Systematic Status of a South American Frog, Allophryne ruthveni Gaige
Author: John D. Lynch; Howard L. Freeman
Date: 2010-02-16T00:00:00+00:00
Keywords: Allophryne ruthveni; Frogs -- South America

The Project Gutenberg eBook of Systematic Status of a South American Frog, Allophryne ruthveni Gaige

This eBook is for the use of anyone anywhere in the United States and most other parts of the world 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. If you are not located in the United States, you will have to check the laws of the country where you are located before using this eBook.

Title: Systematic Status of a South American Frog, Allophryne ruthveni Gaige

Author: John D. Lynch

Howard L. Freeman

Release date: February 16, 2010 [eBook #31293]

Language: English

Other information and formats: www.gutenberg.org/ebooks/31293

Credits: Produced by Chris Curnow, Joseph Cooper, Diane Monico, and
the Online Distributed Proofreading Team at
http://www.pgdp.net

*** START OF THE PROJECT GUTENBERG EBOOK SYSTEMATIC STATUS OF A SOUTH AMERICAN FROG, ALLOPHRYNE RUTHVENI GAIGE ***

University of Kansas Publications
Museum of Natural History

Volume 17, No. 10, pp. 493-502, 3 Figs.
October 27, 1966

Systematic Status of a South American Frog,
Allophryne ruthveni Gaige

BY

JOHN D. LYNCH AND HOWARD L. FREEMAN

University of Kansas
Lawrence
1966

University of Kansas Publications, Museum of Natural History

Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Frank B. Cross

Volume 17, No. 10, pp. 493-502, 3 Figs.
Published October 27, 1966

University of Kansas
Lawrence, Kansas

PRINTED BY
ROBERT R. (BOB) SANDERS, STATE PRINTER
TOPEKA, KANSAS
1966

31-5378

Systematic Status of a South American Frog,
Allophryne ruthveni Gaige

BY

JOHN D. LYNCH AND HOWARD L. FREEMAN

Gaige (1926) described Allophryne ruthveni as a new genus and species of diminutive bufonid from British Guiana. Noble (1931) considered A. ruthveni to be a toothless relative of Centrolenella and placed the genus in the Hylidae. Gallardo (1965) suggested that Allophryne is a leptodactylid of uncertain affinities. Other references to the monotypic genus have consisted only of a listing of the name or of its inclusion in a key. To date the holotype and one paratype (both females) have been reported (Gaige, 1926), and the family position of the genus remains unsettled.

A male of Allophryne ruthveni is among the amphibians and reptiles collected in southern British Guiana by William A. Bentley in January, 1962, and deposited in the Museum of Natural History at The University of Kansas (KU). Four additional specimens (females) are in the American Museum of Natural History; only one of the latter has definite locality data.

Acknowledgments.—We are grateful to Dr. Ernest E. Williams, Museum of Comparative Zoology (MCZ) and Dr. Richard G. Zweifel, American Museum of Natural History (AMNH) for the loan of specimens. We are further indebted to Dr. Zweifel for permission to clear and stain one specimen. Dr. William E. Duellman and Linda Trueb offered many constructive criticisms. Miss Trueb executed the drawings of the skull and finger bones. Mr. Martin Wiley provided x-ray photographs of Allophryne.

METHODS AND MATERIALS

Six of the seven known specimens were available for study. Measurements were taken in the manner described by Duellman (1956). One specimen was cleared and stained, using the technique of Davis and Gore (1936), in order to study the skeleton. X-ray photographs were made of another specimen for comparison.

Specimens examined.—Six, as follows: BRITISH GUIANA, Dist. Demarara: Marudi Creek, AMNH 44749; Dist. Equibo: Tumatumari, MCZ 11790 (paratype); Dist. Rupununi (Berbice): Wai Wai Country, N of Acarahy Mountains, west of New River (2°N, 58°W), KU 69890. Also, 3 specimens from "probably British Guiana," AMNH 70108-10 (70110 cleared and stained).

SYSTEMATIC ACCOUNT

The availability of additional material and the new information pertaining to osteology permit an amplification of Gaige's (1926) description.

Genus Allophryne Gaige

Allophryne Gaige, Occas. Papers Mus. Zool., Univ. Michigan, 176:1, Oct. 14, 1926. Crawford, Annals Carnegie Mus., 21(1):29, 32, Nov. 14, 1931. Noble, The biology of the amphibia. McGraw-Hill, p. 510, 1931. Ruthven, Herpetologica, 1:3, July 11, 1936. Gallardo, Papéis Avulsos, 17:79, Jan. 1, 1965.

Type species.—Allophryne ruthveni Gaige.

Diagnosis and definition.—A genus of diminutive frogs; vomers, maxillae, and premaxillae edentate; skin of head strongly anchored to connective tissue on cranium; prepollical spine absent in males; disk of third finger larger than tympanum, smaller than eye; no humeral hook in either sex; ilia extending anteriorly beyond sacral expansions; adults attaining snout-vent length of 31 mm.; male having darkened external subgular vocal sac; skin of dorsum pustulate.

Allophryne ruthveni Gaige

Allophryne ruthveni Gaige, Occas. Papers Mus. Zool., Univ. Michigan, 176:1-3, pl. I, Oct. 14, 1926. Crawford, Annals Carnegie Mus., 21(1):32, Nov. 14, 1931. Ruthven, Herpetologica, 1:3, July 11, 1936. Barbour and Loveridge, Bull. Mus. Comp. Zool., 96(2):64, Feb., 1946. Peters, Occas. Papers Mus. Zool., Univ. Michigan, 539:10, Sept. 19, 1952.

Holotype.—University of Michigan Museum of Zoology 63419, adult female, from Tukeit Hill, below Kaiteur Falls, Equibo District, British Guiana; obtained in May, 1924, by E. N. Clarke.

Diagnosis.—Fingers free; toes two-thirds webbed; no supernumerary tubercles on soles or palms; no tarsal fold; elongate anal sheath, anal opening on lower surface of thighs; head broad, interorbital space 2.5 times width of upper eyelid; snout subacuminate in dorsal profile, strongly sloping in lateral profile; tympanum visible in males, concealed in females; venter areolate.

External Morphology.—(Fig. 1) Additional features not mentioned in diagnoses: Head wider than long, about as wide as body; supratympanic fold present; canthus rostralis rounded, loreal region slightly concave, nearly vertical; nostril at tip of snout; pupil horizontal; no teeth on maxillary, premaxillary, or vomer; tongue small, round, thick, not notched behind, free posteriorly for one-sixth of length; choanae large, only partly visible from directly below; males having darkened subgular vocal sac; vocal slits present in male.

Axillary membrane lacking or but slightly developed; no tubercles or ridge under forearm; two palmar tubercles; subarticular tubercles small, simple, round, flattened; tips of fingers slightly expanded, T-shaped, with prominent transverse groove; first finger shorter than second (stated as longer than second in diagnosis by Gaige, 1926:2); folds extending laterally from anus for a short distance, then downward to venter of thighs; no appendage on heel, no inner or outer tarsal folds or tubercles; inner metatarsal tubercle oval, about twice as long as wide; outer metatarsal tubercle nearly absent; no supernumerary tubercle on sole; subarticular tubercles on foot small, round, simple, and diffuse; toes T-shaped, slightly wider than digit; toes about two-thirds webbed (Fig. 1d).

Skin of venter coarsely areolate; skin of flanks, throat, chest, undersurfaces of arms, tibia, tarsi, dorsal surfaces of thighs, tarsi, hands, and feet smooth; skin of dorsal surfaces of tibia, forearm, back, and top and sides of head having large horny pustules (sharply spinous in male).

Fig. 1. Allophryne ruthveni, male (KU 69890); (a) Dorsum.
(b) Thenar view of right hand. (c) Lateral profile of head.
(d) Plantar view of right foot. × 3.5.

Color.—Dorsum gray with irregular network of black lines and elongate blotches; flanks and labial region black with large white ocelli; dorsal surfaces of limbs gray, marked as follows: two large, elongate white spots on each thigh, concealed white spot on base of upper arm, black-edged gray transverse bars on forearms and shanks, white spot on each knee and elbow; ventral surfaces pale gray; black-edged white spot on ventral surface of thigh on each side of anal opening; chin and throat dark gray with white spots; vocal sac in male black (Fig. 1a and c).

Gaige (1926) briefly described the color, which conforms to the above in all particulars. The paratype (MCZ 11790) has lost the gray color after 40 years in preservation; now (1966) the ground-color is cream-brown, and the dorsal spotting, noted by Gaige as being black, is now brown.

The spots on the feet, tarsi, knees, thighs, flanks and upper arm are white in preservative, but in life possibly were red or yellow. These colors usually fade to white in preservative. Red or yellow spots are common aposematic colors in frogs.

Variation.—Eight measurements were taken on each specimen and four ratios were computed; these are summarized in Table 1. Gaige's illustration of the holotype shows that it has a greatly reduced pattern, whereas the paratype and three of the other five known specimens have relatively large and numerous spots. The male (KU 69890) and one female (AMNH 70108) have a reduced pattern intermediate between that of the holotype and the four other specimens.

TABLE I.—Variation in Measurements and Proportions of Allophryne ruthveni. (Ranges in parentheses below means.)

Character | Male (1) | Females (5)

Snout-vent (in mm.)

20.6

23.6
(18.4-31.0)[A]

Tibia/snout-vent

0.43

0.43
(0.41-0.47)

Tympanum/head width

0.12

0.15
(0.14-0.16)

Eyelid/interorbital space

0.55

0.53
(0.49-0.56)

Tympanum/eye length

0.40

0.46
(0.42-0.50)

[A]Holotype is reported to be 31 mm. snout-vent length (Gaige, 1926). The largest measured by us was 26.2 mm. snout-vent.

The dorsal spinules are most pronounced and extensive on the male (Fig. 1) and less so in all other specimens examined. The illustration of the holotype suggests that it has equally prominent, but fewer, spinules (Gaige, 1926).

The holotype, a gravid female, is the largest known specimen (31 mm., snout-vent length). Another gravid female (AMNH 70108) has a snout-vent length of 26.2 mm.

Distribution.—All known specimens have been found in the foothills of the northeastern face of the Guiana Massif in British Guiana.

FAMILY POSITION

The following characters of Allophryne are those generally held to be useful in determining family relationships:

1. Presacral vertebrae procoelus, eight in number.

2. Parahyoid absent.

3. Free ribs lacking.

4. Bidder's organ absent.

5. Intercalary cartilages present in digits; phalangeal formulae 3-3-4-4 and 3-3-4-5-4.

6. Coccyx articulating with sacrum by two condyles.

7. Tarsal bones not fused.

8. Pectoral girdle arciferal.

9. Epicoracoidal horns present, free.

10. Terminal phalanges T-shaped.

11. Sacrum procoelus and diapophyses expanded.

12. Maxillae, premaxillae, and prevomers edentate.

13. Cranial roofing bones well ossified.

Griffiths (1959) accorded considerable taxonomic weight to the presence or absence of epicoracoidal horns in showing relationships among the genera placed in the Brachycephalidae [= Atelopodidae; Dendrobatidae; and Leptodactylidae (in part)] by Noble (1931). Allophryne possesses well-developed, free epicoracoidal horns, such as those found in the Hylidae, Centrolenidae, Leptodactylidae and Bufonidae.

The presence of intercalary elements in the digits is characteristic of the Centrolenidae, Hylidae, Phrynomeridae, Pseudidae, and the rhacophorine ranids (including the Hyperoliidae). This element is bony in the pseudids and cartilaginous in the other families. Phrynomerids and rhacophorine ranids lack epicoracoidal horns and have firmisternal pectoral girdles. Centrolenids are small, delicate, arboreal frogs having poorly ossified skulls and fused tarsal bones, but agree with Allophryne in having T-shaped terminal phalanges.

Fig. 2. Dorsal (a) and lateral (b) views of distal phalanges of third finger of Allophryne. × 40.

Only the presence of intercalary cartilages (Fig. 2) suggests relationship of Allophryne to the Hylidae. The T-shaped terminal phalanges suggest affinities with centrolenids, elutherodactyline leptodactylids, or certain "brachycephalid" frogs. Griffiths (1959) clearly showed that Noble's Brachycephalidae was a polyphyletic assemblage. No hylid genus is edentate, and none has either T-shaped terminal phalanges or the unusual dorsal spinules. Perhaps the presence of intercalary cartilages is not indicative of relationship but instead is a parallelism (or convergence) in Allophryne and genera of the Centrolenidae.

CRANIAL OSTEOLOGY

The skull of Allophryne (Fig. 3) is distinctive among anurans; it does not closely resemble the skulls of either hylids or centrolenids, both of which have generally more delicate (except for casque-headed hylids, such as Corythomantis, Diaglena, Osteocephalus, Triprion) and generalized skulls. Allophryne on the other hand has a strongly ossified central region (cranial roofing bones and sphenethmoid complex) and a weak peripheral zone. The peripheral elements are reduced (maxilla, pterygoid, and squamosal) or absent (quadratojugal), whereas the frontoparietals, nasals, sphenethmoid, proötics, and exoccipitals form a compact central zone. An elongate frontoparietal fontanelle is present.

Fig. 3. Dorsal view of skull of Allophryne (AMNH 70110). × 12.

Dorsally (Fig. 3), the premaxillae are not visible. The proportionally gigantic septomaxillae are visible anterior to the nasals. The moderate-sized nasals are separated medially and in broad contact with the ethmoid posteriorly. The palatine process of the nasal does not meet the frontal process of the maxilla. A large frontoparietal fontanelle is evident between the frontoparietals. The tegmen tympani are much reduced and maintain only cartilaginous contact with the posterior arms of the squamosals. The foramen magnum, occipital condyles, and exoccipitals show no unusual features. The pars facialis and frontal process of the maxilla are greatly reduced. The maxilla and premaxilla are articulated. The high, narrow alary processes of the premaxillae extend dorsally about two-thirds of the height of the snout. A cartilaginous internasal septum is illustrated (Fig. 3), but sectioning is necessary to determine the true nature and extent of this element.

Ventrally, the skull lacks palatines. The maxillae, premaxillae, and prevomers are edentate. The parasphenoid is large with relatively short, stout alary (lateral) processes. The sphenethmoid is extensive in ventral aspect and forms the major supporting structure in the anterior part of the skull. The pterygoid has a broad articulation with the maxilla, a tenuous contact with the squamosal, but is not attached to the proötic. The anterior (zygomatic) process of the squamosal is greatly reduced (only about one-third the length of the posterior process).

DISCUSSION

The skull of Allophryne is definitely non-hylid. Most of the post-cranial features do not help to clarify relationships. Allophryne shares several osteological features with the Dendrobatidae: T-shaped terminal phalanges, general cranial morphology and procoelus vertebrae. But, the dendrobatids possess firmisternal pectoral girdles and lack epicoracoidal horns. Also, no dendrobatid has intercalary elements in the digits. We are, therefore, left with a taxonomic enigma. In one or more characters generally regarded as important, Allophryne differs from all presently defined families of frogs. The Hylidae and Dendrobatidae are the only currently recognized families in which the genus might be placed.

The function and taxonomic importance of the large septomaxillae are unknown and are probably associated with the modification of the sphenethmoid-prevomer area. A more detailed study of the cranial osteology of Allophryne, especially the structural relationships of the sphenethmoid-prevomer area may elucidate the relationships of Allophryne.

The relationships of Allophryne cannot be understood without a re-analysis of some of the features used as major criteria in frog classification (the nature of an intercalated cartilage; the nature of the sternal complex; the relative value of cranial osteology; the vertebral structure; and the thigh musculature). Some of these features have been investigated by other workers, most notably Griffiths, but others have not and need re-examination. A re-analysis of some of the major criteria used in frog classification is in progress (Callison, Lynch, and Trueb) and upon completion of that study we think the relationships of Allophryne will become apparent.

A more comprehensive study of the cranial anatomy of certain hylids, leptodactylids, dendrobatids, and atelopodids along with that of Allophryne is needed to clarify the relationships of Allophryne, and might indicate that the recognition of a fifth family is necessary.

CONCLUSION

Among currently recognized families of frogs, Allophryne is least different from the Hylidae although it is our opinion that inclusion of this genus in the Hylidae probably represents an unnatural classification. However, the present evidence suggesting that Allophryne should be in another family is less convincing than evidence suggesting it should be in the Hylidae. We tentatively place Allophryne in the Hylidae.

LITERATURE CITED

Davis, D. D. and Gore, U. R.

1936. Clearing and staining skeletons of small vertebrates. Fieldiana: Technique, 4:1-16.

Duellman, W. E.

1956. The frogs of the hylid genus Phrynohyas Fitzinger, 1843. Misc. Publs. Mus. Zool., Univ. Michigan, 96:1-47, February 21.

Gaige, H. T.

1926. A new frog from British Guiana. Occas. Papers Mus. Zool., Univ. Michigan, 176:1-3, October 14.

Gallardo, J. M.

1965. A propósito de los Leptodactylidae (Amphibia Anura). Papéis Avulsos, 17:77-87, January 1.

Griffiths, I.

1959. The phylogeny of Sminthillus limbatus and the status of the Brachycephalidae (Amphibia: Salientia). Proc. Zool. Soc. London, 132:457-87, May.

Noble, G. K.

1931. The biology of the amphibia. McGraw-Hill, New York, vii + 577 pp.

Transmitted August 2, 1966.

31-5378

Transcriber's Notes

Table 1 and Figs. 2 and 3 have been moved slightly to avoid breaking up the paragraphs of text.

*** END OF THE PROJECT GUTENBERG EBOOK SYSTEMATIC STATUS OF A SOUTH AMERICAN FROG, ALLOPHRYNE RUTHVENI GAIGE ***

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

Creating the works from print editions not protected by U.S. copyright law 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™ electronic works to protect the PROJECT GUTENBERG™ concept and trademark. Project Gutenberg is a registered trademark, and may not be used if you charge for an eBook, except by following the terms of the trademark license, including paying royalties for use of the Project Gutenberg trademark. If you do not charge anything for copies of this eBook, complying with the trademark license is very easy. You may use this eBook for nearly any purpose such as creation of derivative works, reports, performances and research. Project Gutenberg eBooks may be modified and printed and given away—you may do practically ANYTHING in the United States with eBooks not protected by U.S. copyright law. 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™ 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 License available with this file or online at www.gutenberg.org/license.

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

1.A. By reading or using any part of this Project Gutenberg 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 electronic works in your possession. If you paid a fee for obtaining a copy of or access to a Project Gutenberg 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 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 electronic works if you follow the terms of this agreement and help preserve free future access to Project Gutenberg 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 electronic works. Nearly all the individual works in the collection are in the public domain in the United States. If an individual work is unprotected by copyright law 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 mission of promoting free access to electronic works by freely sharing Project Gutenberg works in compliance with the terms of this agreement for keeping the Project Gutenberg 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 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 work. The Foundation makes no representations concerning the copyright status of any work in any country other than 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 License must appear prominently whenever any copy of a Project Gutenberg 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 in the United States and most other parts of the world 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. If you are not located in the United States, you will have to check the laws of the country where you are located before using this eBook.

1.E.2. If an individual Project Gutenberg electronic work is derived from texts not protected by U.S. copyright law (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 trademark as set forth in paragraphs 1.E.8 or 1.E.9.

1.E.3. If an individual Project Gutenberg 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 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 License terms from this work, or any files containing a part of this work or any other work associated with Project Gutenberg.

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 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 work in a format other than “Plain Vanilla ASCII” or other format used in the official version posted on the official Project Gutenberg website (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 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 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 electronic works provided that:

- • You pay a royalty fee of 20% of the gross profits you derive from the use of Project Gutenberg works calculated using the method you already use to calculate your applicable taxes. The fee is owed to the owner of the Project Gutenberg 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™ 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™ 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™ works.

1.E.9. If you wish to charge a fee or distribute a Project Gutenberg™ electronic work or group of works on different terms than are set forth in this agreement, you must obtain permission in writing from the Project Gutenberg Literary Archive Foundation, the manager of the Project Gutenberg™ 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 works not protected by U.S. copyright law in creating the Project Gutenberg™ collection. Despite these efforts, Project Gutenberg™ 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™ trademark, and any other party distributing a Project Gutenberg™ 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 1.F.3. 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 MERCHANTABILITY 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™ electronic works in accordance with this agreement, and any volunteers associated with the production, promotion and distribution of Project Gutenberg™ 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 work, (b) alteration, modification, or additions or deletions to any Project Gutenberg work, and (c) any Defect you cause.

Section 2. Information about the Mission of Project Gutenberg

Project Gutenberg 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’s goals and ensuring that the Project Gutenberg 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 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 information page at www.gutenberg.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. 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 business office is located at 41 Watchung Plaza #516, Montclair NJ 07042, USA, +1 (862) 621-9288. Email contact links and up to date contact information can be found at the Foundation’s website and official page at www.gutenberg.org/contact

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

Project Gutenberg™ depends upon and cannot survive without widespread 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 www.gutenberg.org/donate.

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: www.gutenberg.org/donate.

Section 5. General Information About Project Gutenberg electronic works

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

Project Gutenberg eBooks are often created from several printed editions, all of which are confirmed as not protected by copyright 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 website which has the main PG search facility: www.gutenberg.org.

This website includes information about Project Gutenberg, 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.

Table of Contents

- Systematic Status of a South American Frog, Allophryne ruthveni Gaige

- Systematic Status of a South American Frog, Allophryne ruthveni Gaige

- METHODS AND MATERIALS

- SYSTEMATIC ACCOUNT

- Genus Allophryne Gaige

- Allophryne ruthveni Gaige

- FAMILY POSITION
- CRANIAL OSTEOLOGY
- DISCUSSION
- CONCLUSION
- LITERATURE CITED
- Transcriber's Notes

- THE FULL PROJECT GUTENBERG™ LICENSE