Geographic Variation and Systematics of the Middle American Caecilians, Genera Dermophis

and Gymnopis
Author(s): Jay M. Savage and Marvalee H. Wake
Source: Copeia, Vol. 1972, No. 4 (Dec. 29, 1972), pp. 680-695
Published by: American Society of Ichthyologists and Herpetologists (ASIH)
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 GeographicVariationand Systematicsof the Middle American
Caecilians, Genera Dermophis and Gymnopis
ANDMARVALEE
JAY M. SAVAGE H. WAKE

Analysis of variation in Middle American caecilians of the genera

Dermophis and Gymnopis requires substantial revision of their systematics.
Three species of Dermophis are recognized: D. oaxacae Mertens, 1930, of
southern Pacific Mexico; D. mexicanus Dumeril and Bibron, 1841, of both
coasts of southern Mexico southward to Atlantic slope Costa Rica and
Pacific slope western Panama; and D. parviceps Dunn, 1924, known from
both slopes of southern Costa Rica south through Panama to northern
Colombia. The nominal taxa D. clarkii (Barbour, 1926); D. costaricense
Taylor, 1955; D. eburatus Taylor, 1968; D. gracilior (Gunther, 1902); and
D. septentrionalis Taylor, 1968, are placed in the synonymy of D. mexi-
canus. D. balboai Taylor, 1968; D. glandulosus Taylor, 1955; D. occi-
dentalis Taylor, 1955, are regarded as synonyms of D. parviceps. Gymnopis
is a monotypic genus containing the single form G. multiplicata Peters, 1874.
G. oligozona (Cope, 1877) and G. proxima (Cope, 1875) are synonyms.
The nominal monotypic genus Cryptopsophis Boulenger, 1883, is based
upon specimens of G. multiplicata.
Members of the two genera are tropical lowland to premontane in distri-
bution and are found in appropriate habitats from near sea level to alti-
tudes of 900 m in southern Mexico and Guatemala and to 1400 m in Costa
Rica and western Panama. Unquestioned sympatry is known only between
D. parviceps and G. multiplicata in Pacific Costa Rica, but probably occurs
between D. mexicanus and oaxacae in southern Pacific Mexico and be-
tween D. mexicanus and D. parviceps along the Atlantic slopes of southern
Costa Rica.
Both genera are Middle American Element endemics that have under-
gone evolution in isolation from their congeners in South America during
the separation of Central America from South America during Eocene to
early Pliocene. Present distribution patterns of the four Middle American
species reflect effects of recent physiographic changes in the region.

INTRODUCTION and Gymnopis the sensory tentacle and its

HE caecilians (Order Gymnophiona or
Apoda), limbless blind burrowing am-
phibians that superficially resemble large
earthworms, are among the most bizarre of
tropical animals to the extratropical observer
seeing them for the first time. Even ex-
perienced resident tropical scientists rarely
know of their existence in regions where they
occur. In the present paper our aim is to
elucidate patterns of geographic variation in
Middle American solda con soldas or dos
cabezas (local names for caecilians) and relate
these patterns to clarifying their systematics.
Taylor (1968) in the most recent taxonomic
review recognized six genera in Mexico and
Central America: Caecilia, Copeotyphlinus,
Cryptopsophis, Dermophis, Gymnopis and
Oscaecilia. In Cryptopsophis, Dermophis
680
foramen in the maxilla are located well pos-
terior to the nostril, as compared to the other
three genera which have the tentacle and its
foramen immediately below the nostril.
Caecilia and Oscaecilia are South American
genera that range into eastern parts of
western Panama (Dunn, 1942; Taylor, 1968,
1969). No attempt is made here to review
their status. Copeotyphlinus, a genus pro-
posed by Taylor (1968) for a form left
insertae sedis by Dunn (1942) is probably
based on an Asian specimen, as discussed
below. Within the Middle American genera
with the sensory tentacle and its foramen
far posterior of the nostril, Taylor included
ten species and one subspecies of Dermophis
(Colombia to Mexico) and three species of
Gymnopis (Panama to Mexico). He further

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 SAVAGE AND WAKE-MIDDLE
re-established an additional monotypic genus,
Cryptopsophis (Costa Rica and Nicaragua),
intermediate between the other two genera
in lacking splenial teeth, as in Dermophis
(present in Gymnopis), and in having the
orbit covered by the squamosal bone, as in
Gymnopis (eye in an open socket in Dermo-
phis). The following account is restricted
to a study of the 14 nominal species included
within these two genera.
PATTERNSOF VARIATION
Characteristics.-The principal features dis-
tinguishing populations of allied caecilians
have been discussed by Dunn (1942) and Tay-
lor (1968). A preliminary survey of Middle
American material indicates that only three
features readily recognizable externally ex-
hibit consistent populational variation: 1)
adult size, total length in millimeters; 2)
number of primary body segments; and 3)
number of body segments marked by a
secondary integumentary division. The num-
ber of primary skin folds is slightly less than
the number of vertebrae, but where evident
they are correlated with vertebral and body
segment limits. The dorsal head musculature
and the hyobranchial and jaw musculature
extend over the anteriormost vertebrae.
The musculature obscures a fold-vertebra
relationship in the neck region, but it is in-
volved in the structure of the broad nuchal
collars. At the posterior end of the body,
folds may not be present, although a series
of two to five tiny vertebrae extends past the
vent. Secondary folds occur mid-segmentally
in both Dermophis and Gymnopis and the
presence of dermal bony scales is usually as-
sociated with the presence of these folds.
The secondary folds encircle the body com-
pletely only posteriorly in Middle American
forms; they are short mid-dorsal folds an-
teriorly and become progressively more com-
plete posteriorly.
Data for size, numbers of primary folds
and numbers of secondary folds have been
taken from Dunn (1942) and Taylor (1968)
and specimens not reported by them, prin-
cipally from Costa Rica. We have followed
Taylor's method of counting primary folds.
Dunn's counts are for the number of primary
grooves and include two anterior grooves
(the nuchal grooves of Taylor) that are not
associated with vertebrae. Taylor's counts
are used for all the specimens he examined
but we have not attempted to revise Dunn's
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AMERICAN CAECILIANS 681
figures. For an animal with 177 primary
folds, Dunn's method records 179 or 180
primary grooves (counting one or two nuchal
grooves and the first groove before the first
primary fold as primary grooves). Neverthe-
less Taylor's counts for primary folds on a
particular specimen frequently are identical
to Dunn's for the same animal. Since a vari-
ation of two or three folds is well within any
expected counting error, or anticipated vari-
ational limits, use of Dunn's counts does not
greatly affect interpretation or comparisons.
The total number of folds, primaries plus
secondaries, may also be used as a character,
and in practice is most useful in keys and
for rapid determination, for discrimination
between primaries and secondaries requires
careful counting, usually under a micro-
scope. Since the number of primaries and
secondaries vary independently of one an-
other, populational comparisons are ex-
pedited by the use of the single combined
count.
Taylor (1968) emphasized two other fea-
tures as useful in distinguishing populations
within this group of genera. The first of
these is coloration which he records variously
as being violet, lilac-slate, dark plumbeous,
brownish-ivory or old ivory. In our experi-
ence all Middle American caecilians observed
alive are very dark purple to purple-black
dorsally, with the venter creamy white.
Freshly preserved material is slate-grey above
and dirty-white below. After a long period
in alcohol the animals turn brown or may
even fade to yellow when exposed to light
for some time. All specimens mentioned by
Taylor as brown or ivory were collected in
the previous century and are faded.
Also used by Taylor were the row counts
of dermal scales imbedded in each fold on
the posterior part of the body. These large,
thin structures form a corselet of scales sev-
eral layers deep around the body posteriorly.
Taylor records from two to eight layers
(rows) for various populations in Middle
America. The difficulty of counting the
imbricate layers is exemplified by Taylor
himself in Dermophis oaxacae. In his key
(p. 460) and text, the range is given as five
to six layers; however, in the table (p. 498)
specimens are indicated as having three to
five rows. There appear to be no consistent
differences among the samples in this feature
and the presumed differences appear to be
682 COPEIA, 1972, NO. 4

Fig. 1. Geographicvariationin primaryfolds in Middle AmericanDermophis.

artifacts of counting, due to the imbricate
arrangement of layers.
In the following sections, geographic vari-
ation patterns are described for Central
American caecilians. We follow the philos-
ophy discussed and supported by Savage and
Heyer (1967), which regards the subspecies
as an outmoded, arbitrary and misleading
taxonomic category in herpetology. For this
reason, we have concentrated on patterns,
not nomenclature in this paper.
We have also followed the advice of Stuart
(1963:12) in purposely avoiding use of dia-
critical marks for Spanish words, because
of the potential erratic and inconsistent
errors from origin, manuscript, and proof to
printed page.
Within the body of the text, locality data
are presented with the primary geographic
area first, followed in succession by each
smaller unit as: Mexico: Oaxaca: nr. Puerto
Angel.
On the maps indicating geographic vari-
ation in segmental counts (Figs. 1-7) we have
indicated values for all examples with ac-
curate locality data. In some instances, ad-
ditional specimens with data for only the
state, department or province of origin have
been used in establishing the extreme limits
of variabiliity. In one case (Figs. 1-2), counts
for an example from Nicaragua, without
further data, are indicated adjacent to the
country and in another (Figs. 4-5), counts
for a caecilian from Panama: Chiriqui, are
shown without a locality dot, since these are
the only specimens known from the two
regions.
Geographic Variation in Dermophis.-The
data on Dermophis indicate that two popu-
lational series are represented on the basis
of total length and annular counts. The first
group (large forms) contains samples in
which the adults are large, robust animals
with sexually mature individuals in excess
of 300 mm (300-600 mm). The second
group (small forms) contains populations of
slenderer, smaller caecilians with maximum
adult sizes to 356 mm (200-356 mm). The
large forms range from western Panama into
Mexico and the smaller forms range from
Colombia to Costa Rica. The two groups are
known to be sympatric only in southeastern
Costa Rica at Moravia de Chirripo (Pro-
vincia de Limon) although sympatry is to
be expected in both Atlantic and Pacific
western Panama.
The geographic variation in the number
of primary and secondary folds for samples

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 SAVAGE AND WAKE-MIDDLE
of the large form is mapped (Figs. 1-2). Six
geographic subdivisions may be recognized
on the basis of discontinuities in character-
istics and distribution.
Pacific Mexico (Guerrero to Chiapis)
Atlantic Mexico (central Veracruz to Ta-
basco)
Pacific Middle America (Oaxaca, Mexico,
south to western Nicaragua)
Honduras (Atlantic drainage in northwest)
Costa Rica (Atlantic slopes)
Panama (Pacific western)
Comparisons of these populations indicate
that two sympatric species occur on the
pacific slopes of Mexico (the nominal species
D. oaxacae and D. mexicanus). The former
has 121-137 primaries and 101-121 second-
aries (total 224-258), the latter 102-109 pri-
maries and 54-78 secondaries (total 164-184).
The remaining populations are more closely
allied to D. mexicanus than to D. oaxacae,
as may be seen from the following summary
of segmental counts:
Primaries Secondaries
Pacific Mexico 121-137 101-121
Atlantic Mexico 104-107 51-72
Pacific Middle 121 103-106
America or or
102-112 54-78 (81?)
Honduras 104-107 40-41
Costa Rica 111-117 78-96
Panama 94-102 68-78
Within the series, it is clear that all non-
Mexican specimens belong to the same taxon.
The caecilians of this group from Pacific
Guatemala, El Salvador, Honduras and
western Nicaragua agree in annular counts
with the Atlantic and Pacific slope popu-
lations in Mexico. The allopatric population
in northwestern Honduras differs from the
Atlantic Mexico and Pacific Middle American
series in having a very low number of sec-
ondaries (40-41). All examples of this popu-
lation are from the same small area in north-
western Honduras at Tela and near San
Pedro Sula. A single specimen (USNM
128084) is known from a few kilometers to
the southwest in Provincia de Santa Barbara:
near Quimistan. This example has 106 pri-
maries and 70 secondaries (total 176) and we
regard it as being allied to the Pacific Middle
American population and have not included
it in our Honduran sample. Although
separated by a distance of about 550 km,
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AMERICAN CAECILIANS 683
the southernmost Atlantic Mexico specimen
(BM 1907.12.19.155) from Tabasco: Teapa,
suggests that a dine in secondary counts
runs from north to south along the Carib-
bean coast. The Teapa caecilian has 104
primaries and 51 secondaries (total 155), only
ten more secondaries than in the Honduras
population. Habitat suitable for Dermophis
occurs between Tabasco and Atlantic Hon-
duras in southern Mexico and Guatemala.
We predict that individuals conspecific with
both Mexican and Honduran populations
but with intermediate secondary counts will
be found in this area.
The southernmost locality for the Pacific
Middle American population is in northern
Nicaragua: Leon: El Polvon. Caecilians of
this series probably occur between this lo-
cality and the next known collecting site
350 km to the southeast on the Atlantic
slopes of the Costa Rican cordilleras. The
latter samples from Heredia: La Cinchona
and Cartago: Moravia de Chirripo, have high
secondary counts (78-96) and average higher
in primaries (114) than any non-Mexican
Totals
sample. Specimens from higher elevations
in El Salvador and Honduras approach these
values: secondaries 67-81, primaries x 107.
Total primary and secondary folds in the
Costa Rican series is 193-212 (x 202) com-
pared to the latter specimens with 176-185
(x 180). These values suggest a clinal increase
in the number of secondaries with altitude
and we regard the Costa Rican sample as a
southern component of the Pacific Middle
American series of populations. Closely
allied to the Costa Rican population is that
of Pacific western Panama. The Panamanian
specimens have fewer secondaries (68-78
versus 78-96 in Costa Rica), primaries (94-
102 versus 111-117 in Costa Rica) and total
counts (170-173 versus 193-212 in Costa
Rica). These values fall well within the
limits of this populational system to the
north and we regard the Panama population
as the southern terminal link in the chain of
populations of large Dermophis in Middle
America.
The following names are based on indi-
viduals from the indicated populations.
Pacific Mexico, Gymnopis multiplicata oax-
acae Mertens, 1930 (holotype: Senckenberg
Mus. 22130); Mexico: Oaxaca: between
Puerto Angel and Salina Cruz: Cafetal Con-
cordia, 900 m.
Pacific Middle America, Dermophis ebu-
ratus Taylor, 1968 (holotype: UM 65674);
684 COPEIA, 1972, NO. 4

Fig. 2. Geographicvariationin secondaryfolds in Middle AmericanDermophis.

Nicaragua; the counts for the type, primaries 1955 (holotype: KU 36343); Costa Rica:
107, secondaries 70 and total 177, place it Heredia: Isla Bonita (Cinchona), 1200 m.
well within the variation in Mexican, Guate- Panama, Gymnopis gracilior Gunther, 1902
malan, and Salvadorean samples. Taylor (holotype: BM 1901.12.19.137); Panama:
referred a second example to this species Chiriqui.
(Acad. Nat. Sci. Phila. 4925) from Salvador: Siphonops mexicanus Dumeril and Bibron,
Volcan Izalco, because of its high counts; 1841 (holotype: Paris Museum 5c); Mexico,
primaries 112, secondaries 67 and total 179. the counts of primaries 105, secondaries 60
These values correspond closely to other and total 165 do not permit allocation to
Salvadorean examples, primaries 102-110, either Atlantic or Pacific populations. Our
secondaries 70-81, and totals 176-183. The conclusion is that two species, Dermophis
presumed color differences of this taxon oaxacae and Dermophis mexicanus are rep-
mentioned by Taylor are due to fading with resented by these populations and names.
age. The samples of small slender Dermophis
Dermophis septentrionalis Taylor, 1968 (Fig. 3) are restricted to Costa Rica, Panama
(holotype: Hamburg Museum 2); unknown and northern Colombia. Data for all avail-
locality, listed as Guatemala by Dunn (1942), able specimens of this series are summarized
said to be labeled Brazil by Taylor (1968); below:
the counts of 103 primaries, 73 secondaries
Total Sec-
and a total of 176 leaves no doubt that it Length Primaries ondaries Total
is conspecific with D. mexicanus; it probably PACIFICCOSTA RICA:
is from this population on the basis of Villa Colon
secondary and total counts. USC7032 235 95 31 126
Honduras, Gymnopis clarkii Barbour, Pozo Azul
1926 (holotype: MCZ 11047); Honduras: At- BM 1907.6.28.27 192 100 32 132
lantida: San Pedro Sula; subsequent workers Dominical Road
have consistently regarded this population as KU 36298 186 111 29 140
a subspecies of D. mexicanus. KU 36296 194 112 37 149
Costa Rica, Dermophis costaricense Taylor; KU 36297 204 112 33 145

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 SAVAGE AND WAKE-MIDDLE AMERICAN CAECILIANS 685

Fig. 3. Geographicvariation in primary and secondaryfolds in Dermophis parviceps in Central

America. At each locality upper value representsprimary folds, lower secondaryfolds.

Cerro de la IMuerteN These specimens form three allopatric

San Isidro dle El General series: a) Atlantic Costa Rica-western Pan-
UCR 1253 339 98 40 138 ama: primaries 93-101, secondaries 13-25,
UCR 1444 305 97 31 128 total 106-126; b) Pacific Costa Rica to
San Isidro de El General northern Colombia: primaries 89-100, sec-
KU 29979 162 93 46 139 ondaries 26-46, totals 118-136; c) Dominical
KU 56070 184 - - - Road, Costa Rica: 111-112, sec-
primaries
El Volcan ondaries 29-37, totals 140-149. Each of these
Minton 532 250 99 37 136 samples has been named: a) Siphonops par-
San Vito viceps Dunn, 1924 (holotype: MCZ 9407);
UCR 2826 351 98 41 139 Panama: Bocas del Toro: La Loma, 366 m;
PACIFICPA]NAMA: b) Dermophis balboai Taylor 1968 (holotype:
Cerro Azul KU 76184); Panama: Darien Tacarcuna, 550
(Cheeseman- m; c) Dermophis occidentalis Taylor, 1955
Dunn) 157 92 26 118 (holotype: KU 36298); Costa Rica: Punta-
Tacarcuna renas: Dominical Road, 15 km SSW San
KU 76184 274 89 41 130 Isidro de El General, 800 m.
Darien In addition, Taylor (1955) described a speci-
KU 108935 356 - - - men (KU 29779) of this series from Costa
ATLANTIC COSTA RICA Rica: Puntarenas: San Isidro de El General,
Moravia 700 m, as Dermophis glandulosus. A second
KU 36276 194 96 16 112 specimen (Minton 532) from Costa Rica:
KU 66828 210 99 17 116 Puntarenas: El Volcan, was referred to this
KU 66827 217 101 25 126 form (Taylor, 1968: 479) and simultaneously
to Dermophis occidentalis (Taylor, 1968:
ATLANTIC PANAMA:
La Loma
501). The counts of the two presumed D.
glandulosus and the examples from Villa
MCZ 9407 189 93 13 106 Colon and Pozo Azul, Costa Rica are inter-
COLOMBIA: mediate between Taylor's D. balboai and D.
Villa Artega occidentalis. There appears to be only a
FM 78139 254 91 43 133 single species represented by the three names.
FM 78140 222 91 45 136 Another specimen called glandulosus by

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686 COPEIA, 1972, NO. 4

Fig. 4. Geographic variation in primary folds in Middle American Gymnopis. Range of variation
indicated for Atlantic and Pacific Costa Rican samples; for detail, see Fig. 6. Triangles indicate lo-
calities for specimens referred to Cryptopsophis simus by Taylor (1968).

Fig. 5. Geographic variation in secondary folds in Middle American Gymnopis. Range of varia-
tion given for Atlantic and Pacific Costa Rican samples; for detail see Fig. 7. Triangles indicate lo-
calities for specimens referred to Cryptopsophis simus by Taylor (1968).

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 SAVAGE AND WAKE-MIDDLE AMERICAN CAECILIANS 687

Taylor (1968: 477; 1969: 612) from San Isidro maps (Figs. 6-7). The data for all available
de El General has had the head and a con- Pacific versant Gymnopis are given below:
siderable number of the anterior vertebrae
PrimariesSecondaries Total
removed. Taylor never recorded the counts COSTA RICA
for this example. 5 km NE Tilaran
We predict that Pacific and Atlantic popu- KU 36667 123 97 220
lations of this series will prove to intergrade Tilaran
in west central Panama, although the At- USNM 70656 129 101 230
lantic samples are distinctive in having low Cafias
secondary counts. For these reasons, we UCR 2902 119 103 222
regard the small Dermophis of lower Central Monteverde
America and northern Colombia as a single USC 7246 122 104 226
species, Dermophis parviceps (Dunn). The San Mateo
numbers of primary annuli in both Atlantic USNM 37761 129 111 240
and Pacific populations of this species are Parrita
similar (Atlantic 93-101, x 97 versus Pacific USC 8255A 123 106 229
89-112, x 99). USC 8255B 125 109 234
Geographic variation in Gymnopis.-As 2 km W San Isidro
conceived by Taylor (1968) the genus Gym- de El General
nopis contained three allopatric species: KU 36668 133 117 250
Primaries Secondaries Total KU 29778 130 110 240
Atlantic Guatemala 5.5 km E San Isidro
G. oligozona 128-131 64-76 193-206 de El General
Atlantic Honduras KU 36667 125 101 226
to Panama KU 36671 125 104 229
G. proxima 112-124 84-107 201-226 Dominical Road
Pacific Costa Rica KU 36670 131 110 241
and Panama El Volcan
G. multiplicata 119-133 97-117 217-250 Minton 533 128 111 239
Isla del Cano
The variation in the numbers of primaries UCR 3138 126 108 234
and secondaries is mapped (Figs. 4-5) for UCR 3139 123 108 231
these populations. In terms of primary UCR 3140 131 112 243
counts, the Guatemala sample agrees closely UCR 3141 127 109 236
with specimens from adjacent Honduras and PANAMA:
Pacific Costa Rica. The very low secondary Veragua
counts (64-76) are approached in southern Berlin 3705 131 105 236
Atlantic lowland Costa Rica and adjacent
western Panama (84-107). We regard the The Pacific versant specimens from the
Guatemalan population as a distinctive allo- vicinity of Tilaran, a series of five caecilians
patric form of the wide ranging Caribbean from El Silencio de Tilaran on the Atlantic
Gymnopis, although future collecting in the slope just to the east and an example from
200 km range gap between Alta Verapaz and Laguna Arenal slightly farther east suggest
Progreso, Honduras, may prove that two spe- that gene flow occurs between the two lowland
cies are involved. populations across the low divide between
The continued recognition of Atlantic and Provincias de Guanacaste and Alajuela in
Pacific lowland species is even more dubious. northwest Costa Rica. The most closely ad-
Primary, secondary, and total counts overlap jacent samples of the Atlantic lowland pop-
between the nominal forms although the ulations (Cariblanco and Puerto Viejo) show
means average about 10 more primaries, 10 an approach to Pacific Gymnopis in counts,
while the Arenal, Silencio, and Tilaran ex-
more secondaries, and 20 more in total for
Pacific samples compared with the Atlantic amples suggest a gradual change from the
upland Atlantic slope (114-95-209 at Arenal
population. The key to solution of this prob- 117-96-218 at Silencio), at the divide (123-97-
lem lies in Costa Rica, and we have mapped 220), and on the Pacific slope (129-101-230)
primary and secondary counts for all samples from one population to another as seen in
from the republic on the accompanying Table 1. The single specimen from Cafias

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688 COPEIA, 1972, NO. 4

Fig. 6. Geographicvariationin primaryfolds in CostaRican Gymnopis.

on the Pacific lowlands just west of Tilaran
has counts (119-103-222) very close to those
for samples from the Atlantic lowlands.
Along the Pacific coast there is a general
trend toward higher counts from Cafias and
Tilaran southward (Figs. 7-8). These data
confirm that intergradation probably takes
place between G. proxima (Atlantic) and G.
multiplicata (Pacific) in much the same
fashion as reported for Atlantic and Pacific
populations of the frog Agalychnis callidryas
in the same area (Savage and Heyer, 1967).
As in the latter case, the Pacific lowland
population resembles more closely Atlantic
Honduranean samples than those of immedi-
ately adjacent Atlantic Costa Rica. The gene
exchange between Atlantic and Pacific Costa
Rican populations of Gymnopis may be
rather recent, after a period of allopatric
isolation. Nevertheless, we conclude that
only a single species, Gymnopis multiplicata
Peters, is involved.

TABLE1. SUMMARY
OF DATA FORCOSTARICANGymnopis.

Number of
Specimens Primaries Secondaries Totals

Atlantic 28 112(119)124 84 (97)107 199(215)226

Puerto Viejo 9 120(121)124 95(100)103 208(221)226
Cariblanco 2 114(116)118 95 (99)103 209(215)221
Laguna Arenal 1 114 95 209
El Silencio 5 112(117)120 91 (96)100 203(218)220
Tilaran 1 123 97 220
Tilaran, 5 km NE 1 129 101 230
Pacific 15 119(126)133 97(108)117 220(234)250

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 SAVAGE AND WAKE-MIDDLE AMERICAN CAECILIANS 689

Fig. 7. Geographic variation in secondary folds in Costa Rican Gymnopis.

The Status of Cryptopsophis simus.-Taylor Isla Boquete

(1969:455) resurrected Boulenger's 1883 genus AMNH 8398
Cryptopsophis for nine individuals he re- 126 106 232
garded as conspecific with Cope's (1878) El Bluff, Blue.fields
Siphonops simus from Costa Rica. The AMNH 8397
locality data and counts for these animals are 119 101 220
given below: Bluefields
JSNM377351
Primaries Secondaries Total
No data 121 100 221
BM 83.7.27.26 San Juan Nor:te
1119 100 219 Genoa 37163
COSTARICA 118 97 215
USNM 29765 VARIATION
1220 101 221 117(120)126 97(101)106 215(222)232
NICARAGUA:
Dunn (1942) referred the type of simus
Eden Mine
AMNH 8399 (USNM 29765) to Gymnopis m. multiplicata
and all the other specimens mentioned by
~120
106 226
Santa Ana, Rio Taylor, except Genoa 37163 and AMNH
Huahuashan 54985, to Gymnopis m. proxima. The latter
AMNH 54985
two specimens were not seen by Dunn. The
117 100 217 counts for simus are almost identical to those
HaciendaValencia, for Gymnopis multiplicata from northeastern
Chontales Costa Rica (Puerto Viejo) and eastern Nica-
AMNH 8396 ragua. Cryptopsophis simus was said to differ
122 106 228 from Gymnopis in lacking a single splenial

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690 COPEIA, 1972, NO. 4
Fig. 8. Distribution of Middle American caecilians with principle late Cenozoic biogeographic
shifts indicated by arrows. See text for full discussion.
tooth (present in the latter) on each lower
jaw ramus.
We have re-examined the holotype and the
five examples at the American Museum
referred to C. simus by Taylor. Splenial teeth
are present in all except AMNH 8396. The
single tooth on each mandible is located at
the lingual margin of the mandibular shelf
just behind the raised median ridge that
forms the articulating surface for the inter-
mandibular joint. The teeth are typical
amphibian bipartite structures with a distal
crown separated by a well-marked transverse
groove from the proximal pedicel. In the
holotype the mandibles apparently have been
cut apart so that a small piece of the right
ramus is missing. On the left ramus the
splenial tooth has been broken off at the
level of the jaw but can be recognized as a
tooth structure under magnification (20X).
In AMNH 8397, the crowns of the splenial
teeth barely pierce the gum, but are obviously
present. AMNH 8398 has the mandibles
partially dissected with an incomplete sple-
nial tooth pedicel on the right ramus and
only the basal bony depression for the miss-
ing tooth on the left. AMNH 8399 has had
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the gum removed and two complete splenial
teeth are present. The tooth is broken on the
left side in AMNH 54985 and the right tooth
apparently has been removed during dissec-
tion of the gum, since a basal bony depression
for the missing tooth is indicated.
The left ramus of AMNH 8396 has been
cut a few millimeters to the left of the
mandibular symphysis, destroying the area
where a splenial tooth might have been lo-
cated. The symphyseal articulation is pecu-
liarly massive in this example and only a
slight depression at its posterior margin on
the right ramus suggests that a splenial tooth
was present. Prior dissection of this animal
has made it impossible to tell if splenial teeth
were once present.
There seems little question that these ex-
amples are conspecific with Gymnopis multi-
plicata. It is very likely that the other three
examples called C. simus by Taylor also have
splenial teeth or their absence is an artifact.
The holotype of the type species Cryptop-
sophis is a large example of Gymnopis multi-
plicata, which requires that the genus and
its only nominal species be placed in synon-
ymy of the earlier name.
 SAVAGE AND WAKE-MIDDLE
SYSTEMATICS
Status of Nominal Forms.-The genera
Dermophis and Gymnopis, as conceived in
this report, are comprised of three and one
species respectively. The genera are mem-
bers of the endemic Middle American Ele-
ment in the sense of Savage (1966). The
genus Dermophis now ranges into Colombia.
The species of Middle American caecilians
may be distinguished by the following key,
which includes the species of Caecilia and
Oscaecilia for the sake of completeness. No
attempt has been made to treat the genus
Copeotyphlinus Taylor, 1968, erected for
Cope's (1866) description of Siphonops syn-
tremus based upon a now lost specimen said
to have been from Honduras or British Hon-
duras. Dunn (1942) and Taylor agree that
the description cannot be that of an Ameri-
can caecilian. We concur and suggest that it
was a mislabeled Uraeotyphlus from Asia.
Taylor (1968) has provided detailed synon-
ymies and descriptions of individual animals.
These are not repeated here and reference
should be made to his work for full literature
citations as we have only listed publications
discussed in the text of the present paper.
The key and species diagnoses for Dermophis
and Gymnopis given below will provide a
means of identification of the Middle Amer-
ican forms.
Taxonomic changes from Taylor's 1968
revision (page citations indicated) as devel-
oped in the previous sections are:
Dermophis balboai Taylor, 1968 = Der-
mophis parviceps (Dunn, 1924); p. 461
Dermophis costaricense Taylor, 1955 = Der-
mophis mexicanus (Dumeril and Bibron,
1841); p. 467
Dermophis eburatus Taylor, 1968 = Der-
mophis mexicanus; p. 473
Dermophis glandulosus Taylor, 1955 = Der-
mophis parviceps; p. 475
Dermophis gracilior (Gunther, 1902) = Der-
mophis mexicanus; p. 479
Dermophis mexicanus clarkii (Barbour, 1926)
= Dermophis mexicanus; p. 483
Dermophis occidentalis Taylor, 1955 = Der-
mophis parviceps; p. 501
Dermophis septentrionalis Taylor, 1968 =
Dermophis mexicanus; p. 513
Gymnopis oligozona (Cope, 1878) = Gym-
nopis multiplicata Peters, 1874; p. 455
Gymnopis proxima (Cope, 1875) = Gymnopis
multiplicata; p. 525
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AMERICAN CAECILIANS 691
Cryptopsophis Boulenger, 1883 = Gymnopis
Peters, 1874; p. 455
Cryptopsophis simus (Cope, 1878) = Gym-
nopis multiplicata; p. 455
A Key to Middle American Solda Con Soldas
A. Tentacle and tentacular foramen located
well posterior to nostril on side of head or
just anterior to eye.
B. Tentacle just anterior to eye, tentacular
foramen at or near center or maxillary
bone; orbit roofed over by squamosal
bone; a single splenial tooth on each
ramus of lower jaw so that there are two
tooth series (dentary and splenial); pri-
mary folds 112-133, secondaries 64-117,
total 193-250. . . . Gymnopis multiplicata
(Guatemala to western Panama on the
Caribbean slope and Costa Rica and
western Panama on the Pacific coast).
BB. Tentacle about halfway between eye
and nostril, tentacular foramen in ante-
rior margin of maxillary bone; orbit not
roofed by bone; no splenial teeth so only
one tooth row in each lower jaw ....
................. Dermophis.
C. Primary folds 121-137, secondaries
101-121, total 222-258.... D. oaxacae
(southern Pacific Mexico).
CC. Primary folds less than 120, second-
aries less than 100, total less than 220.
D. Total number of folds usually less
than 145; adults less than 360 mm
total length; secondary folds less
than 50.... D. parviceps (both slopes
of southern Costa Rica, northwest-
ern Panama, south into northern
Colombia).
DD. Total number of folds usually
more than 144; adults 300-600 mm
total length; secondary folds usually
more than 50. ... D. mexicanus
(both coasts of Mexico, south to
Honduras on the Atlantic and Nica-
ragua on the Pacific; Atlantic slopes
of Costa Rica and Pacific western
Panama).
AA. Tentacle and tentacular foramen im-
mediately below nostril.
E. Orbit roofed over by bone; primary
folds 169-189, secondaries 16-31, total
200-218 .... Oscaecilia ochrocephala
(eastern Panama to northern Colombia).
EE. Orbit not roofed over by bone ....
. . . . . . . . . . . . . . . . . . . Caecilia
692 COPEIA, 1972, NO. 4
F. Primaries 226-231, no secondaries
*... . C. elongata (extreme eastern
Panama).
FF. Primaries less than 200, at least 14
secondaries.
G. Primaries 150-192, secondaries 28-
62, total 209-252. . . . C. nigricans
(extreme eastern Panama south to
northwest Ecuador).
GG. Primaries less than 149.
H. Primaries 131-147, secondaries
12-21, total 143-168 . . . C. ten-
taculata (extreme eastern Panama
south to Surinam and Peru).
HH. Primaries 112-124, secondaries
14-31, total 126-156. . . . C. vol-
cani (El Valle de Anton, Panama).
Species Diagnoses
Dermophis mexicanus-distinguished from
all other Middle American species except D.
parviceps, G. multiplicata and C. volcani by
its low numbers of primary folds (94-117 in D.
mexicanus), the remaining species always
have more than 120 primaries; from C.
volcani this species differs in having the ten-
tacle located on the side of the head, slightly
closer to the eye than the nostril (tentacle
under the nostril in the former); G. multi-
plicata has the tentacle just anterior to the
orbit, the eye under the squamosal bone and
splenial teeth present (tentacle well forward
of orbit, orbit not roofed over by bone and
no splenial teeth in D. mexicanus).
Dermophis oaxacae-immediately distin-
guished from all Middle American caecilians
except G. multiplicata by its high number of
secondary folds (101-121); in the other forms
these folds number less than 100. D. oaxacae
differs from G. multiplicata in having the
orbit not roofed over by bone, the tentacle
well forward of the eye and in lacking sple-
nial teeth (orbit roofed over by squamosal,
tentacle just anterior to orbit and splenial
teeth in Gymnopis); D. oxacae occurs sym-
patrically with D. mexicanus but has consist-
ently higher counts (121-137 primaries, 103-
121 secondaries, 224-258 total in the former
versus 94-117 primaries, 40-96 secondaries,
144-212 total in the latter).
Dermophis parviceps-distinct from all
other Middle American forms except D. mex-
icanus and C. volcani in its low numbers of
primaries (89-112), primaries more than 115
in the other species; from C. volcani the spe-
cies differs in having the tentacle between the
eye and nostril rather than below the nostril;
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in D. mexicanus from the area where the two
species approach one another geographically
(Nicaragua, Costa Rica and Panama) there
are 54-96 secondaries versus 13-46 in parvi-
ceps.
Gymnopis multiplicata-the species over-
laps several other species in segmental counts
but may be distinguished from all other Mid-
dle American caecilians by having the orbit
roofed over by the squamosal, the tentacle
just anterior to the orbit and splenial teeth.
It occurs sympatrically only with D. mexi-
canus and D. parviceps. The generic charac-
ters for Gymnopis cited above serve to
separate it from these two forms (orbit open,
tentacle well forward to orbit and no splenial
teeth in Dermophis). In addition, it may be
separated from D. parviceps by its higher
segmental counts (115-133 primaries, 64-117
secondaries, 193-250 total versus 89-112
primaries, 13-46 secondaries, 106-149 total
in the latter). G. multiplicata differs from
0. ochrocephala, the only other caecilian in
the region with the orbit roofed over by the
squamosal, in its segmental counts (169-189
primaries, 16-31 secondaries for ochroceph-
ala) and tentacle position (under nostril in
ochrocephala).
DISTRIBUTION
Geographic Occurrence
The following lists include only records of
unquestionable identity based upon examina-
tion of specimens or the literature. Der-
mophis oaxacae-MEXICO: Guerrero: Xalti-
angius, 15 km N Acapulco; Oaxaca: Cafetal
Concordia, Mirador; Chiapas: 30 km NE
Escuintla, La Esperanza, E Tonala.
Dermophis mexicanus-MEXICO: Vera-
cruz: Cosamaloapan, Cuatotolapam; Ta-
basco: Tabasco, Teapa; Oaxaca: El Barrio,
Matias Romero, Tehuantepec; Chiapas: 9.6
km NE Escuintla, 4.8 km and 8 km SE
Huixtla, 2.5 km W Soconusco, 3.2 km NW
Tapachula, 1.4 km NW and Tonala, La
Zacualpa; GUATEMALA: Escuintla: Escu-
intla; Retalhuleu: Retalhuleu; Suchitepe-
quez: El Cipres; EL SALVADOR: La
Libertad; El Paraiso de Santa Tecla: Santa
Ana: Volcan Izalco; San Vicente: 16 km E
San Vicente: HONDURAS: Atlantida:
Tela; 7.2 km ENE Villanueva; Cortes: La
Lima; San Pedro Sula, 3.2 km W; Santa Bar-
bara; nr. Tuimistan; Valle: Ampala; NICA-
RAGUA: Leon: Polvon; COSTA RICA:
Heredia: La Cinchona; Cartago: Moravia de
 SAVAGE AND WAKE-MIDDLE
Chirripo; PANAMA: Cerro Azul; Darien:
Tacarcuna; COLOMBIA: Antioquia: Villa
Artega.
Gymnopis multiplicata-GUATEMALA:
Alta Verapaz: Finca El Volcan; HONDU-
RAS: El Paraiso: Arenales; Olancho: 4.5
km SE Catacamas; YORO: Progreso District;
NICARAGUA: El Cabo: Cabo Gracias a
Dios; San Ramon; Chontales: Hacienda Va-
lencia de San Miguelito; Rio San Juan: Isla
Boquete; juncture of Rio Colorado and Rio
San Juan; San Juan Norte; Zelaya: Blue-
fields and El Bluff; Eden; 80 km up Rio
Escondido; COSTA RICA: Alajuela: La-
guna Arenal; Cartago: Peralta; Turrialba;
Guanacaste: Canas; El Silencio; Tilaran;
Taboga; 5 km NE Tilaran; Heredia: Cari-
blanco; Puerto Viejo; Limon: Tortuguero;
Los Diamantes; Guapiles; near Limon;
Monteverde; Parismina; between Puerto
Vargas and Cahuita; Reventazon; Salvadora;
Suretka; Puntarenas: Monteverde; Parrita;
2 km W, 5 km E, 15 km ESE San Isidro de
El General on Dominical Road; Isla del
Cano; 1.6 km E El Volcan; San Jose: San
Mateo; PANAMA: Bocas del Toro: Almi-
rante; Coco Plum Estate; Farm Six; Chiriqui:
"Veragua."
Ecologic Occurrence
Caecilians are strictly tropical in distribu-
tion and their ecologic limits within a region
are determined primarily by humidity. In
Middle America, solda con soldas are found
in appropriate habitats from near sea-level
to altitudes of 900 m in southern Mexico and
Guatemala and to 1400 m in Costa Rica and
Panama. Their ranges are restricted to Trop-
ical Lowland and Premontane altitudinal
zones, according to the system of Holdridge
(1967).
Dermophis oaxacae ranges from 45-900 m
along the southern foothills of the Sierra
Madre del Sur and Sierra Madre de Chiapas.
Most localities are on Lowland Moist or
Premontane Wet Forests back from the dry
formations along the narrow coastal plain.
Dermophis mexicanus occurs from near
sea-level to 350 m in southern Mexico in
Lowland Dry and Moist Forests and in sa-
vanna situations within these associations.
The species is found in the Lowland Moist
Forest belt along the foothills of the southern
slopes of the main highlands in Guatemala
up to 400 m, but not in the Lowland Dry
Forest along the coastal plain. In El Salva-
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AMERICAN CAECILIANS 693
dor, this form is known only from Premon-
tane Moist Forest from 500-1000 m in
altitude. In Honduras all records are from
Lowland Dry Forest associations in both
Atlantic and Pacific drainages below 500 m.
The single precise Nicaraguan record is
from similar conditions near the Pacific
coast. Farther south the species is found in
Premontane Moist, Wet or Rainforests along
the Atlantic slope in Costa Rica and along
the Pacific slope in western Panama.
Dermophis parviceps ranges from 50 to
near 850 m in Lowland Wet, Premontane
Moist, Wet, and Rainforest in Pacific Costa
Rica. In Atlantic Costa Rica and western
Panama, a similar series of bioclimatic asso-
ciations are occupied with a vertical range
from 300-1200 m. The records from eastern
Panama and northern Colombia range be-
tween 100 and 550 m in Lowland Wet Forest.
Gymnopis multiplicata occurs from near
sea-level to nearly 900 m in altitude along the
Atlantic versant in Lowland Dry, Moist, and
Wet Forest and Premontane Moist and Wet
Forest. In Pacific Costa Rica, it ranges from
near sea-level to 1400 m in Tropical Wet,
Premontane Moist, and Wet Forests.
For the species of caecilians under consid-
eration, unquestioned sympatry is known
only between D. parviceps and G. multipli-
cata in Pacific Costa Rica in the vicinity of
San Isidro de El General (Dominical Road
and El Volcan). D. parviceps and D. mexi-
canus are both known from Costa Rica in the
vicinity of Moravia de Chirripo (1116 m)
and probably are sympatric there. D. mexi-
canus and D. oaxacae have both been taken
in the area near Tonala Chiapas (45 m) and
may overlap in several localities in the
Pacific drainage of Oaxaca and Chiapas,
Mexico.
Distribution Patterns and History
In an earlier paper, one of us (Savage,
1966) concluded that Middle American cae-
cilians were derived from ancestors that were
isolated from their South American allies by
the early Eocene-Pliocene submergence of
the Isthmian Link region. The inference by
Taylor (1968) that the two genera, Der-
mophis and Gymnopis, are only distantly
related to their probable South American
cognates Brasilotyphlus, Microcaecilia and
Parvicaecilia further strengthens this inter-
pretation. The differences among the genera,
with Gymnopis similar to Brasilotyphlus and
694 COPEIA, 1972, NO. 4
Microcaecilia and Dermophis close to Parvi-
caecilia (based on eye-tentacle-nostril posi-
tions, orbit or not, splenial teeth or not and
fold counts) suggest that both lines were
already present north and south of the Isth-
mian Link area prior to its inundation in late
Paleocene. Present distribution patterns sug-
gest that the immediate ancestors of living
forms developed in evergreen forest situa-
tions on the Atlantic lowlands. The basic
Dermophis stock seems to have evolved in the
Veracruzian center of differentiation accord-
ing to the terminology of Savage (1966) to the
north of an extensive marine embayment
through the Lago de Izabal region of Guate-
mala. Gymnopis is clearly associated with a
more southern center (Caribbean) in what is
now Honduras and Nicaragua, that was iso-
lated south of the marine barrier until some-
time after early Pliocene.
The ancestral stock of Dermophis appar-
ently crossed over the Isthmus of Tehaunte-
pec to the Pacific slope on two different
occasions. The first invasion probably led to
the allopatric speciation of D. oaxacae. A
second transisthmian movement involved
present-day D. mexicanus which then fol-
lowed the Pacific lowlands south into the
emerging Isthmian Link region of lower
Central America. Examination of the paleo-
geography of this area in early Pliocene
(Savage, 1966) shows a link connected to the
rest of Central America through what is now
western Nicaragua with separation of the
eastern lowland areas to the north by an
extensive marine embayment through the
present-day Rio San Juan basin. As the axis
of the Talamanca range of Costa Rica be-
comes uplifted to its present altitudes later
in Cenozoic, the Pacific population in the
link probably became differentiated to be-
come D. parviceps. One population of this
form has moved north around the eastern
margin of the Talamancas to reach south-
eastern Costa Rica. The population of mexi-
canus along the slopes of Atlantic Costa Rica
also underwent some differentiation (Fig. 1-
2). Late Cenozoic drying along the west
coast has eliminated any contact between
mexicanus and parviceps on the Pacific
versant and isolated Costa Rican and Pana-
manian mexicanus from the main body of the
species ranges. D. mexicanus has apparently
moved around the eastern margin of the
Talamanca-Chiriqui range from Atlantic to
Pacific slopes. It is noteworthy that counts of
secondary folds are most different in the
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populations of D. mexicanus and D. parvi-
ceps at Moravia de Chirripo, where they
approach sympatry. Were more known of
their natural history, one might postulate
character displacement as a result of the
sympatry.
On the Caribbean, D. mexicanus has rather
recently moved southward into Honduras
from the north with a more or less contempo-
raneous northward migration from the Pa-
cific lowlands.
Gymnopis multiplicata has moved north-
ward and southward from lowland Honduras
and Nicaragua after the marine barriers were
removed in the Pleistocene. In Costa Rica,
the species has crossed over into the Pacific
versant and may have been completely iso-
lated in this area from its Atlantic congeners
by the uplift of the Talamanca; only lately
has it secondarily come back into contact
with them. This pattern parallels that de-
scribed in Agalychinis callidryas by Savage
and Heyer (1967). The accompanying map
(Fig. 8) summarizes the apparent patterns and
movements as we have reconstructed them.
ACKNOWLEDGMENTS
The American Museum of Natural History
(AMNH), Charles W. Myers and Richard G.
Zweifel; Museum of Natural History, Uni-
versity of Kansas (KU), William E. Duellman;
University of Costa Rica (UCR), Douglas C.
Robinson; and the United States National
Museum (USNM), James A. Peters, loaned
us critical material and we thank them for
their cooperation. Other abbreviations indi-
cate catalogued collections as follows: BM,
British Museum of Natural History; FM,
Field Museum of Natural History; MCZ, Mu-
seum of Comparative Zoology, Harvard;
USC, University of Southern California.
The maps were executed by Ronald T.
Harris at the University of Southern Cali-
fornia.
Fieldwork in Costa Rica by the senior
author was aided by the John Simon Guggen-
hiem Memorial Foundation, the National
Science Foundation and the Organization for
Tropical Studies. Support for studies on
caecilians by the junior author was provided
by the Committee on Research at the Univer-
sity of California at Berkeley.
LITERATURE
CITED
COPE,E. D. 1866. Fourth contribution to the
herpetology of tropical America. Proc. Acad.
Nat. Sci. Phila. 18:123-132.
 PACKARD AND PACKARD-PARIETAL EYE 695

.1878. Tenth contribution to the TAYLOR,E. H. 1955. Additions to the known

herpetology of tropical America. Proc. Amer. herpetological fauna of Costa Rica with com-
Philos. Soc. 17:85-98. ments on other species. No. II. Univ. Kans.
DUNN,E. R. 1942. The American caecilians. Sci. Bull. 37:499-575.
Bull. Mus. Comp. Zool. 91(6):437-540. .1968. The caecilians of the world.
HOLDRIDGE,L. R. 1967. Life zone ecology. Sec- Univ. Kans.Press.
ond ed. Trop. Sci. Center,SanJose, CostaRica. . 1969. Skulls of Gymnophiona and
SAVAGE, J. M. 1966. The origins and history their significance in the taxonomy of the
of the CentralAmericanherpetofauna. Copeia, group. Univ. Kans. Sci. Bull. 98:585-687.
1966:719-766.
, AND W. R. HEYER. 1967. Variation DEPARTMENT OF BIOLOGICAL SCIENCES AND
and distribution in the tree-frog genus Phyl- ALLAN HANCOCK FOUNDATION, UNIVERSITY
lomedusain CostaRica, CentralAmerica. Beit. OF SOUTHERN CALIFORNIA, LOS ANGELES,
Neotrop. Fauna, 5:111-131. CALIFORNIA90007 AND DEPARTMENTOF IN-
STUART,L. C. 1963. A checklist of the herpeto-
fauna of Guatemala. Misc. Publ. Mus. Zool., STRUCTION IN BIOLOGY, UNIVERSITY OF CALI-
Univ. Michigan, 122:1-150. FORNIA, BERKELEY, CALIFORNIA 94720

Photic Exposure of the Lizard CallisaurusdraconoidesFollowing

Shielding of the ParietalEye
GARYC. PACKARD
ANDMARYJ. PACKARD

Shielding the parietal eye of zebra-tailed lizards (Iguanidae: Callisaurus

draconoides) resulted in increased exposure of males, but not females,
to ambient photic conditions. This finding is consistent with the hypothesis
that the parietal eye of lizards is an important component of a control sys-
tem regulating exposure to solar radiation.

INTRODUCTION the reproductive schedule (Stebbins, 1970;

R EMOVING or shielding the parietal (=
parapineal) eye of lizards may lead to
increased exposure to ambient photic condi-
tions (Stebbins, 1970; Stebbins and Eakin,
1958; Stebbins and Wilhoft, 1966), with in-
creased exposure usually being accompanied
by increased locomotor activity (Glaser, 1958;
Stebbins and Eakin, 1958) and by greater
utilization of those parts of the microenviron-
ment characterized by high light intensity
(Stebbins, 1963; Stebbins and Eakin, 1958).
As a working hypothesis, it has been proposed
that the parietal eye may function as a com-
ponent of a physiological "illuminometer"l
regulating the amount of exposure to solar
radiation (Stebbins and Wilhoft, 1966). In
the absence of this control system, lizards
presumably experience excessive photother-
mal stimulation leading to acceleration of
1 The term "photothermal radiation dosimeter,"
introduced by Glaser (1958) and used by subsequent
authors, is probably inappropriate for use in connection
with function of the parietal eye. Inasmuch as illu-
minance, and not radiation, is the environmental vari-
able for which data have been reported, the term
"illuminometer" is to be preferred.
Licht, 1971). Thus, ecological relevance of
the physiological "illuminometer" may be
manifest in the synchronization of repro-
ductive efforts of the lizards comprising a
population.
In the present investigation we tested the
hypothesis that the lacertilian parietal eye
functions as part of a control system pre-
venting excessive exposure to ambient photic
conditions, the object of study being the
diurnal, heliothermic zebra-tailed lizard,
Callisaurus draconoides (Iguanidae).
MATERIALS AND METHODS
Thirty-six zebra-tailed lizards were cap-
tured on the Mojave Desert approximately 3
km southeast of Boulder City, Clark County,
Nevada, from 12-16 July 1971. The 18 male
and 18 female lizards were divided into 2
groups, respectively, of 9 animals each, the
groups of a particular sex being evenly
matched with respect to size. Lizards in each
group were distinctively marked on the
dorsum with red paint, and were marked
also by toe-clipping to permit positive identi-

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