Nototriton barbouri ( Schmidt, 1936 )

Townsend, Josiah H., 2016, Taxonomic revision of the moss salamander Nototriton barbouri (Schmidt) (Caudata: Plethodontidae), with description of two new species from the Cordillera Nombre de Dios, Honduras, Zootaxa 4196 (4), pp. 511-528 : 516-520

Nototriton barbouri ( Schmidt, 1936)

Yoro Moss Salamander Figure 4

Oedipus barbouri Schmidt (1936: 43) .

? Pseudoeurycea barbouri: Taylor (1944: 209) .

Chiropterotriton barbouri: Meyer (1969: 106) .

Nototriton barbouri : Wake & Elias (1983: 11).

Nototriton (Bryotriton) barbouri: Dubois and Raffaëlli (2012: 141) .

Holotype. MCZ 21247, an adult male from Portillo Grande , Departamento de Yoro, Honduras, 1,524 m elevation, collected by R.E. Stadelman on 9 May 1934.

Paratopotypes. All females from the vicinity of the holotype, 1,524–1,828 m elevation; MCZ 21248–50, FMNH 21866–67.

Genetic reference specimen. UF 156538 ( Fig. 4 ), an adult female from Montaña Macuzal (15.079455°N, 87.352985° W; Fig. 2 ), 1,760 m elevation, above El Panal to the west of Yorito , Departamento de Yoro, Honduras; collected 9 April 2008, by J.M. Butler, L. Ketzler, J. Slapcinsky, N.M. Stewart, J.H. Townsend, and L.D. Wilson; original field number JHT 2420; GenBank accession numbers GU971733 (16S) , GU971734 (cyt b), JN377401 (COI).

Referred specimens. 15; ANSP 28981–85, 28200 , Montaña Macuzal S of Pueblo Viejo ; USNM 339700–08 , from the eastern slope of Pico Pijol (15.175°N, 87.558°W), 1,920 m elevation, Parque Nacional Pico Pijol , Departamento de Yoro, Honduras .

Diagnosis. A member of the genus Nototriton diagnosed by possessing 13 costal grooves (>16 costal grooves in Oedipina ), the presence of a sublingual fold and hands and feet longer than broad (sublingual fold absent and hands and feet broader than long in Bolitoglossa ), and small nares (NL/SL 0.006–0.012; 0.017–0.029 NL/SL in Cryptotriton and Dendrotriton ). Phylogenetic analysis support inclusion of N. barbouri in the subgenus Bryotriton, for which all nominal taxa are represented by molecular data ( Fig. 1 ). Notoriton barbouri can be distinguished from the other constituent species of Bryotriton as follows: from N. oreadorum in having relatively more narrow hind feet (HFW/SVL 0.034–0.052, versus 0.056–0.057 in N. oreadorum ); from N. brodiei in having a relatively shorter tail (TL/SVL 0.802–1.165, versus 1.420–1.440 in N. brodiei ), relatively longer limbs (FLL/SVL 0.156– 0.204 and HLL/SVL 0.185–0.231, versus 0.148–0.151 and 0.166–0.180 in N. brodiei ), and fewer maxillary teeth (41–53, versus 60–62 in N. brodiei ); from N. limnospectator in having a relatively broader head (HW/SVL 0.119– 0.141, versus 0.111–0.118 in N. limnospectator ) and a higher average number of costal grooves between adpressed limbs (6.2 [± 0.9], versus 4.4 [±0.8] in N. limnospectator ); from N. mime in having relatively shorter front limbs (FLL/SVL 0.156–0.204, versus 0.195–0.246 in N. mime ) and a higher average number of costal grooves between adpressed limbs (6.2 [± 0.9], versus 4.8 [±0.4] in N. mime ); from N. nelsoni in having a relatively shorter tail (TL/ SVL 0.802–1.165, versus 1.144–1.532 in N. nelsoni ) and a higher average number of costal grooves between adpressed limbs (6.2 [± 0.9], versus 4.9 [±1.4] in N. nelsoni ); from N. picucha in having a relatively more narrow head (HW/SVL 0.119–0.141, versus 0.140–0.148 in N. picucha ) and a higher average number of costal grooves between adpressed limbs (6.2 [± 0.9], versus 4.0 [±0.7] in N. picucha ); from N. stuarti in having a greater number of maxillary teeth (41–53, versus 36 in N. stuarti ), a relatively shorter tail (TL/SVL 0.802–1.165, versus 1.264 in N. stuarti ), and relatively longer hind limbs (HLL/SVL 0.185–0.231, versus 0.178 in N. stuarti ); and from N. tomamorum in having well-developed digits (syndactylous feet in N. tomamorum ), a greater number of maxillary and vomerine teeth (41–53 maxillaries and 12–23 vomerines, versus 26 maxillaries and 11 vomerines in N. tomamorum ) and smaller nares (NL/SL 0.006–0.012, versus 0.018 in N. tomamorum ). A single species of the subgenus Nototriton , N. saslaya , also occurs in the Chortís Highlands, and differs from N. barbouri in having fewer maxillary and vomerine teeth (17–22 maxillaries and 3–11 vomerines, versus 41–53 maxillaries and 12–23 vomerines in N. barbouri ). Table 3 provides a comparative summary of morphological variation among the species of Nototriton from the Chortís Highlands.

Nototriton barbouri is further distinguished from all other species of Nototriton in the Chortís Highlands by model-corrected genetic distances ( Table 2 ), being 1.3–1.9% (16S) and 7.8–8.5% (cyt b) divergent from its closest relative, N. limnospectator , 3.3% (16S) and 11.7–12% (cyt b) divergent from samples from RVS Texíguat, and 3.3% (16S) and 11.4% (cyt b) divergent from samples from PN Pico Bonito.

McCranie & Wilson (2002) provided a description of the coloration in life of an adult female (USNM 339700), as well as morphological data for two males and 13 females of N. barbouri (as “southern Yoro” populations; McCranie & Wilson 2002: 136–137).

Geographic and ecological distribution. I consider this taxon to be restricted to the Sierra de Sulaco in southwestern Departamento de Yoro, where it is known to occur from 1,520–1,920 m elevation in the Lower Montane Wet Forest formation ( Holdridge 1967) in Broadleaf Cloud Forest ( Townsend 2014). This species is known from two localities: the vicinity of the type locality in an unprotected patch of remant cloud forest on the top of Montaña Macuzal (maximum elevation approximately 1,945 m) at the eastern terminus of the Sierra de Sulaco; and in the cloud forests of PN Pico Pijol (maximum elevation approximately 2,282 m) at the western end of the Sierra de Sulaco. These localities are separated by approximated 26 airline km, and represent the highest portions of the Sierra de Sulaco, with the lowest intervening point on the ridge formation connecting these two areas being> 1,180 m elevation ( Fig. 3 ).

Natural history. This species has been found utilizing daytime refugia inside bromeliads ( Schmidt 1936), rotten logs and abandoned hummingbird nests ( McCranie & Wilson 2002), and in leaf litter packed at the base of a large karst outcrop. McCranie & Wilson (1992) reported finding this species actively foraging at night on low limbs and branches of small trees and shrubs. McCranie & Wilson (1992) provided a detailed account of reproductive behavior in N. barbouri from PN Pico Pijol, in which they found a clutch of five eggs without an attending adult and a second clutch with a single female present inside of an abandoned hummingbird nest suspended from limbs approximately 2 m above the ground. The clutch was removed from the nest along with the attending female, and the nest and female were subsequently placed back on the tree with a cloth collecting bag tied around the nest. The next morning, the female had deposited an additional six eggs, and subsequent examination of the nest revealed four other clutches of eggs in various stages of development ( McCranie & Wilson 1992). Clutch size ranged from 5– 19 eggs, with average egg capsule diameter ranging from 4.7–5.2 mm for the six clutches; no egg guarding behavior was observed ( McCranie & Wilson 1992).

Remarks. A single specimen (AMNH 54949) from a locality near Lago de Yojoa (“El Volcán,” north slopes of Cerro Azul Meámbar, 860 m elevation) was assigned to this taxon ( McCranie & Wilson 2002: 576). Nototriton limnospectator was recently confirmed from localities in the immediate vicinity of the “El Volcán” locality in Parque Nacional Cerro Azul Meámbar ( Townsend et al. 2011b), and I consider AMNH 54949 to be representative of N. limnospectator , and not a disjunct lower elevation locality for N. barbouri sensu stricto. Photographs of the genetic reference specimen ( Fig. 4 ; UF 156538) appear in Köhler (2011: 76) and Rafaëlli (2013: 368).

Description of unassigned populations from the Cordillera Nombre de Dios. Restriction of the taxon Nototriton barbouri to populations inhabiting the highlands of southern Yoro (Montaña de Pijol and Montaña de Macuzal) leaves two allopatric lineages from the Cordillera Nombre de Dios without assignment to an existing taxon. With no referable name present in the synonymy of N. barbouri for either of these populations, putatively endemic to highland forests in the central and western portions of the Cordillera Nombre de Dios in Parque Nacional Pico Bonito and Refugio de Vida Silvestre Texiguat ( Figure 4 ), I herein describe these two cryptic lineages as new taxa. Together, they form a clade with N. brodiei , a species restricted to the Sierra de Omoa and Sierra de Caral in northwestern Honduras and adjacent Guatemala, and N. stuarti , a species endemic to the Montañas del Mico in eastern Guatemala ( Fig. 1 ).

Figures

FIGURE 1. Bayesian phylogram of the genus Nototriton showing phylogenetic relationships based on analysis of a concatenated and partitioned dataset of fragments of the mitochondrial genes 16 S, cyt b, and COI, with Bayesian posterior probabilities and bootstrap support values from a complimentary maximum likelihood analysis shown above supported branches.

FIGURE 2. Views of Montaña Macuzal from the northeast (A) and southwest (B) sides, showing the limited extent of remaining forest along the upper reaches of the highest ridge.

FIGURE 3. Distribution of Nototriton in north-central Honduras; red diamond = Montaña Macuzal, type locality of N. barbouri; black diamond = N. barbouri (PN Pico Pijol); circles = N. nelsoni, RVS Texíguat (red = type locality); squares = N. oreadorum, PN Pico Bonito (red = type locality); crosses = N. lignicola; triangle = N. mime; inverted triangle = N. picucha.

FIGURE 4. Adult female of Nototriton barbouri (UF 156538) from the vicinity of the type locality at Montaña Macuzal, 1,760 m elevation, Departamento de Yoro, Honduras.

Tables

TABLE 3. Morphological and morphometric comparison of species of Nototriton the Chortís Highlands; see Material and Methods section for explanation of abbreviations; SVL is given in mm. Comparative morphological data for N. brodiei and N. stuarti are from Campbell & Smith (1998), Wake & Campbell (2000), respectively.

Taxon	N =	SVL	HL/SVL	HW/SVL	TL/SVL
N. nelsoni n. sp.	4	28.1– 34.6	0.185– 0.226	0.127– 0.142	1.144– 1.532
N. oreadorum n. sp.	2	33.7– 35.1	0.178– 0.182	0.117– 0.122	1.111– 1.169
N. barbouri	10	30.2–39.9	0.175– 0.211	0.119–0.141	0.802–1.165
N. brodiei	3	33.2–34.5		0.120	1.420–1.440
N. lignicola	13	28.6–36.8	0.175– 0.213	0.116–0.147	0.651–1.203
N. limnospectator	12	24.5–38.2	0.164– 0.216	0.111–0.118	1.027–1.147
N. mime	4	20.5–29.8	0.182– 0.020	0.121–0.138	0.698–1.117
N. picucha	3	25.7–27.9	0.197– 0.198	0.140–0.148	1.143
N. saslaya	2	28.1–34.6		0.133–0.155	0.883–1.255
N. stuarti	1	32.6		0.138	1.264
N. tomamorum	1	26.9		0.145	0.911
continued.
Taxon	HLL/SVL	FLL/SVL	HFW/SVL	NL/SVL	MT VT
N. nelsoni n. sp.	0.203– 0.223	0.173– 0.198	0.044– 0.058	0.006– 0.010	38–53 17–24
N. oreadorum n. sp.	0.190– 0.222	0.158– 0.178	0.056– 0.057	0.006– 0.009	47–52 20–22
N. barbouri	0.185–0.231	0.156–0.204	0.034–0.052	0.006–0.012	41–53 12–23
N. brodiei	0.166–0.180	0.148–0.151	0.040–0.060	0.004–0.005	60–62 23–24
N. lignicola	0.158–0.215	0.139–0.192	0.035–0.046	0.003–0.007	32–53 15–22
N. limnospectator	0.183–0.227	0.165–0.188	0.035–0.061	0.003– 0.009	39–55 16–24
N. mime	0.224–0.254	0.195–0.246	0.044–0.050	0.007–0.010	27–46 12–24
N. picucha	0.204–0.218	0.179–0.191	0.042–0.043	0.007–0.008	41 16–19
N. saslaya	0.217–0.244	0.194–0.210	0.075–0.091	0.002–0.003	17–22 3–11
N. stuarti	0.178	0.172	0.049	0.012	36 20
N. tomamorum	0.197	0.160	0.037	0.018	26 11