The Project Gutenberg EBook of Geographic Variation in the North American
Cyprinid Fish, Hybopsis gracilis, by Leonard J. Olund and Frank B. Cross

This eBook is for the use of anyone anywhere at no cost and with
almost no restrictions whatsoever.  You may copy it, give it away or
re-use it under the terms of the Project Gutenberg License included
with this eBook or online at www.gutenberg.org


Title: Geographic Variation in the North American Cyprinid Fish, Hybopsis gracilis

Author: Leonard J. Olund
        Frank B. Cross

Release Date: December 27, 2011 [EBook #38425]

Language: English

Character set encoding: ISO-8859-1

*** START OF THIS PROJECT GUTENBERG EBOOK GEOGRAPHIC VARIATION ***




Produced by Chris Curnow, Joseph Cooper, Erica
Pfister-Altschul and the Online Distributed Proofreading
Team at http://www.pgdp.net

University of Kansas Publications
Museum of Natural History

Volume 13, No. 7, pp. 323-348, pls. 21-24, 2 figs.
February 10, 1961

Geographic Variation
In the North American Cyprinid Fish,
Hybopsis gracilis

BY
LEONARD J. OLUND AND FRANK B. CROSS

University of Kansas
Lawrence
1961

University of Kansas Publications, Museum of Natural History

Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Robert W. Wilson

Volume 13, No. 7, pp. 323-348, pls. 21-24, 2 figs.
Published February 10, 1961

University of Kansas
Lawrence, Kansas

PRINTED IN
THE STATE PRINTING PLANT
TOPEKA, KANSAS
1961

28-5871

Geographic Variation In the North American Cyprinid Fish, Hybopsis gracilis

BY
LEONARD J. OLUND AND FRANK B. CROSS

CONTENTS

•   PAGE
• Introduction 325
• Methods, Materials, and Acknowledgments 326
• Description of the Species Hybopsis gracilis 327
• Hybopsis gracilis gracilis 328
• Hybopsis gracilis gulonella 330
• Intraspecific Variation 333
• Natural History 334
• Habitat 334
• Associated Species 336
• Food 338
• Spawning Season 338
• Discussion 340
• Literature Cited 343

INTRODUCTION

The flathead chub, Hybopsis gracilis (Richardson), occurs in the Plains Region of Canada and the United States, in four major drainage systems: Mackenzie River, which discharges into the Arctic Ocean; Saskatchewan River, which discharges into Hudson Bay via Nelson River; and Missouri-Mississippi System and Rio Grande, both draining into the Gulf of Mexico. Each of these systems is occupied in part only. In the Mackenzie Basin, H. gracilis has been reported as far north as Fort Good Hope (Walters, 1955:347). Flathead chubs occur in the Saskatchewan Basin from Alberta eastward to Lake Winnipeg, Manitoba, but have not been found in other streams that flow into Lake Winnipeg (Red River, Brokenhead River and Whitemouth River) nor in Nelson River downstream from Lake Winnipeg. In the Missouri Basin the species occurs more or less continuously from the high plains adjacent to the Rocky Mountains in Montana and Wyoming down the mainstream of the Missouri River to its mouth, and down the mainstream of the Mississippi River as far as Barfield, Arkansas, but not to the Gulf. The species prob[pg 326]ably attains its greatest abundance in the Missouri Basin, but it is scarce or absent in tributaries north and east of the Missouri mainstream, in the South Platte Basin, and in the central part of the Platte River in Nebraska. The flathead chub is unknown in the Mississippi Basin above the mouth of the Missouri River, and in the Ohio River Basin above its mouth. In the Arkansas River Basin, records are restricted to (1) the headwaters and tributaries of the Arkansas River from eastern Colorado downstream as far as Garden City, Kansas, (2) the Cimarron River at Kenton, Cimarron County, Oklahoma, and (3) the South Canadian River and tributaries from northeastern New Mexico eastward as far as Norman, McClain County, Oklahoma, but rarely there. Thus, the range in the Arkansas Basin seems to consist of three isolated segments. Likewise, isolated populations exist in the Rio Grande System, where flathead chubs are confined to the upper parts of the Rio Grande and Pecos basins, above the confluence of the Rio Grande and Pecos Rivers. Records resulting from introductions have been reported for the Gila River by Koster (1957:62) and from the Snake River, Wyoming, by Simon (1946:72).

Six names apply to the flathead chub, the earliest of which is Cyprinus gracilis Richardson (1836:120). Other names have sometimes been accepted as applicable to valid species and/or subspecies, but usage, diagnoses, and stated ranges have been confusingly inconsistent. For most of the past 100 years, Platygobio Gill has been recognized as the appropriate generic name for the flathead chub, but Bailey (1951:192) places Platygobio and other nominal genera of barbeled minnows having short guts, protractile premaxillae, and four teeth (primary row) in the single genus Hybopsis (Agassiz, 1854). Strangely, the orthotype of Hybopsis, H. gracilis Agassiz, is a junior synonym of H. amblops (Rafinesque) (Hubbs and Ortenburger, 1929b:66) and is a younger name than C. gracilis Richardson.

The purpose of this paper is to redescribe the species and to make known its pattern of geographic variation. Natural history will also be considered, as will habitat, food habits, and breeding season.

METHODS, MATERIALS AND ACKNOWLEDGMENTS

DESCRIPTION OF THE SPECIES

Hybopsis gracilis (Richardson)

Flathead Chub

(Synonymy under accounts of subspecies)

Hybopsis gracilis gracilis (Richardson)

(Plate 22)

Hybopsis gracilis gulonella (Cope)

(Plate 23)

INTRASPECIFIC VARIATION

Two subspecies of H. gracilis are recognized by us: one northern and eastern, characteristically inhabiting large rivers (H. g. gracilis), and one southern and western, characteristically inhabiting small streams (H. g. gulonella). Other scientific names that have been applied to this fish in the past are listed in the synonymy.

H. g. gulonella is a chubby, deep-bodied fish, whereas H. g. gracilis is long and slender. The head of the creek subspecies is deeper and longer than that of H. g. gracilis, being rounded anteriorly when seen in sideview. The head of the large-river subspecies is acutely wedge-shaped in profile. H. g. gracilis has a larger orbit than H. g. gulonella. Fins of H. g. gracilis are more strongly falcate than those of the other subspecies. H. g. gracilis has a greater number of lateral line scales, pectoral rays and post-Weberian vertebrae than the creek subspecies. The large-river subspecies attains much larger size than does the creek subspecies (Plate 24). Except in areas of intergradation, complete separation of the two subspecies can be made on the basis of lateral line scales, pectoral rays, post-Weberian vertebrae and head-depth. The regressions of head-depth on standard length in H. g. gracilis from the Saskatchewan River (several localities) and in H. g. gulonella from Beaver Creek, Arkansas River Drainage (KU 4769) are shown in Plate 24. Although values for the largest specimens of H. g. gracilis are omitted from Plate 24, the regression remains essentially linear to standard lengths of approximately 250 mm. On the basis of head-depth alone, separation of the two subspecies is possible in specimens larger than 40 mm. Similar results were obtained by using the regression of postorbital length on standard length, and could have been obtained by using other proportional measurements.

NATURAL HISTORY

Habitat

The species inhabits alkaline streams with shifting sand bottoms where the waterlevel fluctuates considerably with heavy rains and melting snow. The flathead chub is found in silty water and often is the predominant species in streams that have high turbidity. The remarkable ability of this fish to withstand exceedingly high turbidity is illustrated by its predominance in the Little Missouri River, which has an average concentration of suspended silt two and one-half times that of the Missouri River at Kansas City (Personius and Eddy, 1955:42).

H. g. gracilis is found in large rivers throughout its range, occa[pg 335]sionally migrating into smaller streams, especially in the spawning season. It prefers the main channel of rivers in moderate to strong current. All series examined are from elevations lower than 3,000 feet.

H. g. gulonella occupies small rivers and creeks, preferring pools with moderate currents. In fall, dense concentrations of this subspecies have been found in small pools, where brush, driftwood or [pg 336] other debris deflects the current and prevents filling with drifting sand. Hundreds of flathead chubs were collected in such pools in the Purgatoire and Arkansas rivers. Specimens were also collected with ease in Beaver Creek, Colorado, from pools with murky water and slight flow, over bottoms of gravel and bedrock. No brush or other debris was near the pools. In each case the streams carried little water, although they undoubtedly carry greater volumes of water in spring and early summer after rains and spring thaws. The preferred bottom-type for this subspecies seems to be gently shifting sand.

H. g. gulonella is found in warm-water streams, whereas H. g. gracilis occurs in cooler water. The southwestern subspecies was taken in August in the Mora River drainage at Sapello (temperatures above 80° F.) but not at Mora (temperatures below 70° F.). In the Purgatoire River, a thriving population was found where the water temperature was 92° F., on September 6, 1959. In the Arkansas and Pecos rivers and the Rio Grande this subspecies is most abundant below the mountainous parts of the stream-courses, but at elevations higher than 4,000 feet on the plains.

Associated Species

In the Pecos and Arkansas basins, species commonly taken with H. g. gulonella are Catostomus commersonnii, Hybognathus placita, [pg 337] Notropis lutrensis lutrensis, Notropis stramineus missuriensis, Pimophales promelas, and Campostoma anomalum plumbeum. The only spiny-rayed fishes that we have found with H. g. gulonella are Lepomis cyanellus and L. humilis, both of which are scarce. Associates of H. g. gracilis include the same species, plus other ostariophysan fishes such as species of Carpiodes, Ictiobus, and silt-adapted species of Hybopsis and Notropis.

We failed to find the flathead chub at any of 11 localities in the South Platte drainage, where we collected in September, 1959. Dr. George Baxter, of the Department of Zoology, University of Wyoming, told us that he has never found H. gracilis in that drainage. The fauna of the South Platte includes Catostomus catostomus, Semotilus atromaculatus, Hybopsis biguttata, Hybognathus hankinsoni, Notropis cornutus frontalis, Etheostoma nigrum and E. exile--species rarely if ever found with H. gracilis.

Ecologically, H. g. gulonella seems to be the counterpart of Semotilus atromaculatus in streams where the latter species is absent. Observations of H. g. gulonella in the Purgatoire River indicated [pg 338] that loosely-organized groups of flathead chubs congregated one to four inches above the bottom of pools, and near or under protective cover such as roots of vegetation or debris lodged against shore. Individuals moved about independently within the group (rather than as schools), and occasionally rose to the surface, perhaps for food.

Food

The flathead chub is chiefly carnivorous, but its food includes some aquatic vegetation (Table 1). Most organisms found in specimens (both subspecies) were terrestrial insects (Coleoptera, Diptera, Orthoptera); all insects were adult stages, except those designated as larvae in Table 1. Roundworms probably were parasites, rather than food.

Hubbs (1927:76) states that the food of young flathead chubs that were obtained from the Arkansas River System in New Mexico consisted "almost entirely of crustaceans (small ostracods and cladocerans to the exclusion of all else but an occasional larval or adult insect, etc.)."

Spawning Season

Specimens of H. g. gulonella that have been examined reach sexual maturity at approximately 65 mm standard length. Most specimens of H. g. gracilis less than 85 mm in standard length are immature, but larger specimens probably are mature.

The spawning season is in late summer, beginning in July and extending into September. Specimens from the Peace River, collected on August 10, 1952, include females that were mostly spent and tuberculate males. Males and females in spawning condition were collected in the Milk River in August of 1955. A large prespawning female was obtained in Red Deer River in June of 1952. A male from Fort McMurray had fairly well developed tubercles on August 9, 1955. A prespawning female was taken from the Saskatchewan River at Clarkboro Ferry on June 7, 1957. Tuberculate males were collected in the Powder River on June 30, 1957. Specimens from the White River in South Dakota, collected on July 7, 1934, include tuberculate males. The specimens discussed above are H. g. gracilis or intergrades tending toward that subspecies.

Specimens of H. g. gulonella collected in the Arkansas River at Pueblo and Florence, Colorado, on September 7, 1959, include some tuberculate males, although most females are spent. On August 8, 1957, a series of flathead chubs that includes tuberculate males was collected in the Redwater River, Montana. In the Pecos River on [pg 340] August 25, 1958, spawning seemingly had been completed, although a few males still bore tubercles.

[pg 339] Table 1. Organisms Found in Stomachs of Hybopsis gracilis From Various Locations, Expressed as Percentage of Total Volume.

Spawning apparently occurs when river levels recede to the seasonal lows. In late summer, temperatures of these rivers probably are maximal, their turbidities are reduced, and their sandy bottoms are stable. Underhill (1959) reports that this species is rare in the Vermillion River, a northeastern tributary of the Missouri River, except in autumn when large numbers occur near the mouth of the river. We suspect that this is associated with spawning.

PLATE 21

PLATE 22

PLATE 23

PLATE 24

DISCUSSION

Hybopsis gracilis is highly variable in several morphological characteristics, including size and shape of head, body, and fins, and number of scales, vertebrae, and fin-rays. The variations are correlated in a way that indicates the existence of two subspecies. One of these, H. g. gracilis, attains large size, and has 1) a slender, streamlined body, 2) a depressed head that is acutely wedge-shaped in profile, 3) strongly falcate fins with the dorsal and pelvic fins originating anteriorly, and 4) many scales, vertebrae, and pectoral fin-rays. The second subspecies, for which H. g. gulonella is the oldest applicable name, is small, and has 1) a deep, chubby body, 2) head convex in dorsal contour (less depressed than in H. g. gracilis), 3) fins less falcate than in the latter subspecies, with the dorsal and pelvic fins originating more posteriorly, and 4) fewer [pg 341] scales, vertebrae, and pectoral fin-rays than H. g. gracilis. These differences are consistently expressed throughout the size-ranges of the subspecies, and in series collected at the same or nearby localities in several different years. Considerable variability was found in features other than those mentioned above, but individual variation among specimens from the same locality and adjacent localities is so great that none is diagnostic of subspecies. For example, orbital size and length of fins (but not their falcate shape) are variables that have little diagnostic value, although both features seem to vary in clinal fashion, with the higher values in the north.

Variation in H. gracilis, as shown in the graphic analysis (Figs. 1 and 2) and distribution map (Plate 21), presents two clines: a north-south cline and a large-river to small-river (mainly east-west) cline. The absence of H. gracilis from certain portions of river systems is a matter of concern. The species has not been found in the lower Arkansas River and the Rio Grande, nor in sandy tributary creeks in eastern Kansas and Missouri that appear to provide suitable habitat. It has already been noted that H. g. gulonella seems to be the ecological equivalent of Semotilus atromaculatus in streams in which S. atromaculatus is not found. S. atromaculatus occurs in creeks of eastern Kansas and Missouri, and may provide interspecific competition that prevents establishment of the flathead chub in these creeks. Regardless of cause, the gaps in distribution of H. gracilis tend to limit gene flow.

Many characters used in the separation of the two subspecies are known to be influenced by environmental conditions, especially temperature. Hubbs (1922, 1926, 1941), Schultz (1927), Vladykov (1934), Tåning (1952) and Weisel (1955), among others, have pointed out a correlation between temperature (or developmental rate of fish) and the number of vertebrae, scales, and fin-rays. Likewise, Martin (1949) and Hart (1952) have shown that the proportions of some body-parts vary in response to temperature during early development. In H. gracilis, the general nature of the clines found in a majority of characters (but not all characters) suggests a temperature influence. However, temperature-dependent variability that has so far been demonstrated experimentally in fishes is generally of lesser magnitude than the differences distinguishing H. g. gracilis and H. g. gulonella. To our knowledge, the most extreme differences that have been induced by modification of temperature are those reported for Salmo trutta by Tåning (1952:181-182), who states: "Shock treatment produced by especially great changes in temperature (c. 10-14° C), especially during the super-[pg 342]sensitive period [of somatic differentiation that fixes vertebral number] may produce ... a difference of 3-4 vertebrae ... in offspring of the same parents." The difference cited approximates that which distinguishes natural populations of H. g. gracilis and H. g. gulonella. Although we cannot assume that the sensitivity of the brown trout is the same as that of the flathead chub, the causative conditions in Tåning's study could scarcely be expected in nature; furthermore, it seems significant that extremely high (as well as extremely low) mean numbers of scales and vertebrae were found at southern localities, and that low mean numbers of scales and vertebrae were found as far north as Wyoming and Montana. We think it likely that temperature does influence the expression of characters in H. gracilis, directly in individual development, and indirectly as a selective mechanism in the evolutionary process. The extent to which each kind of influence exists can be proved only by experimental work with both subspecies, which we hope to undertake at a later date.

Other environmental factors that may have selective influence in this species are rate of current, volume of flow, and turbidity. Interaction of these environmental factors could result in genetic fixation of morphological characters through natural selection. The characters that distinguish H. g. gracilis from H. g. gulonella seem adaptive to life in large rivers and small streams. Evidence that these characters are under limited, direct environmental influence is found among populations in the Arkansas River System. Although populations in the Arkansas River have no continuity with populations of H. g. gracilis, upstream-downstream variations like those found in other river systems are apparent, but in lesser degree. The direction of variation in the Arkansas River is the reverse of that in the Platte and other tributaries of the Missouri River. For example, the populations farthest upstream (Florence, Pueblo) have slightly higher mean numbers of lateral line scales than do populations from Kansas, downstream.

A remarkable effect of extreme parasitism in H. gracilis has been described by Hubbs (1927). Very young chubs that harbored numerous tapeworms (Proteocephalus) had unusually large numbers of lateral-line scales, large eyes, short snouts, small fins, small mouths lacking barbels, and coalescent nares (internarial bridge weak or absent). Some of these abnormalities presumably resulted from retention of larval characteristics of the fish, correlated with the degree of infestation by tapeworms. No teratological adults were found, indicating that severe infections prevent survival to maturity.

[pg 343] H. g. gracilis occurs in three separate river systems (Mackenzie, Saskatchewan, Missouri-Mississippi) from latitude 36° N to 66° N, and longitude 89° W to 123° W. H. g. gulonella exists as several seemingly-isolated populations in the upper parts of the Rio Grande, Pecos, South Canadian, Cimarron, Arkansas, Platte, and upper Missouri basins, from latitude 35° N to 48° N, and longitude 97° W to 100° W.

There is evidence of high mobility on the part of both subspecies, based on irregularity of their occurrence in certain localities. Many collections have been made in the Cimarron River in the vicinity of Kenton, Oklahoma, from 1925 to the present, but only one of these (in 1957) contained flathead chubs. Bait dealers who seine the South Canadian River in Dewey County, Oklahoma, have taken flathead chubs in abundance in some seasons, but not at all in others. Seasonal variation in abundance in the lower Vermillion River, South Dakota (Underhill, 1959:100) has been cited, and the number collected in the lower Kansas River near Lawrence has varied similarly. Many rivers occupied by H. g. gulonella (and by intergrades) are intermittent, and in some years their sand-filled channels become wholly dry for many miles. These factors probably promote mixing of the two subspecies, and may account, over long periods of time, for the wide dispersal of H. g. gulonella in the Missouri Basin. Flathead chubs are known from Pleistocene beds at Doby Springs, Oklahoma (the Doby Springs local fauna) (Smith, 1958:177). Drainage connections between the Arkansas, Kansas and Platte river systems existed in Kansan and Nebraskan times (Frye and Leonard, 1952:189-190). Populations that have subsequently become isolated in those rivers could be accounted for in this way. Flathead chubs could have entered the Rio Grande-Pecos system by stream-capture from the Arkansas System, in northeastern New Mexico or southern Colorado. H. g. gracilis undoubtedly entered the Saskatchewan and Mackenzie basins from the upper Missouri Basin, following glacial retreat (Walters, 1955:347).

LITERATURE CITED

Agassiz, L.

Bailey, Reeve M.

Beckman, William C.

Bryant, Francis T.

Churchill, Edward P., and William H. Over.

Cleary, Robert E.

Clemens, W. A., A. MacDonald, H. McAllister, A. Mansfield and D. Rawson.

Cockerell, Theodora D. A.

Cope, E. D.

Cope, E. D., and H. C. Yarrow.

Cragin, F. W.

Cross, Frank B., Walter W. Dalquest and Leo Lewis.

Cuvier, M. le B^on and M. A. Valenciennes.

Dymond, J. R.

[pg 345] Eddy, Samuel.

Eigenmann, Carl H.

Ellis, Max M.

Evermann, Barton W., and Ulysses O. Cox.

Evermann, Barton Warren, and Edmund Lee Goldsborough.

Forbes, Stephen Alfred, and Robert Earl Richardson.

Frye, J. C., and A. B. Leonard.

Gilbert, Chas. H.

Gill, Theodore.

Gerard, Charles.

Graham, I. D.

Hankinson, Thomas L.

Harlan, James R., and Everett B. Speaker.

Harrison, Harry M., and Everett B. Speaker.

Hart, J. S.

[pg 346] Hinks, David.

Hubbs, Carl L.

Hubbs, Carl L., and Karl F. Lagler.

Hubbs, Carl L., and A. I. Ortenburger.

Jordan, David Starr.

Jordan, David Starr, and Barton Warren Evermann.

Jordan, David Starr, Barton Warren Evermann, and Howard Walton Clark.

Jordan, David Starr, and Charles H. Gilbert.

Jordan, David Starr, and Seth E. Meek.

[pg 347] Keleher, J. J.

Keleher, J. J., and B. Kooyman.

Koster, William J.

Lindsey, C. C.

Martin, W. R.

Meek, Seth E.

Miller, R. B., and M. J. Paetz.

Moore, George A.

O'Donnel, D. John.

Ortenburger, A. I., and Carl L. Hubbs.

Personius, Robert G., and Samuel Eddy.

Rawson, D. S.

Richardson, John.

[pg 348] Schultz, Leonard P.

Scott, W. B.

Shoemaker, Hurst H., Quentin H. Pickering and Leonard Durham.

Simon, James R.

Slastenenko, E. P.

Smith, C. Lavett.

Suckley, George.

Tåning, A. Vedel.

Thompson, Ernest Seton.

Underhill, James C.

Vladykov, Vadim D.

Walters, Vladimir.

Weisel, George F.

Wynne-Edwards, V. C.

Transmitted November 8, 1960.

□
28-5871

End of the Project Gutenberg EBook of Geographic Variation in the North
American Cyprinid Fish, Hybopsis gracilis, by Leonard J. Olund and Frank B. Cross

*** END OF THIS PROJECT GUTENBERG EBOOK GEOGRAPHIC VARIATION ***

***** This file should be named 38425-h.htm or 38425-h.zip *****
This and all associated files of various formats will be found in:
        http://www.gutenberg.org/3/8/4/2/38425/

Produced by Chris Curnow, Joseph Cooper, Erica
Pfister-Altschul and the Online Distributed Proofreading
Team at http://www.pgdp.net


Updated editions will replace the previous one--the old editions
will be renamed.

Creating the works from public domain print editions means that no
one owns a United States copyright in these works, so the Foundation
(and you!) can copy and distribute it in the United States without
permission and without paying copyright royalties.  Special rules,
set forth in the General Terms of Use part of this license, apply to
copying and distributing Project Gutenberg-tm electronic works to
protect the PROJECT GUTENBERG-tm concept and trademark.  Project
Gutenberg is a registered trademark, and may not be used if you
charge for the eBooks, unless you receive specific permission.  If you
do not charge anything for copies of this eBook, complying with the
rules is very easy.  You may use this eBook for nearly any purpose
such as creation of derivative works, reports, performances and
research.  They may be modified and printed and given away--you may do
practically ANYTHING with public domain eBooks.  Redistribution is
subject to the trademark license, especially commercial
redistribution.



*** START: FULL LICENSE ***

THE FULL PROJECT GUTENBERG LICENSE
PLEASE READ THIS BEFORE YOU DISTRIBUTE OR USE THIS WORK

To protect the Project Gutenberg-tm mission of promoting the free
distribution of electronic works, by using or distributing this work
(or any other work associated in any way with the phrase "Project
Gutenberg"), you agree to comply with all the terms of the Full Project
Gutenberg-tm License (available with this file or online at
http://gutenberg.org/license).


Section 1.  General Terms of Use and Redistributing Project Gutenberg-tm
electronic works

1.A.  By reading or using any part of this Project Gutenberg-tm
electronic work, you indicate that you have read, understand, agree to
and accept all the terms of this license and intellectual property
(trademark/copyright) agreement.  If you do not agree to abide by all
the terms of this agreement, you must cease using and return or destroy
all copies of Project Gutenberg-tm electronic works in your possession.
If you paid a fee for obtaining a copy of or access to a Project
Gutenberg-tm electronic work and you do not agree to be bound by the
terms of this agreement, you may obtain a refund from the person or
entity to whom you paid the fee as set forth in paragraph 1.E.8.

1.B.  "Project Gutenberg" is a registered trademark.  It may only be
used on or associated in any way with an electronic work by people who
agree to be bound by the terms of this agreement.  There are a few
things that you can do with most Project Gutenberg-tm electronic works
even without complying with the full terms of this agreement.  See
paragraph 1.C below.  There are a lot of things you can do with Project
Gutenberg-tm electronic works if you follow the terms of this agreement
and help preserve free future access to Project Gutenberg-tm electronic
works.  See paragraph 1.E below.

1.C.  The Project Gutenberg Literary Archive Foundation ("the Foundation"
or PGLAF), owns a compilation copyright in the collection of Project
Gutenberg-tm electronic works.  Nearly all the individual works in the
collection are in the public domain in the United States.  If an
individual work is in the public domain in the United States and you are
located in the United States, we do not claim a right to prevent you from
copying, distributing, performing, displaying or creating derivative
works based on the work as long as all references to Project Gutenberg
are removed.  Of course, we hope that you will support the Project
Gutenberg-tm mission of promoting free access to electronic works by
freely sharing Project Gutenberg-tm works in compliance with the terms of
this agreement for keeping the Project Gutenberg-tm name associated with
the work.  You can easily comply with the terms of this agreement by
keeping this work in the same format with its attached full Project
Gutenberg-tm License when you share it without charge with others.

1.D.  The copyright laws of the place where you are located also govern
what you can do with this work.  Copyright laws in most countries are in
a constant state of change.  If you are outside the United States, check
the laws of your country in addition to the terms of this agreement
before downloading, copying, displaying, performing, distributing or
creating derivative works based on this work or any other Project
Gutenberg-tm work.  The Foundation makes no representations concerning
the copyright status of any work in any country outside the United
States.

1.E.  Unless you have removed all references to Project Gutenberg:

1.E.1.  The following sentence, with active links to, or other immediate
access to, the full Project Gutenberg-tm License must appear prominently
whenever any copy of a Project Gutenberg-tm work (any work on which the
phrase "Project Gutenberg" appears, or with which the phrase "Project
Gutenberg" is associated) is accessed, displayed, performed, viewed,
copied or distributed:

This eBook is for the use of anyone anywhere at no cost and with
almost no restrictions whatsoever.  You may copy it, give it away or
re-use it under the terms of the Project Gutenberg License included
with this eBook or online at www.gutenberg.org

1.E.2.  If an individual Project Gutenberg-tm electronic work is derived
from the public domain (does not contain a notice indicating that it is
posted with permission of the copyright holder), the work can be copied
and distributed to anyone in the United States without paying any fees
or charges.  If you are redistributing or providing access to a work
with the phrase "Project Gutenberg" associated with or appearing on the
work, you must comply either with the requirements of paragraphs 1.E.1
through 1.E.7 or obtain permission for the use of the work and the
Project Gutenberg-tm trademark as set forth in paragraphs 1.E.8 or
1.E.9.

1.E.3.  If an individual Project Gutenberg-tm electronic work is posted
with the permission of the copyright holder, your use and distribution
must comply with both paragraphs 1.E.1 through 1.E.7 and any additional
terms imposed by the copyright holder.  Additional terms will be linked
to the Project Gutenberg-tm License for all works posted with the
permission of the copyright holder found at the beginning of this work.

1.E.4.  Do not unlink or detach or remove the full Project Gutenberg-tm
License terms from this work, or any files containing a part of this
work or any other work associated with Project Gutenberg-tm.

1.E.5.  Do not copy, display, perform, distribute or redistribute this
electronic work, or any part of this electronic work, without
prominently displaying the sentence set forth in paragraph 1.E.1 with
active links or immediate access to the full terms of the Project
Gutenberg-tm License.

1.E.6.  You may convert to and distribute this work in any binary,
compressed, marked up, nonproprietary or proprietary form, including any
word processing or hypertext form.  However, if you provide access to or
distribute copies of a Project Gutenberg-tm work in a format other than
"Plain Vanilla ASCII" or other format used in the official version
posted on the official Project Gutenberg-tm web site (www.gutenberg.org),
you must, at no additional cost, fee or expense to the user, provide a
copy, a means of exporting a copy, or a means of obtaining a copy upon
request, of the work in its original "Plain Vanilla ASCII" or other
form.  Any alternate format must include the full Project Gutenberg-tm
License as specified in paragraph 1.E.1.

1.E.7.  Do not charge a fee for access to, viewing, displaying,
performing, copying or distributing any Project Gutenberg-tm works
unless you comply with paragraph 1.E.8 or 1.E.9.

1.E.8.  You may charge a reasonable fee for copies of or providing
access to or distributing Project Gutenberg-tm electronic works provided
that

- You pay a royalty fee of 20% of the gross profits you derive from
     the use of Project Gutenberg-tm works calculated using the method
     you already use to calculate your applicable taxes.  The fee is
     owed to the owner of the Project Gutenberg-tm trademark, but he
     has agreed to donate royalties under this paragraph to the
     Project Gutenberg Literary Archive Foundation.  Royalty payments
     must be paid within 60 days following each date on which you
     prepare (or are legally required to prepare) your periodic tax
     returns.  Royalty payments should be clearly marked as such and
     sent to the Project Gutenberg Literary Archive Foundation at the
     address specified in Section 4, "Information about donations to
     the Project Gutenberg Literary Archive Foundation."

- You provide a full refund of any money paid by a user who notifies
     you in writing (or by e-mail) within 30 days of receipt that s/he
     does not agree to the terms of the full Project Gutenberg-tm
     License.  You must require such a user to return or
     destroy all copies of the works possessed in a physical medium
     and discontinue all use of and all access to other copies of
     Project Gutenberg-tm works.

- You provide, in accordance with paragraph 1.F.3, a full refund of any
     money paid for a work or a replacement copy, if a defect in the
     electronic work is discovered and reported to you within 90 days
     of receipt of the work.

- You comply with all other terms of this agreement for free
     distribution of Project Gutenberg-tm works.

1.E.9.  If you wish to charge a fee or distribute a Project Gutenberg-tm
electronic work or group of works on different terms than are set
forth in this agreement, you must obtain permission in writing from
both the Project Gutenberg Literary Archive Foundation and Michael
Hart, the owner of the Project Gutenberg-tm trademark.  Contact the
Foundation as set forth in Section 3 below.

1.F.

1.F.1.  Project Gutenberg volunteers and employees expend considerable
effort to identify, do copyright research on, transcribe and proofread
public domain works in creating the Project Gutenberg-tm
collection.  Despite these efforts, Project Gutenberg-tm electronic
works, and the medium on which they may be stored, may contain
"Defects," such as, but not limited to, incomplete, inaccurate or
corrupt data, transcription errors, a copyright or other intellectual
property infringement, a defective or damaged disk or other medium, a
computer virus, or computer codes that damage or cannot be read by
your equipment.

1.F.2.  LIMITED WARRANTY, DISCLAIMER OF DAMAGES - Except for the "Right
of Replacement or Refund" described in paragraph 1.F.3, the Project
Gutenberg Literary Archive Foundation, the owner of the Project
Gutenberg-tm trademark, and any other party distributing a Project
Gutenberg-tm electronic work under this agreement, disclaim all
liability to you for damages, costs and expenses, including legal
fees.  YOU AGREE THAT YOU HAVE NO REMEDIES FOR NEGLIGENCE, STRICT
LIABILITY, BREACH OF WARRANTY OR BREACH OF CONTRACT EXCEPT THOSE
PROVIDED IN PARAGRAPH F3.  YOU AGREE THAT THE FOUNDATION, THE
TRADEMARK OWNER, AND ANY DISTRIBUTOR UNDER THIS AGREEMENT WILL NOT BE
LIABLE TO YOU FOR ACTUAL, DIRECT, INDIRECT, CONSEQUENTIAL, PUNITIVE OR
INCIDENTAL DAMAGES EVEN IF YOU GIVE NOTICE OF THE POSSIBILITY OF SUCH
DAMAGE.

1.F.3.  LIMITED RIGHT OF REPLACEMENT OR REFUND - If you discover a
defect in this electronic work within 90 days of receiving it, you can
receive a refund of the money (if any) you paid for it by sending a
written explanation to the person you received the work from.  If you
received the work on a physical medium, you must return the medium with
your written explanation.  The person or entity that provided you with
the defective work may elect to provide a replacement copy in lieu of a
refund.  If you received the work electronically, the person or entity
providing it to you may choose to give you a second opportunity to
receive the work electronically in lieu of a refund.  If the second copy
is also defective, you may demand a refund in writing without further
opportunities to fix the problem.

1.F.4.  Except for the limited right of replacement or refund set forth
in paragraph 1.F.3, this work is provided to you 'AS-IS' WITH NO OTHER
WARRANTIES OF ANY KIND, EXPRESS OR IMPLIED, INCLUDING BUT NOT LIMITED TO
WARRANTIES OF MERCHANTIBILITY OR FITNESS FOR ANY PURPOSE.

1.F.5.  Some states do not allow disclaimers of certain implied
warranties or the exclusion or limitation of certain types of damages.
If any disclaimer or limitation set forth in this agreement violates the
law of the state applicable to this agreement, the agreement shall be
interpreted to make the maximum disclaimer or limitation permitted by
the applicable state law.  The invalidity or unenforceability of any
provision of this agreement shall not void the remaining provisions.

1.F.6.  INDEMNITY - You agree to indemnify and hold the Foundation, the
trademark owner, any agent or employee of the Foundation, anyone
providing copies of Project Gutenberg-tm electronic works in accordance
with this agreement, and any volunteers associated with the production,
promotion and distribution of Project Gutenberg-tm electronic works,
harmless from all liability, costs and expenses, including legal fees,
that arise directly or indirectly from any of the following which you do
or cause to occur: (a) distribution of this or any Project Gutenberg-tm
work, (b) alteration, modification, or additions or deletions to any
Project Gutenberg-tm work, and (c) any Defect you cause.


Section  2.  Information about the Mission of Project Gutenberg-tm

Project Gutenberg-tm is synonymous with the free distribution of
electronic works in formats readable by the widest variety of computers
including obsolete, old, middle-aged and new computers.  It exists
because of the efforts of hundreds of volunteers and donations from
people in all walks of life.

Volunteers and financial support to provide volunteers with the
assistance they need, are critical to reaching Project Gutenberg-tm's
goals and ensuring that the Project Gutenberg-tm collection will
remain freely available for generations to come.  In 2001, the Project
Gutenberg Literary Archive Foundation was created to provide a secure
and permanent future for Project Gutenberg-tm and future generations.
To learn more about the Project Gutenberg Literary Archive Foundation
and how your efforts and donations can help, see Sections 3 and 4
and the Foundation web page at http://www.pglaf.org.


Section 3.  Information about the Project Gutenberg Literary Archive
Foundation

The Project Gutenberg Literary Archive Foundation is a non profit
501(c)(3) educational corporation organized under the laws of the
state of Mississippi and granted tax exempt status by the Internal
Revenue Service.  The Foundation's EIN or federal tax identification
number is 64-6221541.  Its 501(c)(3) letter is posted at
http://pglaf.org/fundraising.  Contributions to the Project Gutenberg
Literary Archive Foundation are tax deductible to the full extent
permitted by U.S. federal laws and your state's laws.

The Foundation's principal office is located at 4557 Melan Dr. S.
Fairbanks, AK, 99712., but its volunteers and employees are scattered
throughout numerous locations.  Its business office is located at
809 North 1500 West, Salt Lake City, UT 84116, (801) 596-1887, email
[email protected].  Email contact links and up to date contact
information can be found at the Foundation's web site and official
page at http://pglaf.org

For additional contact information:
     Dr. Gregory B. Newby
     Chief Executive and Director
     [email protected]


Section 4.  Information about Donations to the Project Gutenberg
Literary Archive Foundation

Project Gutenberg-tm depends upon and cannot survive without wide
spread public support and donations to carry out its mission of
increasing the number of public domain and licensed works that can be
freely distributed in machine readable form accessible by the widest
array of equipment including outdated equipment.  Many small donations
($1 to $5,000) are particularly important to maintaining tax exempt
status with the IRS.

The Foundation is committed to complying with the laws regulating
charities and charitable donations in all 50 states of the United
States.  Compliance requirements are not uniform and it takes a
considerable effort, much paperwork and many fees to meet and keep up
with these requirements.  We do not solicit donations in locations
where we have not received written confirmation of compliance.  To
SEND DONATIONS or determine the status of compliance for any
particular state visit http://pglaf.org

While we cannot and do not solicit contributions from states where we
have not met the solicitation requirements, we know of no prohibition
against accepting unsolicited donations from donors in such states who
approach us with offers to donate.

International donations are gratefully accepted, but we cannot make
any statements concerning tax treatment of donations received from
outside the United States.  U.S. laws alone swamp our small staff.

Please check the Project Gutenberg Web pages for current donation
methods and addresses.  Donations are accepted in a number of other
ways including checks, online payments and credit card donations.
To donate, please visit: http://pglaf.org/donate


Section 5.  General Information About Project Gutenberg-tm electronic
works.

Professor Michael S. Hart is the originator of the Project Gutenberg-tm
concept of a library of electronic works that could be freely shared
with anyone.  For thirty years, he produced and distributed Project
Gutenberg-tm eBooks with only a loose network of volunteer support.


Project Gutenberg-tm eBooks are often created from several printed
editions, all of which are confirmed as Public Domain in the U.S.
unless a copyright notice is included.  Thus, we do not necessarily
keep eBooks in compliance with any particular paper edition.


Most people start at our Web site which has the main PG search facility:

     http://www.gutenberg.org

This Web site includes information about Project Gutenberg-tm,
including how to make donations to the Project Gutenberg Literary
Archive Foundation, how to help produce our new eBooks, and how to
subscribe to our email newsletter to hear about new eBooks.