- omys grapperi) has never been found either, although it is present in southern New England and southern New York and also south of Long Island in the coast white cedar swamps and sphagnum bogs of New Jersey; if it existed on Long Island one would expect it in these same habitats and possibly also in red maple swamps. The bog lemming {Synaptomys cooper i) has appar¬ ently escaped all search for it on Long Island. This species is often local and uncommon where it occurs and there is the possibility of it having been overlooked, but it seems unlikely that Synaptomys remains undiscovered in such a heavily settled area as Long Island. The scat¬ tered sphagnum bogs of the pine region would be the most likely habitat. There are no records of the least shrew {Cryptotis parva), to^ my knowledge, although there is still some chance it may be found here. More than most small mammals it is apt to go unnoticed, and it appears to be rare and local this far north. Cryptotis has been found twice on the Connecticut coast, at Darien and Westbrook (see Jarrell, 1965) and several have been collected on Staten Island (Chapin, 1908). Two small bats, the Indiana myotis {Myotis sodalis) and the small-footed myotis {Myotis leibii) have not been found on Long Island as far as I know. Both are rare in the Northeast and known primarily from winter hibernating sites in caves. If a few individuals occasion¬ ally visit Long Island or even spend the summer here they would be difficult to detect. 60 Figure 1. Woods of large beech, tuliptree, oaks, etc., surviving near North Hills, western Nassau County, but encircled by the expanding metropolitan area. March, 1961. Short-tailed shrew, eastern mole, cottontail, chipmunk, gray squirrel, white¬ footed mouse, pine mouse, and raccoon recorded. Figure 2. Same woodland area as figure 1, with small stream. 61 62 Figure 5. Peconic River, near Calverton. July 1963. Masked shrew, meadow mouse, and muskrat numerous in cattails and other marsh vegetation. Otters have been reported in this general area of the river. Figure 6. Looking towards Great Peconic Bay marshes from pine land east of Flanders. November 1962. Eastern mole, chipmunk, and pine mouse common under the pines, but not venturing out into salt marsh (the home of many meadow mice). Also masked shrew, eastern cottontail, Norway rat, white-footed mouse, raccoon, long-tailed weasel, and white-tailed deer found in the general area. 63 Figure 7. Edge of southern white cedar swamp (pine-oak woods to right), near River- head. November 1962. Flying squirrels observed in the cedars; masked shrew, short¬ tailed shrew, eastern cottontail, gray squirrel, and white-footed mouse collected under the cedars. Eastern mole tunnels in dry ground near edge. Also chipmunk, woodchuck, red fox, and white-tailed deer recorded in the general area. Figure 8. Within cedar swamp shown in figure 7. Masked shrews numerous in cavi¬ ties at the base of cedar trees and mounds. 64 Figure 9. Small sphagnum bog with southern white cedar, leatherleaf, and a large sedge, near Flanders. October 1962. Masked shrew and meadow mouse numerous. Also eastern cottontail, southern flying squirrel, white-footed mouse, muskrat, raccoon, long-tailed weasel, and white-tailed deer recorded. Figure 10. Abandoned farmhouse near Manorville. November 1962. Summer mater¬ nity colony of little brown bats in attic, just under peak of roof; bats flew in and out of upper window. Also a few big brown bats present in fall. 65 Figure 11. Weedy, sandy field with black oak and young pitch pines, Hampton Bays. November 1962. Short-tailed shrew and pine mouse abundant, meadow mouse uncom¬ mon. Also eastern mole tunnels throughout the area. Figure 12. Shinnecock Inlet, from base of jetty on west side. Harbor seals in inlet and vicinity during winter, bottle-nosed dolphin found stranded nearby. Also records of pigmy sperm whale, finback whale, and Atlantic right whale off the inlet in recent years. Norway rats live among the jetty rocks. 66 Figure 13. Red maple swamp with small stream, cinnamon fern, skunk cabbage, vines, etc., near Babylon. April 1963. Several star-nosed moles (very local on Long Island) collected in area shown, in tunnels near the stream. Short-tailed .shrew and meadow mouse used the mole tunnels. Also opossum, cottontail, gray squirrel, white-fooled mouse, and raccoon listed. Figure 14. Star-nosed mole above deep tunnel where caught (in area shown in figure 13). April 1963. 67 Figure 15. Grassland with clumps of shrubs and scattered trees, in Shinnecock Hills. November 1962. Masked shrew, short-tailed shrew, and meadow mouse numerous; eastern mole in some depressions. Red foxes observed and dens found in area shown. Opossum and white-tailed deer common. Figure 16. Salt marsh on outer strip about 1 mile east of Moriches Inlet. November 1962. Meadow mouse abundant, masked shrew present. Also muskrat, long-tailed weasel, and mink recorded in this marsh, with opossum, eastern cottontail, and red fox common in general area. 68 Figure 17. View from outer dunes (wilh beachgrass) across interdune valley to inner dunes, East Hampton Beach. November 1962. Meadow mouse, house mouse, and cot¬ tontail rabbit frequented the outer dunes. More mammals were found in the interdune valley, including the above species and also masked shrew, short-tailed shrew, eastern mole, and white-footed mouse. Opossum, red fox, and long-tailed weasel in the area. Figure 18. Sandy, tail-grass area near the moving dunes (background) at Hither Hills. Meadow jumping mouse, as well as meadow mouse and masked shrew, very common in the tall grass here. 69 Figure 19. View across extensive marshes and Napeague Harbor (left) from summit of high moving dune in Hither Hills State Park. November 1962, Meadow mice everywhere here, even in clumps of beachgrass on the high dunes. Red fox common. Figure 20. Moving dunes, showing active edge of a dune encroaching on a patch of woods, Hither Hills State Park. November 1962. Tracks of cottontail, red fox, rac¬ coon, striped skunk, and white-tailed deer frequent on the smooth sand. 70 Figure 21. Woods of white oak and black oak with bayberry undergrowth, in Hither Hills near shore of Block Island Sound. October 1962. Our easternmost locality for the eastern mole. Also chipmunk and pine mouse present, local this far east. Both shrews, gray squirrel, white-footed mouse, and white-tailed deer common. Figure 22. Fresh Pond, Hither Hills State Park. October 1962. Keen’s bats abundant and active over the pond and vicinity at least from June to October. Red bats com¬ mon in late summer and fall. Both species mist-netted over water and shoreline on left. Red fox and mink observed along the shore. 71 Figure 24. Tubers of groundnut or wild bean, from a large pile accumulated by mead¬ ow mice, near Hauppauge. Collected in March 1961. Individual tubers about 1 inch long. Figure 23. Typical scene under a pitch pine in the pine barrens — husked cones and litter left by squirrels feeding on the pine seeds. 72 Figure 25. Sandy area with beachgrass, bayberry, etc., in foreground, near salt water bay on Shelter Island. June 1963. Meadow jumping mouse present, also short-tailed shrew, white-footed mouse, meadow mouse, and raccoon listed, (osprey nest on dead tree in distance). Figure 26. Field on Slielter Island. June 1963. Black-tailed deer (introduced) and white-tailed deer observed feeding here. Red fox sat on rock under lone tree in center distance. Short-tailed shrew, meadow mouse, and eastern cottontail common. LITERATURE CITED Allen, G. M. 1916. The whalebone whales of New Eng¬ land. Mem. Boston Soc. Nat. Hist., 8(2) :107-322. - 1941. Pygmy sperm whale in the Atlantic. Field I Mus. Nat. Hist., Zool. Ser., 27:17-36. Andrews, J. C. and Mott, P. R. 1967. Gray seals at Nan¬ tucket, Massachusetts. J. Mamm., 48(4) :657-658. Andrews, R. C. 1908. Notes upon the external and in¬ ternal anatomy of Balaena glacialis Bonn. Bull. Amer. Mus. Nat. Hist., 24(10) :171-182. - 1909. Further notes on Eubalaena glacialis (Bonn.) Bull. Amer. Mus. Nat. Hist., 26 (21) : 273-275. - 1916. Whale hunting with gun and camera. D. Appleton and Co., New York. 333 pp. Anon. 1956. Stranded mammals of the sea. N.Y. State Conservationist, 11(1) :4-5. Anon. 1968a. The natural history of Long Island — the land mammals. Sanctuary (publication of the L.L Chapter of the Nature Conservancy), 2-19. Anon. 1968b. Rare and endangered fish and wildlife of the United States. Compiled by Committee on Rare and Endangered Wildlife Species, Bureau of Sport Fisheries and Wildlife (Resource Publ. 34), Wash¬ ington, D.C. Audubon, J. J. and Bachman, J. 1841. Descriptions of new species of quadrupeds inhabiting North Amer¬ ica. Proc. Acad. Nat. Sci. Phila., 1:92-103. - 1849-54. The quadrupeds of North America. V. G. Audubon, New York. 3 Vols. (Vol. 1, 383 pp., 1849; Vol. 2, 334 pp., 1851; Vol. 3, 348 pp., 1854). Bailey, V. 1898. Descriptions of eleven new species and subspecies of voles. Proc. Biol. Soc. Wash., 12:85-90. - 1900. Revision of American voles of the genus Microtus. N. Amer. Fauna, No. 17, 88 pp. Bangs, O. 1896. A review of the squirrels of eastern North America. Proc. Biol. Soc. Wash., 10:145-167. Barbour, R. W. and Davis, W. H. 1969. Bats of America. The University Press of Kentucky. 286 pp. Benton, A. and Atkinson, T. 1964. Notes on the New England cottontail in eastern New York, N.Y. Fish and Game J., 11(2) :154-156. Brodo, I. M. 1968. The lichens of Long Island, New York: a vegetational and floristic analysis. N.Y. State Mus. and Sci. Serv. Bull. 410. 330 pp. Carter, D. T. 1950. On the migration of the red bat, Lasiurus borealis borealis. J. Mamm., 31(3) :349- 350. Chapin, J. 1908. Notes on some Staten Island mammals. Proc. S.I. Assoc. Arts and Sciences, 2(2):65-67. Chapman, F. M. 1908. Camps and cruises of an orni¬ thologist. D. Appleton and Co., New York. 432 pp. Clark, J. 1965. Occurrence of the bottlenose porpoise in a tidal river. Underwater Naturalist, 3(3): 19-20. [Amer. Littoral Soc., Sandy Hook Marine Lab., Highlands, New Jersey] Cline, M. G. 1955, Soils and soil associations of New York. Cornell Ext. Bull. 930. 72 pp. Cook, J. J. and Wisner, W. L. 1963. Killer Whale! Dodd, Mead, and Co., New York. 64 pp. Crowe, P. E. 1939. The least weasel on Staten Island, New York. Proc. S.I. Inst. Arts and Sciences, 8:135. Davis, W. T. 1908. Observations on Staten Island moles. Proc. S.I. Assoc, Arts and Sciences, 2(3) :141. - 1920. A Staten Island whale. Proc. S.I. Inst. Arts and Sciences, 1 :56. DeKay, J. E. 1824. An account of the Phoca cristata, recently taken in the vicinity of this city. Annals Lyceum Nat. Hist. N.Y., 1:94-99. - 1842. Zoology of New York. Pt. 1, Mammalia. Albany. 146 pp. Denton, D. 1670. A brief description of New York; for¬ merly called New Netherlands. London, printed for John Hancock. (1902 reprint, Burrows Bros. Co., Cleveland) . Dutcher, B. H. 1889. Bird notes from Little Gull Island, Suffolk Co., N.Y. Auk. 6(2) : 124-139. Dutcher, W. and Dutcher, B. H. 1893. [MS notes on mammals of Montauk Point, Suffolk County, N.Y.] 7 pp., holograph; original on file in U.S. Nat. Mus. Edwards, E. J. and Rattray, J. E. 1932. Whale Off! — The story of American shore whaling. Frederick A. Stokes Co., New York. 285 pp. (also reprinted 1956, Cowafd-McCann, New York.) Edwards, R. L, and Livingstone, R., Jr. 1960. Observa¬ tions on the behavior of the porpoise Delphinus delphis. Science, 132 (3418) :35-36. Fay, F. H. and Chandler, E. H. 1955. The geographical and ecological distribution of cottontail rabbits in Massachusetts. J. Mamm., 36(3) :415-424. Free, G., Winkelblech, C. and Wilson, H. 1957. Soil erosion on Long Island — its control. Cornell Ext. Bull. 968. 31 pp. Fuller, M. L. 1914. Geology of Long Island. U.S. Geol. Surv., Prof. Paper 82. 231 pp. 73 74 Goodwin, G. G. 1933. Occurrence of a gray seal at At¬ lantic City, New Jersey. J. Mamm., 14(1) ;73. - 1935. The mammals of Connecticut. Conn. State Geol. and Nat. Hist. Surv., Bull. 53. 221 pp. Hall, E. R. and Kelson, K. R. 1959. The mammals of North America. Ronald Press Co., New York. 2 Vols. Hamilton, W. J., Jr. 1935. Notes on food of red foxes in New York and New England. J. Mamm., 16(1): 16-21. - 1943. The mammals of eastern United States. Comstock Publ. Co., Ithaca, New York. 432 pp. - 1949. The mammals of Long Island. In Long Island, edited by Paul Bailey, Lewis Historical Publ. Co., New York. Vol. 2, Chapter 28:151-161. Handley, C. 0., Jr. 1966. A synopsis of the genus Kogia (pygmy sperm whales). In Whales, dolphins, and porpoises, edited by K. S. Norris, Univ. of Califor¬ nia Press. Chapter 4:62-69. Hatt, R. T. 1930. Color varieties of Long Island mam¬ mals. J. Mamm., 11(3) :322-323. Helme, A. H. 1902. Notes on the mammals of Long Island, New York, Abstr. Proc. Linnacan Soc. N.Y., 13, 14:19-30. - 1927. Reminiscences of a Long Island Naturalist. Brooklyn Mus. Quart., 14(3) :98-105. Helmuth, W. T. 1931. Balaennptera acutorostrata off Montauk Point, New York. J. Mamm., 12(1) :72-73. - 1954. The hurricane of 1938 — in retrospect. Birds of Long Island, 8:225-241. Hershkovitz, P. 1966. Catalogue of living whales. U.S. Nat. Mus., Bull. 246. 259 pp. Hinderstein, C. 1969. Variation of cranial characters of two cottontails, Sylvilagus floridanus and S. transi- tionalis in a region of sympatry, with special regard to species status. Abstracts of papers presented at 49th annual meeting of Amer. Soc. Mammalogists at New York, N.Y. No. 86. Mimeo. Howell, N. R. 1941. Long Island whaling. Long Island Forum, Bay Shore, New York. 13 pp. Jackson, H. H. T. 1915. A review of the American moles. N. Amer. Fauna, No. 38. 100 pp. Jackson, L. and Miller, R. 1969. The deer take in 1968-69. The Conservationist, 23(6) :13-14,34. Jarrell, G. H. 1965. A correction on the range of Cryp- totis parva in New England. J. Mamm., 46(4)671. Kellogg, R. 1929. What is known of the migration of some of the whalebone whales. Ann. Rept. Smith¬ sonian Inst., 1928:467-494. Kenyon, K. W., et al. 1965. Report of the standing com¬ mittee, marine mammals, 1964-65, submitted at 45th annual meeting of the Amer. Soc. of Mammalogists ■ at Winnipeg, Manitoba. 58 pp., cyclostyled. Kieran, J. 1959. A natural history of New York City. Houghton Mifflin Co., Boston. 428 pp. Kimball, H. M. and Nichols, J. T. 1940. A Long Island hoary bat. J. Mamm., 21(2) :214-216. Lanyon, W. E. 1961. The vertebrates of the Kalbfleisch Field Research Station. Amer. Mus. Nat. Hist. 28 pp. ; Latham, R. 1940. The natural history of Southold Town. The Suffolk Times, June 27, 1940. - 1954. Nature notes from Orient. The Long Island Naturalist, 3:3-9. [Baldwin Bird Club] Linsley, J. H. 1842. A catalogue of the mammalia of Connecticut. Amer. Jour. Sci., 43(7) :345-354. Mackiewicz J. and Backus, R. H. 1956. Oceanic records of Lasionycteris notivagans and Lasiurus borealis. J. Mamm., 37(3) :442-443. McCulley, H. 1967. The broad-handed mole, Scapanus latimanus, in a marine littoral environment. J. Mamm., 48(3) :480-482. Miller, G. S., Jr. 1899. Preliminary list of the mammals of New York. Bull. N.Y. State Mus., 6(29) :271-390. - and Kellogg, R. 1955. List of North American recent mammals. U.S. Nat. Mus., Bull. 205. 954 pp. Moore, J. C. 1966. Diagnoses and distributions of beaked whales of the genus Mesoplodon known from North American waters. In Whales, dolphins, and por¬ poises, edited by K. S. Norris, Univ. of California Press. Chapter 3:32-61. Murphy, R. C. 1918. A sperm whale ashore near New York. Forest and Stream, 88:288. - 1964. Fish-shape Paumanok — nature and man on Long Island. Amer. Phil. Soc., Philadelphia. 67 pp. - and Nichols, J. T. 1913. Long Island fauna and flora — 1. The bats. Brooklyn Mus., Sci. Bull., 2(1): 1-15. Nelson, E. W. 1909. The rabbits of North America, N. Amer. Fauna, No. 29. 314 pp. Nichols, D. G. and Nichols, J. T. 1934. Notes on Long Island, New York, bats. J. Mamm., 15(2) :156. - 1935. Notes on the New York weasel {Mustela novebnracensis) . J. Mamm., 16(4) :297-299. Nichols, J. T. 1907. The star-nosed mole on Long Island, N.Y. Amer. Naturalist, 41(486) :401. - 1927. Notes on the food habits of the gray squir¬ rel. J. Mamm., 8(1) :55-57. - 1958. Food habits and behavior of the gray squirrel. J. Mamm., 39(3) :376-380. Peters, G. H. 1949. The florai of Long Island. In Long Island, edited by Paul Bailey, Lewis Historical Publ. Co., New York. Vol. 2, Chapter 27:137-150. 75 I 1 Raven, H. C. 1934. Beaked whales. Natural History, I 34(5) :501. i - 1937. Notes on the taxonomy and osteology of ^ two species of Mesoplodon (M. europaeus Gervais, I M. mirus True). Amer. Mus. Novitates, 905:1-30. , - 1942. On the structure of Mesoplodon densirostris, ; a rare beaked whale. Bull. Amer. Mus. Nat. Hist., 80(2) :23-50. - and Gregory, W. K. 1933. The spermaceti organ and nasal passages of the sperm whale [Physeter catodon) and other odontocetes. Amer. Mus. Novi- I tates, 667:1-18. Ray, C. 1961. A question in whale behavior. Natural His¬ tory, 70(6) :46-53. Raynor, G. S. 1960. Three litters in a pine mouse nest. J. Mamm., 41(2) :275. i Reed, J. H. 1898. The terns of Great Gull Island, New York, during 1897. Auk, 15(l):40-43. Rice, D. W. and Scheffer, V. B. 1968. A list of the marine mammals of the world. U.S. Fish and Wild¬ life Service, Special Scientific Report — Fisheries No. 579:-16 pp. Ritchie, W. A. 1969. The archaeology of New York State. Rev. ed. Natural History Press, New York. 357 pp. Rockwell, R.H. 1914. Bottle-nosed whales on the Long Island coast. Brooklyn Mus. Quart., 1(3) :147-150. Rowley, J. 1902. The mammals of Westchester County, New York. Abstr. Proc. Linnaean Soc. N.Y., 13,14: 31-60. Schevill, W. E. 1954. Sight records of the gray grampus. Grampus griseus (Cuvier). J. Mamm., 35(1) :123- 124. - - and Watkins, W. A. 1962. Whale and porpoise voices. A phonographic disk, with 24 pp. and 35 text figs. Woods Hole ' Oceanographic Inst. Woods Hole, Massachusetts. Schuberth, C. J. 1968. The geology of New York City and environs. Natural History Press, New York. 304 pp. Schulte, H. von W. 1917. The skull of Kogia hreviceps Blainv. Bull. Amer. Mus. Nat. Hist., 37(17) :361-404. - and Smith, M. F. 1918. The external characters, skeletal muscles and peripheral nerves of Kogia brevi- ceps. Bull. Amer. Mus. Nat. Hist., 38(2) :7-72. Seagears, C. 1944. The fox in New York. N.Y. State Cons. Dept., Albany. 85 pp. Simon, N. 1966. Red data book. Volume 1 — Mammalia. International Union for Conservation of Nature and Natural Resources. Merges, Switzerland. Sleight, Harry D. 1931. The whale fishery on Long Island. The Hampton Press, Bridgehampton, New York. 231 pp. Stone, S. 1964. Girl on a dolphin. Underwater Naturalist, 2(3) :8-ll. [Amer. Littoral Soc,, Sandy Hook Ma¬ rine Lab., Highlands, New Jersey] Stoner, D. 1938. New York State records for the common dolphin, Delphinus delphis. N.Y. State Mus. Circ. 21. 16 pp. Stout, N. J. 1958. Atlas of forestry in New York. State Univ. of N.Y., Coll, of Forestry at Syracuse, Bull. 41. Svenson, H, K. 1936. The early vegetation of Long Island. Brooklyn Bot. Gard. Record, 25 (3) :207-227. Taylor, N. 1923. The vegetation of Long Island. Part 1. The vegetation of Montauk: a study of grassland and forest. Brooklyn Bot. Gard. Mem., 2:1-107. - 1927. The climate of Long Island. Cornell Univ., Agr. Exp. Sta., Bull. 458. 20 pp. Thompson, B. F. 1839. History of Long Island. E. French, New York. 536 pp. Thorpe, M. R. 1938. Notes on the osteology of a beaked whale. J. Mamm., 19(3) :354-362. Tomilin, A. G. 1967. Mammals of the U.S.S.R. and ad¬ jacent countries. Vol. 9, Cetacea. Translated from Russian, Israel Program for Scientific Translations. 717 pp. Townsend, C. H. 1935. The distribution of certain whales as shown by logbook records of American whale- ships. Zoologica, 19(l):l-50. True, F. W. 1885. Report of a trip to Long Island in search of skeletons of the right whale, Balaena cis- arctica. Bull. U.S. Fish. Comm., 5:131-132. - 1889. Contributions to the natural history of the cetaceans, a review of the family Delphinidae. U.S. Nat, Mus., Bull. 36. 192 pp. - 1904. The whalebone whales of the western North Atlantic. Smithsonian Contrib. to Knowledge, 33:1-332. Turrell, L. W. 1939. The natural history of Smithtown. Arts-Craft Press, St. James, Long Island. 89 pp. Ulmer, F. A., Jr. 1941. Mesoplodon mirus in New Jersey, with additional notes on the New Jersey M. densi¬ rostris, and a list and key to the ziphoid whales of the Atlantic coast of North America. Proc. Acad. Nat. Sci. Phila., 93:107-122. - 1961. New Jersey’s whales and dolphins. N.J. Na¬ ture News, 16(3) :89-93. [N.J. Audubon Soc.] Van der Donk, A. 1656. A description of the New Neth¬ erlands. Edited with an Introduction by T. F. O’Don¬ nell, 1968, Syracuse University Press. 142 pp. (fol¬ lows J. Johnson’s 1841 English translation). Waters, J. H. 1967. Foxes on Martha’s Vineyard, Massa¬ chusetts. J. Mamm., 48 (1) : 137-138. - 1967. Gray seal remains from southern New Eng¬ land archaeological sites. J. Mamm., 48(1) : 139-141. 76 - and Rivard, C. J. 1962. Terrestrial and marine mammals of Massachusetts and other New England states. Standard-Modern Printing Co., Inc., Brockton, Massachusetts. 151 pp. Whitmore, F. C., Emery, K. 0., Cooke, H. B. S. and Swift, D. J. P. 1967. Elephant teeth from the Atlantic continental shelf. Science, 156(3781) :1477-1481. Willey, N. 1949. The whaling era on Long Island. In Long Island, edited by Paul Bailey, Lewis Historical Publ. Co., New York. Vol. 1, Chapter 18:533-556. APPENDIX Allen H. Benton, Danny L. Kelly, State University College, Fredonia, New York 1 4063 Siphonaptera From Long Island, New York During the study of Long Island mammals, Paul Connor collected 421 fleas which form the material for this study. We are grateful to Mr. Connor and the New York State Museum and Science Service for the oppor¬ tunity to study this interesting collection. The 421 specimens include only 11 species, a re¬ markably small number. In comparison, a study of ma¬ terial collected by Connor in Lewis County (Benton, 1966) produced 118 individuals of 13 species, while a study of Connor’s Schoharie County material (Kuczek, unpub.) produced 606 specimens of 19 species. It ap¬ pears that the flea fauna of Long Island is impoverished. In part, this is due to the absence of a number of north¬ ern forms associated with such mammal species as Clethironomys gapped and Tamiasciurus hudsonicus. In other cases, the host species may have been present but not collected or collected in such small numbers that a representative sample of the fleas was not obtained. For example, both Tamias striatus and Glaucomys volans were collected in small numbers, but such flea species as Megabothds acerbus and Tamiophila grandis from the chipmunk, and Epitedia faceta, Conor hinopsylla stanfordi, and Opisodasys pseudarctomys from the flying squirrel were not taken. Geary (1959) lists three additional mam¬ mal fleas from Long Island localities: Oropsylla arctomys, the common flea of woodchucks; Megabothris asio, usu¬ ally found on Microtus, and Odontopsyllus multispinosus, a parasite of rabbits. Possibly, the relatively small num¬ ber of species reflects the very limited altitudinal and ecological variability on the island. Certainly, one might expect more than one- third of New York’s flea fauna to be found on such a large land area. Selective collecting from species poorly represented in the State Museum collection might add several species of fleas to the list of species which follows: FAMILY PULICIDAE Cediopsylla simplex (Baker) Fourteen specimens of this rabbit flea were collected from two specimens of Sylvilagus floridanus. This is a common flea on Leporids, particularly cotton¬ tails, throughout the east except in high areas where cottontails do not exist. (2 males, 12 females) FAMILY HYSTRICHOPSYLLIDAE Atyphloceras bishopi (Jordan) Three specimens were taken from Microtus pennsyl- vanicus at Shinnecock Hills, December 15, 1961. This is primarily a winter flea, usually taken in larger numbers from nests of the field mouse. It is known from only a few localities in New York, but probably occurs throughout the State. (3 females) Stenoponia americana (Baker) Forty specimens of this species were taken, with 35 of them (87.5 percent) having been taken in the months of September-March. Six hosts were repre¬ sented, with the largest numbers being found on Peromyscus leucopus (35 percent), and Microtus pennsylvanicus (20 percent). In New York, this species has been found only on Long Island and up the Hudson Valley as far north as Albany. (15 males, 25 females) Epitedia wenmanni testor (Rothschild) Seven specimens were taken from November-March, four from Peromyscus leucopus, three from Microtus pennsylvanicus. The distribution of this southern subspecies in New York is similar to that of Steno¬ ponia americana. (2 males, 5 females) C tenophthalmus pseudagyrtes pseudagyrtes (Baker) By far the commonest flea species collected, this flea made up 30 percent of the fleas collected, with 128 specimens from 10 host species. It was taken through¬ out the year, with 80 percent occurring in the Sep¬ tember-March collections. (76 males, 52 females) Doratopsylla blarinae (C. Fox) Forty specimens of this species were taken, 39 of them from Blarina brevicauda. This species is thus the most host-specific flea taken during this study. The specimens were quite evenly distributed throughout the year, except for a surprising gap during the months of July-October. Possibly this gap was caused by a low population of the host species, or by limited collections during this time period, since it has been taken commonly during these months in other parts of New York. (18 males, 22 females) 77 78 Nearctopsylla genalis genalis (Baker) Eleven specimens were taken, six from Blarina brevi- cauda, five from Scalopus aquaticus. All were taken between December and March. (5 males, 6 females) FAMILY CERATOPHYLLIDAE Orchopeas howardii hnwardii (Baker) This common squirrel flea is represented by 113 specimens, seven from Glaucornys volans, the rest from Sciurus carolinensis. No specimens were taken from September to February (37 males, 76 females) Orchopeas leucopus (Baker) The 54 specimens of this species were well dis¬ tributed throughout the year, with peaks in Novem- ber-December. Forty-eight (88 percent) were taken from Peromyscus leucopus. (22 males, 32 females) Nosopsyllus fasciatus (Bose) The common rat flea was taken only once, a single speciment from Didelphis marsupialis. This was un¬ doubtedly due to an accidental transfer to the opos¬ sum from a barn or outbuilding where rats were present. ( 1 female) FAMILY LEPTOPSYLLIDAE Peromyscopsylla scotti (1. Fox) Ten specimens of this species were taken from Oc- tober-December, substantiating previous reports that this is an autumn flea. All except one were from Peromyscus leucopus. (4 males, 6 females) REFERENCES CITED Benton, Allen H. 1966. Siphonaptera collected on Tug Hill, Lewis County, New York. In “The Mammals of the Tug Hill Plateau, New York,” by Paul F. Connor, N.Y. State Mus. and Sci. Serv., Bull. 406: 76-78. Geary, John M. 1959. The Fleas of New York. Cornell Univ. Agr. Exp. Sta. Memoir 355: 155 pp. Kuezek, Bernard E. 1959 (Unpublished) . The Siphonap¬ tera of Schoharie County, New York. A Seminar Report presented to the Department of Biology, State Univ. of N.Y., Albany, N.Y. 16 pp. 19050 1 f82X fH .nfirmost Clinton Silurian) Stratigraphy and Petrology East-Central New York Donald H. Zenger Temporary Geologist Geological Survey New York State Museum and Science Service BULLETIN NUMBER 417 NEW YORK STATE MUSEUM AND SCIENCE SERVICE The University of the State of New York ALBANY, NEW YORK AUGUST 1971 The State Education Department Uppermost Clinton (Middle Silurian) Stratigraphy and Petrology East-Central New York Donald H. Zenger Temporary Geologist Geological Survey New York State Museum and Science Service ALBANY, NEW YORK The University of the State of Nezv York The State Education Department AUGUST 1971 THE UNIVERSITY OE THE STATE OF NEW YORK Regents of the University (ivith years ivhen terms expire) 1984 Joseph W McGovern, A.B., LL.B., L.H.D., LL.D., D.C.L., Chancellor . New York 1985 Everett J. Penny, B.C.S., D.C.S. Vice Chancellor . White Plains 1978 Alexander J. Allan, Jr., LL.D., Litt.D. . Troy 1973 Charles W. Millard, Jr., A.B., LL.D., L.H.D. . Buffalo 1972 Carl H. Pforzheimer, Jr., A.B., M.B.A., D.C.S., H.H.D. - Purchase 1975 Edward M. M. Warburg, B.S., L.H.D. . New York 1977 Joseph T. King, LL.B. . Queens 1974 Joseph C. Indelicato, M.D. . Brooklyn 1976 Mrs. Helen B. Power, A.B., Litt.D., L.H.D. . Rochester 1979 Francis W. McGinley, B.S., LL.B., LL.D. . Glens Falls 1980 Max J. Rubin, LL.B., L.H.D. . New York 1986 Kenneth B. Clark, A.B., M.S., Ph.D., Litt.D. . Hastings on Hudson 1982 Stephen K. Bailey, A.B., B.A., M.A., Ph.D., LL.D. - - - Syracuse 1983 Harold E. Newcomb, B.A. . Owego 1981 Theodore M. Black, A.B. . Sands Point President of the University and Commissioner of Education Ewald B. Nyquist Executive Deputy Commissioner of Education Gordon M. Ambach Associate Commissioner for Cultural Education John G. Broughton Assistant Commissioner for State Museum and Science Service State Paleontologist, State Museum and Science Service Donald W. Fisher 17055 CONTENTS PAGE ABSTRACT . 1 ACKNOWLEDGEMENTS . 3 INTRODUCTION . 5 PURPOSE . 5 GENERAL SETTING . 5 METHODS OE STUDY . 6 STRATIGRAPHY AND PALEONTOLOGY . 9 KIRKLAND DOLOSTONE . 9 HERKIMER EORMATION . 12 JOSLIN HILL MEMBER . 14 JORDANVILLE MEMBER . 21 STRATIGRAPHIC RELATIONS AND CORRELATION. . 25 PETROLOGY . 29 GENERAL REMARKS . 29 THIN SECTION PETROLOGY . 31 DIAGENESIS . 37 ECONOMIC ASPECTS . 43 CONCLUSIONS . 45 REEERENCES . 50 APPENDIX — LOCATIONS OE SECTIONS . 55 iii ILLUSTRATIONS PAGE FIGURE 1 . 4 Index and outcrop map of the Herkimer Formation in east-central New York FIGURE 2 . 7 Relations between Upper Clinton units in east-central New York as considered by Fisher (1960). Vertical ruling signifies disconformity ; wavy line indicates physi¬ cal evidence of a disconformity. FIGURES . 10-11 Columnar sections of uppermost Clinton units, east-cen¬ tral New York. Numbers at top of sections refer to lo¬ calities : numbers within sections are field units. FIGURE 4 . 13 Restored section of Herkimer and Kirkland and adjacent units. FIGURE 5 . 14 Interbedded shale, dolostone, and sandstone of Joslin Hill Member at Dawes Quarry Creek. FIGURE 6 . 20 External mold of Rusophycus bilobata, natural size. FIGURE 7 . 20 Para-ripples in coarse dolostone, Joslin Hill Member, Dawes Quarry Creek (loc. 4). FIGURE 8 . 23 Medium-bedded Jordanville Member south of Edicks. FIGURE 9 . 24 Stromatolitic (?) structures in the upper Jordanville Member, south of Edicks. FIGURE 10 . 26 Correlation chart for uppermost Clinton units, east-cen¬ tral New York. Wavy lines represent disconformities ; vertical ruling represents hiatus. iv ILLUSTRATIONS Continued FIGURE 11 . 28 Electron probe photomicrographs of ferroan quartzose dolostone ; a sketch of area of analysis ; B, C, and D, X-ray beam scan images for MgKa, FeKa, and SiKa, re¬ spectively. FIGURE 12 . 30 X-ray diffractograms for (1) slightly quartzose dolo¬ stone, Joslin Hill, (2) “ typical ” dolomitic sandstone, Joslin Hill, (3) orthoquartzite, Jordanville, (4) shale, Joslin Hill. q=quartz, d=dolomite, cl=chlorite, “ i ”= “ illite ” Cu radiation, Ni filter; scanning rate 1 “/minute. FIGURE 13 . 31 Triangular diagram with end members of C|uartz, dolo¬ mite, and chlorite. Shaded area would include nonshale lithologies of the Herkimer. FIGURE 14 . 39 Paragenetic sequence. Dashed line connects diagenetic stages for which there is textural evidence. FIGURE 15 . 47 lA. Microfacies 1 : orthoquartzite ; crossed polarizers. IB. Microfacies 2 ; dolomitic, fossil fragmental sandstone ; note crinoid and bryozoan fragments ; ordinary light. IC. Microfacies 3 ; dolomitic siltstone ; crossed polarizers. ID. Microfacies 4 ; crinoid-bearing dolostone ; crossed po¬ larizers. IE. Microfacies 5 ; hematitic sandstone ; note hematite (after chlorite), outlining relict quartz grains, followed by secondary silica ; plane polarized. IF. Mi¬ crofacies 6 ; dolomitic limestone ; note hematitized crinoid fragments and euhedral quartz crystals replacing brachio- pod shell : plane polarized. FIGURE 16 . 48 I A. Polycrystalline quartz (left center) in conglomerate orthoquartzite ; crossed polarizers. IB. Eoliated quartz clast in microfacies 2, Joslin Hill Member ; crossed polar¬ izers. IC. Corrosion of quartz by chlorite followed by secondary silica, Joslin Hill Member ; plane polarized. ID. Hematitization within fenestrate structure of dolomi- tized pelmatozoan fragment, optically continuous rim cement surrounding clast; Joslin Hill Member, crossed polarizers. IE. Portion of clast with chlorite corroding quartz grains and larger dolomite grain at top of clast ; Joslin Hill Member ; plane polarized. IF. Euhedral dolomite grain surrounded by secondary silica with euhedral facies ; Joslin Hill Member ; crossed polarizers. V ILLUSTRATIONS Concluded FIGURE 17 . lA. Pyrite euhedra replacing dolomite of a crinoid frag¬ ment ; note quartz corroded by dolomite along margin ; Joslin Hill Member; crossed polarizers. IB. Murchi- sonia with whorls filled with finer dolomite than that out¬ side shell ; note abundance of quartz grains within shell ; Joslin Hill Member, ordinary light. IC. Dolomite rhombs within pseudomorphically replaced (dolomite) crinoid fragment (dark, at extinction position) ; Joslin Hill Member ; crossed polarizers. ID. Dolomite rhomb (right center) cutting across hematitized margin of fragment; Joslin Hill Member ; plane polarized. IE. Well pre¬ served, though dolomitized, bryozoan walls with sparry zooecial fillings; Joslin Hill Member; ordinary light. IF. Nearly obliterated bryozoan zoarium; Joslin Hill Member ; plane polarized. TABLES TABLE I . Upper Clinton Units in Area of Type Clinton, Including Ostracode Zones of Gillette (Modified From Gillette, 1947) TABLE H . List of Joslin Hill Fossils PAGE 49 PAGE 6 16-19 Uppermost Clinton (Middle Silurian) Stratigraphy and Petrology East-Central New York^ by Donald H. Zenger- ABSTRACT THE UPPERMOST CLINTON UNITS in the type area in east-central New York include the thin 0-1.2 meters (0-4 feet) Kirkland Dolo- stone, a hematitic, fossil fragmental dolostone, overlain by the Herkimer Formation 0-30.5 meters; (0-100 feet) ; these units pinch out eastward but disconformities are less significant than was previ¬ ously believed. The Herkimer is divided into western and eastern facies. The western facies (Joslin Hill Member) consists of fossil- iferous, quartzose dolostone and dolomitic sandstone, dolomitic silt- stone, shale, ferruginous sandstone, dolostone, and orthoquartzite. Shale-sandstone ratios increase to the west. Ripple marks, cross¬ bedding, bioclasts, and shale clasts are common. The eastern facies (Jordanville Member) is a well rounded and sorted, medium-grained orthoquartzite including conglomeratic and cross-laminated beds, shales being practically absent. An eastern source of terrigenous material is indicated by the distribution of lithologies and paleocurrent directional indicators. The discovery of the ostracode Drepanellina clarki above Paraechmina spinosa permits direct correlation with the Appalachians farther south and supports the contention that the upper Clinton in east-central New York is younger than the same horizon in western New York. The nonshaly beds of the Herkimer are divided into six microfacies ; (1) orthoquartzite, (2) dolomitic, fossil-fragmental sandstone to quart¬ zose biodolarenite, (3) dolomitic siltstone, (4) intraclast- and fossil- 1 Submitted for publication, July 2, 1970. 2 Department of Geology, Pomona College, Claremont, California. [1] bearing dolostone, (5) slightly impure sandstone, and (6) partially dolomitized, hematitic biospar ite. Ferroan dolomite, determined by staining and by electron probe microanalysis, is the major carbonate. Iron-rich chlorite and illite are the predominant clay minerals in the shales. A statistically significant negative correlation exists between density and insoluble content ; no correlation is indicated between dolomite and insoluble content. Fossil preservation suggests that dolomitization was mainly early diagenetic. Many of the postdeposi- tional changes were early diagenetic. The basic paragenetic sequence is calcite-hematite-dolomite-chlorite-secondary silica-pyrite. The Jordanville most probably represents a beach environment. The sandstones and dolostones of the Joslin Hill accumulated above wave base in a shallow-water, near-shore zone. Shales and siltstones formed under conditions of less energy. [2] Acknowledgments Considerable field work was supported by a grant from the Geologi¬ cal Society of America. Acknowledgment is also made to the donors of the Petroleum Research Fund, administered by the American Chemical Society, for partial support of this research. Timothy Tackett performed some of the analyses under NSF-URP GE-1038. I thank D. W. Fisher and A. O. Woodford for their critical reading of the manuscript. Helpful advice or suggestions were given by A. J. Boucot, H. S. Muskatt, and Robert Schoen. I am especially grateful to R. G. C. Bathurst for discussions regarding some of the problems associated with the carbonate fabrics. Drilling information was pro¬ vided by G. C. Borland, S. S. Galpin, and A. M. Van Tyne. J. W. Teeter kindly permitted the examination of a specimen of DrcpancUina from the Lockport Formation at Hamilton, Ontario. [3] FIGURE 1. — Index and outcrop map of the Herkimer Formation in east-central New York. [4] Introduction PURPOSE While studying the stratigraphy of the Lockport Formation (Middle Silurian) in New York, the writer discovered an apparent facies relationship between the uppermost Clinton Herkimer Formation and the overlying Lockport (Zenger, 1965, p. 131-132) instead of the long accepted disconformity between these major units (Swartz, and others, 1942; Fisher, 1960). Because New York provides the stan¬ dard Silurian section for the United States and because most of the evidence for the above-mentioned facies relationships occurs in the area of the type Clinton, a further consideration seemed important. It was decided to focus on the uppermost Clinton, particularly the Herkimer “ Sandstone.” This paper summarizes the stratigraphic and paleontologic relations within the uppermost Clinton in the type area and discusses the petrology of the Herkimer. It is hoped that this will provide the beginning of a long-overdue petrologic analysis of the entire Clinton. GENERAL SETTING Clinton section. — Figure 1 shows the outcrop map of the Herki¬ mer Formation in east-central New York. Upper Clinton units, their ranges in thickness, and corresponding ostracode zones are presented in table 1. All units pinch out eastward within the area shown (fig. 1) or a short distance to the east. Previous work. — An exhaustive review of earlier work is not attempted. Chadwick (1918, p. 351) named the “Herkimer Sand¬ stone,” the name being based on outcrops in southern Herkimer County, and the “ Kirkland Iron Ore ” (1918, p. 349) for exposures in the Town of Kirkland in which Clinton lies (fig. 1). Zenger (1966, [5] TABLE 1. — UPPER CLINTON UNITS IN AREA OE TYPE CLINTON, INCLUDING OSTRACODE ZONES OE GILLETTE (MODIFIED FROM GILLETTE, 1947) Rock Units Ostracode Zones Lockport Formation Ilion Member 0-21.3 meters (0-70') Herkimer Formation 0-30.5 meters (0-100') Upper Clinton P araechmina spinosa Kirkland Dolostone 0-1. S meters (0-5') Willowvale Shale 0-7.6 meters (0-25') Mastigobolbina typiis Westmoreland Hematite 0-0.9 meters (0-3') Note : The Dawes Sandstone, a restricted unit 0-3 meters (0-10 feet) thick found only in the western part of the outcrop belt, lies between the Williowvale and the Kirkland. p. 1159-1161) has presented a summary of work on the Herkimer. Gillette (1947) provided the most thorough stratigraphic treatment of the Clinton Group in New York, but the emphasis was on western and central New York. Fisher compiled Silurian correlations in New York (1960). He showed disconformities between the Willow- vale Shale and overlying Kirkland and between the Herkimer and overlying Ilion Shale of the Lockport. A portion of Fisher’s chart is presented in figure 2. As Gillette had concluded (1947, p. 23, 112), Fisher showed the Herkimer as the time equivalent of the Rochester Shale of western New York. METHODS OF STUDY Approximately 2 months were devoted to the field study of the uppermost Clinton stratigraphy across the outcrop belt shown in figure 1. The 20 localities studied are marked in figure 1 and are l)riefly described in the appendix. Directional features were meas¬ ured in the field and oriented specimens were collected. Quantitative insoluble residue determinations were based on solu¬ tion with 25 percent HCl. The precision of the method is good; based on analyses of 16 splits of a calcareous sandstone, the standard devia¬ tion is 0.46 percent. The relative standard deviation (standard devia¬ tion expressed as a percent of the mean) is 0.70 percent. [6] FIGURE 2. — Relations between Upper Clinton units in east-central New York as considered by Fisher (1960). Vertical ruling signifies disconformity ; wavy line indicates physical evidence of a discon- formity. [7] Densities of rock powders were determined using a Beckman Air Comparison Pycnometer (#930). This instrument and its geologic application was described by McIntyre, and others (1965), and by Zenger (1968). The precision of density determinations is high; 18 replicate measurements gave a standard deviation of 0.0027 units. X-ray diffraction (Philips X-ray Diffractometer) scans were run on all samples over a 20 range of 5° to 60°. Quantitative calcite and dolomite compositions were determined using a ratio of peak heights (fixed counts) of these minerals (at about 20 = 29.4° and 30.9°, respectively). These ratios were referred to a calibration curve of calcite-dolomite ratios vs. dolomite percent similar to those presented by Bromberger and Hayes (1966, p. 360). Whereas the instrumental precision is good (relative deviation = 1.1 percent), the analytical, or within specimen variability is considerably higher (relative devia¬ tion = 4.0 percent) owing mainly to variation through sample prepa¬ ration. Calcite-dolomite determinations are best considered semi- quantitative. The electron probe microanalyzer (Philips Electronic Instruments AMR/3) was used to determine the distribution of iron in the iron- rich carbonates of the Herkimer. A description of the operating con¬ ditions of the probe was presented by Baird and Zenger (1966). All samples were studied in thin section. Acidic solutions of Alizarin Red S (calcite positive) and potassium ferricyanide (ferrous iron positive) were used as stains (Sabins, 1962, p. 1184; Warne, 1962, p. 37). [8] Stratigraphy and Paleontology KIRKLAND DOLOSTONE Figure 3 comprises columnar sections and figure 4 is a restored section showing the stratigraphic relations across the outcrop belt. “ Kirkland Dolostone ” is used for the unit commonly called “ Kirk¬ land Hematite ” and “ Kirkland Iron Ore ” because the dominant lithology is dolostone. Fisher (1960) showed the Kirkland extending across the entire outcrop belt but it is the contention of this writer that as a continuous lithologic unit it does not extend eastward beyond Ilion Gorge (loc. 11) ; thin hematite beds in that interval east of Ilion Gorge are best considered as part of the uppermost Willowvale Shale (figs. 3 and 4). From the Oneida quadrangle to Ilion Gorge, the Kirkland ranges from 0.45-1.2 meters (1.5 to 4 feet) in thickness. The unit is grayish-red to moderate red, mottled, thin- to medium- bedded (commonly weathering more massive), coarse-grained to con¬ glomeratic, fossil-fragmental, calcareous, slightly sandy, very hematitic dolostone. Shaly zones and seams of medium-gray, less hematitic dolostone are commonest in the lower and upper parts. Pyrite is rela¬ tively common in some beds and shale clasts are present. The lower contact with the Willowvale is generally sharp, although at some localities there is a very small-scale interfingering of Kirkland and dolomitic laminations of the Willowvale. The contact is considered to be conformable. In the vicinity of Clinton, the Kirkland overlies the Dawes Sandstone, a very restricted, irregularly bedded unit which this writer considers the equivalent of the upper Willowvale farther east. The presence of a marine fauna including Atrypa, Coolinia [Fardenia], Lcptacna, Calymcnc, Trimcms, and Cornellites indicates that Gillette’s consideration of a continental origin (1947, p. 100) for the Dawes was in error. In all cases, the uppermost Kirkland grades into the Herkimer through at least a few inches of transition. The unit is fossiliferous, the dominant elements being brachiopods, bryozoans, and pehnatozoan (probably crinoids) fragments. Abrasion [9] FIGURE 3. — ■ Columnar sections of uppermost Clinton units, east-central New [10] Numbers at top of sections refer to localities ; numbers within sections are field units. [11] and subsequent hematitization have affected the preservation. Fol¬ lowing is a list of the forms collected by the writer : Brachiopods Coolinia [Fardcnla] subplana (Conrad) C. [F.] sp. Lcptacna “ rhoiiiboidalis ” (Wilckens) Protomegastrophia profunda (Hall) Ainphistrophla striata (Hall) Stegerhynchus ncglectns (Hall) Bryozoans Diploclcma sparsiim (Hall) Eridotrypa striata (Hall) E. solida? (Hall) Eenestrellina clegans (Hall) Locality Number 3,9,10 5,6*,7 5,6* 6* 6*,9 7,9 11 4 10 4 Trilobites Trimcriis (Trimcrus) dclphinoccphalus Green 3,5,10,11 Liocalymcne clintoni (Vanuxem) 7,11 Others Dazosonoccras amcricanum Foord 6* Mastigobolbina? sp. 6* Murchisonia (Hormotoina) sp. 3,11 * collected from loose block of Kirkland The stratigraphic position of the Kirkland is equivocal. This writer acknowledges the close lithologic relationship with the Herkimer but is impressed by the lack of a distinct break between the Willowvale and Kirkland. Consequently, the Kirkland is treated as a unit sep¬ arable from the Herkimer. HERKIMER FORMATION Introduction. — Zenger (1966) considered the “Herkimer Sand¬ stone ” of Chadwick (1918, p. 351) as a formation with western and eastern lithofacies, each of which was designated as a member. “ Jos- lin Hill Member ” was applied to the western facies (Oneida, Rome, and Utica quadrangles; fig. 1) and “ Jordanville Member” to the eastern facies (Winfield and Richfield Springs guadrangle). The abrupt lateral change between these facies occurs in the vicinity of Ilion Gorge (figs. 3 and 4). [12] Meters FIGURE 4. • — ■ Restored section of Herkimer and Kirkland and adja¬ cent units. [13] FIGURE 5. — Interbedclecl shale, dolostone, and sandstone of Joslin Hill Member at Dawes Quarry Creek. Joslin Hill Member. — The type section is along the south branch of Moyer Creek in the western part of the Utica quadrangle (loc. 10'; figs. 1 and 3; Zenger, 1966, p. 1163-1164) where 19.8 meters (65 feet) of Herkimer are exposed above the Kirkland. From its western¬ most outcrop (loc. 1) to Moyer Creek the thickness is consistent, ranging from 18.3-22.8 meters (60 to 75 feet). The lower contact with the Kirkland is gradational. Although Gillette (1947, p. 113) and Fisher (1960; fig. 2) postulated a disconformity between the Herkimer and overlying Ilion Member of the Lockport, this writer found a gradational to interfingering relationship (Zenger, 1965, p. 101-103). The exact contact is difficult to determine at many localities. At South Moyer Creek, a greenish sandstone bed of Herki- [14] mer afifinity occurs 2.1 meters (7 feet) above the base of the Ilion. Shale occurs throughout but becomes more important westward (figs. 3 and 5). Shale-sandstone+dolomite ratios range from 1:1 in the west to 1 :3 at Moyer Creek. The shales range from dark gray, smooth, and fissile or paper thin varieties to more thickly laminated and silty types. Though generally unfossiliferous, some shales con¬ tain a well preserved fauna. Nonshaly lithologies are dominated by gray to brownish-gray, very fine- to commonly medium-grained dolomitic sandstone followed in abundance by gray, fine-grained quartzose dolostone and dolomitic siltstone, light-gray to grayish-green sandstone, medium-gray to dark- gray dolostone, grayish-red hematitic sandstone to dolomitic sandstone, and at the westernmost outcrop some beds of dolomitic, very hematitic calcarenites (fig. 3). Bedding ranges from thin to medium and from irregular to flaggy. These beds are more resistant than the inter¬ calated shales and form small waterfalls. Fossils, particulary frag¬ ments, are abundant. These bioclasts range from sand-size to pebble- size. Pelmatozoan fragments, probably crinoids, are ubiquitous. Preservation of fossils is usually poor owing to abrasion or loss of bdetail as a consequence of fossilization as molds in a relatively coarse matrix. In addition to the pelmatozoans, the dominant elements are brachiopods, pelecypods, and bryozoans. A complete list of Joslin Flill fossils is presented in table II. DrepaneUina clarki and associated ostracodes have been found in the uppermost part of the member. Fucoids occur on the bottom of many beds and Hall (1852, p. 18) considered these to be the remains of marine plants. Rusophyciis bilobata (fig. 6) is one of the more common types. Seilacher (1964, p. 298-299, 310) considered this form to represent the cast of a trilo- bite burrow, a likely suspect in this case being Trimerits. [15] Table II List of Joslin Hill Fossils Locality Numbers [16] Protomegastrophia profunda (Hall)' rhynchonellid indet. Rhynchotreta americana Hall “ Schuchertella” sp. Stegerhynchus acinus (Hall) Brachiopods (continued) Stcgerhynchiis neglectus (Hall) [17} **Dawsonoceras americanum (Foord) [18] Ctenodonta sp. cf. Ctenodonta machacri- Table II — List of Joslin Hill Fossils — concluded Locality Numbers [19] Ichnophycus tridactylus Hall Rusophycus bilobata (Vanuxem) FIGURE 6. — External mold of Rusophycus bilobata, natural size. Some bottom marks appear to be inorganic, resembling flute casts and striation and groove casts. Ripple marks are relatively common as is tabular cross-stratiflcation. Para-ripples with wave lengths up to 3 feet are found in coarse-grained, fossil-fragmental dolostone layers (fig. 7), a situation similar in occurrence to those described by Bucher (1919, p. 258-268). Such ripple marks are symmetrical and there is no apparent difference in sorting between the troughs and the crests. Bucher attributed these strustures to tidal currents. FIGURE 7. — Para-ripples in coarse dolostone, Joslin Hill Member, Dawes Quarry Creek (loc. 4). [20] I An attempt was made to determine paleocnrrent directions using directional indicators including cross-stratification, ripple marks, and fossil lineations (e.g. tentaculitids and the conispiral Murchisonin) . Forty-six determinations were made but the results are not unequiv¬ ocal. Of 27 ripple mark measurements, only one lay within 20° of the east-west line and 78 percent lay within 50° of north-south, sug¬ gesting a shoreline oriented roughly along this line. The azimuths of cross-stratification were even less consistent. Hunter (1960, p. 181), however, on the basis of 21 readings for cross-beds in the “ marine ” Herkimer, recorded a deficiency of eastward directed orientations. He concluded that currents were to the west and northwest. Some generalizations can be made despite the complex association of lithologies. The beds immediately beneath the contact with the Ilion, and representing the DrepaneUina clarki Zone, are low-clastic dolostone containing the gastropod Mitrchisonia and tentaculitids, in addition to the ostracode assemblage. With the exception of ferru¬ ginous sandy dolostone just above the Kirkland at College Hill (loc. 3) west of Clinton, hematitic dolarenites and sandstones are only of importance in the middle or upper part of the Joslin Hill, above a consistently sandy zone containing Mytilarca, Modiolopsis, and other pelecypods. This zone is probably an extension of a westerly directed tongue of the Jordanville Member, and it becomes less impressive westward. At Sauquoit (loc. 7) and along South Moyer Creek, the lower third of the member consists of intercalated, fossiliferous, thin- bedded dolomitic siltstone to very fine-grained sandstone,, fine-grained to coarser, crinoidal sandy and silty dolostone, and dark shale. Brach- iopods, particularly Coolinia [Fardenia] and Amphistrophia, are the dominant faunal elements in this interval followed by gastropods (mainly Miirchisonia) , pelecypods (particularly Modiolopsis sp.), and tentaculitids. Restriction of fossils to certain intervals is generally not striking. The ostracodes associated with DrepaneUina clarki (e.g. DrepaneUina modesta, Paraechmina postica, Aeclimina sp., Disygo- pleura sp. cf. D. swartsi, and V elibeyrichia sp.) are limited to the uppermost few feet of Joslin Hill. Strophomenids such as Amphi¬ strophia, Coolinia [Fardenia], and Protomegastro phia are most com¬ mon in the lower part of the member. Jordanville Member. ■ — The Jordanville was named from outcrops along Flat Creek (loc. 14) and along the road south of Edicks (loc. 15), both exposures being near the town of Jordanville in the Rich¬ field Springs quadrangle (Zenger, 1966, p. 1164—1165). Within the outcrop belt, the unit ranges from about 22.8 meters (75 feet) in [21] thickness at Spinnerville Gulf (loc. 12) to a feather edge to the east (fig. 4). The reduction of the Herkimer eastward is caused by a prominent disconformity which has eliminated all Clinton units 4 miles east of Van Hornesville (Fisher, 1960). The disconformity is so ori¬ ented that although the Jordanville is missing at Ohisa Creek (loc. 18), it reappears at Van Hornesville (loc. 19) before pinching out just east of that village (figs. 3 and 4). At all exposures of the lower contact, the Jordanville rests on the Willowvale Shale. Hematitic beds in the upper part of the Willowvale probably represent the east¬ ern equivalents of the Kirkland but with the exception of a 0.3 meter (1-foot) bed at Ohisa Creek (loc. 18), the thin beds are not litho¬ logically similar to that unit being less hematitic and more argil¬ laceous. This contact is sharp and in places the basal Herkimer is conglomeratic. This situation is taken to represent a slight discon¬ formity. The upper contact with the Salina units is also disconform- able. The Lockport is absent east of Ilion Gorge. The top of the Jordanville, overlain by the Vernon Shale, at Spinnerville Gulf (loc. 12) and at Flat Creek (loc. 14), is weathered, friable, and limonitic. Farther east, south of Deck (loc. 17), the overlying Brayman Shale has a basal clayey zone containing rounded quartz grains presumably derived from the underlying Jordanville. At Ohisa Creek (loc. 18), the section is well exposed and hematitic beds of the upper Willow¬ vale are overlain by the Brayman Shale. On Wiltse Hill (loc. 20) just east of Van Hornesville, the Sauquoit Formation is directly over- lain by the Brayman, both the Willowvale and Herkimer having been removed by Silurian erosion. The Jordanville is a consistent mass of light gray to light brown¬ ish-gray, medium to coarse-grained, mostly medium-bedded (fig. 8) mature sandstone or orthoquartzite. Thinner bedding is found in the lower part at Edicks (loc. 15) and at Deck (loc. 17). Conglom¬ eratic layers are present throughout ; the clasts are well rounded and reach pebble proportions. The unit forms high waterfalls as a con¬ sequence of its greater resistance than underlying and overlying shales. Within the Jordanville, shale is minor, occurring either as thin intercalations between quartzite intervals or as shaly partings on sandstone beds. Shale clasts are common. Greenish-gray quartzites are found at the two eastern exposures (Iocs. 17 and 19) and at Van Hornesville (loc. 19) there are grayish-red hematite stringers. Tab¬ ular cross-stratification occurs at various horizons but only a few ex¬ posures were suitable for tbe determination of their orientation. Four of five measurements showed southwest to northwest directed cur¬ rents. All of Hunter’s (1960, p. 181) 15 cross-bedding determina- [22] tions from the “ nonmarine ” Herkimer are to the southwest or north¬ west. Fossils are very rare in the unit, no definite organic remains having- been reported before this study. At Van Hornesville, the writer has found the pelecypods Modiolopsis, Mytilarca, and a rhynchonellid in the upper unit (unit 3, loc. 19). These forms are typically sand dwellers. Arched bedding in the upper part of the member at Flat Creek (loc. 14) and south of Edicks (loc. 15; fig. 9) may represent stromatolites. Davis (1968) has recently reported, for the first time, stromatolites preserved in quartz sandstone. Transitional aone. — Along Steele Creek in the Ilion Gorge (loc. 11), the Herkimer Formation reaches its maximum thickness of more than 30.5 meters (100 feet). This area includes the type sec¬ tion of the Herkimer as established by Chadwick (1918, p. 351) al¬ though it was neither precisely located nor described. Although a complete section is not exposed, the sequence as compiled shows an interfingering of Jordanville and Joslin Hill and also of the upper sandstones and overlying Ilion. The lower part of the section con¬ sists of 7.9 meters (26 feet) of fine-grained, dolomitic, silty sand¬ stone and crinoidal dolostone containing Miirchisonia and tentaculi- tids, and dark, fissile shale with Modiolopsrs and Stegcrhynchits ; this unit is Joslin Hill in affinity. The middle, main mass of Herkimer is pale yellowish-brown, medium- to coarse-grained sandstone which in places is very pyritic. This portion is more than 15.2 meters (50 feet) thick and represents a westerly directed tongue of the Jordan- [23] FIGURE 9. — Stromatolitic (?) structures in the upper Jordanville Member, south of Edicks. ville. The upper 9.1 meters (30 feet) is dominated by elastics and includes faunal elements that are Joslin Hill is aspect. Gray, argil¬ laceous, locally dolomitic and hematitic sandstone with intercalated dark shales contain Tentaciilites, Miirchisonia, Mytilarca, Ctenodonta, Coelospira (?), and Chasuiatopora. This transitional zone is very limited laterally owing to the abrupt change of facies within the Herki¬ mer. [24] Stratigraphic Relations and Correlation It is the opinion of the writer that the Kirkland should be con¬ sidered a separate unit rather than a member of the Herkimer. Gil¬ lette (1947, p. 23; table I) included the Kirkland with the Herkimer as the Paraechmina spinosa Zone although he had no ostracode evi¬ dence for this. However, elements of the Mastigobolbina typiis Zone (table 1) such as Mastigobolbina (?) and Plcthobolina have been found in the Kirkland or its equivalents east of Ilion Gorge. In addition, the presence of Liocalymcne clintoni (Iocs. 7,11) sug¬ gests an association with the Willowvale. Farther south in the Appala¬ chians, this trilobite does not occur above the Mastigobolbina typus Zone. The time relationship of the Dawes, Kirkland, and Herkimer is not clear, however. The Dawes at Lairdsville contains Trinicrus delphinocephalus, which, so far as I know, has not been recorded below the Herkimer elsewhere in east-central New York. However, Trimerns has been observed in the Irondequoit limestone of western New York (Fisher, 1969, personal communication; Gillette, 1947, p. 152), this unit also representing the Mastigobolbina typus Zone. In addition, a form similar to Trimerns delphinocephalus was reported from the M. typus Zone near Hollidaysburg, Pennsylvania, by Ulrich and Bassler (1923, p. 362). If the ostracode occurrences and pres¬ ence of Liocalymcne properly place the eastern part of the Kirkland in the M. typus Zone, then either Trimcrus in the Dawes at Lairdsville (loc. 2) is also in that zone or the Dawes and overlying Kirkland are younger in a westerly direction and would be the time equivalent of the lower Herkimer to the east. Both Trimcrus and Liocalymcne have been found in the Kirkland. It is felt that the restriction of Liocalymcne clintoni to the M. typus Zone in this area is a more plausible hypothesis than the exclusion of Trimcrus from it. On the accompanying correlation chart (fig. 10), the Kirkland-Joslin Hill contact is shown as essentially parallel with time lines although it is possible that the contact rises to the west with respect to time. [25] ONEIDA (West) RICHFIELD SPRINGS (East) FIGURE 10. — Correlation chart for uppermost Clinton units, east- central New York. Wavy lines represent disconformities ; vertical ruling represents hiatus. Based on stratigraphic relations discussed, as well as the presence of Mytilarca and Modiolopsis in both units, the Joslin Hill and Jor- danville are lateral equivalents. The interfingering of upper Herkimer and Ilion Shale indicates a partial lateral equivalence of the Clinton and Lockport Groups. Because of this relationship and because the Ilion is considered equivalent to the upper Lockport of western New York (Zenger, 1965, p. 128-132), it is possible that the upper Her¬ kimer correlates with the lower Lockport of western New York instead of with the upper Rochester Shale as has been thought. The Drepancllina clarki Zone apparently overlies Paracchmina spinosa (Berdan and Zenger, 1%5, p. 99) in the Herkimer; in the Rochester Shale, however, Paraechtnina ranges practically to the top of the Rochester and Drepanellina is not known. Such relationships suggest that the uppermost Clinton in east-central New York is younger than the uppermost Clinton in western New York. Furthermore, although Drepanellina has not yet been found in the lower Lockport of west¬ ern New York, J. W. Teeter (personal communication) has found the ostracode in the Eramosa Member of the Lockport at Hamilton, Ontario. [26] With regard to more southerly Appalachian sections, the Kirkland, and equivalent beds in the uppermost Willowvale, with Liocalymcne, Mastigobolbina (?), and Plethobolbina would best correlate with the upper part of the Mastigobolbina typiis Zone whereas the overlying Herkimer, based on Joslin Hill fossils, correlates directly with the Drepanellina clarki Zone of Maryland and Pennsylvania. Although lithologically similar to the Keefer Sandstone, the Herkimer appears to be, at least in part, younger than that unit. [27] FIGURE 11. — Electron probe photomicrographs of ferroan quartzose dolostone ; A, sketcli of area of analysis ; B, C, and D, X-ray beam scan images for MgKa, FeKa, and SiKa, respectively. [28] Petrology GENERAL REMARKS Considerable effort has been devoted to the petrology of the Kirk¬ land by Dale (1953, p. 77-79), Hunter (1960, various pages), and others. The emphasis in this petrological study is on the Herkimer, particularly on the sandstones and dolostones. Excluding shales, the range of quantitative insoluble residue deter¬ minations for the Joslin Hill Member is from 8 to 99 percent. The average terrigenous content of the Jordanville, however, is about 99 percent, quartz being the major constituent. Density determina¬ tions, using the air comparison pycnometer, range from 2.68 gm/cc to 2.95 gm/cc, with a mean of 2.84 for the Joslin Hill, whereas the Jordanville averages 2.70 gm/cc. The main end members of the Joslin Hill are quartz, dolomite, and much less chlorite. Calcite is minor. Typical dolomite averages about 2.86 gm/cc in density. The high density values in the Joslin Hill can be accounted for by the sub¬ stitution of the ferrous iron ion primarily in dolomite and also in the chlorite (leptochlorite) . Electron probe microanalysis (fig. 11) shows the coincident distribution of magnesium and iron in a quartzose dolostone. The distribution of potassium ferricyanide staining also indicates the presence of ferroan dolomite in the member. As might be expected, a statistically significant negative correlation exists between insoluble content and density (r = — 0.8670, n = 72), owing to the low density of quartz and the high density of soluble ferroan dolomite. That the ferroan dolomite has not reached the end member ankerite is indicated by the lack of any appreciable dis¬ placement of the major dolomite diffraction peak at 2.89A. Crushed samples of all speciments were scanned with the diffracto¬ meter; figure 12 shows X-ray diffractograms for some characteristic lithologies. Although it is difficult to distinguish between chlorite and kaolinite at normal scanning rates (Biscaye, 1964, p. 1281, 1283), peak positions at about 7.08A (26 = 12.5) and at 3.53A (26 = 25.2) strongly suggest chlorite. Biscaye 1964, p. 1281) has mentioned the [29] DEGREES 20 FIGURE 12. — 'X-ray diffractrograms for (1) slightly quartzose dolo- stone, Joslin Hill, (2) “ typical ” dolomitic sandstone, Joslin Hill, (3) orthoquartzite, Jordanville, (4) shale, Joslin Hill, q = quartz, d = dolomite, cl = chlorite, “ i ” = “ illite ” Cu radiation, Ni filter; scanning rate l°/minute solubility of chlorite, but not kaolinite, in dilute HCl. Residues of the Herk'imer, scanned after such acidization reveals the removal of these chlorite peaks. Optically, the light greenish color and low birefrin¬ gence indicate chlorite. The chlorite is high in ferrous iron, as Schoen ( 1964, p. 859) has shown. In figure 13, the shaded area of the triangular diagram represents the area in which the nonshaly lithologies in the Herkimer would fall with respect to quartz, ferroan dolomite, and chlorite. X-ray analysis of the shales shows the constituents to be mainly quartz, chlorite, and “ illite ” that may include some muscovite. Calcite is very minor occurring in less than 25 percent of the speci¬ mens. There is a lack of a statistically significant correlation between insoluble content and calcite-dolomite ratios determined through X-ray diffraction. [30] Chlorite (mainly dolomite) FIGURE 13. — Triangular diagram with end members of quartz, dolo¬ mite, and chlorite. Shaded area would include nonshale lithologies of the Herkimer. THIN SECTION PETROLOGY Friedman’s (1965) terminology of crystallization textures and fabrics will be used with but slight modification. Friedman (1965, p. 648) proposed “ xenotopic ” for fabrics in which the majority of the constituent crystals are anhedral, “ hypidiotopic ” for fabrics in which the majority of crystals are subhedral, and “ idiotopic ” for those fabrics in which the majority are euhedral. As used by Fried¬ man, the terms anhedral, subhedral, and euhedral refer to the degree of development of crystal faces. As Friedman pointed out (p. 653), however, a plane intercrystalline boundary between two crystals is either a crystal face of one or a compromise boundary between the two. To determine whether a plane intercrystalline boundary is a crystal face is impractical when analyzing large numbers of thin sections and a descriptive scheme to be used with a less rigorous visual inspection of crystal boundaries is valuable. Friedman’s classification is modified so that the concept of “ plane intercrystalline boundaries ” is substi¬ tuted for “ crystal faces ” ; in this way, all straight boundaries are included, whether or not they are true crystal faces. [31] For purposes of environmental interpretation and discussion, it is practical to consider six microfacies. Although shales are vital when discussing environmental characteristics, they were not studied opti¬ cally. The six microfacies are illustrated in figure 15 and are as follows ; (1) orthoquartzite (fig. 15a) ; (2) dolomitic, fossil-fragmental sand¬ stone to c|uartzose biodolorudite or hiodolarenite (fig. 15b) ; (3) dolo¬ mitic siltstone to very fine-grained sandstone (fig. 15c) ; (4) intra¬ clast- and fossil-bearing dolostone (fig. 15d) ; (5) slightly impure sandstone (fig. 15e) ; and (6) partially dolomitized, hematitic bio- sparrudite to biosparite (fig. 15f). The first four microfacies are dominant and together with the shaly intervals comprise more than 95 percent of the Herkimer. (1) Orthoquartsite. — This microfacies (fig. 15a) is represented primarily by the Jordanville Member although some orthoquartzite is found toward the middle of the Joslin Hill. Rounding and sorting of the original detrital quartz grains are excellent. Relic grains are distinguished by “ dusty,” microlitic cores or rims surrounded by clearer silica overgrowths. In practically all samples there is a mode between 0.12 and 0.35 mm, or in the fine or medium-grained range. This matrix also exists in conglomeratic samples which consequently have a bimodal grain-size distribution. Overgrowths result in an interlocking texture. Where the quartz grains are elongate, the long axis usually parallels the bedding. “ Single ” quartz grains dominate; extinction in these ranges from nonundulatory through slightly undu- latory to very undulatory (Folk, 1961, p. 71). Polycrystalline grains are common and include those types mentioned by Blatt and Christie (1963, p. 561). These are: (1) grains composed of elongate ele¬ ments with close optical orientation, (2) those in which each unit has nonundulatory extinction but a different extinction position from that of its neighbors, and (3) those with crystal units having strong undulatory extinction and with sutured contacts (fig. 16a). Blatt and Christie (1963) have presented strong evidence indicating that caution is necessary when using the nature of the extinction and crystallinity to imply a particular source. However, in this and other microfacies, distinctly foliated cj^uartz grains (fig. 16b) indicate a contribution from a metamorphic source. Minor chlorite cement occurs primarily as small (silt-sized) greenish blades. In places these corrode secondary silica whereas in others, secondary silica overgrowths include chloritic material. Hem¬ atite, after chlorite, may form rims around detrital cores and in turn [32] are surrounded by secondary silica. Minor dolomite having the geo¬ metric aspect of cement is present in some samples. Pyrite, muscovite, and biotite, and carbonaceous bodies are uncommon. Shale clasts are present in some sections. Zircon is the predominant heavy mineral. Porosity is highest in this microfacies and exists either as intergran¬ ular voids or along partings. (2) Dolomitic, fossil-fragmental sandstone to quartsose biodolo ritdite or biodolarenite. — This diverse microfacies (fig. 15h) is un¬ questionably the dominant one in the Joslin Hill Member and has no representation in the Jordanville. Terrigenous content ranges from 24 to 80 percent and there is a gradation across the 50 percent bound¬ ary used to separate detrital rocks from carbonates. Transported constituents consist of terrigenous quartz grains plus various allochems (Folk, 1959), including bioclasts and less common intraclasts (few, if any, oolites and no pellets were observed in the Herkimer; oolites are present in the Kirkland). Quartz grains and allochems tend to lie with their long axes parallel with the bedding. Quartz is an important constituent of all samples ; allochems vary from constituting a few percent up to 40 percent of a sample. The crystallinity and nature of extinction of quartz grains are similar to those properties in the orthoquartzite microfacies. A foliated, meta- morphic grain is shown in fig. 16b. The size range of quartz grains is primarily from very fine sand through the sand-sized categories, into granule sized (see Folk, 1961, p. 24). Commonly, two modes exist in the grain-sized distribution permitting a separation of grains from matrix. In such samples there is a lower mode in either the very fine sand or coarse silt range and an accompanying mode in the medium or coarse sand range. Not uncommonly the two modes are represented by alternate laminations. The finer grained quartz is subangular in contrast with the better rounded, coarser grains. Corrosion of the quartz grains by dolomite or chlorite (fig. 16c) is severe in places. Pelmatozoan fragments are the dominant bioclasts followed by fragmentary or intact bryozoa zoaria, many of which are out of growth position. Another common type is the gastropod Murchisonia pre¬ served either as single whorls or as two or three connected whorls. Brachiopod and pelecypod remains are rare. The pelmatozoan frag¬ ments usually occur as pseudomorphic replacement (Lucia, 1962, p. 855) of calcite by dolomite. These bioclasts may be surrounded by optically continuous dolomite (fig. 16d). Bryozoan clasts commonly consist of sparry zooecial walls with micritic or chloritic material in the voids. Many specimens show laminations of bioclast-rich [33] layers alternating with finer, more quartzose, less fossiliferous ones. Bioclasts are most common in the medium and coarse sand sizes but occur also in the pebble range. One kind of intraclast consists of chloritic material in which are embedded corroded quartz grains (fig. 16e) and pelmatozoan frag¬ ments. Another is the “ clay gall ” or shale clast consisting of argil¬ laceous matter and silt-sized quartz grains. Other transported con¬ stituents are minor grains of plagioclase, mica, rare heavy minerals including zircon, tourmaline, and sphene, and carbonaceous flakes. The major orthochem is ferroan dolomite, which constitutes from 10 to 60 percent of the rocks. As groundmass material, dolomite consists of a xenotopic to hypidiotopic fabric ranging from decimicron- size to centimicron-size. Very little dolomite is micritic but is pre¬ dominantly spar. Decimicron-sized dolomite appears as anhedral grains with corroded edges ; many of these may have undergone some transportation. Coarser dolospar has the geometry of cement. Quartz grains may float in the dolomite but more commonly are dominant where the dolomite has the appearance of cement. Coarse to very coarse spar also occurs in pockets, most of which are probably void fillings, and as optically continuous spar around pelmatozoan frag¬ ments. Chlorite occurs either as small, silt-sized blades forming a corrosive selvage around quartz grains or as more homogeneous patches of cement. The mineral forms up to 10 percent of some specimens but is generally less abundant. In places dolomite grains are corroded by chlorite (fig. 16e). Secondary quartz overgrowths are not so common as in the previous microfacies, but become most significant in the dolomitic sandstones where quartz grains are in contact rather than floating in the dolo¬ mitic matrix. There are a few instances in which dolomite grains have been “ engulfed ” by secondary quartz overgrowths, some of which have euhedral faces (fig. 16f). Hematite is a minor but practically ubiquitous mineral. It occurs primarily as rims around zooecial openings of bryozoa, within the fenestrate structure of pelmatozoan fragments (fig. 16d), as replace¬ ment of fossils, and as rims around quartz grains in places followed by secondary quartz overgrowths. Pyrite exists as euhedral grains, averaging 0.38 mm in size, re¬ placing portions of pelmatozoans, bryozoans, or apparently even dolo¬ mitic matrix (fig. 17a). It occurs also as a replacement of irregularly shaped organic flakes. Calcite is very rare, existing as isolated grains of spar in only a few samples. [34] (3) Dolomitic siltstone to very fine-grained sandstone. — This inicrofacies is restricted to the Joslin Hill. Many of the characteristic features are shared with the microfacies just discussed. Quartz grains are commonly angular to subangular and are cor¬ roded by both chlorite and dolomite. Polycrystalline grains are not abundant, probably as a consequence of the smaller grain size ; most fall within the range 0.04—0.15 mm. The average quartz content is between 60 and 70 perecent. Secondary silica is minor. Bioclasts become more important as the quartz content decreases. Where present, they are aligned parallel with the bedding and occur in laminations separated by more quartzose, less fossiliferous layers. The dolomitic “ groundmass ” ranges from less than 10 percent to more than 40 percent. It occurs as subhedral grains which are some¬ what coarser than the associated quartz or as coarser, interlocking crystals around the clastic grains. Both dolomite and quartz are cor¬ roded by argillaceous material, particularly chlorite, which is found most typically as small blades around quartz grains but also exists as homogeneous cement. Minor constituents include plagioclase, muscovite, carbonaceous bodies, zircon, and sphene. (4) Intraclast- and fossil-bearing dolostone. ■ — -This microfacies (fig. 15d), restricted to the upper part of the Joslin Hill Member, is one of two represented primarily by carbonate textures. The terri¬ genous content ranges from 8 to 16 percent. Fossil fragments ob¬ served up to 4 mm in greatest dimension constitute between 35 and 50 percent of the samples ; these are pelmatozoan fragments, whorls of Mnrchisonia (fig. 17b), pieces of bryozoa, fillings of shells, of tenta- culitids, and ostracodes. Preservation of fossils ranges from good to poor. There is not such a noticeable tendency for fossils to be oriented with their long axes parallel with bedding. Argillaceous in¬ traclasts are present and pockets of coarser dolomite mosaic, with vague boundaries, represent either clasts or “ micro pseudobreccia (Bathurst, 1959, p. 371). Dolomite is micritic only in the voids of fossil fragments (fig. 17b). Much of the dolomite is “ dusty ” and occasional grains show a rhomb-shaped area within surrounded by clear spar, similar to such features described by Murray (1964). Quartz is generally silt-sized or in the very fine sand range and usually constitutes less than 10 per¬ cent of each specimen. Authigenic chlorite is very rare. Hematite occurs in fossil voids and pyrite as disseminated flakes or replacing carbonate grains. Calcite, as isolated grains of spar, is minor. [35] (5) Slightly impure sandstone. ■ — -This microfacies includes a vari¬ ety of minor sandstone types mostly in the Joslin Hill but with some representation from the Jordanville. It is very similar to the ortho¬ quartzite microfacies so far as characteristics of the quartz grains is concerned, but differs from it in the presence of more appreciable amounts of dolomitic, calcareous, and strontianitic cement. Chlorite is ubiquitous as a minor constituent. Fossil fragments are practically lackdng. A type having most of the above characteristics is hematitic, dolo¬ mitic sandstone. The hematite occurs as rims around the original quartz grains, in turn followed by secondary silica (fig. I5e). The hematite replaces tiny chlorite blades, and, in places, dolomite in the matrix. Pyrite grains transect hematitic areas. Zircon, biotite, and dark carbonaceous chips are present but uncommon in these impure sandstones. (6) Dolomitic, hematitic biosparrudite to biosparitc. — This micro¬ facies (fig. 15f) occurs as thin beds at the westernmost locality in the Joslin Hill. It is characterized by the abundance of calcite and by the lack of terrigenous quartz. The common bioclasts are pelmato- zoans, brachiopods, and Murchisonia. Intraclasts are rare, and micrite is minor. Void-filling spar is common both within fossils and in the space beneath brachiopod shells where the growth is commonly as prismatic crystals. Most pelmato- zoan fragments consist of a single calcite crystal surrounded by op¬ tically continuous calcite as rim cement. Much of the calcite is fer- roan. Quantitative X-ray diffraction analysis of two samples indicated a dolomite content of 16 and 32 percent. Dolomite occurs as rhomb¬ shaped crystals within the micritic filling of fossil voids, as subhedral to euhedral grains in the calcite matrix or cutting across shell walls (fig. 17d), and together with calcite as secondary deposits in voids. Hematite is very common. It occurs within void spaces in pelmato- zoans and gastropods, within orthid brachiopod shells as concentra¬ tions along the punctae, and along the edges and well into the interiors of fossil fragments. Hematite along the shell margins occurs in micron-sized carbonate that may represent micrite envelopes (Bath¬ urst, 1966). Limonite occurs as a replacement of carbonate material. All quartz is authigenic, consisting of euhedral crystals (fig. I5f) 0.08 to 0.50 mm in length within brachiopod shells. The base of the crystals, which contain calcite inclusions, is along the shell margin, the apices being directed inward. [36] DIAGENESIS IMost studies of Holocene (Recent) carbonate sediments involve low-terrigenous material. However, as Chave ( 1967, p. 200-201 ) pointed out, to “ rock-type ” geologists, a carbonate rock may contain up to 50 percent terrigenous matter. Such compositions, coupled with the nearly complete dolomitization of the Herkimer, make difficult any attempt to discuss diagenesis in the light of present-day processes. The determination of paragenesis is based primarily on textural rela¬ tions. With the exception of the predominantly calcitic microfacies 6, much of the earlier diagenesis is clouded or lost through neomorphism (Folk, 1965, p. 21) or replacement. Neomorphism includes all trans¬ formations between one mineral and itself or a polymorph, with par¬ ticular emphasis on the common carbonate minerals. As is indicated by the dolomitized fossils, that dolomite is of definite replacement origin ; it is probable that the matrix dolomite is also of a replacement variety although this cannot be proven. Microfacies 6, which is but partially dolomitized, reveals some significant paragenetic relations which may be extrapolated to other microfacies although this micro¬ facies has such characteristics that it cannot be considered simply as a less dolomitized equivalent of another microfacies. On the basis of relations seen in microfacies 6, pelmatozoan clasts were transported as calcite cry'stals and overgrowths later developed around them as rim cement. Some hematitization occurred very early in diagenesis, probably prior to rim cementation, as revealed by hema¬ tite within the fenestrate structure of pelmatozoan crystals (fig. 16d) which are surrounded by the clear rim cement. Dolomite rhombs transecting the hematitized margins of brachiopods indicate that this hematitization preceded dolomitization (fig. 17d). For other evidence of the diagenetic sequence, one must turn to the dolomitic microfacies. Because in microfacies 6 pelmatozoan plates are followed by calcite overgrowths, it seems most logical to assume that pseudomorphic replacement by dolomite (Lucia, 1962, p. 861) of the plate, as well as of the rim cement, occurred after de¬ position of the bioclasts and after some hematitization. The occur¬ rence, within the replaced fossil, of euhedral dolomite grains which extinguish at different positions than the main mass suggests later possible “degrading recrystallization” (Folk, 1965, p. 23) of the dolornite (fig. 17c). The common preservation of the original fene- [37] strate structure in dolomitized pelmatozoan plates (fig. 16d) suggests early dolomitization. Early dolomitization is further suggested by the excellent preservation of many bryozoa zooecia (fig. 17e) whieh have walls composed of sparry dolomite crystals that are restricted to the walls. However, in some instances, dolomite crystals cut across the walls indicating subsequent recrystallization and some bryozoa are more obliterated than others (fig. 17f), indicating a more inten¬ sive dolomitization. The origin of the dolomite of the “ groundmass ” presents more of a problem. It is very possible that the subhedral grains of dolomite are detrital. Amsbury (1962) deseribed terri¬ genous dolomite grains in Cretaceous rocks of Texas and concluded (p. 14) that the small detrital grains of dolomite could not be dis¬ tinguished from silt-sized authigenic dolomite. In the case of the Herkimer, it is improbable that the dolomite, or perhaps caleite grains prior to dolomitization, have a terrigenous origin ; the source would more likely be intrabasinal. Much of the dolomite, however, has the geometric aspect of cement, that is, large crystals binding quartz grains or bioclasts. Especially where quartz grains are abundant and commonly in contact, the dolomite is in the form of cement and it seems likely that the dolomite represents replaced calcite cement. On the other hand, in some samples the packing of quartz grains and bio¬ clasts is sufficiently “ loose ” that the original carbonate probably was fine-grained carbonate sediment deposited with the allochemical grains and acting as support for them. This would suggest later aggrading neomorphism (Eolk, 1965, p. 22) resulting in coarser spar. It seems possible, however, that where clastic grains or allochems are slightly floating, the effect could be caused by the growth of cement forcing grains apart (Krinsley and Donahue, 1968, p. 861). Chlorite, as blades or as homogeneous cement, is considered to be authigenic and relatively early diagenetic. Wherever chlorite is in contact with quartz it corrodes it severely (fig. 16c) ; in places it cor¬ rodes dolomite grains (fig. 16e) indicating that the chlorite postdates much of the early diagenetic dolomitization. As dolomite content in¬ creases, chlorite content tends to decrease. The spatial relationships suggest that chlorite was precipitated in voids not previously occupied by dolomite, or, in other words, after dolomitization. Chlorite rims in the quartzites and slightly impure sandstones are commonly fol¬ lowed by secondary quartz overgrowths (fig. 16c) suggesting that the secondary silica followed initial formation of the chlorite. Similar overgrowths commonly “trap” euhedral dolomite (fig. 16f) indi¬ cating that secondary silica postdates dolomitization. [38] Such chlorite rims, outlining the detrital quartz grains, are often partially to completely replaced by hematite (fig. 15e). This later stage hematitization also seems to be represented by the replacement of some dolomitic “ groundmass ” by hematite and more locally by the corrosion of dolomite grains by hematite. Later recrystallization of the chlorite has corroded secondary overgrowths. The last mineral to form in the paragenetic sequence is pyrite, euhedral crystals of which may be seen to transect dolomite, chlorite, and hematite (fig. 17a). Figure 14 is an estimation of the timing of the various dia- genetic processes relative to one another. A significant factor is the presence of intraclasts showing many diagenetic features similar to the rocks in which they are enclosed. For example, chloritic clasts containing corroded quartz and dolomite fragments (fig. 16e) probably represent Joslin Ftill intraclasts. This possibly indicates that many diagenetic changes necessarily occurred while the sediment was at or near the sediment -water interface. On the other hand, some of the diagenesis may well have gone on after incorporation of the clast in the matrix. It is of interest to note that later diagenetic changes, such as secondary silicification and pyritiza- tion were not observed in the clasts. Hunter (1960, p. 195-200) hypothesized that the iron for the Clinton iron ores was provided by thorough weathering of mainly crystalline rocks, of not unusually high iron content. He envisioned deposition as occurring when the acidic river waters reached the sea. Schoen (1965, p. 177) could not accept this theory mainly because the ironstones contain less terrigenous material than most other Clin- co Icite _ _ J? e rp^ti te„ _ „ _ _ dolomite _ _ -ChiqiLte _ _ __ „L>Li£a _ pyrite DIAGENESIS - — ^ FIGURE 14. — Paragenetic sequence. Dashed line connects diagenetic stages for which there is textural evidence. [39] ton rocks, an unlikely situation near the mouths of rivers. He sug¬ gested that submarine discharge of iron-rich ground water could have been the medium. Several writers (Murray, 1964; Murray and Lucia, 1967; Weyl, 1960) stressed the importance of “local source” dolomitization. Weyl (1960, p. 87) pointed out that since most ground waters contain little total carbonate, it could be supplied by local solution of carbonate with a mole-for-mole exchange of calcium by magnesium. In such in¬ stances, porosity would be produced in a dolomitized limestone owing to the fact that excess carbonate is necessary to account for the 12 percent volume decrease when calcite is converted to dolomite (Mur¬ ray, 1964, p. 391). According to Murray and Lucia (1967, p. 29), if the sediment being dolomitized is the only source of carbonate (i.e., local source), porosity can be formed and maintained. If, however, the carbonate is brought in with the dolomitizing water (distant- source), porosity would be destroyed. Porosity certainly is minimal in the dolomitic rock of the Joslin Hill but it is believed that compac¬ tion, recrystallization of dolomite, and precipitation of chlorite and secondary silica could have destroyed any porosity present. Some evidence of compaction exists in the form of some stylolitic contacts between grains, and quartz grains penetrating fossil fragments and intraclasts. A local source of carbonate for dolomitization is con¬ sidered feasible for the Herkimer. Of late there has accumulated a considerable mass of literature on the recent occurrences of supratidal dolostone. In fact, Friedman and Sanders (p. 267) considered most dolostones to be evaporitic deposits. Hypersaline brines would act as the dolomitizing agent, either through capillary concentration, as in caliche formation, or through “ seepage refluxion” (Adams and Rhodes, 1960). This later mechanism must be considered as a possibility for dolomitization of the Herkimer. The main dolomitization, in the Joslin Hill Member, is separated from the younger Salina hypersaline units by the thin I lion Shale. Conse¬ quently, Mg-rich brines may have percolated downward from the Salina evaporites providing the possibility of dolomitization in the underlying sediments. There is little direct evidence for the passage of these fluids through the Joslin Hill. However, gypsum and celes- tite were reported by Vanuxem (1842, p. 81), from what is now the Jordanville Member, and Hunter (1960, p. 20) also reported celestite. It is possible that this occurrence resulted from percolating waters moving downward from the Salina Group above, but the Jordanville has little dolomite and, as mentioned above, there is no indication that the more prominent dolomitization of the Joslin Hill was accomplished [40] by such a means. Furthermore, the I lion Shale, which separates the underlying Joslin Hill and overlying Salina, would have been a rela¬ tively impervious barrier to downward percolating brines. There have been many recent attempts, perhaps overextended at times (Zenger, 1970), to find ancient analogs of recent supratidal dolostones. Joslin Hill carbonates did not accumulate in the supra¬ tidal zone. Not only do the abundant fossils and ripple marks bear out this contention, but there is a complete lack of typical supratidal features such as mud cracks, burrows, and stromatolites. Lindholm (1969) recently proposed a possible terrigeneous origin, combined with subsequent diagenetic overgrowths, for ancient dolo¬ stones that show evidence of subtidal accumulation. In many thin sections of Joslin Hill samples there is at least a rough correlation between the grain sizes of quartz and dolomite. This, coupled with the common occurrence of cross laminations and ripple marks, could constitute evidence in favor of Lindholm’s hypothesis. However, it is more likely that these particular clastic textures and structures re¬ sulted from the movement above wave base of carbonate particles that formed within the basin. It is well known that echinoderms contain high percentages of MgCOg in the calcite lattice (Chave, 1952; Grave, 1964). The abundant high-magnesian pelmatozoan fragments, and grains abraded from them, could act as “starters” (Fairbridge, 1957, p. 125), or nuclei, for further precipitation of IMgCOg from alkaline interstitial waters and lead to the eventual development of the mineral dolomite. For example, Schlanger (1957) described the localization of dolomite crystals within segments of Eocene coralline algae of high-magnesian calcite. Finally, it seems quite likely that his Mg/Ca ratios are essential for dolomitization, although perhaps not hypersalinity. In the Coo- rang of Australia, maximum dolomite formation occurs when salinity is about that of normal sea water (Skinner, 1963). In addition, At¬ wood and Bubb (1968) have reported that the salinity and chemistry of interstitial waters within Holocene supratidal dolomitic sediments at Sugarloaf Key, Florida, are essentially those of sea water. Gaines (1970, personal communication) has suggested that possibly in the ancient, very shallow epeiric seas characterized by high surface/vol¬ ume ratios, the deposition of calcite, or aragonite, would be sufficient to raise the Mg/Ca ratio in the water to the point where dolomitiza¬ tion might become kinetically possible. [41] ! it-- ■! .T^ ^ : I'iv 7“^ s*,.-„i^au-‘"WltVf4^| li. i /»y#«L’ ’dJS' ' .•1,<..-C :r,-.u,ii^«d»|!ite..:,!^.,4^'-fl J pit ♦ ' •'<‘- {^H^: a *• ^u4-r» '^ ■ ..tf^l^pg, 1^' Economic Aspects The interfingering of relatively porous quartzitic sandstone with more impervious dolostone, dolomitic sandstone, and shale would theoretically provide subsurface conditions suitable for oil or gas ac¬ cumulation. Sandstone tongues are directed from the east with a minor dip away from the source. However, it would seem likely that local changes in dip in the subsurface could provide the appropriate attitude for natural gas entrapment along the contact between facies. To date, however, little gas has been reported from the Herkimer (A. M. Van Tyne, personal communication). Of 22 wells in which the Herkimer was recognized, only eight reported some gas. Of these, only one has produced commercially (located near West Eaton south of the Oneida quadrangle; (fig. 1)). For a time, this well pro¬ duced 1 million cubic feet/day (G. P. Borland, personal communi¬ cation) and since 1961 has produced 475 million cubic feet of natural gas. It should be mentioned, however, that many wells bottom in the younger productive Oriskany, there being no information of the pro¬ ductivity of the underlying Herkimer. Thin, even beds within the Joslin Hill Member could provide good flagstones (they have been used for this purpose locally around Clin¬ ton) and some thicker beds might be used as building stone of rather attractive quality. However, as evidenced by weathered rinds on many beds, the ferroan content of much of the dolomite eventually results in oxidation of the iron; the length of time that would be necessary for this reaction is not known. The Joslin Hill probably should not be considered as a good pros¬ pect for aggregate in concrete or in bituminous mixes. Reaction be¬ tween the alkalies in cement and silicate minerals may result in expansion, cracking, and decrease in strength of the concrete (Kry- nine and Judd, 1957, p. 330). Although the most susceptible silicate minerals are opal and chalcedony, quartz, if fractured, could be re¬ active. With but few exceptions, the Joslin Hill samples contain con¬ siderable quartz. The quartz does not appear to be fractured and might not be reactive with the alkalies released from the cement ; this [43] point should be investigated more thoroughly before any attempt is made to use this rock as aggregate. It is known that rocks containing high percentages of silicate minerals, including quartz, are “ hydro¬ philic ” (Krynine and Judd, 1957, p. 338), that is, have a high water- sorption ability but a low bitumen sorption and are generally unsuit¬ able for mixing with bitumens, such as asphalt. Furthermore, in many sections of the Joslin Hill, the intercalated shale would have to be considered in any economic venture. Its re¬ moval could result in prohibitive costs. As mentioned previously, the Jordanville Member consists pre¬ dominantly of orthoquartzite. Such rock would not ordinarily provide an aggregate material for reasons mentioned above and because sand¬ stones generally have a low wearing value (Krynine and Judd, 1957, p. 338). Consisting primarily of quartz and being quite uniform, the Jordanville offers some promise of utilization as a glass sand. The fine- to medium-grain size and the purity of the main, white Jordan¬ ville are important factors in this regard. Iron content is generally low with the exception of some pyritic beds in the zone transitional to the Joslin Hill and hematitic beds at Van Hornesville. The very minor shales and beds containing clay galls do not seem to pose a serious problem. [44] Conclusions The Jorclanville Member, represented primarily by the ortho¬ quartzite microfaces, has been considered as terrestrial (fluvial) by Hunter (1960), although he considered the possibility of a transitional environment. The discovery of marine elements (a rhynchonellid brachiopod and two mytiloid pelecypods) in the member at Van Hornesville do not positively refute the concept of a nonmarine en¬ vironment ; such forms might have existed within or slightly below the intertidal zone and could easily have been washed up on a beach. Their presence certainly attests to a proximity of the shoreline. Al¬ though shells, both intact and broken, may be common on the surfaces of many sandy beaches, it seems to be a rule that fossils are extremely rare in ancient beaches now represented by quartzose sandstone (Dun¬ bar and Rodgers, 1957, p. 68). Even in recent beach deposits with a litter of shell material on the surface, immediately beneath this sur¬ face there is a great reduction in such remains. The occurrence of hemispherical structures that may be stromatolites preserved in ortho¬ quartzite (fig. 9) suggests an intertidal to supratidal environment for the Jordanville. It is here suggested that the Jordanville represents a beach (intertidal) environment with possible infraneritic or supra¬ tidal portions. Evidence in the nonshale intervals of the Joslin Hill indicates a high-energy, near-shore, relatively shallow-water environment west of the Jordanville depositional site. Such evidence consists of elongate marine bioclasts lying with their long axes parallel with the bedding, the large size (up to pebble size) and good rounding of these frag¬ ments, the abundance of rounded cpiartz grains, and the general lack of carbonate mud. Most of these characteristics could be explained individually in ways other than by agitation in shallow water but collectively they seem to be unequivocal. Eield evidence such as para- ripples and cross-stratification further support this contention. Micro¬ facies 3 (dolomitic siltstone to very fine-grained sandstone) as well as the shales probably reflect periods of less energy, although it is conceivable that they accumulated through the action of unpreserved [45] holdfasts. The complex association of the coarse-grained microfacies is taken to represent slight changes in the ratio of bioclasts to terri¬ genous grains, those being richer in bioclasts representing slightly more offshore positions. The Joslin Hill, then, reflects minor oscilla¬ tions of the shoreline that lay just to the east for much of Herkimer time. Although previous workers envisioned a disconformity between the western Herkimer, or Joslin Hill, and overlying Lockport, it has been demonstrated that the relations are those of a facies change (figs. 4, 10). Some interfingering also has been observed between the Ilion and the overlying Vernon Shale of the Salina. Consequently there was a much closer time and space relationship between Clinton, Lock- port, and Salina units than was formerly believed. [46] Figure 15. (length of line = 0.5 mm) lA. Microfacies 1; orthoquartzite; crossed polarizers. IB. Micro¬ facies 2 ; dolomitic, fossil fragmental sandstone ; note crinoid and bryozoan fragments ; ordinary light. IC. Microfacies 3 ; dolomitic siltstone ; crossed polarizers. ID. Microfacies 4 ; crinoid-bearing dolo- stone ; crossed polarizers. IE. Microfacies 5 ; hematitic sandstone ; note hematite (after chlorite), outlining relict quartz grains, followed by secondary silica ; plane polarized. IF. Microfacies 6 ; dolomitic limestone ; note hematitized crinoid fragment and euhedral quartz crystals replacing brachiopod shells plane polarized. [47] Figure 16 (length of line =: 0.5 mm) I A. Polycrystalline quartz (left center) in conglomeratic ortho¬ quartzite; crossed polarizers. IB. Foliated quartz clast in micro¬ facies 2, Joslin Hill Member ; crossed polarizers. IC. Corrosion of quartz by chlorite followed by secondary silica, Joslin Hill Member; plane podarized. ID. Hematitization within fenestrate structure of dolomitized pelmatozoan fragment ; optically continuous rim cement surrounding clast; Joslin Hill Member, crossed polarizers. IE. Por¬ tion of clast with chlorite corroding quartz grains and larger dolomite grain at top of clast ; Joslin Hill Member ; plane polarized. IF. Euhedral dolomite grain surrounded by secondary silica with euhedral faces ; Joslin Hill Member ; crossed polarizers. [48] E F Figure 17 (length of line = 0.5 mm) lA. Pyrite euhedra replacing dolomite of a crinoid fragment ; note quartz corroded by dolomite along margin ; Joslin Hill Member ; crossed polarizers. IB. Miirchisonia with whorls filled with finer dolomite than that outside shell ; note abundance of quartz grains within shell ; Joslin Hill Member ; ordinary light. IC. Dolomite rhombs within pseudomorphically replaced (dolomite) crinoid frag¬ ment (dark, at extinction position); Joslin Hill Member; crossed polarizers. ID. Dolomite rhomb (right center) cutting across he- matitized margin of fragment ; Joslin Hill Member ; plane polarized. IE. Well preserved, though dolomitized, bryozoan walls with sparry zooecial fillings ; Joslin Hill Member ; ordinary light. IF. Nearly obliterated bryozoan zoarium ; Joslin Hill Member; plane polarized. [49] References Adams, J. E., and Rhodes, M. L., 1960, Dolomitization by seep¬ age refluxion ; Am. Assoc. Petroleum Geologists Bull., v. 44, p. 1912-1921. Amsbury, D. L., 1962, Detrital dolomite in central Texas: Jour. Sed. Petrology, v. 32, p. 5-14. Atwood, D. K., and Bubb, J. N., 1968, Distribution of dolomite in a tidal flat environment, Sugarloaf Key, Fla. (abstract) : Abstracts with Program, Geol. Soc. America Annual Meeting, Mexico City, p. 10. Baird, A. K., and Zenger, D. H., 1966, Use of the soft X-ray spectrograph and the electron probe microanalyzer for deter¬ mination of elements carbon through iron in minerals and rocks: Advances in X-ray Analysis, v. 9, Plenum Press, New York, p. 487-503. Bathurst, R. G. C., 1959, Diagenesis in Mississippian calcilutites and pseudobreccias: Jour. Sed. Petrology, v. 29, p. 365-376. - , 1966, Boring algae, micrite envelopes and lithiflcation of molluscan biosparites : Geol. Journal, v. 5, p. 15-32. Berdan, J. M., and Zenger, D. H., 1965, Presence of the ostracode Drepanellina clarki in the type Clinton (Middle Silurian) in New York State: U.S. Geol. Survey Prof. Paper 525-C, p. C96-C100. Biscaye, P. E., 1964, Distinction between kaolinite and chlorite in recent sediments by X-ray diffraction : Am. Mineralogist, V. 49, p. 1281-1289. Blatt, Harvey, and Christie, J. M., 1963, Undulatory extinction in quartz of igneous and metamorphic rocks and its significance in provenance studies of sedimentary rocks: Jour. Sed. Petrol¬ ogy, V. 33, p. 559-579. [50] Bromberger, S. H., and Hayes, J. B., 1966, Quantitative deter¬ mination of calcite-dolomite-apatite mixtures by X-ray diffrac¬ tion : Jour. Sed. Petrology, v. 36, p. 358-361. Bucher, W. H., 1919, On ripples and related sedimentary surface forms and their paleogeographic interpretation: Am. Jour. Science, 4th ser., v. 47, p. 149-210, 241-269. Chave, K. E., 1952, A solid solution between calcite and dolomite : Jour. Geology, v. 60, p. 190-192. - , 1964, Skeletal durability and preservation ivi Newell, N.D., and Imbrie, John, editors. Approaches to Paleoecology : New York, John Wiley and Sons, Inc., p. 377-386. - , 1967, Recent carbonate sediments — an unconventional view: Jour. Geol. Education, v. 15, p. 200-204. Chadwick, G. H., 1918, Stratigraphy of the New York Clinton: Geol. Soc. America Bull., v. 29, p. 327-368. Dale, N. C., 1953, Geology and mineral resources of the Oriskany quadrangle: New York State Museum Bull. 345, 197 p. Davis, R. A., Jr., 1968, Algal stromatolites composed of quartz sandstone: Jour. Sed. Petrology, v. 38, p. 953-955. Dunbar, C. O., and Rodgers, John, 1957, Principles of stratig¬ raphy: New York, John Wiley and Sons, Inc., 356 p. Fairbridge, R. W., 1957, The dolomite question in LeBlanc, R. J., and Breeding, J. G., editors. Regional Aspects of Carbonate Deposition : Soc. Economic Paleontologists and Mineralogists, Spec. Publ. 5, p. 125-178. Fisher, D. W., 1960, Correlation of the Silurian rocks in New York State: New York State Mus. and Sci. Service Geol. Sur¬ vey Map and Chart Ser., No. 1. Folk, R. L., 1959, Practical petrographic classification of lime¬ stones : Am. Assoc. Petroleum Geologists Bull., v. 43, p. 1-38. - , 1961, Petrology of sedimentary rocks: Austin, Hemp¬ hill’s Book Store, 154 p. - , 1965, Some aspects of recrystallization in ancient lime¬ stones in Pray, L. C., and Murray, R. C., editors, Dolomitiza- tion and Limestone Diagenesis : Soc. Economic Paleontologists and Mineralogists Spec. Publ. 13, p. 14—18. [51] Friedman, G. M., 1965, Terminology of crystallization textures and fabrics in sedimentary rocks: Jour. Sed. Fetology, v. 35, p. 643-655. - , and Sanders, J. E., 1967, Origin and occurrence of dolo stones in Chilingar, G. V., Bissell, H. J., and Fairbridge, R. W., editors. Carbonate Rocks: New York, Elsevier Publish¬ ing Company, p. 267-348. Gillette, Tracy, 1947, The Clinton of western and central New York: New York State Museum Bull. 341, 191 p. Hall, James, 1852, Paleontology of New York, vol. II, containing descriptions of the organic remains of the lower middle divi¬ sion of the New York System: Albany, C. van Benthuysen, 362 p. Hunter, R. E., 1960, Iron sedimentation in the Clinton Group of the central Appalachian Basin: Unpublished Ph.D. Disserta¬ tion, The Johns Hopkins University, 416 p. Krinsley, David, and Donahue, Jack, 1968, Diagenetic surface textures on quartz grains in limestone: Jour. Sed. Petrology, V. 38, p. 859-862. Krynine, D. P., and Judd, W. R., 1957, Principles of engineering geology and geotectonics: New York, McGraw-Hill Book Co., Inc., 730 p. Lindholm, R. C., 1969, Detrital dolomite in Onondaga Limestone (Middle Devonian) of New York: its implications to the “dolomite question”: Am. Assoc. Petroleum Geologists Bulk, V. 53, p. 1035-1042. Lucia, F. J., 1962, Diagenesis of a crinoidal sediment: Jour. Sed. Petrology, v. 32, p. 848-865. McIntyre, D. B., Welday, E. E., and Baird, A. K., 1965, Geologic application of the air pycnometer : a study of the precision of measurement: Geol. Soc. America Bull., v. 76, p. 1055-1060. Murray, R. C., 1964, Preservation of primary structures and fab¬ rics in dolomite in Newell, N.D., and Imbrie, John, editors. Approaches to Paleoecology : New York, John Wiley and Sons, Inc., p. 388-403. - , and Lucia, F. J., 1967, Cause and control of dolomite distribution by rock selectivity : Geol. Soc. America Bull., v. 78, p. 21-36. [52] Rickard, L. V., and Zenger, D. H., 1964, Paleontology and strati¬ graphy of the Richfield Springs and Cooperstown quadrangles. New York: New York State Mus. and Sci. Service Bull. 396, 101 p. Sabins, F. F., Jr., 1962, Grains of detrital, secondary, and primary dolomite from Cretaceous strata of the western interior: Geol. Soc. America Bull., v. 73, p. 1183-1196. Schlanger, S. O., 1957, Dolomite growth in coralline algae: Jour. Sed. Petrology, v. 27, p. 181-186. Schoen, Robert, 1964, Clay minerals of the Silurian Clinton iron¬ stones, New York State: Jour. Sed. Petrology, v. 32, p. 5-14. - , 1965, Origin of the Silurian Clinton ironstones (ab¬ stract) : Geol, Soc. America Special Paper 82, p. 177. Seilacher, Adolf, 1964, Biogenic sedimentary structures m New¬ ell, N.D. and Imbrie, John, editors. Approaches to Paleoecol- ogy : New York, John Wiley and Sons, Inc., p. 296-316. Skinner, H. C. W., 1963, Precipitation of calcian dolomites and mag'nesian calcites in the southeast of south Australia: Am. Jour. Sci., V. 261, p. 449-472. Swartz, C. K., and others, 1942, Correlation of the Silurian forma¬ tions of North America: Geol. Soc. America Bull., v. 53, p. 533- 538. Ulrich, E. O., and Bassler, R. S., 1923, Paleozoic ostracodes ; their morphology, classification and occurrence in Maryland Geo¬ logical Survey, Silurian, p. 271-391. Vanuxem, Lardner, 1842, Geology of New York, part III, Com¬ prising the survey of the Third Geological District : Albany, W. and A. White and J. Visscher, 306 p. Warne, S. S. J., 1962, A quick field or laboratory staining scheme for the differentiation of the major carbonate minerals: Jour. Sed. Petrology, v. 32, p. 29-38. Weyl, P. K., 1960, Porosity through dolomitization : conserva- tion-of-mass requirements : Jour. Sed. Petrology, v. 30, p. 85-90. Zenger, D. H., 1965, Stratigraphy of the Lockport Formation (Middle Silurian) in New York State: New York State Mu¬ seum and Science Service Bull. 404, 210 p. [53] - , 1966, Redefinition of the Herkimer Sandstone (Middle Silurian), New York: Geol. Soc. America Bulk, v. 77, p. 1159- 1166. - , 1968, Determination of calcite-dolomite composition using the air comparison pycnometer: Jour. Sed. Petrology, V. 38, p. 373-377. - , 1970, Supratidal dolostones : and overemphasis of their significance in geologic record? (abstract) : Am. Assoc. Petro¬ leum Geologists Bulk, v. 54, p. 561. [54] Appendix Locations of Sections of Uppermost Clinton and Adjacent Units. 1. Abandoned quarries and small exposures of Herkimer along Stony Creek beginning 91.4 meters (300 feet) north of east-west secondary road, and west of Route 365, extending to abandoned quarry just south of creek about 274.3 meters (900 feet) east of Route 365 ; from 3.2 to 4 kilometers (2 to 2.5 miles) SW of Verona; C 1/9 Oneida 15' quadrangle (NW 1/9 Vernon 7\' quadrangle) ; lowest exposure of Herkimer (Joslin Hill Member) at about 470 feet. Farther south, Herkimer-Ilion contact (at about 425 foot elevation) exposed in bank of Sconondoa Creek about 182.8-274.3 meters (600-900 feet) west of Route 365 about 1.6 kilometers (1 mile) east of Oneida, N 1/9 Oneida 15' quadrangle (NE 1/9 Oneida 7^' quadrangle). 2. Section, with considerable covered intervals, along stream flowing north to Lairdsville at which points it bends toward the east ; about 5.6 kilometers (3.5 miles) NW of Clinton; SW 1/9 Rome 15' quad¬ rangle (C 1/9 Clinton 7^' quadrangle) ; Willowvale, Dawes, Kirk¬ land, Herkimer (Joslin Hill Member), and Ilion ; base of Herkimer at about 660 feet. 3. East-flowing tributary of Oriskany Creek on north side of Col¬ lege Hill, 1.1 kilometers (0.7 mile) north of Route 412, just west of Clinton; S 1/9 Rome 15' quadrangle (E 1/9 Clinton 7^' quad¬ rangle) ; Kirkland, Herkimer (Joslin Hill Member), Ilion; Herki- mer-Ilion contact at about 740 feet. 4. Sherman Brook (known also as Dawes Quarry Creek) both east and west of bridge on north-south road (Dawes Ave.) which crosses creek 2.8 kilometers (1.8 miles) east of village square in Clinton; C 1/9 Rome 15' quadrangle (W 1/9 Utica West 7^' quadrangle) ; upper Willowvale, Dawes, Kirkland, lower and middle Herkimer (Joslin Hill Member) ; base of Dawes at about 685 feet. 5. Section along “ The Glen,” an east-flowing tributary of Sauquoit Creek at Chadwicks (formerly known as Willowvale), 3.2 kilometers [55] (2 miles) south of New Hartford; SE 1/9 Rome 15' quadrangle (SE 1/9 Utica West 7^' quadrangle) ; Willowvale, Kirkland, lower Herki¬ mer (Joslin Hill Member) ; base of Herkimer at about 815 feet. 6. Northeast-flowing tributary of Sauquoit Creek parallel to and about 0.8 kilometer (0.5 mile) south of Grange Road in village of Chadwicks; SE 1/9 Rome 15' quadrangle (SE 1/9 Utica West 7^' quadrangle) ; Herkimer, Ilion; base of Herkimer (Joslin Hill Mem¬ ber) at about 790 feet. 7. West-flowing tributary of Sauquoit Creek, midway between and parallel to Roberts Road and Laughlin Road, about 1.6 kilometers (1 mile) NNE of Sauquoit; SE 1/9 Rome 15' quadrangle and SW 1/9 Utica 15' quadrangle (SE 1/9 Utica West 7^' quadrangle and SW 1/9 Utica East 7^' quadrangle, respectively) ; Willowvale, Kirk¬ land, Herkimer (Joslin Hill Member), Ilion; base of Herkimer just below 840 feet. 8. Abandoned quarry in middle (?) Herkimer (Joslin Hill Mem¬ ber) just east of Tilden Road about 0.24 kilometer (0.15 mile) north¬ east of junction of Tilden and Higby Roads on hill overlooking Utica; SW 1/9 Utica 15' quadrangle (W 1/9 Utica East 7^' quadrangle) ; top of quarry at about 1130 feet; similar quarry just west of Tilden Road. 9. Section along Starch Factory Creek and west-flowing tributary from 1.2-1.6 kilometers (0.8 to 1.0 mile) NW of Stewart Corners; SW 1/9 Utica 15' quadrangle (C 1/9 Utica East 7^' quadrangle) ; Kirkland, Herkimer (Joslin Hill Member), and Ilion; base of Herki¬ mer at about 1105 feet. 10. South branch of Moyer Creek about 2.1 kilometers (1.3 miles) ENE of Parker Corners; S 1/9 Utica 15' quadrangle (SE 1/9 Utica East 7J' quadrangle) ; Willowvale, Kirkland, Herkimer (Joslin Hill Member), Ilion; base of Herkimer at about 1000 feet. 11. Composite section along Steele Creek (Ilion Gorge), a northeast¬ flowing tributary of the Mohawk River, between 4.8-5. 6 kilometers (3-3.5 miles) SW of Ilion; data taken from the following : SE-flowing tributary of Steele Creek immediately SW of Ilion Fish and Game club on Barringer Road, uppermost Herkimer at about 1090 feetp abandoned quarry immediately north of bend in Barringer Road 1.6 kilometers (1 mile) ESE of Kinne Corners, top of quarry at about 1080; banks of Steele Creek proper between 915 and 930 feet about one-half mile north of Beckus Gulf ; also ledges on hills between [56] these localities, on northwest side of Steele Creek ; upper Willowvale, Kirkland, Herkimer (Joslin Hill and Jordanville transitional), Ilion. 12. Section along and above 35-foot waterfall in Spinnerville Gulf, 1.1 kilometers (0.7 mile) NW of Spinnerville; NE 1/9 Winfield 15' quadrangle (N 1/9 Millers Mills 7i' quadrangle) ; Herkimer (Jordan¬ ville Member) and lower Vernon, top of Herkimer at about 1190 feet. 12. Discontinuous section along northeast-flowing tributary of Ful¬ mer Creek southwest of Mason Road crossing; NE 1/9 Winfield quadrangle (NE 1/9 Millers Mills 7^' quadrangle) ; upper Willow- vale, Kirkland (?), Herkimer (Jordanville Member) ; top of Herki¬ mer (not exposed) at about 1240 feet. 14. Flat Creek just north of paved road 4 kilometers (2.5 miles) NNW of Jordanville; NW 1/9 Richfield Springs 15' quadrangle (W 1/9 Jordanville 7^' quadrangle) ; Willowvale, Herkimer (Jordanville Member), Vernon; top of Herkimer (Jordanville Member) capping- falls at about 1250 feet. 15. Roadcut along sharp bend in paved road over Rock Hill 1.6 kilometers (1 mile) south of Edicks ; NW 1/9 Richfield Springs 15' quadrangle (Cl/9 Jordanville 7^' quadrangle) ; Herkimer (Jordan¬ ville Member) ; top of exposure (top of Herkimer not exposed) at about 1350 feet. 16. Section along west fork of north-flowing creek at waterfall about 1.9 kilometers (1.2 miles) SSE of Edicks; NW 1/9 Richfield Springs 15' quadrangle (C 1/9 Jordanville 7^ quadrangle) ; upper Willow¬ vale, lower Herkimer (Jordanville Member) ; Willowvale-Herkimer contact at about 1310 feet. 17 . Section along waterfall 0.48 kilometer (0.3 mile) south of settle¬ ment of Deck; N 1/9 Richfield Springs 15' quadrangle (E 1/9 Jor¬ danville 7^' quadrangle) ; Herkimer (Jordanville Member), Bray- man ; top of Herkimer at about 1400 feet. 18. Discontinuous section along Ohisa Creek from just east of road crossing 3.4 kilometers (2.1 miles) SW of Cramer Corners on down¬ stream to about 1300-foot elevation; N 1/9 Richfield Springs 15' quadrangle (W 1/9 Van Hornesville 7^' quadrangle) ; Willowvale, Brayman. 19. Sharp bend in Route 80 on western margin of village of Van Hornesville, El/9 Richfield Springs 15' quadrangle (S 1/9 Van Hornesville 7^' quadrangle) ; Willowvale, Herkimer (Jordanville Member), Brayman; base of Herkimer at about 1215 feet. [57] 20. Sporadic outcrops along west fork of northeast-flowing stream on Wiltse Hill just north of school No. 7 about 3.5 kilometers (2.2 miles) south of Starkville; E 1/9 Richfield Springs 15' quadrangle (SE 1/9 Van Hornesville 7^' quadrangle) ; Sauquoit, Brayman. [58] Printed by the University of the State of New York Press Heckman bindery. INC. SEPT 03 N. MANCHESTER. INDIANA 46962