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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
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send their remittance, preferably money order, along with their orders. Remittances
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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
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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.

[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.

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.

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

[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

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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]

■'. . . 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.

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.

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.

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.

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.

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.

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.

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.

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."

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.

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.

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.

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.

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.

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.

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

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.

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.

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.

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.

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.

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).

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.

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.

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.

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.

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.

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.

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.

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

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

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

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

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

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

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

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

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

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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

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

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

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

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.

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

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.

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

Figures 15-19. Cercaria types XV-XIX developing in southeastern Louisiana ancylids. Scale in microns.

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

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

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

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

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

c
o

•^ 30
O 20

^ 10

C 0

o '^

I 0
5
0
I 5
I 0
5
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0)

100
80

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\^ 40

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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 ^

6 ° ^--o-.o— 0--0

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•

40 "D.E

<i>E

y c

1 0

iJ«J

0
1 0

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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.

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

(/>

40 _

20 ^

a^iB

10 O

0 0)

503

4U ^

if)

20 0

10 C
0

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

■ 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

C 4

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■o

o ?

O) 1

C 0

■ ■^

■ ^^

100

8 0

■ ^IM

100

8 — e

*—-»--©■■

>-i)^-^

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illl

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

E
in

o
o

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.

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

c c

o \

c
o

0
0
0
0
0
9
6
3
0

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 )

I I I I I I I I I I r
JJASONDJFMAMJJ

100

40
20

1 0

^^^

0.8

O.b

0.4

0.2

0.0

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.

Tulane Studies in Zoology and Botany

Vol. 21

2 5

o ^

1 5

1 0

o c

c o

0 -^

2 5

■D O 20

oi>

1 5

c c

1 0
5
0

R 0

■ ^^

8 0

2 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

50 (/)

30 ^ ^

20 O \

10 <D C

° o t

20 lO

10 c 0)

5 Q,

Month

Figure 24. Monthly incidence of infection, population structure, and index of relative abundance for (H.
Hebetancylus excentricus and (F.) Ferrissia fragilis at Head of Island Pond.

No. 2

Limpet Snails

the ecology of the respective trematode
faunas. Seasonahty of infection in L. fus-
cus at RAM was indicative of a single,
annual host generation; however, two
cercarial species were maintained essen-
tially year round in SOR L. fuscus, and
thus reflected relative stability in the L.
fuscus population.

Head of Island Pond (Figure 24) was
productive enough to permit a trivoltine
pattern for Hebetancylus excentricus, but
was faunistically poor in digenetic trema-
todes hosted by ancylids. Only seven host-
parasite combinations were observed, as
compared to 12 at BHF. Five of the seven
involved the single generation of Ferrissia
fragilis.

Differences in species composition of
those trematode faunas hosted by ancylids
can be explained on the basis of ecological
differences in the habitats, i.e. absence or
presence of suitable definitive or other
hosts.

Phytogeny of Southeastern Louisiana
Ancylids

Basch (1963) noted a general similarity
in radular patterns among Ferrissia, Lae-
vapex, and Hebetancylus. He also com-
pared anatomy of certain "soft-parts" and
noted a similar verge for Hebetancylus and
Ferrissia; however, the Hebetancylus
pseudobranch was bilobed like Laevapex.
Basch thus proposed that Hebetancylus
had greater affinity with Laevapex, which
he believed to be more advanced. Hebet-
ancylus and Ferrissia were suggested to
have evolved independently from a com-
mon ancestor that was in turn derived
from the family Planorbidae. However,
Turner (1978) reported that Hebetancyl-
us, like Ferrissia, formed a septum or hor-
izontal calcareous shelf partially closing
the shell aperture. This shell-like epi-
phragm, which was deposited by the pos-
terior margin of the mantle, has not been
reported for Laevapex. Because of this
observation we believe that Hebetancylus
and Ferrissia did not evolve independently
and that Hebetancylus occupies an inter-
mediate phylogenetic position between

Ferrissia and Laevapex.

Digenetic trematodes generally exhibit
greater host specificity for their molluscan
first intermediate host than for subse-
quent hosts, perhaps because of a longer
host-parasite evolutionary association
(Pearson, 1972). In light of this general-
ization, we compared the trematode fau-
nas hosted by each southeastern Louisi-
ana ancylid species. Similarities and dif-
ferences between respective faunas
should reflect the emended phylogeny we
have proposed.

Loevgpex
fuscus

Figure 25. Southeastern Louisiana ancylid species as
circles encompassing hosted cercariae and as over-
lapping circles encompassing "shared" and exclu-
sively hosted cercariae.

Tulane Studies in Zoology and Botany

Vol. 21

Figure 25 depicts each of the three spe-
cies as a circle encompassing hosted cer-
cariae, which for convenience have been
assigned arabic rather than conventional
roman numeral designations. Figure 25 al-
so demonstrates overlap or "sharing" of
cercarial species.

Only an echinostome (type VI), a longi-
furcocercous distome (type IV), and Cer-
caria type III, tentatively identified as
Posthodiplostomum minimum, were
"shared" by all three limpet species. None
were "shared" jointly by Hebetancylus
and Laevapex, or by Hebetancylus and
Ferrissia. Ferrissia and Laevapex jointly
"shared" xiphidiocercaria (type I), Mega-
lodiscus temperatus (type II), Lissorchis
sp. (type XI), and Stichorchis subtrique-
r/-w5 (type XVII).

Ferrissia seems to be the least special-
ized. Even though sympatric with the oth-
er genera, Ferrissia maintained the ability
to host exclusively four times as many cer-
cariae. This fact becomes more significant
when noted that F. fragilis comprised only
15% of all limpets examined. Relative
nearness of Ferrissia to the planorbid an-
cestral stock is suggested because at least
five cercariae, hosted exclusively by F.
fragilis or "shared" jointly with Laevapex
fuscus, are also hosted by planorbids.

The relative positions of Laevapex and
Hebetancylus are less clear. Laevapex and
Ferrissia have more cercariae in common
than Hebetancylus and Ferrissia; howev-
er, one cercaria (type XVII) is so broad in
host specificity as to develop in proso-
branch and both orders of pulmonate
gastropods. Furthermore, on the basis of
our study, and that of Smith (1967), L.
fuscus does not appear to be as suitable
host for Megalodiscus temperatus as does
F. fragilis.

Laevapex fuscus and Ferrissia fragilis
are widely distributed throughout the
U.S. east of the Rocky Mountains (Basch,
1963; Clarke, 1973). Thus, the explana-
tion for this "sharing" of cercariae by
southeastern Louisiana Laevapex and
Ferrissia may be found in the long sym-
patric association of these limpets in

North America. Wheieas, Hebetancylus
excentricus is Caribbean in distribution
with its historical, ecological associations
in Central and South America, H. excen-
tricus has been reported in North America
from southern Florida, coastal Georgia
(Basch, 1963), south central Texas (Pils-
bry, 1889; Walker, 1903), north central
Texas (McMahon and Aldridge, 1976),
southern Oklahoma (McMahon, et al,
1976), and southeastern Louisiana (Tur-
ner, 1978).

Finally, the fact that Laevapex hosts an
allocreadiid (type IX), all others of which
are hosted by sphaeriid clams, would tend
to separate it from other ancylids and sup-
ports its placement in a highly derived
phylogenetic position.

LITERATURE CITED

Basch, P.F. 1963. A review of the recent freshwater
limpet snails of North America (Mollusca: Pul-
monata). Bull. Mus. Comp. Zool., Harvard
Univ. 129:339-461.

Bedinger, C.A., Jr. andT.G. Meade. 1967. Biology
of a new cercaria for Posthodiplostomum mini-
mum (Trematoda: Diplostomidae). J. Parasit
53:985-988.

Bennett. H.J. and R. Allison. 1958. Observations on
the life cycle of the trematode Stichorchis sub-
triquetrus. Proc. La. Acad. Sci. 20:10-13.

Bennett, H.J. and A.G.Humes. 1939. Studies on the
precercarial development of Stichorchis sub-
triquetrus (Trematoda: Paramphistomidae). J
Parasit. 25:223-231.

Cagle, F. R. 1950. The life history of the slider turtle,
Pseudemys scripta troostii (Holbrook). Ecol.
Monogr. 20:31-54.

Clarke, A.H. 1973. The freshwater molluscs of the
Canadian Interior Basin. Malacologia. 13:1-509.

Cort, W.W. 1918. A new cercariaeum from North
America. J. Parasit. 5:86-91.

Duncan, B.L. and D.L. DeGiusti. 1976. Three new
lisorchiid cercariaea of the mutabile group from
Laevapex fuscus (Adams, 1841) and Ferrissia
rivularis (Say, 1917). Proc. Helminthol. Soc.
Wash. 43:1-9.

Goodchild, C.G. and D.E. Kirk. 1960. The life his-
tory of Spirorchis elegans Stunkard, 1923 (Tre-
matoda: Spirorchiidae) from the painted turtle.
J. Parasit. 46:219-229.

Holliman, R.B. andJ.E. Fisher. 1968. Life cycle and
pathology of Spirorchis scripta Stunkard, 1923
(Digenea: Sprirorchiidae) in Chrysemys picta
p/cra. J. Parasit. 54:310-318.

No. 2

Limpet Snails

Krull, W.H. and H.F. Price. 1932. Studies on the life
history of Diplodiscus temperatus Stafford, from
the frog. Occ. Pap. Mus. Zool. Univ. Michigan
237:1-37.

Lowery, G.H., Jr. 1974. The mammals of Louisiana
and its adjacent waters. Louisiana State Univer-
sity Press, Baton Rouge. 565 p.

Malek, E.A. and T.C. Cheng. 1974. Medical and
economic malacology. Academic Press, New
York. 398 p.

McMahon, R.F. 1976. Growth, reproduction and
life cycle in six Texan populations of two species
of fresh-water limpets. Amer. Midi. Nat. 95:174-
185.

and D. W. Aldridge. 1976. New distribu-
tion records for three species of freshwater limpet
(Pulmonata: Ancylidae) from North Central
Texas. Malac. Rev. 9:124-125.

_, andG.L. King. 1976. New

distribution records for two species of freshwater
limpet (Pulmonata: Basommatophora) in south-
ern Oklahoma. Southwestern Nat. 21:241-242.

Miller, J.H. 1954. Studies on the life history of Posr-
hodiplostomum minimum (MacCallum, 1921). J.
Parasit. 40:255-270.

Orloff, T.V. 1941. Investigation of the cycle of de-
velopment of the trematode Stichorchis subtri-
quetrus Rud. parasitic in beavers. Comptes Ren-
dus (Doklady) de I'Acad. des Sci. de I'U.R.S.S.
31:641-643.

Pearson, J.C. 1972. A phylogeny of life cycle pat-
terns of the Digenea. pp. 153-189. in: Advances
in parasitology. Vol. 10. Dawes, B. (Ed.) Aca-
demic Press, New York. 411 p.

Peters, L. and J.T. Self. 1963. An allocreadiid cer-
caria from limpets. (Abstr.) J. Parasit. 49
(Suppl.):41.

Pilsbry, H.A. 1889. Recent additions to the United
States snail fauna. The Nautilus. 3:62-64.

Schell, S.C. 1965. The life history of Haematoloe-
chus breviplexus Stafford, 1902 (Trematoda:
Haplometridae McMullen, 1937), with emphasis
on the development of the sporocysts. J. Parasit.
51:587-593.

Smith, R.J. 1959. Ancylid snails: first intermediate
host to certain trematodes with notes on ancylids
as a new host for Megalodiscus and Haemato-
loechus. Trans. Amer. Micro. Soc. 78:228-231.

1967. Ancylid snails as intermediate

hosts of Megalodiscus temperatus and other di-
genetic trematodes. J. Parasit. 53:287-291.

1968. Ancylid snails as first intermedi-

ate ho^is oi Lissorchismutabile comb. n. (Trema-
toda: Lissorchiidae). J. Parasit. 54:283-285.

Turner, H.M. 1978. Hebetancylus excentricus

(More let) (Pulmonata: Ancylidae) in Louisiana
and a report of septum formation. The Nautilus.
92:83-85.

and K.C. Corkum. 1977. New snail

host for Spirorchis scripta Stunkard, 1923 (Di-
genea: Spirorchiidae) with a note on seasonal
incidence. Proc. Helminthol. Soc. Wash. 44:225-
226.

Underwood, H.T. and N.O. Dronen. 1977. The
molluscan intermediate hosts for species oi Hae-
matoloechus Looss 1899 (Digenea: Plagiorchi-
idae) from raniid frogs of Texas. J. Parasit. 63:
112.

Walker, B. 1903. Notes on eastern American ancyli
The Nautilus. 17:13-19,25-30.

Wallace, H.E. 1941. Life history and embryology of
Triganodistomum mutabile (Cort) (Lissorchi-
idae: Trematoda). Trans. Amer. Micro. Soc
60:309-326.

Yamaguti, S. 1975. A synoptical review of life histor-
ies of digenetic trematodes of vertebrates. Kei-
gaku Publ. Co. , Ltd. , Tokyo. 1 100 p.

90 Tulane Studies in Zoology and Botany Vol. 21

ECOLOGICAL MORPHOLOGY OF FRESHWATER STREAM FISHES

A. JOHN GATZ, JR.

Department of Zoology, Duke University,
Durham, North Carolina 27706*

ABSTRACT

Variation and interrelations of 56 morphological
features were studied in 44 species of stream-living
fishes. The morphological characters are interpreted
ecologically based on information in the literature,
character correlations, factor analysis, gut content
analyses, and field observations.

The results show extensive significant linear corre-
lations among many of the characters. A number of
multiple associations were identified and ranked in
the factor analysis. The ecological meanings of these
latter results are that the major morphological trends
in these fishes are: (1) differentiation in feeding
strategy between "lie-in-wait" biting predators and
cruising suction feeders; (2) habitat separation ac-
cording to water velocity; (3) vertical habitat separa-
tion; and (4) feeding differentiation between small
insectivores and large omnivores.

Gut content data were also examined by factor
analysis. The results showed that for these fresh-
water stream fishes, size of prey is the most impor-
tant component in separating the diets of different
species and vertical position of the prey is the second
most important component. Various correlations be-
tween the size and location of the prey and several
morphological characteristics are also discussed.

How much about the biological roles of
a species is determinable from its ana-
tomical features and what particular
functions are regularly associated with
which features? These twin questions are
at the heart of much current work in the
areas of functional and ecological mor-
phology (Bock and von Wahlert, 1965;
Hespenheide, 1973). The purpose of the
research presented here is to provide

*Present Address: Department of Zoology, Ohio
Wesleyan University, Delaware, Ohio 43015

some answers to these questions for cer-
tain North Carolina freshwater stream
fishes.

The relationship between the form of
fishes and their life habits has been the
subject of casual writings from at least the
time of the ancient Hindus (Hora, 1935).
More recently, Hubbs (1941) provided a
good generalized background on this
topic, and subsequent works by Alexan-
der (1967), Aleev (1969), and Gosline
(1971) provide modem and comprehen-
sive summaries of various aspects.

In this paper I (1) summarize earlier
theoretical and experimental studies in
the literature relevant to probable
functional interpretation of various mor-
phological features of fishes and (2)
present the results of my own research
using correlation analysis, factor analysis,
and gut content analysis on these same
morphological features. Additional dis-
cussion of the structure of stream fish
communities based on this morphological
assessment of ecological roles is presented
elsewhere (Gatz, 1979).

MATERIALS AND METHODS

Fishes and Streams. S^inQS of various
lengths were used to collect fishes at six
stations in each of three streams in the
Piedmont of North Carolina. The streams
sampled were East Prong Little Yadkin
(Stokes County, Pee Dee drainage). Mud
Creek (Durham County, Cape Fear
drainage), and Maho Creek (Person
County, Roanoke drainage). Collections
were made at all seasons over a two year

EDITORIAL COMMITTEE FOR THIS PAPER:

DR. ALLEN KEAST, Professor of Biology, Queen's University, Kingston,

Ontario, Canada K7L 3N6
DR. THOMAS M. ZARET, Research Assistant Professor, Institute for
Environmental Studies, University of Washington, Seattle, Washington 98195

Tulane Studies in Zoology and Botany

Vol. 21

period (August, 1972, through August,
1974). All species thus collected were in-
cluded in the analysis of morphology.

Samples of ten adult individuals were
measured for each of the 33 species for
which I had collected sufficient material.
These species were Esox americanus, the
redfin pickerel, Esox niger, the chain
pickerel, Clinostomus funduloides, the
rosyside dace, Hybopsis hypsinotus, the
highback chub, Nocomis leptocephalus,
the bluehead chub, Notemigonus cryso-
leucas, the golden shiner, Notropis al-
borus, the whitemouth shiner, Notropis
altipinnis, the highfin shiner, Notropis an-
alostanus, the satinfin shiner, Notropis ar-
dens, the rosefin shiner, Notropis ceras-
inus, the crescent shiner, Notropis chili-
ticus, the redlip shiner, Notropis procne,
the swallowtail shiner, Phoxinus oreas,
the mountain redbelly dace, Semotilus
atromaculatus, the creek chub, Castosto-
mus commersoni, the white sucker, Eri-
myzon oblongus, the creek chubsucker,
Ictalurus nebulosus, the brown bullhead,
Noturus insignis, the margined madtom,
Aphredoderus sayanus, the pirate perch,
Gambusia affinis, the mosquitofish, Cen-
trarchus macropterus, the flier, Lepomis
auritus, the redbreast sunfish, Lepomis
cyanellus, the green sunfish, Lepomis gib-
bosus, the pumpkinseed, Lepomis gulo-
sus, the warmouth, Lepomis macrochirus,
the bluegill, Micropterus salmoides, the
largemouth bass, Pomoxis nigromacula-
tus, the black crappie, Etheostoma flabel-
lare, the fantail darter, Etheostoma fusi-
forme, the swamp darter, Etheostoma
olmstedi, the tessellated darter, and Perca
flavescens, the yellow perch. For the 11
rarer species, all individuals collected
were used. These 11 species were y4n^w/7/fl
rostrata, the American eel. Umbra pyg-
maea, the eastern mudminnow, Notropis
hudsonius, the spottail shiner, Moxosto-
ma anisurum, the silver redhorse, Moxo-
stoma ery thru rum, the golden redhorse,
Moxostoma pappillosum, the sucker-
mouth redhorse, Moxostoma robustum,
the smallfin redhorse, Ictahirus catus, the
white catfish, Ictahirus punctatus, the

channel catfish, Enneacanthus gloriosus,
the bluespotted sunfish, and Percina cras-
sa, the Piedmont darter. The conclusions
drawn concerning the interpretation of
the morphological features should not,
however, be generalized to groups of fish-
es with grossly different structure or hab-
itats.

Morphological Characteristics.

— Fifty-six morphological characteristics
were chosen for study. Determinations of
50 of these characters were made on each
individual fish. In addition, six characters
related to superficial brain morphology
were measured on a single individual of
each species. A preliminary study of the
brains of several individuals of a single
species (Notropis altipinnis) had indicated
very low variance in these characters.

Each of the 56 characters is listed below
along with the method of determination.
All length measurements less than 120
mm were made with vernier calipers to the
nearest 0.1 mm and length measurements
longer than this were made with a milli-
meter rule. All area measurements were
made from outline drawings using a plan-
imeter. For all qualitative characters, the
various manifestations or states of the char-
acteristic were coded numerically using in-
tegers. These character state codes for
qualitative characters are indicated in
parentheses below where relevant.

Superficial body and body shape charac-
ters

1. Standard length in this study is the
straight line distance from the most anter-
ior part of the head to the terminus of the
vertebral column.

2. Pigmentation pattern was visually
classified into one of four qualitative cate-
gories: (1) silvery or reflective sides; (2)
countershading with a dark lateral band;

(3) mottled pattern or vertical bars; and

(4) countershading without either silver-
iness or a lateral band.

3. Completeness of the lateral line
canal was recorded as being complete (2),
incomplete (1), or lacking (0).

No. 2

Ecological Morphology of Fishes

4. Position of the lateral line canal was
recorded as lacking (0), curving dorsally
(1), horizontal (2), or curving ventrally

(3).

5. Relative head length was measured

as the straight line distance from the most
anterior point on the upper lip to the pos-
terior margin of the opercular membrane
divided by the standard length. In large
adult Lepomis auritus, the redbreast sun-
fish, excessive hypertrophy of the opercu-
lar membrane occurs. For this species the
posterior limit for the head length meas-
urement was taken at a point where a sub-
jectively "reasonable" opercular mem-
brane might end.

6. Flatness index is the maximum
body depth divided by the maximum body
width.

7. Relative body depth is the maxi-
mum body depth divided by the standard
length.

8. Index of trunk shape is the perpen-
dicular distance from the anterior tip of
the head to an imaginary vertical line at
the point of maximum body depth divided
by the standard length.

9. Relative peduncle length is the hor-
izontal distance from a vertical line at the
level of the posterior margin of the base of
the most posterior median fin to the ter-
minus of the vertebral column divided by
the standard length.

10. Caudal peduncle flatness index is
the depth of the peduncle at its midpoint
divided by the width at the same point.

Caudal fin characters

11. Aspect ratio of the caudal fin may
be calculated as span squared divided by
area (Nursall, 1958; Ovchinnikov, 1971).
Span or maximum vertical distance and
area were measured from an outline
drawing of a maximally extended caudal
fin.

12. Caudal span / body depth ratio is

the span of the caudal fin divided by the
maximum body depth.

13. Number of caudal fin rays is the
total count of all rays reaching further
than one half the distance from the base of
the caudal fin to its distal edge. For fishes

in the families Cyprinidae, Catostomidae
and Centrarchidae, this count is the same
as the count for caudal rays given by
Hubbs and Lagler (1958), i.e., this count
is the number of branched rays plus two
(principal ray count). For fishes in other
families such as Esocidae and Ictaluridae,
some "rudimentary rays" (Hubbs and
Lagler, 1958) were also counted. The
number of "rudimentary rays" counted
was determined by the length criterion set
in this study. In the case of Anguilla, in
which the rays of the dorsal, anal and
caudal fins all form a continuous series,
this character was not used.

Paired fin characters

14. Pectoral fin length is the distance
from the base of the pectoral fin to the
extreme tip of the fin at its longest point
divided by the standard length of the fish.

15. Aspect ratio of the pectoral fin is
estimated as a length to width ratio.

16. Relative pectoral fin area is the sur-
face area of the pectoral fin divided by the
surface area of the fish. Outline drawings
of one pectoral fin and of the entire fish in
lateral view were made by tracing around
the objects. Relative pectoral fin area was
taken to be the ratio of the areas of these
two drawings.

17. Relative distance of the pectoral fin
from the center of gravity of the fish was
measured from the medial base of the pec-
toral fin to the point of center of gravity
(CG) on the lateral surface of the fish and
divided by standard length. This point CG
was determined by balancing the fish on
the tip of a dissecting needle. Notice that
this distance can be small both when the
fin is slightly anterior to the CG or poster-
ior to the CG and thus is a different char-
acter than #19 below.

18. Pectoral fin shape was coded based
on a subjective evaluation of whether the
fins were (1) rounded, (2) intermediate,
or (3) pointed.

19. Position of the pectoral fin relative
to the center of gravity is an assignment of
how the pectoral fin when adpressed
against the lateral surface of the body re-
lates to a transverse plane through the

94 Tulane Studies in Zoology and Botany Vol. 21

point CG as defined above. Four possible 31. Eye pigmentation was a categoriza-

relationships were recognized: (1) fin tion according to the presence or absence

wholly anterior to the plane; (2) fin orig- and orientation of any dark bands of pig-

inating anterior to CG and extending pos- mentation running across the cheek and

terior to the plane; (3) fin originating at through the eye. Possible categories and

the level of the CG plane; and (4) fin their character state codes were: (0) no

originating posterior to the CG. such pigment; (1) horizontal line through

20. Number of pectoral fin rays were the eye; (2) vertical line through the eye;
counted on alizarin stained fins. All rays and (3) both horizontal and vertical lines
(including spines), no matter what length, through the eye.

were counted. 32. Position of the mouth was coded

2 1 . Pelvic fin length was measured in a according to the position of the opening of
manner strictly analogous to pectoral fin the mouth when closed. Character states
length (#14, above). were (1) supraterminal if the lower jaw

22. Aspect ratio of the pelvic fin was extended anterior to the upper, (2) ter-
estimated as a length to width ratio. minal if the jaws were subequal, (3) sub-

23. Relative pelvic fin area was meas- terminal if the lower jaw ended slightly
ured analogously to relative pectoral fin posterior to the upper, (4) inferior if the
area (#16, above). opening was clearly recessed from the an-

24. Relative distance of the pelvic fin terior edge of the head, and (5) ventral if
from the center of gravity of the fish was the mouth was positioned along the ven-
measured analogously to the similar char- tral surface of the body.

acter, number 17, for the pectoral fin. 33. Orientation of the mouth was cod-

25. Pelvic fin shape was categorized ed according to the orientation of an imag-
similarly to pectoral fin shape. inary plane perpendicular to the longitud-

26. Position of the pelvic fin relative to inal axis of the body and tangential to both
the center of gravity was an assignment lips of the open mouth. Character states
using the same four character states as were: (1) dorsal if the plane faced above
were used in the analogous character for the vertical, (2) anterior if the plane was
the pectoral fin (#19, above). vertical, (3) oblique if the plane faced ob-

27. Number of pelvic fin rays was liquely downward, and (4) ventral if the
counted like the number of pectoral fin plane was horizontal.

rays. 34. Relative width of the mouth was

28. Position of the dorsal fin relative to the interior lateral dimension of the open-
the center of gravity was an assignment ing when the mouth was fully opened, div-
using the same four character states as ided by the standard length of the fish,
were used in the analogous character for 35. Relati\^ height of the mouth was
the pectoral fin (#19, above). Assignment the interior dorso-ventral dimension of
was made according to the position of the the opening when the mouth was fully
entire dorsal fin base relative to the CG opened, divided by the standard length of
plane. the fish.

Head characters 36 Index of protrusion was measured

29. Relative eye size was the diameter as the ratio of snout length with the mouth
of the eye between fleshy orbits along an open to snout length with the mouth
anterior-posterior axis divided by the closed where snout length is the distance
standard length. from the interior surface of the anterior

30. Position of the eyes involved as- edge of the bony orbit of the eye to the
signing character states depending upon anterior margin of the upper jaw at its
whether the eyes were placed laterally on midpoint.

the head (1), or were oriented dorso- 37. Numberof barbels was the count of

laterally either slightly (2) or greatly (3). these sensory structures about the mouth.

No. 2

Ecological Morphology of Fishes

38. Number of branchiostegal rays was
a count of these structures which support
the gill coverings. Alizarin staining was
used to facilitate counting.

39. Presence of jaw teeth was coded (0)
if none were present, (1) if teeth were on
the mandible and premaxillary and/or
maxillary only, and (2) if teeth were on
both jaws and additional bones of the roof
of the mouth such as the palatine or
vomer.

40. Shape of jaw teeth was coded (0) if
such teeth were absent, (1) if all teeth
were small sharp points, and (2) if both
small pointed and larger canine teeth were
present.

41. Hypertrophy of teeth on the phar-
yngeal arches was coded (0) if all teeth on
the fifth pharyngeal arch were small, and
( 1 ) if the teeth were hypertrophied.

42. Shape of the pharyngeal teeth was
coded (1) if all teeth were short pointed
structures, (2) if the teeth were hooked,
(3) if the teeth had cutting edges, and (4) if
the teeth had flattened grinding surfaces.

43. Number of gill rakers was a count of
the total number of rakers which were
visible after staining with alizarin on both
ascending and descending limbs of the la-
ternal surface of the first arch.

44. Shape of the middle gill raker was
measured as the length divided by the
width of that raker. In the case of picker-
els, genus Esox, in which the gill rakers
are represented by flat plates only, the
length was taken to be zero and hence the
value for this character was also zero.

45. Finer structure of the gill rakers was
coded (0) if each raker was smooth, (1) if
each raker had a ridged surface, (2) if each
raker had fewer than 50 fine sharp teeth
on it, and (3) if each raker had more than
50 fine teeth on it.

Internal body characters

46. Relative volume of the swim blad-
der was estimated by dividing the volume
of the swim bladder by the volume of the
fish. The latter volume was determined by
measuring displacement volume. Swim
bladder volume was measured in different
ways in different families of fishes. In fish-

es such as cyprinids and catostomids in
which the swim bladder could be removed
intact, it was and then filled with water
using a syringe and its displacement vol-
ume determined. In fishes which have
swim bladders with rather rigid walls and
definite median partitions such as cen-
trarchids and ictalurids, the lateral portion
of the left side of the bladder was removed
and the volume of water required to fill
the right half determined. Swim bladder
volume was estimated to be twice this vol-
ume. For fishes like esocids which have
thin walled, non-removeable swim blad-
ders, the volume was estimated mathe-
matically using the formula for the volume
of a cylinder. The estimate of radius used
was obtained from measurements of both
lateral and dorso-ventral diameters of the
swim bladder in a series of transverse sec-
tions of the entire fish.

47. Relative length of the swim bladder
was the ratio of the length of the swim
bladder to the standard length of the fish.

48. Relative gut length was measured
as the length of the entire alimentary canal
posterior to the pharynx divided by the
standard length. The digestive tract was
dissected from the fish and mesenteries
and was then stretched slightly to straight-
en bends when the measurement was tak-
en.

49. Number of pyloric caeca was a
count of the caeca at the junction of the
stomach and the intestine.

50. Percentage of red muscle in the
caudal peduncle was estimated as a per-
centage of total muscle in transverse sec-
tions made near the middle of the pedun-
cle. Sections were made using a freezing
microtome, stained for fats (which are
higher in concentration in red muscle than
in white), and mounted on slides. Plan-
imeter estimates of the areas of the red
and white muscle were made on drawings
of the slides traced from the image pro-
jected by a microprojector.

Brain characters

51. Relative size of the forebrain was
determined by measurement of the length
and the width of both forebrain lobes and

Tulane Studies in Zoology and Botany

Vol. 21

expressing the average of these two dim-
ensions as a proportion of the same two
measurements of the optic lobes. Brains
were exposed dorsally, and thus this char-
acter was an estimate of the amount of the
dorsal surface of the brain which was oc-
cupied by the forebrain relative to the
amount occupied by the optic lobes.

52. Relative size of the optic lobes was
determined by expressing the average of
the length and width of these lobes as a
proportion of the total length of the brain
from the anterior end of the forebrain to
the posterior end of the vagal lobes.

53. Relative size of the cerebellum is,
as are all further brain characters, analo-
gous to the character on the forebrain
(#51, above) in that the average of the
length and width of the lobe(s) under con-
sideration is expressed in relation to the
average of the length and width of the
optic lobes.

54. Relative size of the vagal lobes, as
indicated above, compares the dimen-
sions of the vagal and optic lobes.

55. Relative size of the facial lobes
compares the size of these lobes to the
optic lobes.

56. Relative size of the acoustic tub-
ercles compares the size of these lobes to
the optic lobes.

Gut Content Analysis. — As an aid to the
interpretation of the morphological char-
acteristics, gut contents were examined
throughout the entire length of the diges-
tive tract of all individuals that were stud-
ied morphologically. Replacements were
substituted for all fishes which had either
completely empty tracts or tracts with
wholly digested material. Each replace-
ment was selected to be of as nearly as
possible the same size and the same date
of collection as the original fish.

Items found in the gut were identified as
completely as possible. Grouped cate-
gories of food items were used in the final
analysis of foods taken. These categories
were: (1) fishes, (2) copepods, (3) ostra-
cods, (4) aquatic insects, (5) terrestrial in-
sects, (6) crayfishes, (7) isopods and am-

phipods, (8) diatoms, (9) filamentous al-
gae, (10) molluscs, (11) non-insect terres-
trial invertebrates, and (12) vascular
plants. Size of food items (greatest linear
dimension) was measured directly from
intact items and was estimated from the
dimensions of an identified part in the case
of broken or digested organisms. The
presence of sand and silt in the gut was
recorded also.

Gut content data were coded to facili-
tate numerical treatment. Following in the
tradition of fishery biology (Lagler, 1956),
two ways of considering prey were recog-
nized: number of items of a given cate-
gory, and percentage volume of all food
belonging to a given category. These two
types of information were incorporated
into the gut content characters by forming
two characters for each of the 12 previous-
ly listed categories of food items. Fre-
quency characters were given character
state codes according to the scheme: (1)
never taken, (2) present in up to 209f of
tracts , (3) 21-40%, (4) 41-60%, (5) 61-
80%, and (6) 81-100%. Percentage vol-
umetric importance character state values
were based on the percentage of digestive
tracts examined in which item(s) of the
given category formed a subjectively de-
termined largest percentage volume of
food of a single type, i.e., primary food.
Character state values for these characters
were: (1) not taken at all, (2) taken, but
never the primary food category, (3)
sometimes the primary food category, (4)
taken more often as the primary food than
items in any other category, but still not in
the majority of individuals of the species,
(5) primary food category in the majority
of individuals, and (6) the only food util-
ized. The importance character and fre-
quency character can be identical for any
given category. This was the case for cate-
gories 7, 10, and 12 above, and in these
cases I used only a single character repre-
senting both frequency and importance in
subsequent analyses.

The prey size data were coded as two
characters: absolute prey size and relative
prey size. For both, the coded value for

No. 2

Ecological Morphology of Fishes

each species was derived from an estimat-
ed overall mean size of an item of prey.
Character states for the absolute prey size
character were: (1) < 2 mm, (2) 5 mm, (3)
10 mm, (4) 20 mm, (5) 40 mm, and (6)
>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.

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

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

<
in

O m
z -I

<
X

u.
O

>
X

a
O

I- z

cc LU

uj u

Q
lU
CC

liJ
O

cc
o

2 A

<
ILI
CC
O

-- Efia

/
1 --Upyg

/
-■ Nins

/
Efus

-- Cmac
•- Eglo
0--Lgul

Eolm
Pfia
Pcra

Msal
Icat
Lcya

A say
Ineb
Lmac Pnig

Ipun
Laur Lgib
Gaff

-I

trt

(/)

<
CC

EobI Nard

Ccom Nalb

Pore Cfun

-■ Ncry

/
-)- Mery

/
+ Nana

Nhud Satr

Mani Nchi

\
+ Mpap Nlep

Npro
Nalt
Mrob

z
o

Ncer

Hhyp

LU
LU

z
<
cc

cc
liJ

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

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

< UI

0 <

-J

1 S

Z K
UJ O.
_l UJ

a
x'

Ui >■

Q Q

Z O

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U) UJ

« >

UJ -

z t-

I- <

< -•

_J UJ

U. (£

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

z
<

a.
O

5^
o '^ il

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.

o
a

<
—I
m

u.
O

3--
■■ Pcra
■ • Eolm
■ EfIa Efus

0--:-

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

Nalt
Pore
Nana
Eame

Lmac

Eglo Laur

Lgib
Pnig
Enig

oc
u

^ UJ

u. <

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

< cc

_i UJ
UJ ^

c <

z o

< ^
Z "

V) z

<
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

cc
o

>-
<

-I

z
o

UJ
0.

3
O
III

UJ
CD

3
O

-I

-J

4--

3 -h Icat

Ineb
Ipun

2--

cc
u

1 -

— 1

■ Cmac
Hhyp

Eglo
Pnig

Gaff
Lgib

Lmac

-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

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

U. Z

£ o
iE

3 LU

Z a.

<
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

<
cc

LLI
CL

< o

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

^£

U. Z

O LU

O 5

z o

< a.

I- LL

w
o

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

<£

tsl

DC
O

Z UJ

O a

Z
<

(A
(A

r z

t- I Q

< I- z

-I 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

cc
o

•■ Efus

< 1-

oc <

UJ -I

•• EobI

•• Upyg

• Pore

V) ^

w <n

UJ o

z o.

• Gaff

2 z

O UJ
o >

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

to
c

-a

u o

10 O (0 c

■p. 5 5 I

u
o

-p
c
(d

U
nS

3
o

s s

p< p<

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

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

East Prong

Mud Cr.

Maho Cr.

F value

Signif.

Lepomis aiiritus

0.397

0.444

9.806

<0.01

Lepomis cyanellus

0.379

0.395

5.804

<0.05

Pomoxis nigromaculatus

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.
Canada 28:1259-1268.

Ashmole, N.P. 1968. Body size, prey size, and ecol-
ogical segregation in five sympatric tropical terns
(Aves: Laridae). Syst. Zool. 17:292-304.

Bainbridge, R. 1960. Speed and stamina in three
fish. J. Exp. Biol. 37:129-153.

Barlow, G.W. 1972. The attitude offish eye-lines in
relation to body shape and to stripes and bars.
Copeia 1972:4-12.

Beamish, F.W.H. 1972. Ration size and digestion in
largemouth bass, Micropterus salmoides Lace-
pede. Can. J. Zool. 50:153-164.

Bhimachar, B.S. 1935. A study on the correlation
between the feeding habits and the structure of
the hind brain in the South Indian cyprinoid fish-
es. Proc. Roy. Soc. Lond., B 117:258-272.

No. 2

Ecological Morphology of Fishes

121

Bock, W.J., and G. von Wahlert. 1965. Adaptation
and the foim-function complex. Evolution 19:
269-299.

Boddoke. R. , E. J. Slijper and A. van der Stelt. 1959.
Histological characteristics of the body muscula-
ture of fishes in connexion with their mode of life.
Akademie van Wetenshappen Koninkl. Nederl.
Proceedings C 62:576-588.

Branson, B. A. , and G. A. Moore. 1962. The lateralis
component of the acoustico-lateralis system in
the sunfish family Centrarchidae. Copeia 1962:1-
108.

Breder, CM. Jr. 1926. The locomotion of fishes.
Zoologica (New York) 4:159-297.

Brett, J.R. 1965. The relation of size to rate of oxy-
gen consumption and sustained swimming speed
of sockeye salmon (Oncorhynchus nerka). J.
Fish. Res. Bd. Canada 22:1491-1501.

1967. Swimming performance of sock-
eye salmon (Oncorhynchus nerka) in relation to
fatigue time and temperature. J. Fish. Res. Bd.
Canada 24:1731-1741.

. 1973. Energy expenditure of sockeye

salmon, Oncorhynchus nerka, during sustained
performance. J. Fish. Res. Bd. Canada 30:1799-
1809.

Bridge, T.W., and A.C. Haddon. 1889. Contribu-
tions to the anatomy of fishes. I. The airbladder
and Weberian ossicles in the Siluridae. Proc.
Roy. Soc. London 46:309-328.

Carpenter, K.E. 1940. The feeding of salmon parr in
the Cheshire Dee. Proc. Zool. Soc. London
110:81-96.

Cole, C.F. 1967. A study of the eastern johnny dart-
er, Etheostoma olmsledi Storer (Teleostei, Per-
cidae).Ches.Sci. 8:28-51.

Cole, C.F. 1972. Status of the darters, Etheostoma
nigrum, E. longimanus and E. pondostomone in
Atlantic drainages (Teleostei, Percidae, subge-
nus Boleosoma), p. 119-138. In: The distribu-
tional history of the biota of the southern Appa-
lachians. Part IIL Vertebrates. P.C. Holt, R.A.
Paterson, and J. P. Hubbard (eds.) Virginia Poly-
tech Inst, and State Univ. Res. Div. Monograph
4, Blacksburg, Va.

Disler, N.N. 1971. Lateral line sense organs and
their importance in fish behavior. Israel Program
for Scientific Translations, Jerusalem. (Transl.
from Russian. ) TT 70-54021 .

Dobbin, C.N. 1941. A comparative study of the
gross anatomy of the air bladders of ten families
of fishs of New York and other eastern states. J.
Morphol. 68:1-29.

Evans, H.E. 1950. Structural modifications correlat-
ed with feeding habits in four species of cyprinid
fishes. Unpubl. Ph.D. Dissertation, Cornell
Univ., Ithaca.

1952. The correlation of brain pattern

and feeding habits in four species of cyprinid
fishes. J. Comp. Neur. 97:133-142.

Evans, H.M. 1940. Brian and body of fish. A study of
brain pattern in relation to hunting and feeding in
fish. The Blakiston Company, Philadelphia.

Forbes, S.A. 1880. The food of fishes. Bull. 111. State
Lab. Nat. Hist. l(3):19-70.

1888. On the food relations of fresh-
water fishes: a summary and discussion. Bull. 111.
State Lab. Nat. Hist. 2(8):475-538.

, and R.E. Richardson. 1920. The fishes

of Illinois. 2nd. ed. Natural History Survey of
Illinois. Vol. III. Ichthyology. State of Illinois.

Fry, F.E.J. 1958. Approaches to the measurement of
performance in fish, p. 93-97. In: The investiga-
tion of fish power problems. P. A. Larkin (ed.)
H.R. MacMillan Lectures in Fisheries, Univ. of
British Columbia.

Gatz, A.J. Jr. 1973. Speed, stamina, and muscles in
fishes. J. Fish. Res. Bd. Canada 30:325-328.

1975. The ecology of fishes in three

streams in North Carolina. Unpubl. Ph.D. Dis-
sertation, Duke University, Durham.

_. 1979. Community organization in fishes

as indicated by morphological features. Ecology
(in press).

Gee, J.H. 1968. Adjustment of buoyancy by long-
nose dace (Rhinichthys cataractae) in relation to
velocity of water. J. Fish. Res. Bd. Canada 25:
1485-1496.

1970. Adjustment of buoyancy in

blacknose dace, Rhinichthys atratulus. J. Fish.
Res. Bd. Canada 27:1855-1859.

_. 1972. Adaptive variation in swimblad-

der length and volume in dace, genus Rhinich-
thys. J. Fish. Res. Bd. Canada 29:119-127.

1974. Behavioral and developmental

plasticity of buoyancy in the longnose, Rhinich-
thys cataractae, and blacknose, R. atratulus,
(Cyprinidae) dace. J. Fish. Res. Bd. Canada 31:
35-41.
, K. Machniak and S.M. Chalanchuk.

1974. Adjustment of buoyancy and excess inter-
nal pressure of swimbladder gases in some North
American freshwater fishes. J. Fish. Res. Bd.
Canada31:1139-1141.

Gibbons, J.R.H., and J.H. Gee. 1972. Ecological
segregation between longnose and blacknose
dace (genus Rhinichthys) in the Mink River,
Manitoba. J. Fish. Res. Bd. Canada 29:1245-
1252.

Gilbert, C.R. and R.M. Bailey. 1972. Systematics
and zoogeography of the American cyprinid fish
Notropis (Opsopoeodus) emiliae. Occ Pap. Mus.
Zool., Univ. Michigan No. 664:1-35.

Gordon, M.S. 1968. Oxygen consumption of red and
white muscles from tuna fishes. Science 159:87-
90.

Gorman, O.T., and J.R. Karr. 1978. Habitat struc-
ture and stream fish communities. Ecology 59:
507-515.

Gosline, W.A. 1971. Functional morphology and
classification of teleostean fishs. Univ. Press of
Hawaii, Honolulu.

1973. Considerations regarding the

phylogeny of cypriniform fishes, with special ref-
erence to structures associated with feeding.
Copeia 1973:761-776.

122

Tulane Studies in Zoology and Botany

Vol. 21

Gray, J. 1968. Animal locomotion. Weidenfeld and
Nicolson, London.

Griffith, J.S. Jr. 1974. Utilization of invertebrate
drift by brook trout (Salvelinus fontinalis) and
cutthroat trout (Salmo clarki) in small streams in
Idaho. Trans. Amer. Fish. Soc. 103:440-447.

Gromov, I. A. 1973. Morphological characteristics
and local populations of the eastern carp [Cypri-
nus carpio haematopterus (Temm. et Schlegel)].
J. Ichthyology 13:335-347.

Hall, D.J., W.E. Cooper and E.E. Werner. 1970.
An experimental approach to the production
dynamics and structure of freshwater animal
communities. Limnol. Oceanogr. 15:839-928.

Hammond, B.R., and C.P. Hickman, Jr. 1966. The
effect of physical conditioning on the metabolism
of lactate, phosphate, and glucose in rainbow
trout, Salmo gairdneri. J. Fish. Res. Bd. Canada
23:65-83.

Harman, H.H. 1967. Modem factor analysis. 2nd.
ed. Univ. Chicago Press, Chicago.

Harris, J.E. 1937. The mechanical significance of the
position and movements of the paired fins in the
Teleostei. Pap. TortugasLab. 31:171-189.

1938. The role of fins in the equilibrium

of the swimming fish: 2. The role of the pelvic
fins. J. Exp. Biol. 15:32-47.

-. 1953. Fin patterns and mode of life of

fishes, p. 17-28. In: Essays in marine biology.
S.M. Marshall and A. P. Orr (eds.) Oliver and
Boyd, Edinburgh.

Herrick, C.J. 1924. Neurological foundations of an-
imal behaviour. H. Holt and Company, New
York.

Hespenheide, H.A. 1973. Ecological inferences
from morphological data. Ann. Rev. Ecol. Syst
4:213-229.

Himberg, K.-J.M. 1970. A systematic and zooge-
ographic study of some North European corego-
nids, p. 219-250. In: Biology of coregonid fishes.
C.C. Lindsey and C.S. Woods (eds. ) Univ. Mani-
toba Press, Winnipeg.

Hochachka, P.W. 1961. The effect of physical train-
ing on oxygen debt and glycogen reserves in
trout. Can. J. Zool. 39:767-776.

Hora, S.L. 1922. Structural modifications in the fish
of mountain torrents. Rec. Indian Mus., Calcutta
24:31-61.

1935. Ancient Hindu conception of cor-
relation between form and locomotion of fishes.
J. Asiatic Soc. Bengal. Science 1:1-7.

Horak, D.L. 1969. The effect of fin removal on
stamina of hatchery-reared rainbow trout. Progr.
Fish-Cuh. 31:217-220.

. 1972. Survival of hatchery-reared rain-
bow trout {Salmo gairdneri) in relation to stam-
ina tunnel ratings. J. Fish. Res. Bd. Canada 29:
1005-1009.

Houde, E.D. 1%9. Sustained swimming ability of
larvae of walleye {Stizostedion vitreum vitreum)
and yellow perch (Perca flavescens) . J. Fish. Res.
Bd. Canada 26:1646-1659.

Hubbs, C.L. 1941. The relation of hydrological con-
ditions to speciation in fishes, p. 182-195. In: A
symposium on hydrobiology. Univ. of Wisconsin
Press, Madison.

, and K.F. Lagler. 1958. Fishes of the

Great Lakes region. Rev. ed. Cranbrook Insti-
tute of Science Bull. No. 26.

Hynes, H.B.N. 1970. The ecology of running waters.
Univ. Toronto Press, Toronto.

Ivlev, V.S. 1%1. Experimental ecology of the feed-
ing of fishes. Yale Univ. Press, New Haven.
(Transl. from the Russian by D. Scott.)

Jones, A.N. 1975. A preliminary study of fish segre-
gation in salmon spawning streams. J. Fish. Biol.
7:95-104.

Jones, D.R. 1971. The effect of hypoxia and anaemia
on the swimming performance of rainbow trout
{Salmo gairdneri). J. Exp. Biol. 55:541-551.

Jones, F.R.H. 1951. The swimbladder and the ver-
tical movements of teleostean fishes. 1. Physical
factors. J. Exp. Biol. 28:553-566.

Kanep, S.V. 1971. Some aspects of the functional
morphology of the lake pelyad [Coregonus peled
(Gmelin)]. J. Ichthyology 11:855-866.

Keast, A. 1966. Trophic interrelationships in the fish
fauna of a small stream. Proc. 8th Conf. Great
LakesRes. Div., Univ. Michigan Publ. 15:51-79.

, and D. Webb. 1966. Mouth and body

form relative to feeding ecology in the fish fauna
of a small lake. Lake Opinicon, Ontario. J. Fish.
Res. Bd. Canada 23:1845-1874.

Keenleyside, M.H.A. 1%2. Skin-diving observa-
tions of Atlantic salmon and brook trout in the
Miramichi River, New Brunswick. J. Fish. Res.
Bd. Canada 19:624-634.

Kliewer, E.V. 1970. Gillraker variation and diet in
lake whitefish Coregonus clupeaformis in north-
em Manitoba, p. 147-165. In: Biology of core-
gonid fishes. C.C. Lindsey and C.S. Woods
(eds.) Univ. Manitoba Press, Winnipeg.

Kramer, E. 1960. Zur Form und Funktion des Lo-
komotionsapparates der Fische. Z. Wiss. Zool.
163:1-36.

Lazier, K.F. 1956. Freshwater fishery biology. 2nd.
ed. Wm. C. Brown Company, Dubuque, Iowa.

, J.E. Bardach and R.R. Miller. 1%2.

Ichthyology. John Wiley and Sons, Inc., New
York.

Larimore, R.W., and M.J. Duever. 1968. Effects of
temperature acclimation on the swimming ability
of smallmouth bass fry. Trans. Amer. Fish. Soc.
97:175-184.

Laurence, G.C. 1972. Comparative swimming abili-
ties of fed and starved larval largemouth bass
{Micropterus salmoides). J. Fish. Biol. 4:73-78.

Lin, Y., G.H. Dobbs, III and A.L. Devries. 1974.
Oxygen consumption and lipid content in red and
white muscles of Antarctic fishes. J. Exp. Zool.
189:379-385.

Lindsey, C.C. 1%3. Sympatric occurrence of two
species of humpback whitefish in Squanga Lake,
Yukon Territory. J. Fish. Res. Bd. Canada 20:
749-767.

No. 2

Ecological Morphology of Fishes

123

Lindstrom, T. 1955. On the relation fish size - food
size. Fish. Bd. Sweden. Freshwater Res. Rep.
No. 36:133-147.

Love, R.M. 1970. The chemical biology of fishes.
Academic Press, New York.

Lundberg, J.G., and E. Marsh. 1976. Evolution and
functional anatomy of the pectoral fin rays in
cyprinoid fishes, with emphasis on the suckers
(Family Catostomidae). Amer. Midi. Nat. 96:
332-349.

Machniak, K., and J.H. Gee. 1975. Adjustment of
buoyancy by tadpole madtom, Noturus gyrinus,
and black bullhead, Ictalurus melas, in response
to a change in water velocity. J. Fish. Res. Bd.
Canada 32:303-307.

Marshall, N.B. 1971. Explorations in the life of fish-
es. Harvard Univ. Press, Cambridge, Mass.

Martin, W.R. 1949. The mechanics of environmen-
tal control of body form in fishes. Publ. Ontario
Fish. Res. Lab. No. 70.

Miller, R.J., and H.E. Evans. 1965. External mor-
phology of the brain and lips of catostomid fishes.
Copeia 1965:467-487.

Nelson, E.M. 1961. The comparative morphology of
the definitive swim bladder in the Catostomidae.
Amer. Midi. Nat. 65:101-110.

Nichols, J.T. 1915. On one or two common structur-
al adaptations in fishes. Copeia 1915:19-21.

Nikolskii. G.V. 1933. On the influence of the rate of
flow on the fish fauna of the rivers of central Asia.
J. Anim. Ecol. 2:266-281.

1963. The ecology of fishes. Academic

Press, New York. (Transl. from Russian by L.
Birkett.)

Nilsson, N.-A. 1955. Studies on the feeding habits of
trout and char in north Swedish lakes. Fish. Bd.
Sweden. Freshwater Res. Rep. Drottningholm
No. 36:163-225.

Nilsson, N.-A. 1958. On the food competition be-
tween two species of Coregonus in a north-
Swedish lake. Fish. Bd. Sweden. Freshwater
Res. Rep. Drottningholm No. 39:146-161.

1960. Seasonal fluctuations in the food

segregation of trout, char and whitefish in four-
teen north-Swedish lakes. Fish. Bd. Sweden.
Freshwater Res. Rep. Drottningholm No. 41:
185-205.
• 1963. Interaction between trout and

char in Scandinavia. Trans. Amer. Fish. Soc.
92:276-285.

1965. Food segregation between sal-

monid species in north Sweden. Fish. Bd. Swe-
den. Freshwater Res. Rep. Drottningholm No.
46:58-78.

1967. Interactive segregation between

fish species, p. 295-313. In: The biological basis
of freshwater fish production. S.D. Gerking(ed.)
John Wiley and Sons, Inc. , New York.
Northcote, T.G. 1954. Observations on the compar-
ative ecology of two species of fish, Cottus asper
and Cottiis rhotheus, in British Columbia. Copeia
1954:25-28.

Nursall, J.R. 1958. The caudal fin as a hydrofoil.
Evolution 12:116-120.

Oseid, D., and L.L. Smith, Jr. 1972. Swimming en-
durance and resistance to copper and malathion
of bluegills treated by long-term exposure to sub-
lethal levels of hydrogen sulfide. Trans. Amer.
Fish. Soc. 101:620-625.

Ovchinnikov, V.V. 1971. Swordfishes and billfishes
in the Atlantic Ocean. Ecology and functional
morphology. Israel Program for Scientific Trans-
lations, Jerusalem. (Transl. from Russian.) TT
71-50011.

Pianka, E.R. 1969. Sympatry of desert lizards (Cte-
notus) in western Australia. Ecology 50:1012-
1030.

Phillips, A.M. Jr. 1969. Nutrition, digestion, and
energy utilization, p. 391-432. In: Fish physiol-
ogy. Vol. I. W.S. Hoar and D.J. Randall (eds.)
Academic Press, New York.

Protasov, V.R. 1970. Vision and near orientation of
fish. Israel Program for Scientific Translations,
Jerusalem. (Transl. from Russian. )TT 70-50065.

Reddy, S.R., and T.J. Pandian. 1974. Effect of run-
ning water on the predatory efficiency of the
larvivorous fish Gambusia affinis. Oecologia
(Berl.) 16:253-256.

Roberts, J.L., and J.B. Graham. 1974. Swimming
and body temperature of mackerel. Amer. Zool.
14:1258 (Abstr.)

Roberts, N.J., and H.E. Winn. 1962. Utilization of
the senses in feeding behavior of the johnny dart-
er, Etheostoma nigrum. Copeia 1962:567-570.

Schmitz, E.H., and CD. Baker. 1969. Digestive
anatomy of the gizzard shad, Dorosorna cepedi-
anum and the threadfin shad, D. petenense.
Trans. Amer. Microsc. Soc. 88:525-546.

Schnitzlein, H.N. 1964. Correlation of habitat and
structure in the fish brain. Amer. Zool. 4:21-32.

Schoener, T. W. 1968. The Anolis lizards of Bimini:
resource partitioning in a complex fauna. Ecol-
ogy 49:704-726.

, and G.C. Gorman. 1968. Some niche

differences among three species of Lesser Antil-
lean anoles. Ecology 49:819-830.

Schutz, D.C., and T.G. Northcote. 1972. An exper-
imental study of feeding behavior and interaction
of coastal cutthroat trout (Salmo clarki clarki)
and Dolly Varden (Salvelinus malma). J. Fish.
Res. Bd. Canada 29:555-565.

Schwartz, F.J., and B.W. Dutcher. 1962. Tooth re-
placement and food of the cyprinid, Notropis
cerasinus, from the Roanoke River, Virginia.
Amer. Midi. Nat. 68:369-375.

Schwassmann, H.O., andL. Kruger. 1968. Anatomy
of visual centers in teleosts, p. 3-16. In: The cen-
tral nervous system and fish behavior. E. Ingle
(ed.) Univ. Chicago Press, Chicago.

Sokal, R.R., and F.J. Rohlf. 1969. Biometry. W.H.
Freeman and Company, San Francisco.

Starck, W.A. II, and R.E. Schroeder. 1970. Investi-
gations on the gray snapper, Lutjanus griseus.
Stud. Trop. Oceanogr. Miami 10.

Stevens, E.D. 1968. The effect of exercise on the
distribution of blood to various organs in rainbow

124

Tulane Studies in Zoology and Botany

Vol.21

trout. Comp. Biochem. Physiol. 25:615-625.

Swynnerton, G.H., and E.B. Worthington. 1940.
Note on the food offish in Haweswater (Westmor-
land). J. Anim. Ecol. 9:183-187.

Thomas, J.D. 1962. 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

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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

Rana pipiens

Pseudacris triseriata

Hyla versicolor

Bufo

Tadpoles

Rodentia

Oligochaeta

Hirudinea

Gastropoda

No. 2

Niche Relationships of Thamnophis

133

H
CO

¥mu

+-E^

i crys^

4 — ^

C^im^ -t 'il

\'.}y\'*'\

—

1 » Il

1 — 1 — 1|

12 13

1 i, :Ak -. 1

1 •. "J 1

1 — -, — 1 1

1 — ? — 1 1

o
O

1 I ■ <"Ti 1

|li^(.".")

r • 1 1

t — ?— M

21 22 23 24 25 2*6

(' - l*''j- rl

27 28

2'9

30 31

I .'-I* . • ■ .1

t-h-d —

PL.
Eh

^::'.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—

/^/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

().

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

li^'' v»:'>:'l|

20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35

34-

o
O

-J
<

<
o

■^

f^^^vH^

1 : h

' — ' 1

19 20 21

24 25 26 27 28 29

32 33

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

6.941

-1.620

0.259

0.222

-0.485

-0.525

0.626*

0.358*

0.448*

0.513*

0.165*

0.164*

-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
\

1 i T sirtalis

67 8 910 11 12 1234
AM TIME OF DAY

SYMPATRIC

5 6

8 9
PM

D
O
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

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