diff options
| -rw-r--r-- | .gitattributes | 3 | ||||
| -rw-r--r-- | 37809-8.txt | 2535 | ||||
| -rw-r--r-- | 37809-8.zip | bin | 0 -> 40689 bytes | |||
| -rw-r--r-- | 37809-h.zip | bin | 0 -> 866974 bytes | |||
| -rw-r--r-- | 37809-h/37809-h.htm | 3542 | ||||
| -rw-r--r-- | 37809-h/images/bar_double.png | bin | 0 -> 162 bytes | |||
| -rw-r--r-- | 37809-h/images/bar_single.png | bin | 0 -> 160 bytes | |||
| -rw-r--r-- | 37809-h/images/cover.jpg | bin | 0 -> 18181 bytes | |||
| -rw-r--r-- | 37809-h/images/fig_1.png | bin | 0 -> 46867 bytes | |||
| -rw-r--r-- | 37809-h/images/fig_10.png | bin | 0 -> 34413 bytes | |||
| -rw-r--r-- | 37809-h/images/fig_11.png | bin | 0 -> 16617 bytes | |||
| -rw-r--r-- | 37809-h/images/fig_12.png | bin | 0 -> 42202 bytes | |||
| -rw-r--r-- | 37809-h/images/fig_13.png | bin | 0 -> 74348 bytes | |||
| -rw-r--r-- | 37809-h/images/fig_14.png | bin | 0 -> 25564 bytes | |||
| -rw-r--r-- | 37809-h/images/fig_15.png | bin | 0 -> 4997 bytes | |||
| -rw-r--r-- | 37809-h/images/fig_16.png | bin | 0 -> 45757 bytes | |||
| -rw-r--r-- | 37809-h/images/fig_17.png | bin | 0 -> 14431 bytes | |||
| -rw-r--r-- | 37809-h/images/fig_18.png | bin | 0 -> 42146 bytes | |||
| -rw-r--r-- | 37809-h/images/fig_19.png | bin | 0 -> 49192 bytes | |||
| -rw-r--r-- | 37809-h/images/fig_2.png | bin | 0 -> 19329 bytes | |||
| -rw-r--r-- | 37809-h/images/fig_20.png | bin | 0 -> 27732 bytes | |||
| -rw-r--r-- | 37809-h/images/fig_21.png | bin | 0 -> 29395 bytes | |||
| -rw-r--r-- | 37809-h/images/fig_22.png | bin | 0 -> 11275 bytes | |||
| -rw-r--r-- | 37809-h/images/fig_3.png | bin | 0 -> 59319 bytes | |||
| -rw-r--r-- | 37809-h/images/fig_4.png | bin | 0 -> 40944 bytes | |||
| -rw-r--r-- | 37809-h/images/fig_5.png | bin | 0 -> 40966 bytes | |||
| -rw-r--r-- | 37809-h/images/fig_6.png | bin | 0 -> 63780 bytes | |||
| -rw-r--r-- | 37809-h/images/fig_7.png | bin | 0 -> 19471 bytes | |||
| -rw-r--r-- | 37809-h/images/fig_8.png | bin | 0 -> 50684 bytes | |||
| -rw-r--r-- | 37809-h/images/fig_9.png | bin | 0 -> 41802 bytes | |||
| -rw-r--r-- | 37809.txt | 2535 | ||||
| -rw-r--r-- | 37809.zip | bin | 0 -> 40566 bytes | |||
| -rw-r--r-- | LICENSE.txt | 11 | ||||
| -rw-r--r-- | README.md | 2 |
34 files changed, 8628 insertions, 0 deletions
diff --git a/.gitattributes b/.gitattributes new file mode 100644 index 0000000..6833f05 --- /dev/null +++ b/.gitattributes @@ -0,0 +1,3 @@ +* text=auto +*.txt text +*.md text diff --git a/37809-8.txt b/37809-8.txt new file mode 100644 index 0000000..91c382c --- /dev/null +++ b/37809-8.txt @@ -0,0 +1,2535 @@ +The Project Gutenberg EBook of A Taxonomic Revision of the Leptodactylid +Frog Genus Syrrhophus Cope, by John D. Lynch + +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: A Taxonomic Revision of the Leptodactylid Frog Genus Syrrhophus Cope + +Author: John D. Lynch + +Release Date: October 21, 2011 [EBook #37809] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK A TAXONOMIC REVISION OF THE *** + + + + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper and +the Online Distributed Proofreading Team at +http://www.pgdp.net + + + + + + + + + + UNIVERSITY OF KANSAS + MUSEUM OF NATURAL HISTORY + + Vol. 20, No. 1, pp. 1-45, 22 figs. + + February 20, 1970 + + A Taxonomic Revision + of the Leptodactylid Frog Genus + Syrrhophus Cope + + BY + + JOHN D. LYNCH + + UNIVERSITY OF KANSAS + LAWRENCE + 1970 + + + + + UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY + + Editors of this number: + Frank B. Cross, Philip S. Humphrey, William E. Duellman + + Volume 20, No. 1, pp. 1-45, 22 figs. + Published February 20, 1970 + + UNIVERSITY OF KANSAS + Lawrence, Kansas + + PRINTED BY + THE UNIVERSITY OF KANSAS PRINTING SERVICE + LAWRENCE, KANSAS + 1970 + + + + +A Taxonomic Revision of the Leptodactylid Frog Genus Syrrhophus Cope + +BY + +JOHN D. LYNCH + + + + +INTRODUCTION + + +Cope (1878) proposed the genus _Syrrhophus_ for a medium-sized +leptodactylid frog from central Texas; in the ensuing 75 years the genus +was expanded to include a heterogeneous group of frogs ranging from +Texas to Peru. Taylor (1952) and Firschein (1954) limited the genus to +several species of frogs occurring in Guatemala, México, and Texas. +Lynch (1968) provided a definition of the previously loosely-defined +genus. + +With the exception of Taylor (1952), who treated the Costa Rican +species, none of these authors dealt with the present status of the +nineteen species erroneously assigned to _Syrrhophus_. These species are +listed in Tables 1 and 2 with the name currently applied. Some of them +are new combinations and their justifications will be published +elsewhere. Gorham (1966) is the most recent author to include South +American species in the genus _Syrrhophus_. + +Smith and Taylor (1948) recognized two species groups of the genus in +México, an eastern and a western group (here termed complexes for +purposes of discussion), separated on the basis of the number of palmar +(metacarpal) tubercles (three palmar tubercles in the members of the +eastern complex and two in those of the western complex). Duellman +(1958) reviewed the species of the genus occurring in western México and +concluded that there were five species (two polytypic). Dixon and Webb +(1966) described an additional species from Jalisco, México. The +distributions of some species have been extended, but otherwise the +western complex of species remains unchanged since Duellman's review. + +Smith and Taylor (1948) recognized seven species of the genus in eastern +México. Firschein revised the eastern complex (as then understood), and +in so doing added one new species and treated _Syrrhophus verruculatus_ +as a _nomen dubium_. Dixon (1957) redefined the related genus +_Tomodactylus_ and transferred _T. macrotympanum_ Taylor to the genus +_Syrrhophus_. Neill (1965) described a new subspecies of _S. leprus_ +from British Honduras. Two species (_S. gaigeae_ and _S. marnockii_) +were recognized in Texas until Milstead, Mecham, and McClintock (1950) +synonymized _S. gaigeae_ with _S. marnockii_. Thus, at present, nine +species (one polytypic) are recognized on the eastern slopes and +lowlands from central Texas to British Honduras. These are currently +placed on one species group equivalent to the western complex reviewed +by Duellman (1958). + + + TABLE 1--Species Described as Members of the Genus _Syrrhophus_ but + Now Placed in Other Genera. + + ======================================================================= + Trivial name and author Current combination + ----------------------------------------------------------------------- + _areolatus_ Boulenger, 1898 _Eleutherodactylus areolatus_ + _calcaratus_ Andersson, 1945 _Eleutherodactylus anderssoni_ + _caryophyllaceus_ Barbour, 1928 _Eleutherodactylus caryophyllaceus_ + _coeruleus_ Andersson, 1945 _Eleutherodactylus coeruleus_ + _ineptus_ Barbour, 1928 _Eleutherodactylus diastema_ + _juninensis_ Shreve, 1938 _Eupsophus juninensis_ + _lutosus_ Barbour and Dunn, 1921 _Eleutherodactylus lutosus_ + _molinoi_ Barbour, 1928 _Eleutherodactylus molinoi_ + _montium_ Shreve, 1938 _Niceforonia montia_ + _mystaceus_ Barbour, 1922 _Eleutherodactylus rhodopis_ + _obesus_ Barbour, 1928 _Eleutherodactylus punctariolus_ + _omiltemanus_ Gunther, 1900 _Eleutherodactylus omiltemanus_[1] + _pardalis_ Barbour, 1928 _Eleutherodactylus pardalis_ + ======================================================================= + + [1] New combination. + + + TABLE 2--Species Incorrectly Regarded as Members of the Genus _Syrrhophus_ + but Described as Members of Other Genera. + + ========================================================================== + Trivial name, original generic + assignment, and author Current combination + -------------------------------------------------------------------------- + _chalceus_ (_Phyllobates_) Peters, 1873 _Eleutherodactylus chalceus_ + _festae_ (_Paludicola_) Peracca, 1904 _Niceforonia festae_ + _hylaeformis_ (_Phyllobates_) Cope, 1875 _Eleutherodactylus hylaeformis_ + _palmatus_ (_Phyllobates_) Werner, 1899 _Colostethus palmatus_ + _ridens_ (_Phyllobates_) Cope, 1866 _Eleutherodactylus ridens_ + _simonsii_ (_Paludicola_) Boulenger, 1900 _Niceforonia simonsii_ + ========================================================================== + + + TABLE 3--Nominal Species of _Syrrhophus_ (_sensu strictu_) and the Name + Used Herein. + + ======================================================================= + Original combination Current combination + ----------------------------------------------------------------------- + _campi_, _Syrrhophus_ _cystignathoides campi_ + _cholorum_, _Syrrhophus leprus_ _leprus_ + _cystigathoides_, _Phyllobates_ _cystignathoides cystignathoides_ + _dennisi_, _Syrrhophus_ _dennisi_ new species + _gaigeae_, _Syrrhophus_ _guttilatus_ + _guttilatus_, _Malachylodes_ _guttilatus_ + _interorbitalis_, _Syrrhophus_ _interorbitalis_ + _latodactylus_, _Syrrhophus_ _longipes_ + _leprus_, _Syrrhophus_ _leprus_ + _longipes_, _Batrachyla_ _longipes_ + _macrotympanum_, _Tomodactylus_ _verrucipes_ + _marnockii_, _Syrrhophus_ _marnockii_ + _modestus_, _Syrrhophus_ _modestus_ + _nebulosus_, _Syrrhophus_ _pipilans nebulosus_ + _nivocolimae_, _Syrrhophus_ _nivocolimae_ + _pallidus_, _Syrrhophus modestus_ _pallidus_ + _petrophilus_, _Syrrhophus_ _guttilatus_ + _pipilans_, _Syrrhophus_ _pipilans pipilans_ + _rubrimaculatus_, _Syrrhophus_ _rubrimaculatus_ + _smithi_, _Syrrhophus_ _guttilatus_ + _teretistes_, _Syrrhophus_ _teretistes_ + _verrucipes_, _Syrrhophus_ _verrucipes_ + _verruculatus_, _Phyllobates_ _Nomen dubium_ + ======================================================================= + + +In the course of preparing an account of the species of +_Eleutherodactylus_ occurring in México and northern Central America, it +became necessary to reëxamine the status of the genus _Syrrhophus_ and +its nominal species. It soon became evident that there were more names +than species, that some previously regarded species were geographic +variants, and that the eastern and western groups (complexes here) were +artificial divisions of the genus. I conclude that there are seven +species (one polytypic) of _Syrrhophus_ in eastern México, Texas, and El +Petén of Guatemala, and seven species (one polytypic) in western México. +The current status of each of the 23 names correctly assigned to the +genus is presented in Table 3. + +The fourteen species recognized by me are placed in five species groups. +Two of these groups are presently placed in the western complex +(_modestus_ and _pipilans_ groups) and three in the eastern complex +(_leprus_, _longipes_ and _marnockii_ groups). The two complexes do +not correspond exactly with the eastern and western groups of Smith +and Taylor (1948), Firschein (1954), and Duellman (1958) since +_S. rubrimaculatus_ is now associated with the eastern _leprus_ group. + +The definitions and contents of the five species groups are as follows: + + _leprus_ group: digital pads not or only slightly expanded, rounded + in outline; first finger longer or shorter than second; snout + acuminate or subacuminate, not rounded; outer metatarsal tubercle + conical; digits lacking distinct lateral fringes. + + content: _cystignathoides_, _leprus_ and _rubrimaculatus_. + + _longipes_ group: digital pads widely expanded, triangular in outline; + first finger shorter than second; snout acuminate; outer metatarsal + tubercle not conical; digits bearing lateral fringes. + + content: _dennisi_ and _longipes_. + + _marnockii_ group: digital pads expanded, rounded to truncate in + outline; first finger equal in length to second or slightly shorter; + snout rounded; outer metatarsal tubercle not conical; digits lacking + lateral fringes; generally stout-bodied frogs. + + content: _guttilatus_, _marnockii_, and _verrucipes_. + + _modestus_ group: digital pads expanded, truncate in outline; first and + second fingers subequal in length, first usually slightly shorter + than second; snout subacuminate; inner metatarsal tubercle twice as + large (or larger) as outer metatarsal tubercle; digits bearing + poorly-defined lateral fringes. + + content: _interorbitalis_, _modestus_, _nivocolimae_, _pallidus_, + and _teretistes_. + + _pipilans_ group: digital pads not or only slightly expanded, + truncate in outline; first finger equal in length to second; snout + subacuminate; metatarsal tubercles subequal in size; digits lacking + lateral fringes. + + content: _pipilans_. + + +_Acknowledgments._--For loan of specimens, I am indebted to Richard J. +Baldauf, Texas A & M University (TCWC); W. Frank Blair, University of +Texas (TNHC); Charles M. Bogert and Richard G. Zweifel, American Museum +of Natural History (AMNH); James E. Böhlke and Edmond V. Malnate, +Academy of Natural Sciences of Philadelphia (ANSP); Robert F. Inger and +Hymen Marx, Field Museum of Natural History (FMNH); Ernest A. Liner +(EAL); Michael Ovchynnyk, Michigan State University collection (MSU); +James A. Peters, United States National Museum (USNM); Douglas A. +Rossman, Louisiana State University Museum of Zoology (LSUMZ); Hobart M. +Smith, University of Illinois Museum of Natural History (UIMNH); Charles +F. Walker, University of Michigan Museum of Zoology (UMMZ); and John W. +Wright, Los Angeles County Museum (LACM). Specimens in the collection at +the University of Kansas Museum of Natural History are identified as KU. +The abbreviations EHT-HMS refer to the Edward H. Taylor-Hobart M. Smith +collection and FAS to the Frederick A. Shannon collection. The +type-specimens from these collections are now in the Field Museum of +Natural History and the University of Illinois Museum of Natural +History. + +I have profited from discussions concerning this problem with several +persons, most notably William E. Duellman, Hobart M. Smith, Edward H. +Taylor and Charles F. Walker. Nevertheless, the ideas and conclusions +presented here should not be construed as necessarily reflecting their +opinions. + +David M. Dennis executed all of the figures, and my wife, Marsha, typed +the manuscript. + + +_Materials and Methods._--In the course of this study, 1003 specimens +of the genus were examined. The holotypes of 21 of the 23 +nominal species are extant; I have examined 19 of these. Nine +measurements were taken, and five ratios computed for each of 338 +specimens. Females are available for all species but one; thus, +measurements were taken on individuals of both sexes. + + + + +ANALYSIS OF CHARACTERS + + +_Size and proportions._--Frogs of this genus range in size from 16 +to 40 mm. in snout-vent length. Five species are relatively small: +_S. cystignathoides_, _modestus_, _nivocolimae_, _pallidus_ and +_rubrimaculatus_; one, _S. longipes_, is relatively large, and the +remaining eight species are intermediate in size (22-30 mm.). + +Males are generally smaller than females and have proportionately longer +heads and usually larger tympani. No significant differences were found +among proportions, except that _S. longipes_ has a larger tympanum/eye +ratio than any other species. Frogs in the _Syrrhophus marnockii_ group +tend to have shorter shanks and feet, thereby giving those species a +more stocky appearance. However, the differences are not significant. + +A summary of the data on size and proportions for the frogs of the genus +_Syrrhophus_ is given in Tables 4, 5, and 6. + + +_Hands and Feet._--Taylor and Smith (1945), Smith and Taylor (1948), +Firschein (1954) and Duellman (1958) discussed the value of the palmar +tubercles in identifying frogs of this genus. The eastern complex in +general has a well-developed outer palmar tubercle (Fig. 1) in +distinction to the western complex in which the outer palmar tubercle is +reduced or absent (Fig. 2). Dixon and Webb (1966) imply that the outer +palmar tubercle is rarely absent but is usually smaller than the first +supernumerary tubercle of the fourth finger. My study of the western +species demonstrates that the outer palmar tubercle is indeed usually +present and smaller than the first supernumerary tubercle. + +Differences in interpretation of the terms "unexpanded" and "narrow," as +well as differences in techniques of preservation, have led to confusion +of the reported digital shapes in various species. Constant specific +differences are evident in the hands (Fig. 1). Except in the cases of +excessive uptake of fluids, all species have a terminal transverse +groove at the tip of each digit. Taylor (1940b) stated that _S. smithi_ +lacked grooves, but examination of the holotype reveals faint grooves at +the tops of the digits. _Syrrhophus guttilatus_, _leprus_, _pipilans_, +and _verrucipes_ lack lateral fringes on the fingers. Lateral fringes +are well developed in the _longipes_ and _modestus_ groups but poorly +defined or absent in the other members of the genus. The digital pads of +the frogs of the _longipes_ group are much broader than those of the +other species and are narrowest in the frogs of the _leprus_ group. +Supernumerary tubercles are present on the palmar surfaces of all +species of the genus. + + + TABLE 4--Size and Proportions in the Frogs of the _Syrrhophus leprus_ + Group. + + A: _cystignathoides campi_ + B: _c. cystignathoides_ + C: _leprus_ + D: _rubrimaculatus_ + + ========================================================================== + Snout-vent Tibia Head Tympanum/ Eyelid/ + length length/ width/ Eye Interorbital + Species Sex N (SVL) SVL SVL + -------------------------------------------------------------------------- + A [M] 33 16.3-23.5 41.3-49.6 34.0-40.1 43.7-66.5 43.2-89.6 + (45.8) (37.0) (56.2) (61.5) + [F] 12 16.0-25.8 41.5-51.0 33.0-38.0 42.8-60.0 48.2-69.2 + (45.8) (35.0) (51.2) (60.1) + B [M] 15 16.8-22.1 45.1-50.4 33.2-40.7 44.3-68.7 44.6-65.4 + (47.3) (37.8) (54.8) (60.0) + [F] 6 19.6-24.2 46.4-50.0 34.1-38.1 43.3-56.5 53.2-65.4 + (47.6) (36.2) (46.9) (59.2) + C [M] 14 20.6-26.4 42.3-52.3 35.0-40.3 47.5-62.5 58.2-72.5 + (46.8) (37.4) (56.5) (67.3) + [F] 15 22.1-29.2 43.4-53.3 32.6-38.9 38.6-57.9 50.2-86.9 + (47.1) (35.8) (47.1) (68.1) + D [M] 12 18.2-23.5 40.4-46.2 31.8-35.5 35.5-46.5 65.1-78.5 + (43.4) (33.8) (41.7) (71.7) + ========================================================================== + + + TABLE 5--Size and Proportions in the Frogs of the _Syrrhophus longipes_ + and _S. marnockii_ Groups. + + A: _dennisi_ + B: _longipes_ + C: _guttilatus_ + D: _marnockii_ + E: _verrucipes_ + + ========================================================================== + Snout-vent Tibia Head Tympanum/ Eyelid/ + length length/ width/ Eye Interorbital + Species Sex N (SVL) SVL SVL + -------------------------------------------------------------------------- + A [M] 16 22.8-28.4 43.9-49.7 35.3-41.2 53.9-64.2 55.3-74.0 + (47.4) (38.8) (58.9) (65.1) + [F] 10 25.9-32.0 46.3-50.8 35.6-40.3 50.6-58.7 58.1-70.9 + (48.2) (37.7) (54.9) (63.6) + B [M] 22 22.1-33.2 45.8-51.7 38.7-44.4 61.1-87.2 61.5-83.0 + (48.4) (41.8) (72.0) (72.0) + [F] 19 26.8-39.6 44.3-51.0 36.3-40.8 49.5-72.1 55.3-85.9 + (47.2) (39.1) (59.5) (67.9) + C [M] 19 20.6-29.0 41.2-48.1 36.9-44.9 55.1-75.7 53.3-79.5 + (44.5) (40.6) (64.1) (66.0) + [F] 5 25.7-31.0 41.4-46.8 35.9-42.3 47.6-61.7 62.3-79.8 + (43.6) (38.5) (54.0) (72.9) + D [M] 14 18.4-28.9 42.3-47.2 36.1-43.0 47.2-68.3 51.6-74.4 + (44.1) (39.6) (61.2) (66.3) + [F] 29 20.4-35.4 38.7-46.4 35.9-41.3 45.8-73.3 52.1-70.5 + (42.7) (38.2) (60.3) (60.7) + E [M] 29 17.5-29.2 42.7-49.5 36.2-42.4 56.1-82.2 56.8-82.8 + (46.3) (39.1) (67.8) (70.4) + [F] 6 26.5-31.7 42.4-47.7 36.0-38.1 45.8-57.8 61.0-77.9 + (44.6) (37.0) (53.9) (69.0) + ========================================================================== + + + TABLE 6--Size and Proportions in the Frogs of the _Syrrhophus pipilans_ + and _S. modestus_ Groups. + + A: _pipilans nebulosus_ + B: _pipilans pipilans_ + C: _modestus_ + D: _pallidus_ + E: _teretistes_ + F: _nivocolimae_ + G: _interorbitalis_ + + ========================================================================== + Snout-vent Tibia Head Tympanum/ Eyelid/ + length length/ width/ Eye Interorbital + Species Sex N (SVL) SVL SVL + -------------------------------------------------------------------------- + A [M] 17 22.9-28.5 38.1-42.0 34.4-37.2 36.6-47.8 56.1-82.4 + (40.0) (35.4) (43.6) (68.2) + [F] 3 21.1-22.7 42.1-44.5 33.2-35.8 36.6-47.6 64.3-65.4 + + B [M] 18 22.6-27.8 37.9-44.0 32.2-36.5 38.0-54.0 56.1-79.5 + (41.4) (33.0) (46.2) (67.3) + [F] 1 29.4 38.4 32.5 44.6 55.0 + + C [M] 8 15.8-20.1 38.5-42.6 32.1-38.1 26.8-39.3 57.0-86.9 + (40.6) (34.2) (31.5) (69.1) + [F] 1 18.5 44.2 36.0 24.0 52.1 + + D [M] 6 17.9-19.3 41.0-44.9 32.6-36.2 27.0-35.6 59.4-67.7 + (43.4) (35.2) (30.9) (65.2) + + E [M] 18 19.2-23.2 41.5-45.3 32.5-36.4 28.6-43.8 51.2-75.0 + (43.7) (34.0) (33.7) (62.2) + [F] 1 24.8 41.8 30.8 37.9 60.5 + + F [M] 15 18.9-21.1 42.2-48.6 30.9-37.1 30.0-39.3 42.6-69.1 + (45.0) (33.7) (34.7) (55.0) + [F] 1 24.1 40.9 33.5 27.6 56.5 + + G [M] 1 25.6 43.0 ---- 39.4 57.6 + [F] 9 20.2-26.7 39.9-47.1 32.6-39.3 29.1-41.2 58.2-76.9 + (43.2) (35.8) (36.4) (69.2) + ========================================================================== + + + [Illustration: FIG. 1: Palmar views of hands of six species of the + eastern complex of _Syrrhophus_. (A) _verrucipes_ (UIMNH 15995), + (B) _rubrimaculatus_ (KU 58911), (C) _dennisi_ sp. nov. (holotype, + UMMZ 101121), (D) _guttilatus_ (UIMNH 55520), (E) _marnockii_ + (TCWC 4782), and (F) _longipes_ (TCWC 12179). All ×6.5.] + + + [Illustration: FIG. 2: Palmar views of hands of two species of the + western complex of _Syrrhophus_. _pipilans_ (left, KU 58908, ×6) + and _teretistes_ (center, KU 75269, and right, KU 75263, + respectively, ×9).] + + +In _S. cystignathoides_ and _leprus_, the first finger is longer than +the second, and the first two fingers are equal in length in +_guttilatus_ and _marnockii_. In the other species the first finger is +shorter than the second. + +Supernumerary tubercles are well developed on the plantar surfaces in +all species, except _S. guttilatus_, in which they are poorly defined +(Fig. 3). The relative sizes of the metatarsal tubercles has been used +in the classification of the species and species groups of _Syrrhophus_. +The metatarsal tubercles are similar in all species of the eastern +complex (including _rubrimaculatus_); the outer tubercle is always about +one-half the size of the ovoid inner metatarsal tubercle. In the +_leprus_ group the outer tubercle is conical and compressed. The +metatarsal tubercles of _pipilans_ are about the same size, or the outer +is slightly smaller than the inner. In the _modestus_ group the outer +metatarsal tubercle is about one-third the size of the inner. + +All species, except _guttilatus_, have well-defined to poorly defined +lateral fringes on the toes. All species have expanded toe pads. The +fifth toe is usually shorter than the third, but the second is equal +in length to the fifth in some specimens of _S. cystignathoides_ and +_S. marnockii_. _Syrrhophus nivocolimae_ is the only species with +tubercles along the outer edge of the tarsus; this is merely a +reflection of the highly tuberculate nature of the skin in this species. + + +_Skin texture._--The skin of the dorsum is smooth or very weakly +pustular in all species of the genus except _nivocolimae_ and +_verrucipes_. The dorsal surfaces of _nivocolimae_ are warty; in +_verrucipes_ the skin is pustular. The skin of the venter is areolate in +_cystignathoides cystignathoides_, _dennisi_ and _verrucipes_ but is +smooth in all other species of the genus. + + [Illustration: FIG. 3: Plantar views of feet of four species of the + eastern complex of _Syrrhophus_. (A) _guttilatus_ (UIMNH 55519, ×6), + (B) _leprus_ (UIMNH 42726, ×6), (C) _verrucipes_ (UIMNH 15995, ×6), + and (D) _longipes_ (TCWC 12179, ×4.6).] + + +_Color pattern._--As is evident in the diagnoses, the color patterns of +given populations have been regarded as useful in separating the +species and subspecies. Duellman (1958) suggested that the coloration, +with the exception of _modestus_, was a dark ground color with pale +markings. It is a moot point whether the frogs have light spots on a +dark background or have a light background with an extensive reticulate +dark pattern. The venters are gray or white, and the vocal sac is nearly +black in some species. Interorbital dark bars or triangles are absent in +only two species of the eastern complex, _cystignathoides campi_ and +_marnockii_; the latter lacks a supratympanic stripe, which is present +in the other members of the eastern complex. _Syrrhophus interorbitalis_ +and _nivocolimae_ have light interorbital bars; these bars occur in only +one other population of the genus (_S. c. cystignathoides_). Bars on the +thighs are ill defined or absent in the members of the _marnockii_ and +part of the _modestus_ groups. The color in life is noted in the species +accounts. + + +_Voice._--The voices of all _Syrrhophus_ can be described as a +single short chirp or peep; without audiospectrographic analyses +the significance of the differences between a chirp, peep, or short +whistle cannot be appreciated. Martin (1958) and Wright and +Wright (1949) reported multi-noted calls, and one collector of +_S. verrucipes_ noted the frog "trilled." + +Fouquette (1960) presented analyses of two species (_marnockii_ +and _pipilans nebulosus_). The voices were very similar; both frogs +were reported to "trill" and "chirp." + + + + +SYSTEMATIC ACCOUNT + + +The genus _Syrrhophus_ has been defined (Lynch, 1968) and limited to the +group of species occurring in Guatemala, México and the United States. +The closest relatives of _Syrrhophus_ are the frogs of the genus +_Tomodactylus_ (Dixon, 1957; Firschein, 1954). Lynch (1968) +implied there were no osteological bases for the separation of +_Eleutherodactylus_, _Syrrhophus_, and _Tomodactylus_. At that time, I +believed such to be the case and derived _Syrrhophus_ and _Tomodactylus_ +from the _rhodopis_ complex of _Eleutherodactylus_, with which they +share terrestrial habits and relatively short limbs. In the _rhodopis_ +complex there is a tendency for the loss of the outer palmar tubercle, a +not uncommon condition in _Syrrhophus_ and _Tomodactylus_. + +However, the skulls of _Syrrhophus_ and _Tomodactylus_ show departures +from the pattern observed in the Middle American _Eleutherodactylus_, as +well as many of those species in western South America. Baldauf and +Tanzer (1965) reported that the frontoparietals and prootics were fused +in _Syrrhophus marnockii_ and that the prootics and exoccipitals +appeared to be one bone (otoccipital). The otoccipital is not uncommon +in eleutherodactyline frogs, but the fusion of the frontoparietals with +the prootics (regardless of the fusion of the latter with the +exoccipital) is uncommon in the family. I have found the +frontoparietal-prootic fusion only in _Syrrhophus_ (all species), +_Tomodactylus_ (all species), and _Eleutherodactylus_ (West Indies +species). None of the Middle American _Eleutherodactylus_ has the two +bones fused. Examination of the character is difficult in dried skeletal +preparations. Cleared and stained or macerated preparations are +satisfactory for checking this character. + +Thus, in addition to the presence of numerous plantar supernumerary +tubercles in the frogs of the genera _Syrrhophus_ and _Tomodactylus_, +these two genera can be separated from other Middle American +eleutherodactylines by the fusion of the frontoparietals and prootics. +This character not only further strengthens the argument that the two +genera are closely related but poses a problem of zoogeographic analysis +of the distribution of the character, which will be discussed fully +elsewhere. + + +Key to the Species of the Frog Genus _Syrrhophus_ + + 1. Three large, well-developed palmar tubercles 2 + + Two large palmar tubercles; outer (third) palmar tubercle reduced + in size or absent 9 + + 2. Digital pads more than twice (usually three or more) times width + of digit 3 + + Digital pads less than twice width of digit 4 + + 3. Males having vocal slits; dorsum vermiculate; diameter of + tympanum in males about one-half diameter of eye _S. dennisi_ + + Males lacking vocal slits; dorsum flecked, spotted, or + blotched; diameter of tympanum in male about three-fourths + that of eye _S. longipes_ + + 4. First finger longer than second 5 + + First finger shorter than or equal to second 7 + + 5. Venter smooth; dorsum spotted or vermiculate _S. leprus_ + + Venter areolate, or if smooth, dorsum flecked and interorbital + bar lacking 6 + + 6. Venter areolate; interorbital bar present; ground color + yellowish _S. cystignathoides cystignathoides_ + + Venter smooth; interorbital bar absent; ground color + brown _S. cystignathoides campi_ + + 7. First finger shorter than second; digital tips only slightly + dilated; green in life with darker green spots _S. verrucipes_ + + First finger equal to second; digital tips slightly to moderately + expanded 8 + + 8. Dorsum vermiculate; interorbital bar present; ground color + cream to brown in life _S. guttilatus_ + + Dorsum punctate or flecked; interorbital bar absent; + ground color green in life _S. marnockii_ + + 9. Dorsum dark with pale (red in life) spots; digital pads + not expanded _S. rubrimaculatus_ + + Dorsum pale with dark markings and digital pads slightly to widely + expanded 10 + + 10. Digital tips not widely expanded; tympanum well-defined; + outer metatarsal tubercle more than one-half size of inner 11 + + Digital tips widely expanded, truncate in outline; tympanum + poorly defined; outer metatarsal tubercle less than one-half + size of inner 12 + + 11. Dorsum dark brown with large light spots or blotches; tympanum/eye + ratio usually greater than 43 percent _S. pipilans pipilans_ + + Dorsum dark brown with small light spots; tympanum/eye + ratio less than 48 percent _S. pipilans nebulosus_ + + 12. Light interorbital bar present 13 + + Light interorbital bar absent 14 + + 13. Adults small, less than 22 mm. snout-vent length with a + broad mid-dorsal stripe; dark bands on shank narrower than + light interspaces _S. nivocolimae_ + + Adults larger, more than 22 mm. snout-vent length; dorsum + vermiculate; dark bands on shank broader than light + interspaces _S. interorbitalis_ + + 14. Dorsum spotted with discrete black spots; pattern + definite _S. modestus_ + + Dorsum reticulate or vermiculate, pattern poorly defined 15 + + 15. Adults small, less than 21 mm. snout-vent length; upper arm + not banded _S. pallidus_ + + Adults larger, usually greater than 21 mm. snout-vent length; + upper arm banded _S. teretistes_ + + + + +SPECIES ACCOUNTS + + +The following accounts do not include complete descriptions of each +taxon, because a more than adequate number of descriptions is available +in the recent (1940-1966) literature. An abbreviated synonymy, in which +are listed all combinations and emendations of names and significant +contributions to our knowledge of the taxon, is given for each. For each +species and subspecies the following are given: descriptive diagnosis, +statement of range, remarks on taxonomy, list of specimens examined, +illustration of color pattern, and distribution map. + + +=Syrrhophus cystignathoides= (Cope) + + _Phyllobates cystignathoides_ Cope, 1877:89-90 + [Syntypes.--Originally USNM 32402-32409, (32405 now in MCZ) + from Potrero, near Córdoba, Veracruz, México, Francis Sumichrast + collector.] + +_Diagnosis._--Adults small, males 16.0 to 23.5 mm. in snout-vent length, +females 16.0-25.8 mm. in snout-vent length; vocal slits present in +males; finger tips slightly expanded; first finger longer than second; +outer metatarsal tubercle one-half size of inner, conical, compressed; +skin of dorsum weakly pustular, that of venter smooth to areolate; +tympanum 44 to 69 per cent diameter of eye (mean 55.5 per cent); ground +color yellow to brown in life with brown to black fleckings on dorsum +and flanks; limbs banded; interorbital bar present or not. + +_Remarks._--Two geographic races (subspecies) are herein recognized; +previously these were held by various authors to be species (_campi_ and +_cystignathoides_). Intergradation occurs in southern Tamaulipas and +eastern San Luis Potosí, México. The two subspecies can be distinguished +on the basis of color pattern and the condition of the skin of the +venter. + +_Distribution._--Low to moderate elevations from the Río Grande +embayment to central Veracruz, México (Fig. 5). + + +=Syrrhophus cystignathoides campi= Stejneger, New combination + + _Syrrhophus campi_ Stejneger, 1915:131-32. [Holotype.--USNM 52290, + from Brownsville, Cameron Co., Texas; R. D. Camp collector, + March 31, 1915]. Smith and Taylor, 1948:52. Martin, 1958:50. + +_Diagnosis._--Venter smooth; usually no interorbital light and dark bars +present; ground color brown in life (Fig. 4a). + +_Remarks._--Martin (1958) was the first author to point out +that _S. campi_ was probably a subspecies of the more southern +_S. cystignathoides_. Various references in the literature might lead +one to believe that the two were sympatric over much of northeastern +México; this error was created by the use of a single character +(condition of the skin of the venter) to characterize the two +populations. Specimens from southern Texas have a smooth venter, lack +interorbital bars and have, in general, a brown ground color, whereas +specimens from central Veracruz have an areolate venter, interorbital +light and dark bars and a yellow ground color. In southern Tamaulipas +and eastern San Luis Potosí, these characters vary discordantly, thereby +strongly suggesting that the two populations intergrade. Both +populations agree in other morphological characters; therefore, they are +here treated as geographic variants. + +_Etymology._--Named for the collector of the type specimens, Mr. R. D. +Camp of Brownsville, Texas. + +_Distribution._--Lower Río Grande embayment in Texas to central Nuevo +León and Tamaulipas, México. Intergrades are known from southern +Tamaulipas and adjacent San Luis Potosí, México (Fig. 5). + +_Specimens examined._--(113) TEXAS, Cameron Co.: MCZ 10277-85, 10286 +(10); Brownsville, AMNH 3215, 3218-20, 3221 (3), 5376, 62117, FMNH +105336, KU 8135-39, MCZ 3738-42, 3743 (10), TCWC 5908, 7139, TNHC 92-94, +20909, UMMZ 51760, 54031 (5), USNM 52290 (holotype); 22 mi. SE +Brownsville, TNMC 14223; 8 mi. SW Brownsville, UMMZ 101127 (3); +Harlingen, AMNH 62118, UMMZ 105200-205, 105206 (5), 105207 (4). _Hidalgo +Co._: Bentsen-Río Grande State Park, UMMZ 114378; 6 mi. S McAllen, TNHC +7136-39; Santa Ana Refuge, TCWC 13495-96; Weslaco, TCWC 17658-60. + +MEXICO, _Nuevo León_: Salto Cola de Caballo, AMNH 57953-54, FMNH +30644-45, 37169-70; Monterrey, UIMNH 13324; 40 km. SE Monterrey, UIMNH +3686. _Tamaulipas_: 80 km. Matamoros, FMNH 27150 (13). + +Intergrades [_S. c. cystignathoides_ × _S. c. campi_ (88)] MÉXICO, _San +Luis Potosí_: 5 km. E Ciudad del Maiz, UMMZ 106435; 16 km. W Naranjo, +FMNH 104584; Salto de Agua, 34 km. WSW Antigua Morelos, TCWC 6980. +_Tamaulipas_: 5 km. W Acuña, 1060 m., UMMZ 101172, 101173 (16), +101174-76, 101177 (6); 14.5 km. NNW Chamal, 430 m., UMMZ 111337 (2); 20 +km. NNW Chamal, 700 m., UMMZ 111338 (11); 8 km. N Gómez Farías, 450 m., +UMMZ 101165; 8 km. NE Gómez Farías, Pano Ayuctle, UMMZ 102264, 102924 +(6); 8 km. NW Gómez Farías, 1060 m., LSUMZ 11084, UMMZ 101199, 102928 +(5), 102929-32, 110124 (3); Río Guayala, near Magiscatzin, MCZ 24138-42, +85071-81, UMMZ 88242 (2); Magiscatzin, TCWC 6981; Las Yucas, north of +Aldama, MCZ 29665-68; 16 km. NE Zamorina, UMMZ 101124. + + [Illustration: FIG. 4: _Syrrhophus cystignathoides campi_ + (left, TCWC 13490) and _S. c. cystignathoides_ (right, KU 105500). + Dorsal views ×2, sides of heads ×3.] + + +=Syrrhophus cystignathoides cystignathoides= (Cope), New combination + + _Phyllobates cystignathoides_ Cope, 1877:89-90 [Syntypes.--USNM + 32402-32409, from Potrero, near Córdoba, Veracruz, México, + collected by Francis Sumichrast]. Boulenger, 1882:196. + + _Syrrhophus cystignathoides_: Cope, 1879:268. Kellogg, 1932: + 126-27. Taylor and Smith, 1945: 582-83. Smith and Taylor, 1948:50. + Martin, 1958:49. + + _Syrrhaphus cystignathoides_: Günther, 1900:218. + + _Syrraphus cystignathoides_: Díaz de León, 1904:10. + + _Syrrhopus cystignathoides_: Barbour and Loveridge, 1946:170. + + [Illustration: FIG. 5: Distribution of _Syrrhophus cystignathoides + campi_ (solid symbols) and the nominate subspecies (open symbols).] + +_Diagnosis._--Venter areolate; interorbital light and dark bars present; +ground color yellow to brownish-yellow in life (Fig. 4b). + +_Remarks._--Firschein (1954) briefly considered the status of Peters' +(1871) _Phyllobates verruculatus_ and noted that if it was a +_Syrrhophus_ it would probably be referrable to _S. cystignathoides_. +Peters' (1871) original description corresponds well with +_S. cystignathoides_, and the type-locality ("Huanusco" = Huatusco) is +within the range of that species. Firschein (1954) expressed doubt that +_verruculatus_ was a _Syrrhophus_, because Peters placed it in another +genus. However, Peters described _verruculatus_ a decade before Cope +diagnosed the genus Syrrhophus. Most frogs now called _Syrrhophus_, plus +a number of lower Central American frogs now placed in a variety of +genera were placed in _Phyllobates_ by Boulenger, Cope, and Peters. + +The types of _Phyllobates verruculatus_ were destroyed during World War +II (Günther Peters, _in litt._); the specimens subsequently assigned to +the taxon by Kellogg (1932) are _Syrrhophus cystignathoides_. Because +the type specimens are lost and because the name antedates the more +established name, _cystignathoides_, I favor retaining _Phyllobates +verruculatus_ Peters as a _nomen dubium_. + +Smith and Taylor (1948) reported _S. verruculatus_ from Tianguistengo, +Hidalgo, México. These specimens are examples of _verrucipes_. Smith +(1947) reported a specimen of _verruculatus_ from San Lorenzo, Veracruz. +Firschein (1954) referred it to _cystignathoides_, and Duellman (1960) +concluded that both authors were in error and that the specimen (USNM +123530) was a _leprus_. + +_Etymology._--The trivial name is the diminutive of _Cystignathus_, a +once-used generic name for several leptodactylid frogs. + +_Distribution._--Low and moderate elevations in the foothills along the +Sierra Madre Oriental from eastern San Luis Potosí to Central Veracruz, +México (Fig. 5). + +_Specimens examined._--(130), MÉXICO, _Puebla_: Necaxa, UMMZ 69519-20. +_San Luis Potosí_: 5 km. W Aguismón, LSUMZ 4962-63; along Río Axtla, +road to Xilitla, UMMZ 105500; Tamazunchale, UIMNH 3199; 6.5 km. N +Tamazunchale, UMMZ 104039; 8 km. N Tamazunchale, UMMZ 119490. +_Veracruz_: Coatepec, 1210 m., FMNH 704966-67; 11 km. SE Coatepec, 850 +m., FMNH 70468-70; below Córdoba, FMNH 104588, UIMNH 13321; Cuautlapam, +1000 m., FMNH 106477-80, KU 100364, UIMNH 58200-03, UMMZ 105392; Fortín +de las Flores, UIMNH 13322, 13339; 1.6 km. N Fortín de las Flores, UIMNH +42799-808, UMMZ 105389; 3.2 km. N Fortín de las Flores, UIMNH 26633-35; +4.8 km. N Fortín de las Flores, UIMNH 71967-68; 3.2 km. W Fortín de las +Flores (Barranca Metlac), 910 m., UIMNH 49294-95, UMMZ 115444-46, +118221, 119893 (2); Huatusco, KU 100363; Jalapa, 1400 m., FMNH 70440, +70443-51, 70454-65; 16 km. NE Jalapa, 1300 m., FMNH 70452-53; 8 km. E +Jalapa, UIMNH 13338; 9.5 km. S Jalapa, UMMZ 122083 (2); Mirador, KU +23967; Paraja Nuevo, El Suchil, UMMZ 85490 (7), 85491 (2), 90315; La +Passa, UIMNH 49293, 49297; 1 km. E Plan del Río, 240 m., UMMZ 102067 +(2); Potrero Viejo, FMNH 104583, 104586, 105326-27, KU 26789, 100357-62, +UIMNH 13323, 13340-43; USNM 32402 (lectotype), 32403-04, 32406-09; 9.6 +km. S Santa Rosa, TCWC 12785; 24 km. NE Tezuitlán (Puebla), UMMZ 105388; +Teocelo, FMNH 70437-38, KU 26080, 26790; 3.2 km. N Teocelo, FMNH 70439, +70441-42; 9.6 km. NW Tihuatlán, UIMNH 3684-85; 15 km. ENE Tlacotepec, KU +23966; 26 km. NW Tuxpan, UMMZ 126419. + + +=Syrrhophus leprus= Cope + + _Syrrhophus leprus_ Cope, 1879:268-69 [Holotype.--USNM 10040, from + Santa Efigena, Oaxaca, México, Francis Sumichrast collector]. + Kellogg, 1932:124-5, 128. Taylor and Smith, 1945:582. Smith and + Taylor, 1948:50-51. Duellman, 1958:8, pl. 1, Fig. 2; 1960:56-57. + Gorham, 1966:165. + + _Syrrhaphus leprus_: Günther, 1900:217. + + _Syrrhophus leprus leprus_: Neill, 1965:85-86. + + _Syrrhophus leprus cholorum_ Neill, 1965:85-86 [Holotype.--Wilfred + T. Neill collection 1525, from 3.9 mi. N San Antonio, Toledo + District, British Honduras, collected October 28, 1959, by + R. A. Allen, T. C. Allen, and W. T. Neill]. + +_Diagnosis._--Medium-sized frogs, males 20.5-26.5 mm. in snout-vent, +females 22.0-29.3 mm. in snout-vent length; vocal slits present in +males; tips of fingers dilated slightly; first finger longer than +second; inner metatarsal tubercle twice size of small, conical outer +metatarsal tubercle; skin of dorsum pustular, that of venter smooth; +snout subacuminate; diameter of tympanum 47.5-62.5 per cent of eye in +males, 38.6-57.9 per cent in females; dorsum yellowish-green with +chocolate brown blotches or spots forming reticulations in most +specimens; venter white to gray; flanks brown, spotted with white or +not; limbs banded; interorbital bar obscured by dorsal pattern. + + [Illustration: FIG. 6: Dorsal views of _Syrrhophus leprus_ showing + variation in dorsal pattern (left, UMMZ 121244, ×2; right, KU 26106, + ×1.7). Side of head (UIMNH 42726, ×7).] + + [Illustration: FIG. 7: Distribution of three species of eastern + complex _Syrrhophus_: _leprus_ (circles), _rubrimaculatus_ + (triangles), and _verrucipes_ (squares).] + +_Remarks._--My distribution map (Fig. 7) differs somewhat from that of +Duellman (1958), who was unaware of specimens reported by Taylor and +Smith (1945) from central Veracruz, México. + +Duellman (1958, 1960) regarded _S. leprus_ as having a gray venter. +Neill (1965) characterized his new subspecies on the basis of white +venter and spots on the dorsum. Some specimens from throughout the range +have only small round spots, instead of vermiculations (Fig. 6). The +gray ventral coloration is largely restricted to the population in Los +Tuxtlas, Veracruz, but only about 80 per cent of the specimens from the +Los Tuxtlas have gray venters, whereas specimens from Guatemala, Oaxaca, +Tabasco, and central Veracruz, México, have white venters (rarely gray). +Since the specimens from British Honduras are not distinct from +specimens throughout most of the range, there is no reason to recognize +them as a subspecies. + +_Etymology._--Greek, _lepra_, leprosy, in reference to the mottled color +pattern. + +_Distribution._--Discontinuous; central Veracruz to British Honduras to +low elevations in the foothills of the Sierra Madre Oriental, Los +Tuxtlas, Sierra Madre de Chiapas (Isthmus of Tehuantepec (Fig. 7)). + +_Specimens examined._--(84). GUATEMALA, _Alta Verapaz_: Chinajá, KU +55961-62. _El Petén_: 15 km. NW Chinajá, KU 55963; Piedras Negras, USNM +114085-92; Tikal, UMMZ 117035; Uaxactún, AMNH 55121-22. + +MÉXICO, _Oaxaca_: Cerro San Pedro del Isthmo, UIMNH 35510; Finca La +Gloria, USNM 114093; 30.5 km. N Matías Romero, UIMNH 39459, 71969; Santa +Efigenia, USNM 10040 (holotype). _Tabasco_: Teapa, UMMZ 113799-800; 13.5 +km. W Teapa, UMMZ 120253. _Veracruz_: 27.5 km. N Acayucan, UIMNH 42726; +Atoyac, UIMNH 13331, 49296; 3.2 km. N Catemaco, UIMNH 71976-77; Coyame, +UIMNH 38995, 38998, 40342; Dos Amates, TCWC 21211; Fortín de Las Flores, +FMNH 113751, 113753; Paraja Nuevo, El Suchil, UMMZ 90315; Potrero Viejo, +FMNH 113743-50, 126114-18, KU 26104-06, UIMNH 13332-37, UMMZ 88837; San +Andrés Tuxtla, UIMNH 27123-31, 28611, 71975, UMMZ 115450 (5); San +Lorenzo, USNM 123530; 4.5 km. NW Santiago Tuxtla, JDL 992 (skeleton), +UIMNH 27122; 32 km. S Sayula, EAL 1696; Tepalapan, 1.6 km. S Catemaco, +UMMZ 118222 (2); Volcán San Martín, south slope, UMMZ 118223; Volcán San +Martín, Rancho El Tular, UIMNH 35399-400, 40340-41. + + +=Syrrhophus rubrimaculatus= Taylor and Smith + + _Syrrhophus rubrimaculatus_ Taylor and Smith, 1945:583-85 + [Holotype.--USNM 114070, from La Esperanza, near Escuintla, + Chiapas, México, collected May 13, 1940, by H. M. and R. Smith]. + Duellman, 1958:1-4, 7, 12, 14. Gorham, 1966:167. + + _Syrrhophus rubrimaculata_: Smith and Taylor, 1948:48-49. + + [Illustration: FIG. 8: _Syrrhophus rubrimaculatus_ (upper right, + KU 58911, ×1.6; lower right, KU 58910, ×4) and _S. verrucipes_ + (upper left, UIMNH 15995, ×1.6; lower left, UIMNH 15989, ×3.7).] + +_Diagnosis._--Small frogs, males 18.2-23.5 mm. snout-vent, females +19.0-22.5 mm. snout-vent length (small sample); vocal slits in males; +digital tips scarcely expanded (Fig. 1); first finger shorter than +second; outer palmar tubercle reduced in size; inner metatarsal tubercle +elongate, twice the size of small, conical outer metatarsal tubercle; +diameter of tympanum 35.5-46.5 per cent that of eye in both sexes; +dorsum brown with small pale spots (red in life); venter gray. + +_Remarks._--Previous authors who treated _Syrrhophus_ placed this +species in the western complex, because it occurs on the Pacific versant +and has a reduced outer palmar tubercle. Duellman (1958) placed +_rubrimaculatus_ apart from the other western species, because of its +relatively unexpanded digital tips and coloration. The digital tips are +like those in _leprus_, which _rubrimaculatus_ resembles. Except for the +reduction of the outer palmar tubercle, _rubrimaculatus_ could be a +member of the _leprus_ group. + +_Syrrhophus rubrimaculatus_ is probably best treated as a Pacific +derivative of the _leprus_ group, even though the palmar tubercles do +not agree. The removal of _rubrimaculatus_ from the western complex +results in a more homogeneous remainder and does not greatly increase +the heterogeneity of the eastern complex. + +_Etymology._--Latin, meaning spotted with red; in reference to the +colors in life. + +_Distribution._--Low to moderate elevations on the Pacific versant of +southeastern Chiapas, México (Fig. 7); probably extending into +adjacent Guatemala. + +_Specimens examined._--(48) MÉXICO, _Chiapas_: Escuintla, UMMZ 88283; 6 +km. NE Escuintla, UMMZ 87876-80; La Esperanza, UIMNH 13285, UMMZ +88496-97, USNM 114070 (holotype), 114054-69, 114072; Monte Cristo, UMMZ +88353; 1.3 km. N Puerto Madero, KU 58910-11; Finca San Jerónimo, 600-650 +m., UIMNH 55299-312, 55313-16 (cleared and stained). + + +=Syrrhophus guttilatus= (Cope) + + _Malachylodes guttilatus_ Cope, 1879:264 [Holotype.--USNM 9888, + from Guanajuato, Guanajuato, México; collected in 1877 by + Alfredo Duges]. + + _Syrrhopus guttulatus_: Boulenger, 1888:204-06. + + _Syrrhaphus guttulatus_: Günther, 1900:317. + + _Syrraphus guttulatus_: Díaz de León, 1904:11. + + _Syrrhophus guttilatus_: Nieden, 1923:399-400. Kellogg, 1932:125, + 127-28. Smith and Taylor, 1948:49, 51. Firschein, 1954:52-54. + Gorham, 1966:164. + + _Syrrhophus smithi_ Taylor, 1940b:43-45, pl. 1 [Holotype.--USNM + 108594, from 15 mi. SW Galeana, Nuevo León, México, 1575 m.; + collected on October 13, 1939, by Hobart M. Smith]. Smith and + Taylor, 1948:49, 51. Firschein, 1954:54-55. Martin, 1958:50. + Gorham, 1966:167. + + _Syrrhophus gaigeae_ Schmidt and Smith, 1944:80 [Holotype.--FMNH + 27361, from the Basin, Chisos Mountains, Brewster Co., Texas; + collected on July 24, 1937, by Walter L. Necker]. + + _Syrrhophus petrophilus_ Firschein, 1954:50-52 [Holotype.--UIMNH + 7807, from 5 km. SW San Luis Potosí, San Luis Potosí, México; + collected on July 18, 1949, by David Langebartel]. Gorham, + 1966:166. + + _Syrrhophus marnocki_: Milstead, Mecham, and McClintock, 1950:548 + (in part). + +_Diagnosis._--Medium-sized frogs, males 20.6-29.0 mm. snout-vent, +females 25.7-31.0 mm. snout-vent length; vocal slits in males; digital +tips slightly expanded (Fig. 1); first and second fingers equal; skin of +dorsum smooth to moderately pustular, that of venter smooth; snout +blunt; diameter of tympanum 55.1-75.7 per cent that of eye in males, +47.6-61.7 in females; dorsum and flanks cream to gray with light brown +to black flecking and vermiculations; thighs usually not banded; +interorbital bar present (Fig. 8). + + [Illustration: FIG. 9: _Syrrhophus guttilatus_ (upper left, UIMNH + 55519, ×1.4; lower left, UIMNH 55519, ×2.3) and _S. marnockii_ + (upper right, TCWC 9317, ×1.4; lower right, TCWC 13510, ×2.1).] + +_Remarks._--Cope (1879) distinguished _Malachylodes_ from _Syrrhophus_ +on the basis of the presence of a frontoparietal fontanelle in the +holotype of _guttilatus_. The holotype is a juvenile female and as is +the case in the juveniles of nearly all leptodactylids, a frontoparietal +fontanelle is present. Firschein (1954) used the presence of the +fontanelle to distinguish _guttilatus_ from his _petrophilus_. + +As is clearly evident from the length of the synonymy, I consider a +number of currently used names to be synonymous with _guttilatus_. I +have seen the holotypes of all four names and am unable to recognize +more than a single species. The holotype of _petrophilus_ is a male, +whereas that of _smithi_ is a female. The supposed differences are a +reflection of sexual dimorphism in the size of the eye (Table 5). The +two holotypes, as well as those of _gaigeae_ and _Malachylodes +guttilatus_ agree in color pattern. + +Schmidt and Smith (1944) named _Syrrhophus gaigeae_ from the Chisos +Mountains of the Big Bend region of Texas and compared it only with +_S. marnockii_. Milstead, Mecham and McClintock (1950) synonymized +_gaigeae_ and _marnockii_ because they were unable to verify the +characters Wright and Wright (1949) used to separate them. Specimens +from the Big Bend region differ from those of the Edward and Stockton +Plateaus in having a vermiculate pattern, an interorbital bar, and a +supratympanic stripe. In these respects they agree with specimens from +northern México. Based on limited observations, the Mexican population +is yellowish to brownish in life whereas the central Texas population is +green in life. Lacking evidence of genetic exchange, the two are held to +be specifically distinct. + +Nearly every specimen examined was infested with chiggers of the genus +_Hannemania_. The greatest concentrations are on the venter, in the +groin, and on the thighs. Many specimens have chiggers on the digits and +tarsi. The same, or a related, chigger was found on many specimens of +_Syrrhophus marnockii_ and a few _S. verrucipes_, but on no other +species of the genus. Mr. Willy Wrenn told me that he has seen heavy +infestations of _Hannemania_ on _Syrrhophus pallidus_. Infestation by +_Hannemania_ probably reflects similar ecologies rather than close +relationships. + + [Illustration: FIG. 10: Distribution of _Syrrhophus guttilatus_.] + +_Etymology._--Latin, _guttula_, meaning spotting or flecking, in +reference to the color pattern. + +_Distribution._--Moderate to intermediate elevations (600 to 2000 m.) +along the Sierra Madre Oriental from the Big Bend Region of Texas to +Guanajuato, México (Fig. 10). + +_Specimens examined._--(32) TEXAS, _Brewster Co._: Juniper Canyon, +Chisos Mts., FMNH 27361 (holotype of _S. gaigeae_), 27360, 27362-63, MCZ +15346, 27801, UMMZ 66080, 66082, 66085-91, USNM 76876; Upper Green +Gulch, TCWC 15943. + +MÉXICO: _Coahuila_: 8 km. S Saltillo, UIMNH 55518-21. _Guanajuato_: +Guanajuato, USNM 9888 (holotype of _Malachulodes guttilatus_); 8 km. E +Guanajuato, AMNH 73425; Cerro Cubilete, AMNH 73424. _Nuevo León_: 3 km. +S Galeana, JDL 1215 (skeleton), UIMNH 58204; 24 km. SW Galeana. 1575 m., +USNM 108594 (holotype of _Syrrhophus smithi_). _San Luis Potosí_: 5 km. +SW San Luis Potosí, UIMNH 7807 (holotype of _S. petrophilus_). +_Tamaulipas_: 1.6 km. NW La Joya de Salas, 1530 m., UMMZ 110736 (4). + + +=Syrrhophus marnockii= Cope + + _Syrrhophus marnockii_ Cope, 1878:253 [Syntypes.--ANSP 10765-68, + from "near San Antonio," Bexar Co., Texas; collected by G. W. + Marnock]. + + _Syrrhophus marnocki_: Yarrow, 1882:24, 193. Milstead, Mecham, + and McClintock, 1950:550. + +_Diagnosis._--Medium-sized frogs, males 18.4-28.9 mm. snout-vent, +females 20.4-35.4 mm. snout-vent length; vocal slits in males; digital +tips widened (Fig. 1); first and second fingers equal; skin of dorsum +smooth to weakly pustular, that of venter smooth; snout blunt, rounded; +diameter of tympanum 47.2-68.3 per cent that of eye in males, 45.8-73.3 +in females; dorsum tan to light brown in preservative with rusty-brown +flecks, venter white; ground color green in life; thighs banded; +interorbital bar absent. + +_Remarks._--Specimens from the southern edge of the Edwards Plateau and +the eastern edge of the Stockton Plateau have larger flecks on the back +that tend to form a vermiculate pattern like that of _S. guttilatus_. +The vermiculation is never well developed (see plate 38 in Conant, +1958). Most of the specimens from the Edwards Plateau have a punctate +pattern (Fig. 9). + +Fossils are known from the Sangamon interglacial deposits in Foard and +Knox Counties, Texas (Lynch, 1964; Tihen, 1960). + +_Etymology._--A patronym for the collector of the type specimens. + +_Distribution._--The Edwards Plateau and the extreme eastern edge of the +Stockton Plateau in Texas (Fig. 11). The fossil records lie some 200 +miles to the north. Two specimens (FMNH 103216-17) from Brownsville, +Cameron Co., Texas, were formerly in the EHT-HMS collection (nos. +31348-49). Data given in Taylor's field catalogue (housed in the +Division of Reptiles, Field Museum) are "Brownsville, A. J. Kirn +collector, April 15, 1934." Until verification by recently collected +material is available, this record must be disregarded. + +_Specimens examined._--(103) TEXAS, _Bandera Co._: 10 mi. SW Medina, +TCWC 13508-10; 8 mi. W Medina, KU 60243; 13 mi. W Medina, KU 60242, TCWC +13506-07. _Bexar Co._: UIMNH 34694; Classen ranch, near San Antonio. +UMMZ 98891; Helotes, EAL 1560, MCZ 11837 (2), UMMZ 64045, USNM 13635; 2 +mi. N Helotes, TCWC 9234-35; 3.5 mi. N Helotes, LSUMZ 10363; 8 mi. N +Helotes, TCWC 1549, 4364; San Antonio, FMNH 15553-56, TCWC 13497-99. +_Blanco Co._: 8 mi. NE Blanco, TCWC 4782. _Comal Co._: New Braunfels, +TCWC 13500-05; 5 mi. NE New Braunfels, UMMZ 71016 (10). _Hays Co._: San +Marcos, AMNH 22661-64, 32700, FMNH 15245-46, 26250, 26253-57, 37617, +37665, MCZ 15649-50, 23268-69; 6 mi. SW San Marcos, TCWC 5070-71, 7140, +9232-33, 9236, 9316-17, 9320. _Kendall Co._: 11 mi. E Boerne, AMNH +54660-61, 54662 (2); 10 mi. W Boerne, KU 18441; Kendalia, UIMNH 21434. +_Kerr Co._: Kerr W. M. Area, TCWC 15859; 40 mi. NW Kerrville, TCWC 6555. +_Medina Co._: UIMNH 13287-88; 12 mi. N Castroville, UIMNH 21423; 14 mi. +N Castroville, UIMNH 21424-25; 16 mi. N Castroville, UIMNH 21421-22; 17 +mi. N Castroville, UIMNH 21428-29; 18 mi. N Castroville, UIMNH 21426-27, +21430-33; 6.5 mi. NW Rio Medina, KU 18440. _Real Co._: Rio Frio, FMNH +55156-57. _Travis Co._: Austin, AMNH 44221-22; Mount Bonnell, 5 mi. S +Austin, UMMZ 101453 (10). _Uvalde Co._: 13 mi. from Uvalde, UIMNH 62322. +_Val-Verde Co._: 40 mi. N Del Rio, JDL 214 (skeleton). + + [Illustration: FIG. 11: Distribution of _Syrrhophus marnockii_ + (circles). Starred localities are late Pleistocene records.] + + +=Syrrhophus verrucipes= Cope + + _Syrrhophus verrucipes_ Cope, 1885:383 [Holotype.--ANSP 11325, from + near Zacualtipán, Hidalgo, México (1800 feet lower in a rocky gorge + of a stream near its junction with the Río San Miguel), collected + by Dr. Santiago Bernard]. Kellogg, 1932:126-29. Smith and Taylor, + 1948:52-53. Firschein, 1954:55-57. Gorham, 1966:167. + + _Syrrhaphus verrucipes_: Günther, 1900:216-17. + + _Tomodactylus macrotympanum_ Taylor, 1940e:496-99, pl. 55, + figs. 2a-b. [Holotype.--FMNH 100049 (formerly EHT-HMS 6838), + from La Placita, 8 km. S Jacala, Hidalgo, México, 1850 m.; + collected on July 2, 1936, by Edward H. Taylor]. Smith and + Taylor, 1948:47-48. + + _Syrrhophus macrotympanum_: Dixon, 1957:384. Gorham, 1966:165. + +_Diagnosis._--Medium-sized frogs, males 17.5-26.1 mm. snout-vent, +females 28.0-31.7 mm. snout-vent length; vocal slits in males; digital +tips slightly expanded; first finger shorter than second; skin of dorsum +pustular, that of venter areolate; snout elongate, subacuminate; +diameter of tympanum 56.1-76.7 per cent that of eye in males, 54.3-56.8 +in females; in preservative, dorsum reddish brown with numerous small +black or dark brown spots (Fig. 8); venter white to cream; in life +dorsum green with darker green spots, belly white; iris gold above, +bronze below. + +_Remarks._--Cope's (1885) original description was not sufficiently +clear to enable subsequent authors to recognize this species. Taylor +(1940e) described it as a _Tomodactylus_, but Dixon (1957) pointed out +that _T. macrotympanum_ differed from the other species of the genus in +having a poorly developed lumbo-inguinal (inguinal) gland, and placed +the species in the genus _Syrrhophus_. Comparison of the holotypes of +_S. verrucipes_ and _T. macrotympanum_ leaves no doubt in my mind that a +single species is involved. This same species was reported by Smith and +Taylor (1948) as _S. verruculatus_. + +_Syrrhophus verrucipes_ bears resemblance to members of both the +_leprus_ and _marnockii_ groups. In snout shape it is closer to the +_leprus_ group, whereas in digital pad, the shape of the general body +form, and contiguity of habitat it is most similar to the _marnockii_ +group (_S. guttilatus_). + +_Etymology._--Latin, meaning warty foot, probably in reference to the +numerous plantar supernumerary tubercles. + +_Distribution._--Moderate elevations in southeastern San Luis Potosí, +Queretaro, and northwestern Hidalgo, México (Fig. 7). + +_Specimens examined_--(43) MÉXICO, _Hidalgo_: Jacala, UMMZ 106434; 9.6 +km. NE Jacala, Puerto de la Zorra, 1820 m., KU 60240-41, TCWC 11090, +11147; 8 km. S Jacala, La Placita, 1850 m., FMNH 100049 (holotype of +_Tomodactylus macrotympanum_), 100791-803, 105334-35, 114287, UIMNH +15989-92, 15995-96, UMMZ 117252, USNM 137202; Tianguistengo, FMNH +113705-09, UIMNH 13328-30; near Zacualtipán, ANSP 11325 (holotype of +_Syrrhophus verrucipes_). _Queretaro_: 3.5 km. S San Juan del Río, EAL +1343. _San Luis Potosí_: 9.6 km. W Ahuacatlán, LSUMZ 4968-70. + + +=Syrrhophus dennisi= new species + + _Syrrhophus latodactylus_: Martin, 1958:49 (in part). + +_Holotype._--UMMZ 101121, adult male from a cave near El Pachón, 8 km. N +Antiguo Morelos, Tamaulipas, México, 250 m., collected on March 13, +1949, by Paul S. Martin. + +_Paratopotypes._--(26). UMMZ 101122 (10), 101123 (2), 101126, 126993 +(12). + +_Diagnosis._--Medium-sized frogs, males 22.8-28.4 mm. snout-vent, +females 25.9-32.0 mm. snout-vent; vocal slits in males; digital tips +greatly expanded, more than twice width of digit; first finger shorter +than second; skin of dorsum shagreened to pustular, that of venter +weakly to moderately areolate; toes webbed basally; dorsum light brown +to tan with brown vermiculations; venter white; diameter of tympanum +53.9 to 64.2 per cent that of eye in males, 50.6 to 58.7 per cent in +females. + +_Description and variation._--(Fig. 12). Head wider than body; head as +wide or wider than long in males, sometimes longer than wide in females; +snout acuminate in dorsal view, elongate and rounded in lateral profile; +canthus rostralis rounded but distinct; loreal region slightly concave, +sloping abruptly to lip; lips not flared; eyelid about two-thirds +interorbital distance; length of eye less than distance between eye and +nostril; diameter of tympanum 53.9 to 64.2 per cent that of eye in +males, 50.6 to 58.7 per cent in females; tympanum round and distinct in +both sexes; supratympanic fold moderately distinct; choanae within +border of jaws, completely visible from directly below, rounded to +slightly oval; dentigerous processes of prevomers and teeth absent; +tongue free for posterior one-half, generally oval in outline; vocal +slits present in males. + +Many scattered pustules on dorsum; flanks areolate; skin of venter +areolate or not (variability may be due to differences in preservation); +ventral disc distinct on chest and lower abdomen; inguinal gland present +or not, when present varying from very large and distinct to poorly +defined; axillary gland absent. + +First finger shorter than second; all fingers bearing truncate tips with +pads, each pad having a terminal groove; fingers fringed; fingers three +and four having dilated pads two to three times width of digit; +subarticular tubercles large, conical, rounded, simple; supernumerary +tubercles numerous on thenar surface, none on digits; three palmar +tubercles, outer slightly smaller than largest supernumerary tubercles; +row of tubercles on outer edge of forearm variable, weak to very +distinct; tips of toes wider than digits, rounded to truncate at tips, +each pad having terminal groove; toes having lateral fringes, bases of +toes united by web, web not extending to basal subarticular tubercle; +subarticular tubercles smaller than those of hand, round, conical, +simple; supernumerary tubercles numerous on plantar surfaces, extending +between metatarsal tubercles, present on toes between basal two +subarticular tubercles in some specimens; outer metatarsal tubercle +round, conical, one-half as large as ovoid, non-compressed inner +metatarsal tubercle; tarsal tubercles or folds absent. + +Ground color pale reddish-brown to tan dorsally, creamy on flanks; +dorsal pattern consisting of reddish-brown to brown vermiculations +extending onto flanks; distinct interorbital light bar present; loreal +region darker than snout, reddish-brown compared to tan or pale +reddish-brown; arms colored like dorsum; thighs banded, unicolor brown +on posterior surfaces; shanks and tarsi banded; venter white to cream +punctated with brown in some specimens. + +The variation in proportions is summarized in Table 5. + +_Remarks._--Martin (1958) expressed some doubt that this series of 26 +specimens was identical with "_S. latodactylus_." My study indicates +that the specimens from El Pachón represent a distinctive but allied +species. Males of the two species can be readily separated by the +relative sizes of the tympani, presence or absence of vocal slits, and +color pattern. Females of the two species can be separated by color +pattern. Within the type-series, the pattern varies from weakly to +strongly vermiculate but is always recognizable as vermiculate rather +than spotted as in _S. longipes_ (= _S. latodactylus_ of Taylor and +Martin). + + [Illustration: FIG. 12: _Syrrhophus dennisi_ sp. nov., holotype, + UMMZ 101121 (dorsum ×1.8, side of head ×6.1).] + +_Etymology._--The specific name is a patronym for David M. Dennis, whose +drawings greatly enhance the worth of this paper. + +_Distribution._--Known only from the type series. + + +=Syrrhophus longipes= (Baird), New combination + + _Batrachyla longipes_ Baird, 1859:35, pl. 37, fig. 1-3 + [Holotype.--apparently USNM 3237 (cited as 3207 by Cope, 1887:16), + now lost, from 40 Leagues from (probably north) México City; + collected by John Potts]. Kellogg, 1932:107. + + _Epirhexis longipes_: Cope, 1866:96. + + _Eleutherodactylus longipes_: Kellogg, 1932:107 (part). Smith and + Taylor, 1948:61. Lynch, 1963:580-581. Gorham, 1966:82. + + _Syrrhophus latodactylus_ Taylor, 1940d:396-401, pl. 43, figs. A-F, + text fig. 7 [Holotype.--FMNH 100063 (formerly EHT-HMS 6807), from + Huasteca Canyon, 15 km. W Monterrey, Nuevo León, México, 680 m.; + collected on June 20, 1936, by Edward H. Taylor]. Smith and + Taylor, 1948:50-52. Martin, 1958:48-50. Gorham, 1966:165. + +_Diagnosis._--Large frogs, males 22.1-33.2 mm. snout-vent, females +26.8-39.6 mm. snout-vent length; vocal slits lacking in males; digital +tips greatly expanded (more than twice the width of digit); first finger +shorter than second; skin of dorsum pustular, that of venter smooth; +diameter of tympanum in males 61.1-87.2 per cent that of eye, 49.5-72.1 +per cent in females; dorsum tan with large or small spots and blotches; +limbs banded; interorbital bar or triangle present. + +_Remarks._--I have applied Baird's _Batrachyla longipes_ to the frog +Taylor (1940d) called _Syrrhophus latodactylus_ because the color +pattern (Fig. 13) predominant in the southern part of the range agrees +with that described (figured) for _Batrachyla longipes_. + +The color pattern of individuals in the southern part of the range of +this species consists of large spots or blotches, whereas in the +northwestern part the pattern is made up of smaller spots. In the +northeastern part of the range, the pattern is more reduced and tends to +consist of heavy flecking. The interorbital bar is narrower in specimens +from Nuevo León and Tamaulipas and is triangular in specimens from +Hidalgo and Queretaro. + +The status of the name _Batrachyla longipes_ is currently that of a +_nomen dubium_ (Lynch, 1963). At that time, I was unaware of the +geographic variation in color pattern in _Syrrhophus latodactylus_. + +The exact type-locality of _Batrachyla longipes_ is not known. If it is +40 Leagues north of México City, the locality would be in an area where +the species has a blotched instead of a flecked or spotted pattern. No +justifiable evidence was presented to place _Batrachyla longipes_ in +_Eleutherodactylus_ instead of _Syrrhophus_. Barbour (1923) and Kellogg +(1932) associated another species (_E. batrachylus_) with _longipes_. +Taylor (1940a) noted this as a case of misidentification and corrected +the error but left _longipes_ in the genus _Eleutherodactylus_. Lynch +(1963) noted several points of morphological agreement between +_Syrrhophus_ and _B. longipes_ but did not place _longipes_ in +_Syrrhophus_. + +Baird's (1859) figures of the holotype do not illustrate prevomerine +teeth, but according to Cope (1866) they were present in the holotype. +The digital tips of the frog in the figure are somewhat narrower than +those typically seen in _S. latodactylus_. If the specimen was slightly +desiccated, as possibly was the case, the digits would appear narrower. +There is no evidence contrary to placing _Syrrhophus latodactylus_ in +the synonymy of _Batrachyla longipes_. + + [Illustration: FIG. 13: Dorsal views of _Syrrhophus longipes_ + illustrating geographic variation in pattern (left, TCWC 12179, + ×1.5; right, KU 92572, ×1.8); side of head (TCWC 10966, ×6).] + +Application of Baird's name _Batrachyla longipes_ to the species of frog +heretofore called _Syrrhophus latodactylus_ poses one serious problem. +_Batrachyla longipes_ is the type-species (by original designation) of +the genus _Epirhexis_ Cope, 1866, which has priority over _Syrrhophus_ +Cope, 1878. If _Batrachyla longipes_ is left in the status of a _nomen +dubium_, _Epirhexis_ can be forgotten, for the two names are tied +together. However, since it seems almost certain that _Batrachyla +longipes_ and _Syrrhophus latodactylus_ are conspecific, the former name +should not be left as a _nomen dubium_. _Epirhexis_ never came into +general usage (Cope cited the name four times, but no one else has used +it), whereas _Syrrhophus_ is well established in the zoological +literature. It would serve only to confuse the literature to adhere +strictly to the Law of Priority and replace _Syrrhophus_ with +_Epirhexis_. Therefore, _Syrrhophus_ is used in this paper, even though +_Epirhexis_ has priority. A request for the suppression of _Epirhexis_ +Cope, 1866, has been submitted to the International Commission of +Zoological Nomenclature (Lynch, 1967). + +_Etymology._--Latin, meaning long-footed; Taylor's _latodactylus_ refers +to the wide digital pads. + + [Illustration: FIG. 14: Distribution of _Syrrhophus dennisi_ + (triangle) and _S. longipes_ (circles).] + +_Distribution._--Moderate elevations (650 to 2000 meters) along the +Sierra Madre Oriental from central Nuevo León to northern Hidalgo, +México (Fig. 14). + +_Specimens examined._--(122) MÉXICO, _Hidalgo_: 3 km. NE Jacala, AMNH +52977; 9.6 km. NE Jacala, 1800 m., TCWC 10966-70, 12179; 8 km. S Jacala, +La Placita, 1850 m., FMNH 100266-68, 103244, UIMNH 13291, 13327. _Nuevo +León_: Salto Cola de Caballo, KU 92572; Huasteca Canyon, 15 km. W +Monterrey, 680 m., FMNH 100063 (holotype of _S. latodactylus_), UIMNH +13290; 6.5 km. N Pablillo, EAL 1319; Sabinas Hidalgo, USNM 139728. +_Queretaro_: Cueva de los Riscos, 8 km. SW Jalpan, KU 106300. _San Luis +Potosí_: 13 km. E Santa Barberita, LSUMZ 2295; second camp, San Luis +Potosí road, UIMNH 13326; Xilitla, Cueva sin nombre, UMMZ 125892. +_Tamaulipas_: 4 km. W El Carrizo, 500 m., UMMZ 111343 (31); 8 km. N +Chamal, Bee Cave, KU 106299; 14.5 km. NNW Chamal, 420 m., UMMZ +111339-40, 111342 (4), 111344 (11); 19 km. NNW Chamal, 700 m., UMMZ +111341 (3); El Chihue, 1880 m., UMMZ 111289 (4); 11 km. N Gómez Farías, +1060 m., UMMZ 101166; 11 km. WNW Gómez Farías, 1800 m., UMMZ 108507 (3); +8 km. NW Gómez Farías, 1060-1400 m., LSUMZ 11085, UMMZ 101167 (3), +101168 (4), 101169 (2), 101170 (3), 101171 (2), 101360-61, 102860, +102933 (4), 102934 (2), 102935-38, 102939 (2), 102940-43, 108800 (3), +110735, 111345-46. + + +=Syrrhophus pipilans= Taylor + + _Syrrhophus pipilans_ Taylor, 1940c:95-97, pl. 1 [Holotype.--FMNH + 100072 (formerly EHT-HMS 6843), 14.6 km. S Mazatlán, Guerrero, + México; collected on July 22, 1936, by Edward H. Taylor]. + +_Diagnosis._--Medium sized frogs, males 22.6-28.5 mm. snout-vent, +females 21.1-29.4 mm. snout-vent length; vocal slits present in males; +finger tips slightly expanded, truncate in outline; inner metatarsal +tubercle less than twice the size of outer; skin of dorsum smooth to +shagreened, that of venter smooth; tympanum 36.5-54.0 per cent diameter +of eye; dorsum dark brown with large or small light brown, orange-brown, +or yellowish spots or blotches; limbs banded; interorbital bar absent. + + [Illustration: FIG. 15: Dicegrams of ear size relative to eye + diameter in the two subspecies of _Syrrhophus pipilans_. N = 17 + in _nebulosus_, 18 in _pipilans_.] + +_Remarks._--Two subspecies were recognized by Duellman (1958). +Previously both had been treated as species. The two populations were +distinguished on the basis of color pattern and the size of the +tympanum. Measurements of 17 males of _S. p. nebulosus_ from central +Chiapas and 18 males of _S. p. pipilans_ from southcentral Oaxaca and +Guerrero, México, demonstrates that the supposed difference in tympanum +size is not significant (Fig. 15). There is, however, a tendency for +the western population of _S. pipilans_ to have larger tympani. Based on +the present examination of 112 specimens of this species the two +populations are held to be sufficiently distinct to warrant taxonomic +recognition as subspecies (Fig. 16). + + [Illustration: FIG. 16: _Syrrhophus pipilans nebulosus_ (left, + KU 58908) and _S. p. pipilans_ (right, KU 86885). ×2.7.] + +The parotoid glands attributed to this species by Taylor (1940c:95) are +merely the superficial expression of the _m. depressor mandibulae_ and +scapula. No true glands are present in the parotoid region. + + +=Syrrhophus pipilans nebulosus= Taylor + + _Syrrhophus nebulosus_ Taylor, 1943:353-55, pl. 27, figs. 3-5 + [Holotype.--FMNH 100095 (formerly EHT-HMS 3774), near Tonolá, + Chiapas, México; collected on August 27, 1935, by Hobart M. + Smith and Edward H. Taylor]. Smith and Taylor, 1948:49, 51. + + _Syrrhophus pipilans nebulosus_: Duellman, 1958:2-4, 9, 12, 14. + Stuart, 1963:32-33. Gorham, 1966:166-67. + +_Diagnosis._--Diameter of tympanum 36.6-47.8 per cent that of eye; +dorsum dark brown with numerous small light brown to yellowish spots. + +_Remarks._--The distribution of this subspecies is adequately described +by Duellman (1958). Fouquette (1960) described the vocalization of this +frog. + +_Etymology._--Latin, _nebula_, in reference to the clouded dorsal +pattern. + +_Distribution._--Low to moderate elevations along the Pacific versant of +Chiapas and in the Grijalva valley of Chiapas and Guatemala (Fig. 17). + +_Specimens examined._--(54) GUATEMALA, _Huehuetenango_: Jacaltenango, +UMMZ 117036; 35 km. SE La Mesilla, TNHC 29652. MÉXICO, _Chiapas_: 11.2 +km. N Arriaga, 300 m., UMMZ 125891; 11.8 km. N Arriaga, UMMZ 117279; +12.8 km. N Arriaga, UMMZ 117280; 17.5 km. S Arriaga, UIMNH 57108-109; +1.5 km. S Bochil, 1250 m., KU 58898-908; Cerro Hueco, 7 km. S Tuxtla +Gutierrez, UMMZ 123007; 3.2 km. S Ixtapa, UMMZ 124000; Linda Vista, ca. +2 km. NW Pueblo Nuevo Solistahuacán, KU 58897; Hda. Monserrate, 40 km. +NW Arriaga, UMMZ 102258; near San Ricardo, FMNH 100720; Tapachula, FMNH +75792, 103242, 100695-96, UIMNH 13292; 56 km. E Tapanatepec, Oaxaca, +TNHC 26942, Tonolá, FMNH 100095 (holotype), 100686-92, UIMNH 13293-95; +Tuxtla Gutierrez, FMNH 100693-94, UIMNH 13297; 19 km. N Tuxtla +Gutierrez, TNHC 25229-30; 15.5 km. NE Tuxtla Gutierrez, UMMZ 119892 (3); +19 km. NE Tuxtla Gutierrez, UMMZ 119891 (3); 8 km. NNW Tuxtla Gutierrez, +KU 37809; Unión de Juarez, FMNH 105294. + + +=Syrrhophus pipilans pipilans= Taylor + + _?Syrrhopus verruculatus_: Gadow, 1905:194. + + _Syrrhophus pipilans_ Taylor, 1940c:95-97, pl. 1 [Holotype.--FMNH + 100072 (formerly EHT-HMS 6843), from 14.6 km. S Mazatlán, + Guerrero, México; collected on July 22, 1936, by Edward H. Taylor]. + Taylor and Smith, 1945:581-82. Smith and Taylor, 1948:49, 50-51. + + _Syrrhophus pipilans pipilans_: Duellman, 1958:1-4, 8-9, 13-14, + pl. 2, fig. 1. Gorham, 1966:166. + +_Diagnosis._--Diameter of tympanum 40.6-54.0 per cent that of eye; +dorsum dark brown with large light spots or blotches. + +_Remarks._--Duellman's (1958) synopsis of this subspecies is adequate; +the distribution has not been extended, but several records are now +available which fill in gaps. + + [Illustration: FIG. 17: Distribution of _Syrrhophus pipilans_: + _nebulosus_ (open circles) and _pipilans_ (solid circles).] + +Gadow's (1905) record of _S. verruculatus_ from "Buena Vista, S. +Guerrero" is most likely applicable to this species. Gadow simply +included the name in a list of the species he had collected during his +trip in México (1902-04); no further comment was made on this species +although references to _Syrrhopus_ (sic) appear in several places in the +paper and would appear to apply to the species he had. + +_Etymology._--Latin, _pipilo_, chirping, peeping, in reference to the +call of the male. + +_Distribution._--Sea level to about 1800 meters along the Pacific +versant of western México from central Guerrero to the Isthmus of +Tehuantepec (Fig. 17). + +_Specimens examined._--(62). MÉXICO, _Guerrero_: Acapulco, UMMZ 110125; +6.4 km. N Acapulco, FMNH 100389, 100525; Agua del Obispo, 980-1000 m., +FMNH 75791, 100518-21, 100526, KU 86884-86, UIMNH 13315, UMMZ 119152, +125890 (4); 13.3 km. NW Coyuca, UIMNH 38367, 71982-83; 14.5 km. S +Mazatlán, FMNH 100072 (holotype), 100408, 100511-17, UIMNH 13302-309; +Tierra Colorado, 300 m., KU 67961, UIMNH 13313-14; near El Treinte, FMNH +126639; Xaltinanguis, FMNH 100522-24, 126640. _Oaxaca_: Cacahuatepec, +UIMNH 52853; 8 km. NW Río Canoa, 53 km. ESE Cuajinicuilapa, UIMNH +52852; 6.4 km. N El Candelaria, UIMNH 9501; 11.2 km. S El Candelaria, +UIMNH 9502; 17 km. NE Juchatengo, 1600 m., KU 86887; 31.5 km. N +Pochutla, UMMZ 123999 (2); 32.9 km. N Pochutla, 850 m., UMMZ 123996; +37.1 km. N Pochutla, UMMZ 123998 (2); 41.4 km. N Pochutla, UMMZ 123997 +(2); Cerro Quiengola, FMNH 105653; 3.8 km. N Santiago Chivela, UMMZ +115449; 14.5 km. W Tehuantepec, UMMZ 115448 (2). + + +=Syrrhophus interorbitalis= Langebartel and Shannon + + _Syrrhophus interorbitalis_ Langebartel and Shannon, 1956: 161-65, + figs. 1-2 [Holotype.--UIMNH 67061 (formerly FAS 9378), 36 mi. N + Mazatlán, Sinaloa, México, collected on November 17, 1955, by + E. C. Bay, J. C. Schaffner, and D. A. Langebartel]. Duellman, + 1958:1-4, 10, 12, 14. Gorham, 1966:164-65. + + _Syrrhophis interorbitalis_: Campbell and Simmons, 1962:194, + fig. 1. + + [Illustration: FIG. 18: Left to right. _Syrrhophus interorbitalis_ + (UIMNH 38095, ×1.5), _S. nivocolimae_ (LACM 3203, ×1.3), and + _S. teretistes_ (KU 75263, ×1.5).] + +_Diagnosis._--Medium sized frogs, only known male 25.6 mm. snout-vent, +females 20.0-26.7 mm. snout-vent length (small sample); vocal slits in +males; finger tips expanded; first finger shorter than second; outer +metatarsal tubercle one-third size of inner; skin of dorsum shagreened, +that of venter smooth; diameter of tympanum 37.7-42.4 per cent that of +eye in both sexes; pale yellow-brown ground color mottled with brown; +limb bands broad, much wider than narrow light interspaces; interorbital +bar very long, edged with dark brown to black (Fig. 18). + +_Remarks._--Duellman's (1958) measurements and proportions of +_S. interorbitalis_ were based exclusively on the type series, which is +composed of only females; therefore his _interorbitalis_ data are not +comparable with the data for the other species in his table. Campbell +and Simmons (1962) collected the only known male. The type series was +collected beneath rocks in a stream bed; the collectors heard calling +frogs in the bushes but were unable to obtain specimens (Langebartel and +Shannon, 1956). Campbell and Simmons (1962) reported that their specimen +had a poorly developed interorbital bar in life; in preservative the bar +compares favorably with the bar in the female (Fig. 18). + +_Etymology._--Latin, in reference to the pale interocular band. + +_Distribution._--Pacific lowlands of Sinaloa, México (Fig. 20). + +_Specimens examined._--(10). MÉXICO, _Sinaloa_: 36 mi. N Mazatlán, UIMNH +38094-96, 67061 (holotype), 71970-74; 65 mi. N Mazatlán, LACM 13773. + + +=Syrrhophus modestus= Taylor + + _Syrrhophus modestus_ Taylor, 1942:304-06, pl. 29 [Holotype.--FMNH + 100048 (formerly EHT-HMS 3756), from Hacienda Paso del Río, + Colima, México; collected on July 8, 1935, by Hobart M. Smith]. + Smith and Taylor, 1948:49-50. + + _Syrrhophus modestus modestus_: Duellman, 1958:2-5, 7, 14, pl. 1, + fig. 1. Gorham, 1966:166. + +_Diagnosis._--Small frogs, males 15.8-20.1 mm. snout-vent length, single +female 18.5 mm.; vocal slits present in males; finger tips widely +expanded; first finger shorter than second; inner metatarsal tubercle +about three times size of outer; skin of dorsum shagreened, that of +venter smooth; tympanum concealed; pale cream in preservative with dark +brown spots; limbs banded; bands on forearm and thigh poorly developed +or absent; interorbital bar absent. + +_Remarks._--The tympanum is concealed in _S. modestus_, +_S. nivocolimae_, _S. pallidus_, _S. teretistes_, and to a lesser degree +in _S. interorbitalis_. However, if the specimen is permitted to dry +slightly, the annulus tympanicus becomes visible through the skin and a +tympanum/eye ratio can be computed. + +One of the few cases of sympatry within the genus _Syrrhophus_ involves +this species; _modestus_ and _nivocolimae_ are known to be sympatric at +one locality in southwestern Jalisco, México. + +Duellman (1958) used the trinomial for this population and named a new +subspecies, _pallidus_, from Nayarit. I consider _pallidus_ to be +specifically distinct from _modestus_ because there is no evidence of +genetic exchange, and there is no overlap in the distinguishing +morphological features. I do consider the two populations to be closely +related but feel the interrelationships between _modestus_, _pallidus_, +_nivocolimae_, and _teretistes_ are more complex than would be indicated +by the use of trinomials. The sympatric occurrence of _modestus_ and +_nivocolimae_ is significant; morphologically, they might otherwise be +regarded as subspecies. Although allopatric, similar arguments could be +advanced for the morphologically similar _pallidus_ and _teretistes_. +The four are here afforded species rank since morphological similarity +and allopatry are not sufficient grounds for the assumption of genetic +exchange. + + [Illustration: FIG. 19: _Syrrhophus modestus_ [left, UMMZ 115447 + (WED 11155)] and _S. pallidus_ (right, UMMZ 115453). ×2.2.] + +_Etymology._--Latin, meaning unassuming, modest, in reference to the +small size of the species. + +_Distribution._--Low elevations (up to 700 meters) in the lowlands and +foothills of Colima and southwestern Jalisco, México (Fig. 20). + +_Specimens examined._--(14). MÉXICO, _Colima_: Hda. Paso del Río, FMNH +100048 (holotype), 100167, 100299, UIMNH 13300, UMMZ 110877 (2), USNM +139729; 7.2 km. SW Tecolapa, UMMZ 115477 (4); _Jalisco_: 17.6 km. SW +Autlan, 606 m., KU 102627; 3.2 km. N La Resolana, UMMZ 102100; Bahía +Tenacatita, UMMZ 84264. + + +=Syrrhophus nivocolimae= Dixon and Webb + + _Syrrhophus nivocolimae_ Dixon and Webb, 1966:1-4, Fig. 1 + [Holotype.--LACM 3200, from Nevado de Colima (6 airline miles west + of Atenquique), Jalisco, México, 7800 feet; collected on July 20, + 1964, by Robert G. Webb]. + +_Diagnosis._--Small frogs, males 18.5-21.1 mm. snout-vent length, only +known female 24.1 mm. snout-vent; vocal slits present in males; finger +tips widely expanded; first finger shorter than second; inner metatarsal +tubercle about three times size of outer; skin of dorsum warty, that of +venter smooth; tympanum concealed, its diameter 30.0-39.3 per cent that +of eye in males; mid-dorsal brown band from interorbital bar to anus; +bands on limbs narrow, dark bands less than one-half width of light +bands, upper arm not banded; narrow interorbital light bar. + +_Remarks._--This species is closely related to _S. modestus_ and differs +in color pattern and degree of wartiness of the skin. Dixon and Webb +(1966) held that _nivocolimae_ had no close relatives, but the condition +of the tympanum, size, nature of the outer palmar tubercle, relative +sizes of the metatarsal tubercles, and shape and size of the digital +pads all point to a close relationship between _S. modestus_, +_S. nivocolimae_, and _S. pallidus_. + + [Illustration: FIG. 20: Distribution of the species of the + _modestus_ group: _interorbitalis_ (open circles), _teretistes_ + (solid circles), _modestus_ (open triangles), _pallidus_ (solid + triangles) and _nivocolimae_ (square). Arrow indicates locality of + sympatry between _modestus_ and _nivocolimae_. Solid line about the + localities for _interorbitalis_ is a range estimate based on call + records and specimens examined.] + +Dixon and Webb (1966) reported that _S. nivocolimae_ has a large +tympanum (50.0-59.0 per cent diameter of eye). However, my examination +of the type series and several other specimens from Jalisco reveals that +the largest tympanum/eye ratio is 39.3 per cent. Therefore, the +tympanum/eye ratio in _S. nivocolimae_ is in agreement with those for +_S. modestus_, _S. pallidus_, and _S. teretistes_ (Table 6). + +_Etymology._--_niv_, Latin, and Colima (Nevado de), meaning high on the +volcano, in reference to the higher distribution of this species (around +2000 meters) than other members of the group. + +_Distribution._--Known from southwestern Jalisco, México, at moderate to +high elevations (600-2400 meters). + +_Specimens examined._--(48) MÉXICO, _Jalisco_: 17.6 km. SW Autlán, 606 +m., KU 102626, 102631; 6.4 km. W Atenquique, 2060 m., KU 102628-30, +102632; 8 km. W Atenquique, 1970 m., LACM 3210-12; 9.6 km. W Atenquique, +2360 m., LACM 3200 (holotype), 3201-09; 14.5 km. W Atenquique, 2000 m., +LACM 25424-36, 25439-41, 25446; 15 km. W Atenquique, LACM 37044-46, +37244-47; 16 km. W Atenquique, 2105 m., LACM 25443-45; 17 km. W +Atenquique, 2180 m., LACM 25442. + + +=Syrrhophus pallidus= Duellman, New combination + + _Syrrhophus modestus_: Davis and Dixon, 1957:146. + + _Syrrhophus modestus pallidus_ Duellman, 1958:2-3, 5-7, 14, pl. 3 + [Holotype.--UMMZ 115452, from San Blas, Nayarit, México, sea + level; collected on August 13, 1956, by William E. and Ann S. + Duellman]. Zweifel, 1960:86-88, 91, 93-94, 118, 120-22. Gorham, + 1966:166. + + _Syrrhophis modestus pallidus_: Campbell and Simmons, 1962:194. + +_Diagnosis._--Small frogs, males 17.9-19.3 mm. snout-vent length; vocal +slits in males; finger tips widely expanded; first finger shorter than +second; inner metatarsal tubercle about three times size of outer; skin +of dorsum shagreened, that of venter smooth; tympanum concealed, its +diameter 27.0-35.6 per cent of eye in males; ground color cream +vermiculated with brown, upper arm and thigh lacking, or with few, +indistinct, bands; interorbital bar absent. + +_Remarks._--Considerable debate has been waged relative to the value of +subspecies and to the reasons for recognizing distinct disjunct +populations as species versus subspecies. Lacking evidence of genetic +exchange, I prefer to retain disjunct populations that are distinctive +as species. + +All known specimens of _pallidus_ can be separated from those of +_modestus_ by color pattern. The two nominal species exhibit overlap in +proportions but the same can be said about nearly every species of +_Syrrhophus_; therefore, overlap in proportions can be disregarded in +assessing specific versus subspecific rank. Until contrary evidence is +forthcoming, I consider the disjunct populations heretofore held to be +subspecies of _modestus_ to be specifically distinct. The specimens of +the disjunct population of _pallidus_ on the Tres Marias do not differ +from the mainland population in Nayarit. This evidence, though perhaps +secondary, supports my contention that two species should be recognized. + +_Etymology._--Latin, in reference to the pale ground color in comparison +with that of _S. modestus_. + +_Distribution._--Low elevations in coastal Nayarit and on Islas Tres +Marias (Fig. 20). + +_Specimens examined._--(12) MÉXICO, _Nayarit_: 18.8 mi. NW Ahuacatlán, +UIMNH 7808; San Blas, UMMZ 115452 (holotype), 115453-57; 17 km. NE San +Blas, 150 m., MSU 5085; 12.8 km. E San Blas, UIMNH 71979; 31 km. E San +Blas, UIMNH 71978; 13.5 km. N Tepic, UIMNH 71980-81. + + +=Syrrhophus teretistes= Duellman + + _Syrrhophus teretistes_ Duellman, 1958:2-3, 10-14, pl. 2, fig. 2 + [Holotype.--UMMZ 115451, from 4.8 km. NW Tepic, Nayarit, México, + 840 m.; collected on August 12, 1956, by William E. Duellman]. + Gorham, 1966:167. + +_Diagnosis._--Medium-sized frogs, males 19.2-23.2 mm. snout-vent length, +single known female 24.8 mm. snout-vent; vocal slits in males; finger +tips widely expanded; first finger shorter than second; inner metatarsal +tubercle about three times size of outer; skin of dorsum shagreened, +that of venter smooth; tympanum partially concealed, its diameter +28.6-43.8 per cent of eye in males; ground color brown vermiculated with +dark brown to nearly black; upper arm and thigh banded; interorbital +light bar absent. + +_Remarks._--_S. teretistes_ appears to be most closely related to +_S. pallidus_; I consider it to be an upland derivative of _pallidus_. +Morphologically, the differences between the two are few, but lacking +evidence of genetic exchange they are retained as species. + +_Etymology._--Greek, in reference to the whistle-like nature of the +call. + +_Distribution._--Moderate elevations (840-1200 meters) in the Sierra +Occidental of Nayarit, Sinaloa, and Durango, México (Fig. 20). + +_Specimens examined._--(13) MÉXICO, _Nayarit_: 4.8 km. NW Tepic, 840 m., +UMMZ 115451 (holotype). _Sinaloa_: Santa Lucía, 1090 m., KU 75263-72; 1 +km. NE Santa Lucía, 1156 m., KU 78257; 2.2 km. NE Santa Lucía, 1156 m., +KU 78258. + + + + +DISCUSSION + + +There are relatively few clear-cut morphological differences among the +fourteen species now assigned to _Syrrhophus_. The majority of the +species are allopatric and differ primarily in color patterns. Sympatric +occurrence serves as an indicator of specific distinctness and is one of +the more practical tests of species validity when cross-breeding +experiments are not possible. Two cases of sympatric occurrence are +known for the species of _Syrrhophus_ in western México: _modestus_ and +_nivocolimae_ are sympatric in southern Jalisco and _pipilans nebulosus_ +and _rubrimaculatus_ are sympatric in southeastern Chiapas. In eastern +México, _longipes_ and _verrucipes_ are sympatric in southern Hidalgo, +and _longipes_ is sympatric with _cystignathoides_, _dennisi_, and +_guttilatus_ in southern Tamaulipas. _Syrrhophus cystignathoides_ and +_leprus_ are apparently sympatric in central Veracruz. + +Subspecific assignments have been made only when there is evidence of +intergradation. The sympatric occurrence of morphologically similar +species in this genus has led me to adopt a conservative approach to the +degree of difference philosophy. I have therefore recognized all +morphologically distinct allopatric populations as species. + + [Illustration: FIG. 21: Generic distributions of _Syrrhophus_ + (stipple) and _Tomodactylus_ (hatching). Black areas are zones + of intergeneric sympatry.] + +_Syrrhophus_ is closely allied to another Mexican leptodactylid genus, +_Tomodactylus_, which was revised by Dixon (1957), who along with +numerous other authors noted the close relationship between the two +genera. There is an almost complete lack of sympatry between the two +genera; in very few places in México do they coexist (Fig. 21). +_Tomodactylus_ has its greatest diversity in the Cordillera Volcánica +and Sierra Madre del Sur, whereas _Syrrhophus_ reaches its greatest +diversity in the Sierra Madre Oriental and eastern foothills. The +species of both genera are about the same size and presumably have +similar requirements insofar as food, breeding sites, and habitat +selection. + +Four cases of intergeneric sympatry are known for the two genera: +1) the Chilpancingo region of Guerrero, 2) the lowlands of Colima and +the mountains just inland in Jalisco, 3) the lowlands of central Nayarit, +and 4) the Sierra Madre Occidental on the Durango-Sinaloan border. The +apparent sympatry in the Chilpancingo region involves four species: +_S. pipilans_, _T. albolabris_, _T. dilatus_, and _T. nitidus_. Of the +four, _T. dilatus_ appears to be completely allopatric in that it occurs +at higher altitudes (above 2000 meters), whereas the other three occur +below 1800 meters in the region (Davis and Dixon, 1965). In the +Colima-Jalisco region, _Tomodactylus_ tends to occur higher (Dixon and +Webb, 1966) than some of the _Syrrhophus_, but one subspecies of +_Tomodactylus nitidus_ is a lowland frog, occurring sympatrically with +the lowland _Syrrhophus modestus_. A similar situation is observed in +Nayarit; the lowland _Tomodactylus_ occurs sympatrically with the small +_Syrrhophus pallidus_. In both cases the _Syrrhophus_ is smaller than +the _Tomodactylus_. + + [Illustration: FIG. 22: Altitudinal distributions of _Syrrhophus_ + and _Tomodactylus_. Widths of the columns are proportional to the + numbers of species at a given altitude; narrowest width equals one + species.] + +Frogs of the genus _Syrrhophus_ tend to occur at lower elevations than +do their close relatives of the genus _Tomodactylus_ (Fig. 22). This +generalization is complicated by the occurrence in the Sierra Madre +Oriental in relatively high altitude _Syrrhophus_ (up to 2000 m.) and +the occurrence in Michoacán of low altitude _Tomodactylus_ (to sea +level). There are no _Tomodactylus_ in the Sierra Madre Oriental, +whereas the genus _Syrrhophus_ is represented in the lowlands of western +México (_modestus_ group). _Syrrhophus_ and _Tomodactylus_ exhibit +essentially parapatric distributions. The two genera as now composed can +be characterized as low to moderate elevation frogs (_Syrrhophus_) and +moderate to intermediate elevation frogs (_Tomodactylus_). + + + + +LITERATURE CITED + + +BAIRD, S. F. + + 1859. Reptiles of the Boundary. United States and Mexican Boundary + Survey, pp. 1-35, pls. 1-41. + + +BARBOUR, T. + + 1923. The reappearance of Batrachyla longipes. Proc. New England + Zool. Club, 8:81-83. + + +BARBOUR, T., and A. LOVERIDGE + + 1946. Typical reptiles and amphibians; supplement. Bull. Mus. Comp. + Zool., 96:59-214. + + +BOULENGER, G. A. + + 1882. Catalogue of the Batrachia Salientia ... British Museum., + 2nd ed. + + 1888. Note on the classification of the Ranidae. Proc. Zool. Soc. + London, 1888, pt. 2:204-06. + + +CAMPBELL, H. W., and R. S. SIMMONS + + 1962. Notes on some reptiles and amphibians from western Mexico. + Bull. So. California Acad. Sci., 61:193-203. + + +CONANT, R. + + 1958. A field guide to reptiles and amphibians. Houghton-Mifflin + Co. Boston. 366 pp. + + +COPE, E. D. + + 1866. On the structures and distribution of the genera of the + arciferous Anura. J. Acad. Nat. Sci. Philadelphia, n. ser., + 6:67-112. + + 1877. Tenth contribution to the herpetology of tropical America. + Proc. Amer. Philos. Soc., 17:85-98. + + 1878. New genus of Cystignathidae from Texas. Amer. Nat., + 12:252-53. + + 1879. Eleventh contribution to the herpetology of tropical America. + Proc. Amer. Philos. Soc., 18:261-77. + + 1885. A contribution to the herpetology of Mexico. _Ibid._, + 22:379-404. + + +DAVIS, W. B., and J. R. DIXON + + 1957. Notes on Mexican amphibians, with description of a new + _Microbatrachylus_. Herpetologica, 13:145-47. + + 1965. Amphibians of the Chilpancingo Region, Mexico. _Ibid._, + 20:225-33. + + +DÍAZ DE LEÓN, J. + + 1904. Indice de los Batracios que se enquentran en la Republica + Méxicana. Imprenta de Ricardo Rodriquez Romo. Aguascalientes. + 40 pp. + + +DIXON, J. R. + + 1957. Geographic variation and distribution of the genus + Tomodactylus in Mexico. Texas J. Sci., 9:379-409. + + +DIXON, J. R., and R. G. WEBB + + 1966. A new _Syrrhophus_ from Mexico (Amphibia: Leptodactylidae). + Cont. Sc., Los Angeles Co. Mus., 102:1-5. + + +DUELLMAN, W. E. + + 1958. A review of the frogs of the genus _Syrrhophus_ in western + Mexico. Occ. Pap. Mus. Zool. Univ. Michigan, 594:1-15. + + 1960. A distributional study of the amphibians of the Isthmus of + Tehuantepec, Mexico. Univ. Kansas Publs. Mus. Nat. Hist., + 13:19-72. + + +FIRSCHEIN, I. L. + + 1954. Definition of some little-understood members of the + leptodactylid genus _Syrrhophus_, with a description of a new + species. Copeia, (1):48-58. + + +FOUQUETTE, M. J. + + 1960. Call structure in frogs of the family Leptodactylidae. Texas + J. Sci., 12:201-15. + + +GADOW, H. + + 1905. The distribution of Mexican amphibians and reptiles. Proc. + Zool. Soc. London, 1905, pt. 2:191-244. + + +GORHAM, S. W. + + 1966. Liste der rezenten Amphibien und Reptilien.... Das Tierreich. + Lief, 85:1-222. + + +GÜNTHER, A. C. L. G. + + 1885-1902. Biologia Centrali-Americana. Reptilia and Batrachia. + 326 pp., 76 pls. Syrrhophus section dated 1900. + + +KELLOGG, R. + + 1932. Mexican tailless amphibians in the United States National + Museum. Bull. U.S. Natl. Mus., 160. + + +LANGEBARTEL, D. A., and F. A. SHANNON + + 1956. A new frog (Syrrhophus) from the Sinoloan lowlands of Mexico. + Herpetologica, 12:161-65. + + +LYNCH, J. D. + + 1963. The status of _Eleutherodactylus longipes_ (Baird) of Mexico + (Amphibia: Leptodactylidae). Copeia, (3):580-81. + + 1964. Additional hylid and leptodactylid remains from the + Pleistocene of Texas and Florida Herpetologica. 20:141-42. + + 1967. _Epirhexis_ Cope, 1866 (Amphibia: Salientia): request for + suppression under the plenary powers. I. N. (S). Bull. Zool. + Nomencl., 24:313-15. + + 1968. Genera of leptodactylid frogs in México. Univ. Kansas Publs., + Mus. Nat. Hist., 17:503-15. + + +MARTIN, P. S. + + 1958. A biogeography of reptiles and amphibians in the Gomez Farias + region, Tamaulipas, Mexico. Misc. Publs. Mus. Zool. Univ. + Michigan, 101:1-102. + + +MILSTEAD, W. M., J. S. MECHAM, and H. MCCLINTOCK + + 1950. The amphibians and reptiles of the Stockton Plateau in + northern Terrell County, Texas. Texas J. Sci., 2:543-62. + + +NEILL, W. T. + + 1965. New and noteworthy amphibians and reptiles from British + Honduras. Bull. Florida State Mus., 9:77-130. + + +NIEDEN, F. + + 1923. Anura I ... Das Tierreich. Lief., 46:1-584. + + +PETERS, W. + + 1871. Über neue Amphibien ... des Konigl. Zoologischen Museums. + Monatsb. k. k. Preuss. Akad. Wiss. Berlin, 1870:641-52. + + +SCHMIDT, K. P., and T. F. SMITH + + 1944. Amphibians and reptiles of the Big Bend Region of Texas. + Field Mus. Nat. Hist., zool. ser., 29:75-96. + + +SMITH, H. M. + + 1947. Notes on Mexican amphibians and reptiles. J. Washington Acad. + Sci., 37:408-12. + + +SMITH, H. M., and E. H. TAYLOR + + 1948. An annotated checklist and key to the Amphibia of Mexico. + Bull. U.S. Natl. Mus., 194:1-118. + + +STEJNEGER, L. + + 1915. A new species of tailless batrachian from North America. + Proc. Biol. Soc. Washington, 28:131-32. + + +TAYLOR, E. H. + + 1940a. A new eleutherodactylid frog from Mexico. Proc. New England + Zool. Club, 18:13-16. + + 1940b. Two new anuran amphibians from Mexico. Proc. U.S. Natl. + Mus., 89:43-47, 1 pl. + + 1940c. A new Syrrhophus from Guerrero, Mexico. Proc. Biol. Soc. + Washington, 53:95-98, 1 pl. + + 1940d. New species of Mexican Anura. Univ. Kansas Sci. Bull., + 26:385-405. + + 1940e. Herpetological miscellany no. I. _Ibid._, 26:489-571. + + 1942. New Caudata and Salientia from México. _Ibid._, 28:295-323. + + 1943. Herpetological novelties from Mexico. _Ibid._, 29:343-61. + + 1952. A review of the frogs and toads of Costa Rica. _Ibid._, + 35:577-942. + + +TAYLOR, E. H., and H. M. SMITH + + 1945. Summary of the collections of amphibians made in Mexico under + the Walter Rathbone Bacon Traveling Scholarship. Proc. U.S. + Natl. Mus., 95:521-613. + + +TIHEN, J. A. + + 1960. Notes on Late Cenozoic hylid and leptodactylid frogs from + Kansas, Oklahoma and Texas. Southwest. Nat., 5:66-70. + + +WRIGHT, A. H., and A. A. WRIGHT + + 1949. Handbook of frogs and toads. 3rd ed. Comstock. 640 pp. + + +YARROW, H. C. + + 1882. Checklist of North American Reptilia and Batrachia, with + catalogue of specimens in U.S. National Museum. Bull. U.S. + Natl. Mus., 24:1-249. + + +ZWEIFEL, R. G. + + 1960. Results of the Puritan-American Museum of Natural History + Expedition to Western Mexico. 9. Herpetology of the Tres + Marias Islands. Bull. Amer. Mus. Nat. Hist., 119:81-128. + + + + +TRANSCRIBER'S NOTES + + +Although _Syrrhophus marnocki_ and _Syrrhophus marnockii_ both appear +in this text, a literature search shows that both spellings have been +used and the two instances where there is only one "i" at the end are +in reference to priviously published names. Therefore, they were left +as is. With the exception of the list below and a number of silent +corrections, the text presented is that of the original printed version. + +Typographical Corrections + + Page Correction + ==== ====================== + 3 otherwse => otherwise + 5 poltypic => polytypic + 12 interorbtal => interorbital + 14 neublosus => nebulosus + 16 Cuidad => Ciudad + 16 1946-170 => 1946:170 + 22 rubrimacultaus => rubrimaculatus + 27 resemblence => resemblance + +Text Emphasis + + _Text_ - Italics + + =Text= - Bold + + + + + + +End of the Project Gutenberg EBook of A Taxonomic Revision of the +Leptodactylid Frog Genus Syrrhophus Cope, by John D. Lynch + +*** END OF THIS PROJECT GUTENBERG EBOOK A TAXONOMIC REVISION OF THE *** + +***** This file should be named 37809-8.txt or 37809-8.zip ***** +This and all associated files of various formats will be found in: + http://www.gutenberg.org/3/7/8/0/37809/ + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper 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 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 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 +business@pglaf.org. 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 + gbnewby@pglaf.org + + +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. diff --git a/37809-8.zip b/37809-8.zip Binary files differnew file mode 100644 index 0000000..117739c --- /dev/null +++ b/37809-8.zip diff --git a/37809-h.zip b/37809-h.zip Binary files differnew file mode 100644 index 0000000..11539e2 --- /dev/null +++ b/37809-h.zip diff --git a/37809-h/37809-h.htm b/37809-h/37809-h.htm new file mode 100644 index 0000000..732511c --- /dev/null +++ b/37809-h/37809-h.htm @@ -0,0 +1,3542 @@ +<!DOCTYPE html PUBLIC "-//W3C//DTD XHTML 1.0 Strict//EN" + "http://www.w3.org/TR/xhtml1/DTD/xhtml1-strict.dtd"> + <html xmlns="http://www.w3.org/1999/xhtml"> + <head> + <meta http-equiv="Content-Type" content="text/html;charset=iso-8859-1" /> + <meta http-equiv="Content-Style-Type" content="text/css" /> + <title> + The Project Gutenberg eBook of A Taxonomic Revision Of The Leptodactylid Frog Genus Syrrhophus Cope, by John D. Lynch. + </title> + <style type="text/css"> + + .book {margin-left: 10%; margin-right: 10%;} + p {text-align: justify; text-indent: 1.5em;} + ins {background-color: #e0ffe0; text-decoration: none;} + table {margin-left: auto; padding:4px; margin-right: auto; border-collapse: collapse;} + .brdbt {border-bottom: solid #000 1px;} + .brdbt2 {border-bottom: solid #000 2px;} + .brdtp2 {border-top: solid #000 2px;} + .pagenum {position: absolute; left: 92%; text-indent:0; font-size: 0.75em; text-align: right; color: #b0b0b0;} + .pagenum2 {position: absolute; left: 92%; text-indent:0; text-align: right; color: #b0b0b0;} + .reference {margin-left: 5.5em; text-indent: -3em;} + .vtop {vertical-align: top;} + .center {text-align: center; text-indent:0; } + .text_lf {text-align: left;} + .text_rt {text-align: right;} + .smaller {font-size: 0.75em;} + .smcap {font-variant: small-caps;} + .caption1 {font-weight: bold; font-size:2.00em; margin-top: 1.5em; margin-bottom: 1.5em; text-align: center;} + .caption2 {font-weight: bold; font-size:1.50em; margin-top: 1em; margin-bottom: 1em; text-align: center;} + .caption2nb {font-size:1.50em; margin-top: 1em; margin-bottom: 1em; text-align: center;} + .caption3 {font-weight: bold; font-size:1.15em; margin-top: 1em; margin-bottom: 1em; text-align: center;} + .caption3nb {font-size:1.15em; margin-top: 1em; margin-bottom: 1em; text-align: center;} + .caption4 {font-weight: bold; font-size:1em; margin-top: 1em; margin-bottom: 1em; text-align: center;} + .trans_notes {background:#d0d0d0; padding: 7px; border:solid black 1px;} + .species_ref {margin-left: 2.5em; text-indent: -2.5em; margin-top: 1em; + margin-bottom: 1em; text-align: justify;} + .row_color1 {background-color:#e0e0e0;} + .footnote {margin-left: 10%; margin-right: 10%; font-size: 0.9em;} + .footnote .label {text-align: right;} + .fnanchor {vertical-align: super; font-size: .8em; text-decoration: none;} + .fig_center {margin-left: auto; margin-right: auto; margin-bottom: 1em;} + .fig_left {float: left; clear: left; text-align: center; padding: 3px; margin: 0 4px 0 0;} + .fig_caption {text-align: center;} + + </style> + </head> +<body> + + +<pre> + +The Project Gutenberg EBook of A Taxonomic Revision of the Leptodactylid +Frog Genus Syrrhophus Cope, by John D. Lynch + +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: A Taxonomic Revision of the Leptodactylid Frog Genus Syrrhophus Cope + +Author: John D. Lynch + +Release Date: October 21, 2011 [EBook #37809] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK A TAXONOMIC REVISION OF THE *** + + + + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper and +the Online Distributed Proofreading Team at +http://www.pgdp.net + + + + + + +</pre> + + +<div class="book"><!-- Begin Book --> + +<div class="fig_center" style="width: 233px;"> +<img src="images/cover.jpg" width="233" height="406" alt="" title="" /> +</div> +<br /> +<br /> + +<div class="center"> +<br /> +<br /> +<img src="images/bar_double.png" width="100%" height="15" alt="double bar" /> +<div class="caption2 smcap">University of Kansas Publications</div> +<br /> +<div class="caption2 smcap">Museum of Natural History</div> +<br /> +<img src="images/bar_single.png" width="125" height="15" title="bar" alt="bar" /> +<br /> +<div class="caption2">Volume 20, No. 1, pp. 1-45, 22 figs.</div> +<br /> +<img src="images/bar_single.png" width="250" height="15" title="bar" alt="bar" /> + <span class="caption2">February 20, 1970</span> +<img src="images/bar_single.png" width="250" height="15" title="bar" alt="bar" /> +<br /> +<br /> +<br /> +<br /> +<br /> +<br /> +<div class="caption1"> +A Taxonomic Revision<br /> +of the Leptodactylid Frog Genus<br /> +Syrrhophus Cope<br /> +</div> +<br /> +<br /> +<br /> +<br /> +<div class="caption3">BY</div> +<br /> +<br /> +<div class="caption2">JOHN D. LYNCH</div> +<br /> +<br /> +<br /> +<br /> +<br /> +<br /> +<br /> +<br /> +<br /> +<br /> +<br /> +<br /> +<div class="caption2"> +<span class="smcap">University of Kansas</span><br /> +<span class="smcap">Lawrence</span><br /> +1970<br /> +</div> +<br /> +<br /> +<br /> +</div> + +<p><span class="pagenum"><a name="Page_2" id="Page_2">[Pg 2]</a></span></p> +<div class="center"> +<div class="caption3"> +<span class="smcap">University of Kansas Publications, Museum of Natural History</span><br /> +<br /> +Editors of this number:<br /> +Frank B. Cross, Philip S. Humphrey, William E. Duellman<br /> +<br /> +<br /> +<br /> +<br /> +Volume 20, No. 1, pp. 1-45, 22 figs.<br /> +Published February 20, 1970<br /> +<br /> +<br /> +<br /> +<br /> +<br /> +<br /> +<br /> +<span class="smcap">University of Kansas</span><br /> +Lawrence, Kansas<br /> +<br /> +</div> +<br /> +<br /> +<br /> +<br /> +<div class="caption4"> +PRINTED BY<br /> +THE UNIVERSITY OF KANSAS PRINTING SERVICE<br /> +LAWRENCE, KANSAS<br /> +1970<br /> +</div> +</div> +<br /> +<br /> +<br /> +<br /> + +<p><span class="pagenum"><a name="Page_3" id="Page_3">[Pg 3]</a></span></p> + +<div class="caption1">A Taxonomic Revision of the Leptodactylid Frog<br /> +Genus Syrrhophus Cope</div> + +<div class="caption3">BY</div> + +<div class="caption2">JOHN D. LYNCH</div> +<br /> + +<div class="smcap caption2nb">Introduction</div> + +<p>Cope (1878) proposed the genus <i>Syrrhophus</i> for a medium-sized +leptodactylid frog from central Texas; in the ensuing 75 years the genus +was expanded to include a heterogeneous group of frogs ranging from +Texas to Peru. Taylor (1952) and Firschein (1954) limited the genus to +several species of frogs occurring in Guatemala, México, and Texas. +Lynch (1968) provided a definition of the previously loosely-defined +genus.</p> + +<p>With the exception of Taylor (1952), who treated the Costa Rican +species, none of these authors dealt with the present status of the +nineteen species erroneously assigned to <i>Syrrhophus</i>. These species are +listed in Tables <a href="#Table_1">1</a> and <a href="#Table_2">2</a> with the name currently applied. Some of them +are new combinations and their justifications will be published +elsewhere. Gorham (1966) is the most recent author to include South +American species in the genus <i>Syrrhophus</i>.</p> + +<p>Smith and Taylor (1948) recognized two species groups of the genus in +México, an eastern and a western group (here termed complexes for +purposes of discussion), separated on the basis of the number of palmar +(metacarpal) tubercles (three palmar tubercles in the members of the +eastern complex and two in those of the western complex). Duellman +(1958) reviewed the species of the genus occurring in western México and +concluded that there were five species (two polytypic). Dixon and Webb +(1966) described an additional species from Jalisco, México. The +distributions of some species have been extended, but +<ins title='Correction: was "otherwse"'>otherwise</ins> the western complex of species remains unchanged since +Duellman's review.</p> + +<p>Smith and Taylor (1948) recognized seven species of the genus in eastern +México. Firschein revised the eastern complex (as then understood), and +in so doing added one new species and treated <i>Syrrhophus verruculatus</i> +as a <i>nomen dubium</i>. Dixon (1957) redefined the related genus +<i>Tomodactylus</i> and transferred <i>T. macrotympanum</i> Taylor to the genus +<i>Syrrhophus</i>. Neill (1965) described a new subspecies of <i>S. leprus</i> +from British Honduras. Two species (<i>S. gaigeae</i> and <i>S. marnockii</i>) +were recognized in Texas until Milstead, Mecham, and McClintock (1950) +synonymized <i>S. gaigeae</i> +<span class="pagenum"><a name="Page_4" id="Page_4">[Pg 4]</a></span> +with <i>S. marnockii</i>. Thus, at present, nine +species (one polytypic) are recognized on the eastern slopes and +lowlands from central Texas to British Honduras. These are currently +placed on one species group equivalent to the western complex reviewed +by Duellman (1958).</p> +<br /> + +<div class="center"> +<a name="Table_1" id="Table_1"></a> +<span class="smcap">Table 1</span>—Species Described as Members of the Genus <i>Syrrhophus</i><br /> +but Now Placed in Other Genera.<br /> + +<table style="text-align:left; width:60%;" summary="Species List"> +<tr> + <td class="brdtp2 brdbt">Trivial name and author</td> + <td class="brdtp2 brdbt"> </td> + <td class="brdtp2 brdbt">Current combination</td> +</tr> +<tr> + <td><i>areolatus</i> Boulenger, 1898</td> + <td> </td> + <td><i>Eleutherodactylus areolatus</i></td> +</tr> +<tr> + <td><i>calcaratus</i> Andersson, 1945</td> + <td> </td> + <td><i>Eleutherodactylus anderssoni</i></td> +</tr> +<tr> + <td><i>caryophyllaceus</i> Barbour, 1928</td> + <td> </td> + <td> <i>Eleutherodactylus caryophyllaceus</i></td> +</tr> +<tr> + <td><i>coeruleus</i> Andersson, 1945</td> + <td> </td> + <td><i>Eleutherodactylus coeruleus</i></td> +</tr> +<tr> + <td><i>ineptus</i> Barbour, 1928</td> + <td> </td> + <td><i>Eleutherodactylus diastema</i></td> +</tr> +<tr> + <td><i>juninensis</i> Shreve, 1938</td> + <td> </td> + <td><i>Eupsophus juninensis</i></td> +</tr> +<tr> + <td><i>lutosus</i> Barbour and Dunn, 1921</td> + <td> </td> + <td><i>Eleutherodactylus lutosus</i></td> +</tr> +<tr> + <td><i>molinoi</i> Barbour, 1928</td> + <td> </td> + <td><i>Eleutherodactylus molinoi</i></td> +</tr> +<tr> + <td><i>montium</i> Shreve, 1938</td> + <td> </td> + <td><i>Niceforonia montia</i></td> +</tr> +<tr> + <td><i>mystaceus</i> Barbour, 1922</td> + <td> </td> + <td><i>Eleutherodactylus rhodopis</i></td> +</tr> +<tr> + <td><i>obesus</i> Barbour, 1928</td> + <td> </td> + <td><i>Eleutherodactylus punctariolus</i></td> +</tr> +<tr> + <td><i>omiltemanus</i> Gunther, 1900</td> + <td> </td> + <td><i>Eleutherodactylus omiltemanus</i><a name="FNanchor_1_1" id="FNanchor_1_1"></a><a href="#Footnote_1_1" class="fnanchor">[1]</a></td> +</tr> +<tr> + <td class="brdbt2"><i>pardalis</i> Barbour, 1928</td> + <td class="brdbt2"> </td> + <td class="brdbt2"><i>Eleutherodactylus pardalis</i></td> +</tr> +</table> +<br /> +<div class="center"> +<div class="footnote"> +<a name="Footnote_1_1" id="Footnote_1_1"></a> +<a href="#FNanchor_1_1"><span class="label">[1]</span></a> New combination. +</div> +</div> +</div> +<br /> +<br /> + +<div class="center"> +<a name="Table_2" id="Table_2"></a> +<span class="smcap">Table 2</span>—Species Incorrectly Regarded as Members of the Genus <i>Syrrhophus</i><br /> +but Described as Members of Other Genera.<br /> + +<table style="text-align:left; width:60%;" summary="Species Incorrectly Ided as Genus Syrrhophus"> +<tr> + <td class="brdtp2 brdbt">Trivial name, original generic assignment, and author</td> + <td class="brdtp2 brdbt"> </td> + <td class="brdtp2 brdbt">Current combination</td> +</tr> +<tr> + <td><i>chalceus</i> (<i>Phyllobates</i>) Peters, 1873</td> + <td> </td> + <td><i>Eleutherodactylus chalceus</i></td> +</tr> +<tr> + <td><i>festae</i> (<i>Paludicola</i>) Peracca, 1904</td> + <td> </td> + <td><i>Niceforonia festae</i></td> +</tr> +<tr> + <td><i>hylaeformis</i> (<i>Phyllobates</i>) Cope, 1875</td> + <td> </td> + <td><i>Eleutherodactylus hylaeformis</i></td> +</tr> +<tr> + <td><i>palmatus</i> (<i>Phyllobates</i>) Werner, 1899</td> + <td> </td> + <td><i>Colostethus palmatus</i></td> +</tr> +<tr> + <td><i>ridens</i> (<i>Phyllobates</i>) Cope, 1866</td> + <td> </td> + <td><i>Eleutherodactylus ridens</i></td> +</tr> +<tr> + <td class="brdbt2"><i>simonsii</i> (<i>Paludicola</i>) Boulenger, 1900</td> + <td class="brdbt2"> </td> + <td class="brdbt2"><i>Niceforonia simonsii</i></td> +</tr> +</table> +</div> +<br /> +<br /> + +<div class="center"> +<a name="Table_3" id="Table_3"></a> +<span class="smcap">Table 3</span>—Nominal Species of <i>Syrrhophus</i> (<i>sensu strictu</i>)<br /> +and the Name Used Herein.<br /> +<table style="text-align:left; width:60%;" summary="Nominal Species of Syrrhophus"> +<tr> + <td class="brdtp2 brdbt">Original combination</td> + <td class="brdtp2 brdbt"> </td> + <td class="brdtp2 brdbt">Current combination</td> +</tr> +<tr> + <td><i>campi</i>, <i>Syrrhophus</i></td> + <td> </td> + <td><i>cystignathoides campi</i></td> +</tr> +<tr> + <td><i>cholorum</i>, <i>Syrrhophus leprus</i></td> + <td> </td> + <td><i>leprus</i></td> +</tr> +<tr> + <td><i>cystigathoides</i>, <i>Phyllobates</i></td> + <td> </td> + <td><i>cystignathoides cystignathoides</i></td> +</tr> +<tr> + <td><i>dennisi</i>, <i>Syrrhophus</i></td> + <td> </td> + <td><i>dennisi</i> new species</td> +</tr> +<tr> + <td><i>gaigeae</i>, <i>Syrrhophus</i></td> + <td> </td> + <td><i>guttilatus</i></td> +</tr> +<tr> + <td><i>guttilatus</i>, <i>Malachylodes</i></td> + <td> </td> + <td><i>guttilatus</i></td> +</tr> +<tr> + <td><i>interorbitalis</i>, <i>Syrrhophus</i></td> + <td> </td> + <td><i>interorbitalis</i></td> +</tr> +<tr> + <td><i>latodactylus</i>, <i>Syrrhophus</i></td> + <td> </td> + <td><i>longipes</i></td> +</tr> +<tr> + <td><i>leprus</i>, <i>Syrrhophus</i></td> + <td> </td> + <td><i>leprus</i></td> +</tr> +<tr> + <td><i>longipes</i>, <i>Batrachyla</i></td> + <td> </td> + <td><i>longipes</i></td> +</tr> +<tr> + <td><i>macrotympanum</i>, <i>Tomodactylus</i></td> + <td> </td> + <td><i>verrucipes</i></td> +</tr> +<tr> + <td><i>marnockii</i>, <i>Syrrhophus</i></td> + <td> </td> + <td><i>marnockii</i></td> +</tr> +<tr> + <td><i>modestus</i>, <i>Syrrhophus</i></td> + <td> </td> + <td><i>modestus</i></td> +</tr> +<tr> + <td><i>nebulosus</i>, <i>Syrrhophus</i></td> + <td> </td> + <td><i>pipilans nebulosus</i></td> +</tr> +<tr> + <td><i>nivocolimae</i>, <i>Syrrhophus</i></td> + <td> </td> + <td><i>nivocolimae</i></td> +</tr> +<tr> + <td><i>pallidus</i>, <i>Syrrhophus modestus</i></td> + <td> </td> + <td><i>pallidus</i></td> +</tr> +<tr> + <td><i>petrophilus</i>, <i>Syrrhophus</i></td> + <td> </td> + <td><i>guttilatus</i></td> +</tr> +<tr> + <td><i>pipilans</i>, <i>Syrrhophus</i></td> + <td> </td> + <td><i>pipilans pipilans</i></td> +</tr> +<tr> + <td><i>rubrimaculatus</i>, <i>Syrrhophus</i></td> + <td> </td> + <td><i>rubrimaculatus</i></td> +</tr> +<tr> + <td><i>smithi</i>, <i>Syrrhophus</i></td> + <td> </td> + <td><i>guttilatus</i></td> +</tr> +<tr> + <td><i>teretistes</i>, <i>Syrrhophus</i></td> + <td> </td> + <td><i>teretistes</i></td> +</tr> +<tr> + <td><i>verrucipes</i>, <i>Syrrhophus</i></td> + <td> </td> + <td><i>verrucipes</i></td> +</tr> +<tr> + <td class="brdbt2"><i>verruculatus</i>, <i>Phyllobates</i></td> + <td class="brdbt2"> </td> + <td class="brdbt2"><i>Nomen dubium</i></td> +</tr> +</table> +</div> +<br /> +<br /> + +<p><span class="pagenum"><a name="Page_5" id="Page_5">[Pg 5]</a></span> +In the course of preparing an account of the species of +<i>Eleutherodactylus</i> occurring in México and northern Central America, it +became necessary to reëxamine the status of the genus <i>Syrrhophus</i> and +its nominal species. It soon became evident that there were more names +than species, that some previously regarded species were geographic +variants, and that the eastern and western groups (complexes here) were +artificial divisions of the genus. I conclude that there are seven +species (one <ins title='Correction: was "poltypic"'>polytypic</ins>) +of <i>Syrrhophus</i> in eastern México, +Texas, and El Petén of Guatemala, and seven species (one polytypic) in +western México. The current status of each of the 23 names correctly +assigned to the genus is presented in <a href="#Table_3">Table 3</a>.</p> + +<p>The fourteen species recognized by me are placed in five species groups. +Two of these groups are presently placed in the western complex +(<i>modestus</i> and <i>pipilans</i> groups) and three in the eastern complex +(<i>leprus</i>, <i>longipes</i> and <i>marnockii</i> groups). The two complexes do not +correspond exactly with the eastern and western groups of Smith and +Taylor (1948), Firschein (1954), and Duellman (1958) since <i>S. +rubrimaculatus</i> is now associated with the eastern <i>leprus</i> group.</p> + +<p>The definitions and contents of the five species groups are as follows:</p> + +<div class="species_ref"><i>leprus</i> group: digital pads not or only slightly expanded, rounded +in outline; first finger longer or shorter than second; snout +acuminate or subacuminate, not rounded; outer metatarsal tubercle +conical; digits lacking distinct lateral fringes.<br /> +content: <i>cystignathoides</i>, <i>leprus</i> and <i>rubrimaculatus</i>.</div> + +<div class="species_ref"><i>longipes</i> group: digital pads widely expanded, triangular in outline; +first finger shorter than second; snout acuminate; outer metatarsal +tubercle not conical; digits bearing lateral fringes.<br /> +content: <i>dennisi</i> and <i>longipes</i>.</div> + +<div class="species_ref"><i>marnockii</i> group: digital pads expanded, rounded to truncate in +outline; first finger equal in length to second or slightly shorter; +snout rounded; outer metatarsal tubercle not conical; digits lacking +lateral fringes; generally stout-bodied frogs.<br /> +content: <i>guttilatus</i>, <i>marnockii</i>, and <i>verrucipes</i>.</div> + +<div class="species_ref"><i>modestus</i> group: digital pads expanded, truncate in outline; first and +second fingers subequal in length, first usually slightly shorter than +second; snout subacuminate; inner metatarsal tubercle twice as large (or +larger) as outer metatarsal tubercle; digits bearing poorly-defined +lateral fringes.<br /> +content: <i>interorbitalis</i>, <i>modestus</i>, <i>nivocolimae</i>, <i>pallidus</i>, +and <i>teretistes</i>.</div> + +<p><span class="pagenum"><a name="Page_6" id="Page_6">[Pg 6]</a></span></p> +<div class="species_ref"><i>pipilans</i> group: digital pads not or only slightly expanded, +truncate in outline; first finger equal in length to second; snout +subacuminate; metatarsal tubercles subequal in size; digits lacking +lateral fringes.<br /> +content: <i>pipilans</i>.</div> + +<div class="smaller"> +<p><i>Acknowledgments.</i>—For loan of specimens, I am indebted to Richard +J. Baldauf, Texas A & M University (TCWC); W. Frank Blair, +University of Texas (TNHC); Charles M. Bogert and Richard G. +Zweifel, American Museum of Natural History (AMNH); James E. Böhlke +and Edmond V. Malnate, Academy of Natural Sciences of Philadelphia +(ANSP); Robert F. Inger and Hymen Marx, Field Museum of Natural +History (FMNH); Ernest A. Liner (EAL); Michael Ovchynnyk, Michigan +State University collection (MSU); James A. Peters, United States +National Museum (USNM); Douglas A. Rossman, Louisiana State +University Museum of Zoology (LSUMZ); Hobart M. Smith, University +of Illinois Museum of Natural History (UIMNH); Charles F. Walker, +University of Michigan Museum of Zoology (UMMZ); and John W. +Wright, Los Angeles County Museum (LACM). Specimens in the +collection at the University of Kansas Museum of Natural History +are identified as KU. The abbreviations EHT-HMS refer to the Edward +H. Taylor-Hobart M. Smith collection and FAS to the Frederick A. +Shannon collection. The type-specimens from these collections are +now in the Field Museum of Natural History and the University of +Illinois Museum of Natural History.</p> + +<p>I have profited from discussions concerning this problem with +several persons, most notably William E. Duellman, Hobart M. Smith, +Edward H. Taylor and Charles F. Walker. Nevertheless, the ideas and +conclusions presented here should not be construed as necessarily +reflecting their opinions.</p> + +<p>David M. Dennis executed all of the figures, and my wife, Marsha, +typed the manuscript.</p> +</div> + +<p><i>Materials and Methods.</i>—In the course of this study, 1003 specimens +of the genus were examined. The holotypes of 21 of the 23 +nominal species are extant; I have examined 19 of these. Nine +measurements were taken, and five ratios computed for each of 338 +specimens. Females are available for all species but one; thus, +measurements were taken on individuals of both sexes.</p> +<br /> + +<div class="smcap caption2nb">Analysis of Characters</div> + +<p><i>Size and proportions.</i>—Frogs of this genus range in size from 16 to 40 +mm. in snout-vent length. Five species are relatively small: <i>S. +cystignathoides</i>, <i>modestus</i>, <i>nivocolimae</i>, <i>pallidus</i> and +<i>rubrimaculatus</i>; one, <i>S. longipes</i>, is relatively large, and the +remaining eight species are intermediate in size (22-30 mm.).</p> + +<p>Males are generally smaller than females and have proportionately longer +heads and usually larger tympani. No significant differences were found +among proportions, except that <i>S. longipes</i> has a larger tympanum/eye +ratio than any other species. Frogs in the <i>Syrrhophus marnockii</i> group +tend to have shorter shanks and feet, thereby giving those species a +more stocky appearance. However, the differences are not significant.</p> + +<p>A summary of the data on size and proportions for the frogs of the genus +<i>Syrrhophus</i> is given in Tables <a href="#Table_4">4</a>, <a href="#Table_5">5</a>, and <a href="#Table_6">6</a>.</p> + +<p><span class="pagenum"><a name="Page_7" id="Page_7">[Pg 7]</a></span> +<i>Hands and Feet.</i>—Taylor and Smith (1945), Smith and Taylor (1948), +Firschein (1954) and Duellman (1958) discussed the value of the palmar +tubercles in identifying frogs of this genus. The eastern complex in +general has a well-developed outer palmar tubercle (<a href="#Fig_1">Fig. 1</a>) in +distinction to the western complex in which the outer palmar tubercle is +reduced or absent (<a href="#Fig_2">Fig. 2</a>). Dixon and Webb (1966) imply that the outer +palmar tubercle is rarely absent but is usually smaller than the first +supernumerary tubercle of the fourth finger. My study of the western +species demonstrates that the outer palmar tubercle is indeed usually +present and smaller than the first supernumerary tubercle.</p> + +<p>Differences in interpretation of the terms "unexpanded" and "narrow," as +well as differences in techniques of preservation, have led to confusion +of the reported digital shapes in various species. Constant specific +differences are evident in the hands (<a href="#Fig_1">Fig. 1</a>). Except in the cases of +excessive uptake of fluids, all species have a terminal transverse +groove at the tip of each digit. Taylor (1940b) stated that <i>S. smithi</i> +lacked grooves, but examination of the holotype reveals faint grooves at +the tops of the digits. <i>Syrrhophus guttilatus</i>, <i>leprus</i>, <i>pipilans</i>, +and <i>verrucipes</i> lack lateral fringes on the fingers. Lateral fringes +are well developed in the <i>longipes</i> and <i>modestus</i> groups but poorly +defined or absent in the other members of the genus. The digital pads of +the frogs of the <i>longipes</i> group are much broader than those of the +other species and are narrowest in the frogs of the <i>leprus</i> group. +Supernumerary tubercles are present on the palmar surfaces of all +species of the genus.</p> +<br /> + +<div class="center"> +<a name="Table_4" id="Table_4"></a> +<span class="smcap">Table 4</span>—Size and Proportions in the Frogs +of the <i>Syrrhophus leprus</i> Group.<br /> + +<table style="text-align:center; width:100%;" summary="Frog Sizes and Proportions"> +<tr> + <td class="brdtp2 brdbt">Species</td> + <td class="brdtp2 brdbt">Sex</td> + <td class="brdtp2 brdbt">N</td> + <td class="brdtp2 brdbt">Snout-vent<br />length<br />(SVL)</td> + <td class="brdtp2 brdbt">Tibia<br />length/<br />SVL</td> + <td class="brdtp2 brdbt">Head<br />width/<br />SVL</td> + <td class="brdtp2 brdbt">Tympanum/<br />Eye</td> + <td class="brdtp2 brdbt">Eyelid/<br />Interorbital</td> +</tr> +<tr> + <td class="text_lf"><i>cystignathoides campi</i></td> + <td>♂</td> + <td>33</td> + <td>16.3-23.5</td> + <td>41.3-49.6</td> + <td>34.0-40.1</td> + <td>43.7-66.5</td> + <td>43.2-89.6</td> +</tr> +<tr> + <td> </td> + <td> </td> + <td> </td> + <td>(45.8)</td> + <td>(37.0)</td> + <td>(56.2)</td> + <td>(61.5)</td> +</tr> +<tr> + <td> </td> + <td>♀</td> + <td>12</td> + <td>16.0-25.8</td> + <td>41.5-51.0</td> + <td>33.0-38.0</td> + <td>42.8-60.0</td> + <td>48.2-69.2</td> +</tr> +<tr> + <td> </td> + <td> </td> + <td> </td> + <td>(45.8)</td> + <td>(35.0)</td> + <td>(51.2)</td> + <td>(60.1)</td> +</tr> +<tr> + <td class="text_lf"><i>c. cystignathoides</i></td> + <td>♂</td> + <td>15</td> + <td>16.8-22.1</td> + <td>45.1-50.4</td> + <td>33.2-40.7</td> + <td>44.3-68.7</td> + <td>44.6-65.4</td> +</tr> +<tr> + <td> </td> + <td> </td> + <td> </td> + <td>(47.3)</td> + <td>(37.8)</td> + <td>(54.8)</td> + <td>(60.0)</td> +</tr> +<tr> + <td> </td> + <td>♀</td> + <td>6</td> + <td>19.6-24.2</td> + <td>46.4-50.0</td> + <td>34.1-38.1</td> + <td>43.3-56.5</td> + <td>53.2-65.4</td> +</tr> +<tr> + <td> </td> + <td> </td> + <td> </td> + <td>(47.6)</td> + <td>(36.2)</td> + <td>(46.9)</td> + <td>(59.2)</td> +</tr> +<tr> + <td class="text_lf"><i>leprus</i></td> + <td>♂</td> + <td>14</td> + <td>20.6-26.4</td> + <td>42.3-52.3</td> + <td>35.0-40.3</td> + <td>47.5-62.5</td> + <td>58.2-72.5</td> +</tr> +<tr> + <td> </td> + <td> </td> + <td> </td> + <td>(46.8)</td> + <td>(37.4)</td> + <td>(56.5)</td> + <td>(67.3)</td> +</tr> +<tr> + <td> </td> + <td>♀</td> + <td>15</td> + <td>22.1-29.2</td> + <td>43.4-53.3</td> + <td>32.6-38.9</td> + <td>38.6-57.9</td> + <td>50.2-86.9</td> +</tr> +<tr> + <td> </td> + <td> </td> + <td> </td> + <td>(47.1)</td> + <td>(35.8)</td> + <td>(47.1)</td> + <td>(68.1)</td> +</tr> +<tr> + <td class="text_lf"><i>rubrimaculatus</i></td> + <td>♂</td> + <td>12</td> + <td>18.2-23.5</td> + <td>40.4-46.2</td> + <td>31.8-35.5</td> + <td>35.5-46.5</td> + <td>65.1-78.5</td> +</tr> +<tr> + <td class="brdbt2"> </td> + <td class="brdbt2"> </td> + <td class="brdbt2"> </td> + <td class="brdbt2"> </td> + <td class="brdbt2">(43.4)</td> + <td class="brdbt2">(33.8)</td> + <td class="brdbt2">(41.7)</td> + <td class="brdbt2">(71.7)</td> +</tr> +</table> +</div> +<br /> +<br /> + +<div class="center"> +<span class="pagenum"><a name="Page_8" id="Page_8">[Pg 8]</a></span> +<a name="Table_5" id="Table_5"></a> +<span class="smcap">Table 5</span>—Size and Proportions in the Frogs +of the <i>Syrrhophus longipes</i> and <i>S. marnockii</i> Groups.<br /> + +<table style="text-align:center; width:100%;" summary="Frog Sizes and Proportions"> +<tr> + <td class="brdtp2 brdbt">Species</td> + <td class="brdtp2 brdbt">Sex</td> + <td class="brdtp2 brdbt">N</td> + <td class="brdtp2 brdbt">Snout-vent<br />length<br />(SVL)</td> + <td class="brdtp2 brdbt">Tibia<br />length/<br />SVL</td> + <td class="brdtp2 brdbt">Head<br />width/<br />SVL</td> + <td class="brdtp2 brdbt">Tympanum/<br />Eye</td> + <td class="brdtp2 brdbt">Eyelid/<br />Interorbital</td> +</tr> +<tr> + <td class="text_lf"><i>dennisi</i></td> + <td>♂</td> + <td>16</td> + <td>22.8-28.4</td> + <td>43.9-49.7</td> + <td>35.3-41.2</td> + <td>53.9-64.2</td> + <td>55.3-74.0</td> +</tr> +<tr> + <td> </td> + <td> </td> + <td> </td> + <td> </td> + <td>(47.4)</td> + <td>(38.8)</td> + <td>(58.9)</td> + <td>(65.1)</td> +</tr> +<tr> + <td> </td> + <td>♀</td> + <td>10</td> + <td>25.9-32.0</td> + <td>46.3-50.8</td> + <td>35.6-40.3</td> + <td>50.6-58.7</td> + <td>58.1-70.9</td> +</tr> +<tr> + <td> </td> + <td> </td> + <td> </td> + <td> </td> + <td>(48.2)</td> + <td>(37.7)</td> + <td>(54.9)</td> + <td>(63.6)</td> +</tr> +<tr> + <td class="text_lf"><i>longipes</i></td> + <td>♂</td> + <td>22</td> + <td>22.1-33.2</td> + <td>45.8-51.7</td> + <td>38.7-44.4</td> + <td>61.1-87.2</td> + <td>61.5-83.0</td> +</tr> +<tr> + <td> </td> + <td> </td> + <td> </td> + <td>(48.4)</td> + <td>(41.8)</td> + <td>(72.0)</td> + <td>(72.0)</td> +</tr> +<tr> + <td> </td> + <td>♀</td> + <td>19</td> + <td>26.8-39.6</td> + <td>44.3-51.0</td> + <td>36.3-40.8</td> + <td>49.5-72.1</td> + <td>55.3-85.9</td> +</tr> +<tr> + <td> </td> + <td> </td> + <td> </td> + <td> </td> + <td>(47.2)</td> + <td>(39.1)</td> + <td>(59.5)</td> + <td>(67.9)</td> +</tr> +<tr> + <td class="text_lf"><i>guttilatus</i></td> + <td>♂</td> + <td>19</td> + <td>20.6-29.0</td> + <td>41.2-48.1</td> + <td>36.9-44.9</td> + <td>55.1-75.7</td> + <td>53.3-79.5</td> +</tr> +<tr> + <td> </td> + <td> </td> + <td> </td> + <td> </td> + <td>(44.5)</td> + <td>(40.6)</td> + <td>(64.1)</td> + <td>(66.0)</td> +</tr> +<tr> + <td> </td> + <td>♀</td> + <td> 5</td> + <td>25.7-31.0</td> + <td>41.4-46.8</td> + <td>35.9-42.3</td> + <td>47.6-61.7</td> + <td>62.3-79.8</td> +</tr> +<tr> + <td> </td> + <td> </td> + <td> </td> + <td> </td> + <td>(43.6)</td> + <td>(38.5)</td> + <td>(54.0)</td> + <td>(72.9)</td> +</tr> +<tr> + <td class="text_lf"><i>marnockii</i></td> + <td>♂</td> + <td>14</td> + <td>18.4-28.9</td> + <td>42.3-47.2</td> + <td>36.1-43.0</td> + <td>47.2-68.3</td> + <td>51.6-74.4</td> +</tr> +<tr> + <td> </td> + <td> </td> + <td> </td> + <td> </td> + <td>(44.1)</td> + <td>(39.6)</td> + <td>(61.2)</td> + <td>(66.3)</td> +</tr> +<tr> + <td> </td> + <td>♀</td> + <td>29</td> + <td>20.4-35.4</td> + <td>38.7-46.4</td> + <td>35.9-41.3</td> + <td>45.8-73.3</td> + <td>52.1-70.5</td> +</tr> +<tr> + <td> </td> + <td> </td> + <td> </td> + <td> </td> + <td>(42.7)</td> + <td>(38.2)</td> + <td>(60.3)</td> + <td>(60.7)</td> +</tr> +<tr> + <td class="text_lf"><i>verrucipes</i></td> + <td>♂</td> + <td>29</td> + <td>17.5-29.2</td> + <td>42.7-49.5</td> + <td>36.2-42.4</td> + <td>56.1-82.2</td> + <td>56.8-82.8</td> +</tr> +<tr> + <td> </td> + <td> </td> + <td> </td> + <td> </td> + <td>(46.3)</td> + <td>(39.1)</td> + <td>(67.8)</td> + <td>(70.4)</td> +</tr> +<tr> + <td> </td> + <td>♀</td> + <td> 6</td> + <td>26.5-31.7</td> + <td>42.4-47.7</td> + <td>36.0-38.1</td> + <td>45.8-57.8</td> + <td>61.0-77.9</td> +</tr> +<tr> + <td class="brdbt2"> </td> + <td class="brdbt2"> </td> + <td class="brdbt2"> </td> + <td class="brdbt2"> </td> + <td class="brdbt2">(44.6)</td> + <td class="brdbt2">(37.0)</td> + <td class="brdbt2">(53.9)</td> + <td class="brdbt2">(69.0)</td> +</tr> +</table> +</div> +<br /> +<br /> + +<div class="center"> +<a name="Table_6" id="Table_6"></a> +<span class="smcap">Table 6</span>—Size and Proportions in the Frogs +of the <i>Syrrhophus pipilans</i> and <i>S. modestus</i> Groups.<br /> + +<table style="text-align:center; width:100%;" summary="Frog Sizes and Proportions"> +<tr> + <td class="brdtp2 brdbt">Species</td> + <td class="brdtp2 brdbt">Sex</td> + <td class="brdtp2 brdbt">N</td> + <td class="brdtp2 brdbt">Snout-vent<br />length<br />(SVL)</td> + <td class="brdtp2 brdbt">Tibia<br />length/<br />SVL</td> + <td class="brdtp2 brdbt">Head<br />width/<br />SVL</td> + <td class="brdtp2 brdbt">Tympanum/<br />Eye</td> + <td class="brdtp2 brdbt">Eyelid/<br />Interorbital</td> +</tr> +<tr> + <td class="text_lf"><i>pipilans nebulosus</i></td> + <td>♂</td> + <td>17</td> + <td>22.9-28.5</td> + <td>38.1-42.0</td> + <td>34.4-37.2</td> + <td>36.6-47.8</td> + <td>56.1-82.4</td> +</tr> +<tr> + <td> </td> + <td>♀</td> + <td>♀</td> + <td>♀</td> + <td>(40.0)</td> + <td>(35.4)</td> + <td>(43.6)</td> + <td>(68.2)</td> +</tr> +<tr> + <td> </td> + <td>♀</td> + <td> 3</td> + <td>21.1-22.7</td> + <td>42.1-44.5</td> + <td>33.2-35.8</td> + <td>36.6-47.6</td> + <td>64.3-65.4</td> +</tr> +<tr> + <td class="text_lf"><i>pipilans pipilans</i></td> + <td>♂</td> + <td>18</td> + <td>22.6-27.8</td> + <td>37.9-44.0</td> + <td>32.2-36.5</td> + <td>38.0-54.0</td> + <td>56.1-79.5</td> +</tr> +<tr> + <td> </td> + <td> </td> + <td> </td> + <td> </td> + <td>(41.4)</td> + <td>(33.0)</td> + <td>(46.2)</td> + <td>(67.3)</td> +</tr> +<tr> + <td> </td> + <td>♀</td> + <td> 1</td> + <td>29.4</td> + <td>38.4</td> + <td>32.5</td> + <td>44.6</td> + <td>55.0</td> +</tr> +<tr> + <td class="text_lf"><i>modestus</i></td> + <td>♂</td> + <td> 8</td> + <td>15.8-20.1</td> + <td>38.5-42.6</td> + <td>32.1-38.1</td> + <td>26.8-39.3</td> + <td>57.0-86.9</td> +</tr> +<tr> + <td> </td> + <td> </td> + <td> </td> + <td> </td> + <td>(40.6)</td> + <td>(34.2)</td> + <td>(31.5)</td> + <td>(69.1)</td> +</tr> +<tr> + <td> </td> + <td>♀</td> + <td> 1</td> + <td>18.5</td> + <td>44.2</td> + <td>36.0</td> + <td>24.0</td> + <td>52.1</td> +</tr> +<tr> + <td class="text_lf"><i>pallidus</i></td> + <td>♂</td> + <td> 6</td> + <td>17.9-19.3</td> + <td>41.0-44.9</td> + <td>32.6-36.2</td> + <td>27.0-35.6</td> + <td>59.4-67.7</td> +</tr> +<tr> + <td> </td> + <td> </td> + <td> </td> + <td> </td> + <td>(43.4)</td> + <td>(35.2)</td> + <td>(30.9)</td> + <td>(65.2)</td> +</tr> +<tr> + <td class="text_lf"><i>teretistes</i></td> + <td>♂</td> + <td>18</td> + <td>19.2-23.2</td> + <td>41.5-45.3</td> + <td>32.5-36.4</td> + <td>28.6-43.8</td> + <td>51.2-75.0</td> +</tr> +<tr> + <td> </td> + <td> </td> + <td> </td> + <td> </td> + <td>(43.7)</td> + <td>(34.0)</td> + <td>(33.7)</td> + <td>(62.2)</td> +</tr> +<tr> + <td> </td> + <td>♀</td> + <td>1</td> + <td>24.8</td> + <td>41.8</td> + <td>30.8</td> + <td>37.9</td> + <td>60.5</td> +</tr> +<tr> + <td class="text_lf"><i>nivocolimae</i></td> + <td>♂</td> + <td>15</td> + <td>18.9-21.1</td> + <td>42.2-48.6</td> + <td>30.9-37.1</td> + <td>30.0-39.3</td> + <td>42.6-69.1</td> +</tr> +<tr> + <td> </td> + <td> </td> + <td> </td> + <td> </td> + <td>(45.0)</td> + <td>(33.7)</td> + <td>(34.7)</td> + <td>(55.0)</td> +</tr> +<tr> + <td> </td> + <td>♀</td> + <td> 1</td> + <td>24.1</td> + <td>40.9</td> + <td>33.5</td> + <td>27.6</td> + <td>56.5</td> +</tr> +<tr> + <td class="text_lf"><i>interorbitalis</i></td> + <td>♂</td> + <td> 1</td> + <td>25.6</td> + <td>43.0</td> + <td>——</td> + <td>39.4</td> + <td>57.6</td> +</tr> +<tr> + <td> </td> + <td>♀</td> + <td> 9</td> + <td>20.2-26.7</td> + <td>39.9-47.1</td> + <td>32.6-39.3</td> + <td>29.1-41.2</td> + <td>58.2-76.9</td> +</tr> +<tr> + <td class="brdbt2"> </td> + <td class="brdbt2"> </td> + <td class="brdbt2"> </td> + <td class="brdbt2"> </td> + <td class="brdbt2">(43.2)</td> + <td class="brdbt2">(35.8)</td> + <td class="brdbt2">(36.4)</td> + <td class="brdbt2">(69.2)</td> +</tr> +</table> +</div> +<br /> +<br /> + +<p><span class="pagenum"><a name="Page_9" id="Page_9">[Pg 9]</a></span></p> + +<div class="fig_center" style="width: 451px;"> +<a name="Fig_1" id="Fig_1"></a> +<img src="images/fig_1.png" width="451" height="653" alt="" title="" /> +<div class="fig_caption"><span class="smcap">Fig. 1:</span> Palmar views of hands of six species of the eastern complex of +<i>Syrrhophus</i>. (A) <i>verrucipes</i> (UIMNH 15995), (B) <i>rubrimaculatus</i> (KU +58911), (C) <i>dennisi</i> sp. nov. (holotype, UMMZ 101121), (D) <i>guttilatus</i> +(UIMNH 55520), (E) <i>marnockii</i> (TCWC 4782), and (F) <i>longipes</i> (TCWC +12179). All ×6.5.</div> +</div> + +<p><span class="pagenum"><a name="Page_10" id="Page_10">[Pg 10]</a></span></p> +<div class="fig_center" style="width: 497px;"> +<a name="Fig_2" id="Fig_2"></a> +<img src="images/fig_2.png" width="497" height="218" alt="" title="" /> +<div class="fig_caption"><span class="smcap">Fig. 2:</span> Palmar views of hands of two species of the western complex of +<i>Syrrhophus</i>. <i>pipilans</i> (left, KU 58908, ×6) and <i>teretistes</i> (center, +KU 75269, and right, KU 75263, respectively, ×9).</div> +</div> + +<p>In <i>S. cystignathoides</i> and <i>leprus</i>, the first finger is longer than +the second, and the first two fingers are equal in length in +<i>guttilatus</i> and <i>marnockii</i>. In the other species the first finger is +shorter than the second.</p> + +<p>Supernumerary tubercles are well developed on the plantar surfaces in +all species, except <i>S. guttilatus</i>, in which they are poorly defined +(<a href="#Fig_3">Fig. 3</a>). The relative sizes of the metatarsal tubercles has been used +in the classification of the species and species groups of <i>Syrrhophus</i>. +The metatarsal tubercles are similar in all species of the eastern +complex (including <i>rubrimaculatus</i>); the outer tubercle is always about +one-half the size of the ovoid inner metatarsal tubercle. In the +<i>leprus</i> group the outer tubercle is conical and compressed. The +metatarsal tubercles of <i>pipilans</i> are about the same size, or the outer +is slightly smaller than the inner. In the <i>modestus</i> group the outer +metatarsal tubercle is about one-third the size of the inner.</p> + +<p>All species, except <i>guttilatus</i>, have well-defined to poorly defined +lateral fringes on the toes. All species have expanded toe pads. The +fifth toe is usually shorter than the third, but the second is equal in +length to the fifth in some specimens of <i>S. cystignathoides</i> and <i>S. +marnockii</i>. <i>Syrrhophus nivocolimae</i> is the only species with tubercles +along the outer edge of the tarsus; this is merely a reflection of the +highly tuberculate nature of the skin in this species.</p> + +<p><i>Skin texture.</i>—The skin of the dorsum is smooth or very weakly +pustular in all species of the genus except <i>nivocolimae</i> and +<i>verrucipes</i>. The dorsal surfaces of <i>nivocolimae</i> are warty; in +<i>verrucipes</i> the skin is pustular. The skin of the venter is areolate in +<i>cystignathoides +<span class="pagenum"><a name="Page_11" id="Page_11">[Pg 11]</a></span> +cystignathoides</i>, <i>dennisi</i> and <i>verrucipes</i> but is +smooth in all other species of the genus.</p> + +<div class="fig_center" style="width: 518px;"> +<a name="Fig_3" id="Fig_3"></a> +<img src="images/fig_3.png" width="518" height="680" alt="" title="" /> +<div class="fig_caption"><span class="smcap">Fig. 3:</span> Plantar views of feet of four species of the eastern complex of +<i>Syrrhophus</i>. (A) <i>guttilatus</i> (UIMNH 55519, ×6), (B) <i>leprus</i> (UIMNH +42726, ×6), (C) <i>verrucipes</i> (UIMNH 15995, ×6), and (D) <i>longipes</i> (TCWC +12179, ×4.6).</div> +</div> + +<p><i>Color pattern.</i>—As is evident in the diagnoses, the color patterns +of given populations have been regarded as useful in separating the +<span class="pagenum"><a name="Page_12" id="Page_12">[Pg 12]</a></span> +species and subspecies. Duellman (1958) suggested that the coloration, +with the exception of <i>modestus</i>, was a dark ground color with +pale markings. It is a moot point whether the frogs have light spots +on a dark background or have a light background with an extensive +reticulate dark pattern. The venters are gray or white, and the +vocal sac is nearly black in some species. Interorbital dark bars or +triangles are absent in only two species of the eastern complex, +<i>cystignathoides campi</i> and <i>marnockii</i>; the latter lacks a supratympanic +stripe, which is present in the other members of the eastern +complex. <i>Syrrhophus interorbitalis</i> and <i>nivocolimae</i> have light +<ins title='Correction: was "interorbtal"'>interorbital</ins> +bars; these bars occur in only one other population of the +genus (<i>S. c. cystignathoides</i>). Bars on the thighs are ill defined or +absent in the members of the <i>marnockii</i> and part of the <i>modestus</i> +groups. The color in life is noted in the species accounts.</p> + +<p><i>Voice.</i>—The voices of all <i>Syrrhophus</i> can be described as a +single short chirp or peep; without audiospectrographic analyses +the significance of the differences between a chirp, peep, or short +whistle cannot be appreciated. Martin (1958) and Wright and +Wright (1949) reported multi-noted calls, and one collector of <i>S. +verrucipes</i> noted the frog "trilled."</p> + +<p>Fouquette (1960) presented analyses of two species (<i>marnockii</i> +and <i>pipilans nebulosus</i>). The voices were very similar; both frogs +were reported to "trill" and "chirp."</p> +<br /> + +<div class="smcap caption2nb">Systematic Account</div> + +<p>The genus <i>Syrrhophus</i> has been defined (Lynch, 1968) and +limited to the group of species occurring in Guatemala, México and +the United States. The closest relatives of <i>Syrrhophus</i> are the frogs +of the genus <i>Tomodactylus</i> (Dixon, 1957; Firschein, 1954). Lynch +(1968) implied there were no osteological bases for the separation +of <i>Eleutherodactylus</i>, <i>Syrrhophus</i>, and <i>Tomodactylus</i>. At that time, +I believed such to be the case and derived <i>Syrrhophus</i> and <i>Tomodactylus</i> +from the <i>rhodopis</i> complex of <i>Eleutherodactylus</i>, with +which they share terrestrial habits and relatively short limbs. In the +<i>rhodopis</i> complex there is a tendency for the loss of the outer palmar +tubercle, a not uncommon condition in <i>Syrrhophus</i> and <i>Tomodactylus</i>.</p> + +<p>However, the skulls of <i>Syrrhophus</i> and <i>Tomodactylus</i> show departures +from the pattern observed in the Middle American <i>Eleutherodactylus</i>, +as well as many of those species in western South +America. Baldauf and Tanzer (1965) reported that the frontoparietals +<span class="pagenum"><a name="Page_13" id="Page_13">[Pg 13]</a></span> +and prootics were fused in <i>Syrrhophus marnockii</i> and that +the prootics and exoccipitals appeared to be one bone (otoccipital). +The otoccipital is not uncommon in eleutherodactyline frogs, but +the fusion of the frontoparietals with the prootics (regardless of the +fusion of the latter with the exoccipital) is uncommon in the family. +I have found the frontoparietal-prootic fusion only in <i>Syrrhophus</i> +(all species), <i>Tomodactylus</i> (all species), and <i>Eleutherodactylus</i> +(West Indies species). None of the Middle American <i>Eleutherodactylus</i> +has the two bones fused. Examination of the character is +difficult in dried skeletal preparations. Cleared and stained or +macerated preparations are satisfactory for checking this character.</p> + +<p>Thus, in addition to the presence of numerous plantar supernumerary +tubercles in the frogs of the genera <i>Syrrhophus</i> and <i>Tomodactylus</i>, +these two genera can be separated from other Middle +American eleutherodactylines by the fusion of the frontoparietals +and prootics. This character not only further strengthens the argument +that the two genera are closely related but poses a problem +of zoogeographic analysis of the distribution of the character, which +will be discussed fully elsewhere.</p> +<br /> + +<div class="caption3">Key to the Species of the Frog Genus <i>Syrrhophus</i></div> +<table width="100%" summary="Species Key"> +<tr> + <td class="vtop"> 1. </td> + <td>Three large, well-developed palmar tubercles</td> + <td class="text_rt">2</td> +</tr> +<tr class="row_color1"> + <td> </td> + <td>Two large palmar tubercles; outer (third) palmar tubercle reduced + in size or absent</td> + <td class="text_rt">9</td> +</tr> +<tr> + <td class="vtop"> 2. </td> + <td>Digital pads more than twice (usually three or more) times width of + digit</td> + <td class="text_rt">3</td> +</tr> +<tr class="row_color1"> + <td> </td> + <td>Digital pads less than twice width of digit</td> + <td class="text_rt">4</td> +</tr> +<tr> + <td class="vtop"> 3. </td> + <td>Males having vocal slits; dorsum vermiculate; diameter of tympanum + in males about one-half diameter of eye</td> + <td class="text_rt"><a href="#Syrrhophus_dennisi"><i>S. dennisi</i></a></td> +</tr> +<tr class="row_color1"> + <td> </td> + <td>Males lacking vocal slits; dorsum flecked, spotted, or blotched; diameter + of tympanum in male about three-fourths that of eye</td> + <td class="text_rt"><a href="#Syrrhophus_longipes"><i>S. longipes</i></a></td> +</tr> +<tr> + <td class="vtop"> 4. </td> + <td>First finger longer than second</td> + <td class="text_rt">5</td> +</tr> +<tr class="row_color1"> + <td> </td> + <td>First finger shorter than or equal to second</td> + <td class="text_rt">7</td> +</tr> +<tr> + <td class="vtop"> 5. </td> + <td>Venter smooth; dorsum spotted or vermiculate</td> + <td class="text_rt"><a href="#Syrrhophus_leprus"><i>S. leprus</i></a></td> +</tr> +<tr class="row_color1"> + <td> </td> + <td>Venter areolate, or if smooth, dorsum flecked and interorbital bar + lacking</td> + <td class="text_rt"> 6</td> +</tr> +<tr> + <td class="vtop"> 6. </td> + <td>Venter areolate; interorbital bar present; ground color yellowish</td> + <td class="text_rt"><a href="#Syrrhophus_cystignathoides_cystignathoides"><i>S. cystignathoides cystignathoides</i></a></td> +</tr> +<tr class="row_color1"> + <td> </td> + <td>Venter smooth; interorbital bar absent; ground color brown</td> + <td class="text_rt"><a href="#Syrrhophus_cystignathoides_campi"><i>S. cystignathoides campi</i></a></td> +</tr> +<tr> + <td class="vtop"> 7. </td> + <td>First finger shorter than second; digital tips only slightly dilated; green + in life with darker green spots</td> + <td class="text_rt"><a href="#Syrrhophus_verrucipes"><i>S. verrucipes</i></a></td> +</tr> +<tr class="row_color1"> + <td> </td> + <td>First finger equal to second; digital tips slightly to moderately expanded</td> + <td class="text_rt"> 8</td> +</tr> +<tr> + <td class="vtop"> 8. </td> + <td>Dorsum vermiculate; interorbital bar present; ground color cream to + brown in life + <span class="pagenum"><a name="Page_14" id="Page_14">[Pg 14]</a></span></td> + <td class="text_rt"><a href="#Syrrhophus_guttilatus"><i>S. guttilatus</i></a></td> +</tr> +<tr class="row_color1"> + <td> </td> + <td>Dorsum punctate or flecked; interorbital bar absent; ground color green + in life</td> + <td class="text_rt"><a href="#Syrrhophus_marnockii"><i>S. marnockii</i></a></td> +</tr> +<tr> + <td class="vtop"> 9. </td> + <td>Dorsum dark with pale (red in life) spots; digital pads not expanded</td> + <td class="text_rt"><a href="#Syrrhophus_rubrimaculatus"><i>S. rubrimaculatus</i></a></td> +</tr> +<tr class="row_color1"> + <td> </td> + <td>Dorsum pale with dark markings and digital pads slightly to widely + expanded</td> + <td class="text_rt">10</td> +</tr> +<tr> + <td class="vtop">10. </td> + <td>Digital tips not widely expanded; tympanum well-defined; outer metatarsal + tubercle more than one-half size of inner</td> + <td class="text_rt">11</td> +</tr> +<tr class="row_color1"> + <td> </td> + <td>Digital tips widely expanded, truncate in outline; tympanum poorly defined; + outer metatarsal tubercle less than one-half size of inner</td> + <td class="text_rt">12</td> +</tr> +<tr> + <td class="vtop">11. </td> + <td>Dorsum dark brown with large light spots or blotches; tympanum/eye + ratio usually greater than 43 percent</td> + <td class="text_rt"><a href="#Syrrhophus_pipilans_pipilans"><i>S. pipilans pipilans</i></a></td> +</tr> +<tr class="row_color1"> + <td> </td> + <td>Dorsum dark brown with small light spots; tympanum/eye ratio less + than 48 percent</td> + <td class="text_rt"><a href="#Syrrhophus_pipilans_nebulosus"><i>S. pipilans + <ins title='Correction: was "neublosus"'>nebulosus</ins></i></a></td> +</tr> +<tr> + <td class="vtop">12. </td> + <td>Light interorbital bar present</td> + <td class="text_rt">13</td> +</tr> +<tr class="row_color1"> + <td> </td> + <td>Light interorbital bar absent</td> + <td class="text_rt">14</td> +</tr> +<tr> + <td class="vtop">13. </td> + <td>Adults small, less than 22 mm. snout-vent length with a broad mid-dorsal + stripe; dark bands on shank narrower than light interspaces</td> + <td class="text_rt"><a href="#Syrrhophus_nivocolimae"><i>S. nivocolimae</i></a></td> +</tr> +<tr class="row_color1"> + <td> </td> + <td>Adults larger, more than 22 mm. snout-vent length; dorsum vermiculate; + dark bands on shank broader than light interspaces</td> + <td class="text_rt"><a href="#Syrrhophus_interorbitalis"><i>S. interorbitalis</i></a></td> +</tr> +<tr> + <td class="vtop">14. </td> + <td>Dorsum spotted with discrete black spots; pattern definite</td> + <td class="text_rt"><a href="#Syrrhophus_modestus"><i>S. modestus</i></a></td> +</tr> +<tr class="row_color1"> + <td> </td> + <td>Dorsum reticulate or vermiculate, pattern poorly defined</td> + <td class="text_rt">15</td> +</tr> +<tr> + <td class="vtop">15. </td> + <td>Adults small, less than 21 mm. snout-vent length; upper arm not + banded</td> + <td class="text_rt"><a href="#Syrrhophus_pallidus"><i>S. pallidus</i></a></td> +</tr> +<tr class="row_color1"> + <td> </td> + <td>Adults larger, usually greater than 21 mm. snout-vent length; upper + arm banded</td> + <td class="text_rt"><a href="#Syrrhophus_teretistes"><i>S. teretistes</i></a></td> +</tr> +</table> +<br /> +<br /> + +<div class="smcap caption2nb">Species Accounts</div> + +<p>The following accounts do not include complete descriptions of each +taxon, because a more than adequate number of descriptions is available +in the recent (1940-1966) literature. An abbreviated synonymy, in which +are listed all combinations and emendations of names and significant +contributions to our knowledge of the taxon, is given for each. For each +species and subspecies the following are given: descriptive diagnosis, +statement of range, remarks on taxonomy, list of specimens examined, +illustration of color pattern, and distribution map.</p> +<br /> + +<div class="caption3nb"> +<a name="Syrrhophus_cystignathoides" id="Syrrhophus_cystignathoides"></a> +<b>Syrrhophus cystignathoides</b> (Cope)</div> + +<div class="species_ref"><i>Phyllobates cystignathoides</i> Cope, 1877:89-90 +[Syntypes.—Originally USNM 32402-32409, (32405 now in MCZ) from +Potrero, near Córdoba, Veracruz, México, Francis Sumichrast +collector.]</div> +<p><span class="pagenum"><a name="Page_15" id="Page_15">[Pg 15]</a></span></p> +<p><i>Diagnosis.</i>—Adults small, males 16.0 to 23.5 mm. in snout-vent length, +females 16.0-25.8 mm. in snout-vent length; vocal slits present in +males; finger tips slightly expanded; first finger longer than second; +outer metatarsal tubercle one-half size of inner, conical, compressed; +skin of dorsum weakly pustular, that of venter smooth to areolate; +tympanum 44 to 69 per cent diameter of eye (mean 55.5 per cent); ground +color yellow to brown in life with brown to black fleckings on dorsum +and flanks; limbs banded; interorbital bar present or not.</p> + +<p><i>Remarks.</i>—Two geographic races (subspecies) are herein recognized; +previously these were held by various authors to be species (<i>campi</i> and +<i>cystignathoides</i>). Intergradation occurs in southern Tamaulipas and +eastern San Luis Potosí, México. The two subspecies can be distinguished +on the basis of color pattern and the condition of the skin of the +venter.</p> + +<p><i>Distribution.</i>—Low to moderate elevations from the Río Grande +embayment to central Veracruz, México (<a href="#Fig_5">Fig. 5</a>).</p> +<br /> + +<div class="caption3nb"> +<a name="Syrrhophus_cystignathoides_campi" id="Syrrhophus_cystignathoides_campi"></a> +<b>Syrrhophus cystignathoides campi</b> Stejneger, New combination</div> + +<div class="species_ref"><i>Syrrhophus campi</i> Stejneger, 1915:131-32. [Holotype.—USNM 52290, from +Brownsville, Cameron Co., Texas; R. D. Camp collector, March 31, 1915]. +Smith and Taylor, 1948:52. Martin, 1958:50.</div> + +<p><i>Diagnosis.</i>—Venter smooth; usually no interorbital light and dark bars +present; ground color brown in life (<a href="#Fig_4">Fig. 4a</a>).</p> + +<p><i>Remarks.</i>—Martin (1958) was the first author to point out that <i>S. +campi</i> was probably a subspecies of the more southern <i>S. +cystignathoides</i>. Various references in the literature might lead one to +believe that the two were sympatric over much of northeastern México; +this error was created by the use of a single character (condition of +the skin of the venter) to characterize the two populations. Specimens +from southern Texas have a smooth venter, lack interorbital bars and +have, in general, a brown ground color, whereas specimens from central +Veracruz have an areolate venter, interorbital light and dark bars and a +yellow ground color. In southern Tamaulipas and eastern San Luis Potosí, +these characters vary discordantly, thereby strongly suggesting that the +two populations intergrade. Both populations agree in other +morphological characters; therefore, they are here treated as geographic +variants.</p> + +<p><i>Etymology.</i>—Named for the collector of the type specimens, Mr. R. D. +Camp of Brownsville, Texas.</p> + +<p><i>Distribution.</i>—Lower Río Grande embayment in Texas to central Nuevo +León and Tamaulipas, México. Intergrades are known from southern +Tamaulipas and adjacent San Luis Potosí, México (Fig. 5).</p> + +<p><i>Specimens examined.</i>—(113) TEXAS, Cameron Co.: MCZ 10277-85, 10286 +(10); Brownsville, AMNH 3215, 3218-20, 3221 (3), 5376, 62117, FMNH +105336, KU 8135-39, MCZ 3738-42, 3743 (10), TCWC 5908, 7139, TNHC 92-94, +20909, UMMZ 51760, 54031 (5), USNM 52290 (holotype); 22 mi. SE +Brownsville, TNMC 14223; 8 mi. SW Brownsville, UMMZ 101127 (3); +Harlingen, AMNH 62118, UMMZ 105200-205, 105206 (5), 105207 (4). <i>Hidalgo +Co.</i>: Bentsen-Río Grande State Park, UMMZ 114378; 6 mi. S McAllen, TNHC +7136-39; Santa Ana Refuge, TCWC 13495-96; Weslaco, TCWC 17658-60.</p> + +<p>MEXICO, <i>Nuevo León</i>: Salto Cola de Caballo, AMNH 57953-54, FMNH +30644-45, 37169-70; Monterrey, UIMNH 13324; 40 km. SE Monterrey, UIMNH +3686. <i>Tamaulipas</i>: 80 km. Matamoros, FMNH 27150 (13).</p> + +<p><span class="pagenum"><a name="Page_16" id="Page_16">[Pg 16]</a></span></p> + +<div class="fig_center" style="width: 516px;"> +<a name="Fig_4" id="Fig_4"></a> +<img src="images/fig_4.png" width="516" height="445" alt="" title="" /> +<div class="fig_caption"><span class="smcap">Fig. 4:</span> <i>Syrrhophus cystignathoides campi</i> (left, TCWC +13490) and <i>S. c. cystignathoides</i> (right, KU 105500). Dorsal views ×2, +sides of heads ×3.</div> +</div> + +<p>Intergrades [<i>S. c. cystignathoides</i> × <i>S. c. campi</i> (88)] MÉXICO, <i>San +Luis Potosí</i>: 5 km. E <ins title='Correction: was "Cuidad"'>Ciudad</ins> del Maiz, UMMZ 106435; 16 km. W +Naranjo, FMNH 104584; Salto de Agua, 34 km. WSW Antigua Morelos, TCWC +6980. <i>Tamaulipas</i>: 5 km. W Acuña, 1060 m., UMMZ 101172, 101173 (16), +101174-76, 101177 (6); 14.5 km. NNW Chamal, 430 m., UMMZ 111337 (2); 20 +km. NNW Chamal, 700 m., UMMZ 111338 (11); 8 km. N Gómez Farías, 450 m., +UMMZ 101165; 8 km. NE Gómez Farías, Pano Ayuctle, UMMZ 102264, 102924 +(6); 8 km. NW Gómez Farías, 1060 m., LSUMZ 11084, UMMZ 101199, 102928 +(5), 102929-32, 110124 (3); Río Guayala, near Magiscatzin, MCZ 24138-42, +85071-81, UMMZ 88242 (2); Magiscatzin, TCWC 6981; Las Yucas, north of +Aldama, MCZ 29665-68; 16 km. NE Zamorina, UMMZ 101124.</p> +<br /> + +<div class="caption3nb"> +<a name="Syrrhophus_cystignathoides_cystignathoides" id="Syrrhophus_cystignathoides_cystignathoides"></a> +<b>Syrrhophus cystignathoides cystignathoides</b> (Cope), New combination</div> + +<div class="species_ref"><i>Phyllobates cystignathoides</i> Cope, 1877:89-90 [Syntypes.—USNM +32402-32409, from Potrero, near Córdoba, Veracruz, México, collected by +Francis Sumichrast]. Boulenger, 1882:196.</div> +<div class="species_ref"><i>Syrrhophus cystignathoides</i>: Cope, 1879:268. Kellogg, 1932: 126-27. +Taylor and Smith, 1945: 582-83. Smith and Taylor, 1948:50. Martin, +1958:49.</div> +<div class="species_ref"><i>Syrrhaphus cystignathoides</i>: Günther, 1900:218.</div> +<div class="species_ref"><i>Syrraphus cystignathoides</i>: Díaz de León, 1904:10.</div> +<div class="species_ref"><i>Syrrhopus cystignathoides</i>: Barbour and Loveridge, 1946-170[** +1946:170].</div> +<br /> + +<p><span class="pagenum"><a name="Page_17" id="Page_17">[Pg 17]</a></span></p> + +<div class="fig_center" style="width: 502px;"> +<a name="Fig_5" id="Fig_5"></a> +<img src="images/fig_5.png" width="502" height="659" alt="" title="" /> +<div class="fig_caption"><span class="smcap">Fig. 5:</span> Distribution of <i>Syrrhophus cystignathoides campi</i> (solid +symbols) and the nominate subspecies (open symbols).</div> +</div> + +<p><i>Diagnosis.</i>—Venter areolate; interorbital light and dark bars present; +ground color yellow to brownish-yellow in life (<a href="#Fig_4">Fig. 4b</a>).</p> + +<p><i>Remarks.</i>—Firschein (1954) briefly considered the status of Peters' +(1871) <i>Phyllobates verruculatus</i> and noted that if it was a +<i>Syrrhophus</i> it would probably be referrable to <i>S. cystignathoides</i>. +Peters' (1871) original description corresponds well with <i>S. +cystignathoides</i>, and the type-locality ("Huanusco" = Huatusco) is +within the range of that species. Firschein (1954) expressed +<span class="pagenum"><a name="Page_18" id="Page_18">[Pg 18]</a></span> doubt that +<i>verruculatus</i> was a <i>Syrrhophus</i>, because Peters placed it in another +genus. However, Peters described <i>verruculatus</i> a decade before Cope +diagnosed the genus Syrrhophus. Most frogs now called <i>Syrrhophus</i>, plus +a number of lower Central American frogs now placed in a variety of +genera were placed in <i>Phyllobates</i> by Boulenger, Cope, and Peters.</p> + +<p>The types of <i>Phyllobates verruculatus</i> were destroyed during World War +II (Günther Peters, <i>in litt.</i>); the specimens subsequently assigned to +the taxon by Kellogg (1932) are <i>Syrrhophus cystignathoides</i>. Because +the type specimens are lost and because the name antedates the more +established name, <i>cystignathoides</i>, I favor retaining <i>Phyllobates +verruculatus</i> Peters as a <i>nomen dubium</i>.</p> + +<p>Smith and Taylor (1948) reported <i>S. verruculatus</i> from Tianguistengo, +Hidalgo, México. These specimens are examples of <i>verrucipes</i>. Smith +(1947) reported a specimen of <i>verruculatus</i> from San Lorenzo, Veracruz. +Firschein (1954) referred it to <i>cystignathoides</i>, and Duellman (1960) +concluded that both authors were in error and that the specimen (USNM +123530) was a <i>leprus</i>.</p> + +<p><i>Etymology.</i>—The trivial name is the diminutive of <i>Cystignathus</i>, a +once-used generic name for several leptodactylid frogs.</p> + +<p><i>Distribution.</i>—Low and moderate elevations in the foothills along the +Sierra Madre Oriental from eastern San Luis Potosí to Central Veracruz, +México (<a href="#Fig_5">Fig. 5</a>).</p> + +<p><i>Specimens examined.</i>—(130), MÉXICO, <i>Puebla</i>: Necaxa, UMMZ 69519-20. +<i>San Luis Potosí</i>: 5 km. W Aguismón, LSUMZ 4962-63; along Río Axtla, +road to Xilitla, UMMZ 105500; Tamazunchale, UIMNH 3199; 6.5 km. N +Tamazunchale, UMMZ 104039; 8 km. N Tamazunchale, UMMZ 119490. +<i>Veracruz</i>: Coatepec, 1210 m., FMNH 704966-67; 11 km. SE Coatepec, 850 +m., FMNH 70468-70; below Córdoba, FMNH 104588, UIMNH 13321; Cuautlapam, +1000 m., FMNH 106477-80, KU 100364, UIMNH 58200-03, UMMZ 105392; Fortín +de las Flores, UIMNH 13322, 13339; 1.6 km. N Fortín de las Flores, UIMNH +42799-808, UMMZ 105389; 3.2 km. N Fortín de las Flores, UIMNH 26633-35; +4.8 km. N Fortín de las Flores, UIMNH 71967-68; 3.2 km. W Fortín de las +Flores (Barranca Metlac), 910 m., UIMNH 49294-95, UMMZ 115444-46, +118221, 119893 (2); Huatusco, KU 100363; Jalapa, 1400 m., FMNH 70440, +70443-51, 70454-65; 16 km. NE Jalapa, 1300 m., FMNH 70452-53; 8 km. E +Jalapa, UIMNH 13338; 9.5 km. S Jalapa, UMMZ 122083 (2); Mirador, KU +23967; Paraja Nuevo, El Suchil, UMMZ 85490 (7), 85491 (2), 90315; La +Passa, UIMNH 49293, 49297; 1 km. E Plan del Río, 240 m., UMMZ 102067 +(2); Potrero Viejo, FMNH 104583, 104586, 105326-27, KU 26789, 100357-62, +UIMNH 13323, 13340-43; USNM 32402 (lectotype), 32403-04, 32406-09; 9.6 +km. S Santa Rosa, TCWC 12785; 24 km. NE Tezuitlán (Puebla), UMMZ 105388; +Teocelo, FMNH 70437-38, KU 26080, 26790; 3.2 km. N Teocelo, FMNH 70439, +70441-42; 9.6 km. NW Tihuatlán, UIMNH 3684-85; 15 km. ENE Tlacotepec, KU +23966; 26 km. NW Tuxpan, UMMZ 126419.</p> +<br /> + +<div class="caption3nb"> +<a name="Syrrhophus_leprus" id="Syrrhophus_leprus"></a> +<b>Syrrhophus leprus</b> Cope</div> + +<div class="species_ref"><i>Syrrhophus leprus</i> Cope, 1879:268-69 [Holotype.—USNM 10040, from Santa +Efigena, Oaxaca, México, Francis Sumichrast collector]. Kellogg, +1932:124-5, 128. Taylor and Smith, 1945:582. Smith and Taylor, +1948:50-51. Duellman, 1958:8, pl. 1, Fig. 2; 1960:56-57. Gorham, +1966:165.</div> +<div class="species_ref"><i>Syrrhaphus leprus</i>: Günther, 1900:217.</div> +<div class="species_ref"><i>Syrrhophus leprus leprus</i>: Neill, 1965:85-86.</div> +<div class="species_ref"><i>Syrrhophus leprus cholorum</i> Neill, 1965:85-86 [Holotype.—Wilfred T. +Neill collection 1525, from 3.9 mi. N San Antonio, Toledo District, +British Honduras, +collected October 28, 1959, by R. A. Allen, T. C. +Allen, and W. T. Neill].</div> + +<p><span class="pagenum2"><a name="Page_19" id="Page_19">[Pg 19]</a></span><i>Diagnosis.</i>—Medium-sized frogs, males 20.5-26.5 mm. in snout-vent, +females 22.0-29.3 mm. in snout-vent length; vocal slits present in +males; tips of fingers dilated slightly; first finger longer than +second; inner metatarsal tubercle twice size of small, conical outer +metatarsal tubercle; skin of dorsum pustular, that of venter smooth; +snout subacuminate; diameter of tympanum 47.5-62.5 per cent of eye in +males, 38.6-57.9 per cent in females; dorsum yellowish-green with +chocolate brown blotches or spots forming reticulations in most +specimens; venter white to gray; flanks brown, spotted with white or +not; limbs banded; interorbital bar obscured by dorsal pattern.</p> + + +<div class="fig_center" style="width: 493px;"> +<a name="Fig_6" id="Fig_6"></a> +<img src="images/fig_6.png" width="487" height="581" alt="" title="" /> +<div class="fig_caption"><span class="smcap">Fig. 6:</span> Dorsal views of <i>Syrrhophus leprus</i> showing variation in dorsal +pattern (left, UMMZ 121244, ×2; right, KU 26106, ×1.7). Side of head +(UIMNH 42726, ×7).</div> +</div> + +<p><span class="pagenum"><a name="Page_20" id="Page_20">[Pg 20]</a></span></p> +<div class="fig_center" style="width: 487px;"> +<a name="Fig_7" id="Fig_7"></a> +<img src="images/fig_7.png" width="493" height="293" alt="" title="" /> +<div class="fig_caption"><span class="smcap">Fig. 7:</span> Distribution of three species of eastern complex <i>Syrrhophus</i>: +<i>leprus</i> (circles), <i>rubrimaculatus</i> (triangles), and <i>verrucipes</i> +(squares).</div> +</div> + +<p><i>Remarks.</i>—My distribution map (<a href="#Fig_7">Fig. 7</a>) differs somewhat from that of +Duellman (1958), who was unaware of specimens reported by Taylor and +Smith (1945) from central Veracruz, México.</p> + +<p>Duellman (1958, 1960) regarded <i>S. leprus</i> as having a gray venter. +Neill (1965) characterized his new subspecies on the basis of white +venter and spots on the dorsum. Some specimens from throughout the range +have only small round spots, instead of vermiculations (<a href="#Fig_6">Fig. 6</a>). The +gray ventral coloration is largely restricted to the population in Los +Tuxtlas, Veracruz, but only about 80 per cent of the specimens from the +Los Tuxtlas have gray venters, whereas specimens from Guatemala, Oaxaca, +Tabasco, and central Veracruz, México, have white venters (rarely gray). +Since the specimens from British Honduras are not distinct from +specimens throughout most of the range, there is no reason to recognize +them as a subspecies.</p> + +<p><i>Etymology.</i>—Greek, <i>lepra</i>, leprosy, in reference to the mottled color +pattern.</p> + +<p><i>Distribution.</i>—Discontinuous; central Veracruz to British Honduras to +low elevations in the foothills of the Sierra Madre Oriental, Los +Tuxtlas, Sierra Madre de Chiapas (Isthmus of Tehuantepec (<a href="#Fig_7">Fig. 7</a>)).</p> + +<p><i>Specimens examined.</i>—(84). GUATEMALA, <i>Alta Verapaz</i>: Chinajá, KU +55961-62. <i>El Petén</i>: 15 km. NW Chinajá, KU 55963; Piedras Negras, USNM +114085-92; Tikal, UMMZ 117035; Uaxactún, AMNH 55121-22.</p> + +<p>MÉXICO, <i>Oaxaca</i>: Cerro San Pedro del Isthmo, UIMNH 35510; Finca La +Gloria, USNM 114093; 30.5 km. N Matías Romero, UIMNH 39459, 71969; Santa +Efigenia, USNM 10040 (holotype). <i>Tabasco</i>: Teapa, UMMZ 113799-800; 13.5 +km. W Teapa, UMMZ 120253. <i>Veracruz</i>: 27.5 km. N Acayucan, UIMNH 42726; +Atoyac, UIMNH 13331, 49296; 3.2 km. N Catemaco, UIMNH 71976-77; Coyame, +UIMNH 38995, 38998, 40342; Dos Amates, TCWC 21211; Fortín de Las Flores, +FMNH 113751, 113753; Paraja Nuevo, El Suchil, UMMZ 90315; Potrero Viejo, +FMNH 113743-50, 126114-18, KU 26104-06, UIMNH 13332-37, UMMZ 88837; San +Andrés Tuxtla, UIMNH 27123-31, 28611, 71975, UMMZ 115450 (5); San +Lorenzo, USNM 123530; 4.5 km. NW Santiago Tuxtla, JDL 992 (skeleton), +UIMNH 27122; 32 km. S Sayula, EAL 1696; Tepalapan, 1.6 +<span class="pagenum"><a name="Page_21" id="Page_21">[Pg 21]</a></span> +km. S Catemaco, UMMZ 118222 (2); Volcán San Martín, south slope, +UMMZ 118223; Volcán San Martín, Rancho El Tular, UIMNH 35399-400, 40340-41.</p> +<br /> + +<div class="caption3nb"> +<a name="Syrrhophus_rubrimaculatus" id="Syrrhophus_rubrimaculatus"></a> +<b>Syrrhophus rubrimaculatus</b> Taylor and Smith</div> + +<div class="species_ref"><i>Syrrhophus rubrimaculatus</i> Taylor and Smith, 1945:583-85 +[Holotype.—USNM 114070, from La Esperanza, near Escuintla, Chiapas, +México, collected May 13, 1940, by H. M. and R. Smith]. Duellman, +1958:1-4, 7, 12, 14. Gorham, 1966:167.</div> +<div class="species_ref"><i>Syrrhophus rubrimaculata</i>: Smith and Taylor, 1948:48-49.</div> +<br /> + +<div class="fig_center" style="width: 500px;"> +<a name="Fig_8" id="Fig_8"></a> +<img src="images/fig_8.png" width="500" height="516" alt="" title="" /> +<div class="fig_caption"><span class="smcap">Fig. 8:</span> <i>Syrrhophus rubrimaculatus</i> (upper right, KU 58911, ×1.6; lower +right, KU 58910, ×4) and <i>S. verrucipes</i> (upper left, UIMNH 15995, ×1.6; +lower left, UIMNH 15989, ×3.7).</div> +</div> + +<p><i>Diagnosis.</i>—Small frogs, males 18.2-23.5 mm. snout-vent, females +19.0-22.5 mm. snout-vent length (small sample); vocal slits in males; +digital tips scarcely expanded (<a href="#Fig_1">Fig. 1</a>); first finger shorter than +second; outer palmar tubercle reduced in size; inner metatarsal tubercle +elongate, twice the size of small, conical outer metatarsal tubercle; +diameter of tympanum 35.5-46.5 per cent that of +<span class="pagenum"><a name="Page_22" id="Page_22">[Pg 22]</a></span> +eye in both sexes; dorsum brown with small pale spots (red in life); venter gray.</p> + +<p><i>Remarks.</i>—Previous authors who treated <i>Syrrhophus</i> placed this +species in the western complex, because it occurs on the Pacific versant +and has a reduced outer palmar tubercle. Duellman (1958) placed +<i>rubrimaculatus</i> apart from the other western species, because of its +relatively unexpanded digital tips and coloration. The digital tips are +like those in <i>leprus</i>, which +<i><ins title='Correction: was "rubrimacultaus"'>rubrimaculatus</ins></i> +resembles. Except for the reduction of the outer palmar tubercle, +<i>rubrimaculatus</i> could be a member of the <i>leprus</i> group.</p> + +<p><i>Syrrhophus rubrimaculatus</i> is probably best treated as a Pacific +derivative of the <i>leprus</i> group, even though the palmar tubercles do +not agree. The removal of <i>rubrimaculatus</i> from the western complex +results in a more homogeneous remainder and does not greatly increase +the heterogeneity of the eastern complex.</p> + +<p><i>Etymology.</i>—Latin, meaning spotted with red; in reference to the +colors in life.</p> + +<p><i>Distribution.</i>—Low to moderate elevations on the Pacific versant of +southeastern Chiapas, México (<a href="#Fig_7">Fig. 7</a>); probably extending into +adjacent Guatemala.</p> + +<p><i>Specimens examined.</i>—(48) MÉXICO, <i>Chiapas</i>: Escuintla, UMMZ 88283; 6 +km. NE Escuintla, UMMZ 87876-80; La Esperanza, UIMNH 13285, UMMZ +88496-97, USNM 114070 (holotype), 114054-69, 114072; Monte Cristo, UMMZ +88353; 1.3 km. N Puerto Madero, KU 58910-11; Finca San Jerónimo, 600-650 +m., UIMNH 55299-312, 55313-16 (cleared and stained).</p> +<br /> + +<div class="caption3nb"> +<a name="Syrrhophus_guttilatus" id="Syrrhophus_guttilatus"></a> +<b>Syrrhophus guttilatus</b> (Cope)</div> + +<div class="species_ref"><i>Malachylodes guttilatus</i> Cope, 1879:264 [Holotype.—USNM 9888, from +Guanajuato, Guanajuato, México; collected in 1877 by Alfredo Duges].</div> +<div class="species_ref"><i>Syrrhopus guttulatus</i>: Boulenger, 1888:204-06.</div> +<div class="species_ref"><i>Syrrhaphus guttulatus</i>: Günther, 1900:317.</div> +<div class="species_ref"><i>Syrraphus guttulatus</i>: Díaz de León, 1904:11.</div> +<div class="species_ref"><i>Syrrhophus guttilatus</i>: Nieden, 1923:399-400. Kellogg, 1932:125, +127-28. Smith and Taylor, 1948:49, 51. Firschein, 1954:52-54. Gorham, +1966:164.</div> +<div class="species_ref"><i>Syrrhophus smithi</i> Taylor, 1940b:43-45, pl. 1 [Holotype.—USNM 108594, +from 15 mi. SW Galeana, Nuevo León, México, 1575 m.; collected on +October 13, 1939, by Hobart M. Smith]. Smith and Taylor, 1948:49, 51. +Firschein, 1954:54-55. Martin, 1958:50. Gorham, 1966:167.</div> +<div class="species_ref"><i>Syrrhophus gaigeae</i> Schmidt and Smith, 1944:80 [Holotype.—FMNH 27361, +from the Basin, Chisos Mountains, Brewster Co., Texas; collected on July +24, 1937, by Walter L. Necker].</div> +<div class="species_ref"><i>Syrrhophus petrophilus</i> Firschein, 1954:50-52 [Holotype.—UIMNH 7807, +from 5 km. SW San Luis Potosí, San Luis Potosí, México; collected on +July 18, 1949, by David Langebartel]. Gorham, 1966:166.</div> +<div class="species_ref"><i>Syrrhophus marnocki</i>: Milstead, Mecham, and McClintock, 1950:548 (in +part).</div> + +<p><i>Diagnosis.</i>—Medium-sized frogs, males 20.6-29.0 mm. snout-vent, +females 25.7-31.0 mm. snout-vent length; vocal slits in males; digital +tips slightly expanded (<a href="#Fig_1">Fig. 1</a>); first and second fingers equal; skin of +dorsum smooth to moderately pustular, that of venter smooth; snout +blunt; diameter of tympanum 55.1-75.7 per cent that of eye in males, +47.6-61.7 in females; dorsum and flanks cream to gray with light brown +to black flecking and vermiculations; thighs usually not banded; +interorbital bar present (<a href="#Fig_8">Fig. 8</a>).</p> +<br /> + +<p><span class="pagenum"><a name="Page_23" id="Page_23">[Pg 23]</a></span></p> +<div class="fig_center" style="width: 487px;"> +<a name="Fig_9" id="Fig_9"></a> +<img src="images/fig_9.png" width="487" height="441" alt="" title="" /> +<div class="fig_caption"><span class="smcap">Fig. 9:</span> <i>Syrrhophus guttilatus</i> (upper left, UIMNH 55519, ×1.4; lower +left, UIMNH 55519, ×2.3) and <i>S. marnockii</i> (upper right, TCWC 9317, +×1.4; lower right, TCWC 13510, ×2.1).</div> +</div> + +<p><i>Remarks.</i>—Cope (1879) distinguished <i>Malachylodes</i> from <i>Syrrhophus</i> +on the basis of the presence of a frontoparietal fontanelle in the +holotype of <i>guttilatus</i>. The holotype is a juvenile female and as is +the case in the juveniles of nearly all leptodactylids, a frontoparietal +fontanelle is present. Firschein (1954) used the presence of the +fontanelle to distinguish <i>guttilatus</i> from his <i>petrophilus</i>.</p> + +<p>As is clearly evident from the length of the synonymy, I consider a +number of currently used names to be synonymous with <i>guttilatus</i>. I +have seen the holotypes of all four names and am unable to recognize +more than a single species. The holotype of <i>petrophilus</i> is a male, +whereas that of <i>smithi</i> is a female. The supposed differences are a +reflection of sexual dimorphism in the size of the eye (<a href="#Table_5">Table 5</a>). The +two holotypes, as well as those of <i>gaigeae</i> and <i>Malachylodes +guttilatus</i> agree in color pattern.</p> + +<p>Schmidt and Smith (1944) named <i>Syrrhophus gaigeae</i> from the Chisos +Mountains of the Big Bend region of Texas and compared it only with <i>S. +marnockii</i>. Milstead, Mecham and McClintock (1950) synonymized <i>gaigeae</i> +and <i>marnockii</i> because they were unable to verify the characters Wright +and Wright (1949) used to separate them. Specimens from the Big Bend +region differ from those of the Edward and Stockton Plateaus in having a vermiculate +<span class="pagenum"><a name="Page_24" id="Page_24">[Pg 24]</a></span> +pattern, an interorbital bar, and a supratympanic stripe. +In these respects they agree with specimens from northern México. Based +on limited observations, the Mexican population is yellowish to brownish +in life whereas the central Texas population is green in life. Lacking +evidence of genetic exchange, the two are held to be specifically +distinct.</p> +<br /> + +<div class="fig_center" style="width: 520px;"> +<a name="Fig_10" id="Fig_10"></a> +<img src="images/fig_10.png" width="520" height="748" alt="" title="" /> +<div class="fig_caption"><span class="smcap">Fig. 10:</span> Distribution of +<i>Syrrhophus guttilatus</i>.</div> +</div> + +<p><span class="pagenum"><a name="Page_25" id="Page_25">[Pg 25]</a></span> +Nearly every specimen examined was infested with chiggers of the genus +<i>Hannemania</i>. The greatest concentrations are on the venter, in the +groin, and on the thighs. Many specimens have chiggers on the digits and +tarsi. The same, or a related, chigger was found on many specimens of +<i>Syrrhophus marnockii</i> and a few <i>S. verrucipes</i>, but on no other +species of the genus. Mr. Willy Wrenn told me that he has seen heavy +infestations of <i>Hannemania</i> on <i>Syrrhophus pallidus</i>. Infestation by +<i>Hannemania</i> probably reflects similar ecologies rather than close +relationships.</p> + +<p><i>Etymology.</i>—Latin, <i>guttula</i>, meaning spotting or flecking, in +reference to the color pattern.</p> + +<p><i>Distribution.</i>—Moderate to intermediate elevations (600 to 2000 m.) +along the Sierra Madre Oriental from the Big Bend Region of Texas to +Guanajuato, México (<a href="#Fig_10">Fig. 10</a>).</p> + +<p><i>Specimens examined.</i>—(32) TEXAS, <i>Brewster Co.</i>: Juniper Canyon, +Chisos Mts., FMNH 27361 (holotype of <i>S. gaigeae</i>), 27360, 27362-63, MCZ +15346, 27801, UMMZ 66080, 66082, 66085-91, USNM 76876; Upper Green +Gulch, TCWC 15943.</p> + +<p>MÉXICO: <i>Coahuila</i>: 8 km. S Saltillo, UIMNH 55518-21. <i>Guanajuato</i>: +Guanajuato, USNM 9888 (holotype of <i>Malachulodes guttilatus</i>); 8 km. E +Guanajuato, AMNH 73425; Cerro Cubilete, AMNH 73424. <i>Nuevo León</i>: 3 km. +S Galeana, JDL 1215 (skeleton), UIMNH 58204; 24 km. SW Galeana. 1575 m., +USNM 108594 (holotype of <i>Syrrhophus smithi</i>). <i>San Luis Potosí</i>: 5 km. +SW San Luis Potosí, UIMNH 7807 (holotype of <i>S. petrophilus</i>). +<i>Tamaulipas</i>: 1.6 km. NW La Joya de Salas, 1530 m., UMMZ 110736 (4).</p> +<br /> + +<div class="caption3nb"> +<a name="Syrrhophus_marnockii" id="Syrrhophus_marnockii"></a> +<b>Syrrhophus marnockii</b> Cope</div> + +<div class="species_ref"><i>Syrrhophus marnockii</i> Cope, 1878:253 [Syntypes.—ANSP 10765-68, from +"near San Antonio," Bexar Co., Texas; collected by G. W. Marnock].</div> +<div class="species_ref"><i>Syrrhophus marnocki</i>: Yarrow, 1882:24, 193. Milstead, Mecham, and +McClintock, 1950:550.</div> + +<p><i>Diagnosis.</i>—Medium-sized frogs, males 18.4-28.9 mm. snout-vent, +females 20.4-35.4 mm. snout-vent length; vocal slits in males; digital +tips widened (<a href="#Fig_1">Fig. 1</a>); first and second fingers equal; skin of dorsum +smooth to weakly pustular, that of venter smooth; snout blunt, rounded; +diameter of tympanum 47.2-68.3 per cent that of eye in males, 45.8-73.3 +in females; dorsum tan to light brown in preservative with rusty-brown +flecks, venter white; ground color green in life; thighs banded; +interorbital bar absent.</p> + +<p><i>Remarks.</i>—Specimens from the southern edge of the Edwards Plateau and +the eastern edge of the Stockton Plateau have larger flecks on the back +that tend to form a vermiculate pattern like that of <i>S. guttilatus</i>. +The vermiculation is never well developed (see plate 38 in Conant, +1958). Most of the specimens from the Edwards Plateau have a punctate +pattern (<a href="#Fig_9">Fig. 9</a>).</p> + +<p>Fossils are known from the Sangamon interglacial deposits in Foard and +Knox Counties, Texas (Lynch, 1964; Tihen, 1960).</p> + +<p><i>Etymology.</i>—A patronym for the collector of the type specimens.</p> + +<p><i>Distribution.</i>—The Edwards Plateau and the extreme eastern edge of the +Stockton Plateau in Texas (<a href="#Fig_11">Fig. 11</a>). The fossil records lie some 200 +miles to the north. Two specimens (FMNH 103216-17) from Brownsville, +Cameron Co., Texas, were formerly in the EHT-HMS collection (nos. +31348-49). Data given in Taylor's field catalogue (housed in the +Division of Reptiles, Field Museum) are "Brownsville, A. J. Kirn +collector, April 15, 1934." Until verification +<span class="pagenum"><a name="Page_26" id="Page_26">[Pg 26]</a></span> +by recently collected material is available, this record must be disregarded.</p> + +<p><i>Specimens examined.</i>—(103) TEXAS, <i>Bandera Co.</i>: 10 mi. SW Medina, +TCWC 13508-10; 8 mi. W Medina, KU 60243; 13 mi. W Medina, KU 60242, TCWC +13506-07. <i>Bexar Co.</i>: UIMNH 34694; Classen ranch, near San Antonio. +UMMZ 98891; Helotes, EAL 1560, MCZ 11837 (2), UMMZ 64045, USNM 13635; 2 +mi. N Helotes, TCWC 9234-35; 3.5 mi. N Helotes, LSUMZ 10363; 8 mi. N +Helotes, TCWC 1549, 4364; San Antonio, FMNH 15553-56, TCWC 13497-99. +<i>Blanco Co.</i>: 8 mi. NE Blanco, TCWC 4782. <i>Comal Co.</i>: New Braunfels, +TCWC 13500-05; 5 mi. NE New Braunfels, UMMZ 71016 (10). <i>Hays Co.</i>: San +Marcos, AMNH 22661-64, 32700, FMNH 15245-46, 26250, 26253-57, 37617, +37665, MCZ 15649-50, 23268-69; 6 mi. SW San Marcos, TCWC 5070-71, 7140, +9232-33, 9236, 9316-17, 9320. <i>Kendall Co.</i>: 11 mi. E Boerne, AMNH +54660-61, 54662 (2); 10 mi. W Boerne, KU 18441; Kendalia, UIMNH 21434. +<i>Kerr Co.</i>: Kerr W. M. Area, TCWC 15859; 40 mi. NW Kerrville, TCWC 6555. +<i>Medina Co.</i>: UIMNH 13287-88; 12 mi. N Castroville, UIMNH 21423; 14 mi. +N Castroville, UIMNH 21424-25; 16 mi. N Castroville, UIMNH 21421-22; 17 +mi. N Castroville, UIMNH 21428-29; 18 mi. N Castroville, UIMNH 21426-27, +21430-33; 6.5 mi. NW Rio Medina, KU 18440. <i>Real Co.</i>: Rio Frio, FMNH +55156-57. <i>Travis Co.</i>: Austin, AMNH 44221-22; Mount Bonnell, 5 mi. S +Austin, UMMZ 101453 (10). <i>Uvalde Co.</i>: 13 mi. from Uvalde, UIMNH 62322. +<i>Val-Verde Co.</i>: 40 mi. N Del Rio, JDL 214 (skeleton).</p> +<br /> + +<div class="fig_center" style="width: 502px;"> +<a name="Fig_11" id="Fig_11"></a> +<img src="images/fig_11.png" width="502" height="449" alt="" title="" /> +<div class="fig_caption"><span class="smcap">Fig. 11:</span> Distribution of <i>Syrrhophus marnockii</i> (circles). Starred +localities are late Pleistocene records.</div> +</div> + +<p><span class="pagenum"><a name="Page_27" id="Page_27">[Pg 27]</a></span></p> + +<div class="caption3nb"> +<a name="Syrrhophus_verrucipes" id="Syrrhophus_verrucipes"></a> +<b>Syrrhophus verrucipes</b> Cope</div> + +<div class="species_ref"><i>Syrrhophus verrucipes</i> Cope, 1885:383 [Holotype.—ANSP 11325, from near +Zacualtipán, Hidalgo, México (1800 feet lower in a rocky gorge of a +stream near its junction with the Río San Miguel), collected by Dr. +Santiago Bernard]. Kellogg, 1932:126-29. Smith and Taylor, 1948:52-53. +Firschein, 1954:55-57. Gorham, 1966:167.</div> +<div class="species_ref"><i>Syrrhaphus verrucipes</i>: Günther, 1900:216-17.</div> +<div class="species_ref"><i>Tomodactylus macrotympanum</i> Taylor, 1940e:496-99, pl. 55, figs. 2a-b. +[Holotype.—FMNH 100049 (formerly EHT-HMS 6838), from La Placita, 8 km. +S Jacala, Hidalgo, México, 1850 m.; collected on July 2, 1936, by Edward +H. Taylor]. Smith and Taylor, 1948:47-48.</div> +<div class="species_ref"><i>Syrrhophus macrotympanum</i>: Dixon, 1957:384. Gorham, 1966:165.</div> + +<p><i>Diagnosis.</i>—Medium-sized frogs, males 17.5-26.1 mm. snout-vent, +females 28.0-31.7 mm. snout-vent length; vocal slits in males; digital +tips slightly expanded; first finger shorter than second; skin of dorsum +pustular, that of venter areolate; snout elongate, subacuminate; +diameter of tympanum 56.1-76.7 per cent that of eye in males, 54.3-56.8 +in females; in preservative, dorsum reddish brown with numerous small +black or dark brown spots (<a href="#Fig_8">Fig. 8</a>); venter white to cream; in life +dorsum green with darker green spots, belly white; iris gold above, +bronze below.</p> + +<p><i>Remarks.</i>—Cope's (1885) original description was not sufficiently +clear to enable subsequent authors to recognize this species. Taylor +(1940e) described it as a <i>Tomodactylus</i>, but Dixon (1957) pointed out +that <i>T. macrotympanum</i> differed from the other species of the genus in +having a poorly developed lumbo-inguinal (inguinal) gland, and placed +the species in the genus <i>Syrrhophus</i>. Comparison of the holotypes of +<i>S. verrucipes</i> and <i>T. macrotympanum</i> leaves no doubt in my mind that a +single species is involved. This same species was reported by Smith and +Taylor (1948) as <i>S. verruculatus</i>.</p> + +<p><i>Syrrhophus verrucipes</i> bears +<ins title='Correction: was "resemblence"'>resemblance</ins> to members of +both the <i>leprus</i> and <i>marnockii</i> groups. In snout shape it is closer to +the <i>leprus</i> group, whereas in digital pad, the shape of the general +body form, and contiguity of habitat it is most similar to the +<i>marnockii</i> group (<i>S. guttilatus</i>).</p> + +<p><i>Etymology.</i>—Latin, meaning warty foot, probably in reference to the +numerous plantar supernumerary tubercles.</p> + +<p><i>Distribution.</i>—Moderate elevations in southeastern San Luis Potosí, +Queretaro, and northwestern Hidalgo, México (<a href="#Fig_7">Fig. 7</a>).</p> + +<p><i>Specimens examined</i>—(43) MÉXICO, <i>Hidalgo</i>: Jacala, UMMZ 106434; 9.6 +km. NE Jacala, Puerto de la Zorra, 1820 m., KU 60240-41, TCWC 11090, +11147; 8 km. S Jacala, La Placita, 1850 m., FMNH 100049 (holotype of +<i>Tomodactylus macrotympanum</i>), 100791-803, 105334-35, 114287, UIMNH +15989-92, 15995-96, UMMZ 117252, USNM 137202; Tianguistengo, FMNH +113705-09, UIMNH 13328-30; near Zacualtipán, ANSP 11325 (holotype of +<i>Syrrhophus verrucipes</i>). <i>Queretaro</i>: 3.5 km. S San Juan del Río, EAL +1343. <i>San Luis Potosí</i>: 9.6 km. W Ahuacatlán, LSUMZ 4968-70.</p> +<br /> + +<div class="caption3nb"> +<a name="Syrrhophus_dennisi" id="Syrrhophus_dennisi"></a> +<b>Syrrhophus dennisi</b> new species</div> + +<div class="species_ref"><i>Syrrhophus latodactylus</i>: Martin, 1958:49 (in part).</div> + +<p><i>Holotype.</i>—UMMZ 101121, adult male from a cave near El Pachón, 8 km. N +Antiguo Morelos, Tamaulipas, México, 250 m., collected on March 13, +1949, by Paul S. Martin.</p> + +<p><span class="pagenum"><a name="Page_28" id="Page_28">[Pg 28]</a></span> +<i>Paratopotypes.</i>—(26). UMMZ 101122 (10), 101123 (2), 101126, 126993 +(12).</p> + +<p><i>Diagnosis.</i>—Medium-sized frogs, males 22.8-28.4 mm. snout-vent, +females 25.9-32.0 mm. snout-vent; vocal slits in males; digital tips +greatly expanded, more than twice width of digit; first finger shorter +than second; skin of dorsum shagreened to pustular, that of venter +weakly to moderately areolate; toes webbed basally; dorsum light brown +to tan with brown vermiculations; venter white; diameter of tympanum +53.9 to 64.2 per cent that of eye in males, 50.6 to 58.7 per cent in +females.</p> + +<p><i>Description and variation.</i>—(<a href="#Fig_12">Fig. 12</a>). Head wider than body; head as +wide or wider than long in males, sometimes longer than wide in females; +snout acuminate in dorsal view, elongate and rounded in lateral profile; +canthus rostralis rounded but distinct; loreal region slightly concave, +sloping abruptly to lip; lips not flared; eyelid about two-thirds +interorbital distance; length of eye less than distance between eye and +nostril; diameter of tympanum 53.9 to 64.2 per cent that of eye in +males, 50.6 to 58.7 per cent in females; tympanum round and distinct in +both sexes; supratympanic fold moderately distinct; choanae within +border of jaws, completely visible from directly below, rounded to +slightly oval; dentigerous processes of prevomers and teeth absent; +tongue free for posterior one-half, generally oval in outline; vocal +slits present in males.</p> + +<p>Many scattered pustules on dorsum; flanks areolate; skin of venter +areolate or not (variability may be due to differences in preservation); +ventral disc distinct on chest and lower abdomen; inguinal gland present +or not, when present varying from very large and distinct to poorly +defined; axillary gland absent.</p> + +<p>First finger shorter than second; all fingers bearing truncate tips with +pads, each pad having a terminal groove; fingers fringed; fingers three +and four having dilated pads two to three times width of digit; +subarticular tubercles large, conical, rounded, simple; supernumerary +tubercles numerous on thenar surface, none on digits; three palmar +tubercles, outer slightly smaller than largest supernumerary tubercles; +row of tubercles on outer edge of forearm variable, weak to very +distinct; tips of toes wider than digits, rounded to truncate at tips, +each pad having terminal groove; toes having lateral fringes, bases of +toes united by web, web not extending to basal subarticular tubercle; +subarticular tubercles smaller than those of hand, round, conical, +simple; supernumerary tubercles numerous on plantar surfaces, extending +between metatarsal tubercles, present on toes between basal two +subarticular tubercles in some specimens; outer metatarsal tubercle +round, conical, one-half as large as ovoid, non-compressed inner +metatarsal tubercle; tarsal tubercles or folds absent.</p> + +<p>Ground color pale reddish-brown to tan dorsally, creamy on flanks; +dorsal pattern consisting of reddish-brown to brown vermiculations +extending onto flanks; distinct interorbital light bar present; loreal +region darker than snout, reddish-brown compared to tan or pale +reddish-brown; arms colored like dorsum; thighs banded, unicolor brown +on posterior surfaces; shanks and tarsi banded; venter white to cream +punctated with brown in some specimens.</p> + +<p>The variation in proportions is summarized in <a href="#Table_5">Table 5</a>.</p> + +<p><i>Remarks.</i>—Martin (1958) expressed some doubt that this series of +26 specimens was identical with "<i>S. latodactylus</i>." My study indicates +that the specimens from El Pachón represent a distinctive but allied +species. Males of the two species can be readily separated by the +relative sizes of the tympani, presence or absence of vocal slits, and +color pattern. Females of the two species can be separated by color +pattern. Within the type-series, the pattern varies from weakly to +strongly vermiculate but is always recognizable as vermiculate rather +than spotted as in <i>S. longipes</i> (= <i>S. latodactylus</i> of Taylor and +Martin).</p> +<br /> + +<p><span class="pagenum"><a name="Page_29" id="Page_29">[Pg 29]</a></span></p> +<div class="fig_center" style="width: 336px;"> +<a name="Fig_12" id="Fig_12"></a> +<img src="images/fig_12.png" width="336" height="660" alt="" title="" /> +<div class="fig_caption"><span class="smcap">Fig. 12:</span> <i>Syrrhophus dennisi</i> sp. nov., holotype, UMMZ 101121 (dorsum +×1.8, side of head ×6.1).</div> +</div> + +<p><span class="pagenum"><a name="Page_30" id="Page_30">[Pg 30]</a></span> +<i>Etymology.</i>—The specific name is a patronym for David M. Dennis, whose +drawings greatly enhance the worth of this paper.</p> + +<p><i>Distribution.</i>—Known only from the type series.</p> +<br /> + +<div class="caption3nb"> +<a name="Syrrhophus_longipes" id="Syrrhophus_longipes"></a> +<b>Syrrhophus longipes</b> (Baird), New combination</div> + +<div class="species_ref"><i>Batrachyla longipes</i> Baird, 1859:35, pl. 37, fig. 1-3 +[Holotype.—apparently USNM 3237 (cited as 3207 by Cope, 1887:16), now +lost, from 40 Leagues from (probably north) México City; collected by +John Potts]. Kellogg, 1932:107.</div> +<div class="species_ref"><i>Epirhexis longipes</i>: Cope, 1866:96.</div> +<div class="species_ref"><i>Eleutherodactylus longipes</i>: Kellogg, 1932:107 (part). Smith and +Taylor, 1948:61. Lynch, 1963:580-581. Gorham, 1966:82.</div> +<div class="species_ref"><i>Syrrhophus latodactylus</i> Taylor, 1940d:396-401, pl. 43, figs. A-F, text +fig. 7 [Holotype.—FMNH 100063 (formerly EHT-HMS 6807), from Huasteca +Canyon, 15 km. W Monterrey, Nuevo León, México, 680 m.; collected on +June 20, 1936, by Edward H. Taylor]. Smith and Taylor, 1948:50-52. +Martin, 1958:48-50. Gorham, 1966:165.</div> + +<p><i>Diagnosis.</i>—Large frogs, males 22.1-33.2 mm. snout-vent, females +26.8-39.6 mm. snout-vent length; vocal slits lacking in males; digital +tips greatly expanded (more than twice the width of digit); first finger +shorter than second; skin of dorsum pustular, that of venter smooth; +diameter of tympanum in males 61.1-87.2 per cent that of eye, 49.5-72.1 +per cent in females; dorsum tan with large or small spots and blotches; +limbs banded; interorbital bar or triangle present.</p> + +<p><i>Remarks.</i>—I have applied Baird's <i>Batrachyla longipes</i> to the frog +Taylor (1940d) called <i>Syrrhophus latodactylus</i> because the color +pattern (<a href="#Fig_13">Fig. 13</a>) predominant in the southern part of the range agrees +with that described (figured) for <i>Batrachyla longipes</i>.</p> + +<p>The color pattern of individuals in the southern part of the range of +this species consists of large spots or blotches, whereas in the +northwestern part the pattern is made up of smaller spots. In the +northeastern part of the range, the pattern is more reduced and tends to +consist of heavy flecking. The interorbital bar is narrower in specimens +from Nuevo León and Tamaulipas and is triangular in specimens from +Hidalgo and Queretaro.</p> + +<p>The status of the name <i>Batrachyla longipes</i> is currently that of a +<i>nomen dubium</i> (Lynch, 1963). At that time, I was unaware of the +geographic variation in color pattern in <i>Syrrhophus latodactylus</i>.</p> + +<p>The exact type-locality of <i>Batrachyla longipes</i> is not known. If it is +40 Leagues north of México City, the locality would be in an area where +the species has a blotched instead of a flecked or spotted pattern. No +justifiable evidence was presented to place <i>Batrachyla longipes</i> in +<i>Eleutherodactylus</i> instead of <i>Syrrhophus</i>. Barbour (1923) and Kellogg +(1932) associated another species (<i>E. batrachylus</i>) with <i>longipes</i>. +Taylor (1940a) noted this as a case of misidentification and corrected +the error but left <i>longipes</i> in the genus <i>Eleutherodactylus</i>. Lynch +(1963) noted several points of morphological agreement between +<i>Syrrhophus</i> and <i>B. longipes</i> but did not place <i>longipes</i> in +<i>Syrrhophus</i>.</p> + +<p>Baird's (1859) figures of the holotype do not illustrate prevomerine +teeth, but according to Cope (1866) they were present in the holotype. +The digital +<span class="pagenum"><a name="Page_31" id="Page_31">[Pg 31]</a></span> +tips of the frog in the figure are somewhat narrower than +those typically seen in <i>S. latodactylus</i>. If the specimen was slightly +desiccated, as possibly was the case, the digits would appear narrower. +There is no evidence contrary to placing <i>Syrrhophus latodactylus</i> in +the synonymy of <i>Batrachyla longipes</i>.</p> +<br /> + +<div class="fig_center" style="width: 486px;"> +<a name="Fig_13" id="Fig_13"></a> +<img src="images/fig_13.png" width="486" height="652" alt="" title="" /> +<div class="fig_caption"><span class="smcap">Fig. 13:</span> Dorsal views of <i>Syrrhophus longipes</i> illustrating geographic +variation in pattern (left, TCWC 12179, ×1.5; right, KU 92572, ×1.8); +side of head (TCWC 10966, ×6).</div> +</div> + +<p>Application of Baird's name <i>Batrachyla longipes</i> to the species of frog +heretofore called <i>Syrrhophus latodactylus</i> poses one serious problem. <i>Batrachyla +<span class="pagenum"><a name="Page_32" id="Page_32">[Pg 32]</a></span> +longipes</i> is the type-species (by original designation) of the genus <i>Epirhexis</i> +Cope, 1866, which has priority over <i>Syrrhophus</i> Cope, 1878. If <i>Batrachyla +longipes</i> is left in the status of a <i>nomen dubium</i>, <i>Epirhexis</i> can be forgotten, +for the two names are tied together. However, since it seems almost certain +that <i>Batrachyla longipes</i> and <i>Syrrhophus latodactylus</i> are conspecific, the former +name should not be left as a <i>nomen dubium</i>. <i>Epirhexis</i> never came into general +usage (Cope cited the name four times, but no one else has used it), whereas +<i>Syrrhophus</i> is well established in the zoological literature. It would serve only +to confuse the literature to adhere strictly to the Law of Priority and replace +<i>Syrrhophus</i> with <i>Epirhexis</i>. Therefore, <i>Syrrhophus</i> is used in this paper, even +though <i>Epirhexis</i> has priority. A request for the suppression of <i>Epirhexis</i> Cope, +1866, has been submitted to the International Commission of Zoological +Nomenclature (Lynch, 1967).</p> + +<p><i>Etymology.</i>—Latin, meaning long-footed; Taylor's <i>latodactylus</i> refers to +the wide digital pads.</p> + +<div class="fig_center" style="width: 481px;"> +<a name="Fig_14" id="Fig_14"></a> +<img src="images/fig_14.png" width="481" height="530" alt="" title="" /> +<div class="fig_caption"><span class="smcap">Fig. 14:</span> Distribution of <i>Syrrhophus dennisi</i> (triangle) and <i>S. +longipes</i> (circles).</div> +</div> + +<p><span class="pagenum"><a name="Page_33" id="Page_33">[Pg 33]</a></span> +<i>Distribution.</i>—Moderate elevations (650 to 2000 meters) along the +Sierra Madre Oriental from central Nuevo León to northern Hidalgo, +México (<a href="#Fig_14">Fig. 14</a>).</p> + +<p><i>Specimens examined.</i>—(122) MÉXICO, <i>Hidalgo</i>: 3 km. NE Jacala, AMNH +52977; 9.6 km. NE Jacala, 1800 m., TCWC 10966-70, 12179; 8 km. S Jacala, +La Placita, 1850 m., FMNH 100266-68, 103244, UIMNH 13291, 13327. <i>Nuevo +León</i>: Salto Cola de Caballo, KU 92572; Huasteca Canyon, 15 km. W +Monterrey, 680 m., FMNH 100063 (holotype of <i>S. latodactylus</i>), UIMNH +13290; 6.5 km. N Pablillo, EAL 1319; Sabinas Hidalgo, USNM 139728. +<i>Queretaro</i>: Cueva de los Riscos, 8 km. SW Jalpan, KU 106300. <i>San Luis +Potosí</i>: 13 km. E Santa Barberita, LSUMZ 2295; second camp, San Luis +Potosí road, UIMNH 13326; Xilitla, Cueva sin nombre, UMMZ 125892. +<i>Tamaulipas</i>: 4 km. W El Carrizo, 500 m., UMMZ 111343 (31); 8 km. N +Chamal, Bee Cave, KU 106299; 14.5 km. NNW Chamal, 420 m., UMMZ +111339-40, 111342 (4), 111344 (11); 19 km. NNW Chamal, 700 m., UMMZ +111341 (3); El Chihue, 1880 m., UMMZ 111289 (4); 11 km. N Gómez Farías, +1060 m., UMMZ 101166; 11 km. WNW Gómez Farías, 1800 m., UMMZ 108507 (3); +8 km. NW Gómez Farías, 1060-1400 m., LSUMZ 11085, UMMZ 101167 (3), +101168 (4), 101169 (2), 101170 (3), 101171 (2), 101360-61, 102860, +102933 (4), 102934 (2), 102935-38, 102939 (2), 102940-43, 108800 (3), +110735, 111345-46.</p> +<br /> + +<div class="caption3nb"> +<a name="Syrrhophus_pipilans" id="Syrrhophus_pipilans"></a> +<b>Syrrhophus pipilans</b> Taylor</div> + +<div class="species_ref"><i>Syrrhophus pipilans</i> Taylor, 1940c:95-97, pl. 1 [Holotype.—FMNH 100072 +(formerly EHT-HMS 6843), 14.6 km. S Mazatlán, Guerrero, México; +collected on July 22, 1936, by Edward H. Taylor].</div> + +<p><i>Diagnosis.</i>—Medium sized frogs, males 22.6-28.5 mm. snout-vent, +females 21.1-29.4 mm. snout-vent length; vocal slits present in males; +finger tips slightly expanded, truncate in outline; inner metatarsal +tubercle less than twice the size of outer; skin of dorsum smooth to +shagreened, that of venter smooth; tympanum 36.5-54.0 per cent diameter +of eye; dorsum dark brown with large or small light brown, orange-brown, +or yellowish spots or blotches; limbs banded; interorbital bar absent.</p> +<br /> + +<div class="fig_center" style="width: 467px;"> +<a name="Fig_15" id="Fig_15"></a> +<img src="images/fig_15.png" width="467" height="180" alt="" title="" /> +<div class="fig_caption"><span class="smcap">Fig. 15:</span> Dicegrams of ear size relative to eye diameter in the two +subspecies of <i>Syrrhophus pipilans</i>. N = 17 in <i>nebulosus</i>, 18 in +<i>pipilans</i>.</div> +</div> + +<p><i>Remarks.</i>—Two subspecies were recognized by Duellman (1958). +Previously both had been treated as species. The two populations were +distinguished on the basis of color pattern and the size of the +tympanum. Measurements of 17 males of <i>S. p. nebulosus</i> from central +Chiapas and 18 males of <i>S. p. pipilans</i> from south-central Oaxaca and +Guerrero, México, demonstrates that the supposed difference in tympanum +size is not significant (<a href="#Fig_15">Fig. 15</a>). +<span class="pagenum"><a name="Page_34" id="Page_34">[Pg 34]</a></span> +There is, however, a tendency for +the western population of <i>S. pipilans</i> to have larger tympani. Based on +the present examination of 112 specimens of this species the two +populations are held to be sufficiently distinct to warrant taxonomic +recognition as subspecies (<a href="#Fig_16">Fig. 16</a>).</p> +<br /> + +<div class="fig_center" style="width: 495px;"> +<a name="Fig_16" id="Fig_16"></a> +<img src="images/fig_16.png" width="495" height="356" alt="" title="" /> +<div class="fig_caption"><span class="smcap">Fig. 16:</span> <i>Syrrhophus pipilans nebulosus</i> (left, KU 58908) and <i>S. p. +pipilans</i> (right, KU 86885). ×2.7.</div> +</div> + +<p>The parotoid glands attributed to this species by Taylor (1940c:95) are +merely the superficial expression of the <i>m. depressor mandibulae</i> and +scapula. No true glands are present in the parotoid region.</p> +<br /> + +<div class="caption3nb"> +<a name="Syrrhophus_pipilans_nebulosus" id="Syrrhophus_pipilans_nebulosus"></a> +<b>Syrrhophus pipilans nebulosus</b> Taylor</div> + +<div class="species_ref"><i>Syrrhophus nebulosus</i> Taylor, 1943:353-55, pl. 27, figs. 3-5 +[Holotype.—FMNH 100095 (formerly EHT-HMS 3774), near Tonolá, Chiapas, +México; collected on August 27, 1935, by Hobart M. Smith and Edward H. +Taylor]. Smith and Taylor, 1948:49, 51.</div> +<div class="species_ref"><i>Syrrhophus pipilans nebulosus</i>: Duellman, 1958:2-4, 9, 12, 14. Stuart, +1963:32-33. Gorham, 1966:166-67.</div> + +<p><i>Diagnosis.</i>—Diameter of tympanum 36.6-47.8 per cent that of eye; +dorsum dark brown with numerous small light brown to yellowish spots.</p> + +<p><i>Remarks.</i>—The distribution of this subspecies is adequately described +by Duellman (1958). Fouquette (1960) described the vocalization of this +frog.</p> + +<p><i>Etymology.</i>—Latin, <i>nebula</i>, in reference to the clouded dorsal +pattern.</p> + +<p><i>Distribution.</i>—Low to moderate elevations along the Pacific versant of +Chiapas and in the Grijalva valley of Chiapas and Guatemala (<a href="#Fig_17">Fig. 17</a>).</p> + +<p><i>Specimens examined.</i>—(54) GUATEMALA, <i>Huehuetenango</i>: Jacaltenango, +UMMZ 117036; 35 km. SE La Mesilla, TNHC 29652. MÉXICO, <i>Chiapas</i>: 11.2 +km. N Arriaga, 300 m., UMMZ 125891; 11.8 km. N Arriaga, UMMZ 117279; +<span class="pagenum"><a name="Page_35" id="Page_35">[Pg 35]</a></span> +12.8 km. N Arriaga, UMMZ 117280; 17.5 km. S Arriaga, UIMNH 57108-109; +1.5 km. S Bochil, 1250 m., KU 58898-908; Cerro Hueco, 7 km. S Tuxtla +Gutierrez, UMMZ 123007; 3.2 km. S Ixtapa, UMMZ 124000; Linda Vista, ca. +2 km. NW Pueblo Nuevo Solistahuacán, KU 58897; Hda. Monserrate, 40 km. +NW Arriaga, UMMZ 102258; near San Ricardo, FMNH 100720; Tapachula, FMNH +75792, 103242, 100695-96, UIMNH 13292; 56 km. E Tapanatepec, Oaxaca, +TNHC 26942, Tonolá, FMNH 100095 (holotype), 100686-92, UIMNH 13293-95; +Tuxtla Gutierrez, FMNH 100693-94, UIMNH 13297; 19 km. N Tuxtla +Gutierrez, TNHC 25229-30; 15.5 km. NE Tuxtla Gutierrez, UMMZ 119892 (3); +19 km. NE Tuxtla Gutierrez, UMMZ 119891 (3); 8 km. NNW Tuxtla Gutierrez, +KU 37809; Unión de Juarez, FMNH 105294.</p> +<br /> + +<div class="caption3nb"> +<a name="Syrrhophus_pipilans_pipilans" id="Syrrhophus_pipilans_pipilans"></a> +<b>Syrrhophus pipilans pipilans</b> Taylor</div> + +<div class="species_ref"><i>?Syrrhopus verruculatus</i>: Gadow, 1905:194.</div> +<div class="species_ref"><i>Syrrhophus pipilans</i> Taylor, 1940c:95-97, pl. 1 [Holotype.—FMNH 100072 +(formerly EHT-HMS 6843), from 14.6 km. S Mazatlán, Guerrero, México; +collected on July 22, 1936, by Edward H. Taylor]. Taylor and Smith, +1945:581-82. Smith and Taylor, 1948:49, 50-51.</div> +<div class="species_ref"><i>Syrrhophus pipilans pipilans</i>: Duellman, 1958:1-4, 8-9, 13-14, pl. 2, +fig. 1. Gorham, 1966:166.</div> + +<p><i>Diagnosis.</i>—Diameter of tympanum 40.6-54.0 per cent that of eye; +dorsum dark brown with large light spots or blotches.</p> + +<p><i>Remarks.</i>—Duellman's (1958) synopsis of this subspecies is adequate; +the distribution has not been extended, but several records are now +available which fill in gaps.</p> +<br /> + +<div class="fig_center" style="width: 480px;"> +<a name="Fig_17" id="Fig_17"></a> +<img src="images/fig_17.png" width="480" height="286" alt="" title="" /> +<div class="fig_caption"><span class="smcap">Fig. 17:</span> Distribution of <i>Syrrhophus pipilans</i>: <i>nebulosus</i> (open +circles) and <i>pipilans</i> (solid circles).</div> +</div> + +<p>Gadow's (1905) record of <i>S. verruculatus</i> from "Buena Vista, S. +Guerrero" is most likely applicable to this species. Gadow simply +included the name in a list of the species he had collected during his +trip in México (1902-04); no +<span class="pagenum"><a name="Page_36" id="Page_36">[Pg 36]</a></span> +further comment was made on this species +although references to <i>Syrrhopus</i> (sic) appear in several places in the +paper and would appear to apply to the species he had.</p> + +<p><i>Etymology.</i>—Latin, <i>pipilo</i>, chirping, peeping, in reference to the +call of the male.</p> + +<p><i>Distribution.</i>—Sea level to about 1800 meters along the Pacific +versant of western México from central Guerrero to the Isthmus of +Tehuantepec (<a href="#Fig_17">Fig. 17</a>).</p> + +<p><i>Specimens examined.</i>—(62). MÉXICO, <i>Guerrero</i>: Acapulco, UMMZ 110125; +6.4 km. N Acapulco, FMNH 100389, 100525; Agua del Obispo, 980-1000 m., +FMNH 75791, 100518-21, 100526, KU 86884-86, UIMNH 13315, UMMZ 119152, +125890 (4); 13.3 km. NW Coyuca, UIMNH 38367, 71982-83; 14.5 km. S +Mazatlán, FMNH 100072 (holotype), 100408, 100511-17, UIMNH 13302-309; +Tierra Colorado, 300 m., KU 67961, UIMNH 13313-14; near El Treinte, FMNH +126639; Xaltinanguis, FMNH 100522-24, 126640. <i>Oaxaca</i>: Cacahuatepec, +UIMNH 52853; 8 km. NW Río Canoa, 53 km. ESE Cuajinicuilapa, UIMNH +52852; 6.4 km. N El Candelaria, UIMNH 9501; 11.2 km. S El Candelaria, +UIMNH 9502; 17 km. NE Juchatengo, 1600 m., KU 86887; 31.5 km. N +Pochutla, UMMZ 123999 (2); 32.9 km. N Pochutla, 850 m., UMMZ 123996; +37.1 km. N Pochutla, UMMZ 123998 (2); 41.4 km. N Pochutla, UMMZ 123997 +(2); Cerro Quiengola, FMNH 105653; 3.8 km. N Santiago Chivela, UMMZ +115449; 14.5 km. W Tehuantepec, UMMZ 115448 (2).</p> +<br /> + +<div class="caption3nb"> +<a name="Syrrhophus_interorbitalis" id="Syrrhophus_interorbitalis"></a> +<b>Syrrhophus interorbitalis</b> Langebartel and Shannon</div> + +<div class="species_ref"><i>Syrrhophus interorbitalis</i> Langebartel and Shannon, 1956: 161-65, figs. +1-2 [Holotype.—UIMNH 67061 (formerly FAS 9378), 36 mi. N Mazatlán, +Sinaloa, México, collected on November 17, 1955, by E. C. Bay, J. C. +Schaffner, and D. A. Langebartel]. Duellman, 1958:1-4, 10, 12, 14. +Gorham, 1966:164-65.</div> +<div class="species_ref"><i>Syrrhophis interorbitalis</i>: Campbell and Simmons, 1962:194, fig. 1.</div> +<br /> + +<div class="fig_center" style="width: 516px;"> +<a name="Fig_18" id="Fig_18"></a> +<img src="images/fig_18.png" width="516" height="328" alt="" title="" /> +<div class="fig_caption"><span class="smcap">Fig. 18:</span> Left to right. <i>Syrrhophus interorbitalis</i> (UIMNH 38095, ×1.5), +<i>S. nivocolimae</i> (LACM 3203, ×1.3), and <i>S. teretistes</i> (KU 75263, +×1.5).</div> +</div> + +<p><i>Diagnosis.</i>—Medium sized frogs, only known male 25.6 mm. snout-vent, +females 20.0-26.7 mm. snout-vent length (small sample); vocal slits in males; +<span class="pagenum"><a name="Page_37" id="Page_37">[Pg 37]</a></span> +finger tips expanded; first finger shorter than second; outer +metatarsal tubercle one-third size of inner; skin of dorsum shagreened, +that of venter smooth; diameter of tympanum 37.7-42.4 per cent that of +eye in both sexes; pale yellow-brown ground color mottled with brown; +limb bands broad, much wider than narrow light interspaces; interorbital +bar very long, edged with dark brown to black (<a href="#Fig_18">Fig. 18</a>).</p> + +<p><i>Remarks.</i>—Duellman's (1958) measurements and proportions of <i>S. +interorbitalis</i> were based exclusively on the type series, which is +composed of only females; therefore his <i>interorbitalis</i> data are not +comparable with the data for the other species in his table. Campbell +and Simmons (1962) collected the only known male. The type series was +collected beneath rocks in a stream bed; the collectors heard calling +frogs in the bushes but were unable to obtain specimens (Langebartel and +Shannon, 1956). Campbell and Simmons (1962) reported that their specimen +had a poorly developed interorbital bar in life; in preservative the bar +compares favorably with the bar in the female (<a href="#Fig_18">Fig. 18</a>).</p> + +<p><i>Etymology.</i>—Latin, in reference to the pale interocular band.</p> + +<p><i>Distribution.</i>—Pacific lowlands of Sinaloa, México (<a href="#Fig_20">Fig. 20</a>).</p> + +<p><i>Specimens examined.</i>—(10). MÉXICO, <i>Sinaloa</i>: 36 mi. N Mazatlán, UIMNH +38094-96, 67061 (holotype), 71970-74; 65 mi. N Mazatlán, LACM 13773.</p> +<br /> + +<div class="caption3nb"> +<a name="Syrrhophus_modestus" id="Syrrhophus_modestus"></a> +<b>Syrrhophus modestus</b> Taylor</div> + +<div class="species_ref"><i>Syrrhophus modestus</i> Taylor, 1942:304-06, pl. 29 [Holotype.—FMNH +100048 (formerly EHT-HMS 3756), from Hacienda Paso del Río, Colima, +México; collected on July 8, 1935, by Hobart M. Smith]. Smith and +Taylor, 1948:49-50.</div> +<div class="species_ref"><i>Syrrhophus modestus modestus</i>: Duellman, 1958:2-5, 7, 14, pl. 1, fig. +1. Gorham, 1966:166.</div> + +<p><i>Diagnosis.</i>—Small frogs, males 15.8-20.1 mm. snout-vent length, single +female 18.5 mm.; vocal slits present in males; finger tips widely +expanded; first finger shorter than second; inner metatarsal tubercle +about three times size of outer; skin of dorsum shagreened, that of +venter smooth; tympanum concealed; pale cream in preservative with dark +brown spots; limbs banded; bands on forearm and thigh poorly developed +or absent; interorbital bar absent.</p> + +<p><i>Remarks.</i>—The tympanum is concealed in <i>S. modestus</i>, <i>S. +nivocolimae</i>, <i>S. pallidus</i>, <i>S. teretistes</i>, and to a lesser degree in +<i>S. interorbitalis</i>. However, if the specimen is permitted to dry +slightly, the annulus tympanicus becomes visible through the skin and a +tympanum/eye ratio can be computed.</p> + +<p>One of the few cases of sympatry within the genus <i>Syrrhophus</i> involves +this species; <i>modestus</i> and <i>nivocolimae</i> are known to be sympatric at +one locality in southwestern Jalisco, México.</p> + +<p>Duellman (1958) used the trinomial for this population and named a new +subspecies, <i>pallidus</i>, from Nayarit. I consider <i>pallidus</i> to be +specifically distinct from <i>modestus</i> because there is no evidence of +genetic exchange, and there is no overlap in the distinguishing +morphological features. I do consider the two populations to be closely +related but feel the inter-relationships between <i>modestus</i>, <i>pallidus</i>, +<i>nivocolimae</i>, and <i>teretistes</i> are more complex than would be indicated +by the use of trinomials. The sympatric occurrence of <i>modestus</i> and +<i>nivocolimae</i> is significant; morphologically, they might otherwise be +regarded as subspecies. Although allopatric, similar arguments could be +advanced for +<span class="pagenum"><a name="Page_38" id="Page_38">[Pg 38]</a></span> +the morphologically similar <i>pallidus</i> and <i>teretistes</i>. +The four are here afforded species rank since morphological similarity +and allopatry are not sufficient grounds for the assumption of genetic +exchange.</p> +<br /> + +<div class="fig_center" style="width: 527px;"> +<a name="Fig_19" id="Fig_19"></a> +<img src="images/fig_19.png" width="527" height="405" alt="" title="" /> +<div class="fig_caption"><span class="smcap">Fig. 19:</span> <i>Syrrhophus modestus</i> [left, UMMZ 115447 (WED 11155)] and <i>S. +pallidus</i> (right, UMMZ 115453). ×2.2.</div> +</div> + +<p><i>Etymology.</i>—Latin, meaning unassuming, modest, in reference to the +small size of the species.</p> + +<p><i>Distribution.</i>—Low elevations (up to 700 meters) in the lowlands and +foothills of Colima and southwestern Jalisco, México (<a href="#Fig_20">Fig. 20</a>).</p> + +<p><i>Specimens examined.</i>—(14). MÉXICO, <i>Colima</i>: Hda. Paso del Río, FMNH +100048 (holotype), 100167, 100299, UIMNH 13300, UMMZ 110877 (2), USNM +139729; 7.2 km. SW Tecolapa, UMMZ 115477 (4); <i>Jalisco</i>: 17.6 km. SW +Autlan, 606 m., KU 102627; 3.2 km. N La Resolana, UMMZ 102100; Bahía +Tenacatita, UMMZ 84264.</p> +<br /> + +<div class="caption3nb"> +<a name="Syrrhophus_nivocolimae" id="Syrrhophus_nivocolimae"></a> +<b>Syrrhophus nivocolimae</b> Dixon and Webb</div> + +<div class="species_ref"><i>Syrrhophus nivocolimae</i> Dixon and Webb, 1966:1-4, Fig. 1 +[Holotype.—LACM 3200, from Nevado de Colima (6 airline miles west of +Atenquique), Jalisco, México, 7800 feet; collected on July 20, 1964, by +Robert G. Webb].</div> + +<p><i>Diagnosis.</i>—Small frogs, males 18.5-21.1 mm. snout-vent length, only +known female 24.1 mm. snout-vent; vocal slits present in males; finger +tips widely expanded; first finger shorter than second; inner metatarsal +tubercle about three times size of outer; skin of dorsum warty, that of +venter smooth; tympanum concealed, its diameter 30.0-39.3 per cent that +of eye in males; mid-dorsal brown band from interorbital bar to anus; +bands on limbs narrow, dark bands +<span class="pagenum"><a name="Page_39" id="Page_39">[Pg 39]</a></span> +less than one-half width of light +bands, upper arm not banded; narrow interorbital light bar.</p> + +<p><i>Remarks.</i>—This species is closely related to <i>S. modestus</i> and differs +in color pattern and degree of wartiness of the skin. Dixon and Webb +(1966) held that <i>nivocolimae</i> had no close relatives, but the condition +of the tympanum, size, nature of the outer palmar tubercle, relative +sizes of the metatarsal tubercles, and shape and size of the digital +pads all point to a close relationship between <i>S. modestus</i>, <i>S. +nivocolimae</i>, and <i>S. pallidus</i>.</p> +<br /> + +<div class="fig_center" style="width: 529px;"> +<a name="Fig_20" id="Fig_20"></a> +<img src="images/fig_20.png" width="529" height="598" alt="" title="" /> +<div class="fig_caption"><span class="smcap">Fig. 20:</span> Distribution of the species of the <i>modestus</i> group: +<i>interorbitalis</i> (open circles), <i>teretistes</i> (solid circles), +<i>modestus</i> (open triangles), <i>pallidus</i> (solid triangles) and +<i>nivocolimae</i> (square). Arrow indicates locality of sympatry between +<i>modestus</i> and <i>nivocolimae</i>. Solid line about the localities for +<i>interorbitalis</i> is a range estimate based on call records and specimens +examined.</div> +</div> + +<p>Dixon and Webb (1966) reported that <i>S. nivocolimae</i> has a large +tympanum (50.0-59.0 per cent diameter of eye). However, my examination of +the type series and several other specimens from Jalisco reveals that the +<span class="pagenum"><a name="Page_40" id="Page_40">[Pg 40]</a></span> +largest tympanum/eye ratio is 39.3 per cent. Therefore, the +tympanum/eye ratio in <i>S. nivocolimae</i> is in agreement with those for +<i>S. modestus</i>, <i>S. pallidus</i>, and <i>S. teretistes</i> +(<a href="#Table_6">Table 6</a>).</p> + +<p><i>Etymology.</i>—<i>niv</i>, Latin, and Colima (Nevado de), meaning high on the +volcano, in reference to the higher distribution of this species (around +2000 meters) than other members of the group.</p> + +<p><i>Distribution.</i>—Known from southwestern Jalisco, México, at moderate to +high elevations (600-2400 meters).</p> + +<p><i>Specimens examined.</i>—(48) MÉXICO, <i>Jalisco</i>: 17.6 km. SW Autlán, 606 +m., KU 102626, 102631; 6.4 km. W Atenquique, 2060 m., KU 102628-30, +102632; 8 km. W Atenquique, 1970 m., LACM 3210-12; 9.6 km. W Atenquique, +2360 m., LACM 3200 (holotype), 3201-09; 14.5 km. W Atenquique, 2000 m., +LACM 25424-36, 25439-41, 25446; 15 km. W Atenquique, LACM 37044-46, +37244-47; 16 km. W Atenquique, 2105 m., LACM 25443-45; 17 km. W +Atenquique, 2180 m., LACM 25442.</p> +<br /> + +<div class="caption3nb"> +<a name="Syrrhophus_pallidus" id="Syrrhophus_pallidus"></a> +<b>Syrrhophus pallidus</b> Duellman, New combination</div> + +<div class="species_ref"><i>Syrrhophus modestus</i>: Davis and Dixon, 1957:146.</div> +<div class="species_ref"><i>Syrrhophus modestus pallidus</i> Duellman, 1958:2-3, 5-7, 14, pl. 3 +[Holotype.—UMMZ 115452, from San Blas, Nayarit, México, sea level; +collected on August 13, 1956, by William E. and Ann S. Duellman]. +Zweifel, 1960:86-88, 91, 93-94, 118, 120-22. Gorham, 1966:166.</div> +<div class="species_ref"><i>Syrrhophis modestus pallidus</i>: Campbell and Simmons, 1962:194.</div> + +<p><i>Diagnosis.</i>—Small frogs, males 17.9-19.3 mm. snout-vent length; vocal +slits in males; finger tips widely expanded; first finger shorter than +second; inner metatarsal tubercle about three times size of outer; skin +of dorsum shagreened, that of venter smooth; tympanum concealed, its +diameter 27.0-35.6 per cent of eye in males; ground color cream +vermiculated with brown, upper arm and thigh lacking, or with few, +indistinct, bands; interorbital bar absent.</p> + +<p><i>Remarks.</i>—Considerable debate has been waged relative to the value of +subspecies and to the reasons for recognizing distinct disjunct +populations as species versus subspecies. Lacking evidence of genetic +exchange, I prefer to retain disjunct populations that are distinctive +as species.</p> + +<p>All known specimens of <i>pallidus</i> can be separated from those of +<i>modestus</i> by color pattern. The two nominal species exhibit overlap in +proportions but the same can be said about nearly every species of +<i>Syrrhophus</i>; therefore, overlap in proportions can be disregarded in +assessing specific versus subspecific rank. Until contrary evidence is +forthcoming, I consider the disjunct populations heretofore held to be +subspecies of <i>modestus</i> to be specifically distinct. The specimens of +the disjunct population of <i>pallidus</i> on the Tres Marias do not differ +from the mainland population in Nayarit. This evidence, though perhaps +secondary, supports my contention that two species should be recognized.</p> + +<p><i>Etymology.</i>—Latin, in reference to the pale ground color in comparison +with that of <i>S. modestus</i>.</p> + +<p><i>Distribution.</i>—Low elevations in coastal Nayarit and on Islas Tres +Marias (<a href="#Fig_20">Fig. 20</a>).</p> + +<p><i>Specimens examined.</i>—(12) MÉXICO, <i>Nayarit</i>: 18.8 mi. NW Ahuacatlán, +UIMNH 7808; San Blas, UMMZ 115452 (holotype), 115453-57; 17 km. NE San +Blas, 150 m., MSU 5085; 12.8 km. E San Blas, UIMNH 71979; 31 km. E San +Blas, UIMNH 71978; 13.5 km. N Tepic, UIMNH 71980-81.</p> +<br /> + +<p><span class="pagenum"><a name="Page_41" id="Page_41">[Pg 41]</a></span></p> + +<div class="caption3nb"> +<a name="Syrrhophus_teretistes" id="Syrrhophus_teretistes"></a> +<b>Syrrhophus teretistes</b> Duellman</div> + +<div class="species_ref"><i>Syrrhophus teretistes</i> Duellman, 1958:2-3, 10-14, pl. 2, fig. 2 +[Holotype.—UMMZ 115451, from 4.8 km. NW Tepic, Nayarit, México, 840 m.; +collected on August 12, 1956, by William E. Duellman]. Gorham, 1966:167.</div> + +<p><i>Diagnosis.</i>—Medium-sized frogs, males 19.2-23.2 mm. snout-vent length, +single known female 24.8 mm. snout-vent; vocal slits in males; finger +tips widely expanded; first finger shorter than second; inner metatarsal +tubercle about three times size of outer; skin of dorsum shagreened, +that of venter smooth; tympanum partially concealed, its diameter +28.6-43.8 per cent of eye in males; ground color brown vermiculated with +dark brown to nearly black; upper arm and thigh banded; interorbital +light bar absent.</p> + +<p><i>Remarks.</i>—<i>S. teretistes</i> appears to be most closely related to <i>S. +pallidus</i>; I consider it to be an upland derivative of <i>pallidus</i>. +Morphologically, the differences between the two are few, but lacking +evidence of genetic exchange they are retained as species.</p> + +<p><i>Etymology.</i>—Greek, in reference to the whistle-like nature of the +call.</p> + +<p><i>Distribution.</i>—Moderate elevations (840-1200 meters) in the Sierra +Occidental of Nayarit, Sinaloa, and Durango, México (<a href="#Fig_20">Fig. 20</a>).</p> + +<p><i>Specimens examined.</i>—(13) MÉXICO, <i>Nayarit</i>: 4.8 km. NW Tepic, 840 m., +UMMZ 115451 (holotype). <i>Sinaloa</i>: Santa Lucía, 1090 m., KU 75263-72; 1 +km. NE Santa Lucía, 1156 m., KU 78257; 2.2 km. NE Santa Lucía, 1156 m., +KU 78258.</p> +<br /> + + +<div class="smcap caption2nb">Discussion</div> + +<p>There are relatively few clear-cut morphological differences among the +fourteen species now assigned to <i>Syrrhophus</i>. The majority of the +species are allopatric and differ primarily in color patterns. Sympatric +occurrence serves as an indicator of specific distinctness and is one of +the more practical tests of species validity when cross-breeding +experiments are not possible. Two cases of sympatric occurrence are +known for the species of <i>Syrrhophus</i> in western México: <i>modestus</i> and +<i>nivocolimae</i> are sympatric in southern Jalisco and <i>pipilans nebulosus</i> +and <i>rubrimaculatus</i> are sympatric in southeastern Chiapas. In eastern +México, <i>longipes</i> and <i>verrucipes</i> are sympatric in southern Hidalgo, +and <i>longipes</i> is sympatric with <i>cystignathoides</i>, <i>dennisi</i>, and +<i>guttilatus</i> in southern Tamaulipas. <i>Syrrhophus cystignathoides</i> and +<i>leprus</i> are apparently sympatric in central Veracruz.</p> + +<p>Subspecific assignments have been made only when there is evidence of +intergradation. The sympatric occurrence of morphologically similar +species in this genus has led me to adopt a conservative approach to the +degree of difference philosophy. I have therefore recognized all +morphologically distinct allopatric populations as species.</p> + +<p><span class="pagenum"><a name="Page_42" id="Page_42">[Pg 42]</a></span></p> + +<div class="fig_center" style="width: 500px;"> +<a name="Fig_21" id="Fig_21"></a> +<img src="images/fig_21.png" width="500" height="392" alt="" title="" /> +<div class="fig_caption"><span class="smcap">Fig. 21:</span> Generic distributions of <i>Syrrhophus</i> (stipple) and +<i>Tomodactylus</i> (hatching). Black areas are zones of intergeneric +sympatry.</div> +</div> + +<div class="fig_left" style="width: 295px;"> +<a name="Fig_22" id="Fig_22"></a> +<img src="images/fig_22.png" width="295" height="437" alt="" title="" /> +<div class="fig_caption"><span class="smcap">Fig. 22:</span> Altitudinal distributions of <i>Syrrhophus</i> and <i>Tomodactylus</i>. +Widths of the columns are proportional to the numbers of species at a +given altitude; narrowest width equals one species.</div> +</div> + +<p><i>Syrrhophus</i> is closely allied to another Mexican leptodactylid genus, +<i>Tomodactylus</i>, which was revised by Dixon (1957), who along with +numerous other authors noted the close relationship between the two +genera. There is an almost complete lack of sympatry between the two +genera; in very few places in México do they coexist (<a href="#Fig_21">Fig. 21</a>). +<i>Tomodactylus</i> has its greatest diversity in the Cordillera Volcánica +and Sierra Madre del Sur, whereas <i>Syrrhophus</i> reaches its greatest +diversity in the Sierra Madre Oriental and eastern foothills. The +species of both genera are about the same size and presumably have +similar requirements insofar as food, breeding sites, and habitat +selection.</p> + +<p>Four cases of intergeneric sympatry are known for the two genera: 1) +the Chilpancingo region of Guerrero, 2) the lowlands of Colima and the +mountains just inland in Jalisco, 3) the lowlands of central Nayarit, +and 4) the Sierra Madre Occidental on the Durango-Sinaloan border. The +apparent sympatry in the Chilpancingo region involves four species: <i>S. +pipilans</i>, <i>T. albolabris</i>, <i>T. dilatus</i>, and <i>T. nitidus</i>. Of the four, +<i>T. dilatus</i> appears to be completely allopatric in that it occurs at +higher altitudes (above 2000 meters), whereas the other three occur +below 1800 meters in the region (Davis and Dixon, 1965). In the +Colima-Jalisco region, <i>Tomodactylus</i> tends +<span class="pagenum"><a name="Page_43" id="Page_43">[Pg 43]</a></span> +to occur higher (Dixon and Webb, 1966) than some of the <i>Syrrhophus</i>, +but one subspecies of <i>Tomodactylus nitidus</i> is a lowland frog, +occurring sympatrically with the lowland <i>Syrrhophus modestus</i>. A +similar situation is observed in Nayarit; the lowland <i>Tomodactylus</i> +occurs sympatrically with the small <i>Syrrhophus pallidus</i>. In both cases +the <i>Syrrhophus</i> is smaller than the <i>Tomodactylus</i>.</p> + +<p>Frogs of the genus <i>Syrrhophus</i> tend to occur at lower elevations than +do their close relatives of the genus <i>Tomodactylus</i> (<a href="#Fig_22">Fig. 22</a>). This +generalization is complicated by the occurrence in the Sierra Madre +Oriental in relatively high altitude <i>Syrrhophus</i> (up to 2000 m.) and +the occurrence in Michoacán of low altitude <i>Tomodactylus</i> (to sea +level). There are no <i>Tomodactylus</i> in the Sierra Madre Oriental, +whereas the genus <i>Syrrhophus</i> is represented in the lowlands of western +México (<i>modestus</i> group). <i>Syrrhophus</i> and <i>Tomodactylus</i> exhibit +essentially parapatric distributions. The two genera as now composed can +be characterized as low to moderate elevation frogs (<i>Syrrhophus</i>) and +moderate to intermediate elevation frogs (<i>Tomodactylus</i>).</p> +<br /> +<br /> + +<div style="display: block; clear: both;"> +<br /> +<br /> +</div> +<div class="smcap caption2nb">Literature Cited</div> +<br /> +<br /> + +<div class="smcap">Baird, S. F.</div> +<div class="reference">1859. Reptiles of the Boundary. United States and Mexican Boundary +Survey, pp. 1-35, pls. 1-41.</div> +<br /> + +<div class="smcap">Barbour, T.</div> +<div class="reference">1923. The reappearance of Batrachyla longipes. Proc. New England Zool. +Club, 8:81-83.</div> +<br /> + +<div><span class="smcap">Barbour, T.</span>, and <span class="smcap">A. Loveridge</span></div> +<div class="reference">1946. Typical reptiles and amphibians; supplement. Bull. Mus. Comp. +Zool., 96:59-214.</div> +<br /> + +<div class="smcap">Boulenger, G. A.</div> +<div class="reference">1882. Catalogue of the Batrachia Salientia ... British Museum., 2nd ed.</div> +<div class="reference">1888. Note on the classification of the Ranidae. Proc. Zool. Soc. London, +1888, pt. 2:204-06.</div> +<br /> + +<p><span class="pagenum"><a name="Page_44" id="Page_44">[Pg 44]</a></span></p> + +<div><span class="smcap">Campbell, H. W.</span>, and <span class="smcap">R. S. Simmons</span></div> +<div class="reference">1962. Notes on some reptiles and amphibians from western Mexico. Bull. +So. California Acad. Sci., 61:193-203.</div> +<br /> + +<div class="smcap">Conant, R.</div> +<div class="reference">1958. A field guide to reptiles and amphibians. Houghton-Mifflin Co. +Boston. 366 pp.</div> +<br /> + +<div class="smcap">Cope, E. D.</div> +<div class="reference">1866. On the structures and distribution of the genera of the arciferous +Anura. J. Acad. Nat. Sci. Philadelphia, n. ser., 6:67-112.</div> +<div class="reference">1877. Tenth contribution to the herpetology of tropical America. Proc. +Amer. Philos. Soc., 17:85-98.</div> +<div class="reference">1878. New genus of Cystignathidae from Texas. Amer. Nat., 12:252-53.</div> +<div class="reference">1879. Eleventh contribution to the herpetology of tropical America. Proc. +Amer. Philos. Soc., 18:261-77.</div> +<div class="reference">1885. A contribution to the herpetology of Mexico. <i>Ibid.</i>, 22:379-404.</div> +<br /> + +<div><span class="smcap">Davis, W. B.</span>, and <span class="smcap">J. R. Dixon</span></div> +<div class="reference">1957. Notes on Mexican amphibians, with description of a new <i>Microbatrachylus</i>. +Herpetologica, 13:145-47.</div> +<div class="reference">1965. Amphibians of the Chilpancingo Region, Mexico. <i>Ibid.</i>, 20:225-33.</div> +<br /> + +<div class="smcap">Díaz de León, J.</div> +<div class="reference">1904. Indice de los Batracios que se enquentran en la Republica Méxicana. +Imprenta de Ricardo Rodriquez Romo. Aguascalientes. 40 pp.</div> +<br /> + +<div class="smcap">Dixon, J. R.</div> +<div class="reference">1957. Geographic variation and distribution of the genus Tomodactylus +in Mexico. Texas J. Sci., 9:379-409.</div> +<br /> + +<div><span class="smcap">Dixon, J. R.</span>, and <span class="smcap">R. G. Webb</span></div> +<div class="reference">1966. A new <i>Syrrhophus</i> from Mexico (Amphibia: Leptodactylidae). +Cont. Sc., Los Angeles Co. Mus., 102:1-5.</div> +<br /> + +<div class="smcap">Duellman, W. E.</div> +<div class="reference">1958. A review of the frogs of the genus <i>Syrrhophus</i> in western Mexico. +Occ. Pap. Mus. Zool. Univ. Michigan, 594:1-15.</div> +<div class="reference">1960. A distributional study of the amphibians of the Isthmus of Tehuantepec, +Mexico. Univ. Kansas Publs. Mus. Nat. Hist., 13:19-72.</div> +<br /> + +<div class="smcap">Firschein, I. L.</div> +<div class="reference">1954. Definition of some little-understood members of the leptodactylid +genus <i>Syrrhophus</i>, with a description of a new species. Copeia, +(1):48-58.</div> +<br /> + +<div class="smcap">Fouquette, M. J.</div> +<div class="reference">1960. Call structure in frogs of the family Leptodactylidae. Texas J. Sci., +12:201-15.</div> +<br /> + +<div class="smcap">Gadow, H.</div> +<div class="reference">1905. The distribution of Mexican amphibians and reptiles. Proc. Zool. +Soc. London, 1905, pt. 2:191-244.</div> +<br /> + +<div class="smcap">Gorham, S. W.</div> +<div class="reference">1966. Liste der rezenten Amphibien und Reptilien.... Das Tierreich. +Lief, 85:1-222.</div> +<br /> + +<div class="smcap">Günther, A. C. L. G.</div> +<div class="reference">1885-1902. Biologia Centrali-Americana. Reptilia and Batrachia. 326 pp., +76 pls. Syrrhophus section dated 1900.</div> +<br /> + +<div class="smcap">Kellogg, R.</div> +<div class="reference">1932. Mexican tailless amphibians in the United States National Museum. +Bull. U.S. Natl. Mus., 160.</div> +<br /> + +<div><span class="smcap">Langebartel, D. A.</span>, and <span class="smcap">F. A. Shannon</span></div> +<div class="reference">1956. A new frog (Syrrhophus) from the Sinoloan lowlands of Mexico. +Herpetologica, 12:161-65.</div> +<br /> + +<div class="smcap">Lynch, J. D.</div> +<div class="reference">1963. The status of <i>Eleutherodactylus longipes</i> (Baird) of Mexico +(Amphibia: Leptodactylidae). Copeia, (3):580-81.</div> +<div class="reference">1964. Additional hylid and leptodactylid remains from the Pleistocene of +Texas and Florida Herpetologica. 20:141-42.</div> +<div class="reference">1967. <i>Epirhexis</i> Cope, 1866 (Amphibia: Salientia): request for suppression +under the plenary powers. I. N. (S). Bull. Zool. Nomencl., +24:313-15.</div> +<p><span class="pagenum"><a name="Page_45" id="Page_45">[Pg 45]</a></span></p> +<div class="reference">1968. Genera of leptodactylid frogs in México. Univ. Kansas Publs., Mus. +Nat. Hist., 17:503-15.</div> +<br /> + +<div class="smcap">Martin, P. S.</div> +<div class="reference">1958. A biogeography of reptiles and amphibians in the Gomez Farias +region, Tamaulipas, Mexico. Misc. Publs. Mus. Zool. Univ. Michigan, +101:1-102.</div> +<br /> + +<div><span class="smcap">Milstead, W. M.</span>, <span class="smcap">J. S. Mecham</span>, and <span class="smcap">H. McClintock</span></div> +<div class="reference">1950. The amphibians and reptiles of the Stockton Plateau in northern +Terrell County, Texas. Texas J. Sci., 2:543-62.</div> +<br /> + +<div class="smcap">Neill, W. T.</div> +<div class="reference">1965. New and noteworthy amphibians and reptiles from British Honduras. +Bull. Florida State Mus., 9:77-130.</div> +<br /> + +<div class="smcap">Nieden, F.</div> +<div class="reference">1923. Anura I ... Das Tierreich. Lief., 46:1-584.</div> +<br /> + +<div class="smcap">Peters, W.</div> +<div class="reference">1871. Über neue Amphibien ... des Konigl. Zoologischen Museums. +Monatsb. k. k. Preuss. Akad. Wiss. Berlin, 1870:641-52.</div> +<br /> + +<div><span class="smcap">Schmidt, K. P.</span>, and <span class="smcap">T. F. Smith</span></div> +<div class="reference">1944. Amphibians and reptiles of the Big Bend Region of Texas. Field +Mus. Nat. Hist., zool. ser., 29:75-96.</div> +<br /> + +<div class="smcap">Smith, H. M.</div> +<div class="reference">1947. Notes on Mexican amphibians and reptiles. J. Washington Acad. +Sci., 37:408-12.</div> +<br /> + +<div><span class="smcap">Smith, H. M.</span>, and <span class="smcap">E. H. Taylor</span></div> +<div class="reference">1948. An annotated checklist and key to the Amphibia of Mexico. Bull. +U.S. Natl. Mus., 194:1-118.</div> +<br /> + +<div class="smcap">Stejneger, L.</div> +<div class="reference">1915. A new species of tailless batrachian from North America. Proc. Biol. +Soc. Washington, 28:131-32.</div> +<br /> + +<div class="smcap">Taylor, E. H.</div> +<div class="reference">1940a. A new eleutherodactylid frog from Mexico. Proc. New England +Zool. Club, 18:13-16.</div> +<div class="reference">1940b. Two new anuran amphibians from Mexico. Proc. U.S. Natl. Mus., +89:43-47, 1 pl.</div> +<div class="reference">1940c. A new Syrrhophus from Guerrero, Mexico. Proc. Biol. Soc. Washington, +53:95-98, 1 pl.</div> +<div class="reference">1940d. New species of Mexican Anura. Univ. Kansas Sci. Bull., 26:385-405.</div> +<div class="reference">1940e. Herpetological miscellany no. I. <i>Ibid.</i>, 26:489-571.</div> +<div class="reference">1942. New Caudata and Salientia from México. <i>Ibid.</i>, 28:295-323.</div> +<div class="reference">1943. Herpetological novelties from Mexico. <i>Ibid.</i>, 29:343-61.</div> +<div class="reference">1952. A review of the frogs and toads of Costa Rica. <i>Ibid.</i>, 35:577-942.</div> +<br /> + +<div><span class="smcap">Taylor, E. H.</span>, and <span class="smcap">H. M. Smith</span></div> +<div class="reference">1945. Summary of the collections of amphibians made in Mexico under +the Walter Rathbone Bacon Traveling Scholarship. Proc. U.S. +Natl. Mus., 95:521-613.</div> +<br /> + +<div class="smcap">Tihen, J. A.</div> +<div class="reference">1960. Notes on Late Cenozoic hylid and leptodactylid frogs from Kansas, +Oklahoma and Texas. Southwest. Nat., 5:66-70.</div> +<br /> + +<div><span class="smcap">Wright, A. H.</span>, and <span class="smcap">A. A. Wright</span></div> +<div class="reference">1949. Handbook of frogs and toads. 3rd ed. Comstock. 640 pp.</div> +<br /> + +<div class="smcap">Yarrow, H. C.</div> +<div class="reference">1882. Checklist of North American Reptilia and Batrachia, with catalogue +of specimens in U.S. National Museum. Bull. U.S. Natl. Mus., +24:1-249.</div> +<br /> + +<div class="smcap">Zweifel, R. G.</div> +<div class="reference">1960. Results of the Puritan-American Museum of Natural History Expedition +to Western Mexico. 9. Herpetology of the Tres Marias +Islands. Bull. Amer. Mus. Nat. Hist., 119:81-128.</div> +<br /> +<br /> +<br /> +<br /> + +<div class="trans_notes"> +<div class="caption2">Transcriber's Notes</div> + +<p>Although <i>Syrrhophus marnocki</i> and <i>Syrrhophus marnockii</i> +both appear in this text, a literature search shows that both +spellings have been used and the two instances where there is only +one "i" at the end are in reference to priviously published names. +Therefore, they were left as is. With the exception of the list below +and a number of silent corrections, the text presented is that of the +original printed version. The original cover was modified to +include graphics from the article.</p> + +<div class="caption2">Typographical Corrections</div> + +<table summary="typos"> +<tr> + <td class="brdbt2">Page</td> + <td> </td> + <td class="brdbt2">Correction</td> +</tr> +<tr> + <td> 3</td> + <td> </td> + <td>otherwse => otherwise</td> +</tr> +<tr> + <td> 5</td> + <td> </td> + <td>poltypic => polytypic</td> +</tr> +<tr> + <td>12</td> + <td> </td> + <td>interorbtal => interorbital</td> +</tr> +<tr> + <td>14</td> + <td> </td> + <td>neublosus => nebulosus</td> +</tr> +<tr> + <td>16</td> + <td> </td> + <td>Cuidad => Ciudad</td> +</tr> +<tr> + <td>16</td> + <td> </td> + <td>1946-170 => 1946:170</td> +</tr> +<tr> + <td>22</td> + <td> </td> + <td>rubrimacultaus => rubrimaculatus</td> +</tr> +<tr> + <td>27</td> + <td> </td> + <td>resemblence => resemblance</td> +</tr> +</table> +<br /> +<br /> +</div> +</div><!-- End book --> + + + + + + + + +<pre> + + + + + +End of the Project Gutenberg EBook of A Taxonomic Revision of the +Leptodactylid Frog Genus Syrrhophus Cope, by John D. Lynch + +*** END OF THIS PROJECT GUTENBERG EBOOK A TAXONOMIC REVISION OF THE *** + +***** This file should be named 37809-h.htm or 37809-h.zip ***** +This and all associated files of various formats will be found in: + http://www.gutenberg.org/3/7/8/0/37809/ + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper 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 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 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 +business@pglaf.org. 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 + gbnewby@pglaf.org + + +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. + + +</pre> + +</body> +</html> diff --git a/37809-h/images/bar_double.png b/37809-h/images/bar_double.png Binary files differnew file mode 100644 index 0000000..f2422e6 --- /dev/null +++ b/37809-h/images/bar_double.png diff --git a/37809-h/images/bar_single.png b/37809-h/images/bar_single.png Binary files differnew file mode 100644 index 0000000..1496c61 --- /dev/null +++ b/37809-h/images/bar_single.png diff --git a/37809-h/images/cover.jpg b/37809-h/images/cover.jpg Binary files differnew file mode 100644 index 0000000..6ac09d9 --- /dev/null +++ b/37809-h/images/cover.jpg diff --git a/37809-h/images/fig_1.png b/37809-h/images/fig_1.png Binary files differnew file mode 100644 index 0000000..a2e00bf --- /dev/null +++ b/37809-h/images/fig_1.png diff --git a/37809-h/images/fig_10.png b/37809-h/images/fig_10.png Binary files differnew file mode 100644 index 0000000..4781ba7 --- /dev/null +++ b/37809-h/images/fig_10.png diff --git a/37809-h/images/fig_11.png b/37809-h/images/fig_11.png Binary files differnew file mode 100644 index 0000000..729f613 --- /dev/null +++ b/37809-h/images/fig_11.png diff --git a/37809-h/images/fig_12.png b/37809-h/images/fig_12.png Binary files differnew file mode 100644 index 0000000..d1934b5 --- /dev/null +++ b/37809-h/images/fig_12.png diff --git a/37809-h/images/fig_13.png b/37809-h/images/fig_13.png Binary files differnew file mode 100644 index 0000000..2a7fd51 --- /dev/null +++ b/37809-h/images/fig_13.png diff --git a/37809-h/images/fig_14.png b/37809-h/images/fig_14.png Binary files differnew file mode 100644 index 0000000..c8f8958 --- /dev/null +++ b/37809-h/images/fig_14.png diff --git a/37809-h/images/fig_15.png b/37809-h/images/fig_15.png Binary files differnew file mode 100644 index 0000000..2d55fd4 --- /dev/null +++ b/37809-h/images/fig_15.png diff --git a/37809-h/images/fig_16.png b/37809-h/images/fig_16.png Binary files differnew file mode 100644 index 0000000..368d84a --- /dev/null +++ b/37809-h/images/fig_16.png diff --git a/37809-h/images/fig_17.png b/37809-h/images/fig_17.png Binary files differnew file mode 100644 index 0000000..7aded32 --- /dev/null +++ b/37809-h/images/fig_17.png diff --git a/37809-h/images/fig_18.png b/37809-h/images/fig_18.png Binary files differnew file mode 100644 index 0000000..fe335d4 --- /dev/null +++ b/37809-h/images/fig_18.png diff --git a/37809-h/images/fig_19.png b/37809-h/images/fig_19.png Binary files differnew file mode 100644 index 0000000..2cd997e --- /dev/null +++ b/37809-h/images/fig_19.png diff --git a/37809-h/images/fig_2.png b/37809-h/images/fig_2.png Binary files differnew file mode 100644 index 0000000..2f52bd2 --- /dev/null +++ b/37809-h/images/fig_2.png diff --git a/37809-h/images/fig_20.png b/37809-h/images/fig_20.png Binary files differnew file mode 100644 index 0000000..746f2e7 --- /dev/null +++ b/37809-h/images/fig_20.png diff --git a/37809-h/images/fig_21.png b/37809-h/images/fig_21.png Binary files differnew file mode 100644 index 0000000..e92ef40 --- /dev/null +++ b/37809-h/images/fig_21.png diff --git a/37809-h/images/fig_22.png b/37809-h/images/fig_22.png Binary files differnew file mode 100644 index 0000000..b3690ba --- /dev/null +++ b/37809-h/images/fig_22.png diff --git a/37809-h/images/fig_3.png b/37809-h/images/fig_3.png Binary files differnew file mode 100644 index 0000000..1d45335 --- /dev/null +++ b/37809-h/images/fig_3.png diff --git a/37809-h/images/fig_4.png b/37809-h/images/fig_4.png Binary files differnew file mode 100644 index 0000000..fe48e67 --- /dev/null +++ b/37809-h/images/fig_4.png diff --git a/37809-h/images/fig_5.png b/37809-h/images/fig_5.png Binary files differnew file mode 100644 index 0000000..2c41dc4 --- /dev/null +++ b/37809-h/images/fig_5.png diff --git a/37809-h/images/fig_6.png b/37809-h/images/fig_6.png Binary files differnew file mode 100644 index 0000000..9525b46 --- /dev/null +++ b/37809-h/images/fig_6.png diff --git a/37809-h/images/fig_7.png b/37809-h/images/fig_7.png Binary files differnew file mode 100644 index 0000000..045911d --- /dev/null +++ b/37809-h/images/fig_7.png diff --git a/37809-h/images/fig_8.png b/37809-h/images/fig_8.png Binary files differnew file mode 100644 index 0000000..5339d77 --- /dev/null +++ b/37809-h/images/fig_8.png diff --git a/37809-h/images/fig_9.png b/37809-h/images/fig_9.png Binary files differnew file mode 100644 index 0000000..24c4435 --- /dev/null +++ b/37809-h/images/fig_9.png diff --git a/37809.txt b/37809.txt new file mode 100644 index 0000000..d5f3521 --- /dev/null +++ b/37809.txt @@ -0,0 +1,2535 @@ +The Project Gutenberg EBook of A Taxonomic Revision of the Leptodactylid +Frog Genus Syrrhophus Cope, by John D. Lynch + +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: A Taxonomic Revision of the Leptodactylid Frog Genus Syrrhophus Cope + +Author: John D. Lynch + +Release Date: October 21, 2011 [EBook #37809] + +Language: English + +Character set encoding: ASCII + +*** START OF THIS PROJECT GUTENBERG EBOOK A TAXONOMIC REVISION OF THE *** + + + + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper and +the Online Distributed Proofreading Team at +http://www.pgdp.net + + + + + + + + + + UNIVERSITY OF KANSAS + MUSEUM OF NATURAL HISTORY + + Vol. 20, No. 1, pp. 1-45, 22 figs. + + February 20, 1970 + + A Taxonomic Revision + of the Leptodactylid Frog Genus + Syrrhophus Cope + + BY + + JOHN D. LYNCH + + UNIVERSITY OF KANSAS + LAWRENCE + 1970 + + + + + UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY + + Editors of this number: + Frank B. Cross, Philip S. Humphrey, William E. Duellman + + Volume 20, No. 1, pp. 1-45, 22 figs. + Published February 20, 1970 + + UNIVERSITY OF KANSAS + Lawrence, Kansas + + PRINTED BY + THE UNIVERSITY OF KANSAS PRINTING SERVICE + LAWRENCE, KANSAS + 1970 + + + + +A Taxonomic Revision of the Leptodactylid Frog Genus Syrrhophus Cope + +BY + +JOHN D. LYNCH + + + + +INTRODUCTION + + +Cope (1878) proposed the genus _Syrrhophus_ for a medium-sized +leptodactylid frog from central Texas; in the ensuing 75 years the genus +was expanded to include a heterogeneous group of frogs ranging from +Texas to Peru. Taylor (1952) and Firschein (1954) limited the genus to +several species of frogs occurring in Guatemala, Mexico, and Texas. +Lynch (1968) provided a definition of the previously loosely-defined +genus. + +With the exception of Taylor (1952), who treated the Costa Rican +species, none of these authors dealt with the present status of the +nineteen species erroneously assigned to _Syrrhophus_. These species are +listed in Tables 1 and 2 with the name currently applied. Some of them +are new combinations and their justifications will be published +elsewhere. Gorham (1966) is the most recent author to include South +American species in the genus _Syrrhophus_. + +Smith and Taylor (1948) recognized two species groups of the genus in +Mexico, an eastern and a western group (here termed complexes for +purposes of discussion), separated on the basis of the number of palmar +(metacarpal) tubercles (three palmar tubercles in the members of the +eastern complex and two in those of the western complex). Duellman +(1958) reviewed the species of the genus occurring in western Mexico and +concluded that there were five species (two polytypic). Dixon and Webb +(1966) described an additional species from Jalisco, Mexico. The +distributions of some species have been extended, but otherwise the +western complex of species remains unchanged since Duellman's review. + +Smith and Taylor (1948) recognized seven species of the genus in eastern +Mexico. Firschein revised the eastern complex (as then understood), and +in so doing added one new species and treated _Syrrhophus verruculatus_ +as a _nomen dubium_. Dixon (1957) redefined the related genus +_Tomodactylus_ and transferred _T. macrotympanum_ Taylor to the genus +_Syrrhophus_. Neill (1965) described a new subspecies of _S. leprus_ +from British Honduras. Two species (_S. gaigeae_ and _S. marnockii_) +were recognized in Texas until Milstead, Mecham, and McClintock (1950) +synonymized _S. gaigeae_ with _S. marnockii_. Thus, at present, nine +species (one polytypic) are recognized on the eastern slopes and +lowlands from central Texas to British Honduras. These are currently +placed on one species group equivalent to the western complex reviewed +by Duellman (1958). + + + TABLE 1--Species Described as Members of the Genus _Syrrhophus_ but + Now Placed in Other Genera. + + ======================================================================= + Trivial name and author Current combination + ----------------------------------------------------------------------- + _areolatus_ Boulenger, 1898 _Eleutherodactylus areolatus_ + _calcaratus_ Andersson, 1945 _Eleutherodactylus anderssoni_ + _caryophyllaceus_ Barbour, 1928 _Eleutherodactylus caryophyllaceus_ + _coeruleus_ Andersson, 1945 _Eleutherodactylus coeruleus_ + _ineptus_ Barbour, 1928 _Eleutherodactylus diastema_ + _juninensis_ Shreve, 1938 _Eupsophus juninensis_ + _lutosus_ Barbour and Dunn, 1921 _Eleutherodactylus lutosus_ + _molinoi_ Barbour, 1928 _Eleutherodactylus molinoi_ + _montium_ Shreve, 1938 _Niceforonia montia_ + _mystaceus_ Barbour, 1922 _Eleutherodactylus rhodopis_ + _obesus_ Barbour, 1928 _Eleutherodactylus punctariolus_ + _omiltemanus_ Gunther, 1900 _Eleutherodactylus omiltemanus_[1] + _pardalis_ Barbour, 1928 _Eleutherodactylus pardalis_ + ======================================================================= + + [1] New combination. + + + TABLE 2--Species Incorrectly Regarded as Members of the Genus _Syrrhophus_ + but Described as Members of Other Genera. + + ========================================================================== + Trivial name, original generic + assignment, and author Current combination + -------------------------------------------------------------------------- + _chalceus_ (_Phyllobates_) Peters, 1873 _Eleutherodactylus chalceus_ + _festae_ (_Paludicola_) Peracca, 1904 _Niceforonia festae_ + _hylaeformis_ (_Phyllobates_) Cope, 1875 _Eleutherodactylus hylaeformis_ + _palmatus_ (_Phyllobates_) Werner, 1899 _Colostethus palmatus_ + _ridens_ (_Phyllobates_) Cope, 1866 _Eleutherodactylus ridens_ + _simonsii_ (_Paludicola_) Boulenger, 1900 _Niceforonia simonsii_ + ========================================================================== + + + TABLE 3--Nominal Species of _Syrrhophus_ (_sensu strictu_) and the Name + Used Herein. + + ======================================================================= + Original combination Current combination + ----------------------------------------------------------------------- + _campi_, _Syrrhophus_ _cystignathoides campi_ + _cholorum_, _Syrrhophus leprus_ _leprus_ + _cystigathoides_, _Phyllobates_ _cystignathoides cystignathoides_ + _dennisi_, _Syrrhophus_ _dennisi_ new species + _gaigeae_, _Syrrhophus_ _guttilatus_ + _guttilatus_, _Malachylodes_ _guttilatus_ + _interorbitalis_, _Syrrhophus_ _interorbitalis_ + _latodactylus_, _Syrrhophus_ _longipes_ + _leprus_, _Syrrhophus_ _leprus_ + _longipes_, _Batrachyla_ _longipes_ + _macrotympanum_, _Tomodactylus_ _verrucipes_ + _marnockii_, _Syrrhophus_ _marnockii_ + _modestus_, _Syrrhophus_ _modestus_ + _nebulosus_, _Syrrhophus_ _pipilans nebulosus_ + _nivocolimae_, _Syrrhophus_ _nivocolimae_ + _pallidus_, _Syrrhophus modestus_ _pallidus_ + _petrophilus_, _Syrrhophus_ _guttilatus_ + _pipilans_, _Syrrhophus_ _pipilans pipilans_ + _rubrimaculatus_, _Syrrhophus_ _rubrimaculatus_ + _smithi_, _Syrrhophus_ _guttilatus_ + _teretistes_, _Syrrhophus_ _teretistes_ + _verrucipes_, _Syrrhophus_ _verrucipes_ + _verruculatus_, _Phyllobates_ _Nomen dubium_ + ======================================================================= + + +In the course of preparing an account of the species of +_Eleutherodactylus_ occurring in Mexico and northern Central America, it +became necessary to reexamine the status of the genus _Syrrhophus_ and +its nominal species. It soon became evident that there were more names +than species, that some previously regarded species were geographic +variants, and that the eastern and western groups (complexes here) were +artificial divisions of the genus. I conclude that there are seven +species (one polytypic) of _Syrrhophus_ in eastern Mexico, Texas, and El +Peten of Guatemala, and seven species (one polytypic) in western Mexico. +The current status of each of the 23 names correctly assigned to the +genus is presented in Table 3. + +The fourteen species recognized by me are placed in five species groups. +Two of these groups are presently placed in the western complex +(_modestus_ and _pipilans_ groups) and three in the eastern complex +(_leprus_, _longipes_ and _marnockii_ groups). The two complexes do +not correspond exactly with the eastern and western groups of Smith +and Taylor (1948), Firschein (1954), and Duellman (1958) since +_S. rubrimaculatus_ is now associated with the eastern _leprus_ group. + +The definitions and contents of the five species groups are as follows: + + _leprus_ group: digital pads not or only slightly expanded, rounded + in outline; first finger longer or shorter than second; snout + acuminate or subacuminate, not rounded; outer metatarsal tubercle + conical; digits lacking distinct lateral fringes. + + content: _cystignathoides_, _leprus_ and _rubrimaculatus_. + + _longipes_ group: digital pads widely expanded, triangular in outline; + first finger shorter than second; snout acuminate; outer metatarsal + tubercle not conical; digits bearing lateral fringes. + + content: _dennisi_ and _longipes_. + + _marnockii_ group: digital pads expanded, rounded to truncate in + outline; first finger equal in length to second or slightly shorter; + snout rounded; outer metatarsal tubercle not conical; digits lacking + lateral fringes; generally stout-bodied frogs. + + content: _guttilatus_, _marnockii_, and _verrucipes_. + + _modestus_ group: digital pads expanded, truncate in outline; first and + second fingers subequal in length, first usually slightly shorter + than second; snout subacuminate; inner metatarsal tubercle twice as + large (or larger) as outer metatarsal tubercle; digits bearing + poorly-defined lateral fringes. + + content: _interorbitalis_, _modestus_, _nivocolimae_, _pallidus_, + and _teretistes_. + + _pipilans_ group: digital pads not or only slightly expanded, + truncate in outline; first finger equal in length to second; snout + subacuminate; metatarsal tubercles subequal in size; digits lacking + lateral fringes. + + content: _pipilans_. + + +_Acknowledgments._--For loan of specimens, I am indebted to Richard J. +Baldauf, Texas A & M University (TCWC); W. Frank Blair, University of +Texas (TNHC); Charles M. Bogert and Richard G. Zweifel, American Museum +of Natural History (AMNH); James E. Boehlke and Edmond V. Malnate, +Academy of Natural Sciences of Philadelphia (ANSP); Robert F. Inger and +Hymen Marx, Field Museum of Natural History (FMNH); Ernest A. Liner +(EAL); Michael Ovchynnyk, Michigan State University collection (MSU); +James A. Peters, United States National Museum (USNM); Douglas A. +Rossman, Louisiana State University Museum of Zoology (LSUMZ); Hobart M. +Smith, University of Illinois Museum of Natural History (UIMNH); Charles +F. Walker, University of Michigan Museum of Zoology (UMMZ); and John W. +Wright, Los Angeles County Museum (LACM). Specimens in the collection at +the University of Kansas Museum of Natural History are identified as KU. +The abbreviations EHT-HMS refer to the Edward H. Taylor-Hobart M. Smith +collection and FAS to the Frederick A. Shannon collection. The +type-specimens from these collections are now in the Field Museum of +Natural History and the University of Illinois Museum of Natural +History. + +I have profited from discussions concerning this problem with several +persons, most notably William E. Duellman, Hobart M. Smith, Edward H. +Taylor and Charles F. Walker. Nevertheless, the ideas and conclusions +presented here should not be construed as necessarily reflecting their +opinions. + +David M. Dennis executed all of the figures, and my wife, Marsha, typed +the manuscript. + + +_Materials and Methods._--In the course of this study, 1003 specimens +of the genus were examined. The holotypes of 21 of the 23 +nominal species are extant; I have examined 19 of these. Nine +measurements were taken, and five ratios computed for each of 338 +specimens. Females are available for all species but one; thus, +measurements were taken on individuals of both sexes. + + + + +ANALYSIS OF CHARACTERS + + +_Size and proportions._--Frogs of this genus range in size from 16 +to 40 mm. in snout-vent length. Five species are relatively small: +_S. cystignathoides_, _modestus_, _nivocolimae_, _pallidus_ and +_rubrimaculatus_; one, _S. longipes_, is relatively large, and the +remaining eight species are intermediate in size (22-30 mm.). + +Males are generally smaller than females and have proportionately longer +heads and usually larger tympani. No significant differences were found +among proportions, except that _S. longipes_ has a larger tympanum/eye +ratio than any other species. Frogs in the _Syrrhophus marnockii_ group +tend to have shorter shanks and feet, thereby giving those species a +more stocky appearance. However, the differences are not significant. + +A summary of the data on size and proportions for the frogs of the genus +_Syrrhophus_ is given in Tables 4, 5, and 6. + + +_Hands and Feet._--Taylor and Smith (1945), Smith and Taylor (1948), +Firschein (1954) and Duellman (1958) discussed the value of the palmar +tubercles in identifying frogs of this genus. The eastern complex in +general has a well-developed outer palmar tubercle (Fig. 1) in +distinction to the western complex in which the outer palmar tubercle is +reduced or absent (Fig. 2). Dixon and Webb (1966) imply that the outer +palmar tubercle is rarely absent but is usually smaller than the first +supernumerary tubercle of the fourth finger. My study of the western +species demonstrates that the outer palmar tubercle is indeed usually +present and smaller than the first supernumerary tubercle. + +Differences in interpretation of the terms "unexpanded" and "narrow," as +well as differences in techniques of preservation, have led to confusion +of the reported digital shapes in various species. Constant specific +differences are evident in the hands (Fig. 1). Except in the cases of +excessive uptake of fluids, all species have a terminal transverse +groove at the tip of each digit. Taylor (1940b) stated that _S. smithi_ +lacked grooves, but examination of the holotype reveals faint grooves at +the tops of the digits. _Syrrhophus guttilatus_, _leprus_, _pipilans_, +and _verrucipes_ lack lateral fringes on the fingers. Lateral fringes +are well developed in the _longipes_ and _modestus_ groups but poorly +defined or absent in the other members of the genus. The digital pads of +the frogs of the _longipes_ group are much broader than those of the +other species and are narrowest in the frogs of the _leprus_ group. +Supernumerary tubercles are present on the palmar surfaces of all +species of the genus. + + + TABLE 4--Size and Proportions in the Frogs of the _Syrrhophus leprus_ + Group. + + A: _cystignathoides campi_ + B: _c. cystignathoides_ + C: _leprus_ + D: _rubrimaculatus_ + + ========================================================================== + Snout-vent Tibia Head Tympanum/ Eyelid/ + length length/ width/ Eye Interorbital + Species Sex N (SVL) SVL SVL + -------------------------------------------------------------------------- + A [M] 33 16.3-23.5 41.3-49.6 34.0-40.1 43.7-66.5 43.2-89.6 + (45.8) (37.0) (56.2) (61.5) + [F] 12 16.0-25.8 41.5-51.0 33.0-38.0 42.8-60.0 48.2-69.2 + (45.8) (35.0) (51.2) (60.1) + B [M] 15 16.8-22.1 45.1-50.4 33.2-40.7 44.3-68.7 44.6-65.4 + (47.3) (37.8) (54.8) (60.0) + [F] 6 19.6-24.2 46.4-50.0 34.1-38.1 43.3-56.5 53.2-65.4 + (47.6) (36.2) (46.9) (59.2) + C [M] 14 20.6-26.4 42.3-52.3 35.0-40.3 47.5-62.5 58.2-72.5 + (46.8) (37.4) (56.5) (67.3) + [F] 15 22.1-29.2 43.4-53.3 32.6-38.9 38.6-57.9 50.2-86.9 + (47.1) (35.8) (47.1) (68.1) + D [M] 12 18.2-23.5 40.4-46.2 31.8-35.5 35.5-46.5 65.1-78.5 + (43.4) (33.8) (41.7) (71.7) + ========================================================================== + + + TABLE 5--Size and Proportions in the Frogs of the _Syrrhophus longipes_ + and _S. marnockii_ Groups. + + A: _dennisi_ + B: _longipes_ + C: _guttilatus_ + D: _marnockii_ + E: _verrucipes_ + + ========================================================================== + Snout-vent Tibia Head Tympanum/ Eyelid/ + length length/ width/ Eye Interorbital + Species Sex N (SVL) SVL SVL + -------------------------------------------------------------------------- + A [M] 16 22.8-28.4 43.9-49.7 35.3-41.2 53.9-64.2 55.3-74.0 + (47.4) (38.8) (58.9) (65.1) + [F] 10 25.9-32.0 46.3-50.8 35.6-40.3 50.6-58.7 58.1-70.9 + (48.2) (37.7) (54.9) (63.6) + B [M] 22 22.1-33.2 45.8-51.7 38.7-44.4 61.1-87.2 61.5-83.0 + (48.4) (41.8) (72.0) (72.0) + [F] 19 26.8-39.6 44.3-51.0 36.3-40.8 49.5-72.1 55.3-85.9 + (47.2) (39.1) (59.5) (67.9) + C [M] 19 20.6-29.0 41.2-48.1 36.9-44.9 55.1-75.7 53.3-79.5 + (44.5) (40.6) (64.1) (66.0) + [F] 5 25.7-31.0 41.4-46.8 35.9-42.3 47.6-61.7 62.3-79.8 + (43.6) (38.5) (54.0) (72.9) + D [M] 14 18.4-28.9 42.3-47.2 36.1-43.0 47.2-68.3 51.6-74.4 + (44.1) (39.6) (61.2) (66.3) + [F] 29 20.4-35.4 38.7-46.4 35.9-41.3 45.8-73.3 52.1-70.5 + (42.7) (38.2) (60.3) (60.7) + E [M] 29 17.5-29.2 42.7-49.5 36.2-42.4 56.1-82.2 56.8-82.8 + (46.3) (39.1) (67.8) (70.4) + [F] 6 26.5-31.7 42.4-47.7 36.0-38.1 45.8-57.8 61.0-77.9 + (44.6) (37.0) (53.9) (69.0) + ========================================================================== + + + TABLE 6--Size and Proportions in the Frogs of the _Syrrhophus pipilans_ + and _S. modestus_ Groups. + + A: _pipilans nebulosus_ + B: _pipilans pipilans_ + C: _modestus_ + D: _pallidus_ + E: _teretistes_ + F: _nivocolimae_ + G: _interorbitalis_ + + ========================================================================== + Snout-vent Tibia Head Tympanum/ Eyelid/ + length length/ width/ Eye Interorbital + Species Sex N (SVL) SVL SVL + -------------------------------------------------------------------------- + A [M] 17 22.9-28.5 38.1-42.0 34.4-37.2 36.6-47.8 56.1-82.4 + (40.0) (35.4) (43.6) (68.2) + [F] 3 21.1-22.7 42.1-44.5 33.2-35.8 36.6-47.6 64.3-65.4 + + B [M] 18 22.6-27.8 37.9-44.0 32.2-36.5 38.0-54.0 56.1-79.5 + (41.4) (33.0) (46.2) (67.3) + [F] 1 29.4 38.4 32.5 44.6 55.0 + + C [M] 8 15.8-20.1 38.5-42.6 32.1-38.1 26.8-39.3 57.0-86.9 + (40.6) (34.2) (31.5) (69.1) + [F] 1 18.5 44.2 36.0 24.0 52.1 + + D [M] 6 17.9-19.3 41.0-44.9 32.6-36.2 27.0-35.6 59.4-67.7 + (43.4) (35.2) (30.9) (65.2) + + E [M] 18 19.2-23.2 41.5-45.3 32.5-36.4 28.6-43.8 51.2-75.0 + (43.7) (34.0) (33.7) (62.2) + [F] 1 24.8 41.8 30.8 37.9 60.5 + + F [M] 15 18.9-21.1 42.2-48.6 30.9-37.1 30.0-39.3 42.6-69.1 + (45.0) (33.7) (34.7) (55.0) + [F] 1 24.1 40.9 33.5 27.6 56.5 + + G [M] 1 25.6 43.0 ---- 39.4 57.6 + [F] 9 20.2-26.7 39.9-47.1 32.6-39.3 29.1-41.2 58.2-76.9 + (43.2) (35.8) (36.4) (69.2) + ========================================================================== + + + [Illustration: FIG. 1: Palmar views of hands of six species of the + eastern complex of _Syrrhophus_. (A) _verrucipes_ (UIMNH 15995), + (B) _rubrimaculatus_ (KU 58911), (C) _dennisi_ sp. nov. (holotype, + UMMZ 101121), (D) _guttilatus_ (UIMNH 55520), (E) _marnockii_ + (TCWC 4782), and (F) _longipes_ (TCWC 12179). All x6.5.] + + + [Illustration: FIG. 2: Palmar views of hands of two species of the + western complex of _Syrrhophus_. _pipilans_ (left, KU 58908, x6) + and _teretistes_ (center, KU 75269, and right, KU 75263, + respectively, x9).] + + +In _S. cystignathoides_ and _leprus_, the first finger is longer than +the second, and the first two fingers are equal in length in +_guttilatus_ and _marnockii_. In the other species the first finger is +shorter than the second. + +Supernumerary tubercles are well developed on the plantar surfaces in +all species, except _S. guttilatus_, in which they are poorly defined +(Fig. 3). The relative sizes of the metatarsal tubercles has been used +in the classification of the species and species groups of _Syrrhophus_. +The metatarsal tubercles are similar in all species of the eastern +complex (including _rubrimaculatus_); the outer tubercle is always about +one-half the size of the ovoid inner metatarsal tubercle. In the +_leprus_ group the outer tubercle is conical and compressed. The +metatarsal tubercles of _pipilans_ are about the same size, or the outer +is slightly smaller than the inner. In the _modestus_ group the outer +metatarsal tubercle is about one-third the size of the inner. + +All species, except _guttilatus_, have well-defined to poorly defined +lateral fringes on the toes. All species have expanded toe pads. The +fifth toe is usually shorter than the third, but the second is equal +in length to the fifth in some specimens of _S. cystignathoides_ and +_S. marnockii_. _Syrrhophus nivocolimae_ is the only species with +tubercles along the outer edge of the tarsus; this is merely a +reflection of the highly tuberculate nature of the skin in this species. + + +_Skin texture._--The skin of the dorsum is smooth or very weakly +pustular in all species of the genus except _nivocolimae_ and +_verrucipes_. The dorsal surfaces of _nivocolimae_ are warty; in +_verrucipes_ the skin is pustular. The skin of the venter is areolate in +_cystignathoides cystignathoides_, _dennisi_ and _verrucipes_ but is +smooth in all other species of the genus. + + [Illustration: FIG. 3: Plantar views of feet of four species of the + eastern complex of _Syrrhophus_. (A) _guttilatus_ (UIMNH 55519, x6), + (B) _leprus_ (UIMNH 42726, x6), (C) _verrucipes_ (UIMNH 15995, x6), + and (D) _longipes_ (TCWC 12179, x4.6).] + + +_Color pattern._--As is evident in the diagnoses, the color patterns of +given populations have been regarded as useful in separating the +species and subspecies. Duellman (1958) suggested that the coloration, +with the exception of _modestus_, was a dark ground color with pale +markings. It is a moot point whether the frogs have light spots on a +dark background or have a light background with an extensive reticulate +dark pattern. The venters are gray or white, and the vocal sac is nearly +black in some species. Interorbital dark bars or triangles are absent in +only two species of the eastern complex, _cystignathoides campi_ and +_marnockii_; the latter lacks a supratympanic stripe, which is present +in the other members of the eastern complex. _Syrrhophus interorbitalis_ +and _nivocolimae_ have light interorbital bars; these bars occur in only +one other population of the genus (_S. c. cystignathoides_). Bars on the +thighs are ill defined or absent in the members of the _marnockii_ and +part of the _modestus_ groups. The color in life is noted in the species +accounts. + + +_Voice._--The voices of all _Syrrhophus_ can be described as a +single short chirp or peep; without audiospectrographic analyses +the significance of the differences between a chirp, peep, or short +whistle cannot be appreciated. Martin (1958) and Wright and +Wright (1949) reported multi-noted calls, and one collector of +_S. verrucipes_ noted the frog "trilled." + +Fouquette (1960) presented analyses of two species (_marnockii_ +and _pipilans nebulosus_). The voices were very similar; both frogs +were reported to "trill" and "chirp." + + + + +SYSTEMATIC ACCOUNT + + +The genus _Syrrhophus_ has been defined (Lynch, 1968) and limited to the +group of species occurring in Guatemala, Mexico and the United States. +The closest relatives of _Syrrhophus_ are the frogs of the genus +_Tomodactylus_ (Dixon, 1957; Firschein, 1954). Lynch (1968) +implied there were no osteological bases for the separation of +_Eleutherodactylus_, _Syrrhophus_, and _Tomodactylus_. At that time, I +believed such to be the case and derived _Syrrhophus_ and _Tomodactylus_ +from the _rhodopis_ complex of _Eleutherodactylus_, with which they +share terrestrial habits and relatively short limbs. In the _rhodopis_ +complex there is a tendency for the loss of the outer palmar tubercle, a +not uncommon condition in _Syrrhophus_ and _Tomodactylus_. + +However, the skulls of _Syrrhophus_ and _Tomodactylus_ show departures +from the pattern observed in the Middle American _Eleutherodactylus_, as +well as many of those species in western South America. Baldauf and +Tanzer (1965) reported that the frontoparietals and prootics were fused +in _Syrrhophus marnockii_ and that the prootics and exoccipitals +appeared to be one bone (otoccipital). The otoccipital is not uncommon +in eleutherodactyline frogs, but the fusion of the frontoparietals with +the prootics (regardless of the fusion of the latter with the +exoccipital) is uncommon in the family. I have found the +frontoparietal-prootic fusion only in _Syrrhophus_ (all species), +_Tomodactylus_ (all species), and _Eleutherodactylus_ (West Indies +species). None of the Middle American _Eleutherodactylus_ has the two +bones fused. Examination of the character is difficult in dried skeletal +preparations. Cleared and stained or macerated preparations are +satisfactory for checking this character. + +Thus, in addition to the presence of numerous plantar supernumerary +tubercles in the frogs of the genera _Syrrhophus_ and _Tomodactylus_, +these two genera can be separated from other Middle American +eleutherodactylines by the fusion of the frontoparietals and prootics. +This character not only further strengthens the argument that the two +genera are closely related but poses a problem of zoogeographic analysis +of the distribution of the character, which will be discussed fully +elsewhere. + + +Key to the Species of the Frog Genus _Syrrhophus_ + + 1. Three large, well-developed palmar tubercles 2 + + Two large palmar tubercles; outer (third) palmar tubercle reduced + in size or absent 9 + + 2. Digital pads more than twice (usually three or more) times width + of digit 3 + + Digital pads less than twice width of digit 4 + + 3. Males having vocal slits; dorsum vermiculate; diameter of + tympanum in males about one-half diameter of eye _S. dennisi_ + + Males lacking vocal slits; dorsum flecked, spotted, or + blotched; diameter of tympanum in male about three-fourths + that of eye _S. longipes_ + + 4. First finger longer than second 5 + + First finger shorter than or equal to second 7 + + 5. Venter smooth; dorsum spotted or vermiculate _S. leprus_ + + Venter areolate, or if smooth, dorsum flecked and interorbital + bar lacking 6 + + 6. Venter areolate; interorbital bar present; ground color + yellowish _S. cystignathoides cystignathoides_ + + Venter smooth; interorbital bar absent; ground color + brown _S. cystignathoides campi_ + + 7. First finger shorter than second; digital tips only slightly + dilated; green in life with darker green spots _S. verrucipes_ + + First finger equal to second; digital tips slightly to moderately + expanded 8 + + 8. Dorsum vermiculate; interorbital bar present; ground color + cream to brown in life _S. guttilatus_ + + Dorsum punctate or flecked; interorbital bar absent; + ground color green in life _S. marnockii_ + + 9. Dorsum dark with pale (red in life) spots; digital pads + not expanded _S. rubrimaculatus_ + + Dorsum pale with dark markings and digital pads slightly to widely + expanded 10 + + 10. Digital tips not widely expanded; tympanum well-defined; + outer metatarsal tubercle more than one-half size of inner 11 + + Digital tips widely expanded, truncate in outline; tympanum + poorly defined; outer metatarsal tubercle less than one-half + size of inner 12 + + 11. Dorsum dark brown with large light spots or blotches; tympanum/eye + ratio usually greater than 43 percent _S. pipilans pipilans_ + + Dorsum dark brown with small light spots; tympanum/eye + ratio less than 48 percent _S. pipilans nebulosus_ + + 12. Light interorbital bar present 13 + + Light interorbital bar absent 14 + + 13. Adults small, less than 22 mm. snout-vent length with a + broad mid-dorsal stripe; dark bands on shank narrower than + light interspaces _S. nivocolimae_ + + Adults larger, more than 22 mm. snout-vent length; dorsum + vermiculate; dark bands on shank broader than light + interspaces _S. interorbitalis_ + + 14. Dorsum spotted with discrete black spots; pattern + definite _S. modestus_ + + Dorsum reticulate or vermiculate, pattern poorly defined 15 + + 15. Adults small, less than 21 mm. snout-vent length; upper arm + not banded _S. pallidus_ + + Adults larger, usually greater than 21 mm. snout-vent length; + upper arm banded _S. teretistes_ + + + + +SPECIES ACCOUNTS + + +The following accounts do not include complete descriptions of each +taxon, because a more than adequate number of descriptions is available +in the recent (1940-1966) literature. An abbreviated synonymy, in which +are listed all combinations and emendations of names and significant +contributions to our knowledge of the taxon, is given for each. For each +species and subspecies the following are given: descriptive diagnosis, +statement of range, remarks on taxonomy, list of specimens examined, +illustration of color pattern, and distribution map. + + +=Syrrhophus cystignathoides= (Cope) + + _Phyllobates cystignathoides_ Cope, 1877:89-90 + [Syntypes.--Originally USNM 32402-32409, (32405 now in MCZ) + from Potrero, near Cordoba, Veracruz, Mexico, Francis Sumichrast + collector.] + +_Diagnosis._--Adults small, males 16.0 to 23.5 mm. in snout-vent length, +females 16.0-25.8 mm. in snout-vent length; vocal slits present in +males; finger tips slightly expanded; first finger longer than second; +outer metatarsal tubercle one-half size of inner, conical, compressed; +skin of dorsum weakly pustular, that of venter smooth to areolate; +tympanum 44 to 69 per cent diameter of eye (mean 55.5 per cent); ground +color yellow to brown in life with brown to black fleckings on dorsum +and flanks; limbs banded; interorbital bar present or not. + +_Remarks._--Two geographic races (subspecies) are herein recognized; +previously these were held by various authors to be species (_campi_ and +_cystignathoides_). Intergradation occurs in southern Tamaulipas and +eastern San Luis Potosi, Mexico. The two subspecies can be distinguished +on the basis of color pattern and the condition of the skin of the +venter. + +_Distribution._--Low to moderate elevations from the Rio Grande +embayment to central Veracruz, Mexico (Fig. 5). + + +=Syrrhophus cystignathoides campi= Stejneger, New combination + + _Syrrhophus campi_ Stejneger, 1915:131-32. [Holotype.--USNM 52290, + from Brownsville, Cameron Co., Texas; R. D. Camp collector, + March 31, 1915]. Smith and Taylor, 1948:52. Martin, 1958:50. + +_Diagnosis._--Venter smooth; usually no interorbital light and dark bars +present; ground color brown in life (Fig. 4a). + +_Remarks._--Martin (1958) was the first author to point out +that _S. campi_ was probably a subspecies of the more southern +_S. cystignathoides_. Various references in the literature might lead +one to believe that the two were sympatric over much of northeastern +Mexico; this error was created by the use of a single character +(condition of the skin of the venter) to characterize the two +populations. Specimens from southern Texas have a smooth venter, lack +interorbital bars and have, in general, a brown ground color, whereas +specimens from central Veracruz have an areolate venter, interorbital +light and dark bars and a yellow ground color. In southern Tamaulipas +and eastern San Luis Potosi, these characters vary discordantly, thereby +strongly suggesting that the two populations intergrade. Both +populations agree in other morphological characters; therefore, they are +here treated as geographic variants. + +_Etymology._--Named for the collector of the type specimens, Mr. R. D. +Camp of Brownsville, Texas. + +_Distribution._--Lower Rio Grande embayment in Texas to central Nuevo +Leon and Tamaulipas, Mexico. Intergrades are known from southern +Tamaulipas and adjacent San Luis Potosi, Mexico (Fig. 5). + +_Specimens examined._--(113) TEXAS, Cameron Co.: MCZ 10277-85, 10286 +(10); Brownsville, AMNH 3215, 3218-20, 3221 (3), 5376, 62117, FMNH +105336, KU 8135-39, MCZ 3738-42, 3743 (10), TCWC 5908, 7139, TNHC 92-94, +20909, UMMZ 51760, 54031 (5), USNM 52290 (holotype); 22 mi. SE +Brownsville, TNMC 14223; 8 mi. SW Brownsville, UMMZ 101127 (3); +Harlingen, AMNH 62118, UMMZ 105200-205, 105206 (5), 105207 (4). _Hidalgo +Co._: Bentsen-Rio Grande State Park, UMMZ 114378; 6 mi. S McAllen, TNHC +7136-39; Santa Ana Refuge, TCWC 13495-96; Weslaco, TCWC 17658-60. + +MEXICO, _Nuevo Leon_: Salto Cola de Caballo, AMNH 57953-54, FMNH +30644-45, 37169-70; Monterrey, UIMNH 13324; 40 km. SE Monterrey, UIMNH +3686. _Tamaulipas_: 80 km. Matamoros, FMNH 27150 (13). + +Intergrades [_S. c. cystignathoides_ x _S. c. campi_ (88)] MEXICO, _San +Luis Potosi_: 5 km. E Ciudad del Maiz, UMMZ 106435; 16 km. W Naranjo, +FMNH 104584; Salto de Agua, 34 km. WSW Antigua Morelos, TCWC 6980. +_Tamaulipas_: 5 km. W Acuna, 1060 m., UMMZ 101172, 101173 (16), +101174-76, 101177 (6); 14.5 km. NNW Chamal, 430 m., UMMZ 111337 (2); 20 +km. NNW Chamal, 700 m., UMMZ 111338 (11); 8 km. N Gomez Farias, 450 m., +UMMZ 101165; 8 km. NE Gomez Farias, Pano Ayuctle, UMMZ 102264, 102924 +(6); 8 km. NW Gomez Farias, 1060 m., LSUMZ 11084, UMMZ 101199, 102928 +(5), 102929-32, 110124 (3); Rio Guayala, near Magiscatzin, MCZ 24138-42, +85071-81, UMMZ 88242 (2); Magiscatzin, TCWC 6981; Las Yucas, north of +Aldama, MCZ 29665-68; 16 km. NE Zamorina, UMMZ 101124. + + [Illustration: FIG. 4: _Syrrhophus cystignathoides campi_ + (left, TCWC 13490) and _S. c. cystignathoides_ (right, KU 105500). + Dorsal views x2, sides of heads x3.] + + +=Syrrhophus cystignathoides cystignathoides= (Cope), New combination + + _Phyllobates cystignathoides_ Cope, 1877:89-90 [Syntypes.--USNM + 32402-32409, from Potrero, near Cordoba, Veracruz, Mexico, + collected by Francis Sumichrast]. Boulenger, 1882:196. + + _Syrrhophus cystignathoides_: Cope, 1879:268. Kellogg, 1932: + 126-27. Taylor and Smith, 1945: 582-83. Smith and Taylor, 1948:50. + Martin, 1958:49. + + _Syrrhaphus cystignathoides_: Guenther, 1900:218. + + _Syrraphus cystignathoides_: Diaz de Leon, 1904:10. + + _Syrrhopus cystignathoides_: Barbour and Loveridge, 1946:170. + + [Illustration: FIG. 5: Distribution of _Syrrhophus cystignathoides + campi_ (solid symbols) and the nominate subspecies (open symbols).] + +_Diagnosis._--Venter areolate; interorbital light and dark bars present; +ground color yellow to brownish-yellow in life (Fig. 4b). + +_Remarks._--Firschein (1954) briefly considered the status of Peters' +(1871) _Phyllobates verruculatus_ and noted that if it was a +_Syrrhophus_ it would probably be referrable to _S. cystignathoides_. +Peters' (1871) original description corresponds well with +_S. cystignathoides_, and the type-locality ("Huanusco" = Huatusco) is +within the range of that species. Firschein (1954) expressed doubt that +_verruculatus_ was a _Syrrhophus_, because Peters placed it in another +genus. However, Peters described _verruculatus_ a decade before Cope +diagnosed the genus Syrrhophus. Most frogs now called _Syrrhophus_, plus +a number of lower Central American frogs now placed in a variety of +genera were placed in _Phyllobates_ by Boulenger, Cope, and Peters. + +The types of _Phyllobates verruculatus_ were destroyed during World War +II (Guenther Peters, _in litt._); the specimens subsequently assigned to +the taxon by Kellogg (1932) are _Syrrhophus cystignathoides_. Because +the type specimens are lost and because the name antedates the more +established name, _cystignathoides_, I favor retaining _Phyllobates +verruculatus_ Peters as a _nomen dubium_. + +Smith and Taylor (1948) reported _S. verruculatus_ from Tianguistengo, +Hidalgo, Mexico. These specimens are examples of _verrucipes_. Smith +(1947) reported a specimen of _verruculatus_ from San Lorenzo, Veracruz. +Firschein (1954) referred it to _cystignathoides_, and Duellman (1960) +concluded that both authors were in error and that the specimen (USNM +123530) was a _leprus_. + +_Etymology._--The trivial name is the diminutive of _Cystignathus_, a +once-used generic name for several leptodactylid frogs. + +_Distribution._--Low and moderate elevations in the foothills along the +Sierra Madre Oriental from eastern San Luis Potosi to Central Veracruz, +Mexico (Fig. 5). + +_Specimens examined._--(130), MEXICO, _Puebla_: Necaxa, UMMZ 69519-20. +_San Luis Potosi_: 5 km. W Aguismon, LSUMZ 4962-63; along Rio Axtla, +road to Xilitla, UMMZ 105500; Tamazunchale, UIMNH 3199; 6.5 km. N +Tamazunchale, UMMZ 104039; 8 km. N Tamazunchale, UMMZ 119490. +_Veracruz_: Coatepec, 1210 m., FMNH 704966-67; 11 km. SE Coatepec, 850 +m., FMNH 70468-70; below Cordoba, FMNH 104588, UIMNH 13321; Cuautlapam, +1000 m., FMNH 106477-80, KU 100364, UIMNH 58200-03, UMMZ 105392; Fortin +de las Flores, UIMNH 13322, 13339; 1.6 km. N Fortin de las Flores, UIMNH +42799-808, UMMZ 105389; 3.2 km. N Fortin de las Flores, UIMNH 26633-35; +4.8 km. N Fortin de las Flores, UIMNH 71967-68; 3.2 km. W Fortin de las +Flores (Barranca Metlac), 910 m., UIMNH 49294-95, UMMZ 115444-46, +118221, 119893 (2); Huatusco, KU 100363; Jalapa, 1400 m., FMNH 70440, +70443-51, 70454-65; 16 km. NE Jalapa, 1300 m., FMNH 70452-53; 8 km. E +Jalapa, UIMNH 13338; 9.5 km. S Jalapa, UMMZ 122083 (2); Mirador, KU +23967; Paraja Nuevo, El Suchil, UMMZ 85490 (7), 85491 (2), 90315; La +Passa, UIMNH 49293, 49297; 1 km. E Plan del Rio, 240 m., UMMZ 102067 +(2); Potrero Viejo, FMNH 104583, 104586, 105326-27, KU 26789, 100357-62, +UIMNH 13323, 13340-43; USNM 32402 (lectotype), 32403-04, 32406-09; 9.6 +km. S Santa Rosa, TCWC 12785; 24 km. NE Tezuitlan (Puebla), UMMZ 105388; +Teocelo, FMNH 70437-38, KU 26080, 26790; 3.2 km. N Teocelo, FMNH 70439, +70441-42; 9.6 km. NW Tihuatlan, UIMNH 3684-85; 15 km. ENE Tlacotepec, KU +23966; 26 km. NW Tuxpan, UMMZ 126419. + + +=Syrrhophus leprus= Cope + + _Syrrhophus leprus_ Cope, 1879:268-69 [Holotype.--USNM 10040, from + Santa Efigena, Oaxaca, Mexico, Francis Sumichrast collector]. + Kellogg, 1932:124-5, 128. Taylor and Smith, 1945:582. Smith and + Taylor, 1948:50-51. Duellman, 1958:8, pl. 1, Fig. 2; 1960:56-57. + Gorham, 1966:165. + + _Syrrhaphus leprus_: Guenther, 1900:217. + + _Syrrhophus leprus leprus_: Neill, 1965:85-86. + + _Syrrhophus leprus cholorum_ Neill, 1965:85-86 [Holotype.--Wilfred + T. Neill collection 1525, from 3.9 mi. N San Antonio, Toledo + District, British Honduras, collected October 28, 1959, by + R. A. Allen, T. C. Allen, and W. T. Neill]. + +_Diagnosis._--Medium-sized frogs, males 20.5-26.5 mm. in snout-vent, +females 22.0-29.3 mm. in snout-vent length; vocal slits present in +males; tips of fingers dilated slightly; first finger longer than +second; inner metatarsal tubercle twice size of small, conical outer +metatarsal tubercle; skin of dorsum pustular, that of venter smooth; +snout subacuminate; diameter of tympanum 47.5-62.5 per cent of eye in +males, 38.6-57.9 per cent in females; dorsum yellowish-green with +chocolate brown blotches or spots forming reticulations in most +specimens; venter white to gray; flanks brown, spotted with white or +not; limbs banded; interorbital bar obscured by dorsal pattern. + + [Illustration: FIG. 6: Dorsal views of _Syrrhophus leprus_ showing + variation in dorsal pattern (left, UMMZ 121244, x2; right, KU 26106, + x1.7). Side of head (UIMNH 42726, x7).] + + [Illustration: FIG. 7: Distribution of three species of eastern + complex _Syrrhophus_: _leprus_ (circles), _rubrimaculatus_ + (triangles), and _verrucipes_ (squares).] + +_Remarks._--My distribution map (Fig. 7) differs somewhat from that of +Duellman (1958), who was unaware of specimens reported by Taylor and +Smith (1945) from central Veracruz, Mexico. + +Duellman (1958, 1960) regarded _S. leprus_ as having a gray venter. +Neill (1965) characterized his new subspecies on the basis of white +venter and spots on the dorsum. Some specimens from throughout the range +have only small round spots, instead of vermiculations (Fig. 6). The +gray ventral coloration is largely restricted to the population in Los +Tuxtlas, Veracruz, but only about 80 per cent of the specimens from the +Los Tuxtlas have gray venters, whereas specimens from Guatemala, Oaxaca, +Tabasco, and central Veracruz, Mexico, have white venters (rarely gray). +Since the specimens from British Honduras are not distinct from +specimens throughout most of the range, there is no reason to recognize +them as a subspecies. + +_Etymology._--Greek, _lepra_, leprosy, in reference to the mottled color +pattern. + +_Distribution._--Discontinuous; central Veracruz to British Honduras to +low elevations in the foothills of the Sierra Madre Oriental, Los +Tuxtlas, Sierra Madre de Chiapas (Isthmus of Tehuantepec (Fig. 7)). + +_Specimens examined._--(84). GUATEMALA, _Alta Verapaz_: Chinaja, KU +55961-62. _El Peten_: 15 km. NW Chinaja, KU 55963; Piedras Negras, USNM +114085-92; Tikal, UMMZ 117035; Uaxactun, AMNH 55121-22. + +MEXICO, _Oaxaca_: Cerro San Pedro del Isthmo, UIMNH 35510; Finca La +Gloria, USNM 114093; 30.5 km. N Matias Romero, UIMNH 39459, 71969; Santa +Efigenia, USNM 10040 (holotype). _Tabasco_: Teapa, UMMZ 113799-800; 13.5 +km. W Teapa, UMMZ 120253. _Veracruz_: 27.5 km. N Acayucan, UIMNH 42726; +Atoyac, UIMNH 13331, 49296; 3.2 km. N Catemaco, UIMNH 71976-77; Coyame, +UIMNH 38995, 38998, 40342; Dos Amates, TCWC 21211; Fortin de Las Flores, +FMNH 113751, 113753; Paraja Nuevo, El Suchil, UMMZ 90315; Potrero Viejo, +FMNH 113743-50, 126114-18, KU 26104-06, UIMNH 13332-37, UMMZ 88837; San +Andres Tuxtla, UIMNH 27123-31, 28611, 71975, UMMZ 115450 (5); San +Lorenzo, USNM 123530; 4.5 km. NW Santiago Tuxtla, JDL 992 (skeleton), +UIMNH 27122; 32 km. S Sayula, EAL 1696; Tepalapan, 1.6 km. S Catemaco, +UMMZ 118222 (2); Volcan San Martin, south slope, UMMZ 118223; Volcan San +Martin, Rancho El Tular, UIMNH 35399-400, 40340-41. + + +=Syrrhophus rubrimaculatus= Taylor and Smith + + _Syrrhophus rubrimaculatus_ Taylor and Smith, 1945:583-85 + [Holotype.--USNM 114070, from La Esperanza, near Escuintla, + Chiapas, Mexico, collected May 13, 1940, by H. M. and R. Smith]. + Duellman, 1958:1-4, 7, 12, 14. Gorham, 1966:167. + + _Syrrhophus rubrimaculata_: Smith and Taylor, 1948:48-49. + + [Illustration: FIG. 8: _Syrrhophus rubrimaculatus_ (upper right, + KU 58911, x1.6; lower right, KU 58910, x4) and _S. verrucipes_ + (upper left, UIMNH 15995, x1.6; lower left, UIMNH 15989, x3.7).] + +_Diagnosis._--Small frogs, males 18.2-23.5 mm. snout-vent, females +19.0-22.5 mm. snout-vent length (small sample); vocal slits in males; +digital tips scarcely expanded (Fig. 1); first finger shorter than +second; outer palmar tubercle reduced in size; inner metatarsal tubercle +elongate, twice the size of small, conical outer metatarsal tubercle; +diameter of tympanum 35.5-46.5 per cent that of eye in both sexes; +dorsum brown with small pale spots (red in life); venter gray. + +_Remarks._--Previous authors who treated _Syrrhophus_ placed this +species in the western complex, because it occurs on the Pacific versant +and has a reduced outer palmar tubercle. Duellman (1958) placed +_rubrimaculatus_ apart from the other western species, because of its +relatively unexpanded digital tips and coloration. The digital tips are +like those in _leprus_, which _rubrimaculatus_ resembles. Except for the +reduction of the outer palmar tubercle, _rubrimaculatus_ could be a +member of the _leprus_ group. + +_Syrrhophus rubrimaculatus_ is probably best treated as a Pacific +derivative of the _leprus_ group, even though the palmar tubercles do +not agree. The removal of _rubrimaculatus_ from the western complex +results in a more homogeneous remainder and does not greatly increase +the heterogeneity of the eastern complex. + +_Etymology._--Latin, meaning spotted with red; in reference to the +colors in life. + +_Distribution._--Low to moderate elevations on the Pacific versant of +southeastern Chiapas, Mexico (Fig. 7); probably extending into +adjacent Guatemala. + +_Specimens examined._--(48) MEXICO, _Chiapas_: Escuintla, UMMZ 88283; 6 +km. NE Escuintla, UMMZ 87876-80; La Esperanza, UIMNH 13285, UMMZ +88496-97, USNM 114070 (holotype), 114054-69, 114072; Monte Cristo, UMMZ +88353; 1.3 km. N Puerto Madero, KU 58910-11; Finca San Jeronimo, 600-650 +m., UIMNH 55299-312, 55313-16 (cleared and stained). + + +=Syrrhophus guttilatus= (Cope) + + _Malachylodes guttilatus_ Cope, 1879:264 [Holotype.--USNM 9888, + from Guanajuato, Guanajuato, Mexico; collected in 1877 by + Alfredo Duges]. + + _Syrrhopus guttulatus_: Boulenger, 1888:204-06. + + _Syrrhaphus guttulatus_: Guenther, 1900:317. + + _Syrraphus guttulatus_: Diaz de Leon, 1904:11. + + _Syrrhophus guttilatus_: Nieden, 1923:399-400. Kellogg, 1932:125, + 127-28. Smith and Taylor, 1948:49, 51. Firschein, 1954:52-54. + Gorham, 1966:164. + + _Syrrhophus smithi_ Taylor, 1940b:43-45, pl. 1 [Holotype.--USNM + 108594, from 15 mi. SW Galeana, Nuevo Leon, Mexico, 1575 m.; + collected on October 13, 1939, by Hobart M. Smith]. Smith and + Taylor, 1948:49, 51. Firschein, 1954:54-55. Martin, 1958:50. + Gorham, 1966:167. + + _Syrrhophus gaigeae_ Schmidt and Smith, 1944:80 [Holotype.--FMNH + 27361, from the Basin, Chisos Mountains, Brewster Co., Texas; + collected on July 24, 1937, by Walter L. Necker]. + + _Syrrhophus petrophilus_ Firschein, 1954:50-52 [Holotype.--UIMNH + 7807, from 5 km. SW San Luis Potosi, San Luis Potosi, Mexico; + collected on July 18, 1949, by David Langebartel]. Gorham, + 1966:166. + + _Syrrhophus marnocki_: Milstead, Mecham, and McClintock, 1950:548 + (in part). + +_Diagnosis._--Medium-sized frogs, males 20.6-29.0 mm. snout-vent, +females 25.7-31.0 mm. snout-vent length; vocal slits in males; digital +tips slightly expanded (Fig. 1); first and second fingers equal; skin of +dorsum smooth to moderately pustular, that of venter smooth; snout +blunt; diameter of tympanum 55.1-75.7 per cent that of eye in males, +47.6-61.7 in females; dorsum and flanks cream to gray with light brown +to black flecking and vermiculations; thighs usually not banded; +interorbital bar present (Fig. 8). + + [Illustration: FIG. 9: _Syrrhophus guttilatus_ (upper left, UIMNH + 55519, x1.4; lower left, UIMNH 55519, x2.3) and _S. marnockii_ + (upper right, TCWC 9317, x1.4; lower right, TCWC 13510, x2.1).] + +_Remarks._--Cope (1879) distinguished _Malachylodes_ from _Syrrhophus_ +on the basis of the presence of a frontoparietal fontanelle in the +holotype of _guttilatus_. The holotype is a juvenile female and as is +the case in the juveniles of nearly all leptodactylids, a frontoparietal +fontanelle is present. Firschein (1954) used the presence of the +fontanelle to distinguish _guttilatus_ from his _petrophilus_. + +As is clearly evident from the length of the synonymy, I consider a +number of currently used names to be synonymous with _guttilatus_. I +have seen the holotypes of all four names and am unable to recognize +more than a single species. The holotype of _petrophilus_ is a male, +whereas that of _smithi_ is a female. The supposed differences are a +reflection of sexual dimorphism in the size of the eye (Table 5). The +two holotypes, as well as those of _gaigeae_ and _Malachylodes +guttilatus_ agree in color pattern. + +Schmidt and Smith (1944) named _Syrrhophus gaigeae_ from the Chisos +Mountains of the Big Bend region of Texas and compared it only with +_S. marnockii_. Milstead, Mecham and McClintock (1950) synonymized +_gaigeae_ and _marnockii_ because they were unable to verify the +characters Wright and Wright (1949) used to separate them. Specimens +from the Big Bend region differ from those of the Edward and Stockton +Plateaus in having a vermiculate pattern, an interorbital bar, and a +supratympanic stripe. In these respects they agree with specimens from +northern Mexico. Based on limited observations, the Mexican population +is yellowish to brownish in life whereas the central Texas population is +green in life. Lacking evidence of genetic exchange, the two are held to +be specifically distinct. + +Nearly every specimen examined was infested with chiggers of the genus +_Hannemania_. The greatest concentrations are on the venter, in the +groin, and on the thighs. Many specimens have chiggers on the digits and +tarsi. The same, or a related, chigger was found on many specimens of +_Syrrhophus marnockii_ and a few _S. verrucipes_, but on no other +species of the genus. Mr. Willy Wrenn told me that he has seen heavy +infestations of _Hannemania_ on _Syrrhophus pallidus_. Infestation by +_Hannemania_ probably reflects similar ecologies rather than close +relationships. + + [Illustration: FIG. 10: Distribution of _Syrrhophus guttilatus_.] + +_Etymology._--Latin, _guttula_, meaning spotting or flecking, in +reference to the color pattern. + +_Distribution._--Moderate to intermediate elevations (600 to 2000 m.) +along the Sierra Madre Oriental from the Big Bend Region of Texas to +Guanajuato, Mexico (Fig. 10). + +_Specimens examined._--(32) TEXAS, _Brewster Co._: Juniper Canyon, +Chisos Mts., FMNH 27361 (holotype of _S. gaigeae_), 27360, 27362-63, MCZ +15346, 27801, UMMZ 66080, 66082, 66085-91, USNM 76876; Upper Green +Gulch, TCWC 15943. + +MEXICO: _Coahuila_: 8 km. S Saltillo, UIMNH 55518-21. _Guanajuato_: +Guanajuato, USNM 9888 (holotype of _Malachulodes guttilatus_); 8 km. E +Guanajuato, AMNH 73425; Cerro Cubilete, AMNH 73424. _Nuevo Leon_: 3 km. +S Galeana, JDL 1215 (skeleton), UIMNH 58204; 24 km. SW Galeana. 1575 m., +USNM 108594 (holotype of _Syrrhophus smithi_). _San Luis Potosi_: 5 km. +SW San Luis Potosi, UIMNH 7807 (holotype of _S. petrophilus_). +_Tamaulipas_: 1.6 km. NW La Joya de Salas, 1530 m., UMMZ 110736 (4). + + +=Syrrhophus marnockii= Cope + + _Syrrhophus marnockii_ Cope, 1878:253 [Syntypes.--ANSP 10765-68, + from "near San Antonio," Bexar Co., Texas; collected by G. W. + Marnock]. + + _Syrrhophus marnocki_: Yarrow, 1882:24, 193. Milstead, Mecham, + and McClintock, 1950:550. + +_Diagnosis._--Medium-sized frogs, males 18.4-28.9 mm. snout-vent, +females 20.4-35.4 mm. snout-vent length; vocal slits in males; digital +tips widened (Fig. 1); first and second fingers equal; skin of dorsum +smooth to weakly pustular, that of venter smooth; snout blunt, rounded; +diameter of tympanum 47.2-68.3 per cent that of eye in males, 45.8-73.3 +in females; dorsum tan to light brown in preservative with rusty-brown +flecks, venter white; ground color green in life; thighs banded; +interorbital bar absent. + +_Remarks._--Specimens from the southern edge of the Edwards Plateau and +the eastern edge of the Stockton Plateau have larger flecks on the back +that tend to form a vermiculate pattern like that of _S. guttilatus_. +The vermiculation is never well developed (see plate 38 in Conant, +1958). Most of the specimens from the Edwards Plateau have a punctate +pattern (Fig. 9). + +Fossils are known from the Sangamon interglacial deposits in Foard and +Knox Counties, Texas (Lynch, 1964; Tihen, 1960). + +_Etymology._--A patronym for the collector of the type specimens. + +_Distribution._--The Edwards Plateau and the extreme eastern edge of the +Stockton Plateau in Texas (Fig. 11). The fossil records lie some 200 +miles to the north. Two specimens (FMNH 103216-17) from Brownsville, +Cameron Co., Texas, were formerly in the EHT-HMS collection (nos. +31348-49). Data given in Taylor's field catalogue (housed in the +Division of Reptiles, Field Museum) are "Brownsville, A. J. Kirn +collector, April 15, 1934." Until verification by recently collected +material is available, this record must be disregarded. + +_Specimens examined._--(103) TEXAS, _Bandera Co._: 10 mi. SW Medina, +TCWC 13508-10; 8 mi. W Medina, KU 60243; 13 mi. W Medina, KU 60242, TCWC +13506-07. _Bexar Co._: UIMNH 34694; Classen ranch, near San Antonio. +UMMZ 98891; Helotes, EAL 1560, MCZ 11837 (2), UMMZ 64045, USNM 13635; 2 +mi. N Helotes, TCWC 9234-35; 3.5 mi. N Helotes, LSUMZ 10363; 8 mi. N +Helotes, TCWC 1549, 4364; San Antonio, FMNH 15553-56, TCWC 13497-99. +_Blanco Co._: 8 mi. NE Blanco, TCWC 4782. _Comal Co._: New Braunfels, +TCWC 13500-05; 5 mi. NE New Braunfels, UMMZ 71016 (10). _Hays Co._: San +Marcos, AMNH 22661-64, 32700, FMNH 15245-46, 26250, 26253-57, 37617, +37665, MCZ 15649-50, 23268-69; 6 mi. SW San Marcos, TCWC 5070-71, 7140, +9232-33, 9236, 9316-17, 9320. _Kendall Co._: 11 mi. E Boerne, AMNH +54660-61, 54662 (2); 10 mi. W Boerne, KU 18441; Kendalia, UIMNH 21434. +_Kerr Co._: Kerr W. M. Area, TCWC 15859; 40 mi. NW Kerrville, TCWC 6555. +_Medina Co._: UIMNH 13287-88; 12 mi. N Castroville, UIMNH 21423; 14 mi. +N Castroville, UIMNH 21424-25; 16 mi. N Castroville, UIMNH 21421-22; 17 +mi. N Castroville, UIMNH 21428-29; 18 mi. N Castroville, UIMNH 21426-27, +21430-33; 6.5 mi. NW Rio Medina, KU 18440. _Real Co._: Rio Frio, FMNH +55156-57. _Travis Co._: Austin, AMNH 44221-22; Mount Bonnell, 5 mi. S +Austin, UMMZ 101453 (10). _Uvalde Co._: 13 mi. from Uvalde, UIMNH 62322. +_Val-Verde Co._: 40 mi. N Del Rio, JDL 214 (skeleton). + + [Illustration: FIG. 11: Distribution of _Syrrhophus marnockii_ + (circles). Starred localities are late Pleistocene records.] + + +=Syrrhophus verrucipes= Cope + + _Syrrhophus verrucipes_ Cope, 1885:383 [Holotype.--ANSP 11325, from + near Zacualtipan, Hidalgo, Mexico (1800 feet lower in a rocky gorge + of a stream near its junction with the Rio San Miguel), collected + by Dr. Santiago Bernard]. Kellogg, 1932:126-29. Smith and Taylor, + 1948:52-53. Firschein, 1954:55-57. Gorham, 1966:167. + + _Syrrhaphus verrucipes_: Guenther, 1900:216-17. + + _Tomodactylus macrotympanum_ Taylor, 1940e:496-99, pl. 55, + figs. 2a-b. [Holotype.--FMNH 100049 (formerly EHT-HMS 6838), + from La Placita, 8 km. S Jacala, Hidalgo, Mexico, 1850 m.; + collected on July 2, 1936, by Edward H. Taylor]. Smith and + Taylor, 1948:47-48. + + _Syrrhophus macrotympanum_: Dixon, 1957:384. Gorham, 1966:165. + +_Diagnosis._--Medium-sized frogs, males 17.5-26.1 mm. snout-vent, +females 28.0-31.7 mm. snout-vent length; vocal slits in males; digital +tips slightly expanded; first finger shorter than second; skin of dorsum +pustular, that of venter areolate; snout elongate, subacuminate; +diameter of tympanum 56.1-76.7 per cent that of eye in males, 54.3-56.8 +in females; in preservative, dorsum reddish brown with numerous small +black or dark brown spots (Fig. 8); venter white to cream; in life +dorsum green with darker green spots, belly white; iris gold above, +bronze below. + +_Remarks._--Cope's (1885) original description was not sufficiently +clear to enable subsequent authors to recognize this species. Taylor +(1940e) described it as a _Tomodactylus_, but Dixon (1957) pointed out +that _T. macrotympanum_ differed from the other species of the genus in +having a poorly developed lumbo-inguinal (inguinal) gland, and placed +the species in the genus _Syrrhophus_. Comparison of the holotypes of +_S. verrucipes_ and _T. macrotympanum_ leaves no doubt in my mind that a +single species is involved. This same species was reported by Smith and +Taylor (1948) as _S. verruculatus_. + +_Syrrhophus verrucipes_ bears resemblance to members of both the +_leprus_ and _marnockii_ groups. In snout shape it is closer to the +_leprus_ group, whereas in digital pad, the shape of the general body +form, and contiguity of habitat it is most similar to the _marnockii_ +group (_S. guttilatus_). + +_Etymology._--Latin, meaning warty foot, probably in reference to the +numerous plantar supernumerary tubercles. + +_Distribution._--Moderate elevations in southeastern San Luis Potosi, +Queretaro, and northwestern Hidalgo, Mexico (Fig. 7). + +_Specimens examined_--(43) MEXICO, _Hidalgo_: Jacala, UMMZ 106434; 9.6 +km. NE Jacala, Puerto de la Zorra, 1820 m., KU 60240-41, TCWC 11090, +11147; 8 km. S Jacala, La Placita, 1850 m., FMNH 100049 (holotype of +_Tomodactylus macrotympanum_), 100791-803, 105334-35, 114287, UIMNH +15989-92, 15995-96, UMMZ 117252, USNM 137202; Tianguistengo, FMNH +113705-09, UIMNH 13328-30; near Zacualtipan, ANSP 11325 (holotype of +_Syrrhophus verrucipes_). _Queretaro_: 3.5 km. S San Juan del Rio, EAL +1343. _San Luis Potosi_: 9.6 km. W Ahuacatlan, LSUMZ 4968-70. + + +=Syrrhophus dennisi= new species + + _Syrrhophus latodactylus_: Martin, 1958:49 (in part). + +_Holotype._--UMMZ 101121, adult male from a cave near El Pachon, 8 km. N +Antiguo Morelos, Tamaulipas, Mexico, 250 m., collected on March 13, +1949, by Paul S. Martin. + +_Paratopotypes._--(26). UMMZ 101122 (10), 101123 (2), 101126, 126993 +(12). + +_Diagnosis._--Medium-sized frogs, males 22.8-28.4 mm. snout-vent, +females 25.9-32.0 mm. snout-vent; vocal slits in males; digital tips +greatly expanded, more than twice width of digit; first finger shorter +than second; skin of dorsum shagreened to pustular, that of venter +weakly to moderately areolate; toes webbed basally; dorsum light brown +to tan with brown vermiculations; venter white; diameter of tympanum +53.9 to 64.2 per cent that of eye in males, 50.6 to 58.7 per cent in +females. + +_Description and variation._--(Fig. 12). Head wider than body; head as +wide or wider than long in males, sometimes longer than wide in females; +snout acuminate in dorsal view, elongate and rounded in lateral profile; +canthus rostralis rounded but distinct; loreal region slightly concave, +sloping abruptly to lip; lips not flared; eyelid about two-thirds +interorbital distance; length of eye less than distance between eye and +nostril; diameter of tympanum 53.9 to 64.2 per cent that of eye in +males, 50.6 to 58.7 per cent in females; tympanum round and distinct in +both sexes; supratympanic fold moderately distinct; choanae within +border of jaws, completely visible from directly below, rounded to +slightly oval; dentigerous processes of prevomers and teeth absent; +tongue free for posterior one-half, generally oval in outline; vocal +slits present in males. + +Many scattered pustules on dorsum; flanks areolate; skin of venter +areolate or not (variability may be due to differences in preservation); +ventral disc distinct on chest and lower abdomen; inguinal gland present +or not, when present varying from very large and distinct to poorly +defined; axillary gland absent. + +First finger shorter than second; all fingers bearing truncate tips with +pads, each pad having a terminal groove; fingers fringed; fingers three +and four having dilated pads two to three times width of digit; +subarticular tubercles large, conical, rounded, simple; supernumerary +tubercles numerous on thenar surface, none on digits; three palmar +tubercles, outer slightly smaller than largest supernumerary tubercles; +row of tubercles on outer edge of forearm variable, weak to very +distinct; tips of toes wider than digits, rounded to truncate at tips, +each pad having terminal groove; toes having lateral fringes, bases of +toes united by web, web not extending to basal subarticular tubercle; +subarticular tubercles smaller than those of hand, round, conical, +simple; supernumerary tubercles numerous on plantar surfaces, extending +between metatarsal tubercles, present on toes between basal two +subarticular tubercles in some specimens; outer metatarsal tubercle +round, conical, one-half as large as ovoid, non-compressed inner +metatarsal tubercle; tarsal tubercles or folds absent. + +Ground color pale reddish-brown to tan dorsally, creamy on flanks; +dorsal pattern consisting of reddish-brown to brown vermiculations +extending onto flanks; distinct interorbital light bar present; loreal +region darker than snout, reddish-brown compared to tan or pale +reddish-brown; arms colored like dorsum; thighs banded, unicolor brown +on posterior surfaces; shanks and tarsi banded; venter white to cream +punctated with brown in some specimens. + +The variation in proportions is summarized in Table 5. + +_Remarks._--Martin (1958) expressed some doubt that this series of 26 +specimens was identical with "_S. latodactylus_." My study indicates +that the specimens from El Pachon represent a distinctive but allied +species. Males of the two species can be readily separated by the +relative sizes of the tympani, presence or absence of vocal slits, and +color pattern. Females of the two species can be separated by color +pattern. Within the type-series, the pattern varies from weakly to +strongly vermiculate but is always recognizable as vermiculate rather +than spotted as in _S. longipes_ (= _S. latodactylus_ of Taylor and +Martin). + + [Illustration: FIG. 12: _Syrrhophus dennisi_ sp. nov., holotype, + UMMZ 101121 (dorsum x1.8, side of head x6.1).] + +_Etymology._--The specific name is a patronym for David M. Dennis, whose +drawings greatly enhance the worth of this paper. + +_Distribution._--Known only from the type series. + + +=Syrrhophus longipes= (Baird), New combination + + _Batrachyla longipes_ Baird, 1859:35, pl. 37, fig. 1-3 + [Holotype.--apparently USNM 3237 (cited as 3207 by Cope, 1887:16), + now lost, from 40 Leagues from (probably north) Mexico City; + collected by John Potts]. Kellogg, 1932:107. + + _Epirhexis longipes_: Cope, 1866:96. + + _Eleutherodactylus longipes_: Kellogg, 1932:107 (part). Smith and + Taylor, 1948:61. Lynch, 1963:580-581. Gorham, 1966:82. + + _Syrrhophus latodactylus_ Taylor, 1940d:396-401, pl. 43, figs. A-F, + text fig. 7 [Holotype.--FMNH 100063 (formerly EHT-HMS 6807), from + Huasteca Canyon, 15 km. W Monterrey, Nuevo Leon, Mexico, 680 m.; + collected on June 20, 1936, by Edward H. Taylor]. Smith and + Taylor, 1948:50-52. Martin, 1958:48-50. Gorham, 1966:165. + +_Diagnosis._--Large frogs, males 22.1-33.2 mm. snout-vent, females +26.8-39.6 mm. snout-vent length; vocal slits lacking in males; digital +tips greatly expanded (more than twice the width of digit); first finger +shorter than second; skin of dorsum pustular, that of venter smooth; +diameter of tympanum in males 61.1-87.2 per cent that of eye, 49.5-72.1 +per cent in females; dorsum tan with large or small spots and blotches; +limbs banded; interorbital bar or triangle present. + +_Remarks._--I have applied Baird's _Batrachyla longipes_ to the frog +Taylor (1940d) called _Syrrhophus latodactylus_ because the color +pattern (Fig. 13) predominant in the southern part of the range agrees +with that described (figured) for _Batrachyla longipes_. + +The color pattern of individuals in the southern part of the range of +this species consists of large spots or blotches, whereas in the +northwestern part the pattern is made up of smaller spots. In the +northeastern part of the range, the pattern is more reduced and tends to +consist of heavy flecking. The interorbital bar is narrower in specimens +from Nuevo Leon and Tamaulipas and is triangular in specimens from +Hidalgo and Queretaro. + +The status of the name _Batrachyla longipes_ is currently that of a +_nomen dubium_ (Lynch, 1963). At that time, I was unaware of the +geographic variation in color pattern in _Syrrhophus latodactylus_. + +The exact type-locality of _Batrachyla longipes_ is not known. If it is +40 Leagues north of Mexico City, the locality would be in an area where +the species has a blotched instead of a flecked or spotted pattern. No +justifiable evidence was presented to place _Batrachyla longipes_ in +_Eleutherodactylus_ instead of _Syrrhophus_. Barbour (1923) and Kellogg +(1932) associated another species (_E. batrachylus_) with _longipes_. +Taylor (1940a) noted this as a case of misidentification and corrected +the error but left _longipes_ in the genus _Eleutherodactylus_. Lynch +(1963) noted several points of morphological agreement between +_Syrrhophus_ and _B. longipes_ but did not place _longipes_ in +_Syrrhophus_. + +Baird's (1859) figures of the holotype do not illustrate prevomerine +teeth, but according to Cope (1866) they were present in the holotype. +The digital tips of the frog in the figure are somewhat narrower than +those typically seen in _S. latodactylus_. If the specimen was slightly +desiccated, as possibly was the case, the digits would appear narrower. +There is no evidence contrary to placing _Syrrhophus latodactylus_ in +the synonymy of _Batrachyla longipes_. + + [Illustration: FIG. 13: Dorsal views of _Syrrhophus longipes_ + illustrating geographic variation in pattern (left, TCWC 12179, + x1.5; right, KU 92572, x1.8); side of head (TCWC 10966, x6).] + +Application of Baird's name _Batrachyla longipes_ to the species of frog +heretofore called _Syrrhophus latodactylus_ poses one serious problem. +_Batrachyla longipes_ is the type-species (by original designation) of +the genus _Epirhexis_ Cope, 1866, which has priority over _Syrrhophus_ +Cope, 1878. If _Batrachyla longipes_ is left in the status of a _nomen +dubium_, _Epirhexis_ can be forgotten, for the two names are tied +together. However, since it seems almost certain that _Batrachyla +longipes_ and _Syrrhophus latodactylus_ are conspecific, the former name +should not be left as a _nomen dubium_. _Epirhexis_ never came into +general usage (Cope cited the name four times, but no one else has used +it), whereas _Syrrhophus_ is well established in the zoological +literature. It would serve only to confuse the literature to adhere +strictly to the Law of Priority and replace _Syrrhophus_ with +_Epirhexis_. Therefore, _Syrrhophus_ is used in this paper, even though +_Epirhexis_ has priority. A request for the suppression of _Epirhexis_ +Cope, 1866, has been submitted to the International Commission of +Zoological Nomenclature (Lynch, 1967). + +_Etymology._--Latin, meaning long-footed; Taylor's _latodactylus_ refers +to the wide digital pads. + + [Illustration: FIG. 14: Distribution of _Syrrhophus dennisi_ + (triangle) and _S. longipes_ (circles).] + +_Distribution._--Moderate elevations (650 to 2000 meters) along the +Sierra Madre Oriental from central Nuevo Leon to northern Hidalgo, +Mexico (Fig. 14). + +_Specimens examined._--(122) MEXICO, _Hidalgo_: 3 km. NE Jacala, AMNH +52977; 9.6 km. NE Jacala, 1800 m., TCWC 10966-70, 12179; 8 km. S Jacala, +La Placita, 1850 m., FMNH 100266-68, 103244, UIMNH 13291, 13327. _Nuevo +Leon_: Salto Cola de Caballo, KU 92572; Huasteca Canyon, 15 km. W +Monterrey, 680 m., FMNH 100063 (holotype of _S. latodactylus_), UIMNH +13290; 6.5 km. N Pablillo, EAL 1319; Sabinas Hidalgo, USNM 139728. +_Queretaro_: Cueva de los Riscos, 8 km. SW Jalpan, KU 106300. _San Luis +Potosi_: 13 km. E Santa Barberita, LSUMZ 2295; second camp, San Luis +Potosi road, UIMNH 13326; Xilitla, Cueva sin nombre, UMMZ 125892. +_Tamaulipas_: 4 km. W El Carrizo, 500 m., UMMZ 111343 (31); 8 km. N +Chamal, Bee Cave, KU 106299; 14.5 km. NNW Chamal, 420 m., UMMZ +111339-40, 111342 (4), 111344 (11); 19 km. NNW Chamal, 700 m., UMMZ +111341 (3); El Chihue, 1880 m., UMMZ 111289 (4); 11 km. N Gomez Farias, +1060 m., UMMZ 101166; 11 km. WNW Gomez Farias, 1800 m., UMMZ 108507 (3); +8 km. NW Gomez Farias, 1060-1400 m., LSUMZ 11085, UMMZ 101167 (3), +101168 (4), 101169 (2), 101170 (3), 101171 (2), 101360-61, 102860, +102933 (4), 102934 (2), 102935-38, 102939 (2), 102940-43, 108800 (3), +110735, 111345-46. + + +=Syrrhophus pipilans= Taylor + + _Syrrhophus pipilans_ Taylor, 1940c:95-97, pl. 1 [Holotype.--FMNH + 100072 (formerly EHT-HMS 6843), 14.6 km. S Mazatlan, Guerrero, + Mexico; collected on July 22, 1936, by Edward H. Taylor]. + +_Diagnosis._--Medium sized frogs, males 22.6-28.5 mm. snout-vent, +females 21.1-29.4 mm. snout-vent length; vocal slits present in males; +finger tips slightly expanded, truncate in outline; inner metatarsal +tubercle less than twice the size of outer; skin of dorsum smooth to +shagreened, that of venter smooth; tympanum 36.5-54.0 per cent diameter +of eye; dorsum dark brown with large or small light brown, orange-brown, +or yellowish spots or blotches; limbs banded; interorbital bar absent. + + [Illustration: FIG. 15: Dicegrams of ear size relative to eye + diameter in the two subspecies of _Syrrhophus pipilans_. N = 17 + in _nebulosus_, 18 in _pipilans_.] + +_Remarks._--Two subspecies were recognized by Duellman (1958). +Previously both had been treated as species. The two populations were +distinguished on the basis of color pattern and the size of the +tympanum. Measurements of 17 males of _S. p. nebulosus_ from central +Chiapas and 18 males of _S. p. pipilans_ from southcentral Oaxaca and +Guerrero, Mexico, demonstrates that the supposed difference in tympanum +size is not significant (Fig. 15). There is, however, a tendency for +the western population of _S. pipilans_ to have larger tympani. Based on +the present examination of 112 specimens of this species the two +populations are held to be sufficiently distinct to warrant taxonomic +recognition as subspecies (Fig. 16). + + [Illustration: FIG. 16: _Syrrhophus pipilans nebulosus_ (left, + KU 58908) and _S. p. pipilans_ (right, KU 86885). x2.7.] + +The parotoid glands attributed to this species by Taylor (1940c:95) are +merely the superficial expression of the _m. depressor mandibulae_ and +scapula. No true glands are present in the parotoid region. + + +=Syrrhophus pipilans nebulosus= Taylor + + _Syrrhophus nebulosus_ Taylor, 1943:353-55, pl. 27, figs. 3-5 + [Holotype.--FMNH 100095 (formerly EHT-HMS 3774), near Tonola, + Chiapas, Mexico; collected on August 27, 1935, by Hobart M. + Smith and Edward H. Taylor]. Smith and Taylor, 1948:49, 51. + + _Syrrhophus pipilans nebulosus_: Duellman, 1958:2-4, 9, 12, 14. + Stuart, 1963:32-33. Gorham, 1966:166-67. + +_Diagnosis._--Diameter of tympanum 36.6-47.8 per cent that of eye; +dorsum dark brown with numerous small light brown to yellowish spots. + +_Remarks._--The distribution of this subspecies is adequately described +by Duellman (1958). Fouquette (1960) described the vocalization of this +frog. + +_Etymology._--Latin, _nebula_, in reference to the clouded dorsal +pattern. + +_Distribution._--Low to moderate elevations along the Pacific versant of +Chiapas and in the Grijalva valley of Chiapas and Guatemala (Fig. 17). + +_Specimens examined._--(54) GUATEMALA, _Huehuetenango_: Jacaltenango, +UMMZ 117036; 35 km. SE La Mesilla, TNHC 29652. MEXICO, _Chiapas_: 11.2 +km. N Arriaga, 300 m., UMMZ 125891; 11.8 km. N Arriaga, UMMZ 117279; +12.8 km. N Arriaga, UMMZ 117280; 17.5 km. S Arriaga, UIMNH 57108-109; +1.5 km. S Bochil, 1250 m., KU 58898-908; Cerro Hueco, 7 km. S Tuxtla +Gutierrez, UMMZ 123007; 3.2 km. S Ixtapa, UMMZ 124000; Linda Vista, ca. +2 km. NW Pueblo Nuevo Solistahuacan, KU 58897; Hda. Monserrate, 40 km. +NW Arriaga, UMMZ 102258; near San Ricardo, FMNH 100720; Tapachula, FMNH +75792, 103242, 100695-96, UIMNH 13292; 56 km. E Tapanatepec, Oaxaca, +TNHC 26942, Tonola, FMNH 100095 (holotype), 100686-92, UIMNH 13293-95; +Tuxtla Gutierrez, FMNH 100693-94, UIMNH 13297; 19 km. N Tuxtla +Gutierrez, TNHC 25229-30; 15.5 km. NE Tuxtla Gutierrez, UMMZ 119892 (3); +19 km. NE Tuxtla Gutierrez, UMMZ 119891 (3); 8 km. NNW Tuxtla Gutierrez, +KU 37809; Union de Juarez, FMNH 105294. + + +=Syrrhophus pipilans pipilans= Taylor + + _?Syrrhopus verruculatus_: Gadow, 1905:194. + + _Syrrhophus pipilans_ Taylor, 1940c:95-97, pl. 1 [Holotype.--FMNH + 100072 (formerly EHT-HMS 6843), from 14.6 km. S Mazatlan, + Guerrero, Mexico; collected on July 22, 1936, by Edward H. Taylor]. + Taylor and Smith, 1945:581-82. Smith and Taylor, 1948:49, 50-51. + + _Syrrhophus pipilans pipilans_: Duellman, 1958:1-4, 8-9, 13-14, + pl. 2, fig. 1. Gorham, 1966:166. + +_Diagnosis._--Diameter of tympanum 40.6-54.0 per cent that of eye; +dorsum dark brown with large light spots or blotches. + +_Remarks._--Duellman's (1958) synopsis of this subspecies is adequate; +the distribution has not been extended, but several records are now +available which fill in gaps. + + [Illustration: FIG. 17: Distribution of _Syrrhophus pipilans_: + _nebulosus_ (open circles) and _pipilans_ (solid circles).] + +Gadow's (1905) record of _S. verruculatus_ from "Buena Vista, S. +Guerrero" is most likely applicable to this species. Gadow simply +included the name in a list of the species he had collected during his +trip in Mexico (1902-04); no further comment was made on this species +although references to _Syrrhopus_ (sic) appear in several places in the +paper and would appear to apply to the species he had. + +_Etymology._--Latin, _pipilo_, chirping, peeping, in reference to the +call of the male. + +_Distribution._--Sea level to about 1800 meters along the Pacific +versant of western Mexico from central Guerrero to the Isthmus of +Tehuantepec (Fig. 17). + +_Specimens examined._--(62). MEXICO, _Guerrero_: Acapulco, UMMZ 110125; +6.4 km. N Acapulco, FMNH 100389, 100525; Agua del Obispo, 980-1000 m., +FMNH 75791, 100518-21, 100526, KU 86884-86, UIMNH 13315, UMMZ 119152, +125890 (4); 13.3 km. NW Coyuca, UIMNH 38367, 71982-83; 14.5 km. S +Mazatlan, FMNH 100072 (holotype), 100408, 100511-17, UIMNH 13302-309; +Tierra Colorado, 300 m., KU 67961, UIMNH 13313-14; near El Treinte, FMNH +126639; Xaltinanguis, FMNH 100522-24, 126640. _Oaxaca_: Cacahuatepec, +UIMNH 52853; 8 km. NW Rio Canoa, 53 km. ESE Cuajinicuilapa, UIMNH +52852; 6.4 km. N El Candelaria, UIMNH 9501; 11.2 km. S El Candelaria, +UIMNH 9502; 17 km. NE Juchatengo, 1600 m., KU 86887; 31.5 km. N +Pochutla, UMMZ 123999 (2); 32.9 km. N Pochutla, 850 m., UMMZ 123996; +37.1 km. N Pochutla, UMMZ 123998 (2); 41.4 km. N Pochutla, UMMZ 123997 +(2); Cerro Quiengola, FMNH 105653; 3.8 km. N Santiago Chivela, UMMZ +115449; 14.5 km. W Tehuantepec, UMMZ 115448 (2). + + +=Syrrhophus interorbitalis= Langebartel and Shannon + + _Syrrhophus interorbitalis_ Langebartel and Shannon, 1956: 161-65, + figs. 1-2 [Holotype.--UIMNH 67061 (formerly FAS 9378), 36 mi. N + Mazatlan, Sinaloa, Mexico, collected on November 17, 1955, by + E. C. Bay, J. C. Schaffner, and D. A. Langebartel]. Duellman, + 1958:1-4, 10, 12, 14. Gorham, 1966:164-65. + + _Syrrhophis interorbitalis_: Campbell and Simmons, 1962:194, + fig. 1. + + [Illustration: FIG. 18: Left to right. _Syrrhophus interorbitalis_ + (UIMNH 38095, x1.5), _S. nivocolimae_ (LACM 3203, x1.3), and + _S. teretistes_ (KU 75263, x1.5).] + +_Diagnosis._--Medium sized frogs, only known male 25.6 mm. snout-vent, +females 20.0-26.7 mm. snout-vent length (small sample); vocal slits in +males; finger tips expanded; first finger shorter than second; outer +metatarsal tubercle one-third size of inner; skin of dorsum shagreened, +that of venter smooth; diameter of tympanum 37.7-42.4 per cent that of +eye in both sexes; pale yellow-brown ground color mottled with brown; +limb bands broad, much wider than narrow light interspaces; interorbital +bar very long, edged with dark brown to black (Fig. 18). + +_Remarks._--Duellman's (1958) measurements and proportions of +_S. interorbitalis_ were based exclusively on the type series, which is +composed of only females; therefore his _interorbitalis_ data are not +comparable with the data for the other species in his table. Campbell +and Simmons (1962) collected the only known male. The type series was +collected beneath rocks in a stream bed; the collectors heard calling +frogs in the bushes but were unable to obtain specimens (Langebartel and +Shannon, 1956). Campbell and Simmons (1962) reported that their specimen +had a poorly developed interorbital bar in life; in preservative the bar +compares favorably with the bar in the female (Fig. 18). + +_Etymology._--Latin, in reference to the pale interocular band. + +_Distribution._--Pacific lowlands of Sinaloa, Mexico (Fig. 20). + +_Specimens examined._--(10). MEXICO, _Sinaloa_: 36 mi. N Mazatlan, UIMNH +38094-96, 67061 (holotype), 71970-74; 65 mi. N Mazatlan, LACM 13773. + + +=Syrrhophus modestus= Taylor + + _Syrrhophus modestus_ Taylor, 1942:304-06, pl. 29 [Holotype.--FMNH + 100048 (formerly EHT-HMS 3756), from Hacienda Paso del Rio, + Colima, Mexico; collected on July 8, 1935, by Hobart M. Smith]. + Smith and Taylor, 1948:49-50. + + _Syrrhophus modestus modestus_: Duellman, 1958:2-5, 7, 14, pl. 1, + fig. 1. Gorham, 1966:166. + +_Diagnosis._--Small frogs, males 15.8-20.1 mm. snout-vent length, single +female 18.5 mm.; vocal slits present in males; finger tips widely +expanded; first finger shorter than second; inner metatarsal tubercle +about three times size of outer; skin of dorsum shagreened, that of +venter smooth; tympanum concealed; pale cream in preservative with dark +brown spots; limbs banded; bands on forearm and thigh poorly developed +or absent; interorbital bar absent. + +_Remarks._--The tympanum is concealed in _S. modestus_, +_S. nivocolimae_, _S. pallidus_, _S. teretistes_, and to a lesser degree +in _S. interorbitalis_. However, if the specimen is permitted to dry +slightly, the annulus tympanicus becomes visible through the skin and a +tympanum/eye ratio can be computed. + +One of the few cases of sympatry within the genus _Syrrhophus_ involves +this species; _modestus_ and _nivocolimae_ are known to be sympatric at +one locality in southwestern Jalisco, Mexico. + +Duellman (1958) used the trinomial for this population and named a new +subspecies, _pallidus_, from Nayarit. I consider _pallidus_ to be +specifically distinct from _modestus_ because there is no evidence of +genetic exchange, and there is no overlap in the distinguishing +morphological features. I do consider the two populations to be closely +related but feel the interrelationships between _modestus_, _pallidus_, +_nivocolimae_, and _teretistes_ are more complex than would be indicated +by the use of trinomials. The sympatric occurrence of _modestus_ and +_nivocolimae_ is significant; morphologically, they might otherwise be +regarded as subspecies. Although allopatric, similar arguments could be +advanced for the morphologically similar _pallidus_ and _teretistes_. +The four are here afforded species rank since morphological similarity +and allopatry are not sufficient grounds for the assumption of genetic +exchange. + + [Illustration: FIG. 19: _Syrrhophus modestus_ [left, UMMZ 115447 + (WED 11155)] and _S. pallidus_ (right, UMMZ 115453). x2.2.] + +_Etymology._--Latin, meaning unassuming, modest, in reference to the +small size of the species. + +_Distribution._--Low elevations (up to 700 meters) in the lowlands and +foothills of Colima and southwestern Jalisco, Mexico (Fig. 20). + +_Specimens examined._--(14). MEXICO, _Colima_: Hda. Paso del Rio, FMNH +100048 (holotype), 100167, 100299, UIMNH 13300, UMMZ 110877 (2), USNM +139729; 7.2 km. SW Tecolapa, UMMZ 115477 (4); _Jalisco_: 17.6 km. SW +Autlan, 606 m., KU 102627; 3.2 km. N La Resolana, UMMZ 102100; Bahia +Tenacatita, UMMZ 84264. + + +=Syrrhophus nivocolimae= Dixon and Webb + + _Syrrhophus nivocolimae_ Dixon and Webb, 1966:1-4, Fig. 1 + [Holotype.--LACM 3200, from Nevado de Colima (6 airline miles west + of Atenquique), Jalisco, Mexico, 7800 feet; collected on July 20, + 1964, by Robert G. Webb]. + +_Diagnosis._--Small frogs, males 18.5-21.1 mm. snout-vent length, only +known female 24.1 mm. snout-vent; vocal slits present in males; finger +tips widely expanded; first finger shorter than second; inner metatarsal +tubercle about three times size of outer; skin of dorsum warty, that of +venter smooth; tympanum concealed, its diameter 30.0-39.3 per cent that +of eye in males; mid-dorsal brown band from interorbital bar to anus; +bands on limbs narrow, dark bands less than one-half width of light +bands, upper arm not banded; narrow interorbital light bar. + +_Remarks._--This species is closely related to _S. modestus_ and differs +in color pattern and degree of wartiness of the skin. Dixon and Webb +(1966) held that _nivocolimae_ had no close relatives, but the condition +of the tympanum, size, nature of the outer palmar tubercle, relative +sizes of the metatarsal tubercles, and shape and size of the digital +pads all point to a close relationship between _S. modestus_, +_S. nivocolimae_, and _S. pallidus_. + + [Illustration: FIG. 20: Distribution of the species of the + _modestus_ group: _interorbitalis_ (open circles), _teretistes_ + (solid circles), _modestus_ (open triangles), _pallidus_ (solid + triangles) and _nivocolimae_ (square). Arrow indicates locality of + sympatry between _modestus_ and _nivocolimae_. Solid line about the + localities for _interorbitalis_ is a range estimate based on call + records and specimens examined.] + +Dixon and Webb (1966) reported that _S. nivocolimae_ has a large +tympanum (50.0-59.0 per cent diameter of eye). However, my examination +of the type series and several other specimens from Jalisco reveals that +the largest tympanum/eye ratio is 39.3 per cent. Therefore, the +tympanum/eye ratio in _S. nivocolimae_ is in agreement with those for +_S. modestus_, _S. pallidus_, and _S. teretistes_ (Table 6). + +_Etymology._--_niv_, Latin, and Colima (Nevado de), meaning high on the +volcano, in reference to the higher distribution of this species (around +2000 meters) than other members of the group. + +_Distribution._--Known from southwestern Jalisco, Mexico, at moderate to +high elevations (600-2400 meters). + +_Specimens examined._--(48) MEXICO, _Jalisco_: 17.6 km. SW Autlan, 606 +m., KU 102626, 102631; 6.4 km. W Atenquique, 2060 m., KU 102628-30, +102632; 8 km. W Atenquique, 1970 m., LACM 3210-12; 9.6 km. W Atenquique, +2360 m., LACM 3200 (holotype), 3201-09; 14.5 km. W Atenquique, 2000 m., +LACM 25424-36, 25439-41, 25446; 15 km. W Atenquique, LACM 37044-46, +37244-47; 16 km. W Atenquique, 2105 m., LACM 25443-45; 17 km. W +Atenquique, 2180 m., LACM 25442. + + +=Syrrhophus pallidus= Duellman, New combination + + _Syrrhophus modestus_: Davis and Dixon, 1957:146. + + _Syrrhophus modestus pallidus_ Duellman, 1958:2-3, 5-7, 14, pl. 3 + [Holotype.--UMMZ 115452, from San Blas, Nayarit, Mexico, sea + level; collected on August 13, 1956, by William E. and Ann S. + Duellman]. Zweifel, 1960:86-88, 91, 93-94, 118, 120-22. Gorham, + 1966:166. + + _Syrrhophis modestus pallidus_: Campbell and Simmons, 1962:194. + +_Diagnosis._--Small frogs, males 17.9-19.3 mm. snout-vent length; vocal +slits in males; finger tips widely expanded; first finger shorter than +second; inner metatarsal tubercle about three times size of outer; skin +of dorsum shagreened, that of venter smooth; tympanum concealed, its +diameter 27.0-35.6 per cent of eye in males; ground color cream +vermiculated with brown, upper arm and thigh lacking, or with few, +indistinct, bands; interorbital bar absent. + +_Remarks._--Considerable debate has been waged relative to the value of +subspecies and to the reasons for recognizing distinct disjunct +populations as species versus subspecies. Lacking evidence of genetic +exchange, I prefer to retain disjunct populations that are distinctive +as species. + +All known specimens of _pallidus_ can be separated from those of +_modestus_ by color pattern. The two nominal species exhibit overlap in +proportions but the same can be said about nearly every species of +_Syrrhophus_; therefore, overlap in proportions can be disregarded in +assessing specific versus subspecific rank. Until contrary evidence is +forthcoming, I consider the disjunct populations heretofore held to be +subspecies of _modestus_ to be specifically distinct. The specimens of +the disjunct population of _pallidus_ on the Tres Marias do not differ +from the mainland population in Nayarit. This evidence, though perhaps +secondary, supports my contention that two species should be recognized. + +_Etymology._--Latin, in reference to the pale ground color in comparison +with that of _S. modestus_. + +_Distribution._--Low elevations in coastal Nayarit and on Islas Tres +Marias (Fig. 20). + +_Specimens examined._--(12) MEXICO, _Nayarit_: 18.8 mi. NW Ahuacatlan, +UIMNH 7808; San Blas, UMMZ 115452 (holotype), 115453-57; 17 km. NE San +Blas, 150 m., MSU 5085; 12.8 km. E San Blas, UIMNH 71979; 31 km. E San +Blas, UIMNH 71978; 13.5 km. N Tepic, UIMNH 71980-81. + + +=Syrrhophus teretistes= Duellman + + _Syrrhophus teretistes_ Duellman, 1958:2-3, 10-14, pl. 2, fig. 2 + [Holotype.--UMMZ 115451, from 4.8 km. NW Tepic, Nayarit, Mexico, + 840 m.; collected on August 12, 1956, by William E. Duellman]. + Gorham, 1966:167. + +_Diagnosis._--Medium-sized frogs, males 19.2-23.2 mm. snout-vent length, +single known female 24.8 mm. snout-vent; vocal slits in males; finger +tips widely expanded; first finger shorter than second; inner metatarsal +tubercle about three times size of outer; skin of dorsum shagreened, +that of venter smooth; tympanum partially concealed, its diameter +28.6-43.8 per cent of eye in males; ground color brown vermiculated with +dark brown to nearly black; upper arm and thigh banded; interorbital +light bar absent. + +_Remarks._--_S. teretistes_ appears to be most closely related to +_S. pallidus_; I consider it to be an upland derivative of _pallidus_. +Morphologically, the differences between the two are few, but lacking +evidence of genetic exchange they are retained as species. + +_Etymology._--Greek, in reference to the whistle-like nature of the +call. + +_Distribution._--Moderate elevations (840-1200 meters) in the Sierra +Occidental of Nayarit, Sinaloa, and Durango, Mexico (Fig. 20). + +_Specimens examined._--(13) MEXICO, _Nayarit_: 4.8 km. NW Tepic, 840 m., +UMMZ 115451 (holotype). _Sinaloa_: Santa Lucia, 1090 m., KU 75263-72; 1 +km. NE Santa Lucia, 1156 m., KU 78257; 2.2 km. NE Santa Lucia, 1156 m., +KU 78258. + + + + +DISCUSSION + + +There are relatively few clear-cut morphological differences among the +fourteen species now assigned to _Syrrhophus_. The majority of the +species are allopatric and differ primarily in color patterns. Sympatric +occurrence serves as an indicator of specific distinctness and is one of +the more practical tests of species validity when cross-breeding +experiments are not possible. Two cases of sympatric occurrence are +known for the species of _Syrrhophus_ in western Mexico: _modestus_ and +_nivocolimae_ are sympatric in southern Jalisco and _pipilans nebulosus_ +and _rubrimaculatus_ are sympatric in southeastern Chiapas. In eastern +Mexico, _longipes_ and _verrucipes_ are sympatric in southern Hidalgo, +and _longipes_ is sympatric with _cystignathoides_, _dennisi_, and +_guttilatus_ in southern Tamaulipas. _Syrrhophus cystignathoides_ and +_leprus_ are apparently sympatric in central Veracruz. + +Subspecific assignments have been made only when there is evidence of +intergradation. The sympatric occurrence of morphologically similar +species in this genus has led me to adopt a conservative approach to the +degree of difference philosophy. I have therefore recognized all +morphologically distinct allopatric populations as species. + + [Illustration: FIG. 21: Generic distributions of _Syrrhophus_ + (stipple) and _Tomodactylus_ (hatching). Black areas are zones + of intergeneric sympatry.] + +_Syrrhophus_ is closely allied to another Mexican leptodactylid genus, +_Tomodactylus_, which was revised by Dixon (1957), who along with +numerous other authors noted the close relationship between the two +genera. There is an almost complete lack of sympatry between the two +genera; in very few places in Mexico do they coexist (Fig. 21). +_Tomodactylus_ has its greatest diversity in the Cordillera Volcanica +and Sierra Madre del Sur, whereas _Syrrhophus_ reaches its greatest +diversity in the Sierra Madre Oriental and eastern foothills. The +species of both genera are about the same size and presumably have +similar requirements insofar as food, breeding sites, and habitat +selection. + +Four cases of intergeneric sympatry are known for the two genera: +1) the Chilpancingo region of Guerrero, 2) the lowlands of Colima and +the mountains just inland in Jalisco, 3) the lowlands of central Nayarit, +and 4) the Sierra Madre Occidental on the Durango-Sinaloan border. The +apparent sympatry in the Chilpancingo region involves four species: +_S. pipilans_, _T. albolabris_, _T. dilatus_, and _T. nitidus_. Of the +four, _T. dilatus_ appears to be completely allopatric in that it occurs +at higher altitudes (above 2000 meters), whereas the other three occur +below 1800 meters in the region (Davis and Dixon, 1965). In the +Colima-Jalisco region, _Tomodactylus_ tends to occur higher (Dixon and +Webb, 1966) than some of the _Syrrhophus_, but one subspecies of +_Tomodactylus nitidus_ is a lowland frog, occurring sympatrically with +the lowland _Syrrhophus modestus_. A similar situation is observed in +Nayarit; the lowland _Tomodactylus_ occurs sympatrically with the small +_Syrrhophus pallidus_. In both cases the _Syrrhophus_ is smaller than +the _Tomodactylus_. + + [Illustration: FIG. 22: Altitudinal distributions of _Syrrhophus_ + and _Tomodactylus_. Widths of the columns are proportional to the + numbers of species at a given altitude; narrowest width equals one + species.] + +Frogs of the genus _Syrrhophus_ tend to occur at lower elevations than +do their close relatives of the genus _Tomodactylus_ (Fig. 22). This +generalization is complicated by the occurrence in the Sierra Madre +Oriental in relatively high altitude _Syrrhophus_ (up to 2000 m.) and +the occurrence in Michoacan of low altitude _Tomodactylus_ (to sea +level). There are no _Tomodactylus_ in the Sierra Madre Oriental, +whereas the genus _Syrrhophus_ is represented in the lowlands of western +Mexico (_modestus_ group). _Syrrhophus_ and _Tomodactylus_ exhibit +essentially parapatric distributions. The two genera as now composed can +be characterized as low to moderate elevation frogs (_Syrrhophus_) and +moderate to intermediate elevation frogs (_Tomodactylus_). + + + + +LITERATURE CITED + + +BAIRD, S. F. + + 1859. Reptiles of the Boundary. United States and Mexican Boundary + Survey, pp. 1-35, pls. 1-41. + + +BARBOUR, T. + + 1923. The reappearance of Batrachyla longipes. Proc. New England + Zool. Club, 8:81-83. + + +BARBOUR, T., and A. LOVERIDGE + + 1946. Typical reptiles and amphibians; supplement. Bull. Mus. Comp. + Zool., 96:59-214. + + +BOULENGER, G. A. + + 1882. Catalogue of the Batrachia Salientia ... British Museum., + 2nd ed. + + 1888. Note on the classification of the Ranidae. Proc. Zool. Soc. + London, 1888, pt. 2:204-06. + + +CAMPBELL, H. W., and R. S. SIMMONS + + 1962. Notes on some reptiles and amphibians from western Mexico. + Bull. So. California Acad. Sci., 61:193-203. + + +CONANT, R. + + 1958. A field guide to reptiles and amphibians. Houghton-Mifflin + Co. Boston. 366 pp. + + +COPE, E. D. + + 1866. On the structures and distribution of the genera of the + arciferous Anura. J. Acad. Nat. Sci. Philadelphia, n. ser., + 6:67-112. + + 1877. Tenth contribution to the herpetology of tropical America. + Proc. Amer. Philos. Soc., 17:85-98. + + 1878. New genus of Cystignathidae from Texas. Amer. Nat., + 12:252-53. + + 1879. Eleventh contribution to the herpetology of tropical America. + Proc. Amer. Philos. Soc., 18:261-77. + + 1885. A contribution to the herpetology of Mexico. _Ibid._, + 22:379-404. + + +DAVIS, W. B., and J. R. DIXON + + 1957. Notes on Mexican amphibians, with description of a new + _Microbatrachylus_. Herpetologica, 13:145-47. + + 1965. Amphibians of the Chilpancingo Region, Mexico. _Ibid._, + 20:225-33. + + +DIAZ DE LEON, J. + + 1904. Indice de los Batracios que se enquentran en la Republica + Mexicana. Imprenta de Ricardo Rodriquez Romo. Aguascalientes. + 40 pp. + + +DIXON, J. R. + + 1957. Geographic variation and distribution of the genus + Tomodactylus in Mexico. Texas J. Sci., 9:379-409. + + +DIXON, J. R., and R. G. WEBB + + 1966. A new _Syrrhophus_ from Mexico (Amphibia: Leptodactylidae). + Cont. Sc., Los Angeles Co. Mus., 102:1-5. + + +DUELLMAN, W. E. + + 1958. A review of the frogs of the genus _Syrrhophus_ in western + Mexico. Occ. Pap. Mus. Zool. Univ. Michigan, 594:1-15. + + 1960. A distributional study of the amphibians of the Isthmus of + Tehuantepec, Mexico. Univ. Kansas Publs. Mus. Nat. Hist., + 13:19-72. + + +FIRSCHEIN, I. L. + + 1954. Definition of some little-understood members of the + leptodactylid genus _Syrrhophus_, with a description of a new + species. Copeia, (1):48-58. + + +FOUQUETTE, M. J. + + 1960. Call structure in frogs of the family Leptodactylidae. Texas + J. Sci., 12:201-15. + + +GADOW, H. + + 1905. The distribution of Mexican amphibians and reptiles. Proc. + Zool. Soc. London, 1905, pt. 2:191-244. + + +GORHAM, S. W. + + 1966. Liste der rezenten Amphibien und Reptilien.... Das Tierreich. + Lief, 85:1-222. + + +GUeNTHER, A. C. L. G. + + 1885-1902. Biologia Centrali-Americana. Reptilia and Batrachia. + 326 pp., 76 pls. Syrrhophus section dated 1900. + + +KELLOGG, R. + + 1932. Mexican tailless amphibians in the United States National + Museum. Bull. U.S. Natl. Mus., 160. + + +LANGEBARTEL, D. A., and F. A. SHANNON + + 1956. A new frog (Syrrhophus) from the Sinoloan lowlands of Mexico. + Herpetologica, 12:161-65. + + +LYNCH, J. D. + + 1963. The status of _Eleutherodactylus longipes_ (Baird) of Mexico + (Amphibia: Leptodactylidae). Copeia, (3):580-81. + + 1964. Additional hylid and leptodactylid remains from the + Pleistocene of Texas and Florida Herpetologica. 20:141-42. + + 1967. _Epirhexis_ Cope, 1866 (Amphibia: Salientia): request for + suppression under the plenary powers. I. N. (S). Bull. Zool. + Nomencl., 24:313-15. + + 1968. Genera of leptodactylid frogs in Mexico. Univ. Kansas Publs., + Mus. Nat. Hist., 17:503-15. + + +MARTIN, P. S. + + 1958. A biogeography of reptiles and amphibians in the Gomez Farias + region, Tamaulipas, Mexico. Misc. Publs. Mus. Zool. Univ. + Michigan, 101:1-102. + + +MILSTEAD, W. M., J. S. MECHAM, and H. MCCLINTOCK + + 1950. The amphibians and reptiles of the Stockton Plateau in + northern Terrell County, Texas. Texas J. Sci., 2:543-62. + + +NEILL, W. T. + + 1965. New and noteworthy amphibians and reptiles from British + Honduras. Bull. Florida State Mus., 9:77-130. + + +NIEDEN, F. + + 1923. Anura I ... Das Tierreich. Lief., 46:1-584. + + +PETERS, W. + + 1871. Ueber neue Amphibien ... des Konigl. Zoologischen Museums. + Monatsb. k. k. Preuss. Akad. Wiss. Berlin, 1870:641-52. + + +SCHMIDT, K. P., and T. F. SMITH + + 1944. Amphibians and reptiles of the Big Bend Region of Texas. + Field Mus. Nat. Hist., zool. ser., 29:75-96. + + +SMITH, H. M. + + 1947. Notes on Mexican amphibians and reptiles. J. Washington Acad. + Sci., 37:408-12. + + +SMITH, H. M., and E. H. TAYLOR + + 1948. An annotated checklist and key to the Amphibia of Mexico. + Bull. U.S. Natl. Mus., 194:1-118. + + +STEJNEGER, L. + + 1915. A new species of tailless batrachian from North America. + Proc. Biol. Soc. Washington, 28:131-32. + + +TAYLOR, E. H. + + 1940a. A new eleutherodactylid frog from Mexico. Proc. New England + Zool. Club, 18:13-16. + + 1940b. Two new anuran amphibians from Mexico. Proc. U.S. Natl. + Mus., 89:43-47, 1 pl. + + 1940c. A new Syrrhophus from Guerrero, Mexico. Proc. Biol. Soc. + Washington, 53:95-98, 1 pl. + + 1940d. New species of Mexican Anura. Univ. Kansas Sci. Bull., + 26:385-405. + + 1940e. Herpetological miscellany no. I. _Ibid._, 26:489-571. + + 1942. New Caudata and Salientia from Mexico. _Ibid._, 28:295-323. + + 1943. Herpetological novelties from Mexico. _Ibid._, 29:343-61. + + 1952. A review of the frogs and toads of Costa Rica. _Ibid._, + 35:577-942. + + +TAYLOR, E. H., and H. M. SMITH + + 1945. Summary of the collections of amphibians made in Mexico under + the Walter Rathbone Bacon Traveling Scholarship. Proc. U.S. + Natl. Mus., 95:521-613. + + +TIHEN, J. A. + + 1960. Notes on Late Cenozoic hylid and leptodactylid frogs from + Kansas, Oklahoma and Texas. Southwest. Nat., 5:66-70. + + +WRIGHT, A. H., and A. A. WRIGHT + + 1949. Handbook of frogs and toads. 3rd ed. Comstock. 640 pp. + + +YARROW, H. C. + + 1882. Checklist of North American Reptilia and Batrachia, with + catalogue of specimens in U.S. National Museum. Bull. U.S. + Natl. Mus., 24:1-249. + + +ZWEIFEL, R. G. + + 1960. Results of the Puritan-American Museum of Natural History + Expedition to Western Mexico. 9. Herpetology of the Tres + Marias Islands. Bull. Amer. Mus. Nat. Hist., 119:81-128. + + + + +TRANSCRIBER'S NOTES + + +Although _Syrrhophus marnocki_ and _Syrrhophus marnockii_ both appear +in this text, a literature search shows that both spellings have been +used and the two instances where there is only one "i" at the end are +in reference to priviously published names. Therefore, they were left +as is. With the exception of the list below and a number of silent +corrections, the text presented is that of the original printed version. + +Typographical Corrections + + Page Correction + ==== ====================== + 3 otherwse => otherwise + 5 poltypic => polytypic + 12 interorbtal => interorbital + 14 neublosus => nebulosus + 16 Cuidad => Ciudad + 16 1946-170 => 1946:170 + 22 rubrimacultaus => rubrimaculatus + 27 resemblence => resemblance + +Text Emphasis + + _Text_ - Italics + + =Text= - Bold + + + + + + +End of the Project Gutenberg EBook of A Taxonomic Revision of the +Leptodactylid Frog Genus Syrrhophus Cope, by John D. Lynch + +*** END OF THIS PROJECT GUTENBERG EBOOK A TAXONOMIC REVISION OF THE *** + +***** This file should be named 37809.txt or 37809.zip ***** +This and all associated files of various formats will be found in: + http://www.gutenberg.org/3/7/8/0/37809/ + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper 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 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 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 +business@pglaf.org. 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 + gbnewby@pglaf.org + + +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. diff --git a/37809.zip b/37809.zip Binary files differnew file mode 100644 index 0000000..63baf42 --- /dev/null +++ b/37809.zip diff --git a/LICENSE.txt b/LICENSE.txt new file mode 100644 index 0000000..6312041 --- /dev/null +++ b/LICENSE.txt @@ -0,0 +1,11 @@ +This eBook, including all associated images, markup, improvements, +metadata, and any other content or labor, has been confirmed to be +in the PUBLIC DOMAIN IN THE UNITED STATES. + +Procedures for determining public domain status are described in +the "Copyright How-To" at https://www.gutenberg.org. + +No investigation has been made concerning possible copyrights in +jurisdictions other than the United States. Anyone seeking to utilize +this eBook outside of the United States should confirm copyright +status under the laws that apply to them. diff --git a/README.md b/README.md new file mode 100644 index 0000000..2c765f8 --- /dev/null +++ b/README.md @@ -0,0 +1,2 @@ +Project Gutenberg (https://www.gutenberg.org) public repository for +eBook #37809 (https://www.gutenberg.org/ebooks/37809) |
