Chiasmocleis albopunctata ( Boettger, 1885 )

Peloso, Pedro L. V., Sturaro, Marcelo José, Forlani, Mauricio C., Gaucher, Philippe, Motta, Ana Paula & Wheeler, Ward C., 2014, Phylogeny, Taxonomic Revision, And Character Evolution Of The Genera Chiasmocleis And Syncope (Anura, Microhylidae) In Amazonia, With Descriptions Of Three New Species, Bulletin of the American Museum of Natural History 2014 (386), pp. 1-1 : 26-33

Chiasmocleis albopunctata ( Boettger, 1885) Figure 13 , plate 1A–B

Engystoma albopunctatum: ( Boettger, 1885) . Chiasmocleis albopunctata: ( Méhely, 1904) .

TABLE 2

Summary of acoustic data from advertisement calls of Chiasmocleis examined for this study

A review of data from literature is available in Santana et al. (2009)

Gastrophryne albopunctata: ( Stejneger, 1910) . Chiasmocleis bicegoi: ( Miranda-Ribeiro, 1920) ;

synonymy by Cruz et al. (1997).

HOLOTYPE: ZMB 10588 (fig. 13) according to Bauer et al. (1966). An adult specimen, examined from photographs; in very good state of preservation.

TYPE LOCALITY: Paraguay, Amer[ica]. Merid[ional].

DIAGNOSIS. A large species for the genus; SVL in males 23.2–32.2 mm, in females 28.2– 38.0 mm (SVL data from Caramaschi and Cruz, 1997). Body ovoid and robust; snout rounded in dorsal and lateral views. Four distinctive fingers, all but FI slightly fringed in both sexes; fingers not webbed; FI well developed with a distinct, well-developed subarticular tubercle between the proximal phalanges; subarticular tubercles present on all fingers; adpressed FI does not reach the subarticular tubercle of FII; adpressed FIV reach but do not pass distal margin of proximal tubercle of FIII; palmar tubercles protuberant, divided; relative finger lengths I,IV,II,III. Five distinctive, well-developed toes present; toes fringed in both sexes; toes free of webbing; TI with a distinct, welldeveloped subarticular tubercle; adpressed TI touch or barely touch subarticular tubercle of TII; adpressed TV does not touch or reaches the middle of the middle subarticular tubercle of TIV; toes lack terminal discs; relative toe lengths I, II,V,III,IV. Males may present few, small dermal spines on hands. Femoral line absent. Dorsum usually grayish or dark brown with white spots and stains, especially on snout and dorsolateral region; venter (belly and undersurfaces of thigh) usually brownish or grayish with many white spots.

VARIATION: Number of specimens examined in detail is insufficient to present conclusive data on geographic variation. Forlani et al. (2011: fig. 1) reported significant variation in dorsal color pattern between distinct populations along the species’ range, but did not comment further on it. Dorsal white spots vary in number, but are always present.

CALL AND TADPOLE: The call of Chiasmocleis albopunctata was described twice, first from a population in Bolivia ( De la Riva et al., 1996) and then from a population in southeastern Brazil ( Oliveira-Filho and Giaretta, 2006). De la Riva et al. (1996) reported a call with a fast series (584.4–907.7 notes/ min) of multipulsed notes (5–8 pulses/note) with a mean dominant frequency of 4311.0– 4664.4 Hz. Oliveira-Filho and Giaretta (2006) reported a slower call rate (423–501 notes/ min), with more pulses per call (9) and a very similar dominant frequency (4306 Hz). Unlike Oliveira-Filho and Giaretta (2006: 67–68), we consider these differences trivial. A larger series of calls from distinct populations is needed to further assess variation in the species.

The tadpole of Chiasmocleis albopunctata was described and illustrated by Oliveira- Filho and Giaretta (2006). External morphology is similar to other species of Chiasmocleis ( McDiarmid and Altig, 1999) , but differs from known larvae in having wartlike ornamentations on the oral flaps.

REMARKS. Genetic distances between all specimens of Chiasmocleis albopunctata included in the phylogenetic analysis are given in table 3.

DISTRIBUTION: This species is common in open habitats and its distribution in the Amazon is restricted to transitional Cerrado/Amazonia. The species is widely distributed in open areas in Paraguay, Bolivia, and central and northern Brazil. Forlani et al. (2011) recently discussed the range of the species, provided an updated map, and

19.0 mm. Photo M. Bustamante- AmphibiaWebEcuador (see Ron et al., 2014).

commented on a dubious record of a specimen from Pará, Brazil. We have no additional information on the distribution of this species and refer to Forlani et al. (2011) for data on this matter. The species is common in the Rio Tocantins basin with several records in the state of Tocantins, Brazil ( Forlani et al., 2011).

Chiasmocleis anatipes Walker and Duellman, 1974 Figure 14 , plate 1C–H

HOLOTYPE: KU 146035 (fig. 14); adult male, examined from photographs; in very good state of preservation.

TYPE LOCALITY: Santa Cecília, Província Napo, Ecuador. Coordinates not given in the original. We used the following coordinates, from Paynter (1993): 00 ° 039N / 76 ° 589W.

DIAGNOSIS: A medium-sized species for the genus; SVL in males 18.0– 24.2 mm ( Walker and Duellman, 1974; our data), sole female examined 24.9 mm. Body moderately slender; snout rounded in dorsal and lateral views. Four distinctive fingers, all but FI fringed; fingers not webbed; FI well developed with a subarticular tubercle present between the proximal phalanges; tubercles present on all fingers; adpressed FI surpasses the subarticular tubercle of FII; adpressed FIV does not touch distal tubercle of FIII; palmar tubercles visible, divided; relative finger lengths I,IV,II,III. Five distinctive, well-developed toes present; toes fringed and extensively webbed in males (see remarks for comments on females); TI with a distinct well-developed subarticular tubercle; adpressed TI does not touch or barely reaches subarticular tubercle of TII; adpressed TV reaches the middle of the middle subarticular tubercle of TIV; TII–V with terminal discs; relative toe lengths I,II,V,III,IV. Many dermal spines on dorsum and cloacal region in both sexes; spines on chin of males present, absent in females; spines always more abundant and developed in males. Inguinal blotch and femoral line absent. Venter, belly, and undersurfaces of thigh cream colored with large dark brown irregular spots or blotches.

VARIATION: The number of specimens analyzed do not allow for a detailed analysis of variation. Intensity of marbling on dorsum may vary and a dorsal midline may be absent or present (pl. 1C–H). Number and shape of ventral dark spots vary among specimens (figs. 14–15, pl. 1F, H), being almost entirely absent in the specimen depicted in plate 1F. Detailed description of color in life was given by Walker and Duellman (1974).

CALL AND TADPOLE: The call of Chiasmocleis anatipes was never described; reported to consist of a ‘‘short buzz’’ ( Rodriguez and Duellman, 1994). A recording and a brief sonogram are available at from Ron et al. (2014), but the recording seems to be unsuitable for a detailed bioacoustic analysis. From the sonogram published by Ron et al. (2014) it is clear that the call consists of a short series of multipulsed notes.

The tadpole of Chiasmocleis anatipes was first described by Walker and Duellman (1974) and briefly redescribed by Duellman (1978) and Rodriguez and Duellman (1994). According to those descriptions the tadpole of C. anatipes presents paired spiracles located ventrolaterally and a dextral vent tube, a character state present only in pipoids ( Orton, 1953; Haas, 2003). Duellman apparently noticed the mistake, but the correction never appeared in print; the following is taken from Donnelly et al. (1990): ‘‘ Duellman (1978) described paired ventral spiracles in C. anatipes but the spiracle is single and midventral (W.E. Duellman, personal commun. to R. Altig).’’ Note, however, that Ronn Altig was not an author of the Donnelly et al. paper, but likely acted as a reviewer for the manuscript. We have not examined the mentioned specimens and cannot provide further clarification on the issue. Nonetheless, it is clear that the tadpoles of C. anatipes need to be accurately redescribed.

REMARKS: Walker and Duellman (1974) described Chiasmocleis anatipes based on a series of seven specimens, all males, collected at the type locality. The species was diagnosed based mostly on the amount of toe webbing, extending ‘‘to the bases of the terminal phalanges of each digit’’ ( Walker and Duellman, 1974). Females were unknown at that time. Rodriguez and Duellman (1994) reported a population of C. anatipes from northern Peru (Iquitos region), including males and females, and reinforced in their diagnosis that ‘‘the toes are fully webbed’’ ( Rodriguez and Duellman, 1994: 73). We examined two of the female specimens from Iquitos (MHNSM 15697, 15700, from photographs only: fig. 15A, B) and they clearly do not show full webbing on toes. From the photos it is impossible to discern, however, whether webbing is basal or even totally absent. One of those specimens (MHNSM 15700) shows a ventral pattern very similar to that of the holotype of C. ventrimaculata (fig. 38) and agrees with the description given by Andersson (1945). The other specimen (MHNSM 15979), however, shows a ventral pattern intermediate between the holotype of C. devriesi (MHNSM 21540, not examined, but see figs. 15D, pl. 5G) and that of a specimen of C. ventrimaculata (MHNSM 21539: fig. 15C) from the same region. Both the holotypes of C. ventrimaculata and C. devriesi are females. We had access to a series of specimens from two localities in Napo, Ecuador, that we refer to C. anatipes . All have ventral color patterns very similar to examined types (holotype examined only from photographs; see fig. 14). Most of specimens of this series from Napo (LACM specimens) are juveniles and cannot be unambiguously identified, but two are adults, one male (QCAZ 51042), and one female (QCAZ 51041). The toes of the male specimen are fully webbed and it is unambiguously identified as C. anatipes . The female specimen is not fully webbed, but webbing can be easily seen between TII–TIII, TIII–TIV, and TIV–TV (see also pl. 1G). The two adult specimens (QCAZ 51041–51042) were also included in the genetic analyses and they have identical sequences for 16S and only two SNPs in the COI sequence. These two specimens are sister terminals in the phylogenetic analysis, and therefore we confirm that the female specimen is unambiguously assigned to C. anatipes , clearly showing that full webbing is not a diagnosis of the species as a whole, but solely of its males (contra Rodriguez and Duellman, 1994). In addition to the specimens from Napo, two specimens from Orellana, Ecuador, were also examined for morphology and included in the phylogenetic analyses (QCAZ 44341–44342). These are juvenile specimens (sex could not be determined), but basal webbing is present in both. They would, therefore, be assignable to C. anatipes . In the phylogeny, these two specimens cluster with the single representative of

TABLE 4 Uncorrected pairwise distances between 16S sequences of Chiasmocleis anatipes and C. devriesi

C. devriesi (MHNSM 21540, from Genbank). Genetic distances in the 16S sequences between all specimens of C. anatipes , as well as C. devriesi , is given in table 4.

DISTRIBUTION (fig. 16): Known from northeastern Ecuador, at Santa Cecilia, Provincia Sucumbios (type locality), and at the neighboring provinces of Orellana and Napo. Rodriguez and Duellman (1994) reported the species from the Iquitos region, Peru, but we consider the record questionable (see Remarks). Reports of this species for the Manaus region are based on misidentified specimens (P.L.V.P., personal obs.).

Figures

Fig. 13. Holotype of Chiasmocleis albopunctata from Paraguay (ZMB 10588) in (A) dorsal and (B) ventral views. SVL 5 25 mm.

Fig. 14. Holotype of Chiasmocleis anatipes (KU 146035) in (A) dorsal and (B) ventral views. SVL 5