Amphibia: Caudata: Plethodontidae: Nototriton) from the Montaiias Del Mico of Guatemala
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1 November 2000 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 113(3):815-819. 2000. A new species of diminutive salamander (Amphibia: Caudata: Plethodontidae: Nototriton) from the Montaiias del Mico of Guatemala David B. Wake and Jonathan A. Campbell (DB W) Museum of Vertebrate Zoology and Department of Integrative Biology, University of California, Berkeley, California 94720-3160, U.S.A.; (JAC) Department of Biology, University of Texas at Arlington, Arlington, Texas 76019-0498, U.S.A. Abstract.-Nototriton stuarti, a new species of salamander belonging to the tribe Bolitoglossini, family Plethodontidae, is described from the Montaiias del Mico, north of the Motagua Valley, in eastern Guatemala. The species is geo- graphically isolated from congeneric species, and represents the northwestern- most record for the genus, which ranges from eastern Guatemala to central Costa Rica. This poorly known species is morphologically most similar to Nototriton barbouri, which occurs in northwestern Honduras. Homoplastic evolution has produced di- Chiropterotriton and all endemic to this minutive, attenuate salamanders in many area, are also diminutive. To the south and different clades of plethodontid salaman- east of the Isthmus of Tehuantepec, extend- ders in Middle America. These tiny species ing as far as central Costa Rica, diminutive are difficult taxonomically and it has taken species are found in one well-supported much effort to recognize that there are clade (Garcia-Paris & Wake 2000). This many more species than have been named clade includes the genera Cryptotriton, throughout this region. New discoveries Dendrotriton, Nototriton and Oedipina, and and new data have led to taxonomic revi- possibly the poorly known Bradytriton sions that are gradually identifying mono- Wake & Elias 1983 (G. Parra-Olea, pers. phyletic groups, and as this process goes comm.). Oedipina Keferstein 1868 includes forward the generic level taxonomy has some diminutive species, but also some changed repeatedly. This is especially true large and relatively robust, although elon- of the genus Chiropterotriton Taylor, 1944, gated, species. The phylogenetic diversity which as formerly recognized ranged from of this clade is now understood, at least in northeastern Mexico to Costa Rica, but broad outlines. Oedipina and Nototriton ap- which was not monophyletic and now has pear to be the most deeply nested groups been fragmented into four genera (Chirop- (Garcia-Pm’s & Wake 2000), and they are terotriton, Cryptotriton Garcia-Paris & also the most southerly in distribution, ex- Wake, 2000, Dendrotriton Wake & Elias, tending to Ecuador (moderately large spe- 1983, and Nototriton Wake & Elias, 1983). cies of Oedipina occur in South America, At present diminutive plethodontids found Brame & Wake 1963) and central Costa to the north and west of the Isthmus of Te- Rica, respectively. Both occur as far north huantepec are assigned to Chiropterotriton as Guatemala, and Oedipina ranges into (Darda, 1994), but, in addition, all members Chiapas, Mexico. Cryptotriton and Dendro- of the genera Pawimolge Taylor, 1944, Li- triton range from Honduras into Mexico, neatriton Tanner, 1950, and especially the former possibly even north and west of Thorius Cope, 1869, none closely related to the Isthmus of Tehuantepec [the species 816 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Pacific Ocean - 14‘ i - 8’ 92‘ 100 km Fig. 1. Distribution of certain taxa discussed in the text from southern Mexico, Guatemala, and Honduras, showing the location of the type-locality of Nototriton siuarti. The map shows the location and degree of isolation of morphologically and ecologically similar, diminutive salamanders that have proven to represent several dif- ferent smaller clades within a larger clade. Shaded areas on map represent regions with elevations in excess of 2000 m. Cryptotriton adelos (Papenfuss & Wake, clades in different analyses, but the consen- 1987) from northern Oaxaca is tentatively sus view (allozymes plus mtDNA sequenc- assigned to this genus], whereas Bradytri- es) is that it is most closely related to the ton is known only from its type-locality in other Costa Rican species. The second western Guatemala (Fig. 1). clade includes one species described long As revised by Garcia-Paris & Wake ago but still little known biologically, No- (2000), Nototriton includes 11 nominal spe- totriton barbouri (Schmidt, 1936a), from cies, all diminutive (none exceeds 40 mm, northwestern Honduras, and three recently snout-vent length), long-tailed forms that described taxa, Nototriton lignicola Mc- have small to moderately enlarged nostrils. Cranie & Wilson, 1997 and Nototriton lim- Based on molecular data, the genus itself is nospectator McCranie, Wilson, & Polisar, well supported as a monophyletic group, 1998, the first from north-central and the and there are two well-supported internal second from western Honduras, and Noto- clades, one including Costa Rican species, triton brodiei Campbell & Smith, 1998, and the other including species from Hon- from extreme eastern Guatemala (Fig. 1). A duras and Guatemala. The species Nototri- single specimen representing another spe- ton richardi (Taylor, 1949) from Costa Rica cies belonging to this clade was collected (and presumably its Costa Rican sister spe- in the Montaiias del Mico in eastern Gua- cies Nototriton tapanti Good & Wake, temala and at first confused with small 1993) is basal to one or the other of these specimens of sympatric Oedipina elongata VOLUME 113, NUMBER 3 817 (Schmidt, 1936b). Fewer than 50 specimens adult male) of moderate length. The naso- are available for the second clade, but ear- labial protuberances are modest in size and lier taxonomic judgements based on mor- barely protrude below the upper lip. The phology (McCranie & Wilson 1997, Camp- narrow head has eyes of moderate size that bell & Smith 1998, McCranie et al. 1998) are only slightly protuberant, barely extend- have been supported by subsequent molec- ing to the margin of the jaw. The parotoid ular studies (Garcia-Paris & Wake 2000), so glands are distinct but relatively small. The we are confident that this morphologically head is rounded rather than flattened, and is distinct form represents another new spe- little differentiated from the neck and an- cies. terior body. Maxillary teeth are relatively numerous (36 in the holotype, with 4 pre- Nototriton stuarti, new species maxillary teeth), and there are moderate Stuart's Moss Salamander numbers of vomerine teeth (20). The limbs are short (combined limb length SL = Holotype.-UTA A-33686, an adult male 0.35), with narrow hands and feet that bear from 11.6 km (road) WSW Puerto Santo short, poorly developed digits. The digits Tom&, Montaiias del Mico, Depto. Izabal, are partly fused basally and the outer digits Guatemala, 88"40'W, 15'38'N, 744 m elev., are very short. Digits 2 and 3 of the hand collected on 6 Jan 1991 by Jonathan A, and 2, 3, and 4 of the foot are narrow and Campbell. have pointed tips. The tail is long (1.26 Diagnosis.-Nototriton stuarti is a mem- times SVL) and tapered, with little evidence ber of a clade of northern species of Noto- of basal constriction. The tail is stout ba- triton. It differs from all other members of sally and is of the same diameter as the pos- the clade in having a broader head (0.14 terior end of the body. times SL, from tip of snout to posterior an- Measurements of holotype (in mm).- gle of vent, whereas males of other northern Snout-vent length 32.6, tail length 41.2, tail species have the following values: barbouri depth at base 3.3, tail width at base 3.2, 0.12, lignicola 0.1 1, limnospectator 0.1 1) distance between forelimb and hind limb and fewer maxillary teeth (36, versus 48- insertions 19.4, trunk width just posterior to 60 in other northern species). It is further forelimbs 3.7, distance from tip of snout to distinguished from lignicola, limnospecta- gular fold 6.2, head width 4.5, head depth tor, and barbouri by having a larger nostril 2.5, distance between eye and nostril 0.9, (0.012 times SL, versus 0.0054.011 in elliptical nostril 0.4 X 0.2, distance between barbouri, 0.005-0.01 6 in brodiei, 0.006- nostrils I. 1, eye diameter 1.6, interorbital 0.009 in lignicola and 0.003 in limnospec- distance 1.8, forelimb length 5.6, hindlimb tator), although the nostril is small in com- length 5.8, width of hind foot 1.6, third toe parison with some Costa Rican members of length 0.4, fifth toe length 0.1. the genus. It differs from some species in Osteology.-Information has been ob- having longer limbs (combined limb length tained from radiographs of the holotype. 0.35 times SL, versus 0.31-0.33 in brodiei The skull is relatively well developed, and and 0.30-0.34 in lignicola) and narrower in general resembles the skulls illustrated feet (foot width 0.05 times SL, versus 0.06 for Honduran species of the genus by in barbouri and brodiei and in male limno- McCranie et al. (1998), but with some im- spectator) . portant distinctions. The nasals are relative- Description.-This species is known ly large and are fragmented along the an- from a single individual that is about av- terior margin. The nasolacrimal foramen is erage in size for the genus (32.6 mm, SL). clearly evident and is surrounded by bone, It has small nostrils (0.012 times SL) and a but it is unclear if the nasolacrimal duct broadly rounded snout (but narrow for an pierces the prefrontal bone (as in the genus 8 18 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Cryptotriton) or if the nasal forms the an- sum and are relatively uniform pale gray, terior margin of the foramen and the pre- but there are some obscure whitish spots on frontal the posterior margin (as in some oth- the throat.