Chapter III: Part 3

In the Sierra de los Tuxtlas in southern Veracruz and in the cloud forests along the eastern slopes of the Sierra Madre Oriental northward to Nuevo Leon, _Hyla miotympanum_ seems to be the ecological replacement of _Ptychohyla_. On the Pacific slopes north of Guerrero, Mexico, humid forests in which there are cascading mountain streams are absent; consequently, no _Ptychohyla_ are known from that region. In the mountains of El Salvador _Ptychohyla euthysanota euthysanota_ occurs sympatrically with another small stream-breeding hylid, _Hyla salvadorensis_. To the south of Honduras the highlands diminish into the lowlands of Nicaragua, where habitat suitable for _Ptychohyla_ apparently does not exist. In the mountains of Costa Rica and Panama, the habitats occupied by _Ptychohyla_ in northern Central America are filled by a variety of stream-breeding _Hyla_, such as _Hyla legleri_, _H. rivularis_, _H. rufioculis_, _H. alleei_, and _H. uranochroa_.

Although members of the genus _Ptychohyla_ occur in the southern part of the Mexican Highlands to the west of the Isthmus of Tehuantepec, the greater distribution and differentiation in the genus is in the Chiapan-Guatemalan Highlands. In this respect _Ptychohyla_ is a counterpart of _Plectrohyla_.

Habitat Preference

Frogs of the genus _Ptychohyla_ are ecologically associated with mountain streams at elevations between 650 and 2200 meters; in the geographic region where these frogs occur the vegetation between those elevations consists of cloud forest or pine-oak forest. In some places the frogs have been found in a mixture of oak and semi-deciduous scrub forest. At Vista Hermosa, Oaxaca, _P. leonhardschultzei_ and _P. ignicolor_ were found in cloud forest, whereas at Agua del Obispo, Guerrero, the former species was found in pine-oak forest. _Ptychohyla schmidtorum_ is known only from cloud forest; _P. euthysanota euthysanota_ and _P. spinipollex_ generally are found in cloud forest, but in some places they live in pine-oak forest. _Ptychohyla euthysanota macrotympanum_ has been found in pine-oak forest and in a mixture of oak and semi-deciduous scrub forest. With the possible exception of the members of the _Ptychohyla schmidtorum_ group, which has been found only in cloud forest, it seems as though the type of vegetation is not the controlling factor in the ecological distribution of these frogs.

_Ptychohyla_ has been found only where there are clear, cascading streams overhung by vegetation, on which adults and young perch at night, or even by day. The presence of these streams, in which the tadpoles live, seems to be an important factor in the distribution of _Ptychohyla_. As has been shown previously, the tadpoles of _Ptychohyla_ are adapted for existence in torrential streams, where the water is cool, and the amount of oxygen is high. Clearly these tadpoles are unsuited for life in ponds or sluggish streams in the lowlands, where the temperature of the water is high, a layer of silt on the bottom is deep, and the amount of oxygen is low. The tadpoles cling to rocks on the bottom of the streams; there they move slowly across the rocks, apparently feeding on the thin covering of algae. Tadpoles were not observed on rocks having a thick covering of algae or moss. The tadpoles were observed to swim against the current in torrential streams, in which no fishes were found. Therefore, it seems as though the presence of the stream-habitat for the tadpoles is a significant factor in the ecological distribution of the species of _Ptychohyla_.

Interspecific Competition

At localities where two species of _Ptychohyla_ occur sympatrically (_P. ignicolor_ and _P. leonhardschultzei_ at Vista Hermosa, Oaxaca, and _P. euthysanota euthysanota_ and _P. schmidtorum schmidtorum_ at Finca La Paz, Depto. San Marcos, Guatemala) effort was made to determine what, if any, ecological interspecific relationships existed. Although adults of the sympatric species were found on adjacent leaves or branches of bushes overhanging the streams at both localities, segregation at the time of breeding seems to be maintained by the notably different breeding calls in sympatric species (see discussion of breeding calls). Thus, as has been shown by Blair (1956), Fouquette (1960), and others working on a variety of pond-breeding frogs and toads, the breeding call in _Ptychohyla_ acts as an important reproductive isolating mechanism.

At no locality were _Ptychohyla_ and associated species of hylids found so abundantly as were species of pond-breeding hylids in the lowlands. Apparently reproductive activity is not concentrated in a short breeding season, and it is highly doubtful if the populations of these frogs are as large as those of the lowland pond-breeders. The continual humid conditions and abundance of insect food throughout the year in the cloud forest are perhaps indicative of little interspecific competition among adults of _Ptychohyla_ and other sympatric hylids.

At Finca La Paz, Guatemala, tadpoles of two species of _Ptychohyla_ were ecologically segregated. The tadpoles of _P. euthysanota euthysanota_ were found in riffles in the streams, whereas those of _P. schmidtorum schmidtorum_ were found in slower water, chiefly in small pools in the streams. At Vista Hermosa, Oaxaca, tadpoles of _P. leonhardschultzei_ were found in riffles, and tadpoles of the sympatric _P. ignicolor_ were found in a small pool in a stream. Similar ecological relationships were observed for several species of Costa Rican hylids. Throughout the range of _Ptychohyla_ east of the Isthmus of Tehuantepec, members of the genus occur with species of _Plectrohyla_, all of which are larger than _Ptychohyla_, and all of which have tadpoles that live in torrential streams. Tadpoles of _Ptychohyla spinipollex_ have been found in streams inhabited by the tadpoles of _Plectrohyla guatemalensis_ and _P. quecchi_; tadpoles of _Ptychohyla euthysanota euthysanota_ and _P. schmidtorum schmidtorum_ were found in streams inhabited by tadpoles of _Plectrohyla guatemalensis_, _P. matudai_, and _P. sagorum_. In some streams great numbers of tadpoles occur. The habitat is rather restricted, and the food supply is limited. Consequently, interspecific competition among the various species of hylids whose tadpoles live in the torrential streams probably is highest during the larval stage. Unfortunately, this aspect of salientian population ecology has received no intensive study.

Reproduction and Development

Since the cloud forests inhabited by _Ptychohyla_ are daily bathed in clouds and have a fairly evenly distributed rainfall throughout the year, the frogs living in these forests are active throughout the year. At least some of the species evidently have a long breeding season, for I found calling males of _P. leonhardschultzei_ in February, March, and August, and found tadpoles in February, March, June, and August. Tadpoles of the various species have been obtained throughout much of the year, as follows: _P. euthysanota euthysanota_, February, March, May, and July; _P. euthysanota macrotympanum_, March, June, and August; _P. spinipollex_, February, March, April, June, July, and August; _P. schmidtorum schmidtorum_, March, May, June, July, and August; _P. schmidtorum chamulae_, June and August; _P. ignicolor_, June. I suspect that this temporal distribution more accurately reflects the seasonal activities of collectors than of the frogs.

Calling frogs usually are on vegetation adjacent to or overhanging streams; some calling males of _P. spinipollex_ were on rocks in or by streams. Clasping pairs of _P. euthysanota_ and _P. schmidtorum_ were observed on vegetation by streams. Despite intensive search, no eggs were found. It is doubtful if _Ptychohyla_ deposit eggs on vegetation overhanging streams, as do centrolenids and _Phyllomedusa_, for egg-clutches of these frogs are easily found. Possibly the eggs are laid separately on vegetation above the stream, in which case they could be overlooked easily. In streams where _Ptychohyla_ and other hylid tadpoles occur, empty egg capsules have been found on the lee sides of rocks, but there is no way to determine which species laid the eggs.

Numbers of eggs were counted in gravid females; the largest eggs have diameters ranging from 2.5 to 3.0 mm. The smaller species, comprising the _Ptychohyla schmidtorum_ group, have fewer eggs than do the larger species. Numbers of eggs found in females of the various species are: _P. euthysanota euthysanota_, 108; _P. euthysanota macrotympanum_, 136, 160; _P. leonhardschultzei_, 141; _P. spinipollex_, 119, 134, 143; _P. schmidtorum schmidtorum_, 59, 61, 90; _P. schmidtorum chamulae_, 60, 71, 89.

Duration of the larval stage is unknown. Metamorphosing young have been found from May through August. From two to six completely metamorphosed young are available for each of the species, except for _P. ignicolor_ of which none is available. The smallest young frog is a _P. euthysanota_ having a snout-vent length of 14.2 mm.; the largest young frog is a _P. schmidtorum schmidtorum_ having a snout-vent length of 17.0 mm.

PHYLOGENY OF PTYCHOHYLA

The preceding data on morphology, life histories, and behavior form the basis for the following interpretation of the phylogeny of _Ptychohyla_. Additional data are needed to support some of the ideas discussed below; many of the data that are available for _Ptychohyla_ are lacking for other, possibly related, hylids. The family Hylidae is composed of several hundred species, and most of the species are poorly known. Consequently, any attempt to place _Ptychohyla_ in the over-all scheme of hylid phylogeny would be premature at this time. But, as between the five species of two species-groups here recognized as constituting the genus _Ptychohyla_, some estimate of relationships can be made. First, it is necessary to determine the validity of the genus itself.

Ptychohyla as a Natural Assemblage

As stated in the diagnosis of the genus, the only character that sets this group of species apart from other hylids is the presence of ventrolateral glands in the breeding males. To many systematists the thought of being able to identify to genus only breeding males is sufficiently disturbing to cause them to view with disfavor the recognition of the genus. Nonetheless, the question is raised: Do the five species herein placed in the genus _Ptychohyla_ constitute a natural assemblage? If the genus is considered to be more than a category of convenience, that is to say, a group of related species having a common origin, the primary problem is to determine whether or not the five species form a phylogenetic unit.

The species of _Ptychohyla_ are divided into two groups on the basis of external morphology, breeding calls, and tadpoles. The _Ptychohyla euthysanota_ group seems to be a natural group composed of three species, all of which are more closely related to one another than to any other hylid. Likewise, the species comprising the _Ptychohyla schmidtorum_ group seem to represent a natural unit. If the presence of ventrolateral glands in breeding males is ignored, a student of salientian systematics might derive the _Ptychohyla euthysanota_ group from a hylid stock containing _Hyla miotympanum_ and _Hyla mixomaculata_. Likewise, _Ptychohyla schmidtorum_ could be placed with _Hyla uranochroa_ and related species in Costa Rica. Nonetheless, the fact remains that all of the species assigned to the genus _Ptychohyla_ have ventrolateral glands in the breeding males; furthermore, ventrolateral glands are unknown in other hylids. If the _P. schmidtorum_ group and the _P. euthysanota_ group each evolved from separate hylid stocks, then the ventrolateral glands must have developed independently in both groups. That ventrolateral glands developed independently in five species of frogs in southern Mexico and northern Central America and not in any of the other approximately 500 species of hylids in the world is untenable. It is more logical to assume that the development of the glands took place only once in a stock of hylids that gave rise to the five species herein recognized as members of the genus _Ptychohyla_.

Generic Relationships

The affinities of _Ptychohyla_ apparently are not with any of the other groups that have been generically separated from _Hyla_. Of the daughter genera in Middle America only _Plectrohyla_ has stream-adapted tadpoles, but these large frogs are not closely related to _Ptychohyla_. Stuart (1954:169) suggested that certain montane species of _Hyla_ in lower Central America and _Hyla salvadorensis_ in El Salvador may be related to _Ptychohyla_ or even congeneric. I have had experience with most of these species in the field and believe that Stuart was correct in his suggestion of relationships. The species concerned are four red-eyed stream-breeding _Hyla_ in Costa Rica--_H. alleei_, _H. legleri_, _H. rufioculis_, and _H. uranochroa_, plus _Hyla salvadorensis_ in the mountains of El Salvador. Morphologically all of the species are similar; _Hyla uranochroa_, _H. legleri_, and _H. rufioculis_ have a lateral white stripe that is expanded to form a spot beneath the eye, as in _Ptychohyla schmidtorum_. The tadpoles of _Hyla rufioculis_ and _H. uranochroa_ have large funnel-shaped mouths and long slender tails like those of _Ptychohyla schmidtorum_. Lips of the tadpoles of _H. legleri_ and _H. salvadorensis_ are folded laterally, in this respect resembling those of the _Ptychohyla euthysanota_ group. I do not know the tadpoles and the breeding call of _Hyla alleei_. The breeding calls of _Hyla rufioculis_ and _H. uranochroa_ consist of high melodious notes; the calls of _H. legleri_ and _H. salvadorensis_ consist of series of short notes that have the general characteristics of the call of _Ptychohyla schmidtorum_. Affinities of the genus _Ptychohyla_ seem to me to be with the red-eyed species forming the _Hyla uranochroa_ group in Costa Rica. All of the species in the _Hyla uranochroa_ group have large frontoparietal fontanelles, rather small ethmoids, and small nasals that are not in contact with one another or with the ethmoid. Some species have a complete quadratojugal-maxillary arch; others do not. Assuming that the parental stock that gave rise both to the _Hyla uranochroa_ group and to _Ptychohyla_ was widespread in Central America at a time of cooler, more humid conditions, it is possible that with subsequent warming temperatures and seasonal rainfall in the lowlands the parental stock was restricted to the Costa Rican highlands, where the _Hyla uranochroa_ group developed, and to the Chiapas-Guatemala highlands, where _Ptychohyla_ evolved.

Interspecific Relationships

_Ptychohyla schmidtorum_ is thought to resemble more closely the parental stock of the genus than does any other species of _Ptychohyla_ now extant. This parental stock is discussed above in the account of the generic relationships. _Ptychohyla schmidtorum_ has a red eye, white lateral stripe, frontoparietal fontanelle, funnel-shaped mouth in tadpoles, and lacks nuptial spines; in all of these characters it resembles members of the _Hyla uranochroa_ group. Probably during times of glaciation during the Pleistocene, when climates in Mexico and Central America were depressed, the _Ptychohyla_ stock was more widespread than it is now. Subsequent elevation of climatic zones during interglacial periods would have isolated populations as they are today in regions of cloud forests. Thus, through geographic isolation populations could have differentiated and evolved into the present species. Climatic fluctuation in the Pleistocene must have been of sufficient magnitude to permit the spread of cool, moist forests containing _Ptychohyla_ across the Isthmus of Tehuantepec into the mountains of Oaxaca.

Because of its small nuptial spines, small triangular vomers, coloration, and absence of a rostral keel, _Ptychohyla euthysanota_, more than any of the other species in the _P. euthysanota_ group, resembles _P. schmidtorum_. At the present time _P. euthysanota_ and _P. schmidtorum_ are sympatric.

As I have mentioned previously, ecological segregation and interspecific competition probably is highly developed in the tadpoles of _Ptychohyla_. If this ecological segregation resulted from intraspecific competition in a stock of _Ptychohyla_, possibly _P. euthysanota_ and _P. schmidtorum_ differentiated sympatrically in this way. Specific identity is maintained, at least in part, by different breeding calls in males.

_Ptychohyla spinipollex_ and _P. leonhardschultzei_ seem to be more closely related to one another than either is to _P. euthysanota_. Probably a stock of _P. euthysanota_ was isolated on the Atlantic slopes of northern Central America from _P. euthysanota_ on the southern slopes. The frogs on the Atlantic slopes differentiated and spread into the mountains of Oaxaca, where through isolation by the barrier of the Isthmus of Tehuantepec they developed into _P. leonhardschultzei_, while the stock on the northern slopes of Central America evolved into _P. spinipollex_. Subsequent to the differentiation of _P. leonhardschultzei_ and _P. spinipollex_ from _P. euthysanota_ and during a time of cooler more equable climate than exists now, _P. euthysanota_ and _P. schmidtorum_ invaded the Central Highlands of Chiapas. Subsequent climatic changes isolated populations of each in the Central Highlands, where _P. euthysanota macrotympanum_ and _P. schmidtorum chamulae_ evolved. _Ptychohyla ignicolor_ probably represents stock of _P. schmidtorum_ that crossed the Isthmus of Tehuantepec and became isolated in Oaxaca on the western side of the isthmus.

LITERATURE CITED

AHL, E.
1934. Ueber eine sammlung von Reptilien und Amphibien aus
Mexiko, Zool. Anz., 106:184-186, April 15.

BLAIR, W. F.
1956. Call difference as an isolation mechanism in
southwestern toads (genus _Bufo_). Texas Jour. Sci.,
8:87-106, March.

DUELLMAN, W. E.
1956. The frogs of the hylid genus _Phrynohyas_ Fitzinger,
1843. Misc. Publ. Mus. Zool. Univ. Michigan, 96:1-47,
pls. 1-6, February 21.

1960. Synonymy, variation, and distribution of _Ptychohyla
leonhardschultzei_ Ahl. Studies of American Hylid Frogs,
IV. Herpetologica, 16:191-197, September 23.

1961. Descriptions of two species of frogs, genus Ptychohyla.
Studies of American Hylid Frogs, V. Univ. Kansas Publ.
Mus. Nat. Hist., 13:349-357, pl. 25, April 27.

FOUQUETTE, M. J.
1960. Isolating mechanisms in three sympatric treefrogs in the
Canal Zone. Evolution, 14:484-497, December.

KELLOGG, R.
1928. An apparently new _Hyla_ from El Salvador. Proc. Biol.
Soc. Washington, 41:123-124, June 29.

MERTENS, R.
1952. Die Amphibien und Reptilien von El Salvador.
Senckenbergischen Naturf. Gesell., 487:1-120, pls. 1-16,
December 1.

RIDGWAY, R.
1912. Color standards and color nomenclature. Washington,
D. C., 44 pp., 53 pls.

SHANNON, F. A.
1951. Notes on a herpetological collection from Oaxaca and
other localities in Mexico. Proc. U. S. Nat. Mus.,
101:465-484, May 17.

STUART, L. C.
1954. Descriptions of some new amphibians and reptiles from
Guatemala. Proc. Biol. Soc. Washington, 67:159-178,
August 5.

TANNER, W. W.
1957. Notes on a collection of amphibians and reptiles from
southern Mexico, with a description of a new _Hyla_.
Great Basin Nat., 17:52-56, July 31.

TAYLOR, E. H.
1937. New species of hylid frogs from Mexico with comments on
the rare _Hyla bistincta_ Cope. Proc. Biol. Soc.
Washington, 50:43-54, pls. 2-3, April 21.

1942. New tailless amphibia from Mexico. Univ. Kansas Sci.
Bull., 28: 67-89, May 15.

1944. A new genus and species of Mexican frogs. Univ. Kansas
Sci. Bull., 30:41-45, June 12.

1949. New or unusual Mexican amphibians. Amer. Mus. Novitates,
1437:1-21, December 7.

_Transmitted December 27, 1962._

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Vol. 1. Nos. 1-26 and index. Pp. 1-638, 1946-1950.

*Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest.
Pp. 1-444, 140 figures in text. April 9, 1948.

Vol. 3. *1. The avifauna of Micronesia, its origin, evolution,
and distribution. By Rollin H. Baker. Pp. 1-359,
16 figures in text. June 12, 1951.

*2. A quantitative study of the nocturnal migration of
birds. By George H. Lowery, Jr. Pp. 361-472,
47 figures in text. June 29, 1951.

3. Phylogeny of the waxwings and allied birds. By M.
Dale Arvey. Pp. 473-530, 49 figures in text,
13 tables. October 10, 1951.

*4. Birds from the state of Veracruz, Mexico. By George
H. Lowery, Jr., and Walter W. Dalquest. Pp. 531-649,
7 figures in text, 2 tables. October 10, 1951.

Index. Pp. 651-681.

*Vol. 4. (Complete) American weasels. By E. Raymond Hall.
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Vol. 8. Nos. 1-10 and index. Pp. 1-675, 1954-1956.

Vol. 9. *1. Speciation of the wandering shrew. By James S.
Findley. Pp. 1-68, 18 figures in text.
December 10, 1955.

2. Additional records and extension of ranges of mammals
from Utah. By Stephen D. Durrant, M. Raymond Lee, and
Richard M. Hansen. Pp. 69-80. December 10, 1955.

3. A new long-eared myotis (Myotis evotis) from
northeastern Mexico. By Rollin H. Baker and Howard J.
Stains. Pp. 81-84. December 10, 1955.

4. Subspeciation in the meadow mouse, Microtus
pennsylvanicus, in Wyoming. By Sydney Anderson.
Pp. 85-104, 2 figures in text. May 10, 1956.

5. The condylarth genus Ellipsodon. By Robert W. Wilson.
Pp. 105-116, 6 figures in text. May 19, 1956.

6. Additional remains of the multituberculate genus
Eucosmodon. By Robert W. Wilson. Pp. 117-123,
10 figures in text. May 19, 1956.

7. Mammals of Coahulia, Mexico. By Rollin H. Baker.
Pp. 125-335, 75 figures in text. June 15, 1956.

8. Comments on the taxonomic status of Apodemus
peninsulae, with description of a new subspecies
from North China. By J. Knox Jones, Jr. Pp. 337-346,
1 figure in text, 1 table. August 15, 1956.

9. Extensions of known ranges of Mexican bats. By
Sydney Anderson. Pp. 347-351. August 15, 1956.

10. A new bat (Genus Leptonycteris) from Coahulia. By
Howard J. Stains. Pp. 353-356. January 21, 1957.

11. A new species of pocket gopher (Genus Pappogeomys)
from Jalisco, Mexico. By Robert J. Russell.
Pp. 357-361. January 21, 1957.

12. Geographic variation in the pocket gopher, Thomomys
bottae, in Colorado. By Phillip M. Youngman.
Pp. 363-387, 7 figures in text. February 21, 1958.

13. New bog lemming (genus Synaptomys) from Nebraska.
By J. Knox Jones, Jr. Pp. 385-388. May 12, 1958.

14. Pleistocene bats from San Josecito Cave, Nuevo Leon,
Mexico. By J. Knox Jones, Jr. Pp. 389-396.
December 19, 1958.

15. New subspecies of the rodent Baiomys from Central
America. By Robert L. Packard. Pp. 397-404.
December 19, 1958.

16. Mammals of the Grand Mesa, Colorado. By Sydney
Anderson. Pp. 405-414, 1 figure in text,
May 20, 1959.

17. Distribution, variation, and relationships of the
montane vole, Microtus montanus. By Sydney Anderson.
Pp. 415-511, 12 figures in text, 2 tables.
August 1, 1959.

18. Conspecificity of two pocket mice, Perognathus
goldmani and P. artus. By E. Raymond Hall and Marilyn
Bailey Ogilvie. Pp. 513-518, 1 map. January 14, 1960.

19. Records of harvest mice, Reithrodontomys, from
Central America, with description of a new subspecies
from Nicaragua. By Sydney Anderson and J. Knox Jones,
Jr. Pp. 519-529. January 14, 1960.

20. Small carnivores from San Josecito Cave (Pleistocene),
Nuevo Leon, Mexico. By E. Raymond Hall. Pp. 531-538,
1 figure in text. January 14, 1960.

21. Pleistocene pocket gophers from San Josecito Cave,
Nuevo Leon, Mexico. By Robert J. Russell.
Pp. 539-548,1 figure in text. January 14, 1960.

22. Review of the insectivores of Korea. By J. Knox
Jones, Jr., and David H. Johnson. Pp. 549-578.
February 23, 1960.

23. Speciation and evolution of the pygmy mice, genus
Baimoys. By Robert L. Packard. Pp. 579-670, 4 plates,
12 figures in text. June 16, 1960.

Index. Pp. 671-690

Vol. 10. 1. Studies of birds killed in nocturnal migration. By
Harrison B. Tordoff and Robert M. Mengel. Pp. 1-44,
6 figures in text, 2 tables. September 12, 1956.

2. Comparative breeding behavior of Ammospiza caudacuta
and A. maritima. By Glen E. Woolfenden. Pp. 45-75,
6 plates, 1 figure. December 20, 1956.

3. The forest habitat of the University of Kansas
Natural History Reservation. By Henry S. Fitch and
Ronald R. McGregor. Pp. 77-127, 2 plates, 7 figures
in text, 4 tables. December 31, 1956.

4. Aspects of reproduction and development in the
prairie vole (Microtus ochrogaster). By Henry S.
Fitch. Pp. 129-161, 8 figures in text, 4 tables.
December 19, 1957.

5. Birds found on the Arctic slope of northern Alaska.
By James W. Bee. Pp. 163-211, plates 9-10, 1 figure
in text. March 12, 1958.

*6. The wood rats of Colorado: distribution and ecology.
By Robert B. Finley, Jr. Pp. 213-552, 34 plates,
8 figures in text, 35 tables. November 7, 1958.

7. Home ranges and movements of the eastern cottontail
in Kansas. By Donald W. Janes. Pp. 553-572, 4 plates,
3 figures in text. May 4, 1959.

8. Natural history of the salamander, Aneides hardyi.
By Richard F. Johnston and Gerhard A. Schad.
Pp. 573-585. October 8, 1959.

9. A new subspecies of lizard, Cnemidophorus sacki,
from Michoacan, Mexico. By William E. Duellman.
Pp. 587-598, 2 figures in text. May 2, 1960.

10. A taxonomic study of the middle American snake,
Pituophis deppei. By William E. Duellman.
Pp. 599-610, 1 plate, 1 figure in text. May 2, 1960.

Index. Pp. 611-626.

Vol. 11. Nos. 1-10 and index. Pp. 1-703, 1958-1960.

Vol. 12. 1. Functional morphology of three bats: Sumops, Myotis,
Macrotus. By Terry A. Vaughan. Pp. 1-153, 4 plates,
24 figures in text. July 8, 1959.

*2. The ancestry of modern Amphibia: a review of the
evidence. By Theodore H. Eaton, Jr. Pp. 155-180,
10 figures in text. July 10, 1959.

3. The baculum in microtine rodents. By Sydney Anderson.
Pp. 181-216, 49 figures in text. February 19, 1960.

*4. A new order of fishlike Amphibia from the
Pennsylvanian of Kansas. By Theodore H. Eaton, Jr.,
and Peggy Lou Stewart. Pp. 217-240, 12 figures in
text. May 2, 1960.

5. Natural history of the bell vireo. By Jon C. Barlow.
Pp. 241-296, 6 figures in text. March 7, 1962.

6. Two new pelycosaurs from the lower Permian of
Oklahoma. By Richard C. Fox. Pp. 297-307, 6 figures
in text. May 21, 1962.

7. Vertebrates from the barrier island of Tamaulipas,
Mexico. By Robert K. Selander, Richard F. Johnston,
B. J. Wilks, and Gerald G. Raun. Pp. 309-345,
pls. 5-8. June 18, 1962.

8. Teeth of Edestid sharks. By Theodore H. Eaton, Jr.
Pp. 347-362, 10 figures in text. October 1, 1962.

More numbers will appear in volume 12.

Vol. 13. 1. Five natural hybrid combinations in minnows
(Cyprinidae). By Frank B. Cross and W. L. Minckley.
Pp. 1-18. June 1, 1960.

2. A distributional study of the amphibians of the
Isthmus of Tehuantepec, Mexico. By William E.
Duellman. Pp. 19-72, pls. 1-8, 3 figures in text.
August 16, 1960.

3. A new subspecies of the slider turtle (Pseudemys
scripta) from Coahulia, Mexico. By John M. Legler.
Pp. 73-84, pls. 9-12, 3 figures in text.
August 16, 1960.

4. Autecology of the copperhead. By Henry S. Fitch.
Pp. 85-288, pls. 13-20, 26 figures in text.
November 30, 1960.

5. Occurrence of the garter snake, Thamnophis sirtalis,
in the Great Plains and Rocky Mountains. By Henry S.
Fitch and T. Paul Maslin. Pp. 289-308, 4 figures in
text. February 10, 1961.

6. Fishes of the Wakarusa river in Kansas. By James E.
Deacon and Artie L. Metcalf. Pp. 309-322, 1 figure
in text. February 10, 1961.

7. Geographic variation in the North American cyprinid
fish, Hybopsis gracilis. By Leonard J. Olund and
Frank B. Cross. Pp. 323-348, pls. 21-24, 2 figures
in text. February 10, 1961.

8. Descriptions of two species of frogs, genus
Ptychohyla; studies of American hylid frogs, V.
By William E. Duellman. Pp. 349-357, pl. 25,
2 figures in text. April 27, 1961.

9. Fish populations, following a drought, in the Neosho
and Marais des Cygnes rivers of Kansas. By James
Everett Deacon. Pp. 359-427, pls. 26-30, 3 figs.
August 11, 1961.

10. Recent soft-shelled turtles of North America (family
Trionychidae). By Robert G. Webb. Pp. 429-611,
pls. 31-54, 24 figures in text, February 16, 1962.

Index. Pp. 613-624.

Vol. 14. 1. Neotropical bats from western Mexico. By Sydney
Anderson. Pp. 1-8. October 24, 1960.

2. Geographic variation in the harvest mouse.
Reithrodontomys megalotis, on the central Great
Plains and in adjacent regions. By J. Knox Jones, Jr.,
and B. Mursaloglu. Pp. 9-27, 1 figure in text.
July 24, 1961.

3. Mammals of Mesa Verde National Park, Colorado.
By Sydney Anderson. Pp. 29-67, pls. 1 and 2,
3 figures in text. July 24, 1961.

4. A new subspecies of the black myotis (bat) from
eastern Mexico. By E. Raymond Hall and Ticul Alvarez.
Pp. 69-72, 1 figure in text. December 29, 1961.

5. North American yellow bats, "Dasypterus," and a list
of the named kinds of the genus Lasiurus Gray.
By E. Raymond Hall and J. Knox Jones, Jr. Pp. 73-98,
4 figures in text. December 29, 1961.

6. Natural history of the brush mouse (Peromyscus
boylii) in Kansas with description of a new
subspecies. By Charles A. Long. Pp. 99-111, 1 figure
in text. December 29, 1961.

7. Taxonomic status of some mice of the Peromyscus
boylii group in eastern Mexico, with description of
a new subspecies. By Ticul Alvarez. Pp. 113-120,
1 figure in text. December 29, 1961.

8. A new subspecies of ground squirrel (Spermophilus
spilosoma) from Tamaulipas, Mexico. By Ticul Alvarez.
Pp. 121-124. March 7, 1962.

9. Taxonomic status of the free-tailed bat, Tadarida
yucatanica Miller. By J. Knox Jones, Jr., and Ticul
Alvarez. Pp. 125-133, 1 figure in text.
March 7, 1962.

10. A new doglike carnivore, genus Cynaretus, from the
Clarendonian Pliocene, of Texas. By E. Raymond Hall
and Walter W. Dalquest. Pp. 135-138, 2 figures in
text. April 30, 1962.

11. A new subspecies of wood rat (Neotoma) from
northeastern Mexico. By Ticul Alvarez. Pp. 139-143.
April 30, 1962.

12. Noteworthy mammals from Sinaloa, Mexico. By J. Knox
Jones, Jr., Ticul Alvarez, and M. Raymond Lee.
Pp. 145-159, 1 figure in text. May 18, 1962.

13. A new bat (Myotis) from Mexico. By E. Raymond Hall.
Pp. 161-164, 1 figure in text. May 21, 1962.

14. The mammals of Veracruz. By E. Raymond Hall and
Walter W. Dalquest. Pp. 165-362, 2 figures.
May 20, 1963.

15. The recent mammals of Tamaulipas, Mexico. By Ticul
Alvarez. Pp. 363-473, 5 figures in text.
May 20, 1963.

More numbers will appear in volume 14.

Vol. 15. 1. The amphibians and reptiles of Michoacan, Mexico.
By William E. Duellman. Pp. 1-148, pls. 1-6,
11 figures in text. December 20, 1961.

2. Some reptiles and amphibians from Korea. By Robert
G. Webb, J. Knox Jones, Jr., and George W. Byers.
Pp. 149-173. January 31, 1962.

3. A new species of frog (Genus Tomodactylus) from
western Mexico. By Robert G. Webb. Pp. 175-181,
1 figure in text. March 7, 1962.

4. Type specimens of amphibians and reptiles in the
Museum of Natural History, the University of Kansas.
By William E. Duellman and Barbara Berg. Pp. 183-204.
October 26, 1962.

5. Amphibians and Reptiles of the Rainforests of
Southern El Peten, Guatemala. By William E. Duellman.
Pp. 205-249, pls. 7-10, 6 figures in text.
October 4, 1963.

6. A revision of snakes of the genus Conophis (Family
Colubridae, from Middle America). By John Wellman.
Pp. 251-295, 9 figures in text. October 4, 1963.

7. A review of the Middle American tree frogs of the
genus Ptychohyla. By William E. Duellman.
Pp. 297-349, pls. 11-18, 7 figures in text.
October 18, 1963.

More numbers will appear in volume 15.

* * * * * * *

Transcriber's Notes.

This file was derived from scanned images. With the exception of the list of typographical errors that were corrected below, the original text is presented.

In the original, the Plates were grouped together between pages 328 and 329. Here the Illustration: block which contains the text associated with the Plates were moved just above the text in their respective Systematic Account listing. The Plate text contain the notation "x 2" after the caption to let the reader know that the image was enlarged by a factor of two.

Typographical Errors Corrected:

Several minor typographical corrections were made (missing periods, commas, misspelling of 'and', etc.); but are not indicated here. More substantial changes are listed below:

References to the Plate 11 (Audiograms):
Pl. 1A, Pl. 1B, etc.=> Pl. 11A, Pl. 11B, etc.
Page 301, Paragraph 1: know => known
Page 302, Paragraph 1: Zoolegy => Zoology
Page 303, Paragraph 5: speces => species
Page 305, Paragraph 1: excresences => excrescences
Page 308, Paragraph 6: xiphisterum => xiphisternum
Page 316, Paragraph 3: with => width
Page 327, Paragraph 1: leonhard-schultzei => leonhardschultzei
to match remaining report text
Page 331, Paragraph 1: skelton => skeleton
Publication List Vol. 13, No. 8.: Decriptions => Descriptions

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