_M. hobartsmithi_, having the anterior and posterior surfaces of the
thighs and the upper arms pale pink in life and a grayish brown dorsum
in preservative. The other specimen (UMMZ 94231) has in preservative a
dark brown dorsolateral line on each side enclosing a pale tan area that
extends from the snout to the vent. One specimen from 29 kilometers east
of Morelia (UIMNH 40338) and 13 specimens from San Jose de la Cumbre
(UMMZ 102111) do not have the prominent tarsal tubercles characteristic
of _M. hobartsmithi_. Also, in these fourteen specimens the palmar
tubercles are larger, and the dark anal patch more distinct, than in
typical _M. hobartsmithi_. Possibly these specimens, which are from the
high mountains in the eastern part of Michoacan, represent another
species of _Microbatrachylus_. However, Taylor (1940d:501) reported a
series of _M. hobartsmithi_ from the mountains 10 miles west of Villa
Victoria in the western part of the state of Mexico.
The largest specimen from Michoacan is a gravid female (UIMNH 16104)
having a snout-vent length of 23.5 mm.
_Microbatrachylus hobartsmithi_ has been found in rocky ravines along
streams in the Cordillera Volcanica and the southwestern escarpment of
these mountains at elevations from 1450 to 2750 meters.
~Microbatrachylus pygmaeus~ (Taylor)
_Eleutherodactylus pygmaeus_ Taylor, Trans. Kansas Acad.
Sci., 39:352, 1937.--1 mile north of Rodriguez Clara,
Veracruz, Mexico.
_Microbatrachylus pygmaeus_ Taylor, Univ. Kansas Sci. Bull.,
26:500, November 27, 1940.
_Microbatrachylus albolabris_ Taylor, Univ. Kansas Sci.
Bull., 26:502, November 27, 1940.--2 miles west of Cordoba,
Veracruz, Mexico.
_Microbatrachylus minimus_ Taylor, Univ. Kansas Sci. Bull.,
26:507, November 27, 1940.--Agua del Obispo, Guerrero,
Mexico.
_Microbatrachylus imitator_ Taylor, Univ. Kansas Sci. Bull.,
28:70, May 15, 1942.--La Esperanza, Chiapas, Mexico.
Arteaga (328).
This large series (UMMZ 119247-8) was collected on June 22 and 23, 1958,
before the onset of the heavy summer rains. The frogs were found in a
shaded ravine at the north edge of Arteaga; they were obtained during
the day, at which time they were actively moving about in the leaf
litter along a small stream.
These frogs are all referred to _M. pygmaeus_, because this is the
earliest name available for frogs showing the variation in
characteristics displayed by this large series. The characters used by
Taylor (1936a, 1940d, 1941a, and 1942b) and Smith and Taylor (1948) to
distinguish the various species of _Microbatrachylus_ include color
pattern, relative length of the hind limb, presence and position of
dorsal dermal folds or pustules, relative size of inner and outer
metatarsal tubercles, and the number of palmar tubercles. All specimens
from Arteaga have two palmar tubercles; the inner and outer metatarsal
tubercles are subequal in size. Furthermore, aside from sexual
difference, there is little variation in the relative length of the hind
limbs (Table 3). However, many color patterns do exist in the series;
each of these color patterns is described below.
TABLE 3.--SNOUT-VENT LENGTH EXPRESSED AS A PERCENTAGE OF TIBIA LENGTH IN
ANIMALS OF SIX COLOR PATTERNS OF MICROBATRACHYLUS PYGMAEUS. (LETTERS
REFER TO THE VARIANTS HAVING THE COLOR PATTERN DISCUSSED IMMEDIATELY
BELOW)
+------------+-----------+---------+---------+----+--------+
| | | Number | Range | | Twice |
| Color | Sex | of | of |Mean|standard|
|Pattern | |specimens|variation| | error |
| | | | | |of mean |
+------------+-----------+---------+---------+----+--------+
| A | Male | 25 |51.4-57.5|55.2| 3.34 |
| | Female | 25 |49.3-54.9|51.6| 3.12 |
| B | Male | 20 |51.0-57.1|55.4| 2.44 |
| | Female | 21 |47.3-54.9|51.2| 3.52 |
| C | Male | 6 |54.5-56.2|55.2| .... |
| | Female | 6 |50.0-52.9|51.6| .... |
| D | Male | 17 |52.9-58.2|55.4| 2.64 |
| | Female | 14 |48.5-56.6|52.1| 4.16 |
| E | Male | 10 |50.9-56.9|55.1| 3.40 |
| | Female | 7 |49.6-54.5|51.6| .... |
| F | Female | 2 |51.9-52.6|52.3| .... |
+------------+-----------+---------+---------+----+--------+
A.--225 specimens: Dorsum mottled brown and cream, usually with a dark
spot between the eyes and one or two dark V-shaped marks with the apex
anteriorly on the back; 55 of these have a narrow cream-colored line
from the tip of the snout to the vent and thence onto the posterior
surfaces of the thighs. All are pustulate above; in most specimens the
pustules form no pattern, but in some they tend to form a V in the
scapular region.
B.--41 specimens: Dorsum pale tan or cream-color with brown mottling on
flanks; a brown interorbital bar and a brown chevron in scapular region.
Dorsum irregularly pustulate; in some specimens the pustules tend to
form a V in the scapular region.
C.--12 specimens: Dorsum colored like "A", but having a broad yellow
stripe narrowly bordered by black from the tip of the snout to the vent;
in some specimens there is a narrow yellow stripe on the posterior
surfaces of the thighs. The dorsum is irregularly pustulate.
D.--31 specimens: Dorsum variably streaked with cream-color or pale tan
and brown; usually a broad cream-colored stripe from eyelid to groin
bordered laterally by a somewhat narrower brown stripe; middorsal area
cream-color and separated from dorsolateral cream-colored stripe by a
brown stripe, or middorsal area brown with a cream-colored or yellow,
narrow stripe from tip of snout to vent; a dark stripe from tympanum to
flank; dorsal surfaces of heels creamy white to pale orange; anal patch
brown. A dermal ridge from posterior edge of eyelid to rump; another
ridge extends posteromedially from the eyelid; scattered pustules on the
dorsum in some specimens.
E.--17 specimens: A narrow dark stripe from snout, through nostril and
eye, over tympanum, to vent, enclosing a unicolor dorsum (reddish tan to
yellowish tan in life); heels pale tan or yellow above; anal patch
black. A faint dermal ridge from posterior edge of eyelid to rump, or
part way to rump.
F.--2 specimens: Mottled brown and cream-color above; upper lips and
upper arms white. A dermal fold from posterior edge of eyelid to rump;
scattered pustules on dorsum.
Some of these color variants are assignable to names proposed by Taylor:
"A" and "B" undoubtedly are _M. pygmaeus_ (Taylor, 1936a); "C" probably
is _M. pygmaeus_; "D" is referable to _M. minimus_ (Taylor, 1940d) in
most characteristics, although the coloration is more nearly like that
of _M. lineatissimus_ (Taylor, 1941a), a larger species characterized by
a relatively long hind limb; "E" apparently is _M. imitator_ (Taylor,
1942b); "F" is _M. albolabris_ (Taylor, 1940d). Examination of series of
these frogs from other parts of Mexico shows a similar composition of
color variants. Of 78 specimens from the Rio Sarabia and the village of
Sarabia in Oaxaca (UMMZ 115428-37), 57 are "A," six are "D," three are
"E," and 12 are "F"; of 22 specimens from Teapa, Tabasco (UMMZ 113829),
11 are "A," five are "D," two are "E," and four are "F"; of 33 specimens
from Potrero Viejo, Veracruz (USNM 115447-58, 115461-71, 116840-2,
116864-70), ten are "A," 13 are "E," and ten are "F"; of 31 specimens
from La Esperanza, Chiapas (USNM 115477-9, 116827-39, 116849-63), 28 are
"A" and four are "F."
It is highly doubtful if these color variants are actually distinct
species. Goin (1950 and 1954) in his studies of inheritance of color
pattern in West Indian species of the genus _Eleutherodactylus_ has
shown that similar color pattern variants come from the same clutch of
eggs; furthermore, Goin has worked out the genetic ratios of certain of
these variants. Heathwole (_in litt._) obtained "normal" specimens and
individuals having a broad middorsal stripe ("C" in figure 9) from a
clutch of eggs of _Eleutherodactylus gollmeri_. The presence of a broad
middorsal yellow stripe is common in _Eleutherodactylus rugulosus_.
Perhaps the most interesting aspect of variability in color pattern in
Mexican eleutherodactylids is the parallelism between members of the
_Eleutherodactylus rhodopis_-group and some members of
_Microbatrachylus_. In the former group there are white-lipped
individuals (_Eleutherodactylus beatae_ Boulenger), individuals having a
unicolor reddish or yellowish dorsum (_E. dorsoconcolor_ Taylor), and
individuals having a dorsal pattern of irregular longitudinal brown and
cream-colored streaks (_E. venustus_ Guenther). In the humid forests of
southern Veracruz, northern Oaxaca, and Chiapas members of both groups
occur sympatrically. A proper understanding of the evolutionary
significance of these variants in the two groups, as well as proper
allocation of the presently recognized species, must await experimental
evidence based on studies of the inheritance of color pattern.
Nevertheless, at present it is apparent that certain characters,
especially the nature of the dermal folds and pustules, and the color
pattern, are of little taxonomic value in distinguishing "species" of
_Microbatrachylus_. The data derived from a study of the large series
from Arteaga, together with that from the other series examined,
suggests that _Microbatrachylus albolabris_, _imitator_, _minimus_, and
_pygmaeus_ are morphotypes of one species. Of these names, _pygmaeus_ is
the oldest. Consequently _Microbatrachylus pygmaeus_ has been used here
for the series from Arteaga.
Although _Microbatrachylus hobartsmithi_, a species distinguished from
all of the above by the presence of tubercles on the outer edge of the
tarsus, is known from Michoacan northward into Nayarit,
_Microbatrachylus pygmaeus_ previously has not been known north of
Guerrero, where it occurs in habitats similar to that in which it was
collected at Arteaga.
~Eleutherodactylus augusti cactorum~ Taylor
_Eleutherodactylus cactorum_ Taylor, Univ. Kansas Sci. Bull.,
25:391, July 10, 1939.--20 miles northwest of Tehuacan,
Puebla, Mexico.
_Eleutherodactylus augusti cactorum_, Zweifel, Amer. Mus.
Novitates, 1813:20, December 23, 1956.
Cheran; Coalcoman; Uruapan.
The few specimens indicate that this species occurs at moderate to high
elevations in the state. The specimens from Cheran and Uruapan were
obtained in pine forests; the specimen from Coalcoman was found on a
rocky hillside covered with dense forest and located about 100 meters
below the lower limits of the pine forest in the area. A specimen from
Rancho Reparto (elevation 1850 meters) on the west slope of Cerro
Barolosa was lost.
The specimen from Coalcoman (UMMZ 104728) is a juvenile having a
snout-vent length of 25.0 mm. In life it was tan above, mottled with
olive-green. The ventral surfaces were gray; the hind limbs were
distinctly barred with yellow and brown, and the lips were barred with
yellow and black.
~Eleutherodactylus occidentalis~ Taylor
_Eleutherodactylus occidentalis_ Taylor, Proc. Biol. Soc.
Washington, 54:91, July 31, 1941.--Hacienda El Florencio,
Zacatecas, Mexico.
Arteaga (2); Cascada Tzararacua; Coalcoman (2); 19 km. SW of
Coire (3); La Placita (7); Los Reyes; Ostula (4); Pomaro
(2).
The locality records for this species suggest that it is a member of a
group of animals, the distribution of which includes the western part of
the Mexican Plateau and the Pacific lowlands. In Michoacan this frog
has been collected in pine-oak forest at Cascada Tzararacua and at Los
Reyes, in arid scrub forest at Arteaga and Coalcoman, and in tropical
semi-deciduous forest on the lower Pacific slopes of the Sierra de
Coalcoman. On July 5, 1950, James Peters (1954:6) found calling males at
La Placita.
Most of the specimens are immature; four adult males have snout-vent
lengths of 30.9-33.0 (32.2) mm. In all specimens the first finger is
noticeably longer than the second; the inner metatarsal tubercle is
large, flat, and cream-colored, contrasting with the dark brown sole of
the foot. When the hind limbs are adpressed, the heels broadly overlap.
Characteristically, a dark line extends from the snout, through the eye,
above the tympanum, to a point above the insertion of the forelimb.
Usually there is a dark bar behind the tympanum, two dark brown bars
from the eye to the mouth and thence onto the lower jaw, and another
dark bar on the upper lip between the eye and nostril. One adult from
Arteaga, an adult and a juvenile from La Placita, and one juvenile each
from Coire, Ostula, and Pomaro, have the lower lip barred with dark
brown and white, and have a white stripe extending the length of the
upper lip. In life the dorsum varies from dark gray or olive-brown to
tan or reddish brown.
This species belongs to a group containing two other populations that
are currently recognized as species--_calcitrans_, known only from
Omiltemi, Guerrero, and _mexicanus_, reported from the mountains of
Oaxaca. Another apparently undescribed member of this group has been
collected in the mountains of northern Puebla. The locality records
indicate that the group inhabits the mountains on the periphery of the
Mexican Plateau, except in western Mexico, where _Eleutherodactylus
occidentalis_ extends to the Pacific lowlands.
~Eleutherodactylus rugulosus vocalis~ Taylor
_Eleutherodactylus vocalis_ Taylor, Univ. Kansas Sci. Bull.,
26:401, November 27, 1940.--Hacienda El Sabino, Michoacan,
Mexico.
Arteaga (10); El Sabino (8); Salitre de Estopilas (3);
Tumbiscatio (2); Tzitzio (2).
The distributional data on this frog in Michoacan indicate that it
inhabits riparian situations in arroyos and canyons in the lower slopes
of the Cordillera Volcanica and the Sierra de Coalcoman, where it has
been taken at elevations only below 1100 meters.
The dorsal color of living individuals from Arteaga varied from dark
gray and olive brown to tan and reddish brown. The iris was grayish
brown. In contrast, individuals from Agua del Obispo, Guerrero, had pale
golden eyes; specimens from Matias Romero, Oaxaca, had gold eyes heavily
flecked with gray; and individuals from Volcan San Martin, Veracruz, had
bronze eyes.
The use of the trinomial here is arbitrary. Frogs of the
_Eleutherodactylus rugulosus_ group in Mexico (_rugulosus_, _avocalis_,
and _vocalis_) exhibit only slight differences in size, proportions, and
coloration (Duellman, 1958c:6). Furthermore, the named populations are
allopatric. _Eleutherodactylus rugulosus vocalis_, as defined by
Duellman (_loc. cit._), occurs in the foothills of the Sierra Madre
Occidental and associated ranges from central Sinaloa southward into
Michoacan.
~Tomodactylus angustidigitorum~ Taylor
_Tomadactylus angustidigitorum_ Taylor, Univ. Kansas Sci.
Bull., 26:494, November 27, 1940.--Quiroga, Michoacan,
Mexico.
Angahuan (6); Apo; Carapan (21); 19 km. S of Carapan (13);
Cerro Tancitaro (12); Cheran; Corupu (14); Cuseno Station
(14); Opopeo (3); Paracho (11); Paricutin (2); Patzcuaro
(3); Quiroga (59); San Juan de Parangaricutiro (16);
Tancitaro (25); Uruapan (8); Zacapu (11).
This species is indigenous to the pine-oak forests on the southern rim
of the Mexican Plateau, and has been collected at elevations from 1500
to 2500 meters. Males have been observed to call from rocks, rock
fences, clumps of grass, and low bushes; the call is a single "peep." At
San Juan de Parangaricutiro numerous specimens were found in the daytime
beneath adobe bricks and lava on the volcanic ash derived from Volcan
Paricutin; at Paracho individuals were found by day beneath rocks in a
pine forest.
In most specimens the dorsum is dark reddish brown, and the prominent
inguinal glands are cream-color or pale orange (Pl. 3, Fig. 1). Of eight
individuals collected at Paracho, one was reddish brown, two were
pinkish tan, three were dark brown, and two were black.
~Tomodactylus fuscus~ Davis and Dixon
_Tomodactylus fuscus_ Davis and Dixon, Herpetologica,
11:157, July 15, 1955.--1.5 miles southeast of Huitzilac,
Morelos, Mexico.
Los Cantiles (2); 28 km. E of Morelia.
The range of this species includes the Sierra Ajusco in Mexico and
Morelos and thence westward to the Serrania Ucareo in Michoacan. The
specimen from 28 kilometers east of Morelia was found in an oak forest
on a steep hillside at an elevation of 2100 meters. One from Los
Cantiles was calling from a steep cliff at an elevation of 2200 meters
in pine-oak forest. This specimen (UMMZ 119156) in life had a pale
olive-brown dorsum with irregular dark brown mottling and transverse
bars on the limbs. The interorbital bar, the upper arms, and the tips of
the dorsal pustules were pale orange; the iris was pale grayish gold
(Pl. 3, Fig. 2).
~Tomodactylus nitidus nitidus~ (Peters)
_Liuperus nitidus_ Peters, Monats. Akad. Wiss. Berlin, p.
878, 1869.--Izucar de Matamoras, Puebla, Mexico.
_Tomodactylus amulae_ Guenther, Biologia Centrali-Americana,
Reptilia and Batrachia, p. 219, April, 1900.--Amula,
Guerrero, Mexico.
_Tomodactylus nitidus nitidus_, Dixon, Texas Jour. Sci.,
9:385, December, 1957.
Copuyo (15); Tuxpan (8); Tzitzio (11).
One specimen from Tzitzio (UMMZ 99155) was referred to _Tomodactylus
nitidus petersi_ by Dixon (1957:390). A re-examination of this specimen,
and examination of ten others from the same locality (UMMZ 121571)
reveals that the relatively small size of the tympanum and absence of
dense ventral spotting place these specimens closer to _T. nitidus
nitidus_ than to _T. nitidus petersi_.
The specimens from Tuxpan (UMMZ 114303-4) had in life a gray to olive
tan ground color with dark olive-green markings, bright yellow thighs
with olive-green transverse bands, yellowish tan shanks with olive-green
bars, yellow groin, white inguinal glands with black markings, grayish
white belly with scattered brownish black spots in some specimens, and a
deep golden iris (Pl. 4, Fig. 1). These specimens were found calling
from bushes in a rocky field at an elevation of 1800 meters. The call is
a high-pitched "pee-ee-eep."
~Tomodactylus nitidus orarius~ Dixon
_Tomodactylus nitidus orarius_ Dixon, Texas Jour. Sci.,
9:392, December, 1957.--4.5 miles southwest of Tecolapa,
Colima, Mexico.
La Placita (3); Pomaro.
These specimens, referred to _Tomodactylus petersi_ by Duellman
(1954b:5), were included in _T. nitidus orarius_ by Dixon (1957:392).
Color notes based on living individuals from Tecolapa, Colima (UMMZ
114312 and 116922), are: gray above mottled with brown; venter dirty
white; anterior and posterior surfaces of thighs bright yellow; iris
pale golden (Pl. 4, Fig. 2). The call is a soft "braa" usually followed
by three high notes: "braaa-eep-ee-eep." In Michoacan this subspecies
has been found only in the coastal region and the lower foothills of the
Sierra de Coalcoman, an area in which it replaces _Tomodactylus nitidus
petersi_. This is the only _Tomodactylus_ known to inhabit coastal
lowlands.
~Tomodactylus nitidus petersi~ Duellman
_Tomodactylus petersi_ Duellman, Occ. Pap. Mus. Zool. Univ.
Michigan, 560:5, October 22, 1954.--Coalcoman, Michoacan,
Mexico.
_Tomodactylus nitidus petersi_, Dixon, Texas Jour. Sci.,
9:390, December, 1957.
Aguililla; Apatzingan (8); Cascada Tzararacua: Charapendo
(5); Coalcoman (5); 18 km. E of Dos Aguas (6); El Sabino
(5); La Playa (2); Jiquilpan; Uruapan (2); Volcan Jorullo;
Zamora.
In life, specimens from Apatzingan (UMMZ 114308-9) varied in dorsal
color from grayish tan to pale brown; the dorsal markings were olive
green. The thighs and groin were yellowish orange; the iris was pale
golden, and the vocal sac was purplish gray (Pl. 5, Fig. 1).
Measurements for 13 adult males from the Tepalcatepec Valley are:
snout-vent length, 21.9-26.8 (24.3); tibia length, 8.4-9.9 (9.3); head
width, 7.2-9.2 (7.8); head length, 7.6-8.7 (8.2).
At Apatzingan and Charapendo in the Tepalcatepec Valley males were found
calling from rocks and bushes in open arid tropical scrub forest. The
call, a triple note "peep-ee-eep," is repeated once every 90 to 135
seconds. _Tomodactylus nitidus petersi_ probably ranges throughout the
Tepalcatepec Valley and surrounding foothills. Dixon (1957:392) referred
the specimens from Zamora, Jiquilpan, and Uruapan to this subspecies.
Uruapan is near the lower limits of the pine forest on the slopes of the
Cordillera Volcanica; Zamora and Jiquilpan are on a low part of the
Mexican Plateau southeast of Lago de Chapala.
~Tomodactylus rufescens~ Duellman and Dixon
_Tomodactylus rufescens_ Duellman and Dixon, Texas Jour.
Sci., 11:78, March, 1959.--Dos Aguas, Michoacan, Mexico.
Dos Aguas (14); 18 km. E of Dos Aguas (6).
Fourteen specimens from the pine-oak forests around Dos Aguas (UMMZ
118503-10, 121498-9) have reddish brown dorsal color and a narrow
cream-colored middorsal line (Pl. 5, Fig. 2). Twelve of these specimens
are adult males having snout-vent lengths of 20.7 to 24.6 (22.5) mm. One
female has a snout-vent length of 24.8 mm., and one juvenile has a
snout-vent length of 14.5 mm. Six specimens are from a region of mixture
of pine-oak forest and arid tropical scrub forest at 18 kilometers east
of Dos Aguas (UMMZ 121497, 121500). All are males having snout-vent
lengths of 18.0 to 22.6 (20.7) mm. The dorsum is tan marked with black;
the thighs are yellowish orange.
The specimens from 18 kilometers east of Dos Aguas were found on July
22, 1960, by Floyd L. Downs and John Winklemann, who collected calling
males of _Tomodactylus rufescens_ and _Tomodactylus nitidus petersi_ at
the same locality. Downs (_personal communication_) stated the call was
a single note. At Dos Aguas I heard _T. rufescens_ give two calls, one a
single "peep," the other a triple note--"pee-ee-eep."
In the higher parts of the Sierra de Coalcoman _Tomodactylus rufescens_
seems to fill the same niche as _T. angustidigitorum_ does in the
Cordillera Volcanica. At lower elevations in their respective mountain
ranges the species occur sympatrically with _T. nitidus petersi_.
~Diaglena reticulata~ Taylor
_Diaglena reticulata_ Taylor, Univ. Kansas Sci. Bull.,
28:60, May 15, 1942.--Cerro Arenal, Oaxaca, Mexico.
Nueva Italia (3); Ostula (7).
Until recently frogs of the genus _Diaglena_ were known only from a few
specimens from southern Sinaloa (_Diaglena spatulata_) and from the
Pacific lowlands of the Isthmus of Tehuantepec (_Diaglena reticulata_).
Peters (1955a) reported specimens from Ostula, Michoacan, and compared
these specimens with one _D. reticulata_ from Tehuantepec, Oaxaca, and
four _D. spatulata_ from Sinaloa. This comparison showed that the
specimens from Michoacan, although showing some minor differences from
_D. reticulata_, are closer to that species than to _D. spatulata_.
Subsequent to Peters' work, series of both species of _Diaglena_,
including additional specimens from Michoacan and from Colima, have been
collected, and a more qualified comparison is now possible.
In comparing specimens of _D. spatulata_ from southern Sinaloa (UMMZ
115322) with specimens of _D. reticulata_ from Tehuantepec, Oaxaca (UMMZ
115321), the differences noted by Taylor (1942c:60) were found to be
constant. But specimens from Ostula, Michoacan (UMMZ 104418), and five
individuals from Colima (TNHC 26379-83) were found to be intermediate in
certain characters. The skin of the dorsum in _D. reticulata_ is
granular; that in _D. spatulata_ is smooth. The skin in specimens from
Ostula and Colima is slightly granular. The dorsal ground color of _D.
reticulata_ is yellowish brown with dark reticulations; the dorsal
ground color of _D. spatulata_ is olive-green. Specimens from Ostula and
Colima most closely resemble those from Tehuantepec in coloration, but
the reticulations are more coarse, and the ground color has an
olive-green tint. _Diaglena reticulata_ also differs from _D.
spatulata_ in having a larger over-all size, slightly broader head, a
narrower interorbital distance, and a more pointed snout with a deeper
labial shelf (Table 4). The specimens from Ostula and Colima are
intermediate between _D. reticulata_ from Oaxaca and _D. spatulata_ from
Sinaloa in body proportions.