Phylogeny, Taxonomic Revision, And Character Evolution Of The Genera Chiasmocleis And Syncope (Anura, Microhylidae) In Amazonia, With Descriptions Of Three New Species Author Peloso, Pedro L. V. Author Sturaro, Marcelo José Author Forlani, Mauricio C. Author Gaucher, Philippe Author Motta, Ana Paula Author Wheeler, Ward C. text Bulletin of the American Museum of Natural History 2014 2014-03-20 2014 386 1 1 http://www.bioone.org/doi/abs/10.1206/834.1 journal article 10.1206/834.1 0003-0090 5378601 Chiasmocleis antenori ( Walker, 1973 ) Syncope antenori : ( Walker, 1973 ) . Figure 17, plate 2A–D HOLOTYPE : KU 124009; adult female, examined from photographs (fig. 17). Specimen well preserved, although a very large incision was made on the belly region. Color pattern only slightly discernible on the venter; cream venter with small whiteish spots. TYPE LOCALITY : Puerto Libre, Río Aguarico, 570 m , Provincia Napo , Ecuador . DIAGNOSIS: A small species for the genus; SVL in females 12.4–13.6 mm , no data collected for males. Body slender to slightly robust, snout rounded in dorsal and lateral views. FI very much reduced without subarticular tubercle, tip slightly pointed; FII and FIII well developed with hardly visible subarticular tubercles on FIII, one on FII and two on FIII; FIV reduced but clearly visible, without subarticular tubercles; palmar tubercles not present; relative finger lengths I,IV,II,III. Four distinctive toes present; TI lost, TV reduced but clearly distinguishable; toes not webbed; tip of toes III–IV rounded with terminal discs present. Relative toe lengths II,V,III,IV. No dermal spines. No femoral stripe present. Venter (belly and undersurfaces of thighs) beige with small light spots. VARIATION: Despite adding several specimens of Chiasmocleis antenori into the genetic analysis, the number of specimens analyzed for morphological characters is fairly limited, restricted to three paratypes and a few specimens collected by P.L.V.P. in Serra do Divisor, Acre , Brazil . The specimens from Acre show variable amount of white spots on dorsum. Color of the iris color varies from golden to orange and red (see pl. 2A–D). CALL AND TADPOLE: Call is unknown. Chiasmocleis antenori has endotrophic larvae developing inside terrestrial bromeliads ( Krügel and Richter, 1995 ). The tadpole was described in detail, including several developmental stages, by Krügel and Ritchter (1995). REMARKS: This species is similar in color pattern and external morphology to Chiasmocleis magnova , from which it differs in having an external tympanum visible (not visible in C. magnova: Moravec and Kohler, 2007 ), by its smaller size (maximum recorded SVL 13.6 mm , versus 18.3 mm in C. magnova ), and in having seven presacral vertebrae (eight in C. magnova ). Phylogenetic analysis found two clades within Chiasmocleis antenori , one with three specimens from Ecuador ( Orellana and Pastaza ) and the other with all of the syntopic samples from Brazil and a sample labeled as C. carvalhoi (as Syncope carvalhoi in de Sá et al., 2012 ) from San Martin , Peru . Genetic distances between specimens of C. antenori vary from 0 % –11.4 % , with as much as 9.1 % distance between syntopic specimens from Acre , Brazil . Genetic distances between all specimens of C. antenori included in the phylogenetic analysis are given in table 5. Fig. 16. Distribution of Chiasmocleis anatipes . Star 5 type locality; circle 5 examined specimens. Unfortunately, we have not examined the Ecuadorian or the Peruvian specimens, making any taxonomic conclusions about the status of these populations unattainable at present. DISTRIBUTION (fig. 18): Ecuador ( Napo , Orellana , and Pastaza ), and western Brazil ( Acre ). Coloma et al. (2004) gave a wider distribution and included a large portion of northeastern Peru . A specimen identified as Chiasmocleis carvalhoi from San Martin , Peru ( de Sá et al., 2012 ), was found inside our samples of C. antenori and is regarded as a confirmed record of the species for Peru . Chiasmocleis avilapiresae Peloso and Sturaro, 2008 Figures 19–20 , plates 2E–H, 3 HOLOTYPE (fig. 19): MPEG 23299 ; adult female, in very good state of preservation. TYPE LOCALITY : Estação Científica do Programa Pró-Biodiversidade da Amazônia (PPBio) (aprox. 01 ° 599S, 51 ° 399W), Floresta Nacional Caxiuanã, Municipality of Portel, state of Pará, Brazil . DIAGNOSIS: A large species for the genus; SVL in males 22.2–28.0 mm ( N 5 28), in females 23.6–37.8 mm ( N 5 82). Body ovoid and robust; head triangular, snout rounded in dorsal and lateral views. Four distinctive fingers, all but FI fringed in males, less fringed in females; fingers not webbed; FI well developed, with a distinct, well-developed subarticular tubercle present between the proximal phalanges; distinctly visible subarticular tubercles present on all fingers; adpressed FI reaches or surpasses the subarticular tubercle of FII; adpressed FIV does not reach or barely reaches distal tubercle of FIII; palmar tubercles protuberant, divided; relative finger lenghs I,II,IV,III. Five distinctive and well-developed toes present; Fig. 17. Holotype of Chiasmocleis antenori (KU 23299) in ( A ) dorsal, and ( B ) ventral views. SVL 5 12.3 mm . Photo M. Bustamante- AmphibiaWebEcuador (see Ron et al. 2013). toes fringed, less distinct in females; toes usually extensively webbed in males, only basally webbed in females; TI with a distinct, well-developed subarticular tubercle; adpressed TI does not touch or barely touches subarticular tubercle of TII; adpressed TV does not touch or reaches only to the middle of subarticular tubercle of TIV; TII–IV with terminal discs, usually more developed in females, but also present in males; relative toe lengths I,II,V,III,IV. Males with many dermal spines on fingers and toes; both sexes may show dermal spines, but they are much more numerous and developed in males; males with many spines on anterior portion of chin, absent in females. Inguinal blotch usually absent. Femoral line present. Color pattern given in the variation section. VARIATION: Ventral (throat and belly) color pattern can vary from almost entirely uniform (cream, white, brown, or black) to densely reticulated (fig. 20; see also Peloso and Sturaro, 2008 ). A single specimen, out of 206 examined, showed strong dark vermiculations on the venter (fig. 20D). Western populations (west of the state of Amazonas , Mato Grosso and Rondônia ) have, in general, fewer spots on the ventral region than those from eastern populations (eastern Amazonas and all Pará ). Additionally, the specimens from Aripuanã, Mato Grosso , are slightly smaller than those of remaining populations. A dorsal midline is usually absent, but is present in some specimens, commonly in contact with the femoral line posteriorly (pl. 3D). The dorsum of Chiasmocleis avilapiresae shows a great degree of color variation in life (see pls. 2E–H and 3). In general individuals have a dark, grayish dorsum (pl. 2E, G) while the dorsal surfaces of limbs are usually brightly colored—yellow, orange, or red. In several specimens the flanks are reddish (pls. 2G, 3G) and in a few specimens the reddish pattern extends through most of the dorsum of specimens (pls. 2H, 3H). Some specimens from Mato Grosso (pl. 3E) and Rondônia (pl. 3A, C, F) have golden yellow or orangeish dorsums. TABLE 5 Uncorrected pairwise distances between 16S sequences of Chiasmocleis antenori

Locality	1	2	3	4	5
1	Brazil: Acre (MTR 28350)
2	Brazil: Acre (MTR 28373)	0.080
3	Brazil: Acre (MTR 28416)	0.080	0.005
4	Ecuador: Pastaza: (QCAZ 38506)	0.065	0.092	0.087
5	Ecuador: Orellana: (QCAZ 38719)	0.072	0.095	0.090	0.007
6	a Peru: San Martin (KU 215720)	0.065	0.087	0.087	0.067	0.065

a This specimen is labeled as Chiasmocleis ‘‘carvalhoi’’ in the phylogenetic tree (figs. 6, 7). Sequences for it derive from Genbank and we did not examine the specimen for phenotypic characters. The snout may be whitish or follow the general dorsal pattern. For additional notes on variation, see Peloso and Sturaro (2008) . CALL AND TADPOLES: The advertisement call of Chiasmocleis avilapiresae was recently described by Barros et al. (2010) . The call consists of a fast series (295.4 notes / min) of multipulsed notes (7–18 pulses/note). Mean note duration was 98.0 ± 9.8 ms, and mean interval between notes 111.1 ± 0.1 ms. Mean dominant frequency of was 3368.2 ± 73.2 kHz. Tadpoles are unknown. DISTRIBUTION (fig. 21): Brazil , widespread in rainforest areas south of the Rio Amazonas and west of the Rio Tocantins . Known in the states of Acre , Amazonas, Rondônia , Mato Grosso , and Para´. We are unaware of records of this species in adjacent Bolivia and Peru . Fig. 18. Distribution of Chiasmocleis antenori and C. carvalhoi . Star 5 type locality of C. antenori . Circles 5 examined specimens of C. antenori ; cross 5 type locality of C. carvalhoi (see text for details). triangles 5 examined specimens of C. carvalhoi . Fig. 19. Holotype of Chiasmocleis avilapiresae (MPEG 23299) in ( A ) dorsal, and ( B ) ventral views. SVL 5 34.9 mm. REMARKS: In the list of paratypes given by Peloso and Sturaro (2008: 42) , collection numbers from MNRJ were mistyped. The publication reads MNRJ 14231–80, but the correct collection numbers for that lot are MNRJ 44231–44280. We thank M. Targino for pointing out this mistake to P.L.V.P. Genetic distances between all specimens of C. avilapiresae included in the phylogenetic analysis are given in table 6.