TUL n// <50 HARVARD UNIVERSITY Library of the Museum of Comparative Zoology VOLUME 21 1979 TULANE UNIVERSITY NEW ORLEANS TULANE STUDIES IN ZOOLOGY AND BOTANY, a publication of the Biology Department of Tulane University, is devoted primarily to the biology of the waters and adjacent land areas of the Gulf of Mexico and the Caribbean Sea, but manuscripts on areas outside this geographic area will be considered. Each number contains an indivi- dual monographic study or several minor studies. Normally two numbers plus an index and a table of contents are issued annually. Preferred citation of the journal is Tulane Stud. Zool. and Bot. INFORMATION FOR AUTHORS: Manuscripts submitted for publications are eval- uated by the editors and by an editorial committee selected for each paper. Contrib- utors need not be members of the Tulane faculty. Manuscripts of 20 or more pages, double-spaced, are preferred. We recommend conformance with the principles stated in CBE Style Manual, 4th ed., 1978. Manuscripts should be typewritten and double spaced. Two additional copies should accompany the original to expedite editing and publication. Legends for figures should appear on a separate page and in sequence. Illustrations should be proportioned for one or two column width corresponding to our printed page size, and should allow for insertion of the legend if occupying a whole page. Guidelines for letter and other extraneous markings should be done with a non-photo blue pencil such as Eagle Prismacolor. Photographs should be on glossy paper. Many tables, if carefully prepared with a carbon ribbon and electric typewriter, can be photographically reproduced, thus helping to reduce publication costs. Lettering in any illustrative or tabular material should be of such a size that no letter will be less than 1 Vi mm high when reduced for publication. An abstract not exceeding three percent of the length of the article must accompany the manuscript. Separates of published articles are available to authors at a nominal cost. Page charges, calculated at $45/page, are solicited from authors who have funds for this purpose through their institutions or grants. Acceptance of papers is not dependent on ability to underwrite costs but excessive illustrations and tabular matter may be charged to the author. EXCHANGES, SUBSCRIPTIONS, ORDERS FOR INDIVIDUAL COPIES: Ex- changes are invited from institutions publishing comparable series. Subscriptions are billed in advance. A price list of back issues is available on request. Individuals should send their remittance, preferably money order, along with their orders. Remittances should be made payable to "Tulane University." Subscription rates: Volume 21, 22 $7.50 ea., $8.50 foreign. Copies of Tulane Studies in Zoology and Botany sent to regular recipients, if lost in the mails, will be replaced if the editorial offices are notified before the second subsequent issue is released. COMMUNICATIONS: Address all queries and orders to: Editor, TSZ&B, Depart- ment of Biology, Tulane University, New Orleans, Louisiana 701 18, U.S.A. Harold A. Dundee, Editor Arthur L. Welden, Associate Editor Samuel Clifford, Assistant to the Editors 3B63 035 CONTENTS OF VOLUME 21 NUMBER PAGE 1. A BIBLIOGRAPHY OF LOUISL\NA BOTANY 1951-1975 Eric Sundell 2 2. A SEASONAL AND ECOLOGICAL SURVEY OF FRESHWATER LIMPET SNAILS (PULMONATA: ANCYLIDAE) AND THEIR DIGENETIC TREMA- TODE PARASITES IN SOUTHEASTERN LOUISIANA Hugh M. Turner and Kenneth C. Corkum 67 ECOLOGICAL MORPHOLOGY OF FRESHWATER STREAM FISHES A. John Gatz, Jr. 91 NICHE RELATIONSHIPS OF THAMNOPHIS RADIX HAYDENI AND THAM- NOPHIS SIRTALIS PARIETALIS IN THE INTERLAKE DISTRICT OF MAN- ITOBA Donald R.Hart 125 I»1US. COMR. ZOOW. LIBRARY JUN2 5^*^ HARVARD UNIVERSITY -JUL Uoh.cx ISSN 0082-6782 FEB 5 1979 Volume 21, Number 1 $4.00 Feb. 1,1979 A BIBLIOGRAPHY OF LOUISIANA BOTANY 1951-1975 ERIC SUNDELL TULANE UNIVERSITY NEW ORLEANS TULANE STUDIES IN ZOOLOGY AND BOTANY, a publication of the Biology Department of Tulane University, is devoted primarily to the biology of the waters and adjacent land areas of the Gulf of Mexico and the Caribbean Sea, but manuscripts on organisms outside this geographic area will be considered. Each number is issued separately and contains an individual monographic study, or several minor studies. As volumes are completed, usually on an annual basis, title pages and tables of contents are distributed to recipients receiving the entire series. Manuscripts submitted for publication are evaluated by the editor or associate editor and by an editorial committee selected for each paper. Contributors need not be members of the Tulane University faculty. When citing this series authors are requested to use the following abbreviations: Tulane Stud. Zool. and Bot. INFORMATION FOR AUTHORS: The editors of Tulane Studies in Zoology and Botany recommend conformance with the principles stated in CBE Style Manual, 3rd ed., published in 1972 by the American Institute of Biological Sciences, Washington, D.C. Manuscripts should be submitted on good paper, as original typewritten copy, double-spaced, and carefully corrected. Two copies, carbon or other suitable reproduction, must accompany the original to expedite editing and assure more rapid publication. Legends for figures should be prepared on a separate page. Illustrations should be proportioned for one or two column width reproductions and should allow for insertion of legend if occupying a whole page. Photographs should be on glossy paper. Many tables, if carefully prepared with a carbon ribbon and electric typewriter, can be photographically reproduced, thus helping to reduce publication costs. Letter- ing in any illustrative or tabular material should be of such a size that it will be no less than 1 V^ mm high when reduced for publication. Manuscripts of 20 or more pages, double-spaced, are preferred. An abstract not exceeding three percent of the length of the original article must accompany each manuscript submitted. This will be transmitted to Biological Abstracts and any other abstracting journal specified by the writer. Authors of contributions will receive a Statement of Page Charges, calculated at $45/page. Partial or complete payment of these charges is solicited from authors who have funds available for this purpose through their institutions or grants. Acceptance of papers is not dependent on ability to underwrite costs. Illustrations and tabular matter in excess of 20 percent of the total number of pages may be charged to the author; this charge is subject to negotiation. EXCHANGES, SUBSCRIPTIONS, ORDERS FOR INDIVIDUAL COPIES: Exchanges are invited from institutions publishing comparable series but subscriptions are available if no exchange agreement can be effected. A price list of back issues is available on request. Remittance, preferably money order, should accompany orders from individuals. Make remittances payable to "Tulane University." Authors may obtain separates of their articles at cost. Subscription rates: Vols. 21, 22 $7.50 ea., domestic. S8.50 foreign. Copies of Tulane Studies in Zoology and Botany sent to regular recipients, if lost in the mails, will be replaced if the editorial offices are notified before the second subsequent issue is released. COMMUNICATIONS: Address all queries and orders to: Editor, Tulane Studies in Zoology and Botany, Department of Biology, Tulane University, New Orleans, Louisiana 70118, U.S.A. Harold A. Dundee, Editor Arthur L. Welden, Associate Editor Samuel Clifford, Assistant to the Editors I saw in Louisiana a live-oak growing, All alone stood it, and the moss hung down from the branches; Without any companion it grew there, uttering joyous leaves of dark green , And its look, rude, unbending, lusty, made me think of myself But I wonder' d how it could utter joyous leaves, standing alone there, without its friend, its lover near — for I knew I could not; And I broke off a twig with a certain number of leaves upon it, and twined around it a little moss, And brought it away — and I have placed it in sight in my room; It is not needed to remind me as of my own dear friends, (For I believe lately I think of little else than of them:) Yet it remains to me a curious token — /"/ makes me think of manly love; For all that, and though the live-oak glistens there in Louisiana, solitary, in a wide flat space. Uttering joyous leaves all its life, without a friend, a lover, near, I know very well I could not. Walt Whitman, 1860 TULANE STUDIES IN ZOOLOGY AND BOTANY Volume 21 . Number 1 $4.00 Feb. 1, 1979 A BIBLIOGRAPHY OF LOUISIANA BOTANY 1951 - 1975 ERIC SUNDELL Department of Biology, Tulane University, New Orleans, Louisiana 70118 This bibliography was undertaken to supplement Joseph Ewan's original "Biblio- graphy of Louisiana Botany" (1968) which compiled references through 1950. The format and style followed here represent an extension of that work. Reprinting of the original bibliography, now virtually unavailable, would of course greatly enhance whatever value the supplement might have, not only because it would spare the reader an unnatural disjunction in the story of Louisiana botany, but because it would con- veniently allow contemporary workers and students a comparison with the character and achievements of the past. Such a reprinting is not now under consideration. The present bibliography includes titles on the algae, fungi, bryophytes, and vascular plants of the state of Louisiana and its coastal waters. Most entries fall into the realms of floristics and plant taxonomy, on the one hand, and plant ecology or environmental analysis, on the other. References from diverse branches of botanical literature are assembled, with the focus on the natural flora and vegetation of the state. Thus, horticultural and agricultural items are excluded unless, like the rich literature on the Louisiana irises, they treat largely of plants growing without cultivation - originally, at least - within our boundaries. Forestry is included only when it contains significant redeeming botanical qualities! General monographs are also out-of-bounds, unless locally pertinent floristically or historically — type material from our area, for instance, is grounds for incorporation. Unpublished theses and dissertations, despite valuable field data that so many carry, are omitted. Occasional citations of works not listed in this bibliography (e.g. Viosca, 1935) refer to Ewan's 1967 bibliography. In addition, a number of pre-1951 items that are not found in the original Louisiana bibliography are entered here. Although effort was made to examine all references, a few were missed; these are marked with an asterisk, and, if annotated, the source of the information is cited. Finally, the bibliography is indexed by author, subject, and botanical name. I wish to express my gratitude to Joseph Ewan on several counts: for suggesting this project, in the first place, for the many references he passed on to me; for his critical reading of the manuscript; and, above all, for the various tools and tricks of the bibliographer's trade that he generously shared with me. EDITORIAL COMMITTEE FOR THIS PAPER: DR. JOSEPH EWAN, Professor Emeritus, Tulane University, New Orleans, Louisiana 70 11 8 ^ CHRONOLOGICAL BIBLIOGRAPHY 1863.* Reizenstein, L. von. Catalogue of the Lepidotera of New Orleans and its [ 1 1 vicinity. New Orleans: Issac T. Hinton. 8 p. The oldest catalogue of these pollinators for our area, but according to Lambremont, 1954, the basis for the author's determination of species in unknown. 1873.* Hilgard, E.W. Supplementary and final report of a geological reconnaissance [2] of the State of Louisiana made under the auspices of the New Orleans Academy of Science and the Bureau of Immigration, May and June, 1869. See originally, Hilgard, 1869. According to Cain, 1974, a copy of the supplement is to be found at University of Mississippi Library, Mississippi Collection. 1875. Maitre, R. Illustrated and descriptive catalogue of vegetable and flower [3] seeds, Holland or Dutch bulbs, tuberous and perennial plants, green-house, hot-house and bedding plants, ornamental and evergreen trees, flowering shrubbery, camelias and roses, garden implements, and other garden requi- sites. Philadelphia. [Private printing]. 129 p. Seed store was located at 631 Magazine Street and the nurseries and green-houses at 976 Magazine, in New Orleans. An impressive variety of edibles and ornamentals, (18 palms and 64 "orchids or parasites"") are offered for sale. Bulbs and roots were annually imported from Holland, but much of the rest was "exclusively our own product"" and consequently "thorough- ly to our climate." Illustrations rather few and eclectically borrowed. 1878. Chapman, A.W. An enumeration of some plants - chiefly from the semi- [4J tropical regions of Florida - which are either new, or which have not hitherto been recorded as belonging to the flora of the Southern States. Botan. Gaz. 3:2-6,9-12, 17-21. Half a dozen plants from New Orleans, chiefly Josiah Hale"s collections, are among the new records. 1888. Albrecht, J. The palms. A physiognomic sketch. Trans. Amer. Hort. Soc. (5] 5:98-108. Pantropical discussion by a widely travelled Louisiana resident. One or two local examples drawn upon. 1888. Renauld, F. and J. Cardot. New mosses of North America. I. Botan. Gaz. [6] 13:197-203. Fontinalis flaccida collected by Langlois m Last Louisiana, "in the branches and roots over- flowed in the Bayou Bonfouca"" (St. Tammany Parish). (Plate XIX). Part II, ibid. 14:91-1(X). Here tallied several other of Langlois" mosses, some of them new records for North America. See also item .123. 1895. Muller, J. Graphideae Eckfeldtianae in Louisiana et Florida, lectae, additis [7] obervationibus in Graphideas Calkinsianas ejusdem regionis. Bull. Herb. Boiss. 3:41-50. Lichens from St. Martinville (St. Martin Parish) and Pointe a la Hache (Plaquemines Parish), the collections of Father Langlois. According to Dix (1945), the lichens were sent by Eckfeldt, cuator at the Acad. Nat. Sci. Phila., to Muller for identification. They are credited to Eckfeldt but carry Langlois' collection numbers. About 45 species, 12 of them newly described. See Briquet, J . , Biographies des Botanistes a Geneve ( 1940), for a life of Muller, who was curator for the de Candolles. 1896. Renauld, F. and J. Cardot. Musci Americae septentrionalis exsiccati. Notes [8] sur quelques especes distribuees dans cette collection. Bull. Herb. Boiss. 4:1-19. New Louisiana records are based uf)on collections of Father Langlois. 1899. Bicknell, E. Studies in Sisyrinchium - I: Sixteen new species from the [9] southern states. Bull. Torrey Botan. Club. 26:217-231. 5. furcatum, the only specimens cited are from Hammond, Tangipahoa Parish, in southeastern Louisiana. 1900. Anon. [Note of A.B. Langlois' death.] Botan. Gaz. 30:359. [10] 1901. Lloyd, F.E. A botanical reconnaissance of the Mississippi Sound islands and [11] delta. J. New York Botan. Gard. 2:26-29. Brief observations on the vegetation of the islands lying between Biloxi, Mississippi, and the Mississippi Delta, and of the Delta proper. 1904. Lloyd, F.E. The delta of the Mississippi. J. Geogr. 3:204-213. [12] Observations were made during a botanical reconnaissance in summer of 1900 (see Lloyd, 1901). Botanical annotations. Six photographs. 1923. Huxley, J.S. lis n'ont que de I'ame: an essay on bird-mind. In Essays of a [13] biologist, pp. 105-130. New York. It was on the Louisiana coastal plain, at Mcllhenney's bird sanctuary on Avery Island (Iberia Parish), that the impassioned courtship and nesting rituals of the Louisiana heron and the "little White Egrets" made a lasting impression on this great English biologist. Anhingas and egrets at play are also among the recollections. Botany minimal. 1926. Abbott, E.V. A survey of the microbiological activities in some Louisiana [14] soils: a preliminary survey. La. State Univ., Agr. Exp. Sta., Bull. No. 194. 25 p. According to Hodges, 1962, one of the earliest surveys for soil fungi (among other things) in the southern states. 1928. Pessin, L.J. Mycorrhiza of southern pines. Ecology 9:28-33. 6 [15] Ectotrophic mycorrhiza of Pinus echinata. P. palustris, P. taeda, and P. caribaea studied at Bogalusa (Washington Parish). Seven figures. 1931. Small, J.K. and E.J. Alexander. Botanical interpretation of the iridaceous [16] plants of the Gulf States. Contrib. New York Botan. Card. 327. "Excerpt from the forthcoming Manual of the Flora of the Southeastern United States." which includes our Florida Parishes. 96 species of Iris, 75 of which are new binomials from southern Louisiana. Viosca later ( 1935) reduced these to four species. 1931. Viosca, P., Jr. Spontaneous combustion in the marshes of southern Louisi- [17] ana. Ecology 12: 439-442. In our enlightenment we may no longer believe in the death of Dickens" Mr. Krook by spontaneous combustion, but the author's interpretation of the more than 100 marsh fires in question is intriguing and convincing. 1932. Viosca, P. , Jr. Irises of Louisiana. Flower Grower 19:386-387. [ 18] The earliest of Viosca's iris publications and a popular forerunner of his definitive taxonomic and ecological study that appeared in 1935. See also Penn, 1962. 1933. de Lesdain, B. Lichens de la Louisiane recueillis par les freres G. Arsene et [19] Neon. Ann. Crypt. Exot. 6:49-58. Introduction in French by Frere Arsene, followed by a list of Arsenes Covington collections (St. Tammany Parish) and Neon's from Lafayette. Three new species are described. Also an annotated list of lichens collected by Langlois, determined by I'abbe' Hue' and published in his Lichenes Extra Europaei. 1935. Brown, C. A. Notes on the distribution of an epiphytic orchid, Epidendrum [20] conopseum Ait. in Louisiana. Proc. La. Acad. Sci. 2:105-106. Brief survey of the literature and parish list of collections known to the author. 1935. Brown, C. A. Some wildflowers of Louisiana. La. Conserv. Rev. 4(5):3-7,44. [2 1 ] First in a series of eight, 1935, '36, "37: II. Woodland flora of the pine flats, ibid. 4(6): 18-24; III. Flora of the alluvial soils, ibid. 4(7):22-29; IV. Trees, shrubs, and vines, ibid. 4(8):32-37; V. Ferns and fern allies, ibid. 5(l):12-23; VI. Weeds of Louisiana, ibid. 5(3):19-24; VII. Poisonous plants, ibid. 5(4):34-39; VIII. Four spring flowers, ibid. 6(1):10-11. In all, a popular running account for the outdoorsman, illustrated with sketches and photographs. First report of the introduction of Caperonia castaneaefolia into Louisiana from the tropics, VI, p. 23, most likely a misidentification. No voucher mentioned. 1935. Russell, R.J. and H.V. ^^owe. Cheniers of southwestern Louisiana. Geogr. [22] Rev. 25:449-461. Long narrow sandy ridges along the Gulf Coast, called "cheniers" by the Creoles because of the great bve oaks that thrive in their well-drained, fertile soil. Paper treats of the Quaternary origin of these formations, a contest between subsidence and delta growth. "Origin of Cameron Marsh" (Cameron Parish), pp. 454-456. Two maps, three photos. 1937. * Frye, T.C. and L. Clark. Hepaticae of North America. Univ. of Wash. Publ. [23] Biol.6(l-5):l-1018. 1937 to 1947. Early Louisiana contributions by such noted hepaticologists as Alexander Evans, Marshall A. Howe, and L. M. Underwood tabulated. 65 species designated from Louisiana. Koch, 1961. 1939. Svihla, R.D. Field notes on a collection of Louisiana Hepaticae. Bryologist [24] 42:118-120. About 20 new records for the state, taken in the marshlands near Morgan City (St. Mary Parish). 1940. Grout, A.J. Moss notes, 1939. Bryologist 43:74-76. [25] Range extensions into Louisiana for Leucobryum antillarum and Polytrichum piliferum, based on collections of Faith Pennebaker [Mackaness]. 1942. Brown, C. A. History of the wild strawberry in Louisiana. Home Gardening [26] 2:168,174,180. The author tracks the elusive wild strawberry of Louisiana, a plant mentioned by Bartram and Rafinesque, Riddell, Langlois and Small, his quest ending successfully on Silver Creek, Washington Parish, with the rediscovery oi Fragaria grayana (F. virginica). Brown's motive for this eleven year search was largely philanthropic rather than botanical: to improve our cultivat- ed variety of strawberry by crossing with the native. 1942. Brown, C.A. Native orchids. Home Gardening 3:6-7, 20. [27] Popular article with information on distribution, especially of Epidendrum conopseum. 1942. Foote, L.B. Bibliography of the official publications of the state of Louisiana, [28] 1803-1934. American Imprints Inventory No. 19. 576 p. Issued by Hill Memorial Library at Louisiana State University as part of the WPA Historical Records Survey Program. PubUcations of La. State Univ., Agr. Exp. Sta. and Agr. Ext. Div., may be found pp. 347-393, but are of extremely limited value for our purposes. 1935-1948 comprises vol. I of "State of Louisiana, Official Pubhcations," also compiled by L.B. Foote. Vol. II. 1948-1953, compiled by M.T. Lane. Both volumes appeared in 1954. 1954-1972, Vols. Ill, IV, and V, are, as stated on the title pages, compiled by W.O. Martin, Jr., Secretary of State, but the more likely candidate is M.T. Lane who remains Recorder of Documents throughout that period. A separate series began in 1949 as "State of Louisiana, Public Documents," appearing biannually and consecutively numbered (no volumes). These numbers are periodically superceded by the volumes of "Official Publications." More recent publica- tions of the Agricultural Experiment Station, the Wild Life and Fisheries Commission, and the Forestry Commission are often of botanical interest. 1942. Mackaness, F.P. Bryophytes of the live oak forest. [Abstract.] Proc. La. [29] Acad. Sci. 6:48-49. A rich abundance of corticolous bryophytes characterizes the live oak forest association. Mention of seven sp)ecies. 1942. Prescott, G.W. The fresh-water algae of southern United States II. The algae [30] of Louisiana, with descriptions of some new forms and notes on distribution. Trans. Amer. Microsc. Soc. 61:109-119. Annotated list of some 60 taxa. 1942. Russell, R.J. Flotant. Geogr. Rev. 32:74-98. [3 1 ] Flotant IS a fitting subject for geographers as well as botanists, for the conversion of open water into marsh creates a zone half plant, half land, a natural process in southern Louisiana that alligator-weed and water-hyacinth have only accelerated. Development and anatomy of flotant is emphasized, but the article will serve as a serious introduction to the entire subject, botanically, geographically, and culturally. 1943. Brown, C.A. Opportunities for paleobotanical research in Louisiana. Proc. [32] La. Acad. Sci. 7:36-38. "1942." Synopsis stresses study of lignite, the richest source of information on past vegetation in Louisiana. See also Brown, 1954. 1944. Eyles, D.E., J.L. Robertson, Jr., and G.W. Jex. A guide and key to the [33] aquatic plants of the southeastern United States. U.S. Public Health Bull. No. 286. 151 pp. Reprinted 1%3 by U.S. Dept. Interior, Fish and Wildlife Serv., as Bureau of Sport Fisheries and Wildlife Circular 158. "Southeastern" U.S. officially bounded on the west by the Mississip- pi-Louisiana line. Illustrated keys to genera and species and annotated species lists. A fairly rigorous guide , but without full species descriptions it is of limited value beyond genus. Supt. of Documents No. I 49.4:158. 1944. Kane, H.T. Deep delta country. New York. 283 p. [34] The account of A.B. Langlois' years as priest at Pointe a la Hache (Plaquemines Parish) where, only a few years before his arrival, his predecessor. Father Savelli, had been killed by a furious mob, is rich in botanical anecdotes: overcoming the problem of drying specimens in so humid a place, for instance - "Callers at the rectory during the worst days of July found a hot fire m the living room, and Father covered with perspiration as he worked at the hearth." And parish- ioners coming to mass from the back canals and lakes, proudly bearing plants for Father, pp. 228-234. The author acknowledges the Right Reverend Monsignor Joseph Langlois (p. 271) for hitherto unpublished material regarding his uncle, Father Langlois. Other sources are not indicated, though a lengthy "Selected Bibliography" lends added reliability to Kane's account of "the Botanist-Priest of Plaquemines." 1945. Cook, M.T. Species of Synchytrium in Louisiana. I. Description of species [35] found in the vicinity of Baton Rouge. Mycologia 37: 284-294. Part one in a scries of eight, ( 194.^, -47, -49, -51. -52, '53): Mycologia 37:571-576, 37:715-740, 39:351-357, 41:24-27, 43: 1()3-1(»7, 44:827-828, 45:101-114. Svnchvirium is a genus of gall forming parasites of higher plants. Part 111: Development and structure of the galls. The other seven entries contribute to a state roster of some 40 species, the vast majority of which arc newly described. Most speciments were collected around Baton Rouge. See also several tollow-up articles of John Karling for supplementary data and some taxonomic adjustments. 1945. Dix, W.L. Langlois' Louisiana Cladoniae. Bryologist 57: 156-159. [36] The collection forming the basis of this report resides at the National Museum in Washington, DC. Langlois wrote in the preface to his Catalogue, "It is hoped in a future new edition to give also a list of the lichens of the region." Apparently he never did so. Biographical sketch and history of the lichen specimens make this a particularly valuable contribution. See also Muller, 1894, and Tucker, 1970. 1946.* Brown, C.A. and W.H. Carter. Weed investigations. La. State Univ., Agr. [37] Exp. Sta., Bull. No. 402. 24 p. 1946. Taft, C.E. Some Oedogoniaceae and Zygnemataceae from Texas and Louisi- [38] ana. Trans. Amer. Microsc. Soc. 65:18-26. Nineteen taxa recorded for the state from collections between Shreveport and Baton Rouge. Oedogonium louisianense sp. nov. from Reeves, Allen Parish. 1947. Earle, T.T. The flowering cycle of water hyacinth. Proc. La. Acad. Sci. [39] 10:27-29. New Orleans study. Timetable of ojjening and closing of flowers, with related data. 1947. Lynch, J. J., J. E. King,T.K. Chamberlain, and A.L. Smith. Effects of aquatic [40] weed infestations on the fish and wildlife of the Gulf States. U.S. Dept. Interior, Fish and Wildlife Serv., Spec. Sci. Rep. No. 39. 71 p. Supt. of Documents No. I 49.15:39. Almost exclusively water-hyacinth and alligator-weed. Appendix A: "Investigations of the effects of the water-hyacinth on the fish and fish habits of Louisiana waters," esp. pp. 30-51. 1947. Robinson, B.B. Minor fiber industries. Econ. Botan. 1:47-56. [41] Spanish-moss gatherers in Louisiana, working the bayous for fallen moss, may average 500 pounds per day! Spartina "grass" as a substitute for broom com. Saw palmetto. 1948.* de las Barras y de Arago, Francisco. Cuando la Luisiana era espanola! Una [42] remesa de plantas. Monies 4:126-131. [USD A Botany Subject Index 14019]. 1948. Olive, L.S. Taxonomic notes on Louisiana fungi - I. Mycologia 40:6-20. [43] Several new genera and species of parasitic fungi found in the state, in addition to n^w distributional and host records Thallnsoora gen. nov. (T. aspera on Veronica peregrina) and Heicomina gen. nov. {H. caperoniae on Caperonia castaneae folia) . But see Item 21). All collections from Baton Rouge. 1948. OHve, L.S. Taxonomic notes on Louisiana fungi - II. Tremellales. Mycolo- [44] gia 40:586-604. 10 27 species of jelly fungi, 4 of which are previously undescribed. 1949. Bodman, M.C., Sr. The genus Heterochaete in the United States. Mycologia [45] 41:527-536. H. sublivida Pat. is known only from the type locality at St. Martinville, where it was collected by Langlois in 1897. The genus was previously unknown in the United States. H. andina, the other species in our area, is more widespread. Four figures. 1949. Viosca, P., Jr. Natural checks on the water-hyacinth. Proc. La. Acad. Sci. [46] 12:67-71. A more comprehensive article than the title indicates. Senescence of water bodies, the result of flotant or large permanent mats of water-hyacinth and many native and introduced aquatic weeds, is here considered the consequence of dramage and flood control measures. Solid observation and comprehension make of this plea for rejuvenation of bayous, swamps, and canals ot the lou country a solemn warnmg about Louisiana's future natural health. 1950. Jung, R.C. An annotated Hst of the Lepidoptera of the New Orleans area. [47] Proc. La. Acad. Sci. 13:42-48. A few cultivated plants identified as larval hosts. 1950. Moore, W.G. Limnological studies of Louisiana lakes. I. Lake Providence [48] [East Carroll Parish]. Ecology 31:86-99. "Aquatic Vegetation" discusses briefly the unanticipated paucity of the higher aquatic flora of this hard-water lake. '•Plankton": primarily blue-greens, greens and diatoms. 1950. Scott, A.M. New varieties of Staurastrum ophiura Lund. Trans. Amer. [49] Microsc. Soc. 69:248-253. Var. horridum only in Louisiana; var. longiradiatum. La., Miss, and Fla. 1951. Boudreaux, B.H. The insect family Aphididae in Louisiana. Proc. La. Acad. [50] Sci. 14:14-22. List of species includes the known plant hosts in the state. 1951. Brown, C.A. Cypress - the tree unique: the wood eternal. Garden J. New [51] York Botan. Card. 1:36-39. General information on the ecology and uses of baldcypress. Photographs of several extraordi- nary trees. 1951. Cooke, M.T. Distribution of species of Synchytrium in North America. [52] Mycologia 43:590-597. Fourteen spp. described by the author from material collected in our area. That Louisiana is the Synchytrium center of America reflects Cook's intensive work in his own state and a paucity of collections elsewhere, "...possibly all species found in America have a much wider geographic- al distribution than has been reported." No bibliography. 11 1951. Fassett, N.C. Callitriche in the New World. Rhodora 53:137-155, 161-182, [53] 185-194, 209-222. Louisiana collections of Langlois cited: C. terrestris, C. nuttallii, C. peploides var. peploides, C. heterophylla \aT.heterophylla. Drummond's collection of C. peploides from New Orleans probably isotype of C. drummondi, and Hale's Louisiana collection of C. nuttallii cited by Hegebnaier as an isotype. Key to southeastern species. Maps and crucial illustrations. 1951. Flint, L.H. Some winter red algae of Louisiana. Proc. La. Acad. Sci. 14:34- [54] 36. A few freshwater species discussed. 1951. Olive, L.S. Taxonomic notes on Louisiana fungi. III. Additions to the Trem- [55] ellales.Mycologia 43:677-690. Fifteen species of jelly fungi, most of which represent new records for the state. Several new species described. 1951. Penn, G.H. A brief chronology of the history of entomology in Louisiana. [56] Proc. La. Acad. Sci. 14:72-87. Excellent bibliography. Many entries with botanical information. 1951. Shinners, L.H. The North Texas species of Hymenocallis (Amaryllidaceae). [57] Field and Lab. 19:102-104. Pancratium liriosome Raf . (in Florula Ludoviciana) = Hymenocallis liriosome (Raf. ) Shinners. H. eulae, here newly described from only Texas, has subsequently been found in Louisiana. 1952. Brown, C.A. Notes on the occurrence of Ophioglossum crotalophoroides [58] Walt, in Louisiana. Amer. Fern J. 42:90-92. Evidently common on lawns in the Baton Rouge area and easily overlooked. No mention of vouchers. One figure, no scale. 1952. Gleason, H. A. The new Britton and Brown illustrated flora of the northeast- [59] em United States and adjacent Canada. New York and London. 3 vols. Like Fernald's edition of Gray's Manual, this monumental flora of the northeast remains of inestimable service to southern botanists. The fit with our Louisiana flora is perhaps best for the upland oak-hickory hardwoods of East and West Feliciana and East Baton Rouge parishes and for coastal plain elements. Second and third printings, 1958 and 1963, both slightly revised. 1952.* Horn, N.L. A comparative study of two species of Colletotrichum on vetch. [60] Phytopathology 42:670-674. C. sativum described as a new species on Vicia spp. and Pisum sativum in Louisiana. Biol. Abst. 1952.* Langdon, O.G., M.L. Bombard, and J.T. Cassady. Field book of forage 12 [61] plants on longleaf pine-bluestem ranges. U.S. Forest Serv., Southern Forest Exp. Sta.,Occ. Pap. 127. 117 p. 1952. Penfound, W.T. Southern swamps and marshes. Botan. Rev. 18:413-446. [62] "Systematic" study of the major types of swamps and marshes of the Southeast with in-depth discussion of each. Lengthy bibUography. 1952. Phaff, H.J., E.M. Mrak, and O.B. Williams. Yeasts isolated from shrimp. [63] Mycologia 45:698-719. Thirty-five cultures of yeasts were isolated from shrimp {Penaeus setiferus) collected in the Gulf of Mexico off the coast of Texas. 1952. Shanor, L. The characteristics and morphology of a new genus of Laboulbe- [64] niales on an earwig. Amer. J. Botan. 39:498-504. Filariomycesforftculae, collected in Baton Rouge. 25 figures, including photomicrographs. 1953. Cook, T. Louisiana irises, a bibliography. Lafayette: Southwestern Louisiana [65] Institute. 16 p. "This is not a complete bibliography, but it is the first printed work that shows what has been written on Louisiana Irises. " Two pages of newspaper items are particularly valuable. 1953. Duncan, W.H. Taxonomic collections of vascular plants in the southeastern [66] states — their abundance and relation to production of floras. Rhodora 55:353-358. Dot map, based on published records of five genera, indicates relatively poorly collected areas: most of Mississippi, Louisiana, and Tennessee. Table compares all southeastern states: only Miss, scores lower than La. in "Corrected Average Number of Collections per County." 1953. Exner, B. Comparative studies of four rhizoctonias occurring in Louisiana. [67] Mycologia: 45:698-719. Pathogens of considerable importance on beans, sugar cane, rice, figs. Pathology and taxon- omy. 1953. Flint , L.H. Two new species of Batrachospermum. Proc. La. Acad. Sci. [68] 16:10-15. Batrachospermum mikrogyne Flint et Skuja and B. basilare Flint et Skuja, two freshwater red algae described and, thus far, known only from Louisiana. Voucher specimens filed at 6 herbaria, including LSU & US. 1953. Greene, W.F. and H.L. Blomquist. Flowers of the South, native and exotic. [69] Chapel HUl, North Carolina. 208 p. An informative rather than merely descriptive text accompanies useful line drawings (floral 13 details wanting) to some 400 native and 100 exotic plants. 55 color plates, accenting the horticultural, include many excellent and ambitious artistic compositions remarkable for their botanical accuracy. La. well represented in the many "Fla. to Texas" species. 1953. Karling, J.S. Micromyces and Synchytrium. Mycologia 45:276-287. [70] "Many of the new species created by Cook [see item 35] will have to be reinvestigated thoroughly." 1953. Karling, J.S. Synchytrium urticae. Mycologia 45:613-615. [71] Invalidity of Cook's binomial and discussion of its possible identity with some Russian Synchyt- ria. 1953. Karling, J.S. Synchytrium chamaedryoidis . Mycologia 45:976-977. [72] Cook's = S. urticae, though an invalid name, is confirmed as a true species, and a nomenclatural change is proposed. 1953. Lowy, B. Auricularia in Louisiana. Proc. La. Acad. Sci. 16:28-30. [73] " ... to clarify and summarize briefly our knowledge of this genus in the state." Three species considered. 1953. Lowy, B. Myxomycetes of Louisiana. Mycologia 45:926-933. [74] Checklist includes 42 new reports for the state. Collections from Baton Rouge. Ten figures. 1954. Bick, G.H. A bibliography of the zoology of Louisiana. Proc. La. Acad. Sci. [75] 17:5-48. Reflects author's interest in the literature of "Natural history," thus indirectly of even greater use to botanists than a stricter treatment. Headings include "Climate, Geology, Physiography" and "Major Vegetation Studies." 1954. Brown, C. A. Palynological studies on Louisiana Ugnite. [Abstract.] Intemat. [76] Botan. Cong. Proc. 8(6):270. Fifty outcrops of Tertiary lignite are rich in pollen and fern spores. Correlations suggested between these grains and grains described from Eocene brown coals of Germany. 1954. Drechsler, C. Some Hyphomycetes that capture eelworms in southern states. [77] Mycologia 46:762-782. Two of the new species of nematode-capturers collected in Laplace (St. John the Baptist Parish), Louisiana. 1954. Drechsler, C. Two species of Conidiobolus with minutely ridged zygospores. [78] Amer. J. Botan. 41:567-575. C. rhysosporus and C osmodes new species found in decaying plant detritus from southern Louisiana. Close observation of spores and sp)ore germination. Four pages of figures. 14 1954. Flint. L.H. Sirodotia in Louisiana. Proc. La. Acad. Sci. 17:59-65. [791 Freshwater red algae closely allied to Batrchospermum and favoring cold, swift-flowing streams. Six species collected from Louisiana to date. 1954. Karling, J.S. The galls oi Synchytrium modioliensis . Bull. Torrey Botan. Club [80] 81:199-209. Originally described by Cook from Baton Rouge [35] and apparently collected earlier by Langlois (1888) in the state. A few details of interest concerning the Langlois collection filed at NY. 1954. Karling, J.S. Synchytrium brownii, a new species with sexual reproduction. [81] Sydowia 8:27-30. Established for a parasite of Oenothera laciniata in Louisiana and named for Clair A. Brown in appreciation of his help collecting species of Synchytrium. However, see Karling, 1958. 1954. Karling, J.S. Synchytrium modioliensis Cook and Synchytrium australe Speg- [82] azzini. Mycologia 46:529-533. Cook's parasite collected on Modiola caroliniana from Baton Rouge is synonymous with Spegazzini's Argentinian species. The latter has priority. 1954. Lambremont, E.N. The butterflies and skippers of Louisiana. Tulane Stud. [83] Zooi. 1:127-164. Botanical commentary, mostly larval-feeding habits, scattered throughout. References include several other Louisiana Lepidoptera titles. 1954. Lowy, B. A new Dacrymyces. Bull. Torrey Botan. Club 81:300-303. [84] D. nigrescens, a jelly fungus, from the Baton Rouge area. Eleven figures. 1954. Lowy, B. A new species ofP/a/yg/oea from Louisiana. Mycologia 46: 100-104. [85] P. longibasidia on frondose wood, Goodwood, La. Believed to represent "a primitive ancestral type from which the rusts could conceivably have been derived." 1954. Lytle, S.A. and B.N. Dirskell. Physical and chemical characteristics of the [86] peats, mucks and clays of the coastal marsh area of St. Mary Parish, Louisi- ana. La. State Univ., Agr. Exp. Sta., Bull. No. 484. 37 p. Foldout map of soil types. Information on water-holding capacity and organic matter patently of great use to wetlands ecologists. 1954. Penn, G.H. Introduced pitcher plant mosquitoes in Louisiana (Diptera, [87] Culicidae). Proc. La. Acad. Sci. 17:89-90. Scarcity of Sarracenia purpurea in Louisiana is apparently the critical factor preventing perma- nent establishment of Wyeomyia hay net Dodge. 15 1954.* Stroube, W.H. Host range of the Rhizoctonia aerial blight fungus in Louisi- [88] ana. Plant Dist. Reporter 38:789-790. List of the natural plant hosts of Pellicularia filamentosa f. sasakii. Biol. Abst. 1954. * Stroube, W.H. Puccinia oahuensis on Digitaria ischaemum and D. sanguinal- [89] is in Louisiana. Plant Dist. Reporter 38:120. Apparently the first record of this rust fungus in Louisiana, and the first record of its occurrence on D. ischaemum. Biol. Abst. 1954. Taylor, W.R. Sketch of the character of the marine algal vegetation of the [90] shores of the Gulf of Mexico. U.S. Dept. Interior, Fish and Wildlife Serv., Fishery Bull. 89:117-192. Louisiana algal flora, pp. 187-188. Extensive bibliography. Supt. of Documents No. 149.27:89. Bull. 89 is devoted entirely to the Gulf of Mexico and includes other pertinent titles: Phyto- plankton, pp. 163-169; Bacteria, Fungi, and Unicellular Algae, pp. 217-232. See also Thome, 1954. 1954. Thome, R.F. Floweringplantsof the waters and shores of the Gulf of Mexico. [91] U.S. Dept. Interior, Fish and Wildlife Serv., Fishery Bull. 89:193-202. Literature review yields four principal communities of flowering plants: submarine meadow, mangrove swamp, salt marsh, sand-strand vegetation. Floristic and ecological synopsis with extensive bibliography. Supt. of DocimientsNo. 149.27:89. 1954.* Wolff, S.E. A guide to plant names in Texas, Oklahoma, Louisiana, and [92] Arkansas. Rev. by C.A. Rechenthin. U.S. Dept. Agr., Soil Conserv. Serv. 91 p. 1955. Anon. Forests of Louisiana, 1953-1954. U.S. Forest Serv., Southern Forest [93] Exp. Sta. , Forest Survey Release 75. 64 p. Brief descriptions of forest types. Colored maps showing 5 forest types and "non-typed, less than 10% forest." Egler, J 961 1955. Anon. Publications of the Southern Forest Experiment Station, July 1921 [94] through December 1954. U.S. Forest Serv., Southern Forest Exp. Sta., Occ. Pap. 108 (Revised). 128 p. Numerous valuable titles for the plant ecologist, especially in the areas of grazing and fire ecology, can be had here by selective culling. An understanding of fire ecology is no doubt the key to an understanding of the vegetation of the coastal plain, a vast sub-climax zone of pine, and it is our state and federal agencies that have the funds and manpower to investigate the role of fire, botanically as well as commercially. Reprints are often available on request to S.F.E.S. in New Orleans. See also Punch, 1%2. 16 1955.* Benjamin, R.K. New genera of Laboulbeniales. Aliso 3:183-197. [95] //omaromycei newly described from Illinois and Louisiana: H. epjm parasitic on a beetle. Biol. Abst. 1955. Drechsler, C. Additional species of Zoopagaceae subsisting on Rhizopods [96] and eelworms. Mycologia 47:364-388. Four more types (see item 77) from Laplace (St. John the Baptist Parish). 1955. Eggler, W.A. Radial growth of nine species of trees in southern Louisiana. [97] Ecology 36: 130-136. Based on 70 trees, March to March 1951-1952. Two areas, riverfront in Jefferson Parish and cypress-tupelo swamp in St. Charles Parish. Egler, 1961. 1955. Hansford, C.G. Tropical fungi. V. New species and revisions. Sydowia 9:1- [98] 88. Among the many new descriptions of spp. and vars., find (p. 3) Irenopsis quercifolia on Quercus, Louisiana and Florida. Herbarium study. 1955. Karhng, J.S. Synchytrium ranunculi Cook. Mycologia 47:130-139. [99] Originally described as a parasite of Ranunculus pusillus at Baton Rouge. Life cycle with 31 figures. 1955.* Lowy, B. Illustrations and keys to the tremellaceous fungi of Louisiana. [100] Lloydial8(4):149-181. Nine species. Speairs and Lowy, 1957. 1955. Lowy, B. Some Louisiana Gasteromycetes. Proc. La. Acad. Sci. 18:45-53. [101] Keys to orders, families, genera, and species, followed by a checklist of 32 species, 28 of which represent first reports from the state. Eleven photographs help to compensate for the lack of full species, habitat and substrate descriptions. 1955. Wherry, E.T. The identity of Dupratzia Rafinesque. Castanea 20:71. [102] The genus Dupratzia, first proposed by Rafinesque in his Florula Ludoviciana of 1817, is formally placed on record as a later name for Eustoma Salisbury, (Gentianaceae), the "West Indian bluebell." (Index Kewensis refers Dupratzia, with a question, to Phlox). 1955. Wilbur, R.L. A revision of the North American genus Sabatia (Gentianace- [103] ae). Rhodora57:l-33, 43-71, 78-104. Tulane herbarium consulted. Type of 5. macrophylla Hook. var. microphylla collected by Drummond in St. Tammany Parish. 5. brachiata: Louisiana collections by Drummond, Ar- sene. Cocks. 5. grandiflora: single Louisiana collection, by Josiah Hale, filed at Gray. S. brevifolia: also a single state collection, at MO, without locality data: "... should not be included in flora of the state without a better substantiated record." Louisiana vouchers for several other species are numerous. 17 1956. Blake, S.F. The identity of Calyptocarpus blepharolepis. Rhodora 58:275- [104] 278. Some history of C. vialis's eastern march from Texas across Louisiana and some New Orleans collections. 1956. Breen, R.S. and R.A. Pursell. More mosses from Stone Mountain, Georgia [105] and vicinity. Bryologist 59:184-186. "Specimens of Splachnobryum collected by Faith Pennebaker Mackaness in New Orleans are 5. wrightii, a species closely related to 5. bemoullii." This collection of an uncommon, tropical American moss, was published originally as 5. bemoullii. See also Koch, 1957. 1956. Brown, C.A. Commercial trees of Louisiana. Baton Rouge: Louisiana For- [106] estry Commission. 76 p. Paperback. Seventy species described and illustrated with photographs: bark, leaves, fruits or flowers. Interesting tidbits. Map of tree regions divides state into six vegetation regions. 1956. Gunter, G. Land, water, wildlife and flood control in the Mississippi Valley. [107] Proc. La. Acad. Sci. 19:5-11. Careful summary and interpretation of a subject that will always be at the heart of the natural history of Louisiana. The fate of once natural overflow areas currently deprived of both floodwater and its alluvium is one of several topics considered. 1956. Humm, H.J. Sea grasses of the northern Gulf Coast. Bull. Mar. Sci. Gulf and [108] Carib. 6:305-308. Observations and collections in Mississippi Sound off Ocean Springs, Miss. Annotated list of six species with key. 1956. Karling, J.S. Undescribed species oiSynchytrium. Mycologia 48:83-98. [109] 5. nitidum collected by Clair Brown on Specularia (Triodanis) biflora at Baton Rouge. 1956. Koch, L.F. Louisiana muscology 1. Review and summary of literature. Bry- [110] ologist 59:192-203. Details of collectors and collections within the state (Drummond, Featherman, Mohr, Lang- lois, et al.) and subsequent history of the collections and their incorporation into catalogues of broader geographic range. Louisiana citations are culled from a diffuse literature, making the literature cited surely the most complete and useful list to date for Louisiana bryologists. Two appendices: doubtful Louisiana taxa and nomina nuda and other rejected reports. 1956. Lowy, B. A note on Sirobasidium. Mycologia 48:324-327. [Ill] S. sanguineum Lager . & Pat. found on a dead branch of Frormu^mgra near Baton Rouge. Few records for this genus in North America, and this and Olive's 1951 report are our only local collections of the species. 18 1956. Mackin, J.G. Dermocystidium marinum and salinity. Proc. Natl. Shellfish [112] Assoc. 46:116-128. A parasite of oysters studied in Redfish Bay (Plaquemines Parish), Louisiana. The genus is hanging in Ainsworth and Bisby's Dictionary of the Fungi, "? Chytridiales or Protozoa." 1956. Moore, J. A. Notes on fern distribution in Louisiana. Amer. Fern. J. 46:82- [113] 84. Localities in northern Louisiana additional to Brown and Correll, 1942. Six ferns and Selaginel- la apoda. 1956. Moore, J. A. Silene virginica in the Gulf States. Rhodora 58:27-29. [114] Rejwrted by Riddell, 1852, and later by Dormon, 1934, as occurring in Louisiana, but not upheldin Hitchcock and Maguire's revision of North American species of Silene (1947). Vouchers from four parishes. 1956. Penfound, W.T. Primary production of vascular aquatic plants. Limnol. [115] Oceanogr. 1:92-101. Comparison of primary production in vascular aquatics with that of several terrestrial commun- ities emphasized the high productivity of the former. Data on productivity of water-hyacinth supplied by T.T. Earle from a New Orleans population. 1956. Plakidas, A.G. Cercospora leaf spot ofAbelia. Mycologia 48:382-385. [116] A parasite of the popular ornamental shrub, Abelia grandiflora, is described: Cercospora abeliae sp. nov. from Baton Rouge. 1956. Schuster, R.M. Notes on Nearctic Hepaticae X. A study of Cephaloziella [117] rhizantha, C. florkhe and C. ludoviciana. Langlois collected the type of C. ludoviciana in 1888 in St. Martinville. Here incorporated in C. rizantha. 1956. Shinners, L.H. Euthamia pulverulenta Greene (Compositae) in southeastern [ 1 18] Louisiana. Field and Lab. 24:38. 1956. Shinners, L.H. Forestiera autumnalis Buckley (Oleaceae) in eastern Texas [1^9] and western Louisiana. Southwestern Nat. 1:87-88. "Though adequately described as long ago as 1863 (Proc. Acad. Nat. Sci. Phila. 14(1862):7), from 'Eastern Texas and Western Louisiana', this very distinct species has never been included in any flora or list." New collection from Sabine Parish (sterile material). 1956. Shinners, L.H. Hypochoeris glabra L. (Compositae) in Arkansas and Louisi- [120] ana. Southwestern Nat. 1:88. 19 1956. Shinners, L.H. Physostegia serotina (Labiatae), a new species from coastal [121] Louisiana and Texas. Field and Lab. 24:17-19. Type from Calcasieu Parish. Key to the eight species of Physostegia occurring in the "Gulf Southwest." 1956. Shinners, L.H. Tragia smallii Shinners, sp. nov. Field and Lab. 24:37. [122] T. betonicaefolia sensu Small. Paratype from Vernon Parish. 1956. Shinners, L.H. Yellow-flowered Oxalis (Oxalidaceae) of eastern Texas and [123] Louisiana. Field and Lab. 24:39-40. "Though often quite weedy, all the Oxalis of eastern Texas . . . and Louisiana are, in my opinion, undoubtedly native." Three species distinguished; one new variety described. 1956. Stemitzke, H.S. and J. A. Putnam. Forests of the Mississippi Delta. U.S. [124] Forest Serv., Southern Forest Exp. Sta., Forest Survey Release 78. 42 p. Brief descriptions of forest types. No map of forest types. Egler, 1961 . 1956. Ware, G.H. Vegetational zonation on a Red River sand bar near Natchi- [125] toches [Natchitoches Parish], Louisiana. Proc. La. Acad. Sci. 19:21-24. An obscure aerial view supplements this short discussion of a periodically flooded habitat. 1956. Wilson, H.R. Louisiana bryophytes. Bryologist 59:17-21. [126] Excellent prelude to Koch's more comprehensive Louisiana review published the same year. Wilson includes liverworts and adds a list of some fifty species collected in a baldcypress-tupelo gum swamp in Tangipahoa Parish, of which many are first reports for the state. 1957. Bandoni, R.J. The spores and basidia of Sirobasidium. Mycologia 49:250- [127] 255. Recharacterization of the genus based in part upon a Lowy collection of S. sanguineum from near Baton Rouge. 1957. Brown, C.A. Check list of woody plants of Louisiana, native, naturalized, [128] and cultivated. Baton Rouge, La. Forestry Comm. , Bull. No. 8. 16 p. Pamphlet. Second edition appeared in 1959, 76 p., 71 figs. Fourth edition in 1964 as La. Forestry Comm. Bull. 10. 80 p. 1957. Deiler, F.G. Vegetation management of storage water reservoirs at Garden [129] Island Bay [Plaquemines Parish]. Proc. La. Acad. Sci. 20:24-35. A massive invasion of vegetation (Phragmites, Typha, Zizaniopsis, Altemanthera, Eichhornia) and the chemical counterattack documented with excellent tables on species, species aggrega- tions, herbicide treatments and effects. Nutria cited as a natural check on water-hyacinth. 20 1957.* Dillon, O.W. Food habits of wild ducks in the rice-marsh transition area of [130] Louisiana. Proc. Ann. Conf. Southeast Assoc. Game and Fish Comm. 11: 114-119. 1957. Glasgow, L.L. and A. Ensminger. A marsh deer "die-off' in Louisiana. J. [131] Wildlife Mgmt. 21:245-247. Gulf marshes, Vermilion and Iberia Parishes. With information on vegetation types, site types, and extensive vegetation change related to a 1952 salt water inundation. Egler, 1961 . 1957. Hardin, J.W. Studies in the Hippocastanaceae, IV. Hybridization in Aescu- [132] /M5.Rhodora 59: 185-203. A. glabra X pavia has been collected in Bossier Parish. 1957. James, C. W. Notes on the cleistogamous species of Polygala in southeastern [133] United States. Rhodora 59:51-56. P. polygama forma obovata elevated to P. crenata, the type collected in New Orleans, Drummond 1832. 1957. Koch, L.F. Louisiana muscology 2. The herbarium of Tulane University, [134] New Orleans. Brittonia 9:96-71. Twenty-one taxa are added to the known flora of the state, mostly the collections of Penfound andF.P. Mackaness. 1957. Lowy, B. A new Exidia. Mycologia 49:899-902. [135] Type from Baton Rouge. 1957. Rock, H.F.L. A revision of the vernal species of Helenium (Compositae). [136] Rhodora 59:101-116, 128-158, 203-216. Southeastern genus with Louisiana representatives. Drummond and Arsene collections of H. vemale, a Joor collection of//, brevifolium from Slidell (St. Tammany Parish), 1887, and a Ball collection of //. flexuosum from near "Alexander" (Rapides Parish). //. drummondii is endemic to Texas-Louisiana coastal plain. 1957. Shinners, L.H. Polygonum bicorne Raf. instead of P. longistylum Small. [137] Rhodora 59:265-267. "... the much abused Rafmesque ...," "The special condemnation of the Florula Ludoviciana ...," "... the lie that the descriptions were scrappy and inadequate has been monotonously repeated by persons who never saw Robin's 3-volume book ..." More than a defense, this is a counterattack. 21 Shinners, L.H. Wahlenbergia marginata (Thunb.) A. D.C. (Campanulaceae) in Louisiana. Southwestern Nat. 2:44. A single small plant was found on a roadside in Rapides Parish. As Shinners predicted, it is now a common roadside weed in many parts of the state. 1957. Speairs, R.K., Jr. and B. Lowy. Notes on Tremella in North America. Proc. [139] La. Acad. Sci. 20:83-84. Of 19 North American species, 10 have been found in Louisiana. History of collections and species list for the state. 1957. Wherry, E.T. Reminiscences of John K. Small. Castanea 22:126-129. [ 140] Among the memorable anecdotes: Wherry's discovery that Small was color-blind - colonies of bright red clinopodiums or castilleias looked gray to him. Their discovery of the "Louisiana Irises," now internationally celebrated, while waiting for the Lake Pontchartrain ferry-boat to be repaired - they had hoped to be in New Orleans for lunch. 1958. Chabreck, R.H. Beaver-forest relationships in St. Tammany Parish, Louisi- [141] ana. J. Wildlife Mgmt. 22: 179-183. Plot sampling along streams occupied by beaver revealed that beaver used 22 woody plant species, (tabulated). Liquidambar styraciflua, Magnolia virginiana, Pinus glabra, and P. taeda received the most use. During summer months beaver fed heavily on roots and basal portions of herbaceous plants, esp. Leersia. Excellent and intriguing study. Literature cited contains a few related titles for our area. 1958. Clark, L. Frullania gymnotis found in the United States. Bryologist 61 :67. [ 142] Three widely separated locahties. The Louisiana material was collected in 1926 and reported by Svihla as F. obcordata in 1939. Dormon, C. Flowers native to the deep South. Baton Rouge: Claitor's Book Store. 176 p. Updates nomenclature of Wild Flowers of Louisiana, 1942, yet still no authorities cited. 33 color plates (registery often blurred) and 100 excellent line drawings, all by the author. Accurate and with all the fringe benefits of an informal catalogue. Gleason, H.A. Two new stations for Carex picta. Rhodora 60:175. Rare sedge first discovered near New Orleans by Drummond. New stations in Mississippi. Harris, V.T. and R.H. Chabreck. Some effects of Hurricane Audrey on the marsh at Marsh Island [Vermilion Parish], Louisiana. Proc. La. Acad. Sci. 21:47-50. 5400 foot line transect through a Spartina patens - Scirpus olneyi marsh. The island lies in Vermihon Bay. 22 1958. Karling, J.S. Synchytrium fulgens Schroeter. Mycologia 50:373-375, [146] Cook's Lx)uisiana collection: identified as S. fulgens, renamed 5. brownii by Karling in 1954, restored to S. fulgens: what one physiologist has superciliously called the Taxologic Cycle. 1958. Lowy, B. Anomalous phalloids. Mycologia 50:792-794, f 147] From Baton Rouge. 1958. Maisenhelder, L.C. Understory plants of bottomland forests. U.S. Forest [148] Serv., Southern Forest Exp. Sta., Occ. Pap. 165. 40 p. Thirty-six plants identified and illustrated with rather washed-out photographs. Especially for the Mississippi delta. 195; Ownbey, G. Monograph of the genus Argemone for North America and the [149] West Indies. Torrey Botan. Club Mem. 21(1): 1-159. Identity of Rafinesque's A. alba from Louisiana, pp. 138-140. For earher discussion of same, see Prain, D. , 1895, "An account of the genus Argemone," J. Botan. 33:330. 1958. Pursell, R. A. Discovery of Solmsiella kurzii in Louisiana. Bryologist 61:366- [150] 367. On Magnolia grandiflora. Weeks Island (salt dome), Iberia Parish. The first collection of this minute, allegedly rare, liverwort-like moss outside the type locality in Florida. 1958. Shinners, L.H. Carduus nutans L. (Compositae), a European thistle in north- [151] western Louisiana. Southwestern Nat. 3:220. 1958. Welden, A.L. Prodromus fungi ludovicianae. J. Tenn. Acad. Sci. 33:252-257. [152] Ten species of resupinate Homobasidiomycetes classed within the Thelephoraceae from lower Louisiana. Emphasis is on microscopic characters which, at that date, had only recently gained ascendency over gross morphology as an index of true relationships. Several new records for the state. One figure. 1958. Wood, C.E., Jr. The genera of the woody Ranales in the southeastern United [153] States. J. Arnold Arboretum 39:296-346. This is both the introduction and the first contribution to a projected generic study of the seed plants of the Southeast (bounded by and including Arkansas and Louisiana on the west). "... the objectives are toward a review and reorganization of familial and generic lines ... and, especially, toward bringing together at least a part of the vast botanical literature which bears upon the plants of this rich area." We will not pursue the project through this bibliography but we do alert theieader to its existence and great usefulness. 1959. Bick, G.H. Contributionsof Edward Foster to the biology of Louisiana. Proc. [154] Li. Acad. Sci. 22:8-17. 23 Foster's wide range of interests included microcrustaceans, insects, coins and medals, and, for our purposes, horticulture. Louisiana Society of Naturalists was organized in 1897 and Louisi- ana Entomological Society in 1920, both as a result of his efforts. Interesting tidbits about his friend. R.S. Cocks. 1959. Chamberlain, J.L. Gulf Coast marsh vegetation as food of wintering water [155] fowl. J. Wildhfe Mgmt. 23:97-102. Marsh food utilization, based on 1251 gizzard analyses, involving relative abundance of 24 plants, plus 25 others. Rockefeller Wildlife Refuge (Cameron Parish) of 86,000 acres. Egler, 1961. 1959. Humm, H.J. and R.M. Darnell. A collection of marine algae from the [156] Chandeleur Islands [St. Bernard Parish]. Univ. of Tex., Publ. Inst. Mar. Sci. 6:265-276. St. Bernard Parish. Annotated list of 35 species and discussion of ecological and phytogeo- graphic relations. Maps, table, and diagram of relative abundance in littoral zone. 1959. Kimble, R.B. and A. Ensminger. Duck food habits in southwestern Louisi- [157] ana marshes following a hurricane. J. Wildlife Mgmt. 23:453-455. Comparison of the results of this study after with a similar study before the hurricane (Chamberlain, 1959) indicates a great difference in the food eaten by ducks. Description of gross hurricane damage to vegetation amplifies conclusions drawn from duck gizzards. 1959. Lowy, B. and A.L. Welden. Synopsis of Louisiana polypores. Amer. [158] Midi. Nat. 61:329-349. Survey of polypores reported from the literature (Featherman and Langlois especially) and from authors' collections. Keys to genera and species, annotations and photographs of the more interesting species. 1959. Lytle, S.A., B.E. Grafton, A. Ritchie, and H.L. Hill. Soil survey of St. Mary [159] Parish, Louisiana. U.S. Dept. Agr., Soil Conserv. Serv. Soil Survey 1952(3). 45 p. An ongoing, county by county, national undertaking, indexed as Supt. of Documents No. A 57.38, and then alphabetically by county (or parish). Essential ecological data, maps and tables. 1959. Mullahy, J.H. Preliminary survey of the algal flora of the Chandeleur Islands [160] [St. Bernard Parish]. Proc. La. Acad. Sci. 22:62-68. A group of barrier islands protecting the southeastern marshlands of St. Bernard Parish. Twenty-seven species of green , brown, and red algae were collected, indicating that the islands may well contain Louisiana's best growth of marine algae. Good, compact bibliography. Author's abstract, "The algal flora of the Chandeleur Islands of Louisiana," appeared the same year in Internal. Botan. Congr. Proc. 9(2):275. 1959.* Nelson, I.S. Louisiana irises. In L.F. Randolph, ed. Garden irises, pp. [161] 227-235. Ithaca, New York. 24 1959. Reese, W.D. Syrrhopodon parasiticus in the southern United States. Bryolo- [162] gist 62: 182- 186. New state reports for Alabama, Mississippi and Louisiana. 1960. Blair, R.M. Deer forage increased by thinning in a Louisiana loblolly pine [163] plantation. J. Wildlife Mgmt. 24:401-405. Quantitative analysis of herbaceous and woody forage and palatable browse proved production of understory vegetation was directly related to pine-thinning intensity. Application of course to the millions of acres in the south shared by deer and loblolly pine. I960.* Harmon, B.G., C.H. Thomas, and L.L. Glasgow. Waterfowl foods in Lou- [164] isiana rice fields. North Amer. Wildlife Conf. Trans. 25:153-161. 1960. Kubota, J., V.A. Lazar, and K.C. Beeson. The study of cobah status of soils [165] in Arkansas and Louisiana using the black gum as the indicator plant. Proc. Soil Sci. Soc. Amer. 24:527-528. Nyssa sylvatica is a cobalt accumulator. Biol. Abst. 1960. Plakidas, A.G. Angular leaf spot of Magnolia. Mycologia 52:255-259. [ 166] Disease and pathogen (Isariopsis magnoliae) newly described, on M. grandiflora from Folsom (St. Tammany Parish), Louisiana. (Later reduced to synonomy under Cercospora magnoliae. Mycologia 54:448-454). 1960. Reese, W.D. Psilotum in Louisiana. Amer. Fern J. 50:269-270. [167] 1960. Shinners, L.H. Ranunculus trilobus (Ranunculaceae) in southern Louisiana: [ 168] new to the United States. Southwestern Nat. 5: 170. ■ 1960. Welden, A.L. The genus Cymatoderma (Thelephoraeceae) in the Americas. [169] Mycologia 52:856-876. The author resides in New Orleans, and many Louisiana citations document the study of this tropical and sub-tropical group. 1960. Welden, A.L. Prodromus fungorum ludovicianorum IL J. Tenn. Acad. Sci. [170] 35:231-237. Eight of the ten species reported as new records for the state. 1961. Crum, H. and L.E. Anderson. A new Fissidens from Louisiana. Bryologist [171] 64:345-348. F. kochii n. sp., Sarpy Wildlife Refuge, St. Charles Parish, about 15 miles northwest of New Orleans — a favorite haunt for Tulane botanists and zoologists. Four figures. 25 1961 . Darnell, R.M. Trophic spectrum of an estuarine community, based on studies [ 172] of Lake Pontchartrain, Louisiana. Ecology 42:553-568. Sources of primary organic matter, both autochthonous and allochthonous, include bacteria, phytoplankton, submerged and marsh vegetation. 1961. Dean, B.E. Trees and shrubs in the heart of Dixie. Birmingham. 246 p. [173] This field guide may be the most engaging blend of the sacred and profane on the botanist's book shelf. 430 species are described, annotated, and illustrated with line drawings. Supple- mental illustrations are many and range from useful photographs of living plants and herbarium specimens to a series of rather odd, unforgettable color plates, evidently the paintings of the illustrator, Forrest Bonner. His persimmons and pomegranates seem excerpted from a Ce- zanne still life, and the trumpet vine has the charm of a Matisse. Roland Harper aided with identification of some of the specimens and as a general fund of information to the author. 1961. Dukes, G.H., Jr. Some tertiary fossil woods of Louisiana and Mississippi. [174] [Abstract.] Amer. J. Botan. 48:540. Collectionsof petrified wood made in Vernon, Sabine, and De Soto parishes. Four new species reported from the Louisiana material (not enumerated in the abstract). 1961.* Eggler, W.A. Vegetation of the drainage basin of Grand Bayou Blue. In R.J. [175] Russell, ed., Louisiana coastal marsh ecology, pp. 1-23. La. State Univ., Coastal Stud. Inst., Tech. Rep. No. 14. 1961. Eggler, W.A. and W.G. Moore. The vegetation of Lake Chicot [Evangeline [176] Parish], Louisiana, after eighteen years of impoundment. Southwestern Nat. 6:175-183. Comparison made to Penfound's study of the lake (1949) just following impoundment. 1961 . Egler, F.E. Cartographic guide to selected regional vegetational literature. - [177] Where plant communities have been described. Part IL Southeastern United States. Sarracenia6:l-87. Annotated Louisiana bibliography and accompanying map, pp. 75-81, which have been of great service to this and to Ewan's original bibliography. 1961.* Halls, L.K. and T.H. Ripley, eds. Deer browse plants of southern forests. [178] U.S Forest Serv., Southern and Southeastern Forest Exp. Sta. 78 p. Illustrated guide to identification of about 35 species or species groups, with some information on forage value and management. Accounts of the individual species were prepared by authorities from all parts of the South. Punch, 1962. 1961.* Hoffpauir, CM. Methods of measuring and determining the effects of marsh [179] fires. IProc. Ann. Conf. Southeast. Assoc. Game and Fish Comm. 15:142-160. 26 1961. Koch, L.F. Louisiana hepaticology. 1. A list of species. Bryologist 64:54-57. [180] Brief historical sketch precedes list of hepatics: Faith Pennebaker Mackaness, collector and curator, most important contributor to Louisiana hepaticology to date. (Koch worked mostly with collections at Tulane). 1961. Lemaire, R.J. A preliminary annotated checklist of the vascular plants of the [181] Chandeleur and adjacent islands, St. Bernard and Plaquemines Parishes, Louisiana. Proc. La. Acad. Sci. 24:116-122. The Chandeleurs are the easternmost remnants of an abandoned delta of the Mississippi River. 1 19 species, 93 genera, 39 families. 1961. Lemaire, R.J. A preliminary annotated checklist of the vascular plants of the [182] marshesandincludedhigher landsof St. Bernard Parish, Louisiana. Proc. La. Acad. Sci. 24:56-70. In extreme southeastern Louisiana: more than 90*"^^ of the 617 square miles of land area of the parish is marsh. 280 species, 212 genera, 85 families. Habitat and abundance annotations. 1961. Lemaire, R.J. A range extension for Parapholis incurva. Rhodora 63:176- [183] 177. Freemason Island, St. Bernard Parish: a first record of this European grass on the Gulf Coast. 1961. Negus, N.C., E. Gould, and R.K. Chipman. Ecology of the rice rat, Oryzo- [184] mys palustris (Harlan), on Breton Island [St. Bernard Parish], Gulf of Mexi- co, with a critique of the social stress theory. Tulane Stud. Zool. 8:93-123. Island divided into five areas based on vegetational zonation. Description of Breton Island, pp. 95-98, Food habits, p. 109. Two photos, Fimbristylis and Opuntia-Yucca communities. 1961. Randolph, L.F., J. Mitra, and I.S. Nelson. Cytotaxonomic studies of Louisi- [185] ana irises. Botan. Gaz. 123:125-133. Karyotype analysesof /. /w/v«. /. giganticaerulea. I. hrevicaulis, the Abbeville Red population, and natural hybrids gave no evidence in support of Edgar Anderson's assumption of introgres- sive hybridization. See Randolph, 1966 and 1967, 1961. Reese, W.D. A contribution to the bryology of the southern United States. [186] Bryologist 64:50-54. Range extensions on the Gulf Coast for ten species of mosses, six representing new state records for Louisiana. Brief description, including noteworthy ferns and angiosperms, of Week's Island (Iberia Parish), one of the coast salt domes and the locality for three of the new mosses. 1961. Russell, N.H. Keys to Louisiana violets (K/o/a-Violaceae). Southwestern 187] Nat. 6:184-186. Two keys: spring (flowering) and summer (vegetative). Sixteen species. 27 1962. Daubs, E.H. The occurrence of Spirodella oligorrhiza in the United States. [188] Rhodora 64:83-85. New collections of this Asiatic aquatic from widely separated areas include two in Louisiana. New to the state. 1962. Glasgow, L.L. and H.A. Junca. Mallard foods in southwest Louisiana. Proc. [ 189] La. Acad. Sci. 25:63-74. The most important wild water fowl in the northern hemisphere. Grasses prove first in importance to mallards as well as to man. Four tables give exhaustive lists of plant materials, identified to species, taken from mallard crops. Related titles cited. 1962. Harrar, E.S. and J.G. Harrar. Guide to southern trees. New York, 709 p. [190] Dover reprint of the original 1946 publication with updated nomenclature. Ungenerously illustrated and of minimal usefulness for difficult genera like oaks and hickories. Clair Brown supplied the authors with "botanical materials" from our region. 1962.* Harris, V.T. and F. Webert. Nutria feeding activity and its effect on marsh [191] vegetation in southwestern Louisiana. U.S. Dept. Interior, Fish and Wildlife Serv., Spec. Sci. Rep. - Wildlife No. 64. 53 p. Supt. of Documents No. I 49. 15/3:64. 1962. Hodges, C.S. Fungi isolated from southern forest tree nursery soils. Mycolo- [192] gia 54:221-229. 45 fungi documented from Louisiana soils, the majority imperfects. Gonytnchum macrocladi- um a new record from soil in the U.S. 1962.* Jemison, E.S. and R.H. Chabreck. The availability of water fowl foods in [193] coastal marsh impoundments in Louisiana. North Amer. Wildlife Conf. Trans. 27:1-19. 1962. Nelson, I.S. Native Louisiana irises. Plants and Garden 18(l):62-63. [194] /. fulva, I. hrevicaulis, I. giganticaerulea, and their hybrids: culture, morphology, and breeding. Biol. Absi. 1962. Oliver, R.L. and W.H. Lewis. Chromosome numbers of 5/5>'r/>ic/imm (Irida- [195] ceae) in eastern North America. Sida 1:43-48. Vouchers for 5. albidum Raf., 5. sagittiferum Bickn., and .S'. bermudiana L. from Louisiana. We need all the help we can get with this genus! 1962. Penn, G.H. Percy Viosca Jr. - Naturalist. Tulane Stud. Zool. 9:234-237. [196] 28 ■'. . . one of the last of America's great naturalists . . . Viosca was an authority on the wild flowers of Louisiana, particularly the ecology and hybridization of irises ... In 1935 he publishes a lengthy article straightening out the taxonomic jumble in which hitherto a multitude of species had been described in Louisiana. After years of crossbreeding and countless observations in the wetlands his deduction that there were only four species of Louisiana irises was surprising, but remains undisputed by taxonomic botanists." 1962. Penn, G.H. Bibliography of studies by P. Viosca, Jr. Tulane Stud. Zool. [197] 9:239-242. A chain of publications from 1915 to 1%1 leaving no doubt that this was a naturalist of extraordinary scope. 1962. Perdue, R.E. , Jr. , Two new varieties and a new combination in Rudbeckia, [198] Rhodora 64:328-329. R. nilida Nun. var. lexuna Perdue var. nov. restricted to southeastern Texas and western Louisiana. 1962. Punch, L.E. Publications of the Southern Forest Experimental Station, 1955 [199] through 1961. U.S. Forest Serv., Southern Forest Exp.aSta., Occ. Pap. 108 (Supplement 1). 57 p. This list supplements the 1955 revision of Occ. Pap. 108 [Item 94]. Supplement 2, 1962 through 1970, appeared in 1971. Publications from 1971 to present appear in annual lists. All entries are accompanied by brief abstracts, and, again, many excellent papers are available on request to the S.F.E.S. in New Orleans. 1962. Shinners, L.H. Annual sisyrinchiums (Iridaceae) in the United States. Sida [2(K)] 1:32-42. The major area for all three species is in eastern Texas and Louisiana. Doubtful or excluded names: 5. brownii Small, named in honor of Prof. Clair A. Brown of Louisiana State Univer- sity, equals S. exile Bicknell. Many Louisiana collections cited for all species and one hybrid. 1962. Shinners, L.H. Drosera (Droseraceae) in the southeastern United States: an [201] interim report. Sida 1:53-59. Field observations in Louisiana and neighboring states. Collections from the state cited for D. capillaris Poiret, D. annua EL. Reed, and D. leucantha Shinners, sp. nov. Contends with Carroll E. Wood, Jr. over recognition of D. annua. 1962. Shinners, L.H. Evolution of the Gray's and Small's Manual ranges. Sida [202] 1:1-31. ■'Perhaps the first thing that strikes one about the historical record is the abundance of authors and floras in the North, and their paucity in the South - and the further strange fact that ail three authors of Southern floras came from the North ... Plainly the reasons are not botanical." (The author, by the way, is Canadian). This brilliant and intriguing historical synopsis of botany in the North and South is required reading. And there are predictions for the future, too. Regrettably, one of Shinner's works-in-hand, a flora of the Gulf Southwest, which was to include Louisiana west of the Mississippi, never saw the light of publication. 29 1962. Shinners, L.H. M/c/-omma /?roH'«e/ and its allies (Labiatae). Sida 1:94-97. [203] M. hrownei var. pilosiuscula Gray. Collection from St. Bernard Parish, Louisiana, examined. 1962. Shinners, L.H. Ranunculus trachycarpus (Ranunculaceae) in south central [2()4] Louisiana: new to North America. Sida 1:104-105. 1962. Shinners, L.H. Siphonychia transferred to Paronychia (Caryophyllaceae). [205] Sida 1:101-103. P. drummondiij . & G. added to Louisiana flora. 1962. Shinners, L.H. Synopsis of Collinsonia (Labiatae). Sida 1:76-83. [206] What Briquet and Rafinesque have done, Shinners has undone, and restored Collinsonia to its pre- 1897 limits. Louisiana collections cited for C. serotina Walter and C. tuberosa Michaux. 1962.* Simmons, E.G. and W.H. Thomas. Phytoplankton of the eastern Mississippi [207] delta. Univ. Tex., Publ. Inst. Mar. Sci. 8:269. 1963. Ewan, J. Riddell's place in the phytography of Louisiana. [Abstract.] Amer. [208] J. Botan. 50:634. Systematic botanist, 1807-1865. 1963. * Linnartz, N.E. Relation of soil and topographic characteristics to site quality [209] for southern pines in the Florida parishes of Louisiana. J. Forestry 61(6):434- 438. Loblolly, slash, and longleaf pines and their preferred soils. Depth to least permeable layer in soil profile, percent sand, drainage, pH, and slope are the variables that prove most significant in relation to "site index." Biol. Abst. 1963. Logan, L.A. A list of seed plants of Lincoln Parish, Louisiana. Proc. La. [210] Acad. Sci. 26:18-32. The parish lies in the shortleaf pine uplands of north central Louisiana. Background plus bare list: 381 genera m 1 14 families. 1963.* McGinn, L.R. and L.L. Glasgow. Loss of waterfowl foods in rice fields in [211] southwest Louisiana. Proc. Ann. Conf. Southeast Assoc. Game and Fish Comm. 17:30-34. 1963. Owens, A.G., Jr. and S. Riche. Monanthochloe littoralis (Gramineae) in [212] Louisiana. Sida 1:182. 30 1963. Richardson, A.L. Some monocotyledonous plants of East Baton Rouge [213] Parish. Proc. La. Acad. Sci. 26:9-15. Twenty-three species listed (grasses, sedges, rushes excluded) with full annotations. 1963. Shiflet, T.N. Major ecological factors controlling plant communities in Lou- [214] isiana. J. Range Mgmt. 16(5):23 1-235. Salinity of the free soil water and fluctuating water levels are the major factors that control plant communities in Louisiana's four and one-half million acres of marshlands. "All flesh is grass," is the text emphasized ; manipulation of salinity and water levels to change plant composition to better fit a range livestock enterprise. 1963. Shinners, L.H. Southeastern records of Stachys affinis (S. sieboldii) and S. [215] floridana (Labiatae). Castanea 28:44-46. S. floridana is added to the flora of Louisiana by a specimen from Orleans Parish. Now (1977) a fairly common weed in gardens here in New Orleans. Perennial with white, moniliform tubers - very striking. 1963. Solymosy, S.L. Plants of the U.S.L. "In-Use" Arboretum. Lafayette. 40 p. [2 16] Plant list is coded to map of U.S.L. campus, the "In-use" Arboretum. 1964. Barrett, E.R. Variations ofsummer fungi in soil areas in northeast Louisiana. [217] Proc. La. Acad. Sci. 27:53-58. Variations are quantitative, the only taxonomic distinction being Agaricales vs. Polyporales. Four general soil areas investigated: Coastal Plain, Mississippi Terrace and Loessial Hills, Flatwoods, Recent Alluvium; their distribution clearly mapped. 196 Crum H. and L.E. Anderson. Notes on Physcomitrium collenchymatum . [218] Bryologist 67:350-355. Several collections of Mackaness nee Pennebaker, Mackaness, Koch, and Reese firmly enlist this little moss in the state flora. 1964. Gagliano, S.M. An archaeological survey of Avery Island [Iberia Parish]. La. [219] State Univ. , Coastal Stud. Inst. 76 p. An evaluation of the extent of prehistoric occupation of this coast salt dome. Botanical material is slim, but discussion of geological conditions, past and present, has bearing on local phyto- geography. 1964. Johnston, M.C. Scirpus molestus (Cyperaceae), sp. nov. from Arkansas, [220] Louisiana, and Texas. Southwestern Nat. 9:310-312. PrcMously mistaken tor -S. kuilolepis, the type of which was collected in New Orleans. 1964. Lowy, B. Pisolithus in Louisiana. Mycologia 56:319. [221] Infrequently reported from North America, P imcionus is in fact one of the commoner Gasteromycetes of our region. 31 1964. Radford, A.E., H.E. Ahles, and C.R. Bell. Manual of the vascular flora of [222] the Carolinas. Chapel Hill. 1 183 p. Much overlap naturally exists between the Carolina flora and ours, and, despite the fact that Louisiana fails to qualify as one of the "eight of the southeastern states outside the Carolinas" for purposes of range citations, the book is nonetheless possibly the most useful single up-to- date volume for our area in termsof coverage and cost. It is currently used at L.S.U. as text for the introductory plant taxonomy course. A good many taxa are illustrated with excellent line drawings. Keys are artificial and highly serviceable. 1964. Reese, W.D. Notes on Louisiana mosses. Bryologist 67:206-209. [223] Part one of at least six entries (1964, "65. "67, "69, '72, "74): Bryologist 6^:243-245, 70:124-125. 72:68-69, 75:95-97, 77:467-468. Additions and adjustments to the roster of Louisiana mosses, including many range extensions to the author"s original state records. 1964. Richardson, A.L. A botanical report of some Archichlamydeae of East [224] Baton Rouge Parish. Proc. La. Acad. Sci. 27:21-39. 173 species m 97 genera and 31 families. Arrangement of families follows Engler and PrantI through the Umbelliferae. Full annotations. 1964. Shinners, L.H. Scutellaria thieretii (Labiatae), a new species from coastal [225] Louisiana. Sida 1:251-252. Sp. nov. Holotype from Vermilion Parish; also collected from Cameron Parish. Shinners, a splitter, named this new skullcap for "an energetic and productive collector." 1964. Shinners, L.H. Two youngias {''Crepis japonica'' : Compositae) introduced in [226] the southeastern United States. Sida 1 :386-388. First reported from U.S. in Small's Manual as Crepis from only southern Louisiana. Several collections cited from the state for Y. japonica (L.) DC. 1964. Thieret, J.W. Additions to the flora of Louisiana. Castanea 28:169-170. [227] "1963'; Nine new species. 1964. Thieret, J.W. Fatoua villosa (Moraceae) in Louisiana: new to North [228] America. Sida 1:248. From southeast Asia. First U.S. collection is not from Lafayette, however, but most likely New Orleans: G.P. DeWolf 5.n. 11 Oct. 1950, Tulane Campus (NO). By 1977 a common garden weed in Lafayette and New Orleans, and how very unlike a mulberry! 1964. Thieret, J.W. Lysimachia japonica (Primulaceae) and Clinopodium gracile [229] (Labiatae) in Louisiana: new to the United States. Sida 1 :249-250. 32 1964. Thieret, J.W. More additions to the Louisiana flora. Sida 1:294-295. [230] Ten state records, including three Cyperus spp. 1965. Dorman, C. Natives preferred. Baton Rouge: Claitor's Book Store. 217 p. [231] Native trees and flowersof the southeast, where they can and cannot be grown. Author, herself a Louisiana native, tempers her enthusiasm with a caution to "diggin' women and men" against immoderate and indiscriminate transplanting. 1965. Ewan, J. French naturalists in the Mississippi Valley. In J.F. McDermott. The [231A] French in the Mississippi Valley, pp. 159-174. Urbana. Brothers Louis and Jean Prat, Le Page du Pratz, Joseph Lakanal, etc., and their La. travels noticed. 1965. Ewan, J., ed.. Letters from Charles Sprague Sargent to Reginald Somers [231B] Cocks, 1908-1926. J. Arnold Arboretum 46:1-44, 122-159, 324-361, 411-444. Sargent wrote to Cocks in 1911, from the Arnold Arboretum, "To show how poor we are in Louisiana plants I could not find this morning a Louisiana specimen of Liquidambar in our herbarium." Most fascinating are Sargent's struggles with collections (and taxonomy) of the tougher genera: willows, hawthorns, plums, oaks, basswoods, hickories. Reprinted as separate with change of paging the same year. 1%5. Lemmon, B.E. Notes on a recent collection of Fissidens neonii, Bryologist [232] 68:325-331. One of very few Louisiana endemics. The first reappearance of this moss since the original collection by Brother Neon in 1931 near Lafayette, which E.B. Bartram (1932) described as Moenkemeyera. Eleven figures. 1965.* Livingstone, R., Jr. A preliminary bibliography with KWIC index on the [233] ecology of estuaries and coastal areas of the Eastern United States. U.S. Dept. Interior, Fish and Wildlife Serv., Spec. Sci. Rep. - Fisheries No. 507. 352 p. Supt. of Documents No. I 49. 15/2:507. 1965. Lowy, B. and W.B. Cooke. The 1960 Louisiana foray. Mycologia 57:478-483. [234] A good harvest for the Mycological Society of America: Myxomycetes, Ascomycetes, Basidi- omycetes, and Fungi Imperfecti identified from 7 collecting localities near Baton Rouge. One new genus, Gliocephalotrichum, was describ>ed from the Tunica Hills area, West Fehciana Parish. 1965. Reese, W.D. and B.E. Lemmon. A natural hybrid between Weissia and [235] Astomum and notes on the nomenclature of the North American species of Astomum. Bryologist 68:277-283. W. controversa X A. ludovicianum, in southern Louisiana. 33 1965. Rougeou, C.L. A tribute to Professor Ira S. Nelson. La. Soc. Hort. Res. [236] Monthly News Letter 6( 12) :54-56. Professor of Horticulture at University of Southwestern Louisiana, who brought many new and rare species into cultivation, is remembered both as teacher and horticulturalist. 1965. Solymosy, S.L. Limnophila indica (R.Br.) Druce (Scrophulariaceae) in Lou- [237] isiana. Sida 2: 175. 1965. Stemitzke, H.S. Louisiana forests. U.S. Forest Serv., Resource Bull. SO-7. [238] 31 p. Volume in growing stock (trees at least 5 inches in diameter) has risen 43 percent for softwood and declined 20 jsercent for hardwood since the midfifties. Of value not for its botany, for it is pure and unregenerate forestry, but as a comprehensive state survey loaded with tabular data, it does allow botanists an intimate look at foresters' activities without our having to wire-tap. 1966. Banks, D.J. Paspalum minus (Gramineae) in Louisiana and Mississippi. [239] Rhodora 68:94-96. New locations bridge the gap between Texas and Alabama stations. Diploid race indicated by new chromosome counts, n=lO. 1966. Grelen, H.E. and V. Duvall. Common plants of longleaf pine - bluestem [240] range. U.S. Forest Serv., Southern Forest Exp. Sta., Res. Pap. SO-23. 96 p. Descriptions and excellent botanical drawings of more than 80 understory species: Andropogon and Panicum, legumes and composites are the dominants. Notes on values for cattle and wildlife. An elegant field guide. 1966. Krai, R. Eriocaulaceae of continental North America north of Mexico. Sida [241] 2:285-332. Tropical American family with outliers in southeastern U.S. Eriocaulon decangulare, E. lexense, E. compressum, Lachnocaulon anceps on record tor Louisiana, and others so close as to be suspected. 1966. Krai, R. Juncus capitatus Weigel (Juncaceae) in Louisiana: new to the United [242] States. Sida 2:390-392. But see Sida I'Alib — "^Juncus capitatus (Juncaceae) previously reported from the United States," for corrections. 1966. Krai, R. Observations on the flora of the southeastern United States with [243] special reference to northern Louisiana. Sida 2:395-408. Emphasis on northern woodland elements. Approximately a dozen species new to the state may be sifted from a longer list, but almost all entries represent range extensions and elucida- tions. 34 1966. Krai, R. Xyris (Xyridaceae) of the continental United States and Canada. [244] Sida 2: 177-260. A difficult genus of the Gulf and Atlantic Coastal Plains with about half of some twenty species native to the state. Many of the author's voucher specimens for another smears collected in Louisiana. Generously illustrated. 1966. Lemmon, B. A. An ecological and floristic study of the mosses of Lafayette [245] Parish. Proc. La. Acad. Sci. 29:23-36. Annotated checklist includes 91 taxa, of which 45 are reported as "firsts" for the parish. Four bryological habitats and their mosses described. Study based on collections of Brother Neon, William D. Reese, Correll and Correll, and the author. 1966. Lemmon, B.E. Fissidens hyalinus in Louisiana. Bryologist 69:241-243. [246] Disjunct in Japan and eastern U.S. New to the state. One figure. 1966. Maples, R.S., Jr. and D.D. Lutes. A checklist of ferns in Lincoln Parish, [247] Louisiana. Amer. Fern. J. 56:33-36. 16 species, 6 parish records. Annotations. 1966. Moore, W.G. Central Gulf States and the Mississippi Embayment. In David [248] G. Frey, ed.. Limnology in North America, pp. 287-300. Madison, Milwau- kee, and London. 734 p. Arkansas, Louisiana, Mississippi, and Alabama, with emphasis on our area with its rich and varied network of inland waters. Vital background for students of aquatic botany. Map of major drainage systems, p. 289. Extensive references. 1966. Randolph, L.F. Iris nelsonii, a new species of Louisiana iris of hybrid origin. [249] Baileya 14:143-169. Known locally as the Abbeville Reds, the assumed parental species of this stable hybrid are /. fulva and /. giganticaerulea. In-depth cytogenetic and morphological analysis with discussion of hybridization and introgression. Excellent photographs of plants and habitats. 1966. Reese, W.D. fiar/?e//flpen^w/a(Sull.)Fleisch., a review of its distribution and [250] comments on related species. Bryologist 69:208-213. Type locality Lafourche Parish. Occurring in China and Japan, Mexico and in the United States only in Louisiana where it was first collected by Riddell. W.S. Sullivant described it as Meteorium in 1856 in ed. 2 of Gray's Manual. Two figures. 1966. Rickett, H.W. Wildflowers of the United States. II. The southeastern states. [251] 2 vols. New York. 688 p. 241 folio-size, color plates (each with 6 or 7 separate photos) and numerous text figures make this the most sumptuous of modem wildflower picture books. Woody plants excluded, but many obscure herbs normally passed over are admitted and illustrated. Clair Brown is among the collaborators. 35 1966. Society for Louisiana Irises. 25th anniversary publication. Lafayette. 5 1 pp. [25 1 A] Reminiscence of John K. Small, a tribute to Ira S. Nelson, and Caroline Dorman's History of the Louisiana Iris make this a valuable botanical contribution. 1966. Solymosy, S.L. Poisonous plants. La. Soc. Hort. Res. Monthly News Letter [252] 7(3): 1-22. Annotated, well-informed list of both native and cultivated plants from our area. As herbarium curators know well, public inquiries on this subject generally outnumber all others. 1966. Stuckey, R.L. The distribution of Rorippa sylvestris (Cruciferae) in North [253] America. Sida 2:361-376. Evidence for an introduced status includes two Louisiana collections of 1885: W.B. Waite's from Orleans Parish (US) and A.B. Langlois" from Plaquemines Parish (MICH). 1966. Stuckey, R.L. Rorippa walteri and R. obtusa synonyms of R. teres (Crucifer- [254] ae). Sida 2:409-418. Other synonyms include Nasturtium micropetalum Fischer and Meyer and M obtusum Nuttall, holotypes from in and near New Orleans. Photograph of a portion of the latter, from Philadel- phia Academy Herbarium. Information on Henry Little, its obscure collector, is scant. 1966. Thieret, J. W. Additions to the Louisiana flora. Sida 2:264-265. [255] Ten species towards a projected Louisiana Flora. 1966. Thieret, J.W. Habit variation in Myrica pensylvanica and M. cerifera. Casta- [256] nea 31:183-185. Habit variation within M. cerifera - M. pusilla complex studied in Louisiana is a response to habitat and taxonomically of little significance. Specific distinction between them is untenable. 1967. Bamforth, S.S. A microbial comparison of two forest soils of southeastern [257] Louisiana. Proc. La. Acad. Sci. 30:7-16. A pine forest on the Prairie Terrace near Hickory (St. Tammany Parish) vs. a Delta hardwood bottomland forest south of New Orleans. Soils examined for bacteria, fungi, and protozoa. Two tables. 1967. Boudreaux, B.H. In Memorium: Percy Viosca, Jr., 1892-1961. Proc. La. [258] Acad. Sci. 30:5-6. See also Penn's tribute of 1962, from which much of the information here was taken. 1967. Delahoussaye, A.J. and J.W. Thieret. Cyperus subgenus Kyllinga (Cypera- [259] ceae) in the continental United States. Sida 3: 128-136. Three of the four spp. Louisiana residents. Illustrations of achenes and spikelets, not habit; distribution maps, limited synonomy, and species descriptions. Key to subgenera or distinctive features of Kyllinga would have hastened our focus. 36 1967. Duncan, W.H. Woody vines of the southeastern United States. Sida 3: 1-76. [260] Twenty-nine genera treated, 21 occurring in the state. Physical map of study area, distribution maps, excellent drawings, keys, and species descriptions for large genera. NO and LAF material examined. 1967. Ewan, J. Introduction to C.S. Rafinesque, Florula ludoviciana (1817). Clas- [261] sica Botanica Americana 5:i-xl. New York and London. Introduction of historical and taxonomic interest and is, like Shinner's article of 1957, sympa- thetic \o the man tor whom 'much maligned" has become a permanent epithet. 1967. Hutto, J. and R.D. Thomas. Hottonia inflata (Primulaceae) in Ouachita [262] Parish, Louisiana. Sida 3: 187. A second location for the state. 1967. Pinkava, D.J. Biosystematic study of Berlandiera (Compositae). Brittonia [263] 19:285-298. B. X hetonicifolia (Hook.) Small (pro. sp.) = B. pumila X texana to the east and west of our area, respectively. Drummond collected the type in New Orleans in 1833. 1967. Radford, A.E., C.R. Bell, J.W. Hardin, and R.L. Wilbur. Contributor's [264] guide for the "Vascular flora of the southeastern United States. " Chapel Hill, North Carolina. 20 p. A floristic manual to the forested region of the southeastern U.S., the work of some 50 contributors, was projected for publication by 1975. Individual contributions have appeared in the periodical literature while awaiting incorporation into the Manual. For example, J. Elisha Mitchell Soc. (1971 & 72) carried Hardin's "Studies of the Southeastern U.S. Flora ' I, II, and III (X7:.^9-.^(); (S8:3()-.^2). which provides keys, PDmenclature and distribution to the Eietulaeeae. gymnosperms, Magnoliaceae and Iliiciaceae. 1967. Randolph, L.F., I.S. Nelson, and R.L. Plaisted. Negative evidence of intro- [265] gression affecting the stability of Louisiana iris species. Cornell Univ., Agr. Exp. Sta., Memoir 398. 56 p. More information is brought to bear on Anderson's treatise of introgressive hybridization (1949), with comparisons of allopatric and sympatric populations of three species and three groups of natural hybrids. See also Randolph, 1966. 1967. Reese, W.D. The discovery of Tortula vectensis in North America. Bryolo- [266] gist 70:1 12-1 14. A moss whose total known range is southern Louisiana and the Isle of Wight, England! Three figures. 1967. Reese, W.D. and J.W. Thieret. Botanical study of the Five Islands of Louisi- [267] ana [Iberia and St. Mary Parishes]. Castanea 31:251-277. "1966." 37 The "Five Islands" are Louisiana's salt domes on the central Gulf Coast, one attaining a height of more than 150 feet, and all offering habitats not otherwise afforded in the surrounding marshlands and prairie terraces. Geology, climate, phytogeography, and botanical history. Plant list includes mosses and liverworts, and bibliography includes several pertinent geological entries. Ten years later, the powers that be are at loggerheads over the use and misuse of these unique formations. 1967. Thieret, J.W. Thirty additions to the Louisiana flora. Sida 3:123-127. [268] 1967. Thomas, R.D. Burmannia biflora (Burmanniaceae) and Bowlesia incana [269] (Umbelliferae) in Ouachita Parish, Louisiana. Sida 3:183-184. Previous state collections discussed. 1968. Bamforth, S.S. Forest soil protozoa of the Florida Parishes of Louisiana. [270] Proc. La. Acad. Sci. 31:5-15. Seventeen forest sites examined for protozoa, algae, and proteolytic and cellulose decompos- ers. Forest types include oak-hickory, bottomland hardwood-cypress, and pine-deciduous. Two tables. 1968. Chabreck, R.H., T. Joanen, and A.W. Palmisano. Vegetative type map of [271] the Louisiana coastal marshes. La. Wildlife and Fish Comm. New Orleans. Fresh, intermediate, brackish and saline marshes are recognized and plotted. See Chabreck, 1972, for full account of species composition. 1968. Ewan, J. A bibliography of Louisiana botany. Southwest La. J. 7:1-83. [272] "1967.- Reviewed by Earl L. Core, Castanea 33; 154-155, and by F. A. Stafleu in Taxon 17:433-434. 1968. Haynes, R.R. Potamogeton in Louisiana. Proc. La. Acad. Sci. 31:82-90. [273] Descriptions and distribution maps for each of nine species. Habit sketches from herbarium specimens are excellent; fruits also illustrated. P. epihydrus is new to the states flora, known only from one locality in Washington Parish. 1968. Hutchins, R.E. Island of adventure. A naturalist explores a Gulf Coast [274] wilderness. New York. 243 p. Island in the Pascagoula River, Mississippi. Refreshing and accurate popular account relevant to Louisiana offers much first hand, early-morning observation with the dew still on it. Botany accent is on the sublime and the curious; insectivorous and saprophytic plants, Fibonacci numbers of opposing spirals in the florets of Composite heads and the scales of pine cones, pollination tricks in jack-in-the-pulpit. Illustrated with fine black and white photographs by the author. 38 1%8. Jones, R.E. A study of the mosses of Ouachita Parish, Louisiana. Castanea [275] 33:18-30. Bryological habitats discussed at length, plus annotated checklist. 1968. Kniffen, F.B. Louisiana, its land and people. Baton Rouge. 196 p. [276] The basic geography is more valuable than the chapters on soil and vegetation per se. Also, Spanish-moss gathering, a picturesque and nearly-vanished folk economy, pp. 98-99. 1968. Lytle, S.A. The morphological characteristics and relief relationships of [277] representative soils in Louisiana. La. State Univ., Agr. Exp. Sta., Bull. No. 631. 23 p. 1968. Reese. W.D. New moss records for the southeastern United States. Bryolo- [278] gist: 71:138-139. Predominantly Louisiana collections. 1968. Reese, W.D. and R.E. Jones. Diphyscium foliosum in Louisiana. Bryologist [279] 71:142. Of phytogeographic interest, this apparent Pleistocene relict has not previously been found on the coastal plain. 1968. Richards, E.L. A monograph of the genus Ratibida. Rhodora 70:348-393. [280] R. columnifera cited from Texas and Arkansas, not Louisiana. Tulane herbarium holds an R.S. Cocks 1908 collection from R.R. tracks in Shreveport. Clair Brown (1972) calls it "widely distributed in prairie and pinelands" but must be confusing it with the common R. peduncularis. 1968. Robbins, H.C. The genus Pachysandra (Buxaceae). Sida 3:211-248. [28 1 ] The single American species of this southeast Asian genus is native to Louisiana, disjunct in the Tunica Hills. Voucher from West Feliciana Parish. See also item 413. 1968. Thieret, J.W. Additions to the vascular flora of Louisiana. Proc. La. Acad. [282] Sci. 31:91-97. As well as an annotated list of 27 new species, the accomplishments of the Louisiana Flora Project are reviewed, and mention is made of the unsatisfactory state of knowledge of the Louisiana flora. 1968. Welch, W.C. and T.E. Pope. Vines for Louisiana. La. State Univ. and Agr. [283] and Mech. Coll. , Coop. Ext. Pub. 1525. 17 p. Pamphlet consisting mostly of a list and description of some of the native and exotic vines (44 in number) that may be grown in Louisiana, along with horticultural background. Photographs. 39 1969.* Anon. Weeds of the southern United States. La. State Univ. and Agr. and [284] Mech. Coll. , Coop. Ext. Pub. 1516. 42 p. 1969. Bamforth, S.S. Protozoa and algae of the Mississippi deltaic soils. Proc. La. [285] Acad. Sci. 32:68-77. Litters and soils from forests and grasslands of natural levee ridges, swamps, and a marsh. 1969.* Blair, R.M. andE.A. Epps, Jr. Seasonal distribution of nutrients in plants of [286] seven browse species in Louisiana. U.S. Forest Serv., Southern Forest Exp. Sta.,Res. Pap. SO-51.35p. Dry matter, protein, ether extract, crude fiber, N-free extract, ash, Ca, and P contents determined. Biol. Abst. 1969. Depoe, C.E. Bacopa egensis (Poeppig) Pennell (Scrophulariaceae) in the [287] United States. Sida 3:313-318. First U.S. collections by Dr. Josiah Hale in the early 1800's from Louisiana remained the only collections (4 sheets at NY) apparently extant until the plant's rediscovery in the state in 1966. Summary of detective work, drawings of leaves and fruit, herbarium specimen shown. Now firmly established and here added to the state flora. 1969. Eleuterius, L.N. and S.B. Jones, Jr. A floristic and ecological study of pitcher [288] plant bogs in south Mississippi. Rhodora 71:29-34. Ecological brief cites fire in the maintenance of this unique subclimax community. Pitcher plant bog bibliography. 1969. Ewan, J. Historicalproblemsfortheworkingtaxonomist.Taxon 18:194-203. [289] A few pages on deciphering and verifying herbarium labels are illustrated by some samples from Louisiana botany: J.L. Riddell labels are examined and followed into the hands of other botanists and curators such as Joseph F. Joor, Josiah Hale, and John Torrey, with something less than fidelity to the collector's original information as one of the consequences. 1969. Ewan, J. 10 conifers for southern (New Orleans and vicinity) Louisiana. In [290] Handbook on conifers. Plants and Gardens 25 (2). Special Printing, pp. 41-42. S.L. Solymosy's "Conifers for southwestern Louisiana" appears on p. 42. 1969. Freeman, J.D. Trillium gracile (Liliaceae), a new sessile-flowered species [291] from eastern Texas and Louisiana. Sida 3:289-292. Plants previously treated as T. ludovicianum. Type from Texas. 40 1969. Ingram, J. and W.J. Dress. The Louisiana irises and Hortus Third. Baileya [292] 16:93-97. "1968." The objective here is to account for Latin names used for these irises in forthcoming Hortus Third. Ins X vinicolor Small elucidated, and a synoptical history of the Louisiana Ins question becomes the fnnge benefit. Three Ins plates reprmted from Curtis' Botanical Magazine and Addisonia. 1969. Isley, D. Legumes of the United States: I. Native Acacia. Sida 3:365-386. [293] The only Louisiana Acacia is A. smallii Isely. published origmally m Small's Manual as Vachellia denstflora Alexander, type from Bayou La Fourche (La Fourche Parish) nearCut-Ott - a segregate of the A. farnesiana complex. 1969.* Linnartz, N.E., Ed. Ecology of southern forests. La. State Univ. Proc. Ann. [294] Forestry Symp. No. 17. 203 p. Twelve papers discuss the relation of southern forest communities to climate, physiographic features, soil, water, and biotic factors, as well as commercial matters and modification procedures. Biol. Abst. 1969. Miller, L.W. Acalypha graciliens Gray var. delzii L. Miller, var. nov. (Eu- [295] phorbiaceae). Sida 3:447. From Texas east into central Louisiana. 1969. [Nelson, T.C. and W.M. Zillgitt]. A forest atlas of the South. U.S. Forest [296] Serv., Southern and Southeastern Forest Exp. Sta. New Orleans and Ashe- ville. North Carolina. ■Virginia to eastern Texas, strictly maps, and these worth at least a thousand words each: soils and forest types, climate, insect pests, fire occurrence rates and more. Design and printing, the crux of such a visual book, are superb. 1969. Ownbey, G. and W.A. Olson. Cytotaxonomic notes on the species of [297] Cirsium native to the southeastern United States. Rhodora 71:285-296. "C". muticum from Louisiana differs in small ways from the species as we are accustomed to see it in the northern wetlands." Camera lucida metaphase figures for C. muticum and C. horridu- lum from Louisiana collections in Lafayette Parish. 1969. Reese, W.D. and A.W. Westling. Fissidens asplenioides on the Gulf coastal [298] plain. Bryologist 72:71-72. First Louisiana report of this Pleistocene relict moss. 1969. Rollins, R.C. On a weed species of /?or/ppfl. Rhodora 71:552-553. [299] R. indica (\^.)H\Qm. var. apew/a (DC) Hochr, a native of eastern Asia, collected in the U.S. from Louisiana and from near Portland, Oregon. 41 1969. Rylander, M.K. An ecological and floristic study of the vegetation of the [3()0] Delta Regional Primate Research Center, Covington, St. Tammany Parish, Louisiana. Proc. La. Acad. Sci. 32:83-111. 500 acres of second growth loblolly pine forest yield 325 vascular species in ten vegetation associes. Invasion of plants in a cleared area also studied. Six figures painstakingly composed. 1969. Thieret, J.W. Baptisia lactea (Rafinesque) Thieret, comb. nov. (Legumino- [301] sae).Sida 3:446. Based on Dolichos lacteus Rafinesque, Florula Ludoviciana, 1817. The plant is frequent in the prairie region of Louisiana. 1969. Thieret, J.W. Dopatrium junceum (Scrophulariaceae) in Louisiana. Sida [302] 3:448. 1969. Thieret, J.W. Rumex obovatus and Rumex paraguayensis (Polygonaceae) in [303] Louisiana: new to North America. Sida 3:445-446. 1969. Thieret, J.W. Sagittaria guayanensis (Alismataceae) in Louisiana: new to the [304] United States. Sida 3:445. Like so many of the author's discoveries, a weed in a rice field. 1969. Thieret, J.W. Trifolium vesiculosum (Leguminosae) in Mississippi and Lou- [305] isiana: new to North America. Sida 3:446-447. 1969. Thieret, J.W. Twenty-five species of vascular plants new to Louisiana. Proc. [306] La. Acad. Sci. 32:78-82. One is the marine angiosperm Halophila engelmannii Aschers. Celastrus scandens L. (bitter- sweet), originally noted by R.S. Cocks in 1914 from West Feliciana, subsequently disclaimed by Clair Brown in 1945 and, following suit, by O.K. Brizicky in 1964, here readmitted to the state's flora, based on Thieret's verification of Cocks' voucher specimen at the Tulane Herbarium, evidently overlooked by Brown. Status in the wild, however, remains uncertain. (See also item 413). 1969. Thieret, J.W.. R.R. Haynes, and D.H. Dike. Blyxa fl«6em7e(Hydrocharita- [307] ceae) in Louisiana: new to North America. Sida 3:343-344. Another addition to the state's noteworthy aquatic angiosperm flora, this a native of the Old World tropics. 1969. Vogel, E. and A.D. Oliver, Jr. Evaluation oi Arzama densa as an aid in the [308] control of water-hyacinth in Louisiana. J. Econ. Entomol. 62:142-145. 42 The larvae of this common noctuid moth of southern Louisiana were found to feed on water-hyacinth, causing death of some plants and preventing development of seed heads on others. Pickerelweed also eaten. See Ann. Entomol. Soc. Amer. 62; 749-752 for life history. 1970. Anderson, L.C. Studies on Bigelowia (Astereae, Compositae) I. Morphology [309] and taxonomy. Sida 3:451-465. Two species, both Louisiana natives. State collections tabulated for morphological studies. Map and four tables. 1970. Correll, D.S., M.C. Johnston, and collaborators. Manual of the vascular [310] plants of Texas. Renner: Texas Research Foundation. 1881 p. Rising to the occasion of a vascular flora that numbers nearly 5000 species and embraces habitats as diverse as woodland, prairie, and desert, this is a mighty work of industry and accuracy and, for our needs, makes a fine complement to Small's 1933 Manual which extends west only to the Mississippi River. Louisiana is included in range citations, and discussion of the East Texas Forest Region and Gulf Prairies and Marshes Area holds true for our area as well. Species descriptions are exhaustive. 1970. Kuprionis, J. Louisiana Tech Arboretum; its history and development. Rus- [311] ton, Louisiana. 38 p. Includes a list of trees and shrubs in the arboretum, coded to map of same. 1970.* Meyers, S.P., M.E. Nicholson, P. Miles, J.S. Rhee, and D.G. Ahearn. [312] Mycological studies in Barataria Bay, Louisiana, and biodegradation of oys- ter grass, Spartina alterniflora. La. State Univ., Coastal Stud. Bull. 5:111-124. 1970. Mitchell, R.S. A re-evaluation oi Polygonum meisnerianum in North Ameri- [313] ca.Rhodora 72:182-188. Earliest U.S. collections seen by the author all from Louisiana: Drummond, 1832, from New Orleans; Riddell, 1838, from Madisonville (St. Tammany Parish); and Langlois, 1885 and 1892. Collections have been extremely infrequent in North America since the turn of the century, and author notes that there are no specimens at LSU or LAP. 1970. Pursell, R.A. and W.D. Reese. Phytogeographic affinities of the mosses of [314] the Gulf Coastal Plain of the United States and Mexico. J. Hattori. Bot. Lab. 33:115-152. The mosses of the Gulf Coast of the United States are descendents of the Tertiary moss flora of eastern North America, with a limited number of species from regions to the south. 1970. Reese, W.D. and S. Tucker. The 1967 foray of the American Bryological and [315] Lichenological Society in Louisiana and Texas. Bryologist 73:692-701. Collections in western Louisiana yielded 30 lichen taxa new to our state. Check list. 43 1970. Rense, W.C. The perique tobacco industry of St. James Parish, Louisiana: a [316] world monopoly. Econ. Botan. 24:123-130. A delicate and complex curing process yields high quality and flavor and high consumer demand. Circumstances beyond the control of a 300 acre per year small business make perique's future regrettably uncertain. Map and illustrations. 1970. Rhodes, D.C. Psilotum nudum (Psilotaceae) in North Louisiana. Sida 3:525. [317] Lincoln Parish, extending known range 250 miles northward. 1970. Richard, M. A. The swamp maple, Acer rubrum var. drummondii, a neglect- [318] ed native ornamental tree. La. Soc. Hort. Res. J. 11:1-23. Gardeners may not be as interested as taxonomists and ecologists in a few of the topics, for example, morphological variability and climatic influence on date of blooming. 1970.* Smith, W.C. Spartina "die-back" in Louisiana marshlands. La. State Univ., [319] Coastal Stud. Bull. 5 :89-96. 1970. Thieret, J. W. Bacopa repens (Scrophulariaceae) in the conterminous United [320] States. Castanea 35 : 132- 136. Nomenclature; field and laboratory observations on habitat, associates, morphology; citation of U.S. collections. Thieret's field work done in the rice ponds of Acadia Parish. 1970. Thieret, J.W. Nemophila microcalyx, an incorrect name. Rhodora 72:399- [321] 400. Rafinesque again visiting nomenclatural turmoil on future generations. His epithet trilobum predates Nuttall's microcalyx and hence the correct name is N. trilobum (Raf.) Thieret. 1970. Thieret, J.W. Notes on Epifagus. Castanea 34:397-402. "1969." [322] E. virginiana (Orobanchaceae). Field observations were made largely in Evangeline and East FeUciana parishes, Louisiana. 1970. Tucker, S.C. Langlois's collection sites of Louisiana lichens. Bryologist 73: [323] 137-142. Parish locations for about 30 sites mapped and listed, with notes and known dates of collections. A few place names cited remain unlocated: Aurelia Co. (or New Aurelia), Bayou Millien (or MiJlieu), Bois Charmante, and Jardin de Bouchetel. Langlois references gathered from local histories and biographical accounts. 1970. Weniger, D. Cacti of the Southwest: Texas, New Mexico, Oklahoma, Arkan- [324] sas and Louisiana. Austin, Texas; London, England. 249 p. The genus Opuntia is a minor but quite interesting floristic element in Louisiana, and this is one of our few local references. Reviewed by G.D. Rowley, Taxon 20:175-176; "... professional 44 botanists are left baffled by the almost complete omission of literature citations throughout ... one looks in vain for a bibliography ..." Of more than amateur stature nonetheless. 1970. Wherry, E.T. Notes on phloxes in the Gulf States. Castanea 35:198-199. [325] 1. Phlox philosa subsp. fulgida given official status in flora of Louisiana. Earlier, unidentified Louisiana material now falls into this taxon. 2. The source of Phlox divaricata subsp. lamphamii cultivar "Opelousas," offered for sale in southwestern Louisiana, cannot be traced back to the wild in the state and remains a mystery. 1970. Wright, L.D., F.J. Swaye, and J.M. Coleman. Effects of Hurricane Camille [326] on the landscape of the Breton-Chandeleur Island chain [St. Bernard Parish] and the eastern portion of the lower Mississippi Delta. La. State Univ., Coastal Stud. Bull. 4:13-34. Damage to marsh vegetation, pp. 26-30. 1971. Bamforth, S.S. Microbial variations in the semi-tropical soils of St. Mary [327] Parish, Louisiana. Proc. La. Acad. Sci. 33:7-12. "1970." 1971. Bamforth, S.S. Floristic patterns of major groups of soil algae in Louisiana. [328] Proc. La. Acad. Sci. 34:7-11. Extending earlier studies (1%7, '68, '69, '70) to construct a floristic pattern for the state. Synopsis of Lytle 's six major soil regions of Louisiana ( 1968) and their characteristic vegetation. A similar study in Russia yielded comparable algal patterns. 1971. Barrett, E.R. Morchella in Louisiana. Castanea 36:147. [329] M. angusticeps Pk. and M. esculenta (L.) Pers. 1971. Choudhary, M.C. Nitella iyengarii, a new species from the southeastern [330] United States. Castanea 36:209-213. Type specimen collected by John W. Thieret from bottom of "the pool," Lacassine Wildlife Refuge, Cameron Parish. 1971. Guerke, W.R. Notes on Hepaticae from Louisiana and New Mexico. Bryol- [331] ogist 74:202-204. Four new state reports. Literature Cited: for "Notes on Louisiana Hepaticae" read "Southern Hepaticae" - the article contains no reference to our area. 1971. Kiger, R.W. Arthraxon hispidus (Gramineae) in the United States: taxonom- [332] ic and floristic status. Rhodora 73:39-46. Range clarification of Louisiana supercedes Agnes Chase's 1950 revision of Hitchcock's Manual. 45 1971. Krai, R. A treatment of Abildgaardia, Bulbostylis and Fimbristylis (Cypera- [333] ceae) for North America. Sida 4:57-227. Fifty odd tropical to warm temperate taxa, of which at least a dozen are in the state. Many chromosome counts taken from Louisiana specimens. Comprehensive monograph includes maps and fine, detailed illustrations (whole plant, spikelet, achene, leaf sheath). Field and herbarium study, LAF consulted. 1971. Leithead, H.L.,L.L. Yarlett andT.N. Shiflet. 100 native forage grasses in 11 [334] southern states. U.S. Dept. Agr., Soil Conserv. Serv., Agr. Handbook No. 389. 216 p. For each species: description, growth characteristics, distribution, site adaptation, use and management, range map and illustrations which include ligule for identification of sterile material. Bound with glue, the leaves caducous. 1971. Nelson, B. Caroline Dormon [1888-1971]. La. Soc. Hort. Res. J. 12:42-44. [335] Miss Dormon's extensive knowledge of the botany and natural history of her native Louisiana had earned her a reputation as an outstanding naturalist. A world somewhat queasy with overspecialization will greatly miss her strengths. 1971. * Paden, P. , P. Paden, and C.J. Felix. A study of lower and middle Cretaceous [336] spores and pollen from the southeastern United States: II. Pollen. Pollen Spores 13(3):447-473. Sixty-five core samples from southwestern Mississippi and northern Louisiana in which atten- tion was devoted to the lower Tuscalossa Formation. Twenty-four genera containing 33 species described and illustrated, several of both ranks new. Biol. Abst. 1971. Thieret, J.W. Additions to the Louisiana flora. Castanea 36:219-222. [337] A further contribution to the Louisiana Flora Project begun in 1%2. Eighteen new species and varieties, five of which are grasses, mostly from Thieret's own collections. 1971. Thieret, J.W. Eriocaulon cinereum R. Br. in Louisiana. Southwestern Nat. [338] 15:391. 1971. Thieret, J.W. Physalis lagascae (Solanaceae) in Louisiana: new to the conter- [339] minous United States. Sida 4:277. 1971. Thieret, J.W. Quadrat study of a bottomland forest in St. Martin Parish, [340] Louisiana. Castanea 36: 174-181. "Intended to augment our meager knowledge of the plant cover of the state. " This Mississippi River flood plain community dominated by Celtis laevigata, with green ash, sweetgum, and baldcypress subordinate. Three tables. 46 1971. Thomas, R.D. Burmanniaceae in Louisiana. Castanea 35:329-330. "1970." [341] Author, curator of the herbarium at Northeast Louisiana University in Monroe, cites his collections of the three native species, all apparently rare in the state: Apteria aphylla (Nutt.) Barnhart, Burmannia bifloa L., B. capitata (Walt.) Mart. 1971. Thomas, R.D. Collecting vascular plants in the habitat near the ground - or, [342] locating and collecting "Belly Plants." Castanea 36:148-149. All author's belly-work done in Louisiana. Tiny plants may be much more common than extant herbarium specimens indicate: Ophioglossum crotalophoroides , previously considered rare in the state, now documented from twenty-six parishes and probably as common as bracken. Other under-two-inchers include Lepuropetalon spathulatum and Tillaea aquatica. 1972. Baker, L. and H. Beckert. A partial bibliography of the ecology and biology [343] of the coast of the Gulf of Mexico with emphasis on the Louisiana coast. Univ. Southwest. La. Res. Ser. No. 15, Biol. 108 p. Indexes include "Ecology," "Microorganisms," "Vegetation," "Miscellaneous," and "Bibli- ographies." 983 entries. 1972. Batson, W.T. A guide to the genera of native and commonly introduced ferns [344] and seed plants of the southeastern United States, excluding penmsula Florida. Columbia, South Carolina. 151 p. The author fails to delimit his "southeastern United States" though the cover map shows only the Atlantic coastal states. Brief descriptions and little leaf-and-flower sketches accompany each genus. Keys to families and genera. 1972. Brown, C. A. Wildflowers of Louisiana and adjoining states. Baton Rouge. [345] 247 p. Currently the handiest picture book on wiidflower identification in our area. Thirty pages of introduction include crash courses in floral morphology, plant taxonomy, ecology, geography, and a useful color map of the parishes and vegetation regions of Louisiana. Color photographs illustrate some 400 species, including the more spectacular trees and shrubs. Inconsistent registry quality - Spiranthes, Sisyrinchium, Pyrus. Brief descriptions, range citations, stingy condiments. A few errors have come to my attention: Asdepias viridiflora Raf., p. 145, is a misinterpretation of the species illustrated. The correct name is A. obovata Ell. Pictures of Liatris squarrosa (L.) Michx. and L. pycnostachy Michx., pp. 210 & 211, have been trans- posed. And the name Ratibida peduncularis (T. & G.) Bamh. should probably replace R. columnaris (Sims) Don. a more western and northern species not elsewhere reported for Louisiana (see item 280). The picture and description agree with both. And Salvia lyrata is missing! Review by J. Ewan in Quart. Rev. Biol. 49:75-76. 1972. Chabreck, R.H. Vegetation, water and soil characteristics of the Louisiana [346] coastal region. La. State Univ., Agr. Exp. Sta., Bull. No. 664. 72 p. An exhaustive study with much valuable tabular information. Bibliography includes several unpublished Ph.D. dissertations of interest. Paperback, sewn in signatures, well designed and printed - an outstanding contribution in a field flooded with literature. 47 1972. Correll, D.S. and H.B. Correll. Aquatic and wetland plants of the southwest- [347] ern United States. U.S. Environmental Protection Agency. 1777 p. Reissued in two volumes in 1975 by Stanford University Press. Range includes the state of Texas to the Louisiana line. Thus, with Thieret's checklist of Louisiana aquatic and marsh plants (1972) as a companion, this is an invaluable contribution to the botany of our state. Full species descriptions are supplemented with full page drawings, many from other publications. The reader will find the preface to be a most eloquent - and thus controversial - plea for environmental protection. 1972. Lieux, M.H. A melissopalynological study of 54 Louisiana (U.S.A.) honeys. [348] Rev. Palaeobot. Palynol. 13:95-124. Pollen analyses of 54 commercial Louisiana honeys: 58 different pollen types identified. Trifolium repens, Rubus, Berchemia scandens, and Salix were high scorers. Biol. Abst. 1972. Lorio, P.L., Jr., V.K. Howe, and C.N. Martin. Loblolly pine rooting varies [349] with microrelief on wet sites. Ecology 53:1134-1140. A tree whose star is rising in the southern pinelands. Additional data towards an understanding of the dynamics of moisture stress and pine distribution. Tables & figures. 1972. Montz, G.H. A seasonal study of the vegetation on levees. Castanea 37: 140- [350] 146. East bank, St. Charles Parish. Vegetation sampled by 55 line transects at three seasons. Levees host a rich, mostly introduced, weedy flora. 1972.* Murphy, P.K. and R.E. Noble. The monthly availability and use of browse [351] plants by deer on a bottomland hardwood area in Tensas Parish, Louisiana. Proc. Ann. Conf. Southeast. Assoc. Game and Fish Comm. 26:29-57. 1972. Newton, M.B., Jr. Atlas of Louisiana, a guide for students. La. State Univ., [352] Misc. Publ. 72-1. 196 p. Soils and vegetation, pp. 32-44; climate and weather; land forms. 1972. Reese, W.D. List of the mosses of Louisiana. Bryologist 75:290-298. [353] Based on literature reports and herbarium specimens, the moss flora of Louisiana is character- ized as a diminished eastern North American flora enriched by distinct tropical American and coastal plain elements. 240 species and varieties, 90 genera, 39 families. Fissidens neonii is endemic to the state. Bibliography composed to complement that of Koch's 1956 paper - repetitions are minimal and the weight is on reports published later than 1956. 1972. Richard, M.A. Hardy palms for south Louisiana. La. Soc. Hort. Res. J. [354] 13:62-83. Thirteen palms are recommended, described and illustrated. 48 1972. Smith, C.A., Jr. A preliminary survey of the cultivated plants of Ouachita [355] Parish, Louisiana. Northeast La. Univ., M.S. Thesis. Unpubl. Representatives of 1 19 fcimilies, 385 genera, and 658 sf>ecies and varieties were collected from cultivation or found to be on deposit at NLU. This is such an industrious contribution to a field so poorly investigated in Louisicina, that I break a general policy to include it. Needless to say, many meaningful but unpublished theses and dissertations go unnoted in this bibliography. They are often retrievable through the references cited in related, pubhshed articles. 1972. Stuckey, R.L. Taxonomy and distribution of the genus Rorippa (Cruciferae) [356] in North America. Sida 4:279-430. Five species in Louisiana. Nasturtium micropetalum and N. obtusum, types from near New Orleans, synonomous with R. teres. Illustrations limited. 1972. Thieret, J.W. Aquatic and marsh plants of Louisiana: a checklist. La. Soc. [357] Hort. Res. J. 13:1-45. Vascular plants only, with some synonomy. 1972. Thieret, J.W. Checklist of the vascular flora of Louisiana. Part I. Ferns and [358] fern allies, gymnosperms, and monocotyledons. Lafayette Nat. Hist. Mus., Tech. Bull. 2. 48 p. "The policy is simple: no voucher, no listing," but herbaria possessing vouchers not cited. Synonomy included. 728 sjjecies of monocots, 8 gymnosperms and 63 ferns and allies. 1972. Thieret, J.W. Rotala indica (Lythraceae) in Louisiana. Sida 5:45. [359] 1972. Thieret, J.W. The yam sham. Lafayette Nat. Hist. Mus. Newsletter, March. [360] Yams versus sweet potatoes, botanically speaking, and mention of our ornamental air-potato, Dioscorea bulhifera. Chatty one page mimeo. 1972. Thieret, J.W. Zeuxine strateiimatica in Louisiana. Amer. Orchid Soc. Bull. [361] 41:413. An asiatic, weedy orchid that has already spread through much of Florida. Delta National Wildlife Refuge, Plaquemines Parish; new to the state. 1972. Thomas, R.D. Botrychium lunarioides, Ophioglossum crotalophoroides, and [362] Ophioglossum engelmanni in a Louisiana cemetery. Southwestern Nat. 16: 431-432. 1972. Thomas, R.D. Eulophia ecristata (Fernald) Ames (Orchidaceae) in Grant 13631 Parish, Louisiana. Southwestern Nat. 16:431. 49 1972. Wilson, B.H. and M.H. Lieux. Pollen grains in the guts of field collected [364] tabanids in Louisiana. Ann. Entomol. Soc. Amer. 65:1264-1266. Alluvial and salt marsh areas collected. Oak, grass, and composite pollen most abundant, the former two types windbome. 1973. Allen, CM. Additions to the Louisiana flora. Sida 5:130-131. [365] Four new species including Corallorhiza odontorhiza from St. Helena Parish. See also Sida 5:287 (1974) for more information on the same collection of coral-root. 1973. Curry, M.G. and CM. Allen. Alismataceae of Louisiana. Proc. La. Acad. [366] Sci. 36:88-91. Taxonomy of Echinodorus and Sagittaria. Key to species plus distribution maps and illustra- tions. 1973. Feibleman, P.S. and the editors of Time-Life Books. The bayous. New York. [367] 184 p. Photography! And another popular book with bayou flavor is Bayou Backwaters by Allan W. Eckert (1968, New York, 155 p.), nature writing with strictly animal adventures: canebrake rattler vs. spotted skunk, alligator vs. armadillo. 1973. Flock, J.M. Guide to the botanical contributions of Lloyd H. Shinners (1918- [368] 1971). Sida 5:137-179. Many first reports from Louisiana by this indefatigable Texas botanist. 1973. Grelen, H.E. and W.F. Mann, Jr. Distribution oi Senna seymeria (Seymeria [369] cassioides) - a root parasite on southern pines. Econ. Botan. 27:339-342. The plant is an obligate parasite on at least three species of southern pines and is apparently selective for pines only. A southwide survey indicated that it is most common on moist, sandy sites of the lower Gulf and Atlantic Coastal Plains. Punch, 1962 (1974). 1973. Guerke, W.R. Epiphyllous Hepaticae in Louisiana. Bryologist 76:201-202. [370] The first report of this common tropical phenomenon in Louisiana. All of the 7 species listed were collected on Magnolia grandiflora. 1973.* Gunn, CR. and J.V. Dennis. Tropical and temperate stranded seeds and [371] fruits from the Gulf of Mexico. Contrib. Mar. Sci. 17:111-121. Beaches along the western coast of Louisiana receive large numbers of tropical disseminules that have been carried by currents from the Caribbean region, northern South America, and Mexico. Disseminules of 34 tropical and 24 temperate species have been collected on northern and western Gulf beaches. Biol. Abst. 1973.* Halls, L.K. Delicacy for the birds. La. Conserv. 25(3 & 4): 26-29. [372] Seed of spurges (Euphorbiaceae) is valuable for birds. Flowers of several species are pictured in color. Punch. 1962 (1974). 50 1973. Landry, G. and J. W. Thieret. Iso'etes louisianensis (IsoStaceae), a new species [373] from Louisiana. Sida 5:129-130. Holotype from Washington Parish. The only Isoetes previously collected in Louisiana is /. melanopoda, considered rare in the state. 1973. Lasseigne, A. Louisiana legumes. Univ. Southwest. La., Southwest. Stud. [374] Ser.,No. 1.255p. Comprehensive treatment of the family in Louisiana, in which 166 species are recognized. Generic descriptions, keys to species, habitat, dates of flowering. Line drawings partially supplant species descriptions. Range maps to parish. Perfect binding, not in signatures, a misfortune. Muhilithed. 1973. Mahler, W.F. By any other name. Sida 5:180-181. [375] Louisiana botany? A Louisiana botanist, Joseph Ewan, engages Lloyd Shinners in a duel of poems: "Sat Down Be Sida" is parried by "Lines Penned in Retaliation." 1973. Montz, G.M. Psilotum in New Orleans. Castanea 38:207-208. [376] Psilotum nudum collected on University of New Orleans (nee LSU in N.O.) campus; first report from Orleans Parish. Review of previous state citations and collections. (During the 1976 AIBS meeting, several plants were discovered on Tulane campus under a row of azalea bushes.) 1973. Parks, J.C. A revision of North American and Caribbean Melanthera (Com- [377] positae). Rhodora 75:169-210. M. nivea, the only U.S. sp)ecies outside peninsula Florida, is a Louisiana native, and the type of M. carpenleri Small, a synonym, was collected in West Feliciana Parish. 1973. Pecora, R. A. A report on the algal flora of southwestern Louisiana: Phyto- [378] flagellates. Proc. La. Acad. Sci. 36:76-82. List of 34 flagellates, of which 13 are reported for the first time in the state; annotations and more or less brief descriptions. 1973. Solymosy, S.L. Hydrilla verticillata (L. f.) Casp., a new plant for Louisiana. [379] La. Soc. Hort. Res. J. 14:92. 1973. TTiomas, R.D. Dirca palustris (Thymelaeaceae): new to Louisiana. Sida [380] 5:132. 1973. Thomas, R.D., W.H. Wagner, Jr., and M.R. Mesler. Log fern {Dryopteris [381] celsa) and related species in Louisiana. Castanea 38:269-274. Update of Brown and Correll's work of 1942. Morehouse Parish. 51 1974. Allen, CM. Nineteen species of grasses (Poaceae) new to Louisiana. Proc. [382] La. Acad. Sci. 37:18-20. Field and herbarium study. 1974. Allen, CM. Some unusual grasses (Poaceae) of Louisiana. [Abstract.] Proc. [383] La. Acad. Sci. 37:130. Recently discovered hybrid: Sphenopholis obtusata X 5. pensylvanica. Volumes 37 and 38 of the Proceedings include abstracts of papers presented at the 1974 and 1975 Annual Meetings of the Louisiana Academy. Several are here included primarily to link certain workers with certain topics. 1974. Bamforth, S.S. Coastal plankton off Grand Isle [Jefferson Parish], Louisiana. [384] Proc. La. Acad. Sci. 37:64-69. Collections from Freeport Sulphur Company Grand Isle Platform west of the delta. Annotated list and one figure correlating major groups with temperatures and salinity. 1974. Cain, R.H. Pimple mounds: a new viewpoint. Ecology 55:178-182. [385] The character and distribution of pimple mounds in Louisiana and adjoining territory points to a possible pedestal tree-erosion origin. Literature cited brings out of obscurity several related publications and theses. 1974. Delcourt, H.R. and P. A. Delcourt. Primeval magnolia-holly-beech climax in [386] Louisiana. Ecology 55:638-644. Reconstruction of early settlement forest patterns in West Feliciana Parish from an American Land Office Survey of 1821. This study represents the first quantitative evidence to support the existence of a late-successional, or climax, magnolia-beech-holly forest in the original vegeta- tion of upland mesic sites of the Gulf Coastal Plain. Two tables, one parish map of forest types. 1974.* Duffy, LT. Some like it wet. Amer. Forestry 80(6): 32-35. [387] Hardwoods native to the lower Mississippi Valley generally survived the 1973 flood without serious damage. Newly planted stands as well as older trees of species not naturally found on bottomland sites, were frequently lost. Punch, 1962 (1975). 1974. Fruge, D.W. The vegetation of Lacassine Pool, Lacassine National Wildlife [388] Refuge [Cameron Parish], Louisiana. U.S. Dept. Interior, Fish and Wildlife Serv., Lafayette. Mimeo. Two major communities analyzed, emergent and floating-leaved/submergent. Checklist of 138 spp. 1974. Gonsoulin, G.J. A revision of Styrax (Styracaceae) in North America, Cen- [389] tral America, and the Caribbean. Sida 5:191-258. 5. americana and 5. grandifolia in the state, the history of the former going back to Robin's Voyages (1807). Specimens from LAF utilized. 1974. Gregory, P.E. and D.R. Sanders, Sr. Some aspects of the life history and [390] ecology of Cabomba caroliniana Gray. [Abstract.] Proc. La. Acad. Sci. 37:131. 1974. Grelen, H.E. Pinehill bluestem, Andropogon scoparius var. divergens An- [391] derss. ex Hack., an anomaly of the A. scoparius complex. Amer. Midi. Nat. 91:438-444. Review of published accounts of the relation of var. divergens to other taxa, together with morphological investigations of transplanted clones, indicates that separation of southern forms of A. scoparius is taxonomically imprudent. Punch, 1962 (1975). More on the same appeared in 7. Range Mgmt. 27:283-284 (1974). 1974. Jones, S.B., Jr. Mississippi Flora. I. Monocotyledon families with aquatic or [392] wetland species. Gulf Res. Repts. 4:357-379. IV. Dicotyledon families with aquatic or wetland species, ibid, 5:1-22 (1975). Keys, distribution maps (in Miss), habitats, references, nomenclature, and notes are given for some 28 families (grasses, sedges, rushes excluded) many of whose members spill over into Louisiana. 1974. * Kapraun, D.F. Seasonal periodicity and spatial distribution of benthic marine [393] algae in Louisiana. Contrib. Mar. Sci. 18:139-167. A broad investigation providing information on physiology, taxonomy, floristic affinities, and life histories. A list of 85 benthic marine algae including 31 new records for Louisiana is given. Biol. Abst. 1974. Luke, W.J., C.A. Schexnayder, and C.A. Brown. Establishment of Tilland- [394] sia recurvata L. in East Baton Rouge Parish, Louisiana. Proc. La. Acad. Sci. 37:17. Seconding Thieret's provisional inclusion of ball-moss (1%9) in the state flora based on a 1939 Viosca collection from parts unknown. The plant is established on an ornamental yaupon on the LSU campus, and the groundskeeper has agreed to go easy. 1974. Montz, G.M. Linaria vulgaris Hill in Louisiana. Castanea 39:190. [395] 1974. Montz, G.M., and A. Cherubini. An ecological study of a bald-cypress [396] swamp in St. Charles Parish, Louisiana. Castanea 38:378-386. "1973." Quadrat sampling: DBH and frequency-density of cypress. Of particular interest are a compar- ison of township maps of 1860 to quadrangle maps of 1%9 in study of plant succession, documenting baldcypress' extension into adjacent marsh, and discussion of a change from lower to higher salinities in the Mississippi River-Gulf outlet area and Lake Pontchartrain, to the detriment of baldcypress and doubtless many other plants. 53 1974. Oefinger, S.W., Jr. and L.K. Halls. Identifying woody plants valuable to [397] wildlife in southern forests. U.S. Forest Serv., Southern Forest Exp. Sta., Res. Pap. SO-92. 76 p. Twigs, buds, and other key identification features are illustrated in color for 70 browse species common to pine-hardwood forests of the South. Punch, 1962 (1975). 1974. Pecora, R.A. Reports on the algal flora of southwestern Louisiana. II. [398] Collections of Chlorophyceae and Xanthophyceae. Proc. La. Acad. Sci. 37:50-54. Eighteen of the 19 species reported for the first time in the state. 974. Piehl, M. A. Preliminary reports on the flora of Louisiana. No. 1. The families [399] Magnoliaceae, Illiciaceae, and Schisandraceae. [Abstract.] Proc. La. Acad. Sci. 37:13L 1974. Piehl, M.A. Rare or endangered higher plants of Louisiana - a progress [400] report. [Abstract.] Proc. La. Acad. Sci. 37:131. Comprehensive state list has been prepared. 1974. Piehl, M.A. Verification of the occurrence of Tillandsia recurvata, ball-moss, [401] in Louisiana and its endangered status. La. Soc. Hort. Res. J. 15:1-6. Locally abundant on L.S.U. campus! See item 394. 1974. Piehl, M.A. and M.A. Spindler. Preliminary reports on the flora of Louisi- [402] ana. No. 4. The family Lauraceae. [Abstract.] Proc. La. Acad. Sci. 37:131. Range of Lindera benzoin (L.) Bl. considerably wider than an early report by Cocks suggests. 1974. Piehl, M.A. and P. A. Piehl. Preliminary reports on the flora of Louisiana. [403] No. 2. The families Aristolochiaceae, Celastraceae, and Menispermaceae. [Abstract.] Proc. La. Acad. Sci. 37:131. 1974. Piehl, P. A. and M.A. Piehl. Vines of Louisiana, part I. Louisiana lianas, a [404] provisional annotated list of native, naturalized and cultivated woody vines. La. Soc. Hort. Res. J. 15:51-60. 35 families, 60 genera and 128 species. 1974. Reese, W.D. Noteworthy Louisiana mosses. [Abstract.] Proc. La. Acad. Sci. [405] 37:131. 54 1974. Sanders, D.R., Sr. Notes on the appearance of a new aquatic plant in [406] Louisiana. [Abstract.] Proc. La. Acad. Sci. 37:132. Hydrilla verticillata Royle, potentially a dangerous introduction into Louisiana waters. Soly- mosy, however, reported it in 1973. 1974. Sanders, D.R., Sr. and J.H. Mannings. Effects of water fluctuation on the [407] vegetation of Black Lake. [Abstract.] Proc. La. Acad. Sci. 37:132. Effective and, according to authors, ecologically sound method of removing unwanted aquatic plants. 1974. Short, M.S. and M.A. Piehl. Preliminary reports on the flora of Louisiana. [408] No. 3. The family Annonaceae. [Abstract.] Proc. La. Acad. Sci. 37:132. 1974. Spindler, M.A. and R.E. Noble. Fall vegetation on the spoil banks of Superi- [409] or Canal, Cameron Parish, Louisiana. Proc. La. Acad. Sci. 37:74-88. An artificial community that is increasing in a state where a six inch change in elevation can significantly alter the plant life. 1974. Thieret, J.W. and CM. Allen. Setaria pallide-fusca (Gramineae) in Louisi- [410] ana. Castanea 39:290-291. This LSU campus weed proves new to state, to conterminous U.S. and to North America. 1974. Valentine, J.M., Jr. Unpublished administrative reports. U.S. Dept. Interi- [411] or. Fish and Wildlife Serv., Lafayette. 5 p. Mimeo. Titles from 1960 to 1974, mostly grazing and succession studies in local wildlife refuges. No authors. 1975. Allen, CM. Trichomanes petersii in Louisiana. [Abstract.] Proc. La. Acad. [412] Sci. 38:121. 1975. Delcourt, H.R. and P.A. Delcourt. The blufflands: Pleistocene pathway into [413] the Tunica Hills [West Feliciana Parish]. Amer. Midi. Nat. 94:385-400. Numerous plant species of a northern mixed mesic forest reach their southern terminus in the hilly land above the eastern escarpment of the Mississippi River alluvial valley known as the Tunica Hills. The epic of several of these Pleistocene migrants and relicts is here explored in full. 1975. Duncan, W.H. and L.E. Foote. Wildflowers of the southeastern United [414] States. Athens, Georgia. 296 p. 55 In his foreword, ex-govemor of Georgia, Jimmy Carter, praises this reference aa "both comprehensive and beautiful. " Indeed it expands the concept of a popular wildflower book by its conviction that even the amateur botanist deserves the most accurate and thorough botany. Range includes Louisiana. Illustrated with color photographs. Reviewed by Earl L. Core, Castanea 40:364 (1975) and by J. Ewan, Ecology 58:222-223 (1977). 1975. Elliott, P.B. and S.S. Bamforth. Interstitial protozoa and algae of Louisiana [415] salt marshes. J. Protozool. 22:514-519. Sediments of stagnant pools among Spartina sp. and Distichlis spicata constitute a "sulfide biome" in which the abundance of nutrients provides habitable conditions for all the major groups of microflora. Excellent figures. 1975. Gould, E. and J. Ewan. Phytogeographic and ecologic relationships of the [416] flora of Breton Island [St. Bernard Parish], Louisiana. Tulane Stud. Zool. Botan. 19:26-36. Island 3 miles long by V2 mile wide. Salt spray community of 84 species of vascular plants and 3 mosses, predominantly of temperate origin, studied in light of records published 60 years previously. 1975. Graham, S.A. Taxonomy of the Lythraceae in the southeastern United [417] States. Sida 6:80-103. Cuphea glutinosa "first collected in the United States in 1884 in Vermilion Parish, Louisiana." Chromosome count based on Thieret collection from Lafayette Parish. Further information on Rotala indica (see Thieret, 1972). All seven genera treated are in our state. Material from LAF consulted. Illustrations and maps wanting. 1975. Guerke, W.R. A floristic study of the Hepaticae and Anthocerotae of the [418] Florida Parishes. Bryologist 77:593-600. Physical description and vegetation of the study area, floristic comparisons and phytogeogra- phy, as well as a Ust of 83 species in 24 families. 1975.* Johnson, R.L. Natural regeneration and development of Nuttall oak and [419] associated species. U.S. Forest Serv., Southern Forest Exp. Sta., Res. Pap. SO-104. 12 p. Nuttall oaks from a dense seedling catch survived for 5 to 10 years in complete shade and for 15 years with 1 or 2 hours of daily sunlight. They made little growth, but they responded well when released after periods of 1 to 9 years. Green ash, water hickory, and sugarberry survived equally well and outgrew the oaks after the overstory was removed. Punch, 1962 (1975). 1975. Johnston, L. A. Revision of the Rhamnus serrata complex. Sida: 6:67-79. [420] Louisiana un-botany: R. lanceolata "is unknown in Louisiana" though collections are mapped from neighboring east Texas. 56 1975. Jones, R.E. Louisiana Tech University Herbarium: new national plant de- [421] pository forU.S. Corps of Engineers. [Abstract.] Proc. La. Acad. Sci. 38:121. Louisiana Tech chosen as the voucher specimen dep)ository for Corps of Engineers' new Environmental Effects Laboratory Waterways Experiment Station, Vicksburg, Mississippi. 1975. Landry, G.P. Unusual pteridophytes of Louisiana. [Abstract.] Proc. La. [422] Acad. Sci. 38:121. 1975. Lieux, M.H. Dominant pollen types recovered from commercial Louisiana [423] honeys. Econ. Botan. 29:87-96. Same honey samples as Lieux, 1972, and some of the same information. Additionally, electron micrographs and morphological descriptions of pollen grains. Two maps. 1975. Marx, P.S. and R.D. Thomas. A survey of the vascular plants of Caldwell [424] Parish, Louisiana. Proc. La. Acad. Sci. 38:75-85, 924 species. Area near Horseshoe Lake north of Columbia is the only Louisiana site for Dirca palustris (see Thomas, 1973). Two others new to the state: Quercus oglethorpensis and Zan- thoxylum americanum. 1975. Massey, J.R. Fatoua villosa (Moraceae): additional notes on distribution in [425] the southeastern United States. Sida 6:116. See also, Thieret, 1964. 1975. Montz, G.N. The submerged vegetation of Lake Pontchartrain, Louisiana. [426] U.S. Army Corps of Engineers, New Orleans District, Environ. Qual. Sect. 17 p. Mimeo. Vallisneria americana, Ruppia maritima, and Najas guadalupensis most abundant. Distribution maps. 1975. Noble, R.E. and P.K. Murphy. Short term effects of prolonged backwater [427] flooding on understory vegetation. Castanea 40:228-238. Mississippi River bottomland hardwood forest in Tensas Parish sampled before and after flood. Elimination of six taxa, in contrast to the vegetation's general resilience, carries implications on range and tolerance of species in an unstable environment. 1975. Pecora, R.A. Rediscovery of Oscillatoria ludoviciana Flint and Prescott. [428] [Abstract.] Proc. La. Acad. Sci. 38:121. 1975.* Sand, N.H. and P. Koch. Creole carpentry in 1800. Forests and People [429] 25(3): 16-19, 38-45. 57 The Kent Plantation House, in Alexandria (Rapides Parish), Louisiana, exemplifies construc- tion practices in which reliance was almost exclusively on local materials. Though French tradition and culture predominated m Louisiana, carpenter's tools of English design seem to have been preferred. Punch, 1962 (1976). See also where La. compared with St. Louis in scholarly paper by Charles E. Paterson, "Houses of French St. Louis," in J.F. McDermott, French in Mississippi Valley, 1%5, 17-40, illus. 1975. Somers, P. and W.R. Buck. Selaginella ludoviciana, S. apoda and their [430] hybrids in the southeastern United States. Amer. Fern J. 65:76-82. Hybrids are rare but have been reported for Florida and Louisiana. 1975. SulHvan, V.I. Wind pollination in Eupatorium (Compositae). [Abstract.] [431] Proc. La. Acad. Sci. 38:121-122. 1975 Thieret, J.W. Hemigraphis reptans (Acanthaceae), a greenhouse weed in [432] Louisiana. Sida 6: 115. 1975. Thomas, R.D. and R. Overby. A preliminary survey of the vascular flora of [433] Caddo Parish, Louisiana. [Abstract.] Proc. La. Acad. Sci. 38:122. 58 INDEX TO AUTHORS Abbott, E.V. 1926 Aheam, Donald G. 1970 Ahles, Harry E. 1964 Albrecht, Joseph 1888 Alexander, Edward J. 1931 Allen, Charles M. 1973, 1974, 1975 Anderson, Loran C. 1970 Anderson, Lewis E. 1961, 1964 Anonymous 1900, 1955, 1969 Baker, Lynn 1972 Bamforth, Stuart S. 1967, 1968, 1%9, 1971, 1974, 1975 Bandoni, Robert J. 1957 Banks, Donald J. 1966 Barrett, Elton R. 1964, 1971 Batson, WadeT. 1972 Beckert, Heino 1972 Beeson, K.C. 1960 Bell, C. Ritchie 1964, 1967 Benjamin, Richard K. 1955 Bick, George H. 1954, 1959 Bicknell, Eugene 1899 Blair, Robert M. 1960, 1969 Blake, Sidney F. 1956 Blomquist, HugoL. 1953 Bodman, Mary C. , Sr. 1949 Bombard, Miriam L. 1952 Boudreaux, Bruce H. 1951, 1967 Breen, RuthS. 1956 Brown, Clair A. 1935, [1936, 1937], 1942, 1943, 1946, 1951, 1952, 1954, 1956, 1957, 1972, 1974 Buck, William R. 1975 Cain, Robert H. 1974 Cardot, Jules 1888, 1896 Carter, W.H. 1946 Cassady, J.T. 1952 Chabreck, Robert H. 1958, 1962, 1968, 1972 Chamberlain, James L. 1959 Chamberlain, T.K. 1947 Chapman, AlvinW. 1878 Cherubini, Arthur 1974 Chipman, Robert K. 1961 Choudhary, M.C. 1971 Clark, Lois 1937, 1958 Coleman, James M. 1970 Cook, Melville T. 1945, [1947, 1949], 1951, [1952, 1953] Cook, Tressie 1953 Cooke, William B. 1965 Correll, Donovan S. 1970, 1972 Correll, Helen B. 1972 Crum, Howard 1961, 1964 Curry, Mary G. 1973 Darnell, Rezneat M. 1959, 1%1 Daubs, Edwin H. 1962 Dean, Blanche E. 1961 Deiler, Frederick G. 1957 Delahoussaye, A. James 1967 de las Barras y de Aragon, Francisco 1948 Delcourt, Hazel R. 1974, 1975 Delcourt, Paul A. 1974, 1975 de Lesdain, Bouly 1933 Dennis, J. V. 1973 Depoe, Charles E. 1969 Dike, David H. 1969 Dillon, O.W. 1957 Dirskell, B.N. 1954 Dix, W.L. 1945 Dormon, Caroline 1958, 1965 Drechsler, Charles 1954, 1955 Dress, William J. 1969 Duffy, LT. 1974 Dukes, George H. , Jr. 1961 Duncan, Wilbur H. 1953, 1967, 1975 Duvall, Vinson L. 1966 Earle, Thomas T. 1939 Eggler, Willis A. 1955, 1961 Egler, Frank E. 1961 Eleuterius, Lionel N. 1969 Elliott, PriscillaB. 1975 Ensminger, Allen 1957, 1959 Epps, E.A.,Jr. 1969 Ewan, Joseph 1963, 1965, 1967, 1968, 1969, 1975 Exner, Beatrice 1953 Eyles, Don E. 1944 Fassett, Norman C. 1951 Feibleman, Peter S. 1973 Felix, Charles J. 1971 Flint, Lewis H. 1951, 1953, 1954 Flook, Jerry M. 1973 Foote, Leonard E. 1975 Foote, LucyB. 1942 Freeman, John D. 1969 Fruge, David W. 1974 Frye,T.C. 1937 Gagliano, Sherwood M. 1964 Glasgow, Leslie L. 1957, 1960, 1962, 1963 Gleason, Henry A. 1952, 1958 Gonsoulin, Gene J. 1974 Gould, Edwin 1961, 1975 Grafton, B.E. 1959 Graham, Shirley A. 1975 Greene, Wilhelmina F. 1953 Gregory, P.E. 1974 Grelen, Harold E. 1966, 1973, 1974 Grout, Abel J. 1940 Guerke, Wayne . 1971, 1973, 1975 59 Gunn, Charles R. 1973 Gunter, Gordon 1956 Halls, Lowell K. 1%1, 1973, 1974 Hansford, C.G. 1955 Hardin, James W. 1957, 1%7 Harmon, BudG. 1960 Harrar, Ellwood S. 1%2 Harrar, J. George 1%2 Harris, Van T. 1958, 1962 Haynes, Robert R. 1968, 1%9 Hilgard, Eugene W. 1873 Hill, H.L. 1959 Hodges, Charles S. 1%2 Hoffpauir, CM. 1%1 Horn, Norman L. 1952 Howe, H.V. 1935 Howe, Virgil K. 1972 Humm, Harold J. 1956, 1959 Hutchins, Ross E. 1968 Hutto, Joan 1%7 Huxley, Julian S. 1923 Ingram, John 1%9 Isely, Duane 1%9 James, Charles W. 1957 Jemison, E.S. 1%2 Jex, Gamett W. 1944 Joanen, T. 1968 Johnson, R.L. 1975 Johnston, Laveme A. 1975 Johnston, Marshall C. 1964, 1970 Jones, Raymond E. 1968, 1975 Jones, Samuel B., Jr. 1%9, 1974 Junca, Hugh A. 1%2 Jung, Rodney C. 1950 Kane, Harnett T. 1944 Kapraun, Donald F. 1974 Karling, John S. 1953, 1954, 1955, 1956, 1958 Kiger, Robert W. 1971 Kimble, Robert B. 1959 King, J. E. 1947 Kniffen, FredB. 1968 Koch, LeoF. 1956, 1957, 1%1 Koch, Peter 1975 Krai, Robert 1966, 1971 Kubota, Joe 1960 Kuprionis, John 1970 Lambremont, Edward N. 1954 Landry, Garrie P. 1973, 1975 Langdon,O.G. 1952 Lasseigne, Alex 1973 Lazar, V.A. 1960 Leithead, H.L. 1971 Lemaire, Robert J. 1%1 Lemmon, Betty E. 1%5, 1966 Lewis, Walter H. 1%2 Lieux, Meredith H. 1972, 1975 Linnartz, Norwin E. 1%3, 1%9 Livingstone, R., Jr. 1%5 Lloyd, Francis E. 1901, 1904 Logan, Lowell A. 1%3 Lorio, Peter L., Jr. 1972 Lowy, Bernard 1953, 1954, 1955, 1956, 1957, 1958, 1959, 1964, 1965 Luke, W.J. 1974 Lutes, Dallas D. 1966 Lynch, John J. 1947 Lytle, S.A. 1954, 1959, 1968 Mackaness, Faith Pennebaker 1942 Mackin,J.G. 1956 Mahler, William F. 1973 Maisenhelder, Louis C. 1958 Maitre, Reinhardt 1875 Mann, W.F.,Jr. 1973 Manning, J.H. 1974 Maples, Robert S., Jr. 1966 Martin, Carolyn N. 1972 Marx, Paul S. 1975 Massey, Jimmy R. 1975 McGinn, L.R. 1963 Mesler, Michael R. 1973 Meyers, Samuel P. 1970 Miles, P. 1970 Miller, Lillian W. 1969 Mitchell, Richard S. 1970 Mitra, Jyotirmay 1961 Montz, Glen N. 1972, 1973, 1974, 1975 Moore, John A. 1956 Moore, Walter G. 1950, 1961, 1966 Mrak, E.M. 1952 Mullahy, John H. 1959 Muller, Jean 1895 Murphy, Patrick K. 1972, 1975 Negus, Norman C. 1961 Nelson, Barbara 1971 Nelson, IraS. 1959, 1961, 1%2, 1967 Nelson, Thomas C. 1%9 Newton, Milton B., Jr. 1972 Nicholson, M.E. 1970 Noble, Robert E. 1972, 1974, 1975 Oefinger, S.W.,Jr. 1974 Olive, Lindsay S. 1948, 1951 Oliver, Abe D., Jr. 1969 Ohver, Royce L. 1962 Olson, Willis A. 1%9 Overby, R. 1975 Owens, A. G., Jr. 1%3 Ownbey, Gerald B. 1958, 1%9 Paden, P. 1971 Paden, P. 1971 Palmisano, A.W. 1968 Parks, James C. 1973 Pecora, Richard A. 1973, 1974, 1975 Penfound, WUliam T. 1952, 1956 Penn, George H. 1951, 1954, 1%2 60 Perdue, Robert E., Jr. 1962 Pessin, Louis J. 1928 Phaff, Herman J. 1952 Piehl, Martin A. 1974 Piehl, Paula A. 1974 Pinkava, Donald J. 1967 Plaisted, Robert L. 1967 Plakidas, AntoniosG. 1956, 1960 Pope, Thomas E. 1968 Prescott, Gerald W. 1942 Punch, Louis E. 1962 Pursell, Ronald A. 1956, 1958, 1970 Putnam, John A. 1956 Radford, Albert E. 1964, 1967 Randolph, Lowell F. 1%1, 1966, 1%7 Reese, William D. 1959, 1960, 1%1, 1964, 1%5, 1966, 1%7, 1%8, (1%9], 1970, 1972, 1974 Reizenstein, L. von 1863 Renauld, Ferdinand 1888, 1896 Rense, William C. 1970 Rhee, J.S. 1970 Rhodes, Donald G. 1970 Richard, Michael A. 1970, 1972 Richards, Edward L. 1%8 Richardson, Annie L. 1963, 1964 Riche, Sam 1%3 Rickett, Harold W. 1966 Ripley, Thomas H. 1961 Ritchie, Alexander 1959 Robbins, Herbert C. 1968 Robertson, J. Lynne, Jr. 1944 Robinson, Brittain B. 1947 Rock, Howard F.L. 1957 Rollins, Reed C. 1969 Rougeou, Clyde L. 1965 Russell, Norman H. 1%1 Russell, Richard J. 1935, 1942 Rylander, Michael K. 1%9 Sand, N.H. 1975 Sanders, D.R.,Sr. 1974 Schexnayder, C.A. 1974 Schuster, Rudolf M. 1956 Scott, Arthur M. 1950 Shanor, Leland 1952 Shiflet,T.N. 1963,1971 Shinners, Lloyd H. 1951, 1956, 1957, 1958, 1960, 1%2, 1%3, 1964 Short, M.S. 1974 Simmons, Emory G. 1%2 Small, John K. 1931 Smith, A. L. 1947 Smith, Carlos A., Jr. 1972 Smith, W.G. 1970 Solymosy, Sigmond L. 1%3, 1%5, 1966, 1973 Somers, Paul 1975 Speairs, Richard K., Jr. 1957 Spindler, Michael A. 1974 Stemitzke, Herbert S. 1956, 1965 Stroube, William H. 1954 Stuckey, Ronald L. 1966, 1972 Sullivan, Victoria 1. 1975 Svihla, Ruth D. 1939 Swaye, F.J. 1970 Taft, Clarence E. 1946 Taylor, William R. 1954 Thieret, John W. 1964, 1966, 1967, 1968, 1%9, 1970, 1971, 1972, 1973, 1974, 1975 Thomas, C.H. 1960 Thomas, R. Dale 1967, 1971, 1972, 1973, 1975 Thomas, William H. 1962 Thome, Robert F. 1954 Tucker, Shirley C. 1970 Valentine, Jacob M., Jr. 1974 Viosca, Percy, Jr., 1931, 1932, 1949 Vogel, E. 1%9 Wagner, Warren H., Jr. 1973 Ware, George H. 1956 Webert, Fred 1962 Welch, William C. 1%8 Welden, Arthur L. 1958, 1959, 1960 Weniger, Del 1970 Westling, Andrew W. 1%9 Wherry, Edgar T. 1955, 1957, 1970 Wilbur, Robert L. 1955, 1967 Williams, O.B. 1952 Wilson, B.H. 1972 Wilson, HoUis R. 1956 Wolff, S.E. 1954 Wood, Carroll E., Jr. 1958 Wright, L.D. 1970 Yarlett, L.L. 1971 Zillgitt, Walter M. 1969 INDEX TO BOTANICAL NAMES 61 A be lia lib Abildgaardia 333 Acacia 293 Acalypha 295 Acer 7,1^ Aesculus 132 Agaricales217 Alismataceae 366 Andropogon 240, 391 Annonaceae 408 Anthocerotae418 Apteria 341 Argemone 149 Aristolochiaceae 403 Arthraxon332 Ascomycetes 234 Astomum 235 A uricularia 73 Bacopa 287, 320 Baptisia 301 Barbella 250 Basidiomycetes 234 Batrachospermum 68 Berchemia 348 Berlandiera 263 Betulaceae 264 Bigelowia 309 fi/>'jca 307 Botrychium 362 Bowlesia 269 Bryophyta29, 126,315 Bulbostylis 333 Burmannia 269, 341 Burmanniaceae 341 Cabomba 390 Callitriche 53 Calyptocarpus 104 Caperon/a 21, 43 Carduus 151 Career 144 Carya 23 IB Celastraceae 403 Celastrus 306 Ce/r« 340 Cephaloziella 117 Cercospora 116, 166 Chlorophyceae 398 Cirsium 297 Cladonia 36 Clinopodium 229 Colletotrichum 60 Collinsonia 206 Conidiobolus 78 Corallorhiza 365 Crafaegu5 231 B Crep« 226 Cuphea All Cymatoderma 169 Cyperus 230, 259 Dacrymyces 84 Dermocystidium 1 12 Digitaria 89 Dioscorea 360 Diphyscium 279 D/rcfl 380, 424 Dm/c/i/« 415 Dolichos 301 Dopatrium 302 Drosera 201 Dryopteris 381 Dupratzia 102 Echinodorus 366 Epidendrum 20, 27 Epifagus 322 EriocauIaceae241 Eriocaulon 338 Euastrum 30 Eulophia 363 £wpator/Mw431 Eustoma 102 Euthamia 1 18 Exidia 135 Fatoufl 228, 425 Filariomyces 64 Fimbristylis 184, 333 Fissidens 171, 232, 246, 298, 353 Fontinalis 6 Forestiera 119 Fragaria 26 Fraxinus 1 1 1 Frullania 142 Fungi Imperfecti 234 Gasteromycetes 101 Gliocephalotrichum 234 Gonium 30 Gonytrichum 192 Gymnospermae 264, 358 Halophila 306 Helenium 136 Helicomina 43 Hemigraphis 432 Hepaticae 23, 24, 180, 331, 370, 418 Heterochaete 45 Homaromyces 95 Hottonia 262 Hydrilla 379, 406 Hymenocallis 57 Hyphomycetes 77 Hypochoeris 120 lUiciaceae 264, 399 Irenopsis 98 /m 16, 18, 65, 140, 161, 185, 194, 196, 249, 251A, 265, 292 Isariopsis 166 62 Isoetes 373 Juncus 242 Laboulbeniales 64, 95 Lauraceae 402 Leguminosae 374 Leersia 141 Lepuropetalon 342 Leucobryum 25 Limnophila 27)1 Linaria 395 Lindera 402 Liquidambar 141, 23 IB Lysmachia 229 Lythraceae 417 Magnolia 141, 150, 166, 370, 386 Magnoliaceae 264, 399 Manihot Al Melanthera 1,11 Menispermaceae 403 Micromeria 203 Micromyces 70 Meteorium 250 Modiola 82 Moenkemeyera 232 Monanthochloe 212 Monocotyledons 213, 358 Morchella 329 MuscillO, 134 Myrica 256 Myxomycetes 74, 234 jVa/fli 426 Nasturtium 254, 356 Nemophila 321 Nitella 330 Ny^ia 165 Oedogoniaceae 38 Oedogonium 38 Oe«o//jera81 Ophioglossum 58, 342, 362 Opuntia 184, 324 Oscillaloria 428 Oxa//i 123 Pachysandra 281 Panicum 240 Pancratium 57 Paronychia 205 Parpholis 183 Paspalum 239 Pellicularia 88 P/j/o;t 325 Phragmites 129 Physalis 339 Physcomitrium 218 Physostegia 121 A>JMj 15, 141 Pisum 60 Platygloea 85 Polygala 133 Polygonum 137, 313 Polyporaceae 158 Polyporales217 Polytrichum 25 Potamogeton 273 Prunus 231 B P5//omml67, 317, 376 Puccinia 89 Cw^rcz^ 98, 23 IB, 424 Ranales 153 Ranunculus 99, 168, 204 Ratibida 280 Rhamnus 420 Rhizoctonia 67, 88 Rorippa 253, 254, 299, 356 /?o/a/a 359, 417 /?ufru5 348 Rudbeckia 198 Rumex 303 Ruppia 426 Sabatia 103 Sagittaria 304, 366 5fl/tr231B,348 Salvia 345 Sarracenia 87 Schisandraceae 399 5c/rpw5 145, 220 Scutellaria 225 5e/trg/>ie//a 113,430 Senna 369 5e/a/-/a 410 Seymeria 369 Silene 1 14 Siphonychia 205 Sirobasidium 111, 127 Sirodotia 79 Sisyrinchium 9, 195, 200 Solmsiella 150 Spartina 41, 145,312,319,415 Specularia 109 Sphenopholis 383 Spirodella 188 Splachnobryum 105 5rac/i_v5 215 Staurastrum 49 ^o-rojc 389 Synchytrium 35, 52, 70, 71, 72, 80, 81, 82, 99, 109, 146 Syrrhopodon 162 Thallospora 43 Thelephoraceae 152 r///a231B r/Z/acfl 342 Tillandsia 394, 401 Tortula 266 Trag/a 122 63 Tremella 139 Tremellales44,55, 1(X) Trichomanes 412 7n/b////m305,348 Trillium 291 Triodanis 109 Typha 129 Vachellia 293 Vallisneria 426 Veronica 43 V/c/a 60 V/o/a 187 Wahlenbergia 138 VVVw^/fl 235 Xanthophyceae 398 Xyris 244 Youngia 226 KMCffl 184 Zanthoxylum 424 ZfMJtZ>2e361 Zizaniopsis 129 Zoopagaceae 96 Zygnemataceae 38 64 INDEX TO SUBJECTS Algae 30. 48, 54, 90, 156, 160, 207, 285, 328, 378,384,393,398,415 Aquatic plants 33, 40. 46, 48, 91. 108, 115. 172, 176, 188, 347, 357, 388, 392, 407, 426 Arboreta 216. 311 Archaeology 219 Bibliographies 28. 65. 75, 94, 177. 197. 199, 233. 272, 343, 368 Birds 13, 372 Bonner, Forrest 173 Botanists (see also Collectors) Anderson, Edgar 185, 265 Brizicky.G.K. 306 Brown. Clair 81, 109, 190, 200, 251. 280, 306 Cocks, R.S. 103.154.2318,280,306,402 Cook, M.T. 70, 71, 72, 80, 82, 146 Correll. D.S. 245 Correll. H.B. 245 Dormon. Caroline 114, 251 A. 335 Earle.T.T. 115 Evans, Alexander 23 Ewan. Joseph 345. 375, 414 Hardin. J. W. 264 Harper, Roland 173 Howe, Marshall A. 23 Hu^, labbe'l9 Koch. L.F. 218.353 Lowy. Bernard 127 Nelson. Ira S. 236, 251 A Olive, Lindsay 111 Penfound, W.T. 134, 176 Rafinesque, C.S. 102, 137, 149, 206, 261, 280. 321 Reese, WD. 218, 245 Riddell.J.L. 114,208,250.289,313 Robin, C.C. 137,389 Sargent, Charles Sprague 231 B Shinners.L.H. 368,375 Small, J. K. 140, 251 A Solymosy,S.L. 290,406 Svihla, R.D. 142 Thieret,J.W.225,330.417 Torrey, John 289 Underwood, L.M. 23 Viosca. Percy, Jr. 16. 1%, 197, 258 Wood, Carroll E.. Jr. 201 Carter, Jimmy 414 Checklists 128. 181, 182, 210, 224, 245, 247. 275. 353. 357. 358. 388 Collectors mentioned in commentary (see also Botanists and "index to Authors") 66 Arsene, Brother 19. 103. 136 Ball.C.R. 136 Drummond, T. 53. 103, 110, 133. 136, 144,263.313 Featherman, A. 110, 158 Hale, Josiah 4, 53, 103, 287, 289 Joor,J.F. 136,289 Langlois, A.B. 6. 7, 8, 10, 19, 34, 36, 45, 53, 80, 1 10, 1 17, 158, 253, 313, 323 Little. Henry 254 Mackaness, Faith Pennebaker 25, 105, 134, 180,218 Mohr,C.T. 110 Neon, Brother 19, 232.245 Pennebaker. Faith, see Mackaness, F.P. Waite,W.B.253 Colloquial names 92 Abbeville Reds 249 alligator-weed 31, 40, 129 baldcypress 5 1 , 340 ball-moss 401 beech 386 green ash 340, 419 holly 386 loblolly pine 349 log fern 381 Nuttalloak419 pinehill bluestem 391 pitcher plant 288 Spanish-moss 41, 276 strawberry 26 sugarberry419 sweetgum 340 sweet potato 360 water hickory 419 water-hyacinth 31, 39, 40, 46, 1 15, 308 yam 360 Conifers 15, 209, 290 Conservation, see Environmental quality Cultivated plants, see Horticulture Eckert, Allan W. 367 Ecology 294 aquatic habitats 115, 233 communities 177, 214 distribution, factors affecting 209, 214, 413. 427 fire 179, 288 productivity 115 salinity 396, 411 succession 3%, 411 vegetation 125, 145. 175. 300, 340, 346, 350, 388. 396, 409, 416, 426 water relations & flooding 346, 349, 387, 407, 427 Economic botany (see also Forestry) baldcyrpess51 carpentry. Creole 429 fiber 41 honey 348, 423 range plants 214, 334, 411 Spanish-moss 41, 276 65 tobacco 316 Endangered and rare plants 400, 401 Entomology 56, 154 Environmental quality 46. 107, 347, 421 Ferns and allies 21, 1 13, 247, 358, 381, 422 Fish, see Wildlife and fish Flotant31,46 Forestry 93, 94, 106, 199. 238, 294, 2% Foster, Edward 154 Fungi 14, 43, 44, 45, 55, 63, 90, 100, 147, 152, 158, 170, 192 217, 234, 257, 312 Geography 12, 22, 31. 276, 367, 385 Geology2, 219. 385 Generic studies 153, 344 Guides, see Manuals GulfofMexico90, 91, 108. 156. 160.371 Habitats and floristic provinces bayous 46, 367 bottomland forest 21, 97, 257, 340, 364, 387. 427 cemeteries 362 cheniers 22 coastal plain 314, 343 forests 29, 93. 124, 148, 243. 270. 294. 386.397,413.419 levees 350 longleaf pine-bluestem range 61, 240 mangrove swamps 91 marshes 17, 62, 91, 131, 145, 155, 157, 175, 179. 182, 191, 193, 214, 271, 319, 346,364.415 pine flatwoods 21. 300, 349 pitcher plant bog 288 prairie 301 shortleaf pine uplands 210 spoil banks 409 strand 91, 371, 416 swamps 46, 62, 97, 126, 396 History of botany 202. 208. 231 A, 231 B, 261,272,289 Horticuhure 3. 69, 128, 154, 161, 231, 236, 283, 290. 292, 31 1, 318, 54, 355, 404 Hurricanes 145. 157, 326 Insects 50, 64. 87. 95. 154, 364 Lepidoptera 1, 47, 83, 308 Invertebrates (see also Insects) 63, 77, 96, 112, 154 Lane, M.T. 28 Lichens 7, 19.36.315,323 Limnology 48, 172, 176, 248 Localities, Louisiana Alexandria. Rapides Parish 136, 429 Aurelia co. 323 Avery Island, Iberia Parish 13, 219 Barataria Bay 312 Baton Rouge, East Baton Rouge Parish 35, 43, 58, 64, 74, 80. 82, 84, 99, 109, 111,116,127,135,147,324 Bayou Bonfouca, St. Tammany Parish 6 Bayou Lafourche, Lafourche Parish 293 Bayou Millien 323 Black Lake 407 Bogalusa, Washington Parish 15 Bois Charmante 323 Breton Island, St. Bernard Parish 184, 326, 416 Cameron Marsh, Cameron Parish 22 Chandeleur Islands, St. Bernard Parish 156,160,181,326 Covington, St. Tamm^y Parish 19, 300 Delta National Wildlife Refuge, - Plaquemines Parish 361 Five Islands (salt domes), Iberia and St. Mary Parishes 267 Florida Parishes 16, 209, 270, 418 Folsom, St. Tammany Parish 166 Freemason Island, St. Bernard Parish 183 Garden Island Bay, Plaquemines Parish 129 Goodwood 85 Grand Bayou Blue 175 Grand Isle, Jefferson Parish 384 Hammond, Tangipahoa Parish 9 Hickory, St. Tammany Parish 257 Horseshoe Lake, Caldwell Parish 424 Jardin de Bouchetel 323 Lacassine Wildlife Refuge, Cameron Parish 330, 388 Lafayette, Lafayette Parish 19, 232, 425 Lake Chicot, Evangeline parish 176 Lake Pontchartrain 140, 172, 396, 426 Lake Providence, East Carroll Parish 48 Laplace, St. John the Baptist Parish 77. 96 Madisonville. St. Tammany Parish 313 Marsh Island. Vermilion Bay, Vermilion Parish 145 Mississippi Valley 231 A Morgan City, St. Mary Parish 24 Natchitoches, Natchitoches Parish 125 New Orleans. Orleans Parish 1, 3, 4. 39, 47, 53, 104, 115, 133. 140, 144, 220, 254,257,290,313,356,376 northern Louisiana, 243 Pointe a la Hache, Plaquemines Parish 7, 34 Redfish Bay, Plaquemines Parish 112 Reeves, Allen Parish 38 Rockefeller Wildlife Refuge, Cameron Parish 155 Sarpy Wildlife Refuge, St. Charles Parish 171 Silver Creek, Washington Parish 26 Slidell, St. Tammany Parish 136 St. Martinville. St. Martin Parish 7. 45. 117 66 Superior Canal, Cameron Parish 409 Tunica Hills, West Feliciana Parish 234,281, 413 Weeks Island, Iberia Parish 150, 186 Localities, Mississippi Mississippi Sound and islands 11, 108 Pascagoula River 274 Louisiana, additions to state flora algae 30, 38, 49, 68, 330, 378, 393, 398 bryophytes 6, 8, 23, 24, 25, 134, 142, 150, 162, 171, 186, 218, 223, 235, 246, 266, 278,279,298,331.370 fungi 35, 43, 44, 52, 55, 60, 64, 74, 77, 84, 85, 89, 95, 96, 98, 101, 109, 116, 135. 152, 166, 170, 192 lichens 7, 19,315 vascular plants 4, 9,16, 21, 57, 114, 118, 119, 120, 121, 122, 123, 138, 158, 168, 183, 188, 198, 201, 203, 204, 205, 212, 215, 220, 225, 227, 228, 229, 230, 237, 239, 242, 243, 249, 255, 268, 273, 282, 287, 291, 295, 302, 303, 304, 305, 306, 307, 337, 338, 339, 359, 361, 365, 373, 379, 380, 382, 383, 395, 410, 412, 424, 432 Louisiana parishes Acadia 320 Allen 38 Bossier 132 Caddo 433 Calcasieu 121 Caldwell 424 Cameron 22, 155, 225, 330, 388,409 De Soto 174 East Baton Rouge 35, 43, 58, 64, 74, 80, 82, 84, 99, 109, 111, 116, 127, 135, 147 213,224,324,394 East Carroll 48 East Feliciana 322 Evangeline 176, 322 Grant 363 Iberia 13, 131, 150, 186, 219, 267 Jefferson 97, 384 Lafayette 245, 297, 417 Lafourche 250, 293 Lincoln210, 247, 317 Morehouse 381 Natchitoches 125 Orleans 1, 3, 4, 39, 47, 53, 104, 115, 133, 140, 144, 215, 220, 253, 254, 257, 290, 313,356,376 Ouachita 262, 269, 275, 355 Plaquemines 7, 34, 129, 181, 253, 361 Rapides 136, 138, 429 Sabine 119, 174 St. Bernard 156, 160, 181, 182, 183, 184, 326,46 St. Charles 97, 171,350,3% St. James 316 St. John the Baptist 77, 96 St. Martin 7, 45, 112, 117,340 St. Mary 24. 86, 159,267,327 St. Tammany 6, 19, 103, 136, 141, 166, 257, 300, 313 Tangipahoa 9, 126 Tensas 351, 427 Vermilion 131, 145,225,417 Vernon 122, 174 Washington 15,26 273,373 West Feliciana 234, 281, 306, 377 386, 413 Louisiana rivers Mississippi 107, 427 Mississippi delta 11, 12, 34, 124, 148, 207,326 Red 125 Manuals (identification guides) 33, 59, 69 106, 143, 148, 173, 190, 222, 240, 251, 264, 310, 334, 345, 397, 414 Maps 177, 248, 271, 296, 352 Martin,W. 0.,Jr. 28 Mosses 6, 8, 25, 186, 223, 245, 275, 278, 314, 353, 405 Mycorrhiza 15 Natural History 274, 367 New Orleans Academy of Sciences 2 Paleobotany 32, 174,413 Philadelphia Academy of Sciences 7, 254 Phytogeography 314, 371, 413, 416, 418 Pines, see Conifers Plants (genera & families) cacti 324 composites 364 grasses 108, 189, 334, 364, 382, 383 oaks 364 orchids 20, 27 palms 5, 41, 354 spurges 372 Poisonous plants 21, 252 Pollen and palynology 76, 336, 348, 364, 423 Prain, D. 149 Soil microbiology 14, 217, 257, 270, 285, 327,328,415 Soils 86, 159, 165, 209, 217, 277, 346 Spontaneous combustion 17 Trees and shrubs 21, 97, 106, 128, 173, 190, 231, 231 B Vines 21, 260, 283, 404 Weeds 21, 37, 40, 46,284, 350 Wildflowers 21, 69, 143, 231, 251, 345, 414 Wildlife and fish 40, 61, 107, 129, 131, 141, . 184, 191 deer browse 61, 163, 178 286, 351, 397, 411 duck foods 130,155,157, 164, 189, 193, 211 Zoology 75 T^L 770H.'^ ISSN 0082-6782 ,.^1^ c?VARD Volume 21, Number 2 $3.50 JUNE 20, 1979 A SEASONAL AND ECOLOGICAL SURVEY OF FRESHWATER LIMPET SNAILS (PULMONATA: ANCYLIDAE) AND THEIR DIGENETIC TREMATODE PARASITES IN SOUTHEASTERN LOUISL\NA HUGH M. TURNER and KENNETH C. CORKUM p. 61 ECOLOGICAL MORPHOLOGY OF FRESHWATER STREAM FISHES A. JOHN GATZ, JR. p. 91 NICHE RELATIONSHIPS OF THAMNOPHIS RADIX HAYDENI AND THAMNOPHIS SIRTALIS PARIETALIS IN THE INTERLAKE DISTRICT OF MANITOBA DONALD R. HART p. 125 TULANE UNIVERSITY NEW ORLEANS TULANE STUDIES IN ZOOLOGY AND BOTANY, a publication of the Biology Department of Tulane University, is devoted primarily to the biology of the waters and adjacent land areas of the Gulf of Mexico and the Caribbean Sea, but manuscripts on areas outside this geographic area will be considered. Each number contains an indivi- dual monographic study or several minor studies. Normally two numbers plus an index and a table of contents are issued annually. Preferred citation of the journal is Tulane Stud. Zool. and Bot. INFORMATION FOR AUTHORS: Manuscripts submitted for publications are eval- uated by the editors and by an editorial committee selected for each paper. 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Subscription rates: Volume 21, 22 $7.50 ea., $8.50 foreign. Copies of Tulane Studies in Zoology and Botany sent to regular recipients, if lost in the mails, will be replaced if the editorial offices are notified before the second subsequent issue is released. COMMUNICATIONS: Address all queries and orders to: Editor, TSZ&B, Depart- ment of Biology, Tulane University, New Orleans, Louisiana 70118, U.S.A. Harold A. Dundee, Editor Arthur L. Welden, Associate Editor Samuel Clifford, Assistant to the Editors TULANE STUDIES IN ZOOLOGY AND BOTANY Volume 2 1 , Number 2 $3.50 JUNE 20, 1979 A SEASONAL AND ECOLOGICAL SURVEY OF FRESHWATER LIMPET SNAILS (PULMONATA: ANCYLIDAE) AND THEIR DIGENETIC TREMATODE PARASITES IN SOUTHEASTERN LOUISIANA HUGH M. TURNER* and KENNETH C. CORKUM Department of Zoology and Physiology, Louisiana State University, Baton Rouge, Louisiana 70803 ABSTRACT An 18-month survey involving almost 20,000 freshwater limpet snails of the family Ancylidae re- vealed three species, each representing a different genus, occurring in southeastern Louisiana. Laevapex fuscus was collected only from lentic habitats, where it exhibited, at most, a bivoltine pattern of reproduction with two successive genera- tions per year; in habitats of low primary productiv- ity a single annual generation was produced. Ferris- siafragilis was collected from lotic and ientjc habitats and produced a single annual generation. Hebetan- cylus excentricus also displayed a broad habitat pre- ference but exhibited up to a trivoltine pattern of reproduction. Southeastern Louisiana ancylid snails play a major role in the ecology of digenetic trematodes since 19 species of larval trematodes were found developing in three ancylid species. Hosts, descriptions, season- al incidence, and possible identity of these cercariae are given. Seasonal occurrence of trematode intramoUuscan stages was closely related to seasonal occurrence of the ancylid snail hosts. Bivoltine patterns of repro- duction for Hebetancylus excentricus were reflected in two generations of cercarial production. On the basis of comparative ecology, anatomy, and hosted trematodes, we propose that Hebetan- cylus occupies an intermediate phylogenetic position between Ferrisia and Laevapex. Ferrissia is least specialized in terms of its hosted trematode fauna, which bears similarity to that hosted by snails of the family Planorbidae. * Present Address: Department of Biology, Mc- Neese State University, Lake Charles, Louisiana 70605 INTRODUCTION Literature on parasites of ancylid snails is meager, with less being known than for other major families of freshwater gastro- pods (Malek and Cheng, 1974). Smith (1967) reviewed the literature on larval digenetic trematodes developing in ancyl- ids and noted that the six reports from 1827 to 1959 were incomplete in descrip- tion, with no indication as to specific or generic identity of the adult trematodes. None was from North America. Smith (1959) was the first to report North American ancylid snails as first in- termediate hosts for trematodes. He re- ported the occurrence of Megalodiscus temperatus (Stafford, 1905) in Ferrissia fragilis (Tryon, 1863) in Michigan. The life cycle of this rectal parasite of frogs had been elucidated by Krull and Price (1932), who implicated the planorbid snail Heli- soma trivolvis (Say, 1817) as a natural snail host. Peters and Self (1963) reported an oph- thalmoxiphidiocercaria of the family Al- locreadiidae Stossich, 1903 developing in Laevapex fuscus (C.B. Adams, 1841) from southeastern Oklahoma and encys- ting in the mantle of unionid clams and limpets, including limpets hosting the re- diae. These investigators believed the cer- caria to be that of Allocreadium ictaluri EDITORIAL COMMENTS FOR THIS PAPER: DR. DAVID W. FREDERICKSON, Assistant Professor of Biology, Tulane University, New Orleans, Louisiana 701 18 DR. R. JAY SMITH, Associate Professor of Biology, University of Detroit, Detroit, Michigan 48221 67 68 Tulane Studies in Zoology and Botany Vol. 21 Pearse, 1924, an intestinal parasite of cat- bile described by Cort (1918) and reported fish. They noted that miracidia, hatched by Wallace (1941) to be the larva of Tri- from eggs obtained from infected catfish, ganodistomum mutabile (Cort, 1918), an penetrated and remained in the tissues of intestinal parasite of catostomid fishes. L. fuscus. All other members of the tre- Wallace (1941) reported Helisoma trivol- matode family Allocreadiidae, for which vis and H. campanulata (Say, 1821) as life cycles are known, utilize sphaeriid natural snail hosts, with the planarian clams as first intermediate hosts (Yama- Dugesia sp. or the commensal annelid, guti, 1975). Chaetogaster limnaei von Baer, 1827 serv- Basch (1963), in his monograph on ing as second intermediate host for T. North American Ancylidae, noted that of mutabile. 150 ancylid collections taken during the Duncan and DeGiusti (1976) produced summer of 1961, from 23 states, about evidence that the hsorchiid cercariaeum, 15% were heavily infected with one or thought by Smith (1968) to be Cercari- more species of trematode larvae. aeum mutaOile, was m fact a complex of Smith (1967) examined Ferrissia fragil- three cercarial species. These cercariaea is, F. parallela (Haldeman, 1841), and differed in size, arrangement of tegumen- Laevapex fuscus from several marsh tal papillae, tegumental spination pat- pond, stream, and river localities in terns, and species of intermediate hosts, southeastern Michigan. Fifteen different Since Duncan and DeGiusti believed trematode species were found to use one them to be larval stages of described lis- or more of the three limpet species as first sorchiids, they did not formally describe intermediate hosts. Unfortunately, Smith them as new species. They did, however, did not figure or otherwise describe those provide full descriptions while designating cercariae except to note that Megfl/o^wc«5 them as Cercariaeum types I through III. temperatus developed in all three limpet Cercariaeum type I utilized Laevapex fus- species. Identification of the other 14 cer- cus as first intermediate host with Chaeto- cariae was only to the level of superfamily gaster limnaei as second intermediate, or family. Cercariaeum types II and III utilized L. In addition to Megalodiscus temperatus, fuscus and Ferrissia rivularis respectively, Smith (1967) noted another amphistome while Chaetogaster limnaei, Dugesia tigri- developing in Ferrissia parallela; a strige- na (Girard, 1850) and D. dorotocephala oid in F. fragilis; another in F. parallela; (Woodworth, 1897) served as second in- an echinostomatid in F. fragilis; two oth- termediates. ers in F. parallela; a fourth in Laevapex Underwood and Dronen (1977) report- fuscus;2LSTp\rovc\n\d'\nF. fragilis, F. paral- ed an unidentified species of Ferrissia lela, and L. fuscus; a lissorchiid and cyan- from Texas as an experimental host for the thocotylid in L. fuscus; and three xiphidi- frog lung fluke Haematoloechus brevi- ocercariae in F. parallela. Significance of plexus Stafford, 1902. These investigators the apparent specificity exhibited by most noted that xiphidiocercariae obtained cercariae is tempered by Smith's failure to from Ferrissia sp. were smaller at maturity note the occurrence of sympatry among than H. breviplexus cercariae described by the limpet hosts. Schell (1965) from the experimentally in- Smith (1968) reported that lissorchiid [^ l^Vt"'' ^9^'' '^^''""'"' '''"'''''" cercariaea (tailless cercariae), previously ^ - I ^^ j^ ^^^ reported (Smith, 1967) m Laevapex /m.$cw5 „ . ' ,. ^ , •, ^ , r from Michigan, also parasitized Ferrissia Ferrissia fragilis as a natural snail host for rivularis (Say, 1817) in southeastern Mich- ^jie turtle blood f^uke Spirorchis scr^pta igan. According to Smith, this cercari- Stunkard, 1923 in Louisiana. The larva of aeum was identical to Cercariaeum muta- ^- '"^y. ^^^ Previously been reported to develop in certain species oi naturally in- No. Limpet Snails 69 fected planorbid snails (Goodchild and Kirk, 1960; Holliman and Fisher, 1968). From the above review, ancyHds obvious- ly appear to have been a neglected group in most parasitological studies. The objectives of this study were as fol- lows: (1) to survey the freshwater limpet (Ancylidae) fauna of southeastern Louisi- ana, (2) to investigate related ecological aspects including habitat, seasonality, and population dynamics, (3) to investigate its role in the ecology of digenetic trematode parasites, (4) to provide descriptions and figures of those trematode cercariae de- veloping in southeastern Louisiana ancyl- ids, (5) to identify cercariae by life cycle studies and/or a survey of trematode life history literature, and (6) to postulate a phylogenetic relationship for southeast- em Louisiana ancylids based upon simi- larities and differences in hosted trema- todes. MATERIALS AND METHODS Limpets were collected biweekly, for up to 18 consecutive months, from five ecol- ogically dissimilar localities in southeast- em Louisiana. Collecting sites, with sur- vey period in parentheses, were as fol- lows: drainage ditch on Ben Hur Road Experimental Farm, 1 mi S of LSU Baton Rouge campus. East Baton Rouge Parish, (June 1975-November 1976); Beaver Pond Branch, 3 mi SE of Livingston, Liv- ingston Parish, on Interstate Highway 12 (July 1975-November 1976); borrow pit on W side of Atchafalaya floodway levee at Ramah, Iberville Parish, on Interstate Highway 10 (July 1975-August 1976); roadside swamp, 2 mi SE of Sorrento, As- cension Parish, on Rt. 61 (June 1975-July 1976) ; and pond, 2 mi E of Head of Island, Livingston Parish, on Rt. 22 (June 1975- August 1976). Collections were made by removing, with a sharp scalpel blade, limpets attach- ed to submerged or floating debris as well as leaves and stems of floating or sub- merged vegetation. Specimens were placed in glass finger bowls containing water from the collecting site. Bowls rest- ed on an inverted styrofoam ice chest that served as a floating work table. Collecting time for each of the 156 col- lections was a multiple of 15 minutes dura- tion. This was done to establish an index of relative abundance by noting number of limpets collected during 15 minutes. To roughly determine population structure, all individuals seen, regardless of size, were collected. Snails were maintained in their collec- tion containers until identified, graded in- to sizes, and examined for larval digenetic trematodes; this was usually done within 24 hours after collection. Voucher speci- mens of all limpet species were deposited in the mollusc collection of the Delaware Museum of Natural History (DMNH). The smaller limpet species Ferrissia fra- gilis (DMNH 119530) was graded into three size classes based upon shell length: <2 mm, 2-4 mm, and>4 mm. Laevapex fuscus (DMNH 119532) and Hebetancylus excentricus (Morelet, 1851) (DMNH 119531) were graded into classes:<3 mm, 3-5 mm, and>5 mm. Presumably changes, over the course of our survey, in relative numbers of each class should reflect popu- lation dynamics, including reproduction, recmitment, and death. Although collec- tions were biweekly, data on limpet popu- lations were pooled for each month. Visual screening for digenetic trema- tode larvae was possible since most of the limpet's "soft-parts," including the diges- tive gland, were exposed to inspection when the snail was inverted. Identification was made by dissection or by allowing cer- cariae to emerge from live, isolated indi- viduals. Limpets without visible germinal sacs (rediae or sporocysts) were dissected and examined for immature larval stages and/or metacercariae. Biweekly data on incidence of infec- tion, like those used for population biol- ogy, were pooled for each month. Only those individuals of Hebetancylus excen- tricus and Laevapex fuscus greater than 3 mm shell length were considered when computing monthly incidence. Neither cercariae or germinal sacs were evident 70 Tulane Studies in Zoology and Botany Vol. 21 among the smaller individuals; however, this was not true for Ferrissia fragilis, all sizes of which' were considered when com- puting incidence. Cercariae, designated as types I-XIX (Figures 1-19) were studied live and un- stained, vitally stained with nile blue, or fixed in hot 10% formalin. Unless other- wise noted, measurements were taken from 10-20 formalin preserved specimens under light coverslip pressure and are ex- pressed in microns. Ranges are given with mean in parentheses. Drawings are com- posites made from live and preserved specimens, either free hand, with aid of a microprojector, or from photomicro- graphs. RESULTS AND OBSERVATIONS Ecology of Southeastern Louisiana Ancylids After 18 months of collecting and exam- ining almost 20,000 ancylid specimens, three species, each representing a differ- ent genus, were found to occur in our study areas. Laevapex fuscus was collected from submersed stems of emergent vege- tation and from submersed debris at three localities exhibiting a lentic character. Ferrissia fragilis and Hebetancylus excen- tricus occurred on emergent and floating vegetation at all five collecting sites, which along with the seasonality and population dynamics of their ancylid snail faunas, are characterized below. 1. Ben Hur Experimental Farm (BHF) This 3 m wide, mud-bottom ditch, al- though never dry, had an intermittent flow and drained the LSU experimental farm and 20 shallow ponds for experimen- tal rearing of catfish and crayfish. During the 18-month survey of this locality, 3,774 Hebetancylus excentricus and 172 Ferrissia fragilis were collected exclusively off submersed leaves and stems of alligator weed [Alternanthera sp.). Laevapex fus- cus was not seen or collected. Population dynamics and seasonal abundance of both Ferrissia fragilis and Hebetancylus excentricus are presented in Figure 20. F. fragilis was collected inter- mittently and in small numbers during the course of the survey at BHF. Usually few- er than two individuals were collected per 15 minutes of collecting effort. A peak in numbers occurred in April, when about 10 limpets were collected per 15 minutes. This peak was followed by a gradual de- cline to zero in August. Recruitment oc- curred in mid spring as evident by the increasingly greater percentage of the population comprised by individuals of the small size class (<2 mm length). Indi- viduals of the large size class (>4 mm length), probably representing the post- ovigerous segment of the population, were collected only at this time. Basch's (1963) observation of maximum length of 4 mm for F. fragilis may substantiate a post-ovigerous role for those large size class individuals. However, large size class (>5 mm length) Laevapex fuscus and H. excentricus, noted by Basch to have max- imum lengths of 7.75 and 5.8 mm respec- tively, would not necessarily comprise the post-ovigerous segment. Hebetancylus excentricus was present throughout the study and exhibited two distinct population peaks: one in Novem- ber, and another in May. This species dis- played a bivoltine pattern of reproduction with two successive generations per year as evident by the observation that peaks in the index of relative abundance coincide with or closely follow periods of increase in percentage of small size class individu- als (< 3 mm length). In H. excentricus, reproduction and recruitment are preced- ed by a build up in percentage of the large size class sector of the population. 2. Beaver Pond Branch (BPB) This narrow, continuously flowing stream originates near Livingston, Louisi- ana, and empties into Hog Branch, a larg- er, sand-bottomed stream forming part of the drainage for the so-called Florida Par- ishes of southeastern Louisiana. During periods of high water in Hog Branch (two occurred during our survey) BPB received backwater from Hog Branch. This condi- tion persisted for several days before normal flow returned in both streams. No. 2 Limpet Snails 71 Hebetancylus excentricus and Ferrissia fragilis were collected from floating vege- tation and from emergent arrowhead {Sagittaria sp.). Laevapex fuscus was not encountered. Figure 21 shows population biology and seasonal abundance of 1,294 H. excentricus and 1,853 F. fragilis col- lected during 17 months.. Seasonality is evident and perhaps reflects seasonal par- titioning of the habitat by the two limpet species. From January to July, 1976, F. fragilis was present in large numbers at a time when H. excentricus was almost ab- sent from the habitat. H. excentricus be- came the predominant limpet species dur- ing the fall. 3. Ramah (RAM) This site is a borrow pit of the Atchafa- laya River Basin and is subjected to large fluctuations in water level. Because of fluctuations, emergent vegetation that consisted mainly of Sagittaria was season- al, and occurred mostly in summer and fall. Although no measurement of dis- solved oxygen was taken during the study, this habitat would be classified as meso- trophic. All three limpet species were sympatric at RAM. Laevapex fuscus was collected from submersed and floating debris, water hyacinth (Eichornia crassipes), and emergent Sagittaria. Ferrissia fragilis and Hebetancylus excentricus were similarly collected; however, H. excentricus was seldom found on debris. Limpet popula- tion biology and seasonal abundance for 3,663 L. fuscus, 511 F. fragilis, and 425 H. excentricus collected during our 14-month survey are presented in Figure 22. As at BPB, Hebetancylus excentricus and Ferrissia fragilis displayed a pro- nounced seasonality that was interpreted as seasonal partitioning of the habitat. However, seasonality of limpets may also reflect seasonal absence of vegetation on which H. excentricus was most often col- lected. F. fragilis, like the BPB popula- tion, was abundant in early spring through summer, while H. excentricus was most abundant during the fall. Data indicate production of a single H. excentricus and F. fragilis generation per year. Laevapex fuscus was the predominant limpet species at RAM and was collected in relatively high numbers almost year round. The low fall-winter index of rela- tive abundance may have been due to L. fuscus burrowing into the mud, where they were unavailable for collection by our methods. Indications are that a single generation is produced annually. Re- cruitment certainly began in April, be- cause population structure for that month derhonstrates an expanding population. 4. Sorrento (SOR) This is a shallow, former, roadside swamp, with a mud bottom. Water level was maintained at about 0.5 m during the first 12 months of the survey; however, in June 1976 the level began to drop and the site was completely dry by August 1976. The 100 m X 30 m area had been cleared several years earlier as right of way for an underground pipeline, and was free of trees. As a result, a luxuriant growth of emergent grass (Paspalum sp.) was main- tained almost year round and gave the locality a marsh appearance. This site, with its shallowness, abundant emergent vegetation, and high primary productiv- ity, fits the criteria of a eutrophic habitat. All three ancylid species occurred at this locality; however, as seen in Figure 23 the scale for relative abundance of Lae- vapex fuscus is 10 fold greater than for Hebetancylus excentricus and 100 fold greater than for Ferrissia fragilis. Total numbers collected were 4,960, 352, and 40, respectively. Ferrissia fragilis was collected in too small numbers for any determination of population dynamics; however, a pro- nounced seasonality was noted. Hebet- ancylus excentricus appears to have a single generation each year with recruit- ment in the spring. Laevapex fuscus produced a fall and spring generation. Egg hatching and re- cruitment from one biannual cycle ex- tended into the next, and thus gave the appearance of year-round reproduction. Another observation concerning the L. 72 Tulane Studies in Zoology and Botany Vol. 21 fuscus population was the predominance of large-size class individuals in most pooled monthly samples. Predominance of large individuals indicates a rapid growth rate, probably attributable to the eutrophic environment, and is striking when compared to the pooled monthly population structures at the less eutrophic Ramah (Figure 22). As will be shown lat- er, apparent differences in productivity have considerable implications for the ecology of the respective digenetic tre- matode populations. 5. Head of Island Pond (HIP) This permanent pond has a shallow margin at its north end lined with emer- gent vegetation. The remaining margin was deep and without emergent vegeta- tion. Much sunken debris — sheets of plas- tic, metal cans, glass, etc., was found along the front or south margin. Laevapex fuscus occurred on this debris; however, because of depth, collecting was difficult. For this reason, L. fuscus was not included among the ancylids studied at this locality. We mention the occurrence because it demonstrates habitat selection by the three ancylid species, since L. fuscus was found only on this deep water debris, while Hebetancylus excentricus and Ferris- sia fragilis occurred only on debris and emergent vegetation along the shallow margin. Population biology and seasonal abun- dance for 1,836 Hebetancylus excentricus and 182 Ferrissia fragilis collected during our 15-month survey are shown in Figure 24. F. fragilis was abundant only in Febru- ary and March. We believe that presence of individuals classed as large, accom- panied by an increasing percentage of the smaller class, indicates recruitment at this time. Hebetancylus excentricus occurred year round with production of three genera- tions; one in the summer, another in the fall, and the third in the spring. Digenetic Trematodes in Southeastern Louisiana Ancylids Cercariae representing 19 trematode species were found developing in one or more of three ancylid species. Hosts, descriptions, seasonality of incidence, and possible identity of these larvae are given below (Figures 20-24). Cercflr/fl Type I (Figures 1, 21, 22, and 23) Laevapex fuscus collected from SOR, Ferrissia fragilis from BPB, and both spe- cies from RAM were found to harbor in- fections with xiphidiocercariae of the ar- matae group and designated Cercaria type I. Description: Body 228-276(254) long by 84-112(99) wide and covered with small spines that become less dense at the pos- terior end. Caudal pocket present. Tail 184-228(205) long by 29-35(32) wide at base. Oral sucker 53-63(58) long by 56- 64(61) wide. Acetabulum 45-51(47) long by 44-53(50) wide. Stylet shouldered, 28- 33(31) long. Five pairs of pre-acetabular penetration glands with ducts that empty near stylet. Ceca extend to post- acetabular level. Excretory bladder cellu- lar and Y-shaped. Flame cell formula or number not determined. Ferrissia fragilis harboring these cercar- iae were collected only during July of the second summer and comprised 1% of the sampled population at BPB (Figure 21). Infection occurred in 0.8% of the sampled F. fragilis population during the same month at RAM (Figure 22). Laevapex fuscus from this locality, however, har- bored infection throughout spring and summer. At SOR (Figure 23) Cercaria type I was present in L. fuscus almost year round with a peak seasonal incidence of 8.5% in August. Cercaria Type II (Figures 2, 20, 21, 23, and 24) Amphistome cercariae of the diploco- tylea type, designated Cercaria type II, were found developing in both Laevapex fuscus and Ferrissia fragilis collected at SOR. F. fragilis from BHF, HIP, and BPB also harbored infections. Description: Body 230-292(260) long by 102-144(123) wide. Tail 415-450(432) long by 35-46(40) wide. Oral sucker 118- No. 2 Limpet Snails 73 147(128) long. Acetabulum 40-77(57) trunks containing many small concretions, long by 113-149(131) wide. Ceca thick Flame cell formula, or number not deter- w ailed, extend posteriorly to near level of mined, excretory bladder. Paired excretory Figures 1-7. Cercaria types I-VII developing in southeastern Lousiana ancylid snails. Scale in microns. 74 Tulane Studies in Zoology and Botany Vol. 21 Cercaria type II, although smaller, fits of that species developing in Laevapex the description given by Krull and Price fuscus and Ferrissia fragilis from Michi- (1932) for the cercaria of Megalodiscus gan. Smith reported an overall incidence /eArzpera/w5, and by Smith (1967) for larvae of 14% for F. fragilis, whereas an inci- Figures 8-14. Cercaria types VIII-XIV developing in southeastern Louisiana ancylids. Scale in microns. No. 2 Limpet Snails 75 Figures 15-19. Cercaria types XV-XIX developing in southeastern Louisiana ancylids. Scale in microns. 76 Tulane Studies in Zoology and Botany Vol. 21 dence of only 1% was noted for infections in L. fuscus. Cercarial production in Michigan F. fragilis was in two distinct waves, one in spring and the other in au- tumn (Smith, 1967). Like Smith, we noted a disparity in in- cidence of infection among the two limpet species. Of almost 5,000 Laevapex fuscus examined from SOR, only one harbored infection with Megalodiscus temperatus, while eight of 40 Ferrissia fragilis were infected. Two waves of cercarial produc- tion were observed. One began in late fall and continued until early spring, while the other began in early summer. Cercaria Type III (Figures 3, 20, 22, 23. and 24) Longifurcocercous, apharyngeate, mon- ostome cercariae, with small eyespots, were found developing in all three limpet species and were collected at all localities except BPB. Description: Body 179-207( 194) long by 31-39(34) wide, with 12 annulations be- tween oral sucker and acetabular anlagen. Two pairs of lateral setulae on posterior bodv near junction of tail stem. Tail stem 202-246(232) long by 26-37(31) wide, with five pairs of lateral setulae grouped near junction with body, and six or seven pairs grouped near bifurcation. Caudal bodies not observed in tail stem. Furcae 161- 207( 196) long by 16-23( 17) wide, with del- icate finfolds originating at level of excre- tory pores, which emptied on anterior sur- face of furcae. Oral sucker 37-55(46) long by 17-28(21) wide. Three pairs of penetra- tion glands between eyespots and excre- tory bladder, with acetabular anlagen lo- cated immediately posterior to anterior pair. Genital primordium between poster- ior penetration glands and excretory bladder. Extent of ceca not determined. Eleven pairs of tlame cells located in body and two pairs in tail. Cercaria type III resembles the larva of Posthodiplostomum minimum (MacCal- lum, 1921), a parasite of the heron, Nycti- corax nycticorax (Linnaeus, 1758). Two morphologically different cercariae have been described for P. minimum. Miller (1954) described Cercaria minimum, which bears similarity to Cercaria type III; however. Miller noted the presence of five pairs of caudal bodies in the tail stem, and 10 pairs of flame cells in the body and tail. Cercaria minimum Miller, 1954 developed in naturally infected Physa heterostropha Say, 1817 in Michigan. More recently, Bedinger and Meade (1967) reported on the life cycle of Posthodiplostomum min- imum, the cercaria of which developed in naturally infected Physa halei Lea, 1864 in Texas. Those cercariae lack caudal bod- ies, have "flagellets" in two groups on the tail stem, and have 13 pairs of flame cells in the body and tail. On morphological grounds Cercaria type III, although somewhat larger, more nearly resembles the cercaria described by Bedinger and Meade; however, in size, it resembles Cercaria minimum. Cercaria Tvpe IV (Figures 4, 20, 21,22, 23, and 24) Longifurcocercous, pharyngeate, di- stome cercariae were found developing in all three species of limpets and were col- lected at all five localities. Description: Body 1 19- 178( 151 ) long by 41-62(51) wide, with anteriorly directed needle-like spines extending posteriorly to level of pharynx, and with three pairs of lateral papillate setulae. Tail stem 228- 364(3 16) long by 72-98(77) wide, with four pairs of lateral setulae. Furcae 180-292 (239) long by 32-48(39) wide, with excre- tory pores opening at mid-length of pos- terior surface. Oral sucker 35-44 (39) long by 25-32(28) wide. Pharynx 9-12(10) long by 22-28(26) wide, with a single circle of inwardly directed, needle-like spines. Number and arrangement of penetration glands not determined. Ceca extended to postacetabular level. Six pairs of flame cells in body and two pairs in tail. Cercaria Type V (Figures 5 and 20) An echinostome cercaria, with spiny collar, was found developing only in He- betancylus excentricus from BHF. Meas- No. 2 Limpet Snails 77 urements of Cercaria type V used in the following description were taken from a single live specimen. Description: Body 176 long by 148 wide, with an undetermined number of small collar spines. Tail without finfold, 392 long. Oral sucker 44 long by 44 wide. Acetabulum 50 long by 50 wide. Extent of ceca not determined. Many small con- centrations restricted to middle and anter- ior portion of paired excretory trunks. Flame cell formula or number not deter- mined. Cercaria type V was collected only dur- ing June of the first summer and com- prised 4.1% of the sampled Hebetancylus excentricus population (Figure 20). Cercaria Type VI (Figures 6, 20, 21,22, and 23) Echinostome cercariae, without spiny collars, were found developing in all three limpet species at all localities except HIP. Description: Body 168-196(184) long by 86-116(102) wide. Tail without finfold, 364-416(393) long by 43-53(46) wide, with pair of short flagellets near tip. Oral suck- er 35-43(39) long by 38-43(40) wide. Ace- tabulum 37-50(43) long by 51-58(54) wide. Extent of ceca not determined. Paired ex- cretory trunks each contain three or four large concentric concretions restricted to pre-acetabular region. Thirteen pairs of flame cells present in body. Cercaria type VI may be a species of Echinochasmus Dietz, 1909, since cercar- iae of this genus often lack collar spines, have few and large excretory concentra- tions, and have short flagellets on the tail (Yamaguti, 1975). Cercaria Type VII (Figures 7, 20, 21, and 22) A brevifurcocercous, apharyngeate, distome cercaria of the turtle blood fluke family Spirorchiidae Stunkard, 1921 was found developing only in Ferrissia fragilis from BHF, BPB, and RAM. Description: Body 192-256(219) long by 58-72(64) wide at acetabulum, with three pairs of lateral setulae. Anterior third of oral sucker covered with dense, fine spines. Spines around acetabulum large and numerous. Remainder of body cover- ed with minute, sparse spines. Tail stem attached subterminally, 568-680(590) long by 52-67(60) wide. Short lateral setulae on tail stem arranged in two groups of nine pairs each. Anterior group all nonpapil- late. Posterior group with last three pairs papillate. Furcae 200-268(227) long, with finfolds along margin. Oral sucker 44- 69(60) long by 28-37(34) wide. Acetabu- lum protrusible, 24-30(28) long by 26- 32(28) wide. Eyespots in second quarter of body. Five pairs of penetration glands from posterior margin of eyespots to fourth quarter of body. Sixth pair of fused glands in fourth quarter. Ceca extend to level of acetabulum. Five pairs of flame cells in body and one pair of large flame cells in tail. Under laboratory conditions cercariae emerged mostly at night. Life cycle studies (Turner and Corkum, 1977) indicated Cercaria type VII to be the larva of Spirorchis scripta. Cercaria Type VIII (Figures 8 and 22) A xiphidiocercaria of the armatae group parasitized only Ferrissia fragilis from RAM. Measurements from the following description were taken from a single living specimen. Description: Body 252 long by 140 wide. Tail 203 long by 42 wide. Oral suck- er 69 long by 64 wide. Stylet unshoulder- ed, 25 long. Acetabulum 53 long by 64 wide. Extent of ceca not determined. Ex- cretory bladder Y-shaped. Arrangement and number of flame cells or penetration glands not determined. Cercaria Type IX (Figures 9 and 22) An ophthalmoxiphidiocercaria of the family Allocreadiidae was found develop- ing only in Laevapex fuscus collected at RAM. Measurements for the following description were taken from a single living specimen. Description: Body 328 long by 228 wide at acetabulum. Tail 270 long by 43 wide. Oral sucker, with five rows of small, blunt 78 Tulane Studies in Zoology and Botany Vol. 21 spines forming a semicircle below mouth, 56 long by 69 wide. Acetabulum with two alternating circles of small, blunt spines, 64 in diameter. Stylet short and blunt. Eyespots in first quarter of body. Extent of ceca not determined. Number and ar- rangement of flame cells or penetration glands not determined. According to Dr. Lewis E. Peters (1975, pers. comm.), Cercaria type IX ap- pears "very similar to the cercaria that [he and Dr. J. Teague Self] studied in Okla- homa in the early 1960's." Peters also not- ed that he and Dr. Raymond Cable had confirmed identity of the cercaria, which also occurred in Laevapex from Indiana, to be that of Allocreadium ictaluri. Al- though Peters and Self (1963) reported L. fuscus as one of several second intermedi- ate hosts, we were unable to find any har- boring allocreadiid metacercariae among the 3,663 L. fuscus examined from RAM. Cercaria Type X (Figures 10 and 23) A small xiphidiocercaria of the armatae group, designated Cercaria type X, was found developing only in Laevapex fuscus atSOR. Description: Body 152-182(169) long by 74-90(86) wide. Caudal pocket present. Tail 106-140(123) long by 30-37(35) wide. Acetabulum 24-28(26) long by 23-28(25) wide. Stylet slightly shouldered, 23-27(26) long. Extent of ceca not determined. Ex- cretory bladder Y-shaped. Flame cell formula or number not determined. Pen- etration gland number and arrangement not determined. Pharynx, although pres- ent, was not observed. Cercaria Type XI (Figures 1 1 and 22) Tailless cercariae (cercariaea) of the family Lissorchiidae Poche, 1926 were found developing in Laevapex fuscus and Ferrissia fragilis at RAM. Description: Body 324-392(358) long by 80-128( 101) wide, with nine pairs of later- al papillate setulae. Pharynx 24-28(26) long by 26-31(29) wide. Oral sucker 50- 64(54) long by 5 1-61(57) wide, with mouth surrounded on three sides by two alter- nating rows of small, inwardly directed spines. Acetabulum 58-69(63) long by 58- 71(64) wide, with two alternating circles of small, inwardly directed spines surround- ing inner rim, and a third row present on anterior quadrant of inner rim. Tegument with larger spines directed posteriorly; however, spines absent from area be- tween oral sucker and acetabulum. Ceca extend to near base of excretory bladder. Excretory bladder cylindrical and thick walled. Flame cell and penetration gland number and arrangement not determined. Cercaria type XI resembles Cercariae- um type I described by Duncan and De- Giusti (1976); however, it differs in host specificity (not restricted to Laevapex fus- cus), in size (slightly larger than Cercar- iaeum type I), in arrangement of lateral papillate setulae, and in number and ar- rangement of small spines on the oral sucker and acetabulum. Duncan and DeGiusti (1976) noted that papillae pattern and number were vari- able; however, tegumental spination pat- tern was fixed. Differences in size may be attributable to techniques used by these authors, since they first relaxed emergent cercariae in menthol solution prior to fix- ing in hot 10% formalin. They also meas- ured cercariae in groups of twenty under a "floating" coverslip. Cercaria Type XII (Figures 12 and 21) A longifurcocercous, pharyngeate, di- stome cercaria was found developing only in Hebetancylus excentricus from BPB. Description: Body 108-156(131) long by 62-99(83) wide. Tail stem 108-200(149) long by 83-150(108) wide. Furcae 104- 163(124) long. Oral sucker 35-48(41) long by 29-39(34) wide. Acetabulum 23-29(25) long by 23-30(26) wide. Ceca surround acetabulum and extend to post-acetabular position. Tail, as seen in lateral view, ap- pearing as a pair of empty, pressed trous- ers with excretory pores opening on inner surface of furcae. Number and arrange- ment of flame cells and penetration glands not determined. No. 2 Limpet Snails 79 Cercaria Type XIII (Figures 13, 20 and 24) A longifurcocercous, pharyngeate, dis- tome cercaria parasitized only F. fragilis from HIP and BHF. Description: Body 164-220(193) long by 41-58(46) wide. Tail stem 288-316(297) long by 172-196(186) wide at bifurcation. Furcae 220-268(248) long by 40-60(48) wide. Oral sucker 44-58(51) long by 25- 39(28) wide. Pharynx 12-14(13) long by 14-18(17) wide. Acetabulum with three circles of blunt, concentrically placed spines, 25-32(30) long by 30-35(32) wide. Three pairs of penetration glands, two pre- and one post-acetabular. Several smaller pairs of cells scattered throughout the body. Genital primordium between posterior penetration glands and excre- tory bladder. Extent of ceca not deter- mined. Anterior body to level of pharynx covered with small, dense spines. Flame cell formula or number not determined. Tegument on tail stem at bifurcation formed into loose, bladder-like "skirt" continuous with furcae. "Skirt" most ev- ident when cercariae were placed under coverslip pressure. Figure 13 was drawn without coverslip pressure and shows "skirt" in folded configuration. Cercaria Type XIV (Figures 14 and 20) A longifurcocercous, pharyngeate, di- stome cercaria was found developing only in Ferrissia fragilis from BHF. Description: Body 112-153(127) long by 32-51(44) wide, with small dense spines to level of mid-oral sucker. Tail stem 187- 220(207) long, with nine pairs of lateral setulae. Furcae 162-189(179) long by 14- 18(16) wide. Oral sucker 32-41(37) long by 26-34(30) wide. Acetabulum 18-21(20) long by 19-25(23) wide, with two alternat- ing circles of small, blunt spines. Two pairs of penetration glands, one pre- acetabular the other post-acetabular. Genital primordium between posterior pair and excretory bladder. Extent of ceca not determined. Five pairs of flame cells in body and two pairs in tail. Cercaria Type XV (Figures 15 and 24) A single Ferrissia fragilis collected at HIP was found to harbor infection with a longifurcocercous, pharyngeate, distome cercaria. Description: Body 132-188(161) long by 34-52(41) wide, with small, dense spines to level of mid-oral sucker, less densely spined to level of gut bifurcation. Tail stem 280-336(305) long by 124-152(140) wide. Furcae 220-276(252) long by 28- 46(35) wide. Oral sucker 35-52(45) long by 20-30(25) wide. Acetabulum 25-30(27) long by 28-32(30) wide, with three alter- nating circles of short, blunt spines. Ceca extend to mid-acetabulum. Two pairs of penetration glands, one pre- and the other post-acetabular. Muscle fibers from me- dian portion of each fiber tract in tail stem contralateral in furca and form a chiasma anterior to bifurcation. Lateral fibers ip- silateral. Two pairs of flame cells in tail. Flame cell number or arrangement for body not determined. Cercaria Type XVI (Figures 16 and 20) A single Ferrissia fragilis collected at BHF was found to harbor infection with a small xiphidiocercaria of the ornatae group. Description: Body 108-150(130) long by 58-72(64) wide. Tail with finfold, 90- 122(108) long by 18-23(20) wide. Oral sucker 32-35(33) long by 30-35(32) wide. Acetabulum 18-23(21) long by 21-23(22) wide. Stylet shouldered 16-21(19) long. Five pairs of penetration glands and ceca extend to mid-acetabular level. Excretory bladder cellular and Y-shaped. Flame cell formula or number not determined. Cercaria type XVI may be the larva of the frog lung fluke Haemaetoloechus brev- iplexus. In size of body and tail it appears intermediate between that noted by Schell (1965) for H. breviplexus larvae develop- ing in experimentally infected Gyraulus similaris in Idaho, and that reported by Underwood and Dronen (1977) for those in an unknown species of experimentally infected Ferrissia from Texas. All other 80 Tulane Studies in Zoology and Botany Vol. 21 measurements, however, agree with those of Schell. Cercaria Type XVII (Figures 17 and 21) Heavily pigmented amphistome cercar- iae of the pigmentata group were found developing in Ferrissia fragilis from BPB, and Laevapex fuscus from RAM. Description: Body 256-352(281 ) long by 160-273(233) wide, with 12 pairs of lateral setulae arranged along anterior half. Tail 506-800(702) long by 48-68(57) wide. Oral sucker 51-72(61) long by 48-69(54) wide. Acetabulum subterminal, 58-83(73) long by 62-101(85) wide. Eyespots large and oval, located in anterior quarter of body. Extent of ceca not determined. Paired ex- cretory trunks filled with concretions and extending anteriorly with several coils un- til reaching level of oral sucker before turning posteriorly. Flame cell formula or number not determined. Cercariae main- tained under laboratory conditions emerged only in morning, within five minutes after exposure to light, and quick- ly encysted on green vegetation. Cercaria type XVII most nearly re- sembles the cercaria of Stichorchis sub- triquetrus (Rudolphi, 1814), a cecal para- site of the beaver. Castor canadensis Kuhl, 1820. Bennett and Humes (1939) reported on the pre-cercarial development of this species in experimentally infected lym- naeid snails, Lymnaea parva Lea, 1841, in Louisiana. Bennett and Allison (1958) later obtained cercariae from experiment- ally infected L. parva; however, these authors suggested that this snail was not a satisfactory host because of high mortality among infected individuals. Except for a slightly shorter tail, Cercar- ia type XVII fits the description for S. subtriquetrus given by Orloff (1941) for cercariae obtained from naturally infected Planorbis vortex (Linnaeus, 1758), Lym- naea ovata (Draparnaud, 1805), the oper- culate hydrobiid, Bithynia tentaculata (Linnaeus, 1758), and the terrestrial snail, Succinea putris (Linnaeus, 1758), in Rus- sia. Cercaria Type XVIII (Figures 18 and 24) A single Ferrissia fragilis from HIP was found to harbor infection with a longifur- cocercous, pharyngeate, distome cercar- ia. Description: Body 150-184(166) long by 48-62(56) wide, with two pairs of lateral setulae, one pair near mouth and another papillate pair at level of excretory blad- der. Anterior body to level of mid-oral sucker covered with small, dense spines. Tail stem 240-316(297) long by 128- 172(160) wide, with two groups of lateral setulae, two pairs near junction with body and four pairs near bifurcation. Furcae 208-268(243) long, with excretory pores opening on anterior surface of mid-length. Oral sucker 40-44(42) long by 28-35(30) wide. Acetabulum 28-32(30) long by 30- 35(33) wide, with several alternating cir- cles of small, blunt concentrically placed spines. Extent of ceca not determined. Number and arrangement of penetration glands not determined; however, one pair pre-acetabular. Genital primordium be- tween acetabulum and excretory bladder. Nine pairs of flame cells in body and three pairs in tail. Tegument on tail stem at bi- furcation formed into loose, bladder-like "skirt" continuous with furcae. Presence of three pairs of flame cells in the tail stem seems to be a deviation from a maximum of two pairs noted for other cercariae of this type (Yamaguti, 1975). Since only nine pairs were observed in the body, perhaps one pair in the tail may have originally been derived from the body complement. Cercaria Type XIX (Figures 19 and 24) A single Ferrissia fragilis from HIP was found to harbor infection with a longifur- cocercous, pharyngeate, distome cercar- ia. Description: Body 148-204(170) long bv 56-96(76) wide. Tail stem 248-284(256) long by 80-101(90) wide, with 10 pairs of lateral setulae. Furcae 208-220(213) long by 22-32(27) wide, with excretory pores opening on posterior surface. Oral sucker No. 2 Limpet Snails 81 o c o •^ 30 O 20 ^ 10 C 0 20 o '^ I 0 5 0 I 5 I 0 5 0 O C 0) O c 100 80 o ^0 — 40 "cS 20 3 ° Q_IOO O 80 Q. 60 \^ 40 ° 20 0 SMALL *TYPE II (F) ^TYPE IV (F) ^TYPE VKF) Q TYPE XIV(F) MEDIUM wNxxxxxxvvK LARGE -i>-TYPE III (HJ -o TYPE IV (H) * TYPE V(H) ♦ TYPE VI (H ) — TYPE VII (F) o TYPE XIII (F) e TYPE XV(F) « TYPE XVI(F) *\ / V ^ Q 6 ° ^--o-.o— 0--0 ©— e H. excentricus 60 • 50 (A 40 "D.E 30 E 20 y c 1 0 iJ«J 0 1 0 3E o _ 8 in 6 • y^ o . 4 2 Co) a 0 month Figure 20. Monthly incidence of infection, population structure, and index of relative abundance for (H.) Hebetancylus excentricus and {¥.) Ferrissiafragilis at Ben Hur Experimental Farm. Small, medium, and large refer to size classes of limpets. Cercarial type and limpet host combinations are indicated by various symbols. Relative abundance is indicated by circle with unbroken line. 82 Tulane Studies in Zoology and Botany Vol. 21 SMALL + TYPE I (F.) ♦ TYPE VI (H.) ■ TYPE XII (H.) — MEDIUM «»«mLARGE ©TYPE II (F) o TYPEIV(H) -^TYPEVI(F) -TYPE VII (F) c TYPE XV(F) ®TYPEXVII(F) JASONDJFMAMJJASON Month 50 (/> 40 _ C 20 ^ a^iB 10 O c 0 0) (0 503 t 4U ^ 30 if) 20 0 "^ 10 C 0 0 a Figure 2L Monthly incidence of infection, population structure, and index of relative abundance for (H.) Hebetancylus excenlricus and (F. ) Ferrissia fragilis at Beaver Pond Branch. No. 2 Limpet Snails 83 ■ SMALL— MEDIUM «.xv LARGE o TYPE I (L) + TYPE I (F.) >^ TYPE III (L.) oTYPEIV(H.) e TYPE VI (R) o TYPE VI ( L) • TYPE VII (F) »TYPE VIII (F) ^ TYPE IX (L.) « TYPE X I (L) o I 2 0) 8 o C 4 c O 0 0) ■o o ? o O) 1 c C 0 ■ ■^ ■ ^^ 100 80 60 40 o 20 \ o 0 100 8 0 c o 60 ■ ^IM 40 ^^ (U 20 3 0 a 100 o 80 a 60 40 20 0 8 — e *—-»--©■■ >-i)^-^ H. excentricus II illl g jliulm F_ f rgqilis — I I 1 I JASONDJFMAMJJA 60 50 40 30 20 I 0 0 20 I 6 I 2 8 4 0 20 I 6 I 2 H 8 4 0 CO c CD E in (D a 0) o o o o o c month Figure 22. Monthly incidence of infection, population structure, and index of relative abundance for (L.) Laevapex fuscus, (H.) Hebetancylus excentricus, and (F.) Ferrissia fragilis at Ramah. 84 Tulane Studies in Zoology and Botany Vol. 21 39-5 1 (45 ) long by 30-39(35 ) wide. Pharynx 12-15(14) long by 14-17(16) wide. Ace- tabulum 48-57(52) long by 46-62(5 1 ) wide, with six circles of small, blunt spines around inner rim. Extent of ceca not de- termined. Two pairs of penetration glands, one pre- and the other post- acetabular. Ten pairs of flame cells in body and two pairs in tail. DISCUSSION Ecology of Southeastern Louisiana Ancylids Although Ferrissia fragilis accounted for only 15% of the total ancylids collected and examined, it occurred at all sampled habitats. In contrast, Laevapex fuscus, which comprised 45% of the ancylids studied, was restricted to lentic habitats, and thus displayed greater ecological specialization. Hebetancylus excentricus was intermediate in ecological specificity because, although occurring at all habi- tats, it was not as successful, in terms of abundance, as F. fragilis at lotic BPB or L. fuscus at lentic SOR. McMahon (1976) stated this relationship when he noted, "representative species of these three limpet genera from the progressively less euryoecic series: Ferrissia-Hebetancylus- Laevapex." Bivoltine and trivoltine reproductive patterns with overlapping generations, as exhibited by Laevapex fuscus and Hebet- ancylus excentricus, partially account for their predominance over Ferrissia fragilis at most habitats. Although McMahon (1976) noted a trivoltine pattern for a Texas L. fuscus population, we found no more than a bivoltine for southeastern Louisiana populations. Even at eutrophic SOR the reproduction-recruitment cycle occurred only twice during the 14-month survey. Role of Southeastern Louisiana Ancylids in Trematode Ecology Southeastern Louisiana ancylids play a major role in trematode life cycles be- cause collectively they host at least 19 cer- carial species, which in turn utilize five classes of vertebrates as definitive hosts. Expectedly, the ecology of trematode intramolluscan stages was closely related to the ecology of the limpet hosts. No- where was this more evident than BHF (Figure 20), where the bivoltine reproduc- tion of Hebetancylus excentricus was re- flected in two cycles of cercarial produc- tion. The winter discontinuity in cercarial production reflected an interim between limpet cycles. Seasonality of those cercar- iae developing in Ferrissia fragilis did not demonstrate biannual patterns. Rather, individual trematode species were sea- sonal with discrete annual periods of cer- carial production. At BPB (Figure 21) the annual genera- tion of each limpet species is reflected in an annual generation of cercaria produc- tion. Cercaria type XVII (probably Stich- orchis subtriquetrus) from this locality demonstrated a pronounced seasonality, which must also relate to the life history of the beaver. Lowery (1974) noted that, al- though little is known about reproduction in Louisiana beaver, young are thought to be born in April .or May. If true, June cercarial emergence and encystment on aquatic vegetation may be correlated with feeding habits of recently weaned beaver, and thereby insured infection of a pre- sumably more susceptible individual. A similar seasonal correlation between cercarial emergence and life history of de- finitive host may exist for Cercaria type VII {Spirorchis scripta) and the turtle, Chrysemys scripta (Schoepff, 1793). Cagle (1950) reported egg hatching in Louisiana C scripta to occur from early July to early September. Peak cercarial emergence in July probably insures infection of young, susceptible individuals, not previously ex- posed or harboring a current infection with this blood fluke. Differences in population biology of Laevapex fuscus at RAM and SOR were earlier attributed to putative differences in primary productivity between these habitats. As shown in Figures 22 and 23, these differences are in turn reflected in No. 2 Limpet Snails 85 o 0) o c 0) o c c o \ c o o 0) 0 0 0 0 0 9 6 3 0 o o a o SMALL MEDIUM wvmsmmLARGE oTYPEI(L) »TYPEII(L.) ©TYPE 11(F) -TYPEIII(L) oTYPEIlKH.) =^TYPEIV(L) o TYPE IV (H.) o TYPE VI (L.) co. TYPE X (L ) C3 I I I I I I I I I I r JJASONDJFMAMJJ CO c 100 C 80 CD 60 E 40 20 LO 0 ^n 1 0 0) 8 a 6 4 "D 0) 2 0 o 1 0 ^^^ 0.8 o O.b o 0.4 0.2 o 0.0 c month Figure 23. Monthly incidence of infection, population structure, and index of relative abundance for (L.) Laevapexfuscus, (H.) Hebetancylus excentricus and (F.) Ferrissia fragilis at Sorrento. 86 Tulane Studies in Zoology and Botany Vol. 21 2 5 20 o ^ 1 5 o 1 0 0) 5 o c c o U 0 -^ 2 5 ■D O 20 oi> 1 5 c c 1 0 5 0 1 00 R 0 c o bO ■ ^^ 40 ^^ (U 20 3 0 a 00 o 8 0 a 60 o 40 o 2 0 0 — SMALL — MEDIUM..LARGE « TYPE ll (F.) « TYPE III (H.) o TYPE IV (H.) o TYPE XIII (F) ® TYPEXIV(F) t TYPE XVIll (F) || TYPE XIX (F.) o t H excentricus F f rggilis r \ \ " ^ 1 I TTTT7 JJASONDJFMAMJJA 60 50 (/) 30 ^ ^ 20 O \ 10 60 mm. For the relative prey size char- acter, the estimated average absolute prey size was divided by the average standard length for the species sample to give a proportion. This was coded according to the scheme: (1) < 0.05, (2) 0.10, (3) 0.15, (4) 0.20, (5) 0.25, and (6) > 0.30. Mathematical Methods Simple linear correlations.— T'wo types of correlation coefficient were determined between the species means of all charac- ters for all 33 common species. Pearson product-moment correlation coefficients were calculated using the Biomedical Computer Program BMD PIM. Spear- man rank-order correlation coefficients were calculated with procedure Spearman of the Statistical Analysis System, North Carolina State University. The model un- derlying the former type requires that the two variables being correlated each exhib- it a normal distribution. This was not the case for a number of the characters in this study; therefore, the latter type coeffi- cient was also calculated because the model for it does not require that the vari- ables fit any particular distribution. The Bonferonni technique was used in assigning significance to the correlation coefficients. This is a very conservative procedure which allows one to know the probability of making a family Type I er- ror. A Type I error is made if a hypothesis is rejected when it is true (in this case, the recognition of correlations as significant which are not). All statistical tests which employ the same set of observations be- long to the same family. Thus, in the cor- relation analysis described here, the 55 correlations of each morphological char- acter with every other character is a family of tests because the same set of observa- tions is used 55 times. In order for the probability of making a Type I error to equal P< 0.05 in an entire family of tests, the Bonferonni technique requires that each individual test be conducted at a probability of 0.05 / n where n is the num- ber of tests in the family. For the present study, 0.05 / 55 is approximately equal to 0.001. Therefore, Pearson correlation co- efficients were considered to be significant if they were greater than or equal to 0.546 (df = 31, P < 0.001) and Spearman corre- lation coefficients, if they were greater than or equal to 0.571 (df = 31, P <0.001). This technique controls Type I errors because hypotheses of no relation- ship between two variables are rejected only at a very small value of P, i.e., when correlations are quite likely significant. Of course, the probability of making a Type II error (failure to reject a hypothesis when it is false, i.e., in this case, failure to recognize a significant correlation) is, as always, inversely related to the probabil- ity of making a Type I error. Factor Analy sis. -¥diC\or analysis is a multivariate statistical technique that generates a small number of hypothetical factors which reproduce the linear corre- iations between variables in the original data set. Each factor so generated is a linear combination of covarying portions of the original variables. Interpretation of the hypothetical factors is facilitated by orthogonal rotation using the varimax cri- terion. Orthogonal rotation of the factors makes each factor statistically indepen- dent; i.e., the factors are not correlated. The varimax criterion maximizes the vari- ance of each factor by causing as many factor loadings (i.e., coefficients relating the original variables to factors) as pos- sible to tend toward zero or one. Thus each of the originally measured variables tends to make either a large or a negligible contribution to each of the hypothetical factors. The chief value of factor analysis is as an aid in the recognition of overall trends in the variation and covariation of a large number of variables. Additionally, because factor analysis also indicates the proportion of the total pattern of variation accounted for by each factor, it is of value in understanding the relative strength of trends in variation. 98 Tulane Studies in Zoology and Botany Vol. 21 Factor analysis was performed in this study using Biomedical Computer Pro- gram BMD 08M. Orthogonal rotation us- ing the varimax criterion was used. In ac- cordance with the suggestion of Harman (1967), only factors with eigenvalues greater than one were considered. This prevents "overfactorization," i.e., the generation of hypothetical constructs that are explaining trends in error variances rather than in the pattern of variation in the original variables. RESULTS AND DISCUSSION In the first three sections below the re- sults of the three major types of analyses are presented individually. Each analysis by itself provides some useful information in the understanding of fish morphology. In a fourth section, then, all these results are integrated with information from the literature in order to provide a functional, ecological interpretation of each of the morphological features studied. Simple Linear Correlations. — Extensive correlations between characters were de- termined. Using the previously defined criterion for significance, 500 of the 3080 Pearson product-moment coefficients were significant as were 494 of the Spear- man rank-order coefficients. However, 200 of these differed between types of co- efficient. I tested whether part of the dif- ference in result might have been due to differences in character distributions. The total set of characters was divided into two groups: those in which the values showed continuous variation (e.g., proportional measurements) and those in which the values were discrete (e.g., fin ray counts and qualitative characters with integer character state codes). Chi-square analys- is was then used to test the null hypothesis that the two types of correlation coeffi- cient acted at random with regard to these different types of characters. In both cases the hypothesis of randomness was reject- ed. Of correlations unique to the Spear- man rank-order correlation coefficient, significantly more (P < 0.005) involved characters with discrete character state distributions than one should expect at random. Of correlations unique to the Pearson product-moment correlation co- efficient, significantly more (P< 0.05) in- volved characters with continuous varia- tion. However, visual examination of fre- quency plots of character state values in- dicated that characters with discrete char- acter state distributions did not as a class necessarily show the greatest deviation from normal distributions. As a conse- quence of all this, I have somewhat arbi- trarily listed below only the approximately 400 correlations determined to be signifi- cant by both types of coefficient. If the r in both cases was greater than 0.717 so that the coefficient of determination was greater than 50%, the correlation has been marked with an asterisk. Some of these many correlations are discussed in- dividually below in the interpretation of morphological characters. Additional dis- cussion of the methodological problem of which correlation coefficient is appropri- ate for use is found in Gatz (1975). 1. Standard length - Positive correla- tions: number of gill rakers. Negative cor- relations: *relative peduncle length; rela- tive pectoral fin area. 2. Pigmentation pattern - No signifi- cant correlations. 3. Completeness of lateral line - Posi- tive correlations: position of lateral line. No significant negative correlations. 4. Position of lateral line - Positive correlations: completeness of lateral line; aspect ratio of caudal fin; caudal span / body depth; number of caudal fin rays; position of pectoral fin; hypertrophy of pharyngeal teeth; percentage of red muscle. Negative correlations: pelvic fin aspect ratio; presence of jaw teeth; shape of jaw teeth; fine structure of gill rakers. 5. Relative head length - Positive cor- relations: *relative body depth; position of pectoral fin; position of pelvic fin; *width of mouth; *height of mouth; *presence of jaw teeth; *fine structure of gill rakers; number of pyloric caeca. Nega- tive correlations: *caudal span / body iNo. 2 Ecological Morphology of Fishes 99 depth; * distance of pectoral fin from CG; position of dorsal fin; position of the mouth; hypertrophy of pharyngeal teeth. 6. Flatness index — Positive correla- tions: * relative body depth; *pelvic fin length, * relative eye size; protrusibility of the mouth. No significant negative corre- lations. 7. Relative body depth - Positive cor- relations: ^relative head length; *flatness index; *pelvic fin length; *distance of pel- vic fin from CG; relative eye size; protrus- ibility of the mouth. Negative correla- tions: *caudal span / body depth; distance of pectoral fin from CG. 8. Index of trunk shape - No signifi- cant correlations. 9. Relative peduncle length - No sig- nificant positive correlations. Negative correlations: ^standard length; number of gill rakers. 10. Caudal peduncle flatness index — Positive correlations: relative size of fore- brain. No significant negative correla- tions. 1 1 . Aspect ratio of caudal fin - Positive correlations: position of lateral line; num- ber of caudal fin rays; number of pectoral fin rays; position of pelvic fin; position of dorsal fin; hypertrophy of pharyngeal teeth; percentage of red muscle. Negative correlations: pelvic fin aspect ratio; posi- tion of jaw teeth. 12. Caudal span / body depth - Posi- tive'correlations: position of lateral line; number of caudal fin rays ; distance of pec- toral fin from CG; * position of pelvic fin, number of pelvic fin rays; *position of dorsal fin; * hypertrophy of pharyngeal teeth. Negative correlations: *relative head length; *relative body depth; pec- toral fin length; position of pectoral fin; *pelvic fin length; *distance of pelvic fin from CG; * pelvic fin shape; * presence of jaw teeth; *fine structure of gill rakers. 13. Number of caudal fin rays - Posi- tive correlations: position of lateral line; aspect ratio of caudal fin; caudal span / body depth; *position of pelvic fin; *num- ber of pelvic fin rays; hypertrophy of pharyngeal teeth; percentage of red mus- cle. Negative correlations: position of pec- toral fin; aspect ratio of pelvic fin; pelvic fin shape; relative size of optic lobes. 14. Pectoral fin length - Positive corre- lations: position of pectoral fin; *pelvic fin length; * distance of pelvic fin from CG; *pelvic fin shape; relative size of eye. Negative correlations: caudal span / body depth; pelvic fin position; *number of pel- vic fin rays. 15. Aspect ratio of the pectoral fin - Positive correlations: pectoral fin shape. No significant negative correlations. 16. Relative pectoral fin area - No sig- nificant positive correlations. Negative correlations: standard length; *relative length of swim bladder. 17. Distance of pectoral fin from center of gravity - Positive correlations: caudal span / body depth; position of dorsal fin; hypertrophy of pharyngeal teeth. Nega- tive correlations: *relative head length; *relative body depth; position of pectoral fin; *width of the mouth; presence of jaw teeth ; number of pyloric caeca. 18. Pectoral fin shape — Positive corre- lations: pectoral fin aspect ratio. No sig- nificant negative correlations. 19. Position of the pectoral fin - Posi- tive correlations: relative head length; caudal span / body depth; pectoral fin length; position of lateral line; pelvic fin length; pelvic fin aspect ratio; distance of pelvic fin from CG; pelvic fin shape; fine structure of gill rakers. Negative correla- tions: distance of pectoral fin from CG; number of caudal fin rays; *position of pelvic fin; *number of pelvic fin rays; per- centage of red muscle. 20. Number of pectoral fin rays - Posi- tive correlations: aspect ratio of caudal fin; position of the dorsal fin; hypertrophy of pharyngeal teeth. No significant nega- tive correlations. 21. Pelvic fin length - Positive correla- tions: *flatness index; *relative body depth; *pectoral fin length; position of the pectoral fin; *distance of the pelvic fin from CG; pelvic fin shape; relative eye size; fine structure of gill rakers. Negative correlations: *caudal span / body depth; 100 Tulane Studies in Zoology and Botany Vol. 21 position of the pelvic fin ; number of pelvic fin rays; position of the dorsal fin. 22. Aspect ratio of the pelvic fin - Posi- tive correlations: position of the pectoral fin; distance of pelvic fin from CG; pelvic fin shape; presence of jaw teeth; shape of jaw teeth; fine structure of gill rakers; * relative size of optic lobes. Negative cor- relations: position of the lateral line; as- pect ratio of caudal fin; number of caudal fin rays; *position of pelvic fin; number of pelvic fin rays; hypertrophy of pharyngeal teeth; * percentage of red muscle; relative size of facial lobes. 23. Relative pelvic fin area - Positive correlations: shape of jaw teeth. No signif- icant negative correlations. 24. Distance of pelvic fin from center of gravity — Positive correlations: relative head length; *relative body depth; *pec- toral fin length; position of pectoral fin; * pelvic fin length, pelvic fin aspect ratio; * pelvic fin shape; *fine structure of gill rakers; number of pyloric caeca. Negative correlations: *caudal span / body depth; *position of pelvic fin; *number of pelvic fin rays; * position of dorsal fin. 25. Pelvic fin shape - Positive correla- tions: *pectoral fin length; position of pec- toral fin; pelvic fin length; aspect ratio of pelvic fin; * distance of pelvic fin from CG; relative eye size; presence of jaw teeth; *fine structure of gill rakers; *relative size of optic lobes. Negative correlations; *caudal span / body depth; number of caudal fin rays; *number of pelvic fin rays: *position of pelvic fin; *position of dorsal fin; hypertrophy of pharyngeal teeth; percentage of red muscle; relative size of cerebellum; relative size of facial lobes. 26. Position of pelvic fin — Positive cor- relations: position of lateral line; aspect ratio of caudal fin; *caudal span / body depth; *number of pelvic fin rays; *posi- tion of dorsal fin; *hypertrophy of phar- yngeal teeth; percentage of red muscle; relative size of facial lobes. Negative cor- relations: pectoral fin length; *position of pectoral fin; * aspect ratio of pelvic fin; * distance of pelvic fin from CG; * pelvic fin shape; presence of jaw teeth; shape of jaw teeth; relative size of optic lobes; pel- vic fin length; *fine structure of gill rakers; number of pyloric caeca. 27. Number of pelvic fin rays - Positive correlations: caudal span / body depth; * number of caudal fin rays; * position of pelvic fin; position of dorsal fin; hyper- trophy of pharyngeal teeth; relative size of facial lobes. Negative correlations: *pec- toral fin length; * position of pectoral fin; pelvic fin length; aspect ratio of pelvic fin; * distance of pelvic fin from CG; * pelvic fin shape; relative eye size; presence of jaw teeth; fine structure of gill rakers; number of pyloric caeca; relative size of optic lobes. 28. Position of dorsal fin - Positive correlations: aspect ratio of caudal fin; caudal span / body depth; distance of pec- toral fin from CG; number of pectoral fin rays; * position of pelvic fin; number of pelvic fin rays; * hypertrophy of pharyn- geal teeth; percentage of red muscle. Negative correlations: relative head length; position of pectoral fin; pelvic fin length; * distance of pelvic fin from CG; *pelvic fin shape; *presence of jaw teeth; shape of jaw teeth; *fine structures of gill rakers; number of pyloric caeca. 29. Relative eye size - Positive correla- tions: *flatness index; relative body depth; pectoral fin length; pelvic fin length; protrusibility of the mouth. Nega- tive correlations: number of pelvic fin rays; relative size of cerebellum. 30. Position of the eyes - No significant correlations. 31. Eye pigmentation — No significant positive correlations. Negative correla- tions: percentage of red muscle. 32. Position of the mouth - Positive correlations: *orientation of the mouth; * relative size of facial lobes. Negative cor- relations: relative head length; width of the mouth; *height of the mouth. 33. Orientation of the mouth - Positive correlations: *position of the mouth. Negative correlations: *height of the mouth. 34. Relative width of the mouth - Posi- tive correlations: * relative head length; No. 2 Ecological Morphology of Fishes 101 *height of the mouth; presence of jaw teeth. Negative correlations: *distance of pectoral fin from CG; position of the mouth; hypertrophy of pharyngeal teeth. 35. Relative height of the mouth — Posi- tive correlations: *relative head length; *width of the mouth. Negative correla- tions: *position of the mouth; orientation of the mouth. 36. Protriisibility of the mouth - Posi- tive correlations: *flatness index; relative body depth; relative eye size; relative swim bladder volume. No significant neg- ative correlations. 37. Number of barbels — Positive cor- relations: * relative size of forebrain; *re- lative size of cerebellum. No significant negative correlations. 38. Number of branchiostegal rays - Positive correlations: presence of jaw teeth; *shape of jaw teeth. Negative cor- relations: *hypertrophy of pharyngeal teeth; percentage of red muscle. 39. Presence of jaw teeth — Positive correlations: *relative head length; aspect ratio of pelvic fin; pelvic fin shape; width of the mouth; number of branchiostegal rays; *shape of jaw teeth; fine structure of gill rakers; number of pyloric caeca; rela- tive size of optic lobes. Negative correla- tions: position of lateral line; *caudal span / body depth; distance of pectoral fin from CG; position of the pelvic fin; number of pelvic fin rays; *position of dorsal fin; * hypertrophy of pharyngeal teeth; per- centage of red muscle. 40. Shape of jaw teeth — Positive corre- lations: relative head length; aspect ratio of pelvic fin relative pelvic fin area; *num- ber of branchiostegal rays; *presence of jaw teeth; fine structure of gill rakers; re- lative size of optic lobes. Negative correla- tions: position of lateral line; aspect ratio of caudal fin; position of pelvic fin; posi- tion of dorsal fin; * hypertrophy of phar- yngeal teeth; percentage of red muscle. 41. Hypertrophy of pharyngeal teeth - Positive correlations: position of lateral line; aspect ratio of caudal fin; *caudal span / body depth; number of caudal fin rays; distance of pectoral fin from CG; number of pectoral fin rays; *position of pelvic fin; numberof pelvic fin rays; *posi- tion of dorasl fin; percentage of red mus- cle. Negative correlations: relative head length; aspect ratio of pelvic fin; pelvic fin shape; width of the mouth; *number of branchiostegal rays; *presence of jaw teeth; *shape of jaw teeth; fine structure of gill rakers; relative size of optic lobes. 42. Shape of pharyngeal teeth - No sig- nificant correlations. 43. Number of gill rakers — Positive correlations: standard length; fine struc- ture of gill rakers. Negative correlations: relative peduncle length. 44. Shape of gill rakers - No significant correlations. 45. Fine structure of gill rakers - Posi- tive correlations: relative head length; position of pectoral fin; distance of pelvic fin from CG; pelvic fin length; pelvic fin aspect ratio; pelvic fin shape; presence of jaw teeth; shape of jaw teeth; number of gill rakers; number of pyloric caeca; rela- tive size of optic lobes. Negative correla- tions: position of lateral line; caudal span / body depth; position of pelvic fin; number of pelvic fin rays; position of dorsal fin; hypertrophy of pharyngeal teeth; percen- tage of red muscle ; relative size of cerebel- lum; relative size of facial lobes. 46. Relative volume of the swim bladder — Positive correlations: *relative length of swim bladder. No significant negative cor- relations. 47. Relative length of the swim bladder — Positive correlations: * relative volume of swim bladder. Negative correlations: * relative pectoral fin area. 48. Relative gut length - No significant correlations. 49. Number of pyloric caeca - Positive correlations: relative head length; dis- tance of pelvic fin from CG; presence of jaw teeth; fine structure of gill rakers. Negative correlations: distance of pector- al fin from CG; pelvic fin position; num- ber of pelvic fin rays; position of dorsal fin. 50. Percentage of red muscle in the peduncle - Positive correlations: position 102 Tulane Studies in Zoology and Botany Vol. 21 of lateral line; aspect ratio of caudal fin; number of caudal fin rays; position of pel- vic fin; position of dorsal fin; hypertrophy of pharyngeal teeth. Negative correla- tions: position of pectoral fin; *aspect ra- tio of pelvic fin; pelvic fin shape; eye pig- mentation; number of branchiostegal rays; presence of jaw teeth; shape of jaw teeth; fine structure of gill rakers; relative size of optic lobes. 5 1 . Relative size of the forebrain — Pos- itive correlations: caudal peduncle flat- ness index. No significant negative corre- lations. 52. Relative size of the optic lobes - Positive correlations: aspect ratio of pelvic fin; *pelvic fin shape; presence of jaw teeth; shape of jaw teeth; fine structure of gill rakers. Negative correlations: number of caudal fin rays; *position of pelvic fin; number of pelvic fin rays; hypertrophy of pharyngeal teeth; percentage of red mus- cle; relative size of cerebellum; relative size of facial lobes. 53. Relative size of the cerebellum — Positive correlations: * number of barbels; *relative size of vagal lobes; *relative size of facial lobes. Negative correlations: pel- vic fin shape; relative eye size; fine struc- ture of gill rakers; relative size of optic lobes. 54. Relative size of the vagal lobes - Positive correlations: *relative size of cer- ebellum. No significant negative correla- tions. 55. Relative size of the facial lobes - Positive correlations: position of pelvic fin; number of pelvic fin rays; *position of the mouth; relative size of cerebellum. Negative correlations: pelvic fin shape; aspect ratio of pelvic fin; fine structure of gill rakers; relative size of optic lobes. 56. Relative size of the acoustic tuber- cles - No significant correlations. Factor Analysis. — Factor analysis of morphological characters was useful in pointing out some relationships not seen in the linear correlation analyses. More- over, its basic property of ordering the importance of the covarying characters resulted in a ranking of trends which would not otherwise have been possible. The first three factors identify the three primary trends in the ecological differen- tiation among the fishes studied. These factors combine variables which relate to, respectively, technique of predation, maneuverability and utilization of habi- tats, and vertical zonation. These trends and all other trends defined by the factor analysis together with the associated re- sultant separations of species are described below. Altogether, the factor analysis in- dicated nine significant multivariate trends in the covariation of characters which joint- ly accounted for 79% of the total variance. Factor 1 , the major trend accounting for 31% of the character variance, indicated an association of several characters which relate to feeding behavior and technique. It indicates a positive association among having a high number of branchiostegal rays and the presence of jaw teeth (es- pecially canines) and having many small teeth on the gill rakers; and a negative association of these characters with the presence of hypertrophied pharyngeal teeth and much red muscle in the oedun- cle. I interpret this to mean that the major morphological trend in ecological separa- tion of the fishes studied has been in dif- ferentiation in feeding strategy between "iie-and-wait" biting predators (high scores on factor 1) and cruising suction feeders (low scores on factor 1). The or- dination of families which results from consideration of this factor goes from Esocidae (pikes) with high factor scores through Percidae (perches), Ictaluridae (catfishes), and Centrarchidae (sunfishes) to Cyprinidae (minnows) with low scores (Fig. 1). Factor 2 (12% variance) indicated the association among a number of characters relating to habitat separation by differ- ences in body shape and proportions (see section on interpretation of morphologi- cal characters below). This factor associat- ed a high ratio of caudal span to body depth, a large number of pelvic fin rays, and the location of both pelvic and dorsal fins entirely posterior to the center of No. 2 Ecological Morphology of Fishes 103 gravity of the fish with low values for both the flatness index and relative body depth, short paired fins, pelvic fins which are rounded and located near the center of gravity of the fish and small relative eye size. Thus this factor separated the pick- erel-like morphology at one extreme and the sunfish-like morphology at the other. Figure 2 shows the factor scores for the species on this factor. Factor 3 (10'"^^ variance) identified characters associated with a benthic habit. Thus it associated large pectoral fin area with a small swim bladder and dorsally displaced eye position. Darters, naturally, showed especially high scores on this character (Fig. 3) and all suckers except Erimyzon oblongus showed somewhat positive scores. All other fishes showing positive scores on this factor I also infer to have a benthic preference (Fig. 3). Factor 4 (7'^f variance) indicated the next most important trend in character variation among the fishes studied was that of being a small insectivore (see be- low). The major association shown by this factor was between small size and having a short relative gut length. To a lesser ex- tent, these morphological features were also associated with presence of hooked pharyngeal teeth, few gill rakers, a supraterminal or terminal mouth which opens anteriorly, a long relative peduncle length and a large relative eye size. Fishes with high scores included both mid-water minnows and darters, Gambusia affinis and Aphredoderus sayanus; whereas sun- fishes, pickerels, catfishes, and especially suckers had low scores (Fig. 4). Factor 5 (6% variance) and Factor 6 (4% variance) each indicated associations of characters which differentiate two fam- ilies. Factor 5 identified the relationship between having barbels, having a flat- tened peduncle and having few pectoral fin rays. All these characters are special- izations of catfishes (Fig. 5). Factor 6 iden- tified the association of large mouth size with large head size and having rounded pectoral fins placed near the center of gravity of the fish. Thus it identified a suite 3-1- -■ Enig -- Eame ai < in O m z -I < X a. u. O > X a O I- z cc LU uj u Q lU CC liJ O < a >- X o < UJ cc o z 2 A (A < ILI CC O z -- Efia / 1 --Upyg / -■ Nins / Efus -- Cmac •- Eglo 0--Lgul Eolm Pfia Pcra Msal Icat Lcya A say Ineb Lmac Pnig Ipun Laur Lgib Gaff CO > < cc -I < a trt LU cc 1- LU (/) ^ o < CC X EobI Nard Ccom Nalb Pore Cfun -■ Ncry / -)- Mery / + Nana / Nhud Satr \ Mani Nchi \ + Mpap Nlep Npro Nalt Mrob O z o Ncer Hhyp LU LU t- 5 < o z < cc CO u. o cc liJ CO s z FIGURE 1. Ordination of species by mean factor scores on Factor 1 of the factor analysis of mor- phology. See text for interpretation. Abbrevia- tions for species names are: Asay = Aphredoder- us say anus; Ccom = Catostomus commersoni; Cfun = Clinostomus funduloides; Cmac = Cen- trarchus macropterus; Eame = Esox americanus; Efla = Etheostoma flabellare; Efus = Etheo- stomafusiforme; Eglo = Enneacanthus gloriosus; Enig = Esox niger\ EobI = Erimyzon oblongus; Eolm = Etheostoma olmstedi; Gaff = Gambusia affinis; Hhyp = Hybopsis hypsinotus; Icat = Icta- lurus catus; Ineb = Ictalurus nebulosus; Ipun = Ictalurus punctatus; Laur = Lepomis auritus; Lcya = Lepomis cyanellus; Lgib = Lepomis gib- bosus; Lgul = Lepomis gulosus; Lmac = Lepo- mis macrochirus; Mani = Moxostoma anisurum; Mery = Moxostoma erythrurum; Mpap = Moxo- stoma pappillosum; Mrob = Moxostoma robus- tum; Msal = Micropterus salmoides; Nalb = No- tropis alborus; Nalt = Notropis altipinnis; Nana = Notropis analostanus ;^ard = Notropis ardens; Ncer = Notropis cerasinus; Nchi = Notropis chil- iticus; Ncry = Notemigonus chrysoieucas; Nhud = Notropis hudsonius; Nins = Noturus insignis; Nlep = Nocomis leptocephalus; Npro = Notropis procne; Pcra = Percina crassa; PfIa = Perca flavescens; Pnig = Pomoxis nigromaculatus; Pore = Phoxinus oreas; Satr = Semotilus atromacula- tus; and Upyg = Umbra pygmaea. 104 Tulane Studies in Zoology and Botany V( of characters which indicate ecological speciaHzations shared by most sunfishes and which differentiate them from the other fishes (Fig. 6). The remaining three factors, together accounting for only 9% of the variance, did not in fact demonstrate clear cut morphological patterns. Factor 7 identi- fied species which lack a lateral line canal (Fig. 7). Perhaps the most important point here is that although this character was assumed to indicate benthic habitat pre- ferences (see below), the factor analysis did not associate it with the other charac- ters in Factor 3. Factor 8 did not show any easily interpretable result but identified a weak association between small pelvic fin size and a low number of caudal fin rays (Fig. 8). Factor 9 identified species with extremely long, thin gill rakers (Fig. 9). There was also a tendency for a high index of trunk shape to be associated with this factor. Gut Content Analysis. The results of the gut content analyses for the 33 common species are presented in Table 1 according to the coded format described previously. Notably, no two species were found to have the same diet in both size and com- position. This result was obtained even with the relatively small sample size of individuals per species compared to most fish studies (e.g., Keast, 1966; Nilsson, 1960; Schwartz and Dutcher, 1962; al- though see also Carpenter, 1940) and the gross nature of the food categories. Di- etetically, perhaps the two most similar . UJ in N Ui UJ IT < UI 0 < -J 1 S 5x Z K UJ O. _l UJ a x' Ui >■ Q Q Z O - CO U) UJ « > UJ - z t- I- < < -• _J UJ U. (£ o (0 < o 5 t 2-- Enlg ■ ■ Eame ■• Nins 0 -- Upyg Satr Ineb Icat Nard I pun Nlep Npro Cfun Gaff Nchi Pore Ccom Nana Nalt — Mery Mpap Nalb Ncfy Ncer Nhud Mrob Hhyp EobI Man! Asay Efia Efus Msal -■ Pcra Pfla Eolm ■ Lcya Lgul Pnig Laur Cmac Lgib -■ Lmac -. Eglo O z < UI a. O z 5^ o '^ il o o o o FIGURE 2. Ordination of species by mean factor scores on Factor 2 of the factor analysis of mor- phology. See text for interpretation. Abbrevia- tions as in legend for Fig. 1. cc UI o a < —I m (0 u. O UJ o > 3-- ■■ Pcra ■ • Eolm ■ EfIa Efus 0--:- O z < UJ QC O ? i -1 ■ ■ Asay Nins Ccom ■•Hhyp Nlep Mpap Nhud Mery Mani Pfla Mrob Lcya Msal Nchi Nalb Cfun Ncer Upyg Ipun Icat Satr Npro ■■ Lgul EobI • Nard ■• Ineb ■■ Cmac Gaff Ncry I Nalt Pore Nana Eame Lmac Eglo Laur Lgib Pnig Enig O z CO < Ui oc u z UJ > UJ < ^ UJ u. < _J < oc -I o < I- CO UJ O a. a FIGURE 3. Ordination of species by mean factor scores on Factor 3 of the factor analysis of mor- phology. See text for interpretation. Abbrevia- tions as in legend for Fig. 1. No. Ecological Morphology of Fishes 105 sympatric species were Clinostomus fun- duloides and Semotilus atromaculatus . Both of these species fed exclusively on terrestrial insects, but Semotilus took larger prey items in both relative and ab- solute terms. Gambusia affinis and No- tropis ardens similarly differ only in prey size in this study, but were not found in the same stream. A factor analysis with orthogonal rota- tion was performed on the food data in an effort to define trends in types of prey eaten. Nine factors accounting for 80*^ of the total variance were identified. The major trend (Factor 1 - 23% total vari- ance) was one of prey size. Factor 1 separ- ated predators on small items from those on large items, especially fish. Factor 2 (18% total variance) identified the second most important trend in feeding to be one relating to vertical position of prey. It sep- arated bottom feeding as defined by the presence of ostracods, molluscs, filamen- tous algae and silt in the gut from surface feeding as defined primarily by eating ter- restrial insects. All remaining factors merely identified individual food cate- gories in order of their relative importance to numbers of species of fishes. Factor 3 (13% variance) identified the insects as the most important prey type and also in- dicated the tendency of fishes to eat either aquatic insects or terrestrial ones, but not both. Factor 4 (10% variance) related to + Gaff o z UJ m > < cc _i UJ UJ ^ c < u z o UJ _I LL o Q < ^ Z " V) z o z (A < UJ cc o ^ t 1 -- 0-- ■ Nalt Nard Asay Nalb Npro Nhud Satr Laur Lcya Nlep Icat Lgul -1 -- 2-- ■3-- ■ • Nins •• Ipun ■ EobI Cfun Pcra Nana Nchi Lmac Ncry Pnig Lgib Cmac Enig Upyg Efus Efia Ncer Eglo Hhyp Nins Pfia Eolm Msal Eame I neb Pore ■ Mery Mpap •■ Mani Ccom <' Mrob O z < UJ cc o (A >- < < OC O 1- UJ -I o z o o UJ 0. z 1- 3 O III (A o O Q. a cc UJ CD u. I S O H 3 UJ 3 O Z I s H (A o z UJ UJ > _i < -I UJ z z C3 UJ UJ cc Z > > UJ M t- < 1- H < UJ < < oc (C _l -J a o UJ UJ 3 z cc cc (A 4-- 3 -h Icat Ineb Ipun 2-- o z w < UJ cc u z 1 - - (J z Q m 0. — 1 ■ Cmac Hhyp Eglo Pnig Gaff Lgib Lmac CD U. O 0- -rNlep Nana Satr Ncry Lgul ^Mrob '"~~Nard Laur Nchi Pcra Nalt EobI Eolm Asay ^Lcya ^Efus /^Efla ^ Upap Pore Upyg PfIa Ncer Cfun Npro Mery Nalb If) OC iu i< Z U. -1- Msal Mani Ccom Nhud Eame Enig FIGURE 4. Ordination of species by mean factor scores on Factor 4 of the factor analysis of mor- phology. See text for interpretation. Abbrevia- tions as in legend for Fig. 1. FIGURE 5. Ordination of species by mean factor scores on Factor 5 of the factor analysis of mor- phology. See text for interpretation. Abbrevia- tions as in legend for Fig. 1. 106 Tulane Studies in Zoology and Botany Vol. 2: ship, could be demonstrated. Exceptions to this generalization are noted below where the characters are discussed seria- tim. For each character the significance assumed at the outset, support from the literature, and the results of this study are given. Superficial body and body shape charac- ters 1. Standard length was assumed to be an indicator of prey size. Numerous stud- ies of fish usually involving only a single species (e.g., Swynnerton and Worthing- ton, 1940; Lindstrom, 1955 and references cited therein; Nilsson, 1955, 1958; Thom- as, 1962; Hall et al., 1970) have shown prey size to increase as a function of in- crease in body size. I found absolute prey size to be positively correlated (r = 0.690, P < 0.001) with standard length as was predicted. Relative prey size was not so correlated (r = 0.309, 0.10 > P > 0.05). The correlation demonstrated here for this widely assumed relationship in ecol- ogical work (see Schoener and Gorman, 1968; Ashmole, 1968; Pianka, 1969; among others) is unusual in having been established for so broad a range of taxa. 2. Pigmentation pattern was assumed to indicate habitat and behavior according to the scheme of Nikolskii (1963). Nikol- skii indicated the following correlates: (1) ■ ■ Cmac >- < cc ^< Lil O U. Z £ o iE 3 LU Z a. O z to < u DC O - Gaff 2-- Icat Msal I pun Mrob ■ Upyg 1 -- Ineb ■- Ccom Mery Mani Nana Eolm Efus ■■ Satr ■ Nlep Pnig Nard Efia ■• Cfun Mpap Npro Pfia Ncry Lcya 0- - Nchi Lmac -- Lgul Laur ■ Cmac Lgib Enig - Pcra ■ ■ Ncer Nalt Eame Pore -1--"Nalb Nhud • Hhyp Nins ■ EobI Eglo • ■ Asay 2-- 3-- 2-|- PnIg EfIa Ncry 1-- ■• Nins Mery Gaff Upyg Mani ■ ■ Mrob Ncer Ccom Cfun Enig Nlep Eame Nalt EobI Nana Msal Eglo Hhyp Mpap ■■ Pcra Satr Nalb Nard Nchi Npro ■■ Lcya Pore Efus Lgul Asay Icat ■ - Nhud ■• Eolm •1 — Lmac Pf la •• Ineb Ipun ■ ■ Laur •■ Lgib 0--: z < LJ CC o (0 c Ul < cc o u. o LLI CL < o cc z z> ^ cc I- I- Q Q Z z iH FIGURE 8. Ordination of species by mean factor scores on Factor 8 of the factor analysis of mor- phology. See text for interpretation. Abbrevia- tions as in legend for Fig. 1 . FIGURE 9. Ordination of species by mean factor scores on Factor 9 of the factor analysis of mor- phology. See text for interpretation. Abbrevia- tions as in legend for Fig. 1. No. 2 Ecological Morphology of Fishes 107 eating copepods; Factor 5 (8%), diatoms; Factor 6 (6%), terrestrial invertebrates; Factor 7(5%), amphipods and isopods; Factor 8 (4%), filamentous algae; and Factor 9 (3%), crayfish. The results of the gut content analyses for the rare species according to decreas- ing order of importance of categories present are: Anguilla rostrata - empty; Umbra pygmaea - aquatic insects; No- tropis hudsonius - aquatic insects, silt; Moxostoma anisurum — aquatic insects, molluscs, ostracods, copepods, and silt; M. erythrurum - aquatic insects, ostra- cods, diatoms, silt and sand; M. pappillo- sum — ostracods, vascular plants, silt; M. rohustum - aquatic insects, sand; Ictalur- us catus - terrestrial insects, sand (gut nearly empty); /. punctatus - crayfish, aquatic insects, terrestrial insects, vascu- - Lgul 2 -- < _] DC < O a. O u. 1- O O ^£ t- U. Z O LU O LU O 5 z o < a. I- LL w o o z < UJ c o -- Lcya Msal Cfun Asay 1 -HSatr Laur Pfia Pnig Upyg Nlep Ineb Nchi Eame O-f-Ncer Nins Cmac Hhyp Gaff Icat Enig Ipun Nalt Eglo Mpap Nhud Efia Nard /Pcra Mani EobI Efus ^Lgib Ccom Pore Nana Nalb Mery 1 -t-Npro Mrob Ncry • • Eolm Lmac z (0 < UJ <£ u z o 2 UJ tsl v> < DC O UJ - O CC ^ K lar plants; Enneacanthus gloriosus - aquatic insects, terrestrial insects; and Percina crassa - aquatic insects and sand. These results were utilized below in the interpretation of the morphological char- acters. Interpretation of Morphological Char- acters.— ^The initial choice of all characters was based on some inferred functional and/or ecological significance. Although in most cases there was some precedent in the literature for the supposed association between character and function, the ex- perimental substantiation of any particu- lar association has been extremely rare. In fact, seldom have data been compiled (but see Aleev, 1969) so that even a statistical correlation, let alone a causal relation- 0-- Nins Mery Hhyp Msal Eame Ineb Eglo Mpap Mani Pnig ■• ""Laur Lgib Lcya PfIa Lgul ■ ■ Asay Efia 2-- - 3-- Eolm Enig -Ipun ,Nchl Nana Ncer Ncry Nard -Cfun Npro Nhud Mrob Nlep -Nalb Satr Lmac Icat ^Cmac Ccom Pcra Nalt o z V) < UJ cc o z •■ Efus UJ z < < 1- oc < UJ -I LL o •• EobI •• Upyg • Pore V) ^ w FIGURE 6. Ordination of species by mean factor scores on Factor 6 of the factor analysis of mor- phology. See text for interpretation. Abbrevia- tions as in legend for Fig. 1. FIGURE 7. Ordination of species by mean factor scores on Factor 7 of the factor analysis of mor- phology. See text for interpretation. Abbrevia- tions as in legend for Fig. 1. 108 Tulane Studies in Zoology and Botany Vol. 21 silvery or reflective sides - solitary pel- agic; (2) countershading with a dark later- al band - schooling pelagic; (3) mottled pattern or vertical bars - vegetal or ben- thic rocks; and (4) countershading without either silveriness or a lateral band - ben- thic over a sand bottom. 3. Completeness of the lateral line canal was assumed to be an indicator of habitat and behavior also. This sensory structure which detects water movements tends to be best developed in active fishes and reduced or absent in bottom dwellers and sluggish forms (Branson and Moore, 1962). 4. The position of the lateral line canal was assumed to relate to vertical habitat position and to the predation relations of the fish. The character states used and the particular associations were: (0) lacking - benthic or sluggish forms; (1) curving dor- sally - fishes which have predators and/or prey dorsal to them in the water column; (2) horizontal - predators in the upper and midwater region; and (3) curving ven- trally - fishes which feed high in the water column and are subject to predation from below (Marshall, 1971; Disler, 1971). 5. Relative head length was assumed to be related to prey size, a fish with a relatively larger head being able to handle relatively larger prey. To the best of my knowledge this relationship has not previ- ously been studied in fishes, aUhough Schoener (1968) has demonstrated it to hold for certain lizards. My results show that both absolute (r = 0.697, P < 0.001) and relative (r = 0.701, P < 0.001) prey size are positively correlated with relative head length. Thus, large headed little fish like Aphredoderus sayanus took large prey. Both relative head length and stan- dard length are shown in this study to be equally good predictors of absolute prey size, but only the former seems strongly related to relative prey size. 6. Flatness index was assumed to in- dicate habitat water velocity with a lower index being associated with more rapidly flowing water (H.E. Evans, 1950; Hora, 1922; Nikolskii, 1933). As noted above. this character and the next showed a sig- nificant positive correlation with each other as well as being associated in the factor analysis (Factor 2, Fig. 2). Both of these results should be expected given the predicted common dependence of these characters on habitat water velocity. 7. Relative body depth was assumed to be inversely related to habitat water velocity and directly related to capacity for making vertical turns, i.e., about the axis of pitch (Nikolskii, 1933; Aleev, 1969). Significant differences in relative body depth were found between samples of each of three species of centrarchids collected in two different streams (Table 2). For two of the species, an increased sample size was employed to guard against the possibility of a non-random error in the initial measurement or sampling which involved only ten individuals. The intraspecific differences would support the proposed interpretation if (1) habitat water velocity were higher in East Prong Little Yadkin (Pee Dee) than in Maho Creek (Roanoke) and higher in the latter than in Mud Creek (Cape Fear) ; and (2) the morphological variation were due to adaptation to these local physical condi- tions. The required differences in habitat water velocities seem likely. East Prong Little Yadkin has a gradient three to four times greater than the other two streams and hence would be expected to have a mean velocity one and a half to two times greater than either of the others (Hynes, 1970). The few pools which are present are small and isolated backwaters are vir- tually nonexistent. Thus likely both Le- pomis aiiritus and Lepomis cyanellus are subjected to a stronger selective pressure for increased tolerance to flowing water in East Prong Little Yadkin than in Maho Creek. Alternative possibilities, e.g., character displacement, or in the case of the introduced L. cyanellus, founder ef- fect, do exist, but seem less likely. With regard to the comparison of Pomoxis ni- gromaculatus in the two streams, a higher mean velocity is expected in Maho Creek than in Mud Creek because although the No. Ecological Morphology of Fishes 109 TABLE 1 . Results of the gut content analyses for the 33 common species of fishes. The numerical entries In the table for each food category are char- acter codes for, respectively, frequency and Importance of that particular prey type In the diet of the species. For a complete explanation of the cod- ing , see the methods section of the text. t o o o (0 o » to c < o c (0 o (0 -a u o 10 O (0 c ■p. 5 5 I 10 o (0 3 o z: eg m u o > (0 1) a -p c (d U nS rH 3 o (0 cd > O m s s p< p< 3 K Esox amerlcanus Esox niger Cllnostomus funduloldes Hybopsls hypslnotus Npcomls leutocephalus Notemlgonus crysoleucas Notropls alborus Notropls altlplnnls Notropls analostanus Notropls ardens Notropls ceraslnus Notropls chllltlcus Notropls procne Phoxlnus oreas Semotilus atromaculatus Gatostomus commersonl Erlmyzon oblongus Ictalurus nebulosus Noturus Inslgnls Aphredoderus sayanus Gambusla afflnls Centrarchus macropterus Lepomis aurltus Lepomis cyanellus Lepomis gjbbosus Lepomis gulosus Lepomis macrochlirus Mlcropterus salmoides Pomoxis nlgromaculatus Etheostoma flabellare E the o stoma fusiforme 'Etheostoma olmstedl Perca flavescens 23 11 11 23 22 55 1 11 11 1 1 1 22 1 k k 55 22 11 33 n 33 1 11 11 1 1 1 22 1 6 5 11 11 11 11 66 11 1 11 11 1 1 1 11 1 2 2 11 23 11 55 23 22 1 22 11 1 1 5 11 1 1 2 11 11 11 45 23 23 1 23 11 1 I* 1 11 1 2 2 11 22 11 33 3^ 11 1 JVi 23 1 2 4 11 1 1 1 11 11 11 33 23 11 1 34 11 1 1 3 11 1 1 1 11 11 11 11 55 11 1 11 33 1 1 1 11 1 1 2 11 11 11 33 55 11 1 22 23 1 1 2 11 1 1 2 11 11 11 33 55 11 1 11 11 1 1 1 11 1 2 2 11 11 11 65 ^3 11 1 11 22 1 1 3 11 1 3 3 11 11 11 33 ^+5 11 1 11 33 1 2 1 11 1 2 2 11 11 11 ^44 22 11 1 /+4 22 1 1 4 11 1 1 1 11 11 11 23 11 11 1 65 11 1 1 6 11 1 1 1 11 11 11 11 66 11 1 11 11 1 1 1 11 1 3 k 22 11 32 65 23 32 1 11 33 1 6 3 23 1 3 2 11 44 33 43 11 11 1 23 32 1 2 6 22 1 1 1 11 11 ^44 33 11 22 1 11 33 2 3 5 11 2 2 1 11 11 11 65 22 11 1 11 11 1 6 1 11 1 2 1 11 11 11 55 11 11 2 11 11 1 2 2 23 1 3 6 11 11 11 33 55 11 1 11 11 1 1 1 11 1 1 1 23 23 11 43 44 11 1 11 11 1 1 1 11 1 3 3 11 11 11 5^ 33 33 1 11 11 1 1 1 23 2 4 4 11 11 11 43 33 44 2 11 11 1 1 2 11 1 4 5 11 11 33 56 43 11 1 11 22 1 1 4 11 1 2 2 11 11 11 44 43 33 1 11 11 1 1 1 11 1 4 5 11 22 11 55 ^+3 11 1 11 11 1 1 2 11 1 2 2 33 11 11 23 ^ 23 1 11 11 1 1 1 11 1 3 4 33 22 11 11 65 33 1 11 11 1 1 1 22 1 3 2 11 11 11 65 11 11 1 11 11 1 1 1 22 1 2 3 11 43 11 65 11 11 1 11 11 1 1 1 11 1 1 2 11 11 11 65 11 11 1 11 11 1 3 1 11 1 1 2 11 11 11 65 11 33 1 11 11 1 1 1 11 1 2 2 no Tulane Studies in Zoology and Botany Vol. 21 TABLE 2. Intraspeciflc variation in relative bcxiy depth (see text for definition) in three species of centrarchids „ N^t= not measiared. Species n East Prong Mud Cr. Maho Cr. F value Signif. Lepomis aiiritus 15 0.397 NM 0.444 9.806 <0.01 Lepomis cyanellus 10 0.379 NM 0.395 5.804 <0.05 Pomoxis nigromaculatus 28 NM 0.431 0.403 8.874 <0.01 gradients are similar in both, the former is larger (Hynes, 1970). Again, the observed differences correspond to the hypothe- sized interpretation of the character. No other species was found to show intra- specific variation in this character. Such variation has been reported in Gromov (1973) who compared lake and river pop- ulations of carp. 8. Index of trunk shape was assumed to be directly related to hydrodynamic ability in that a high value is associated with a late separation of the boundary lay- er (Aleev, 1969; Ovchinnikov, 1971). Hence higher values are assumed to indi- cate fishes which spend more time cruis- ing. However, Aleev (1969, Table 21) did point out that body height affects the in- dex so that deep bodied fishes are apt to have higher values than one might expect from their swimming habits. My results tend to substantiate Aleev's findings. In the first place, I did not find this character to show a significant corre- lation with any other. Inasmuch as other characters were also assumed to relate to various aspects of hydrodynamics and cruising, this suggests either that this character measures some unique aspect of swimming ability, or that there are other irregularities in the values. The ordination of species means for this character sug- gests the latter to be the case as high values are recorded both for cruising minnows and for several species of laterally flat- tened, deep bodied centrarchids. Thus my results indicate the same ambiguity in the significance of this index as those of Aleev (1969). 9. Relative peduncle length was as- sumed to be directly related to swimming ability (Hora, 1922; Kanep, 1971). 10. Caudal peduncle flatness index was assumed to relate inversely to amplitude of swimming movements (Nursall, 1958) and, like flatness index of the body, to be higher in less active swimmers. In general, a listing of species according to increasing value for this character produced an or- dination of species which, based on litera- ture description of their habits, would be concordant with the assumed significance. Thus, for example, Pomoxis nigromacu- latus showed a higher index than Lepomis auritus and L. cyanellus which in turn showed higher values than Micropterus salmoides. Caudal fin characters 11. Aspect ratio of the caudal fin was assumed to be directly proportional to the amount of swimming the fish does (Nich- ols, 1915; Harris, 1953; Nursall, 1958; Aleev, 1969; Kramer, 1960). The positive correlation of this character with the per- centage of red muscle in the peduncle is the most tangible evidence for the as- sumed function (see discussion of red muscle, below). 12. Caudal span/body depth ratio was assumed to be directly proportional to speed of swimming in that ratios of in- creasing magnitude greater than one in- dicate better hydrodynamics for the caud- al fin as the lobes of the fin are out of the zone of vortices shed from the body (Aleev, 1969). I found low values for sun- fishes and high values for minnows which, according to the hydrodynamic argument No. 2 Ecological Morphology of Fishes 111 presented above, would imply that min- nows are faster swimmers than sunfishes. Experimental substantiation of this is lacking. 13. The number of caudal fin rays was assumed to be directly proportional to the significance of the caudal fin as the princi- pal means of locomotion (Marshall, 1971). My results suggest the separations of species given by this character may well be more phylogenetic than functional or ecological for my species. For instance, all sunfishes had 17 caudal fin rays, suckers had 18, and both minnows and pickerels had 19. Paired fin characters Although the importance of the paired fins in low speed maneuvering has been investigated by a number of persons (e.g. , Aleev, 1969;Breder, 1926; Gosline, 1971; Gray, 1968; Harris, 1953), precise associ- ations between functional specializations and particular morphological features have not always been defined. Thus for some of the following characters, no def- inite functional correlate is given but rath- er an assumed function based on theoret- ical considerations. 14. Pectoral fin length was assumed to increase as a function of amount of low speed maneuvering in the behavior of the fish (Gray, 1968; Starck and Schroeder, 1970; Kanep, 1971). Factor 2 of the factor analysis indicates an association between high values in this character and body shape characters which indicate low habi- tat water velocity (see Fig. 2). One might expect a fish in quiet water to do more low speed maneuvering than one living in fast water. 15. Aspect ratio of the pectoral fin was assumed likely to relate in some manner to the ways the fin might be able to function. The only correlation found for this charac- ter was with the next. 16. Relative pectoral fin area was as- sumed to be directly proportional to the capacity of the fin to function in braking, fanning to maintain position, and acceler- ation from the stop (Gosline, 1971). My results, particularly Factor 3 of the factor analysis (Fig. 3), indicate large relative pectoral fin area is also associated with benthic habits. Recently, A.N. Jones (1975) also reported pectoral fin area to relate to benthic living, particularly in areas of current. Jones found that salmon had larger pectoral fins than trout and were more frequently found in riffles. He suggested that the larger pectoral fin of the salmon enables it to "hold station" in riffles by deflecting water over the fin and thereby enable salmon to occupy terri- tories unavailable to trout. Keenleyside (1962) reported observations made while skin diving which confirm this interpreta- tion. He observed that salmon rest on the bottom with their pectoral fins spread, whereas trout do not maintain contact with the bottom, but rather usually swim actively against the current to maintain position. For additional information, see also Lundberg and Marsh (1976) who dis- cuss how suckers use their pectoral fins on the substrate. 17. Relative distance of the pectoral fin from the center of gravity of the fish was assumed to be directly proportional to the importance of the pectoral fin in the turn- ing and maneuverability of the fish (Aleev, 1969; Breder, 1926) in that all else being equal, a larger distance from the center of gravity provides greater torque. The various correlations reported above for this character do not provide any real support for this argument. 18. Pectoral fin shape, like its aspect ratio, was assumed to relate to its func- tion. One correlate in the literature is that rounded fins are characteristic of fishes which remain motionless in midwater (Aleev, 1969). 19. Position of the pectoral fin relative to the center of gravity was assumed to have something to do with the turning capacity of the fish. 20. Number of pectoral fin rays has been suggested by Hubbs (1941) to be directly proportional to the amount of time the fish spends in the current. 21. Pelvic fin length was assumed to be related to habitat preference, being longer 112 Tulane Studies in Zoology and Botany Vol. 21 in rocky habitat species and shorter in free swimming species (Starck and Schroeder, 1970). 22. Aspect ratio of the pelvic fin was assumed to be small in fishes in which the pelvic fin is used only for a brake or for- ward swimming aid and large if the fins are important for backing and hovering (Har- ris, 1937). 23. Relative pelvic fin area was as- sumed to be largest in fishes with demersal habitat preference (Aleev, 1969). 24. Relative distance of the pelvic fin from the center of gravity of the fish was assumed to be an indication of torque and hence of the importance of the pelvic fin in the turning and maneuverability of the fish (Aleev, 1969; Breder, 1926). 25. Pelvic fin shape was assumed to re- late to its function. One correlate in the literature is that more falcate pelvic fins are found in fishes which are constantly in the current (Hubbs, 1941). 26. Position of the pelvic fin relative to the center of gravity was assumed to lower with increased proportional use of the pectoral fin as brakes because the pelvics must be close to the center of gravity to counteract the pitch induced by such braking (Harris, 1938). Values were as- sumed to be higher with increased impor- tance of the pelvic fins in turning move- ments (Breder, 1926). 27. Number of pelvic fin rays was as- sumed to be directly proportional to the amount of time the fish spends in the cur- rent (Hubbs, 1941). 28. Position of the dorsal fin relative to the center of gravity was assumed to re- flect what functions the abrsal fin was cap- able of performing: (a) anterior rudder function is better the further anterior to the center of gravity the fin is, (b) keel function is best served at the level of the center of gravity, (c) stabilization of for- ward movement is associated with a pos- terior position, and (d) posterior rudder and/or locomotory function are best per- formed the further posterior the fin is po- sitioned (Gosline, 1971). Head characters 29. Relative eye size was assumed to be directly proportional to the development of visual capabilities in the fish (Protasov, 1970) and hence this character was as- sumed directly proportional to the impor- tance of sight in the feeding of the fish (H.E. Evans, 1950). Not all species reputed to be visual predators, e.g., Esox niger, had large rel- ative eye sizes. Part of the explanation for this might be the known negative allo- metry of eye size in fishes (Martin, 1949). A transformation based on regressions of absolute eye size on standard length and using the intercepts as a measure of rela- tive eye size merely gave a second value nearly perfectly correlated (r = 0.9985) with the original character. Somewhat more disconcerting than the partially counterintuitive sequencing of some species in values for relative eye size was the low correlation of this character with relative size of the optic lobes, which was also assumed to be directly propor- tional to the development of visual capa- bilities in the fish. The Pearson coefficient was 0.455 (0.01 > P > 0.005) and the Spearman, 0.476 (P = 0.005). Still, enough of a trend appeared to be evident between visual habits of well known spe- cies and the relative eye size values for me to consider the use of this character to be valid. 30. Position of the eyes was assumed to relate to vertical habitat preference. Lat- eral placement was assumed tc indicate pelagic habit and increased displacement dorsally was associated with the assump- tion of a more sedentary mode of life (Aleev, 1969). My results provide support for this interpretation. Although no signif- icant simple correlations with this charac- ter were identified, the factor analysis (Factor 3, Fig. 3) did indicate it to be associated with several other characters which also were assumed to reflect benthic habitat preference. 31. Eye pigmentation was assumed to reflect habitat and behavior. The assumed correlates were: no pigmentation - slow No. 2 Ecological Morphology of Fishes 113 moving and/or benthic fishes; horizontal habitat a fish obtained its foods. Given Hne of pigmentation - cruising habit; ver- that certain types of prey are apt to be tical line through the eye - fast turning found on or near the bottom (e.g., ostra- habit; and presence of both a horizontal cods, diatoms, aquatic insect larvae) and and a vertical line through the eye - both others only at the surface or in mid-water specializations (Barlow, 1972). (e.g., water striderst, terrestrial insects, 32. Position of the mouth was assumed fishes), the existence of correlations be- to indicate the location of the food eaten relative to the fish (Aleev, 1969; Al- Hussaini, 1949; Schmitz and Baker, tween those types in the gut and mouth position and/or orientation can be deter- mined. Due to the manner in which I cod- 1969). Schutz and Northcote (1972) pro- ed position of the mouth and orientation vided experimental support for this inter- of the mouth, one should expect positive pretation. They found that Dolly Varden correlations between these characters and {Salvelinus malma) which have subter- items of benthic prey and negative corre- minal mouths were more effective bottom feeders than cutthroat trout (Salmo clarki clarki) which have terminal mouths and vice versa when both species were tested in aquaria. This evidence, notably, is only for two closely related species. My results which support the assumed interpretation are discussed below in conjunction with orientation of the mouth. 33. Orientation of the mouth was also assumed to indicate from where in the lations with surface and mid-water prey. Table 3 shows the correlations for all prey categories with these two morphol- ogical features. The expected relations, although weak, do hold for ostracods, fil- amentous algae, aquatic insects and fre- quency of terrestrial insects. They were not demonstrated for diatoms or for fish- es. The strongest positive correlations were with sand and silt, and the strongest negative, with prey size. The former cor- TABLE 3. Correlations of gut contents with position and orientation of the mouth. The first figures for each category present coefficient and significance for frequency. The second fi- gures give coefficient and significance for importance, where applicable (see text/. Gut content category Correlations with position of the mouth Fish Gopepods Ostracods Aquatic insects Terrestrial insects Crayfish Amphipods and isopods Diatoms Filamentous algae Molluscs Sand Silt Terrestrial invertebrates Vascular plants Absolute prey size Relative prey size -.31. NS +.18, NS +.39. P <.05 +.39. P<.05 -,h2,, P <.02 -.29. NS -.28, NS +.28, NS +.45, P <.0l +.12, NS +.63, P <.001 +.47, P <.01 -.12, NS -.05. NS -.55, P <.001 -.60, P <.ooi -.35, P <.05 +.17. NS +.31. NS +.40, P<.02 -.29, NS -.37. P<.05 +.34, NS +.38, P<.05 -.07. NS Correlations with orientation of the mouth -.24, NS +.06, NS +.46, P <.01 +.45, P <.0l -.44, P <.0l -.15, NS -.23. NS +.25. NS + .46, P-C.Ol +.15. NS +.54, P<.002 +.48, P<.005 -.14, NS -.01, NS -.41, P<.02 -.52, P<.002 -.28, NS +.02, NS +.38, P <.05 + .47. P<.01 -.34. NS -.22. NS +.30. NS +.40, P <.05 -.06, NS 114 Tulane Studies in Zoology and Botany Vol. 21 relations are considered very strong evi- dence that fishes with more ventral mouths obtain their prey from the bottom and those with more terminal or anterior mouths do not. The negative prey size correlation also supports the assumed sig- nificance of these characters in that the smallest prey items in this study, diatoms, tend to be benthic organisms in lotic hab- itats. Thus, my results indicate support for the hypothesized significances for both of these characters for a wide range of taxa. 34. Relative width of the mouth was assumed to be directly proportional to the size of food eaten (Aleev, 1969; Forbes and Richardson, 1920; Starck and Schroeder, 1970). My results pertaining to this character are discussed below with those for the next character. 35. Relative height of the mouth was also assumed to indicate size of prey. This is another of the few characters I used for which direct experimental evidence exists (Werner, 1974). Werner showed that op- timal prey size for Lepomis cyanellus and L. macrochirus was a function of mouth size. My results show that both width and height of the mouth are highly significant- ly correlated with both relative and abso- lute prey size (for both Spearman and Pearson coefficients, all r's are between 0.594 and 0.681; all P's are less than 0.001). Thus my correlative evidence sup- ports my having assumed generality of the type results obtained by Werner (op. cit. ). Additionally, my extensive multifamily correlations also indicate that the correla- tions of prey size with gape (of the mouth) found by Thomas (1962) for two salmon- ids and Northcote (1954) for two cottids were not unique to these two families. 36. The index of protrusion was as- sumed to be greatest in fishes with the smallest prey. This is based on the as- sumption that strength of bite is inversely related to the degree to which the pre- maxillary is protruded and that a stronger bite is required to capture a large item of prey than a small one (Al-Hussaini, 1949; Aleev, 1969;Gosline, 1973). My results do not support these assump- tions. This index did not correlate nega- tively with either relative or absolute prey size (r = -0. 14) among all species of fishes as should have been predicted. Five of the eight species taking prey of the largest relative prey sizes {Esox niger, E. ameri- canus, Aphredoderus sayanus, Semotilus atromacidatus, and Micropterus sal- moides) did have mouths with an index of protrusion of one or nearly one (i.e., no or very little protrusibility); however, so did such fishes as all members of the genera Etheostoma (darters) and Ictalurus (cat- fishes) which take prey of very small rela- tive size. The other three predators on large relative size prey {Lepomis auritus, L. cyanellus, and L. gulosus) all had mouths which showed high values for this index. This character therefore seems to be of limited usefulness in comparing fish- es between different families. 37. Number of barbels was assumed to be directly proportional to the importance of non-optic senses in feeding (Gilbert and Bailey, 1972) and the benthic habit of the fish (Gosline, 1973; Aleev, 1969). The barbels are innervated by the facial nerve so that one should expect this character to be strongly correlated with the relative size of the facial lobes. In fact, only the Pearson correlation of these two variables was significant (r = 0.790, P < 0.001) and not the Spearman. 38-40. Number of branchiostegal rays, presence of jaw teeth, and shape of jaw teeth were all assumed to be inversely re- lated to the importance of suction in the capture of prey (Gosline, 1973). In addi- tion, shape of jaw teeth was assumed to indicate size of prey. One would not expect suction to be im- portant in the capture of large prey, i.e., such items as fish or crayfish. In view of this expectation, the positive correlations of absolute prey size with number of branchiostegal rays (P < 0.001), with presence of jaw teeth (P < 0.002) and with the shape of jaw teeth (P < 0.002) provide evidence in support of the func- No. 2 Ecological Morphology of Fishes 115 tion I assumed for these characters. Addi- tionally, my results provide further sup- port for this proposed fimctional signifi- cance in that all three characters showed positive correlations with both frequency and importance offish and crayfish (P's < 0.05 to < 0.001). 41. Hypertrophy of teeth on the phar- yngeal arches was assumed to relate to manner of feeding, especially to be direct- ly related to the importance of suction in capture of prey (Gosline, 1973). My re- sults support this hypothesis in that a neg- ative correlation was found between abso- lute prey size and this character (P < 0.02). 42. Shape of pharyngeal teeth was as- sumed to relate to dominant food type in the diet (H.E. Evans, 1950; Al-Hussaini, 1949). I found a weak positive correlation (P < 0.05) of the character with the fre- quency of occurrence of both filamentous algae and ostracods in the diet (recall the coding for this character provided a high value for teeth with grinding surfaces). No further significant correlations were de- termined. Apparently many different fish species, each with its own type of denti- tion, are able to eat from the same broad prey categories used in this study. 43. Number of gill rakers was assumed to be inversely correlated with the pres- ence of larger and more benthic prey in the diet (Kliewer, 1970; Himberg, 1970; Nilsson, 1958). My results did not support a generalization of this observation on salmonids; in fact, they show the reverse relationship. Absolute prey size and number of gill rakers gave a positive corre- lation (r = 0.519; P < 0.002). Several factors enter into this: (1) considering the many reduced plates in Esox each to rep- resent a gill raker so that those predators on large prey have among the highest gill raker counts; (2) the generally high gill raker counts among sunfishes; and (3) the generally low gill raker counts among minnows. Thus a strong taxonomic com- ponent negates the general utility of this character along lines such as those demon- strated by Kliewer (1970) and Himberg (1970). In this study I varied my interpretation of the number of gill rakers depending on the degree of taxonomic relatednessof the species being compared. For intrafamilial comparisons, I assumed that the usual functional interpretation is correct and that a low number of gill rakers was indic- ative of large benthic prey. For general interfamihal comparisons, I interpreted differences in number of gill rakers by bearing in mind that the two strongest pos- itive correlations with this character for all species are with items of the two largest prey size categories in the diet: (1) fish (P < 0.001) and (2) crayfish (P < 0.01). < 44. Shape of the middle gill raker, as measured by the ratio of length to width, was assumed to be inversely related to food size (Kliewer, 1970; Starck and Schroeder, 1970; Forbes and Richardson, 1920). My results lend little support to this assumption. No correlation between this variable and food size was found. Nor was this character correlated with any of the other morphological characters. The only correlation found was a weak positive as- sociation between shape of gill rakers (high values indicate long, thin gill rakers) and importance of copepods in the diet (r = 0.373, P < 0.05). This gives minor sup- port to the functional interpretation as- sumed in this study, but obviously more parameters are operational in the selec- tive regime for this feature than just the ability to obtain small mobile prey like copepods. 45 . Finer structure of the gill rakers was assumed to relate to food type and food handling. My results supported this as- sumption as a number of significant rela- tionships between prey and gill raker ar- mament were observed. The strongest of these correlations are with prey size. High numbers of fine teeth on the gill rakers are correlated both with absolute size of prey (r = 0.577; P < 0.001) and relative size of prey (r = 0.613; P < 0.001). Weaker cor- relations were observed with individual categories of large prey types: crayfish (r = 0.51; P < 0.002); fish (r = 0.42; P < 116 Tulane Studies in Zoology and Botany Vol. 21 0.02); and non-insect terrestrial inverte- brates (r = 0.39; P < 0.05). Weak nega- tive correlations were found with both dia- toms (r = -0.42; P < 0.02) and filamen- tous algae (r = -0.41; P < 0.05), which indicated a tendency for fishes eating these latter two types of prey to have smooth or ridged gill rakers. Internal body characters 46. Relative volume of the swim blad- der was assumed to be inversely propor- tional to the bottom dwelling preference of the fish (Forbes, 1880; Bridge and Haddon, 1S89) and also negatively corre- lated with water speed in the habitat of the fish (Hora, 1922; Gee, 1968, 1974). This latter relationship is one of the very few in which experimental evidence exists that demonstrates a causal relationship be- tween ecological conditions and morphol- ogy. Gee (1970, 1972) and Gee et al. (1974) have presented data that indicate a number of species of fishes respond to an increase in water velocity in the laboratory by decreasing the volume of their swim bladders. A complete listing of relative volume of the swim bladder (first value) and relative length of the swim bladder (second value) follows. Species are listed in order of in- creasing swim bladder volume and paren- theses around a species name and values indicates that the sample size was 1 or 2 rather than the usual 10. The species mean values are: Etheostoma flabellare 0,0; E. fusiforme 0,0; E. olmstedi 0,0; {Percina crassa 0,0); Aphredoderus say anus 0.036,0.282; (Anguilla rostrata 0.037, 0.200); (Umbra pygmaea 0.38,0.359); No- turus insignis 0.046,0.098; Notropis chiliti- cus 0.054,0.318; (Notropis hudsonius 0.054,0.341); (Moxostoma robustum 0.054,0.462); (Ictalurus punctatus 0.054, 0.182); Nocomis leptocephalus 0.056, 0.304; Notropis ardens 0.056,0.311; Cato- stomus commersoni 0.057,0.360; Clino- stomus funduloides 0.061,0.340; Perca flavescens 0.061,0.330; Erimyzon oblong- us 0.061,0.358; Lepomis cyanellus 0.061, 0.297; Notropis procne 0.062,0.321; Se- motilus atromaculatus 0.063,0.344; No- tropis cerasinus 0.064,0.363; (Ictalurus catus 0.065,0.186); Ictalurus nebulosus 0.066,0.197; Esox niger 0.067,0.415; Mic- ropterus salmoides 0.068,0.314; Phoxinus oreas 0.068,0.336; Hybopsis hypsinotus 0.068,0.343; Notropis analostanus 0.069, 0.356; (Enneacanthus gloriosus 0.071, 0.343); Lepomis gibbosus 0.071,0.336; Lepomis auritus 0.072,0.315; Gambusia affinis 0.074,0.264; Notropis altipinnis 0.074,0.341; (Moxostoma erythrurum 0.075,0.473); Moxostoma anisurum 0.076,0.446; Lepomis gulosus 0.076, 0.341 ; Notropis alborus 0.078, 0.337; Cen- trarchus macropterus 0.079,0.426; Le- pomis macrochirus 0.080,0.346; (Moxo- stoma pappillosum 0.083,0.465); Esox americanus 0.083,0.435; Notemigonus crysoleucas 0.083,0.378; Pomoxis nigro- maculatus 0.085,0.364. A comparison of habitat observations of the fishes with the values obtained in this study confirmed both of the points in the assumed significance of this variable. Known bottom dwellers in fast current re- gions had small swim bladder volumes. For example, all Etheostoma studied lack- ed gas-filled swim bladders and the blad- der of Noturus insignis had a relative vol- ume of only 4.6%. Aphredoderus saya- nus, a bottom dwelling inhabitant of quiet water regions, also had a small swim bladder (3.6%). Pelagic, quiet water in- habitants, e.g., Pomoxis nigromaculatus, Lepomis macrochirus, and Notemigonus crysoleucas had large swim bladders (8.0- 8.5%). Additionally, the factor analysis indicated relative volume of the swim blad- der to covary with several other characters related with benthic habits (Factor 3, Fig. 3). Comparisons of the actual magnitudes of the results listed above with a theoretical value which is assumed to give a fish neutral buoyancy are interesting. F.R.H. Jones (1951) calculated that if the average density of a fish were 1.076 g/cc, then a swim bladder volume equal to 7.06% of the total volume of the fish would be nec- essary to achieve hydrostatic equilibrium. Only a few species of fishes studied here No. 2 Ecological Morphology of Fishes 117 showed mean values significantly differ- ent from this figure. Notropis chiliticus, Noturus insignis, Umbra pygaea, Anguilla rostrata, Aphredoderus sayanus, and all the darters had smaller percentage vol- umes and only Pomoxis nigromaculatus had a larger percentage volume than this theoretical figure. However, among the remaining species many significant differ- ences did exist. These differences seem likely to reflect biological differences as stated above. Previously reported values of the percentage volume of the swim bladder in Cypriniformes have ranged from 5% to 10% (Alexander, 1959). 47. Relative length of the swim bladder was assumed to relate in the same way to the same qualities as relative volume of the same structure (Nelson, 1961; H.E. Evans, 1950). My results show that the ordination of species on this character would be similar to that obtained with the preceding (see listing above) and that the two characters are highly correlated (Pearson r = 0.88; P < 0.001). However, this character tends to have a stronger taxonomic component than the preced- ing. Thus even catfishes with fairly large relative swim bladder volumes (e.g., Ic- talurus punctatus) have small relative swim bladder length. Suckers, especially those of the genus Moxostoma which have a three-part swim bladder, have large rela- tive swim bladder lengths. Studies which encompass several families of teleosts and employ length of the swim bladder (or even length and depth but not width as Dobbin, 1941) as the sole estimator of buoyancy rest on shaky ground. 48. Relative gut length was assumed to be directly related to mud feeding, herbi- vory, and omnivory, and inversely related to camivory and insectivory (H.E. Evans, 1950; Lagier et al., 1962; Forbes, 1888; Schmitz and Baker, 1969). My results support the "mud feeding" aspect espec- ially, as positive correlations exist be- tween this character and the presence of ostracods (P's < 0.02-0.002), diatoms (P's < 0.05-0.01), and sih (P < 0.001) in the gut. 49. Number of pyloric caeca was as- sumed to be correlated with the protein richness of the diet, as these structures function as an enzyme source and an area of absorption for protein nitrogen (Beam- ish, 1972; Phillips, 1969). My results show that the taxonomic component to this character was great. Thus, although some of the species for which I found piscivor- ous food habits (e.g., Micropterus sal- moides, Pomoxis nigromaculatus, and Centrarchus macropterus - see Table 1) do have the most pyloric caeca, the most piscivorous of the fishes studied, Esox niger, lacks these structures. 50. Percentage of red muscle in the caudal peduncle was assumed to be direct- ly proportional to the amount of sustained swimming or cruising that the fish does (Boddoke et al., 1959; Love, 1970; Gatz, 1973). Recently Roberts and Graham (1974) reported on the basis of electro- myographic recordings that red muscle primarily is operating at slow swimming speeds in mackerel and that both red and white muscles increase in activity during acceleration. The following listing of species by in- creasing percentage of red muscle corres- ponds generally to one's intuitive expecta- tions given this function. Note sample size is 10 except for species in parentheses for which n = 1 or 2. Species mean percen- tages are: (Umbra pygmaea — 0%); Ethe- ostoma flabellare — 09^ ; E. fusiforme - 0% ; E. olmstedi - 0% ; (Percina crassa - 0%); Esox niger - 1.8%; (Enneacanthus gloriosus — 2.2%); Esox americanus — 2.2%; Aphredoderus sayanus — 3.2%; Pomoxis nigromaculatus - 3.4% ; Lepom- is cyanellus — 3.9%; Lepomis macrochir- us - 4.2%; Lepomis gibbosus - 4.2%; Lepomis auntus — 4.4%; Micropterus salmoides - 4.4%; Gambusia affinis - 4.5%; {Anguilla rostrata - 4.7%); Eri- myzon oblongus - 4.7%; Lepomis gulo- sus — 4.7%; Centrarchus macropterus — 5.0%; (Moxostoma anisurum - 5.4%); Perca flavescens - 5.5%; Notropis albor- us — 5.8%; Phoxinus oreas - 6.1%; Icta- lurus nebulosus — 6.3%; (Notropis hud- 118 Tulane Studies in Zoology and Botany Vol. 21- sonius — 6.6%); {Moxostoma erythrurum — b.7%); Hybopsis hypsinotus - 6.7%; Notemigonus crysoleucas — 6.7% ; (Ictalu- rus catus — 7.5%); Notropis procne — 7.5%; Catostomus commersoni - 1.1%; {Moxostoma pappillosum - 8.4%); No- turus insignis — 8.5%; Notropis altipinnis — 8.8%; Semotilus atromaculatus - 9.2%; Notropis cerasinus - 9.2%; No- comis leptocephalus — 9.7%; Clinostomus funduloides — 9.8%; Notropis analosta- nus - 9.9%; Notropis ardens - 11.1%; (Moxostoma robustum - 11.3%); No- tropis chiliticus - 12.0%; and (Ictalurus punctatus — 14.2%). Most of the work in the hterature in which actual measurements of stamina for swimming in fishes were made has been for salmonids (e.g., Brett, 1965, 1967, 1973; Hammond and Hickman, 1966; Hochachka, 1961; Horak, 1969, 1972; D.R. Jones, 1971; Stevens, 1968; and Vincent, 1960); none of these fishes occur in my study streams. A few papers do report on such non-salmonids as goldfish (Fry, 1958), smallmouth bass (Larimore and Duever, 1968), largemouth bass (Laurence, 1972) and bluegills (Oseid and Smith, 1972). However, only the work by Houde (1969) and Bainbridge (1960) was of an interspecific comparative nature such that one could compare the swim- ming abilities for two or more species with red muscle content. Houde (op. cit.) found little difference between two spe- cies of percids with the 9 to 15 mm total length larvae that he used. Gatz (1973) speculated on the possible relationship be- tween differences in swimming perfor- mance among the three species tested by Bainbridge (1960) and differences in red muscle content and concluded the rela- tionship assumed in this paper seemed op- erative. Supportive evidence of a somewhat dif- ferent sort comes from the work of Reddy and Pandian (1974) on the predation ef- fectiveness of Gambusia affinis in flowing water. In addition to giving data which indicate a halving of predatory efficiency, they reported that the Gambusia appear- ed to have lost much energy in the flowing water and appeared to have very little available to spare for predation. Such re- sults correspond nicely with the low amount (4.5%) of red muscle I found in this species and the proposed interpreta- tion of this character. A similar experi- ment would be interesting with a species such as Notropis chiliticus or N. ardens, both of which have more than twice the red muscle Gambusia affinis has. A further point of interest about the percentage of red muscle in the peduncle concerns the probable selective forces in- volved in achieving the observed values for this character. An ability to cruise slowly for prolonged periods with low metabolic expenditure readily enough would "explain" selection for red muscle. But why do some species lack it entirely (e.g., Etheostoma) or have very little (e.g., Esox and Umbra)! Two points come to mind. First, some fishes move about slowly, seemingly entirely by the movement of the paired fins. Esox and Umbra are examples, and the fin muscu- lature of these species is red muscle. Sec- ond, if red muscle were metabolically more costly to maintain than white in a resting state, then one would expect its presence to be selected against in those species which do not demonstrate slow cruising behavior. Some evidence of just such a higher metabolic rate for red mus- cle for white has been given by Gordon (1968) and Lin etal. (1974). Brain characters 51. Relative size of the forebrain was assumed to be directly proportional to the importance of olfaction in the life of the fish (Tuge et al., 1968) and also with noc- turnal habits (H.M. Evans, 1940). 52. Relative size of the optic lobes was assumed to be proportional to the impor- tance of vision in the species (Schwassman and Kruger, 1968). 53. Relative size of the cerebellum was assumed to be large in active, quick swimming fishes and in forms which live in fast water, but small in sluggish, slow wat- er fishes (Herrick, 1924; Schnitzlein, No. 2 Ecological Morphology of Fishes 119 1964; Miller and Evans, 1965). 54. Relative size of the vagal lobes was assumed to be positively correlated with "mouth tasting," i.e., eating mud which is then sorted to a degree in the mouth using taste buds of the mucous membrane of the posterior oral cavity (Bhimachar, 1935; H.E. Evans, 1952; H.M. Evans, 1940). 55. Relative size of the facial lobes was assumed to be directly proportional to the importance of lip and barbel tasting in the feeding behavior of the fish (H.M. Evans, 1940; H.E. Evans, 1952; Miller and Ev- ans, 1965). 56. Relative size of the acoustic tuber- cles was assumed to be directly propor- tional to the importance of the lateral line system in the behavior of the fish (H.M. Evans, 1940; Tuge et al., 1968). For all six characters relating to the hypertrophy of various lobes of the brain, my results placed at least some species for which behavior was known at seemingly appropriate positions in the ordinations obtained. For example, Anguilla rostrata and members of the genus Ictalurus have large forebrains; and cyprinids have rela- tively larger cerebellums than centrar- chids. The assumption that other less known species were in appropriate rela- tive positions did not seem unwarranted. Few studies have been done comparing the relative importance of different senses in feeding and thus, by assumption in this study, with the relative sizes of different brain lobes. The only such paper of which I am aware which involved a species in this study was that by Roberts and Winn (1962). They found Etheostoma olmstedi (I changed the nomenclature here to cor- respond with Cole, 1967, 1972) to rely much more heavily on sight than olfac- tion. This corresponds well with my re- sults. I found that E. olmstedi had one of the smallest values for relative size of the forebrain and one of the largest for rela- tive size of the optic lobes of all those species measured. Caveats and Summation Bock and von Wahlert (1965) point out that any particular anatomical feature or structure may serve multiple functions and thus be subject to natural selection as a part of any of several form-function complexes. They suggested that the biol- ogical role of any feature cannot be de- duced from the mere study of the form- function complex, but must be deter- mined by direct observation of the organ- ism. Alternatively, it would seem that if natural selection has operated in a repeat- able manner to cause a regular association between a particular morphological fea- ture and one form-function complex, then evidence for this should exist as a constan- cy of utilized faculties for the particular feature. Specifically, there should be strong correlations between such a feature and some manifestation of its biological role and also there should be correlations between this feature and others associated with the same role. For example, if a prin- cipal selective pressure on mouth size is always prey size, then there should be a correlation between prey size and any measure of mouth size among a group of species of fishes and correlations between the various measures of mouth size. The results presented above attest to the validity of both points of view. First, the many significant correlations indicate that some repeatable selective forces do seem to be governing some portion of the variation in the morphological features measured. This should not be surprising in that each character was chosen on the ba- sis that I felt the strongest single selective force operating on it was recognizable and in that these selective forces were at times the same for more than one character. In a number of cases, e.g., relative width and height of the mouth, strong correlative evidence in support of the hypothesized adaptation has been given. On the other hand, only about 2% of the between char- acter correlations have coefficients of de- termination greater than 50%. This dearth of high correlations, especially be- tween characters assumed to be respond- ing to similar selective forces, gives sup- port to the viewpoint of Bock and von Wahlert (1965). That is, the variability 120 Tulane Studies in Zoology and Botany Vol. 21 seen between features of presumed close functional relation could well be the result of selective pressures for two (or more) different faculties operating on the same feature. Complex rather than simple causative patterns for the correlations ob- served seem likely. All of the support for any particular ecological interpretation of morphological features provided by this study is of a correlative nature. As Sokal and Rohlf (1969, Fig. 15.5) so effectively illustrate, a large number of causal patterns can result in correlations and these patterns need to be sorted out. Certain types of informa- tion would greatly aid in this area. First, experimental studies analogous to those of Gee et al. (1974), Machniak and Gee (1975), and Werner (1974) are necessary for many more features. Second, direct observations of fishes in their natural hab- itats of the sort made by Keeleyside (1962) would be especially useful for many of the habitat related characters. Hopefully the present work will be of heuristic value in indicating some of the features about which observations might be made most gainfully. In answer to the questions posed origi- nally, this study indicates that some signifi- cant portion of the biology of a freshwater stream fish is determinable from its mor- phology. While it is well known that fishes exhibit extreme plasticity of behavior when raised in monospecies cultures, they do tend to specialize in the presence of other species and interspecific competi- tion (Nilsson, 1955, 1960, 1963, 1965, 1967; Ivlev, 1961; Trojnar and Behnke, 1974; Griffith, 1974; Andrusak and Northcote, 1971). Gorman and Karr (1978) report that for stream fish com- munities these specializations occur pri- marily in the selection of specific habitat types and secondarily in the preference for certain food resources if several are readi- ly available within a given habitat. Earlier literature (Thomas, 1962; Gibbons and Gee, 1972; Keast and Webb, 1966; Lind- sey, 1963; Schutz and Northcote, 1972) and the present study both indicate that many of these specializations are related to morphology. The functions and biolog- ical roles that are regularly associated with many morphological features are given above. ACKNOWLEDGMENTS I thank John G. Lundberg for initiating my interest in fishes and providing guid- ance and encouragement throughout this study, Stephen A. Wainwright for pro- voking my interest in morphology, and Daniel A. Livingstone for his influence on my ecological thinking. A number of per- sons aided in the collection of fishes all of whom I thank, but especially Julia B. Lev- erenz and J.G. Lundberg. This research represents a portion of a doctoral disserta- tion which was supported m part by a Co- cos Foundation traineeship in morphol- ogy and by the Department of Zoology of Duke University. LITERATURE CITED Aleev, Yu. G. 1969. Function and gross morphology in fish. Israel Program for Scientific Translations, Jerusalem (Trans!, from Russian.) TT 67-51391. Alexander, R. McN. 1959. The physical properties of the swimbladder in intact Cypriniformes. J. Exp. Biol. 36:315-332. 1967. Functional design in fishes. Hutchinson University Library, London. Al-Hussaini, A.H. 1949. On the functional mor- phology of the alimentary tract of some fishes in relation to differences in their feeding habits. Anatomy and histology. Quart. J. Micr. Sci. 90: 109-139. Andrusak, H., andT.G. Northcote. 1971. Segrega- tion between adult cutthroat trout {Salmo clarki) and Dolly Varden (Salvelinus malma) in small coastal British Columbia lakes. J. Fish. Res. Bd. 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The food and growth of brown trout (Salmo trutta L.) and its feeding relation- ships with the salmon parr (Salmo salar L.) and the eel [Anguilla anguilla (L.)] in the River Teifi, West Wales. J. Anim. Ecol. 31:175-205. Trojnar, J.R., and R.J. Behnke. 1974. Management implications of ecological segregation between two introduced populations of cutthroat trout in a small Colorado lake. Trans. Amer. Fish. Soc. 103:423-430. Tuge, H K. Uchihashi and H. Shimaura. 1968. An atlas of the brains of fishes of Japan. Tsukiji ShokanPubl. Co.. Tokyo. Vincent, R.E. 1960. Some influences of domestica- tion upon three stocks of brook trout {Salvelinus fontinalis Mitchill). Trans. Amer. Fish. Soc. 89: 35-52. Werner, E.E. 1974. The fish size, prey size, handling time relation in several sunfishes and some impli- cations. J. Fish. Res. Bd. Canada 31:1531-1536. JUNE 20, 1979 NICHE RELATIONSHIPS OF THAMNOPHIS RADIX HAYDENI AND THAMNOPHIS SIRTALIS PARIETALIS IN THE INTERLAKE DISTRICT OF MANITOBA DONALD R. HART Department of Biology, Tulane University, New Orleans, Louisiana 70118 ABSTRACT The niches occupied by the western plains garter snake ( Thamnophis radix haydeni) and the red-sided garter snake (Thamnophis sirtalis parietalis) in the Interlake district of Manitoba were compared with respect to feeding habits and six environmental fac- tors. The species were studied both in allopatry and in sympatry. Thamnophis sirtalis occurring in sympatry with T. radix altered its feeding habits, but not in such a way as to produce niche displacement. Its feeding habits did not differ significantly from those of T. radix either in allopatry or in sympatry. Changes in its feeding habits are attributed to changed in the rela- tive availability of food items. Of the environmental factors examined, habitat, air temperature and substrate temperature contrib- uted most strongly to niche discrimination in allo- patry. Thamnophis radix occurred near meadow ponds at high air but low substrate temperatures and T. sirtalis occurred near fen-like marshes of the as- pen forest at lower air but higher substrate temper- atures. In sympatry the contributions of habitat and air temperature to niche discrimination were re- duced, whereas those of light intensity and substrate temperature were increased, T. radix selecting a higher value of both than T. sirtalis. This shift in the importance of environmental factors to niche dis- crimination need not be attributed to species inter- action but may be explained in terms of behavioral compensation for concurrent changes in the avail- able niche. The shift was partially produced by changes in the daily activity pattern, towards mid- day for T. radix and towards morning and evening for T. sirtalis. INTRODUCTION The summer niche requirements of garter snakes are poorly known and the degree of competitive interaction between species is not known at all. Food and habi- tat utilization (Carpenter, 1952; Fleharty, 1967) and food utilization (Fouquette, 1954) have been compared for sympatric populations of garter snakes species in three widely separated localities. None of these authors, however, obtained detailed comparative data for allopatric popula- tions of the same species. Niche displace- ment among sympatric garter snakes, while suggested by these authors as a pos- sible explanation for resource partition- ing, has seldom been properly examined. A notable exception is the work of White and Kolb (1974) on sympatric populations of two garter snake species in California, one of which is represented by a nearby allopatric population. In the Interlake district of Manitoba two garter snake species occur: the wes- tern plains garter snake {Thamnophis ra- dix haydeni Kennicott) and the red-sided garter snake {Thamnophis sirtalis parietal- is Say). While the range of overlap be- tween these species is limited, both occur in sufficiently large numbers to allow the collection of adequate samples from with- in this range. The present study compares the niches occupied by these two species, both within the range of overlap and be- yond it, to determine whether any changes in the relative niche positions were occur- ring that could be attributed to the pres- ence or absence of a congeneric species rather than to changes in the available niche. EDITORIAL COMMITTEE FOR THIS PAPER: DR. CHARLES C. CARPENTER, Professor of Zoology, University of Oklahoma, Norman, Oklahoma 73069 DR. PATRICK T. GREGORY, Assistant Professor of Biology, University of Victoria, Victoria, B.C., Canada V8W 2Y2 125 126 Tulane Studies in Zoology and Botany Vol 21 MATERIALS AND METHODS Field Study. -The study area is located approximately between 50° N and 51° N latitude and between 97° W and 98° W longitude. It is bounded on the east by Lake Winnipeg and on the west by Lake Manitoba as outlined in Figure 1. Snakes were captured within this area from May through August of 1973 and from June through August of 1974. The sampling ef- fort in 1974 was concentrated within a re- gion defined on the basis of the first sum- mer's sampling and included all quarter- townships (23.4 Jcm^) in which both T. radix and T. sirtalis had been captured. This region of sympatry is shown in Figure 1. Severe flooding in May of 1974 pre- vented sampling before June. Snakes were found by searching on foot in the vicinity of water and were captured by hand. Preliminary sampling had shown that garter snakes in the Interlake district were seldom found far from a water body of some sort. Snakes seen crossing roads were not captured since the road habitat is not a natural one and cannot be avoided by snakes in their movements. The samp- ling effort was spread over the daylight hours from sunrise to sunset since prelim- inary attempts to find snakes at night had proven unproductive. No systematic sampling plan was followed due to the difficulty of collecting large samples with- out being somewhat opportunistic in al- location of the sampling effort. Any de- veloping bias in collection with respect to time of day, geography or weather condi- Lake Winnipeg 20 km Fig. 1 . Map of the study area showing major roads, towns (open circles) and approximate lines of continuous (solid) and discontinuous (dashed) aspen forest. Areas included in the sympatric region are stippled. All other areas are included in the allopatric region. The known communal den of T. radix (solid square) and known communal dens of T. sirtalis (solid circles) are located. No. 2 Niche Relationships of Thamnophis 127 tions was soon detected from field records and appropriate corrections were made. The feeding habits of T. radix and T. sirtalis were examined by palpating the stomachs of all snakes captured to force regurgitation of recently eaten food. Due to the difficulty of determining the num- ber of prey represented in partially digest- ed remains, only the presence, and not the number, of food items of each type found in a snake was recorded. Six environmental factors were record- ed at all capture sites to determine the relative importance of each of these fac- tors as niche dimensions. A niche dimen- sion is defined here as any factor serving to separate species ecologically (Levins, 1968) and the importance of an environ- mental factor refers here to its value in separating T. radix and T. sirtalis ecolo- gically. Since biotic, physical and temporal environmental factors were considered, the term "environmental factors" is used in its broadest sense. However, the envir- onmental factors considered were all judged capable, a priori, of eliciting direct response from a snake. Habitat was classified as an environ- mental factor according to the type of water body nearest the capture site of a snake. Water was usually available within several hundred meters of a capture site. Three habitat classes that reflected the proportion of water surface area covered by a sedge mat were recognized. A sedge mat was composed of moss and old sedge decaying just beneath the surface of the water and was overlain by the previous year's growth of sedge, which was folded to form a compact horizontal network at or above the surface of the water. Habitat classes included ponds (no sedge mat; sharp interface between land and water), open marshes (sedge mat at the edges on- ly; open water or sparse emergent vegeta- tion dominant) and closed marshes (dom- inated by sedge mat; little or no open water). The terminology of aquatic com- munities in the aspen forest and parkland has not been standardized. Ponds in the study area conformed to the definition of Radforth (1964). Cover was readily avail- able only in the form of shrubbery or small mammal burrows. Large rocks and logs were rare and did not seem to be used for cover. Heinselman (1963) included the marsh types referred to here as open and closed in his definition of marsh. Closed marshes in the study area resembled fens but were neither sufficiently peaty nor suf- ficiently patterned to conform well to Heinselman's definition of that term. Cover was readily available in the form of sedge mat and encroaching aspen forest. Habitat classes were coded in the above order from 1 through 3. They were judged to be sufficiently discrete in nature to war- rant the use of a discrete variable as a numerical code. A transition from the first habitat class to the last was apparent on a south-west to north-east cline through the study area and corresponded to a transi- tion from parkland to aspen forest. Substrate moisture was classified as an environmental factor on the basis of a simple test performed at the capture site of a snake. If water could not be forced through the fingers by pressing the back of the hand against the substrate with as much force as possible, the substrate was termed dry. If water could just be forced through the fingers by similar action, the substrate was termed damp. If the cup of the hand could be filled with water by similar ac- tion, the substrate was termed water- logged. Two additional moisture classes included standing water (with emergent vegetation) and open water (with no emergent vegetation). Moisture classes were coded in the above order from 1 through 5. This crude method of quantify- ing substrate moisture seemed to correlate . well with the amount of water at the sui^ face although it was certainly not a perfect linear measure. It was considered prefer- able to measurements of water content from soil samples since the latter reflected moisture beneath the surface, which often differed markedly from that found at the surface. Light intensity incident upon the snake at the precise point where it was originally 128 Tulane Studies in Zoology and Botany Vol . 2 1 seen was measured as an environmental The six environmental factors described factor with a Vivitar photographic light above were felt to be of potential impor- meter (Model 43). The rrieter was sensi- tance in characterizing and distinguishing tive only to visible light. Intensity was re- between the niches of T. radix and T. sir- corded in units of exposure value (EV) at talis. For an environmental factor to be an ASA setting of 100 with the light meter important in this sense it is necessary, pointed directly at the sun. though not sufficient, that the snakes be Air temperature at substrate level was (sensitive to it. One way in which all of the recorded at all capture sites with Yellow environmental factors measured seemed Springs Incorporated (YSI) thermistors potentially capable of affecting a snake (Models 43TD and 42SC) and YSI probes ^^s by means of influencing its body tem- (Model 402). The probe was held in the perature. To detect such influences both body shadow of the investigator as close to Qj-al and cloacal body temperatures of the the substrate surface as possible without snakes were taken along with measure- contact, ments of the six environmental factors at Substrate temperature was recorded at g^ch capture site. The same thermistors all capture sites with the same thermistors, used to record air and substrate tempera- Since substrate temperature near the sur- ^ures were used for this purpose. Body face was extremely sensitive to the depth temperatures were always taken immedi- and exact location of the thermistor ately upon capture and with minimal han- probe, temperatures were taken at a jju^g of the animals. The two body tem- depth of 5 cm below the surface. Although peratures were taken consecutively and in this procedure minimized measurement no particular order, error due to minor variations in probe po- Data of the type described above were sition it must be noted that temperatures taken from snakes captured in allopatric at this depth often differ from those exper- ^^d sympatric regions in the summer of ienced by a snake at the contact surface. 1973. An additional sample of snakes was When snakes were found upon a vegeta- collected from within the sympatric region tion mat the surface was defined at the [^ the summer of 1974. Data from this level of the highest part of the snake's sample were pooled with those taken from body that was in contact with the vegeta- ^Yiq same region the year before, tion. Analytical Methods. - Feeding habits of Time of capture of each snake was re- 7 radix and T. sirtalis were compared, corded as an environmental factor on the ^Qth in allopatric and in sympatric re- daylight-saving time scale. It was coded as gions, using a 2xC contingency chi-square an absolute deviation (in hours) from 1400 test for association between species and hours (2 P.M. and the approximate mid- food type utilized. Feeding habits in al- point of the day) so as to reflect the degree lopatric and sympatric regions were also of mid-day activity of the snakes. No dis- compared, both for T. radix and T. sirtal- tinction was made with this time scale be- /^^ using a 2xC contingency chi-square test tween morning and evening, so that the ^Jj. association between region and food scale would correlate roughly with diel type utilized. Columns were pooled in all variations in environmental temperature contingency tables to produce expected and light intensity. Such a scale was desir- values greater than 1 . able in order that the degree of correlation Sampling effort in the field was estimat- of time of capture with temperature or ed for each species in hours spent search- light intensity at capture sites would serve ing for snakes within its range. Daily pro- as an indicator of the extent to which filesof catch per unit effort, calculated for snakes were making use of diel variations hourly intervals throughout the course of in selecting environmental temperatures the day, were compared between T. radix or light intensities. and T. sirtalis both in allopatric and in No. Niche Relationships of Thamnophis 129 sympatric regions. For intervals in which less than three search hours had been spent catch per unit effort was not calcu- lated. Slight biases often developed in the sampling effort before they were noticed and corrected. The effort spent searching for T. radix and T. sirtalis varied slightly throughout the summer in favor of either one species or the other. Since the range of environmental factors available to the snakes also varied throughout the course of the summer, such disparate seasonal allocation of the sampling effort was likely to produce spurious differences between species in the range of environmental fac- tors measured at capture sites. Therefore, seasonal trends in environmental factors were quantified so that environmental data from capture sites could be adjusted for these trends by covariance, using pow- ers of the date (day of the year and day^) as covariates. This method of compensa- ting for seasonal effects in long term data was used by Green (1974) in a similar type of study. It allowed description of season- al trends in the environmental factors as parabolic curves. On the basis of visual inspection of the scatter of points about these curves, this description was consid- ered to be adequate. The frequency distributions of envir- onmental factors were often non-normal. Medians often differed significantly from the means indicating skewness, and 95% confidence intervals on individual obser- vations often exceeded or fell short of the measured range indicating kurtosis. With large samples non-normality has little ef- fect upon comparisons of the means either by univariate (Scheffe, 1959) or by multi- variate (\ioin Krishnaiah, 1969) methods, although it may influence comparisons of variance and covariance more strongly. Heterogeneity of variance, which often accompanies non-normality, was consid- ered more of a problem. Press (1972) cau- tions that although univariate compari- sons of means are little affected by hetero- geneity of variance when sample sizes are large, most assertions about robustness with respect to the assumption of homo- geneity of variance in the muhivariate case are speculative. The reliability of multivariate methods must therefore be considered uncertain to the extent that this assumption is violated. Adjustment of environmental data for seasonal trends, as described above, improved multivariate homogeneity of variance and covariance in the present study. The means of environmental factors tor T. radix and T. sirtalis were compared both by univariate and multivariate meth- ods. Large sample normal deviate tests were used for univariate comparisons. These were applied to environmental data that had not been adjusted for seasonal trends since the seasonal trends shown by two of the environmental factors differed significantly between species. In these cases the description and use of common seasonal trends to adjust the environ- mental data for long term effects was not strictly justified. Discriminant function analysis (Cooley and Lohnes, 1971) was used for multivariate comparison of en- vironmental factors between T. radix and T. sirtalis. Environmental data for use in discriminant function analysis were ad- justed for seasonal trends since this pro- cedure improved multivariate homo- geneity of variance and covariance. The use of seasonally adjusted data may have been somewhat artificial since not all en- vironmental factors showed seasonal trends that were common to both species. However, uncertainty about the robust- ness of multivariate methods to violations of the homogeneity of variance assump- tion made the use of seasonally adjusted environmental data preferable for multi- variate comparison. Multivariate compar- ison was considered more informative than univariate comparison because it takes correlations between the environ- mental factors into consideration. The method of discriminant analysis as used here is analogous to a multiple re- gression of a dependent variable reflecting species membership upon a set of envir- onmental factors. Environmental factors 130 Tulane Studies in Zoology and Botany Vol. 21 which contribute strongly to discrimina- tion between the niches of T. radix and T. sirtalis are weighted heavily in the mul- tiple regression equation, or discriminant function, so as to maximize the ratio of between species to within species variabil- ity in the predicted values of the depen- dent species variable. These predicted values are called discriminant scores. Each snake was therefore assigned a dis- criminant score which was a linear func- tion of the environmental factors meas- ured at its capture site. The difference between the mean discriminant score of T. radix and T. sirtalis is a measure of the overall ecological distance between their niches. An increase in this distance when snakes are collected from a region of sym- patry can be taken as evidence of niche displacement. The magnitude of the weight assigned to an environmental factor in the discrim- inant function depends not only upon the importance of that factor to species dis- crimination but also upon the magnitude of its measurement units. Standardization of all weights equalizes the scale of mea- surement so that the standardized weights represent the relative proportion of spe- cies discrimination contributed to the function by each environmental factor. Multiplication of these standardized weights by the discriminating power of the entire function gives an index of the ecol- ogical importance of each factor which can be compared between different discrim- inant functions. The measure of discrim- inating power used here was the squared canonical correlation coefficeint between the species variable and the environmen- tal factors. This measure represents the fraction of variation in species memberr ship explained by the discriminant func- tion. The index so produced was used to compare the ecological importance of en- vironmental factors between allopatric and sympatric regions. The sign of this index for any one factor indicated the re- lative position of T. radix and T. sirtalis on that environmental gradient. A negative sign indicated that T. radix was found at the lower end of that gradient and T. sir- talis at the upper end. To determine the degree of dependence of body temperatures upon the environ- mental factors, multiple regressions of oral and cloacal temperatures on these fac- tors were calculated separately for T. ra- dix and T. sirtalis. Normal deviate tests were used to compare slopes and inter- cepts between species. RESULTS In the summer of 1973, 137 T. radix and 128 T. sirtalis were captured. Of these 32 T. radix and 36 T. sirtalis were found with- in the sympatric region. A-n additional sample of 73 T. radix and 32 T. sirtalis was collected from within the sympatric region in the summer of 1974. Feeding Habits. -From Table 1 it was apparent that there were no differences in the feeding habits of T. radix and T. sirtal- Js either in the allopatric region (X7- = 9.66" P> .01) or in the sympatric region (X^^ = 10.47; P > .01). The feeding hab- its of T. radix did not differ between al- lopatric and sympatric regions (Xg^ = 11.68; P > .01). The feeding habits of T. sirtalis did differ between allopatric and sympatric regions (X^^ = 13.42; P .01). The change involved an increased utilization in the sympatric region of wood frogs (Rana sylvatica) and a decreased utilization of chorus frogs {Pseudacris triseriata). Environmental Factors. - Seasonal trends in the environmental factors were adequately described as parabolic func- tions of the date. Habitat, as recorded at capture sites, was affected by seasonal movements of the snakes in the allopatric region. As T. sirtalis moved north in the fall towards its denning sites (Gregory and Stewart, 1975) it was found more fre- quently near the closed marshes of the aspen forest. A slight shift in T. radix to- wards the ponds of the open parkland to the south was not significant, but the trends for T. radix and T. sirtalis differed significantly from one another in the alio- No. 2 Niche Relationships of Thamnophis 131 patric region. In the restricted sympatric region seasonal movements of the snakes had less effect on habitat classes and no significant seasonal trends were shown. Moisture, as recorded at capture sites, decreased throughout the summer in the allopatric region as water bodies dried up. In the sympatric region, flooding in the spring of 1974 increased July moisture levels at capture sites. This effect was not apparent until July because the flooding restricted snake collection in June. The effect was more pronounced for T. radix than T. sirtalis because the proportion of T. sirtaUs taken in the sympatric region during the summer of 1974 was severely reduced due to drowning at dens in the spring. This produced significant differ- ences between T. radix and T. sirtalis in the seasonal trends shown by moisture within the sympatric region, even though their individual trends were not signifi- cant. Light intensity decreased through- out the summer as the azimuth of the sun decreased. Air and substrate tempera- tures peaked in mid-summer. Time of cap- ture of the snakes, expressed as a devia- tion from mid-day, showed no significant seasonal trends. Univariate comparisons of environ- mental factor means between the niches of T. radix and T. sirtalis, applied to envir- onmental data that had not been adjusted for seasonal trends, showed interspecific differences in habitat, moisture, air tem- perature and substrate temperature to be significant (P < .01) in the allopatric re- gion (Fig. 2). Similar comparisons in the sympatric region showed significant (P, < .01) interspecific differences in habitat, light intensity and substrate temperature. These differences remained apparent after adjustment of the environmental data for seasonal trends. Discriminant scores derived from seasonally adjusted environmental data gave better discrimi- nation between the niches of T. radix and T. sirtalis than did any single environ- mental factor, both in allopatric and in sympatric regions, but did not give any evidence of niche displacement. On the contrary, discriminant scores became more similar in the sympatric region than in the allopatric region (Fig. 3). Multivari- ate results agreed well with those derived by univariate methods, although some dif- ferences were apparent (Table 2). The in- dex of ecological importance, derived from the weights in the discriminant func- tion, was large for habitat, both in allo- patric and in sympatric regions, although it became smaller in the sympatric region, indicating reduced importance there. The negative sign of this index indicates that T. radix occupied the lower end of the habi- tat gradient (ponds) and T. sirtalis the up- per end (closed marshes). Substrate mois- ture, in contast to univariate results, was not ecologically important in either re- gipn. Light intensity was important only in the sympatric region with T. radix occupy- ing the upper (sunny) end of the gradient whereas air temperature was important only in the allopatric region, with T. radix occupying the upper (warm) end of the gradient. Substrate temperature was im- portant in both allopatric and sympatric regions, although it become more impor- tant in the latter region. Associated with the increased importance of this factor in the sympatric region, in contrast to uni- variate results, was a reversal in the rela- tive species positions on the gradient, T. radix coming to occupy the upper (warm) rather than the lower (cool) end. Time of capture, expressed as a deviation from mid-day, was not an ecologically impor- tant factor in either the allopatric or the sympatric region. These results agree with those based on univariate comparisons, although in the sympatric region they give considerably less importance to the time factor. Thermal Relations. -Oral and cloacal temperatures of T. radix were significant- ly higher (P < .01) than those of T. sirtalis in the allopatric region only. Body tem- peratures of T. sirtalis increased signifi- cantly (oral P < .05; cloacal P< .01) in the sympatric region (Fig. 4). Frequency dis- tributions of body temperatures were negatively skewed, as shown by the fact 132 Tulane Studies in Zoology and Botany Vol.^l that the median temperatures were always higher than the means, indicating that oc- casionally exceptionally cold snakes were encountered. Multiple regression showed oral tem- peratures of T. radix to depend most strongly upon light intensity, air temper- ature and substrate temperature (Table 3). Cloacal temperatures of T. radix de- pended most strongly upon the same three environmental factors. Of these three fac- tors air temperature was most important. Oral temperatures of T. sirtalis also de- pended most strongly upon light intensity, air temperature and substrate tempera- ture. Habitat exerted an additional effect upon oral temperatures of T. sirtalis, over and above that produced by the other en- vironmental factors, oral temperatures being lower in the closed marsh. Cloacal temperatures of T. sirtalis depended most strongly upon air temperature and sub- strate temperature and were independent of light intensity. Cloacal temperatures, like oral temperatures, were lower in the closed marsh, other environmental factors being equal. An additional effect seemed to be exerted upon the cloacal tempera- tures of T. sirtalis by substrate moisture and time of capture, cloacal temperatures being higher at moist capture sites and lower near mid-day, other factors being equal. The relationships of oral and cloac- al temperature to the habitat factor dif- fered significantly (P < .01) between T. radix and T. sirtalis, being negative for T. sirtalis only. The relationships of cloacal temperature to substrate moisture, sub- strate temperature and time of capture differed significantly (P < .01) between T. radix and T. sirtalis, being more positive for T. sirtalis. Activity.- A bimodal pattern of daily activity was suggested by daily profiles of catch per search hour in the field for both T. radix and T. sirtalis in the allopatric region (Fig. 5). Effort in this region ranged from 3 to 22 search hours per hour- ly time interval. In the sympatric region the morning and evening peaks of activity for T. sirtalis were farther apart than in the allopatric region, while T. radix eliminat- ed its morning peak and became more active towards mid-day. Effort in this re- gion ranged from 4 to 41 search hours per hourly interval and catches were generally lower for both species than in the allopat- ric region. DISCUSSION Feeding Habits. -Both T. radix and T. sirtalis appeared to feed opportunistically, taking a wide variety of food items in rough proportion to their apparent abun- dance. This may be considered an adapta- TABLE 1. Number of T. radix and T. sirtalis stomachs containing various food items in allopatric and sympatric regions. FOOD Allopatric Region Sympatric Region CLASS T. radix T. sirtalis T. radix T. sirtalis Rana sylvatica 24 15 36 24 Rana pipiens 4 0 3 0 Pseudacris triseriata 8 11 7 4 Hyla versicolor 2 2 1 3 Bufo 3 3 4 2 Tadpoles 5 0 1 0 Rodentia 1 0 1 0 Oligochaeta 5 2 0 0 Hirudinea 6 6 6 0 Gastropoda 0 0 2 0 No. 2 Niche Relationships of Thamnophis 133 H CO O ¥mu +-E^ I i crys^ 4 — ^ > H o C^im^ -t 'il \'.}y\'*'\ — 1 » Il 1 — 1 — 1| 9 10 11 12 13 1 i, :Ak -. 1 14 15 16 1 •. "J 1 1 — -, — 1 1 1 — ? — 1 1 o O Eh 1 I ■ <"Ti 1 |li^(.".") r • 1 1 t — ?— M 16 17 18 19 20 21 22 23 24 25 2*6 (' - l*''j- rl 27 28 2'9 30 31 I .'-I* . • ■ .1 t-h-d — o PL. Eh n & ^::'.4^ I I I I 12 13 14 15 1^ ?7 18 l'9 2*0 2'l 22 23 24 2^ 26 27" ^ -^ U S cr^^ -2 J i3— 5 6 /^/g. 2. Location of 7. radur (open rectangles) and T. sirtalis (stippled rectangles) on gradients of the six environmental factors measured at capture sites. Rectangles indicate 95% confidence intervals on the means (solid circles). Medians (vertical lines) and 95'7r confidence intervals on the observations (horizontal lines) are also shown. The axis of an environmental factor is broken beyond the measured range of that factor. Statistics above the axis pertain to the allopatric region. Statistics below the axis pertain to the sympatric region. Time is measured in units of hours + 2 P.M. Habitat code 1 = pond; 2 = open marsh; 3 = closed marsh. Moisture code 1 = dry; 2 = damp; 3 = waterlogged; 4 = standing water; 5 = open water. 134 Tulane Studies in Zoology and Botany Vol. 21 tion to the short growing season encoun- tered in the temperate zone. In contrast to the studies of Carpenter (1952), Fou- quette (1954) and Fleharty (1967), who worked with different species combina- tions, no differences were found between the feeding habits of T. radix and T. sirtal- is in sympatry. Gregory's (1977a) limited observations on the feeding habits of T. radix captured largely in sympatry with T. sirtalis invited similar conclusions. Al- though T. sirtalis altered its feeding habits in the sympatric region, it did not do so in such a way as to displace its niche from that of T. radix. The major changes in- volved an increased utilization of wood frogs and a decreased utilization of chorus frogs. These changes may have been the result of variations in the relative availa- bility of wood frogs and chorus frogs. Since there was no niche displacement in sympatry with respect to feeding habits, the changes cannot be attributed to spe- cies interaction. White and Kolb (1974), in their study of two garter snakes species in California, similarly rejected the hy- pothesis of competitive displacement since changes in the feeding habits of T. sirtalis fitchi, when in sympatry with T. elegans, were not in the direction of niche displacement. They felt that feeding hab- its were determined largely by availabil- ity, which interpretation is also suggested in the present study. Enviornmental Factors. -In the allo- patric region, habitat, air temperature and substrate temperature contributed most strongly to niche discrimination (Table 2). Thamnophis radix occurred near ponds at high air but low substrate temperatures while T. sirtalis occurred near marshes at lower air but higher substrate tempera- tures. Although univariate comparisons between T. radix and T. sirtalis also at- tributed importance to substrate mois- ture, multivariate results indicated that this factor was superfluous due to its strong correlation with other environ- mental factors of greater value in dis- criminating between species. Substrate temperature was a particularly strong negative correlate of substrate moisture. The ecological importance of habitat can be related to the thermal responses of the snakes. Habitat exerted an indepen- dent effect upon both oral and cloacal temperatures of T. sirtalis; this was prob- ably due to the shade provided by the sedge mat in the closed marsh. Large ex- panses of sedge mat with associated stands of aspen forest provide a readily accessible thermal refuge. This type of cover may have been particularly important to T. sir- talis, which is more darkly colored than T. TABLE 2. Discriminant function coefficients (unstandardized) and indices of discrim- inating value derived from them in allopatric and sympatric regions. ALLOPATRIC Habitat Moisture Light Air Temp. Sub. Temp. Time Rc^* Coefficient Index** SYMPATRIC -30.729 - .502 Habitat -2.922 2.021 - .066 .085 Moisture Light 1.110 .117 Air Temp. -1.227 - .103 Sub. Temp. -1.703 - .069 Time .54 Rc^* Coefficient Index** -6.117 - .163 -0.243 1.660 - .010 .142 -0.105 - .022 0.577 .104 -0.661 - .055 .25 -T *Rj> = Squared cannonical correlation coefficient (fraction of variation in species membership accounted for by the discriminant function) "Standardized discriminant function coefficients multiphed by R N (). Niche Relationships of Thamnophis 135 radix in the Interlake district and perhaps subject to overheating in the summer (Hart, 1975). Such cover was conspicu- ously absent from the pond habitat util- ized extensively by T. radix. Cover was similarly important to niche discrimina- tion in the studies of Carpenter (1952) and Fleharty(1967). The importance of air and substrate temperature to niche discrimination in the allopatric region also reflects the thermal relations of the snakes. Both factors ex- erted a strong effect upon oral and cloacal temperatures of T. radix and T. sirtalis. Since T. radix is lighter in color than T. sirtalis in the Interlake district, it may have been better adapted to the higher air tem- peratures that characterized its range and able to tolerate them by seeking out low substrate temperatures. Thamnophis sir- talis was apparently unable to tolerate high air temperatures without a thermal refuge in the form of closed marsh or as- pen forest and, probably for this reason, it did not occur much beyond the range of these habitat types. Fleharty (1967) simi- larly found air and substrate temperatures to be useful in defining the niches of garter snake species. In the restricted sympatric region, the available range of environmental factors was probably reduced. As a result, the importance of habitat and air temperature to niche discrimination was also reduced, the latter becoming insignificant. Howev- er, the importance of substrate tempera- ture and light intensity at capture sites was increased (Table 2). This can only be in- terpreted as a behavioral reaction of the snakes, but need not be explained as a reaction to the presence of another spe- cies. It may represent compensation for changes in the available niche, T. radix seeking out higher light intensities and substrate temperatures to compensate for lower available air temperatures in the sympatric region; T. sirtalis seekmg out lower light intensities and substrate tem- peratures to compensate for higher avail- able air temperatures in the sympatric re- gion and for the scarcity of shade normally afforded by the closed marsh and forest of its allopatric habitat. One of the ways in which this selection could have been accomplished is suggest- ed by the daily profiles of catch per search hour. In the allopatric region, both species showed a roughly bimodal activity pattern with morning and evening peaks. In the sympatric region, however, T. radix was most active during mid-afternoon close to the hottest part of the day, and T. sirtalis was active earlier in the morning and later in the evening, apparently avoiding the heat of mid-afternoon. This increased dif- ference between the activity patterns of T. radix and T. sirtalis in the sympatric region was illustrated by univariate comparisons of time to capture, coded to reflect mid- day activity. The difference in time of cap- ture between T. radix and T. sirtalis was increased in the sympatric region, but was not great (.05 > P > .01). Multivariate re- sults virtually eliminated the increased importance of this factor in the sympatric region indicating that the activity shift was entirely due to the selection of higher light intensities and substrate temperatures by T. radix and lower ones by T. sirtalis. Fig. 3. Location of T. radix and T. sirtalis on the axes of the discriminant functions (DF) derived from seasonally adjusted data in allopatric and sympatric regions. Symbols and format as in Fig. 2. 136 Tulane Studies in Zoology and Botany Vol. 21 These changes in selection of light inten- sity and substrate temperature must rep- resent behavioral adjustments other than those reflected in the daily activity pat- tern. MacArthur and Levins (1967) showed that when ecological similarity surpasses a minimal limit, interspecific selective forces act to produce competitive niche displacement. While the relative niche positions of T. radix and T. sirtalis changed in the sympatric region, there was no net displacement of the niches oc- cupied (Fig. 3). This does not preclude a competitive interaction since there was probably an increased similarity of the available niches. It does, however, sug- gest that competitive interaction was of minor significance if it occurred at all. The relationship between limiting similarity and resource availability in competitive interactions has been discussed by May (1974), Riebesell (1974) and Pianka. 1974). Schoener (1974) has pointed out that resource availability is an unmeas- ured variable in most studies of resource partitioning. Measurement of niche sep- aration relative to the available niche breadth was not attempted in the present study due to the difficulty of quantifying an available as opposed to occupied niche with respect to the environmental factors considered. Thermal Relations. -The higher body temperatures of T. radix in the allopatric region (Fig. 4) suggest that this species has a slightly higher preferred temperature than T. sirtalis. Such a difference would correlate well with differences in thermal tolerance. Thamnophis sirtalis is reported to have a slightly lower minimum critical temperature than T. radix (Fitch, 1965). Vincent (1971) found T. Sirtalis hibernat- ing at body temperatures as low as -3°C while Bailey (1949) did not find T. radix to survive hibernation at temperatures lower than -2*-*C. While cloacal body tempera- tures of T. sirtalis upon capture were slightly lower than those reported by Fitch (1965) as "preferred", body temperatures of reptiles captured in the field are often found to be lower than preferred tempera- tures (Licht et al., 1966) due, perhaps, to greater catchability at sub-optimal tem- peratures. In spite of the ecological adjustments made by both species in the sympatric re- gion, in apparent compensation for mar- ginal air temperatures, body temperatures became more similar there. This was pri- marily due to an increase in the body tem- perature of T. sirtalis, which probably in- dicates that T. sirtalis had passed its limit of behavioral thermal adjustment and may explain why it was unable to pene- o U < o li^'' v»:'>:'l| 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 + 34- o O -J < < o u ■^ f^^^vH^ 1 : h 1 ' — ' 1 19 20 21 22 23 24 25 26 27 28 29 3b 31 32 33 34 r-j-'i*!""'-"! 1."* "'•'■'.i-!i;t 1 - 1 • D Fig. 4. Oral and cloacal temperatures of T. radix and T. sirtalis in allopatric and sympatric regions. Symbols and format as in Fig. 2. Ni). Niche Relationships of Thamnophis 13 7 trate further into the range of T. radix. Body temperatures of T. radix were not ahered appreciably in the sympatric re- gion, and in fact showed a very slight in- crease in spite of the fact that T. radix there was at the northern periphery of its range. Thamnophis radix, therefore, had not passed its Umit of behavioral thermal adjustment, but was probably at or very close to this limit in the sympatric region. Neither T. radix nor T. sirtalis would have been able to alter their activity patterns much further since T. radix was already concentrating its activity in the hottest part of the day and T. sirtalis had virtually eliminated its mid-day activity. Negative skewness in the frequency distributions of reptilian body tempera- tures is often reported (Cowles and Bo- gert, 1944; Brattstrom, 1965). As Heath (1964) points out, although this pattern is often interpreted as indirect evidence for a photophobic reaction at body tempera- tures higher than the modal one, similar patterns are produced by inanimate ob- jects incapable of movement, and need not be attributed to behavioral thermo- regulation. The significance of the pattern in the body temperature distributions of T. radix and T. sirtalis is, therefore, uncer- tain. The oral and cloacal body temperatures of T. radix and T. sirtalis depended most strongly upon air temperature (Table 3) and next most strongly upon substrate temperature. Fleharty (1967) related cloacal temperatures of T. elegans, T. cyr- topsis and T. rufipunctatus primarily to substrate temperature and found little effect due to air temperature. However, Fleharty measured air temperature 24 cm above ground level rather than immedi- ately above ground level as in the present study. He measured substrate tempera- ture right at ground level so that it repre- sented the combined effects of air and substrate temperature as measured in the present study. Gregory (pers. comm.) found cloacal temperatures of T. or- dinoides, T. elegans and T. sirtalis picker- ingi to depend strongly upon both air and substrate temperatures; the latter, meas- ured immediately below ground level, exerted the greatest effect. Oral temperatures of both T. radix and T. sirtalis depended strongly upon light intensity. While the cloacal temperatures of T. radix also depended upon this factor, those of T. sirtalis did not. This may be explained by the fact that T. sirtalis was often captured on the sedge mat of a closed marsh, with its head above the mat TABLE 3. Regression equations for dependence of oral and cloacal temperatures on environmental factors [Y = I -H (H x Habitat) -h (M x Moisture) + (L\ Light) + (A x Air Temp. ) + (S x Sub. Temp. ) + (T x Time )] T. radix T. sirtalis Oral Cloacal Oral Cloacal I 6.941 -1.620 H 0.259 0.222 M -0.485 -0.525 L 0.626* 0.358* A 0.448* 0.513* S 0.165* 0.164* T -0.239 -0.210 4.420 -0.113 1.081** -1.090** 1.460 3.048** 0.448* 0.073 0.536* 0.635* 0.273* 0.382** 0.029 0.333** * Partial regression coefficient non-zero (P <.01 ) '* Partial regression coefficient non-zero and different from that of the other species (P <.0!) 138 Tulane Studies in Zoology and Botany Vol.21 and its tail below, shaded from the sun. Light intensity was not measured under the sedge mat. Oral and cloacal temperatures of T. sir- talis were influenced by habitat. The pro- tection from the sun offered the snake by the sedge mat and forest of the closed marsh habitat probably explains this ef- fect. The direct effect of substrate moisture upon the cloaca! temperatures of T. sirtal- is, other factors being equal, may have been due to relative humidity near the substrate, which was not measured, a di- rect effect of relative humidity upon body temperatures of reptiles, due to inhibition of evaporative cooling, was suggested by Cowles and Bogert (1944). Relative hu- midity was probably high under the sedge mat of a closed marsh. Such an effect probably is not completely explained by the habitat factor since there was consid- erable variability in the moisture of a closed marsh between the edges and the center. 3 1 ALLOPATHIC a p o X \ X o < 1 i T sirtalis 67 8 910 11 12 1234 AM TIME OF DAY SYMPATRIC 5 6 8 9 PM D O X \ X o < 1 ■ T. radix 6 7 8 9 10 11 12 1 2 3 4 A M TIME OF DAY 5 6 7 8 9 PM Fig. 5. Profiles of catch per search-hour calculated for hourly intervals throughout the course of the day. Profiles are shown for T. radix (solid squares) and T. sirtalis (solid circles) in allopatric and sympatric re- gions. The direct effect of time of capture, coded as a deviation from mid-day, upon the cloacal temperatures of T. sirtalis is difficult to interpret. Diel variations in other measured environmental factors such as light intensity, air temperature and substrate temperature cannot, in a mul- tiple regression, explain an independent effect due to time of capture. Such varia- tions would, in a simple regression, be expected to result in an inverse rather than direct relationship between body temper- ature and time of capture, with higher body temperatures closer to mid-day. The observed effect of time of capture upon the cloacal temperatures of T. sirtalis over and above those produced by variation in other measured factors suggests either the influence of unmeasured environmental factors exhibiting a diel cycle or an intrin- sic diel cycle in cloacal temperature. Activity. -The. degree of bimodality in the daily activity pattern of T. radix was found by Heckrotte (1962) to,be temper- ature dependent. A similar temperature dependence appears in the daily activity patterns of both T. radix haydeni and T. sirtalis parietalis (Hart, 1975) and explains the opposite adjustments in activity pat- tern made by T. radix and T. sirtalis in the sympatric environment (Fig. 5) and the increased importance of time of capture in discriminating between the niches of T. radix and T. sirtalis in this region. By be- coming most active near mid-day, T. radix was able to select higher light intensities and substrate temperatures. By avoiding the heat of mid-day, T. sirtalis was able to select lower light intensities and substrate temperatures. Jordan (1967), on the basis of a small sample of snakes from roads within a region of sympatry in Minnesota, similarly found T. sirtalis to be active ear- lier in the morning and later in the evening than T. radix. Competitive Interaction and Density- Dependence. —The primary importance of physical as opposed to biotic factors in the ecology and evolution of temperate zone animals and plants has been stressed by Dobzhansky (1950) and Fischer (1961). Bogert (1949) suggested that this was es- pecially true of northern poikilotherms No. 2 Niche Relationships of Thamnophis 139 and noted that stringent thermal require- ments might prevent speciaUzation with respect to density-dependent ecological factors. The importance of thermal factors to northern reptiles has been illustrated by Gregory (1977b) who attributed the dis- appearance of the youngest year class at a communal den of T. sirtalis to reproduc- tive failure brought on by a cool, cloudy summer. It is illustrated in the present study by the fact that ecological adjust- ments made by both T. radix and T. sirtalis at their range peripheries involved those niche parameters most strongly related to body temperatures. Marked reduction in the relative frequency of T. sirtalis cap- tures following spring flooding of many of its dens during the present study further emphasizes the significance of climatic factors in regulating populations of north- ern reptiles. These factors act indepen- dently of density and often catastrophical- ly. The similar feeding habits of T. radix and T. sirtalis in Manitoba and the lack of evidence for niche displacement between them suggest that density-dependent fac- tors such as competitive interaction are probably not important to northern poiki- lotherms enduring a harsh continental climate. However, the food partitioning among three Thamnophis species on Van- couver Island (Carr and Gregory, 1976; Gregory, 1978) is suggestive of competi- tive interaction. Such density-dependent factors may be important to northern poikilotherms in maritime regions where the climate is more moderate. This paper is based upon a thesis sub- mitted to the University of Manitoba in partial fulfillment of the requirements for the degree of Master of Science. I wish to thank Dr. K. W. Stewart of the University of Manitoba for his supervision and advice during all stages of the study. The thesis version of the manuscript was critically reviewed by Dr. W. O. Pruitt, Dr. J. C. Rauch and Dr. R. J. MacKay, all of the University of Manitoba. The manuscript in its present form was critically reviewed by Dr. H. A. Dundee of Tulane Univer- sity and Dr. J. S. Rogers of the University of New Orleans. I was ably assisted in the field by Mr. Graham Latonas. LITERATURE CITED Bailey, R.M. 1949. Temperature toleration of garter snakes in hibernation. Ecology 30: 238-242. Bogert, CM. 1949. Thermoregulation in reptiles: A factor in evolution. Evol. 3: 195-211. Brattstrom, B.H. 1965. Body temperatures of rep- tiles. Amer. Midi. Natur. 73: 376-422. Carr, CM. and Gregory, P.T. 1976. Can tongue flicks be used to measure niche sizes? Can. J. Zool. 54: 1389-1394. Carpenter, CC 1952. Comparative ecology of the common garter snake {Thamnophis sirtalis sirtal- is), the ribbon snake (Thamnophis sauritus) and Butler's garter snake (Thamnophis butleri) in mixed populations. Ecol. Monogr. 22: 235-258. Cooley, W.W. and Lohnes, P.R. 1971. Multivariate data analysis. Wiley and Sons Inc., Toronto. Cowles, R.B. and Bogert, CM. 1944. A preliminary study of the thermal requirements of desert rep>- tiles. Bull. Amer. Mus. Nat. Hist. 83: 261-2%. Dobzhansky, T. 1950. Evolution in the tropics. Amer. Sci. 38:208-221. Fischer, A.G. 1961. Latitudinal variation in organic diversity. Amer. Sci. 49: 50-74. Fitch, H.L. 1965. An ecological study of the garter snake Thamnophis sirtalis. Univ. Kansas Publ. Mus. Nat. Hist. 15: 493-564. Fleharty, E.D. 1967. Comparative ecology of Thamnophis elegans, Thamnophis crytopsis and Thamnophis rufipunctatus in New Mexico. Southwest. Natur. 12: 207-230. Fouquette, M.J. 1954. Food competition among four sympatric species of garter snakes. Texas J. Sci. 6: 172-188. Green, R.H. 1974. Multivariate niche analysis with temporally varying environmental factors. Ecol- ogy 55: 73-83. Gregory, P.T. and Stewart, K.W. 1975. Long-dis- tance dispersal and feeding strategy of the red- sided garter snake (Thamnophis sirtalis parietalis) in the Interlake of Manitoba. Can. J. Zool. 53: 238-245. 1977a. Life history observations of three species of snakes in Manitoba. Can. Field Natur. 91: 19-27. . 1977b. Life-history parameters of the red-sided garter snake ( Thamnophis sirtalis parietalis) in an extreme environment, the Inter- lake region of Manitoba. Nat. Mus. Canada, Pubis. Zool. No. 13,44 pp. 1978. Feeding habits and diet overlap of three species of garter snakes (Thamnophis) on Vancouver Island. Can. J. Zool. 56: 1967-1974 Hart, D.R. 1975. A quantitative niche comparison of the western plains garter snake (Thamnophis ra- dix haydeni) and the red-sided garter snake ( Thamnophis sirtalis parietalis) in allopatric and sympatric regions of Manitoba's Interlake Dis- trict. M.Sc. Thesis, University of Manitoba, Winnipeg, Canada. 140 Tulane Studies in Zoology and Botany Vol. 21 Heath, J.E. 1964. Reptilian thermoregulation: eval- uation of field studies. Science 146: 784-785. Heckrotte, C. 1962. The effect of the environmental factors in the locomotory activity of the plains garter snake (Thamnophis radix radix). An. Be- hav. 10: 193-207. Heinselman, M.L. 1963. Forest sites, bog processes and peatland types in the glacial Lake Agassiz region, Minnesota. Ecol. Monogr. 33: 327-374. Ito, K. in Krishnaiah, P.R. (ed.) 1969. Multivariate Analysis II. Academic Press, New York. Jordan, O.R. 1967. The occurrence of Thamnophis sirtalis and Thamnophis radix in the prairie-forest ecotone west of Itasca State Park, Minnesota. Herpetoiogica 23: 303-308. Levins, R. 1968. Evolution in changing environ- ments. Princeton University Press, Pnnceton, New Jersey. Licht, P., Dawson, W.R., Shoemaker, V.H. and Main, A.R. 1966. Observations on the thermal relations of Western Australian lizards. Copeia 1966:97-110. MacArthur, R. and Levins, R. 1967. The limiting similarity, convergence and divergence of co- existing species. Amer. Nat. 101: 377-385. May, R.M. 1974. On the theory of niche overlap. Theor. Pop. Biol. 5: 297-332. Pianka, E.R. 1974. Niche overlap and diffuse com- petition. Proc. Nat. Acad. Sci. 71: 2141-2145. Press, S.J. 1972. Applied multivariate analysis. Holt, Rhinehart and Winston, Inc., New York. Radforth, N.W. 1964. Prerequisite for design of en- gineering works on organic terrain - a symposi- um. Part II - Definitions and terminology. NRC Assoc. Cettes on Soil and Snow Mechanics, Tech. Memo. 81:24-35. Riebesell, J.F. 1974. Paradox of enrichment in com- petitive systems. Ecology 55: 183-187. Scheffe, H. 1959. The analysis of variance. John Wiley and Sons, New York. Sc*iocner, T.W. 1974. Competition and the form of habitat shift. Theor. Pop. Biol. 6: 265-308. Vincent, T. 1971. Resistance to cold stress in the red-sided garter snake Thamnophis sirtalis pari- etalis. M.Sc. Thesis, University of Manitoba, Winnipeg, Canada. White, M. and Kolb, J. A. 1974. A preliminary study of Thamnophis near Sagehen Creek, California. Copeia 1974: 126-136. JUNE 20, 1979 TULANE STUDIES IN ZOOLOGY AND BOTANY VOLUME 21 INDEX TO SCIENTIFIC NAMES Note: No4. containing a single article, "A Bibliography of Louisiana Botany, 1951- 1975", is completely cross indexed on pages 58-66 of that article. Allocreadiidae, 68, 77 Allocreadium ictaluri, 67, 78 Alternanthera sp. , 70 Ancylidae, 67-88 Anguillarostrata 92-93, 107, 116-117, 119 Aphredoderus sayanus 92, 103-109, 114, 116-117 armatae group, 78 Bithynia tentaculata , 80 Castor canadensis, 80 Castostomus commersoni, 92, 103-107, 109, 116-117 Centrarchidae, 102 Centrarchus macropterus, 92, 103-107, 109, 116-117 Cercaria minimum, 76 type I, 71-73, 82-83, 85 type II, 71-74, 81-82, 85-86 type III, 73, 76, 81, 83, 85-86, 88 type IV, 73, 76, 81-83, 85-86 type V, 73, 76-77, 81 type VI, 73, 77, 81-83, 85 type VII, 74, 77, 83 type VIII, 74, 77, 83 type IX, 74, 77-78, 83 type X, 74, 78, 83 typeXI,74, 78, 83 type XII, 74, 78, 82 typeXIII, 74, 81,86 type XIV, 74,79,81,86 type XV, 75, 79, 81-82 type XVI, 75,79-81 type XVII, 75, 80, 82, 84 typeXVIII, 75,80, 86 typeXIX, 75,80, 84, 86 Cercariaeum mutabile, 68 type I, 78 Chaetogaster limnaei, 68 Chrysemys scripta, 84 Clinostomus funduloides, 92, 103-107, 109, 116, 118 Cyprinidae, 102 Dugesia, 68 doctorocephala, 68 tigrina, 68 Echinochasmus , 11 Echinostomatidae, 68 Eichornia crassipes, 71 Enneacanthus gloriousus, 92, 103-107, 116-117 Erimyzon oblongus, 92, 103-107, 109, 116-117 Esocidae, 102 Esoxamericanus, 92, 103-107, 109, 114, 116-118 niger Etheostoma flabellare fusiforme, 92, 109, 114, 116-118 olmstedi, 92, 103-107, 109, 114, 116-119 Ferrissia, 79, 87-88 fragilis, 67-72, 74, 76-88 parallela, 68 rivularis, 68 Gambusiaaffinis, 92, 103-107, 109, 116-118 Gyraulus similaris, 68, 79 Haematoloechus breviplexus, 68, 79 Hebetancylus, 87-88 excentricus, 69-72, 76-78, 81-88 Helisoma companulata, 68 trivolvis, 67-68 Hybopsis hypsinotus, 92, 103-107, 109, 116, 118 Ictaluridae, 102 Ictalurus cams, 92, 103-107, 114, 116, 118-119 nebulosus, 92, 103-107, 109, 114, 116-117, 119 punctatus, 92, 103-107, 114, 116, 118-119 Laevapex, 78, 87-88 fuscus, 67-72, 74, 76-78, 80, 83-88 Lepomisauritus, 92-93, 103-110, 116-117 cyanellus, 92, 103-110, 114, 116-117 gibbosus, 92, 103-107, 109, 116-117 gulosus, 92, 103-107, 109, 114, 116-117 macrochirus, 92, 103-107, 109, 114, 116-117 Lissorchiidae, 68 Lissorchis sp., 86 Lymnaea ovata, 80 Megalodiscus temperatus, 67-68, 74, 76, 88 Micropterussalmoides, 92, 103-107, 109-110, 114,116,117 Moxostoma anisurum 92, 103-107, 116-117 erythrurum, 92, 103-107, 116, 118 pappillosum, 92, 103-107, 116, 118 robustum, 92, 103-107, 116, 118 Nocomis leptocephalus, 92, 103-107, 109, 116, 118 Notemigonus crysoleucas, 92, 103-107, 109, 116, 118 Notropisalborus, 92, 103-107, 109, 116-117 altipinnis, 92, 103-107, 109, 116, 118 analostanus, 92, 103-107, 109, 116, 118 ardens, 92, 103-107, 109, 116, 118 cerasinus, 92, 103-107, 109, 116, 118 chiliticus, 92, 103-107, 109, 116-118 hudsonius, 92, 103-107, 116-117 procne, 92, 103-107, 109, 116, 117-118 Noturusinsignis, 92, 103-107, 109, 116, 117-118 Nycticorax nycticorax, 76 orantae group, 79 Paspalum sp., 71 PercafJavescens, 92, 103-107, 109, 116-117 Percidae, 102 Percina crassa, 92, 103-107, 116-117 Phoxinus areas, 92, 103-107, 109, 116-117 Physa halei, 76 pigmentata group, 80 Planorbis vortex, 80 Pomoxis nigromaculatus, 80 Posthodiplostomum minimum, 76, 88 Pseudacris triseriata, 130 Ranasylvatica, 130 Sagittaria, 71 Salmoclarki, 113 Salvelinus malma, 113 Semotilus atromaculatus, 92, 103-107, 109, 114, 116,118 Sphaeriidae, 68, 86 Spirorchiidae, 68 Spirorchis scripta, 68, 77, 84 Stichorchis subtriquetrus , 80, 84, 88 Succinea putris, 80 Thamnophis cyrtopsis, 137 elegans, 134, 136-137 ordinoides, 137 radix haydeni, 125-136 rufipunctatus, 137 sirtalisfitchi, 134 sirtalis parietalis, 125-136 sirtalis pickeringi, 137 Triganodistomum mutabile, 68 Umbra pygmaea, 91, 103-107, 116-117 cAcme Bookbindinfl Co.. 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