snout-vent length in males 84.6 mm, in females 104 mm; 2) skin on dorsum
in males bearing many moderately large, spinous tubercles; 3) skin on
flanks smooth; 4) web extending to middle of antepenultimate phalange on
inner edge of third finger; 5) dorsum brown usually with a large medial
dark brown blotch or, less frequently, several dark spots; narrow
middorsal yellow line present in some; 6) venter cream or tan with or
without small, irregular brown flecks; 7) lips brown with vertical cream
bar below eye in some, expanded into pale labial stripe posteriorly in
some females; 8) flanks tan or cream with or without small, irregular
brown spots; 9) dermal roofing bones of skull exostosed, casqued, and
co-ossified (in large adults); 10) dermal sphenethmoid present; 11)
nasals juxtaposed medially; 12) anteromedial margin of frontoparietals
at mid-level of orbit; 13) frontoparietal fontanelle covered; 14)
palatine serrate; 15) parasphenoid bearing odontoids; 16) zygomatic
ramus of squamosal usually articulating with maxillary arch; 17)
transverse processes of third presacral vertebra approximately equal in
width to sacral diapophyses; transverse processes of presacral vertebrae
3-8 subequal in width; 18) intermandibularis and submentalis muscles
connected; 19) supramandibular portion of interhyoideus extensively
developed; associated skin forming everted pouch.
The moderately rugose dorsum (in males), large size, extensive webbing
on the hand, and frontoparietal flanges in adults serve to distinguish
_taurinus_ from other members of the genus.
_Distribution._--The Amazon Basin, the upper Orinoco Basin, and the
Guianas. Most localities are below 500 m, but the species ascends the
lower Amazonian slopes of the Andes to elevations of about 1000 m (Fig.
11). A record from Caracas, Venezuela, and those from Provincia Carchi
and Provincia Esmeraldas, Ecuador, are considered to be erroneous. The
latter specimens were included in a collection sold to the University
of Illinois; contained in the collection are many common Amazonian
species unknown from the Pacific lowlands. 516 specimens from 151
localities.
[Illustration: FIG. 11. Distribution of _Osteocephalus taurinus_.]
_Remarks._--This widespread species is highly variable in size and
coloration. Striking differences in snout-vent length are evident in
series from various parts of the range. The smallest calling males
(CAS-SU 12351-6 from Rio Tapirape, Brasil) have snout-vent lengths of
46.5-60.3 (mean 53.3) mm, whereas the largest (FMNH 140254, KU 92243-6,
WCAB 9997, 10001, 10003-4 from Igarape Marmelo, Brasil) have snout-vent
lengths of 71.5-84.6 (mean 77.6) mm. Mean values of snout-vent lengths
of males from other localities are: Rio Pastaza drainage, Ecuador 73.8
mm, Surinam 67.7 mm, Rio Ucayali drainage, Peru 57.6 mm, and Guyana 55.5
mm. Although the difference between the smallest and largest adults is
highly significant, populations bridging the gap do exist. Furthermore,
the geographic arrangement of small versus large frogs is a confusing
mosaic. We have entertained the thought that we have included more than
one species in _taurinus_, but on the basis of preserved specimens we
are unable to detect consistent differences distinguishing two or more
taxa.
The coloration and pattern of _taurinus_ are so variable that no one
series of statements can describe samples drawn from the entire range of
species. We have been unable to determine geographic trends in color
pattern; instead the variation within a given sample can encompass the
variety known in most other samples. Two minor exceptions do exist. A
narrow middorsal light stripe is present in some individuals from
throughout the range, but striped specimens are most common in the upper
Amazon Basin. The absence of dorsal markings is uncommon in the entire
species, but it is most frequent in individuals from the Guianas. A few
individuals, such as KU 105230, have scattered white spots on the
dorsum.
The coloration of four males in life from Lago Agrio, Ecuador (KU
126652-5) was: "Dorsal ground color tan to dark brown with darker brown
markings. Flanks creamy tan to yellow with brown or black flecks or
mottling. Venter uniform creamy yellow or yellow with brown spots or
reticulations. Iris greenish yellow with radiating black streaks and a
median, horizontal reddish brown streak." (W. E. Duellman, field notes,
12 May 1969.) A female from Santa Cecilia, Ecuador (KU 123173), was:
"Dorsum mottled olive-green and tan. Flanks tan with brown spots. Belly
and throat creamy white, becoming tan posteriorly. Edge of upper jaw
olive-green." (W. E. Duellman, field notes, 16 June 1968.) Another
female from Santa Cecilia (KU 123175), was: "Brown dorsally with
cream-colored mottling. Transverse bars on legs darker brown with
cream-colored edges. Margin of upper lip creamy yellow. Anterior and
posterior surfaces of thighs tan. Flanks white with brown spots. Venter
creamy white. Iris greenish bronze with heavy radiating reticulations of
black." (W. E. Duellman, field notes, 22 July 1968.)
The tendency for females to have a labial stripe posteriorly and the
absence of dorsal tubercles in females has resulted in the
identification of many such specimens as _O. leprieurii_.
Ontogenetic change in coloration is slight in _taurinus_. Most juveniles
(less than 40 mm in snout-vent length) can be identified readily. There
is a tendency for the dorsal markings of juveniles to consist of several
small spots. Apparently with growth the spots usually coalesce, forming
a large median blotch, but some adults retain the juvenile pattern.
Cochran and Goin (1970:251) erroneously identified several juveniles
from Colombia as _Hyla palpebrogranulata_ Andersson.
=Osteocephalus verrucigerus= (Werner)
_Hyla verrucigera_ Werner, 1901:601 [Holotype.--ZMB 16589 from
"Ecuador"; Richard Haensch collector].
_Hyla riopastazae_ Andersson, 1945:72 [Holotype.--NHRM 1960 from
Banos, Rio Pastaza, Provincia Tungurahua, Ecuador; William
Clarke-MacIntyre collector].
_Hyla orcesi_ Funkhouser, 1956:78 [Holotype.--CAS-SU 13150 from Rio
Pacayacu, tributary of Rio Cotapino, Provincia Napo, Ecuador;
collector unknown].
_Osteocephalus orcesi_--Cochran and Goin, 1970:317.
_Osteocephalus verrucigerus_--Trueb and Duellman, 1970:601
[Synonymized _Hyla riopastazae_ Andersson, 1945, and _Hyla orcesi_
Funkhouser, 1956, with _Hyla verrucigera_ Werner, 1901].
_Justification of Synonymy._--Trueb and Duellman (1970:605) discussed
the assignment of the names in the synonymy of _O. verrucigerus_; only a
brief resume is given here.
The extant type of _Hyla verrucigera_ is a juvenile male having a
snout-vent length of 32.0 mm. The dorsum is smooth except for tubercles
on the eyelids; the skin is loose, and the body is soft. The specimen is
faded to a pale brown; indistinct dark spots are present on the back,
and transverse bars are evident on the limbs.
The holotype of _Hyla riopastazae_ is a gravid female having a
snout-vent length of 64.7 mm. The dorsum is smooth. The dorsal ground
color is pale brown with indistinct brown transverse bars on the limbs.
The throat, chest, and belly are cream with brown spots and mottling.
The holotype of _Hyla orcesi_ is an adult male having a snout-vent
length of 52.6 mm. The dorsum is heavily tuberculate. The dorsum is dark
brown with faint transverse bars on the forearms and feet; the ventral
surfaces are creamy brown.
Trueb and Duellman (1970) provided conclusive evidence that the types of
_H. verrucigera_, _riopastazae_, and _orcesi_ are a juvenile, adult
female, and adult male, respectively, of one species, the earliest
available name for which is _Hyla verrucigera_ Werner, 1901.
_Diagnosis._--1) Size moderate, sexual dimorphism evident; maximum
observed snout-vent length in males 54.3 mm, in females 65.8 mm; 2) skin
on dorsum in males bearing large, keratinized tubercles; 3) skin on
flanks smooth; 4) web extending to base of antepenultimate phalange on
inner edge of third finger; 5) dorsum uniformly dark brown or black,
with tan snout in females; 6) venter creamy white, heavily mottled with
black or dark brown, especially in females; 7) lips marked with pale tan
labial stripe and suborbital bar; 8) flanks dull reddish brown; 9)
dermal roofing bones of skull lacking exostosis; 10) dermal sphenethmoid
absent; 11) nasals widely separated medially; 12) anteromedial margin
of frontoparietals at anterior border of orbit; 13) frontoparietal
fontanelle covered; 14) palatine serrate; 15) parasphenoid bearing
odontoids; 16) zygomatic ramus of squamosal extending approximately
one-half of distance to maxillary arch; 17) transverse processes of
third presacral vertebra approximately equal in width to sacral
diapophyses; transverse processes of presacral vertebrae 3-8 subequal in
width; 18) intermandibularis and submentalis muscles connected; 19)
supramandibular portion of interhyoideus forming simple, tubular,
posterolateral extension; associated skin unmodified.
_Osteocephalus verrucigerus_ can be distinguished from other members of
the genus by its uniformly dark dorsum, heavily mottled venter, and
large, spinous tubercles on the dorsum in males.
_Distribution._--Lower Amazonian slopes (500-1840 m) of the Andes and on
the western fringe of the Amazon Basin in Ecuador and Peru; one locality
(Acevedo) in upper Rio Magdalena drainage in Colombia (Fig. 9). 40
specimens from 13 localities.
_Remarks._--In life the dorsum in males is dull olive-green; the groin,
anterior and posterior surfaces of the thighs, inner surfaces of limbs,
and upper arms are dark brown. The ventral surfaces of the limbs are
pinkish tan; the other ventral surfaces are pale creamy tan with reddish
brown spots. The suborbital spot is pale greenish tan, and the iris is
deep reddish brown. In females the dorsum is dull olive-brown; the
anterior part of the head is tan, and the suborbital spot is yellowish
tan. The groin and hidden surfaces of the limbs are dark reddish brown.
The ventral surfaces of the limbs are brown; the throat and chest are
creamy white, and the belly is reddish tan, both with dark brown
mottling.
Considerable ontogenetic change occurs in coloration. Juveniles are pale
above with a dark median dorsal blotch and dark transverse bars on the
limbs. The venter is white. The change consists principally of an
increase in dark pigment and subsequent obliteration of the juvenile
pattern.
Tadpoles of this species have moderately long tails with low fins,
robust bodies, two rows of labial papillae with median part of the upper
lip bare, and two upper and five lower rows of teeth. Trueb and Duellman
(1970) described the eggs, tadpoles, mating call, and variation in the
adults.
GENERIC RELATIONSHIPS
Among the 33 genera currently recognized in the family Hylidae, there
are two basic types of vocal sac structure (Duellman, 1970b), namely
the subgular type and the lateral type. Only four hylid genera, all
Neotropical lowland groups, are known to possess paired lateral vocal
sacs; these are _Osteocephalus_, _Argenteohyla_, _Phrynohyas_, and
_Trachycephalus_. The geographical distributions and morphological
characteristics of these four genera suggest that they are more closely
related to one another than with any other hylid genera.
Of the four genera, _Osteocephalus_ is the most generalized in
morphology, and, like _Phrynohyas_, has no specialized habits.
_Osteocephalus_ and _Argenteohyla_ are similarly distinguished from
_Phrynohyas_ and _Trachycephalus_ on the basis of vocal sac structure.
The vocal sacs of _Osteocephalus_ and _Argenteohyla_ are posterior and
protrude posterolateral to the angles of the jaws when they are
inflated, whereas those of _Phrynohyas_ and _Trachycephalus_ are more
lateral and protrude posterior to the angles of the jaws when inflated.
Although _Osteocephalus_ and _Argenteohyla_ have similar vocal sac
structure, they are obviously distinct. The monotypic _Argenteohyla_ is
a rather specialized, semifossorial frog (Trueb, 1970b), characterized
by smooth skin, moderate-sized digital discs, and a large inner
metatarsal tubercle. The general architecture of the skull is not unlike
that of _Osteocephalus_; the skulls of both are well roofed, broader
than long, and characterized by posterolaterally oriented parasphenoid
alae. _Argenteohyla_ bears small, slightly curved prevomerine
dentigerous processes in contrast to the large, angular processes of
_Osteocephalus_. The skull of _Argenteohyla_ shows specializations,
apparently adaptations to its semifossorial mode of existence, which
further distinguish the genus from _Osteocephalus_. In comparison with
_Osteocephalus_, the cranium of _Argenteohyla_ is slightly depressed
anteriorly, the roofing bones extensively casqued, and the palatines
robust.
Osteologically, _Osteocephalus_ more closely resembles _Phrynohyas_ than
either of the other two genera, but _Osteocephalus_ and _Phrynohyas_ are
clearly distinct on the basis of their respective vocal sac structure.
Like _Osteocephalus_, skulls of the members of the genus _Phrynohyas_
are broader than long, have extensive dermal roofing bones, and have
posterolaterally oriented parasphenoid alae. In contrast to
_Osteocephalus_, the dentigerous processes of the prevomers are curved,
rather than angular in _Phrynohyas_. Furthermore, the latter genus is
singularly distinguished from _Osteocephalus_, _Argenteohyla_, and
_Trachycephalus_ by having extensively developed parotoid glands that
produce a viscous, milky volatile secretion.
_Trachycephalus_ is the most readily identifiable of the four genera
under discussion. Members of this genus are large frogs with heavily
casqued and co-ossified skulls (Trueb, 1970a). The dermal roofing bones
bear ornate and characteristic patterns of sculpturing. The medial ramus
of the pterygoid does not articulate with the otic capsule, and the
parasphenoid alae are laterally, rather than posterolaterally, oriented.
A dermal sphenethmoid is present, and the parasphenoid bears odontoids.
The basic structure of the skull has many characters in common with both
_Osteocephalus_ and _Phrynohyas_. The obvious modifications of dermal
roofing bones and of palatal and suspensory elements seem to be
specializations adapting members of the genus _Trachycephalus_ to their
peculiar phragmotic habits. The vocal sac structure of _Trachycephalus_
is like that of _Phrynohyas_ and therefore further distinguishes it from
_Osteocephalus_.
Morphologically, _Osteocephalus_ seems to be sufficiently diverse and
generalized so as to represent a modern derivative of an ancestral type
which might have given rise to _Phrynohyas_, _Trachycephalus_, and
_Argenteohyla_. The specialized vocal sac structure in _Phrynohyas_ and
_Trachycephalus_ suggests that these two genera may be rather closely
allied and represent a single phyletic line from an ancestral stock
similar to _Osteocephalus_. _Argenteohyla_ is quite distinct from
_Phrynohyas_ and _Trachycephalus_ and apparently represents a distinct
phyletic line from the ancestral stock.
OCCURRENCE OF _OSTEOCEPHALUS_ IN AMAZONIAN ECUADOR
All of our observations on members of this genus have been made at four
localities: 1) Santa Cecilia at an elevation of 340 meters on the Rio
Aguarico, a tributary of the Rio Napo, 2) Lago Agrio, 330 meters, about
14 kilometers east of Santa Cecilia, 3) Puerto Libre, 570 meters, on the
Rio Aguarico just east of its formation by the confluence of the Rio
Cofanes and Rio Chingua, and 4) south slope of the Cordillera del Due,
above the Rio Coca, 1150 meters. _Osteocephalus leprieurii_ was found at
all four localities, and _buckleyi_ was found at all but the last;
_taurinus_ was found at Santa Cecilia and Lago Agrio, and _verrucigerus_
was found only in the Cordillera del Due. Our data are based on
collections of 113 frogs and three lots of tadpoles, as well as
observations on calling sites and young. The observations are summarized
by species, as follows:
_Osteocephalus buckleyi._--No breeding activity was observed. Males were
found only at night in March, June, and July. One was perched on a
_Heliconia_ leaf in a swamp at Puerto Libre, and two were on bushes in
the forest at Santa Cecilia. A gravid female was found on a recently
felled tree at Lago Agrio on the night of 12 May 1969.
_Osteocephalus leprieurii._--Males were heard calling sporadically at
Puerto Libre in July 1968, and at Santa Cecilia in May 1969. A small
chorus was found on the night of 12 May 1969 at Lago Agrio, where the
frogs were perched on branches of fallen trees over a temporary pool.
The call is a soft rattling chuckle. In late April and May many gravid
females and males with well-developed nuptial excrescences were obtained
from trees as they were felled at Lago Agrio. The reproductive condition
of the frogs indicates that they probably breed in May. One individual
called nearly every night from a large tree at Puerto Libre between 4-17
July 1968. The tree was felled on the latter date, but no frog was
found. Two nights later apparently the same individual called from a
bromeliad at a height of about 10 m on a large bamboo adjacent to the
felled tree; the frog was collected when the bamboo was cut down.
Throughout the rainy months that we have worked in Ecuador
(April-August) we have found occasional individuals perched on bushes or
low trees at night. Large numbers of adults were observed only during a
clearing operation which resulted in the felling of many large trees.
Thus, it seems likely that _leprieurii_ is a tree-top inhabitant. A
partially digested adult male was removed from the stomach of a
_Hemiphractus proboscideus_.
At Santa Cecilia many recently metamorphosed young and juveniles were
found in June and July 1968. Most of these were on low bushes or herbs
in swamp forest at night; some were found in unfolded _Heliconia_ leaves
by day, and one was observed on the forest floor by day. Snout-vent
lengths of 18 specimens are 12.3-17.0 (mean 15.1) mm. The smaller frogs
were recently metamorphosed as evidenced by the melanophore deposits
above the vent. The coloration of the young is strikingly different from
that of the adults (see account of _O. leprieurii_), so the association
of the young and adults was not made until individuals with intermediate
patterns were obtained at Lago Agrio in May 1969. Probably juveniles
obtained in June and July are the offspring of an April or May breeding.
We have been unable to associate tadpoles with this species.
_Osteocephalus taurinus._--A small chorus occurred at Lago Agrio on 12
May 1969. Males were calling from the ground adjacent to a small pool
amidst recently felled trees. The males were very wary and, when
approached, jumped onto limbs and ran up branches; this behavior was
noted by Bokermann (1964). The call consists of a series of low-pitched,
short notes--like a slow trill--four to six notes per call group. Call
groups are repeated two, three, or four times followed by a lapse of
several minutes. Although no amplectant pairs were found, several gravid
females were collected at Lago Agrio in May, so it can be safely assumed
that the species breeds in May. From April through July occasional
individuals were observed on bushes and trees at night. During clearing
operations at Lago Agrio several individuals were obtained from the tops
of trees as they were felled.
_Osteocephalus verrucigerus._--Observations were made in a broad,
shallow ravine, in which there was a small stream. On 2-4 August 1968,
males were observed calling from low bushes and rocks at the edge of a
quiet pool in the stream. The call consists of a series of well-pulsed,
low-pitched, guttural notes produced at the rate of 5-10 per minute. One
amplectant pair was found at the base of a bush adjacent to the pool on
3 August. Another female was found on a branch of a tree 2 m above the
ground and 10 m from the stream. Tadpoles of this species were found in
the quiet silt-bottomed pool.
SPECIMENS EXAMINED
The localities for each of the specimens examined are given in the
following paragraphs. The arrangement of the data is as follows:
alphabetically by country, state (department or province), and locality;
alphabetically by the first letter in the abbreviations for the museums,
and numerically after each museum abbreviation. Specimens lacking
precise locality data are listed first in the most restricted political
unit possible; localities which have not been found on maps or the
positions of which are not known to us are given in quotation marks.
Where more than one specimen is included under one museum number, the
number of specimens is given in parentheses after the museum number.
Unless noted otherwise, all specimens are alcoholics.
_Osteocephalus buckleyi_
BOLIVIA: _El Beni_: Ivon, BMNH 1967.2070-1. _Santa Cruz_:
Buenavista, CM 4333, 4339, UMMZ 66563-5.
BRASIL: _Amapa_: No specific locality, WCAB 13284.
COLOMBIA: _Amazonas_: Rio Guacaya, USNM 152759. _Huila_: Acevedo,
Rio Suaza, FMNH 69702. _Narino_: Rumiyacu, FMNH 54756. _Meta_:
Rio Guejar, Campamento La Macarena, USNM 152199.
ECUADOR: No specific locality, NHMW 6209, WCAB 35499. _Chimborazo_:
Pallatanga, BMNH 1947.2.13.46; Santiago, FMNH 42529.
_Morona-Santiago_: "Rio Santiago" (= Rio Zamora), MIZS 2950.
_Napo_: Lago Agrio, KU 126646; Puerto Libre, Rio Aguarico, KU
123172; Santa Cecilia, AUM 8138, KU 105208-9, 109506, 123171.
_Pastaza_: Alpayacu, BMNH 1912.11.1.64; Canelos, BMNH
1947.2.13.40-1, 1947.2.13.43-5; Colonia Mena, Rio Conambo, ZSM
33/1962; Don Tomas, USNM 166014; Guache, Rio Pastaza, AMNH 79986;
Rio Bobonaza, USNM 166005; Rio Capahuari, USNM 166554; Rio Conambo
at Rio Shiona-yacu, USNM 166018; Rio Copataza, upper Rio Pastaza,
USNM 166007-13; Rio Pastaza, NHRM 1946; Rio Pucyacu, USNM 165997
(skeleton), 165998-6001; Rio Rutuno, USNM 166006; Rio Villano,
USNM 166002-4; Sarayacu, BMNH 1947.2.13.36-9, MCZ 26090, ZMB 10166.
GUYANA: _Mazaruni-Potaro_: Kartabo, AMNH 70971; Membaru River,
upper Mazaruni River, UMMZ 85168; Oko Mountains, FMNH 26722-3.
_North West_: Amakura River, Haulover, UMMZ 83558-9. _Rupununi_:
Marudi River, AMNH 46233; Shudi-kar-wau, AMNH 49252. _West
Demerara_: Dunoon, UMMZ 52449, 52508.
PERU: _Junin_: Chanchamayo, BMNH 1911.12.13.79-80. _Loreto_:
Andoas, AMNH 79984-5; Cashiboya, AMNH 43454; San Antonio, Rio
Itaya, AMNH 43218. _Puno_: Yahuaramayo, BMNH 1913.2.25.7.
SURINAM: _Suriname_: Powakka, CM 44217.
SOUTH AMERICA: No specific locality, NHMW 6208.
_Osteocephalus leprieurii_
BRASIL: _Acre_: Tarauaca, FMNH 83247. _Amazonas_: Rio Javari,
Benjamin Constant, CAS-SU 12620; Rio Uaupes, north of Rio Japu,
NHMG 489.
COLOMBIA: _Amazonas_: Gino-goje, lower Rio Apoporis, MCZ 28038,
28040-2, 28044, USNM 152136-8.
ECUADOR: No specific locality, WCAB 35452-3; "Napo-Pastaza," USNM
166571. _Napo_: Avila, UMMZ 92093; south slope Cordillera del Due,
KU 123170; Lago Agrio, KU 125961-2 (skeletons), 126611-44, UMMZ
129326 (2); Limon Cocha, Rio Napo, KU 99210-6, UIMNH 63087-9,
63098, 63106-9, 63118-9, 64802-4, 64858, 87998-9, 88001-30,
88437-8, 88580, 88604-5, 89852-97, 89999-90000; Loreto, CAS-SU
11439, WCAB 36526; Puerto Libre, Rio Aguarico, KU 123190-1; Puerto
Napo, UIMNH 55818-20; Rio Cotapino, UMMZ 92094; Rio Napo, UMMZ
92078; Santa Cecilia, AUM 8099, 8102, 8113-5, 8127-9, 8131, 8137,
8139-46, 8148, KU 105210-20, 109509-11, 111971, 122964-87, 123169,
126645. _Pastaza_: Canelos, BMNH 1947.2.13.42, KU 120915; Rio
Alpayacu, UMMZ 92079; Rio Arajuno, USNM 166560-2, WCAB 40176; Rio
Oglan, USNM 16655203, 166558; Rio Rutuno, USNM 166559; Rio
Shilcayacu, below Puyo, USNM 166557; Rio Villano, USNM 166551.
FRENCH GUIANA: No specific locality, MNHN 4629. _Inini_: Lunier
River, MNHN 98/217.
GUYANA: _Mazaruni-Potaro_: Kartabo, AMNH 70967-8, 70972, 70976.
_Rupununi_: Shudi-kar-wau, AMNH 49255. _West Demerara_: Demerara
Falls, BMNH 72.10.16.23, 72.10.16.37-8.
PERU: _Loreto_: Estiron, Rio Ampiyacu, MZUSP 31033-4; Pebas, CAS-SU
3158, 3160; Roaboya, AMNH 43064.
SURINAM: No specific locality, MCZ 2036, RMNH 11468. _Marowijne_:
Camp 3, RMNH 13045-6; Wane Creek North, RMNH 11469-70. _Saramacca_:
Right Coppename River, RMNH 11467.
_Osteocephalus pearsoni_
BOLIVIA: _El Beni_: upper Rio Beni, below mouth of Rio Mapiri, MCZ
15565, UMMZ 57548, 67464-5; Rurrenbaque, UMMZ 57533.