First record of Pseudopaludicola boliviana Parker , 1927 ( Anura , Leptodactylidae , Leiuperinae ) in the Brazilian state of Amapá , with comments on its advertisement call and distribution

We provide the first record of Pseudopaludicola boliviana from the state of Amapá (municipality of Macapá), northern Brazil. It is also the species easternmost record, extending its known geographic distribution in Brazil about 1,260 km from the state of Roraima, northern Brazil. Additionally, we also provide a call description for this population of P. boliviana and make comparisons with the available acoustic data for the species from Bolivia and Argentina. This new record fills a gap in the distribution of P. boliviana in northern Brazil.

Pseudopaludicola boliviana was described from Santa Cruz, Bolivia (Parker 1927), and assigned to the P. pusilla group mainly by the presence of T-shaped ter- minal phalanges, expanded toe tips (disks or pads), and tubercles on the upper eyelids and heels (Lynch 1989;Cardozo and Suarez 2012).This species has a cis-Andean distribution associated with open environments from northern South America (Colombia and Venezuela; Lynch 1989), across Bolivia (De la Riva et al. 2000), to the south (Paraguay and Argentina;Lobo 1991;1994).The advertisement call of this species has already been described twice before, once at or near the type locality (Santa Cruz, Bolivia;Márquez et al. 1995) and later from Corrientes, Argentina (Duré et al. 2004).These calls diverge in both temporal and spectral call traits.
Calls of one male were recorded using a Sennheiser® K6/ME67 directional microphone and a Marantz® PMD 670 digital recorder set at a sampling rate of 44.1 kHz and a sample size of 16 bits, and analyzed with Raven Pro 32-bit 1.5 beta version (Bioacoustics Research Program 2014).Temporal traits were manually measured from oscillograms, spectral traits from spectrograms The individuals of Pseudopaludicola boliviana from the state of Amapá (Figure 2) exhibited variation in morphology and color, such as the presence or absence of vertebral stripe, interorbital blotch, glandular folds, and the extent of glands on dorsum, whereas the morphological features of conical tubercles on heels, tubercles evident on upper eyelids (but not elongated), and expanded toe tips (disks or pads) were present in all examined specimens, which represent diagnostic characters for this species (Lynch 1989).Calling males had white vocal sacs.
Males of Pseudopaludicola boliviana called during daytime and decreased call activity after nightfall in all three areas.Males mostly called at the margins of shallow temporary pools and floodplains.At these sites, males were in calling activity from 17:30 to 22:40 h during the entire dry season, and maintained their activity as long as the reproductive sites were flooded (usually from July to November).We also observed the following syntopic species in calling activity: Rhinella major (Muller & Helmich, 1936), Dendropsophus cf.nanus (Boulenger, 1889), Scinax nebulosus (Spix, 1824), S. ruber (Laurenti, 1768), and Leptodactylus fuscus (Schneider, 1799).
In Larger sample sizes for both populations are required prior to any assumption so that individual variation and effects of body size on spectral call traits can be assessed and tested.
In comparison with the call described from Argentina (Duré et al. 2004), temporal traits of call duration, call interval, and notes per call mostly overlapped, but duration of 1 st to 4 th notes were considerably shorter than our values.This might be explained due to different analytic procedures.We measured note period (from beginning of one note to beginning of the next; see Jang et al. 2011 for definition) instead of note duration (beginning to end of one note) given that intervals between notes were often barely identified or inexistent in our sample, whereas inter-note intervals in calls analyzed by Duré et al. (2004) seemed to be more discernible based on their Figure 3B.With respect to call frequency, minimum and dominant frequencies had quite similar values, but maximum frequency of their calls presented an expressive lower value range (< 6000 Hz) compared to ours (6966 to 7390 Hz).These authors did not provide their software settings for call analysis.Differences in spectral resolution might explain the diverging values observed for maximum frequency, but the other spectral traits were quite similar.It might be worth for future studies to stress that their Figures 1 and 2 have a time scale problem, being that Figures 1A and 2A portrayed the temporal and spectral structures, respectively, of a series of five calls, and Figures 1B and 2B should probably have provided the temporal and spectral structure, respectively, of one call at higher resolutions, but time scales in all figures (1A-B, 2A-B) 1A and 1B are the same.Plus, spectral structure of the portrayed call in Figure 2B is completely uninformative due to the lack of resolution.
The new distributional record of Pseudopaludicola boliviana is the first for the state of Amapá and represents the easternmost occurrence for this species, whose distribution range in northern South America encompasses (east of the Andes) Colombia, Venezuela, Guyana, Suriname (referred to as P. pusilla in Hoogmoed 1979), as well as northern Brazil in the state of Roraima (Parker 1935;Lobo 1991).The new record extends the known distribution of this species eastward by about 1,260 km in Brazil.
Additional surveys are still required to better understand the distribution pattern of Pseudopaludicola boliviana, whose known range has a considerable gap between Cerrado and Chaco formations (Argentina, Paraguay, and Bolivia) and the savanna and open grassfield formations in northern South America, split by a wide region covered with forested environments (Amazon Rainforest) in northern Brazil (Figure 1).

Figure 1 .
Figure 1.Geographic distribution map of Pseudopaludicola boliviana in South America.New distributional records are for the Brazilian state of Amapá (indicated in green on the map).Distribution records and their respective coordinates are provided in the Appendix (TableA1).

Figure 3 .
Figure 3. (A) Waveform section (ca.815 ms) of three advertisement calls of Pseudopaludicola boliviana, and (B) spectrogram and respective oscillogram of the first call in A. Notice the stereotyped four-note structure, and an emphasized frequency band at approximately 10 kHz.Air temperature: 27.5°C.

Figure 2 .
Figure 2. Pseudopaludicola boliviana pair in amplexus.Notice the distinctive morphological features: conical tubercle on heels, tubercles on upper eyelids, and expanded toe tips.
Check List 12(6): 1991, 15 November 2016 doi: http://dx.doi.org/10.15560/12.6.1991ISSN 1809-127X © 2016 Check List and Authors comparison with the acoustic data described for a population of Pseudopaludicola boliviana from Bolivia (Márquez et al. 1995), our calls (state of Amapá, northern Brazil) had the same four-note call pattern, temporal traits of note duration and pulses per note had similar values as well.Dominant frequency, on the other hand, was remarkably lower (4119 to 4543 Hz; Márquez et al. 1995) compared to our values (4737 to 5297 Hz).