Zootaxa 4661 (1): 197–200 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Correspondence ZOOTAXA Copyright © 2019 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4661.1.12 http://zoobank.org/urn:lsid:zoobank.org:pub:7F9CBF28-6AB1-4BD9-8E9E-001F0C372BAA

The tadpole of boehmei in southeastern

CONNOR M. FRENCH1,2, CARMEN BURKETTE1, STEFFEN REICHLE3 & JASON L. BROWN1,4 1Department of Zoology, Southern Illinois University, 1125 Lincoln Drive, Carbondale, IL, USA, 62901 2Current Address: Biology Department, The Graduate Center, City University of New York, 365 Fifth Avenue, New York, NY, USA, 10016 3Museo de Charupas, Santiago de Chiquitos, Bolivia 4Corresponding author. E-mail: [email protected]

To date, half (16 of 32) of the species of Ameerega have had their tadpoles described: A. altamazonica, A. bassleri, A. bilinguis, A. braccata, A. cainarachi, A. flavopicta, A. hahneli, A. macero, A. parvula, A. petersi, A. picta, A. rubriventris, A shihuemoy. A. silverstonei, A. smaragdina, and A. trivittata (Lescure, 1976; Silverstone, 1976; Duellman, 1978; Myers & Daly, 1979; Rodriguez & Myers, 1993; Haddad & Martins, 1994; Lötters et al., 1997; Duellman, 2005; Costa et al., 2006; Twomey & Brown, 2008; Brown & Twomey, 2009; Poelman et al., 2010; Schulze et al., 2015). Ameerega boehmei is a putative member of a clade containing Ameerega braccata, A. flavopicta, A. berohoka, A. munduruku, all of which in- habit various parts of the ‘dry diagonal’ between the Amazon and Atlantic rainforests (Prado & Gibbs, 1993). Adult in this group are morphologically similar, generally dark-bodied with yellow dorsolateral stripes, orange flash marks and some also possessing bright-yellow dorsal spots. Despite considerable research on their breeding behavior, acoustics and systematics (Lötters et al., 2009; Forti et al., 2013), the tadpole of Ameerega boehmei, the southern-most and western- most distributed species in this tentative group, has not been described. Ameerega boehmei is restricted to the Serranías de Santiago and Chochis, isolated Precambrian sandstone massifs in the Chiquitanía region of Bolivia (Lötters et al. 2009). The cerrado semi-arid open savanna ecosystem where these frogs are found is surrounded by seasonally dry tropical forests that have undergone fire regimes of variable intensity in the past (Power et al. 2016). Ameerega boehmei are found close to slow-flowing streams and are observed calling from rock crevices or small tufts of riparian grasses and shrubs punctuating the edges of streams. We observed six adults emitting territorial calls across the surveyed time period (21 July 2017, 11:30 am to 2:15 pm) and recorded the temperature at two crevice calling sites as 24.1º C and 23.4º C, recorded at midday. No tadpoles were seen in the observed stream. However, between 80-120 tadpoles were observed in an adjacent small pool (Fig. 2). No predatory insects or fish were observed in the pool. It had a thick mixture of sand and silt lining the bottom and was partially shaded by a rocky outcrop and bushes. Its maximum depth was 170 mm and its width at the widest points were 191 mm and 112 mm. Water temperature ranged from 25.3º C to 26.2º C, recorded at 11:10 am. The surrounding relative humidity was 51.4%. Tadpoles exhibited behavior similar to other Ameerega species with benthic tadpoles, settling at the bottom of the pool, mostly in shaded areas, unless disturbed (Schultze et al. 2015). Nearby pools contained no tadpoles and were inhabited by predatory insects. Specimens are housed in Colección Boliviana de Fauna (CBF) in La Paz, Bolivia and were collected by SR in Ser- ranía de Santiago, roughly 6 km east of Santiago de Chiquitos (18.19’ S, 59.34’ W, ca. 800 m above sea level), Chiquitanía region, Provincia San José de Chiquitos, Departamento Santa Cruz, Bolivia, April 2002. Tadpoles were fixed in 10% buff- ered commercial formaldehyde solution. Tadpoles were staged following the Gosner’s table (Gosner, 1960). The descrip- tion is based on an individual in Gosner’s Stage 28 (CBF 7505). Eight measurements were taken with a dial caliper (to the nearest 0.1 mm) following Altig and McDiarmid (1999): total length, body length, tail length, maximum tail height, tail muscle height, tail muscle width, internarial distance, and interocular distance. Morphometric data are given as mean ± standard deviation. The tadpole of A. boehmei belongs to the exotrophic ecomorphological guild, benthic type as defined by Altig & Johnson (1989). This description is based on a tadpole (CBF 7505) at developmental Stage 28 (Gosner 1960; Fig. 1) with a total length 26.8 mm; body length 10.6 mm; maximum width 6.2 mm; depth 5.1 mm; internarial distance 1.8 mm; eye to nares distance 0.9 mm; eye diameter 1.2 mm; interorbital distance 2.2 mm; oral disc width 2.3 mm; tail length 16.2 mm. The body is globular, compressed in lateral view, and ovoid in dorsal view. The snout is rounded in dorsal view

Accepted by E. Twomey: 25 Jul. 2019; published: 27 Aug. 2019 197 (Fig. 1). The oral disc is small and located anteroventrally (Fig. 1). Papillae are laterally emarginated, simple and coni- cal; absent on middle third of anterior labium. Anterior jaw sheath has a medial indentation with serration, posterior jaw sheath V-shaped and has serration throughout. Lateral processes long, extending well past lower jaw sheath. Labial tooth row formula (LTRF) is 2(2)/3. P-1 and P-2 equal width, P-3 shorter (80% of P1 and P2). Nares are oval, small, without projections and inflections, and are located dorsolaterally. Eyes are small and oriented dorsolaterally. The spiracle is single, sinistral, and located just before mid-body. Lateral lines present on either side of the dorsal midline, extending from the nares to the base of the tail. Fins are concave and the posterior end is pointed. Fins reach maximum height at the last third of the tail; the dorsal fin does not extend onto the body. The maximum tail height is reached at about mid-length and is as high as body height. In preservative, the body is dark grey, the belly is slightly translucent with digestive tract slightly visible; caudal musculature creamy white with small, irregular grey flecks and slightly translucent fins. In life, the body is dark brown with black spots, the belly is transparent but slightly pigmented posteriorly, digestive tract well vis- ible through skin, caudal musculature poorly pigmented and transparent tail fin with melanophores in small and irregular clusters along the tail. Ontogenetic variation of 12 characteristics was measured on 6 tadpoles in stages 25–30 (Gosner 1960) and minimal inter-tadpole variation was observed (Table 1). Despite the unique habitat, distribution and natural history, the morphology of the tadpole of Ameerega boehmei ap- pears to be incredibly similar to most other Ameerega species. For a detailed discussion of tadpole variation in context of other members of this group, see Poelman et al. (2010), Schultze et al. (2015), and Menin et al. (2017).

FIGURE 1. Tadpole of Ameerega boehmei. Top to Bottom: dorsal, lateral, and ventral view of tadpole (CBF 7505). Inset: illustration of the oral disc from a preserved specimen (CBF 7505; Gosner stage 28). Scale bar = 5 mm.

198 · Zootaxa 4661 (1) © 2019 Magnolia Press French et al. FIGURE 2. (a) An overview of the habitat for Ameerega boehmei in Serranía de Santiago, about 6 km east of Santiago de Chiquitos in Provincia San José de Chiquitos (b) Habitat where adult males were observed in the small patches of vegetation adjacent to the stream (c) Pool used for tadpole deposition by Ameerega boehmei (d) Adult Ameerega boehmei (e) Arrow points to tadpole of Ameerega boehmei in situ.

TABLE 1. Table of tadpole measurements. Measurements in mm. CBF 7506 CBF 7507 CBF 7505 CBF 7508 CBF 7510 CBF 7509 Gosner Stage 25 25 28 28 28 30 Total Length 23.0 23.5 26.8 20.2 19.6 21.0 Body Length 10.1 9.3 10.6 9.8 8.8 8.7 Maximum Width 5.6 5.2 6.2 4.4 4.4 4.5 Depth 4.4 5.1 5.1 4.4 4.3 4.9 Internarial Distance 2.0 2.1 1.8 1.9 1.7 1.8 Eye to Nares Distance 0.8 0.9 0.9 0.9 0.8 0.9 Eye Diameter 1.0 0.9 1.2 0.8 0.7 0.7 Interorbital Distance 2.3 2.3 2.2 1.9 2.0 2.0 Oral Disc Width 2.5 2.5 2.3 2.2 2.0 2.1 Tail Length 12.9 14.2 16.2 10.4 10.8 12.3

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