Taxonomic review of Physalaemus olfersii (Lichtenstein & Martens, 1856) with revalidation of Physalaemus lateristriga (Steindachner, 1864) and description of two new related species (Anura: Leiuperidae) Author Cassini, Carla Santana Author Cruz, Carlos Alberto Gonçalves Author Caramaschi, Ulisses text Zootaxa 2010 2491 1 33 journal article 10.5281/zenodo.195620 f438d90d-81ab-4d3f-8b90-59e28d2423b6 1175-5326 195620 Physalaemus olfersii ( Lichtenstein & Martens, 1856 ) ( Figs. 4 , 5 , 6 , 7 ) Phryniscus olfersii Lichtenstein & Martens, 1856 . Phrynidium olfersi — Cope, 1867 . Paludicola olfersi — Peters, 1882 . Physalaemus olfersi — Parker, 1927 . Lectotype . ZMB 3375 ( Fig. 4 ), by present designation, female (SVL 29.4; TBL 12.3; HL 8.1; HW 6.5 mm ), collected in an undetermined locality in the Serra do Mar mountain range, from the state of Rio de Janeiro to São Paulo ( Fig. 8 ), by Ignaz Marian Franz Olfers and Frederich Sellow, between the years of 1818 to 1819 . Diagnosis. (1) Body length small to moderate (SVL 26.2–36.4 mm in males, 22.4–41.1 mm in females); (2) texture of skin shagreened; (3) head longer than wide; (4) snout pointed to subelliptical in dorsal view and acute in lateral profile; (5) lateral black stripe, from the posterior corner of the eye to the inguinal region; (6) inguinal gland poorly evident; (7) oblique white line from the posterior corner of the eye to the arm insertion; (8) white line weakly developed on canthus rostralis and well marked on the border of the upper eyelid and on the dorsolateral fold; (9) annulus of tympanum indistinct; (10) vocal sac developed, dark gray; (11) tarsal tubercle weakly developed; (12) tarsal fold weakly developed or absent; (13) advertisement call composed by one multipulsed note (pulse rate 135 to 156 pulses per second); (14) mean dominant frequency 1.92 kHz. FIGURE 4: Physalaemus olfersii , lectotype (ZMB 3375, adult female). A—dorsal, B—ventral, C—lateral views, and D—foot (photographs conceded by Rainer Günther). Comparison with other species. Physalaemus olfersii is distinguished of P. aguirrei and P. insperatus by the absence of a white line outlining the inferior margin of the mandible and by the absence of the supratympanic fold (both present in those species). Additionaly, P. olfersii is distinguished of P. aguirrei by the presence of an oblique white line from the posterior corner of the eye to the arm insertion (absent in P. aguirrei ); ventral region light brown with brown and white dispersed blotches (ventral region light brown, with white blotches, except on the ventral surface of the thigh that is uniformly white in P. aguirrei ); non– harmonic physical structure of the call (harmonic in P. aguirrei ; Pimenta & Cruz 2004 ); and mean dominant frequency 1.92 kHz (3.1 kHz in P. aguirrei ; Pimenta & Cruz 2004 ). From P. insperatus , by the vocal sac not expanded laterally (expanded laterally in P. insperatus ); and ventral region light brown with brown and white dispersed blotches (ventral region light brown with white blotches, except on the ventral surface of the thigh that is uniformly white in P. insperatus ). Physalaemus olfersii is distinguished of P. feioi sp. nov. and P. orophilus sp. nov. by the presence of a well marked white line on the dorsolateral fold (weakly marked in P. feioi sp. nov. and absent in P. orophilus sp. nov. ); vocal sac uniformly dark gray (with concentrated gray blotches in P. feioi sp. nov. and P. orophilus sp. nov. ); and mean dominat frequency of 1.92 kHz (2.56 kHz in P. feioi sp. nov. and 3.18 kHz in P. orophilus sp. nov. ). Physalaemus olfersii is distinguished from P. lateristriga comb. nov. ; by the loreal region oblique (vertical in P. lateristriga comb. nov. ), annulus of tympanum indistinct (distinct, weakly developed in P. lateristriga comb. nov. ); and tarsal fold weakly developed (absent in P. lateristriga comb. nov. ). Moreover, it is distinguished from P. lateristriga comb. nov. by the tibia light gray or light brown weakly marbled, except on dorsal surface (dark gray or dark brown marked marbled in P. lateristriga comb. nov. ); outer metatarsal tubercle rounded (conic in P. lateristriga comb. nov. ); and advertisement call with lower pulse rate (171 to 202 pulses per second in P. lateristriga com. nov.). Physalaemus olfersii has smaller body length than P. maximus (SVL 41.1–46.5 mm in males, 42.2–51.5 mm in females of P. maximus ); and mean dominant frequency of 1.92 kHz (two values: 0.73 or 0.78 KHz in P. maximus ; Baêta et al. 2007 ). Finally, P. olfersii present larger body length than P. soaresi (SVL 17.9–22.8 mm in males, 19.9–21.52 mm in females of P. s o a re s i ); oblique white line from the posterior corner of the eye to the arm insertion (absent in P. s o a re s i ); tarsal tubercle weakly developed (developed in P. soaresi ); tarsal fold absent or weakly developed (developed in P. soaresi ); developed, dark gray vocal sac (weakly developed and light brown in P. soaresi ); head longer than wide (wider than long in P. s o a re s i ); and advertisement call with mean dominant frequency of 1.92 kHz (3.3 kHz in P. s o a re s i ; Weber et al. 2005 ). Description. Body moderately robust ( Figs. 5 , 6 ); head longer than wide; snout pointed to subelliptical in dorsal view, acute in lateral profile; nostrils elliptical, located and expanded laterally, closer to tip of snout than to eye; canthus rostralis distinct; loreal region oblique; eye slightly protuberant; annulus of tympanum and supratympanic fold indistinct; parotoid gland not developed; presence of a dorsolateral fold from the posterior corner of eye, delimiting the dorsal region from the flank, ending at the anterior margin of the inguinal region; vocal sac subgular, well developed, extending to the chest; choanae small, rounded, well distinguished from each other; tongue narrow, long, free posteriorly; maxillary and premaxillary teeth visible; vomerine teeth absent. Arms short, slender; forearms slightly more robust and as short as arms; fingers thin, long, not fringed; finger lengths I <II ≤ IV <III; large nuptial pad divided, one part covering from 1/2 to 2/3 of the inner metacarpal tubercle and other part enclosing the thumb, except the inner side; subarticular tubercles large, elliptical; outer metacarpal tubercle medium–sized, elliptical; inner metacarpal tubercle large, elliptical, approximately three times the size of the outer metacarpal tubercle; absence of supernumerary tubercles; fingers tip slightly expanded. Legs moderately robust; tibia slightly longer than thigh; sum of tibia and thigh lengths shorter than SVL; tarsal fold absent; toes thin, long, not fringed; toe lengths I <II <III = V <IV; subarticular tubercles large, protruding; inner metatarsal tubercle large, elliptical; outer metatarsal tubercle medium–sized, ovoid; supernumerary tubercles absent; toes tip expanded. Dorsal surface shagreened and ventral surfaces smooth. Ventral disc absent. Color in life. Dorsal color pattern varying from light brown ( Fig. 7 ) to gray with small, irregular, and disperse scattered brown blotches; dorsal median line extending on posterior 2/3 of the dorsum; interorbital white line interrupted at midway; loreal region dark brown with some dispersed white dots; a black stripe from the postorbital region to the anterior margin of the inguinal gland; a white line weakly developed on canthus rostralis and well marked on the border of the upper eyelid and on the dorsolateral fold; a dark brown line bordering the internal margin of the ventral surface of the arms; a rectangular blotch on dorsum of the forearm; transversal bars on dorsum of thighs; an irregular blotch on internal side of tibia, near the heel; gular region dark gray with white dots; white dots on the anterior region of the abdomen; uniformly light brown on the posterior region of the abdomen and ventral proximal surfaces of thighs; nuptial pads brown; color of the inguinal gland not distinguishable of the color pattern of the dorsum; posterior region of the abdomen and tubercles of palmar and plantar surfaces light red. Color in preservative (70% ethanol). General color pattern maintained in preservative. Dorsal surface light brown or gray with a light gray blotch on the center; tubercles of plantar and palmar surfaces; and abdomen, white. Variation (n = 151). Males are slightly smaller than females. Range, mean, and standard deviation of SVL of males and females are in table 4. Few specimens observed have a mucronate snout in dorsal view ( sensu Heyer et al. 1990). Tarsal fold is absent in most specimens analyzed, although some specimens have a tarsal fold weakly developed. FIGURE 5: Physalaemus olfersii (MNRJ 56120) from municipality of Teresópolis, state of Rio de Janeiro. A, dorsal, and B, ventral views. Bar = 20 mm. FIGURE 6: Physalaemus olfersii (MNRJ 56120) from municipality of Teresópolis, state of Rio de Janeiro. A—dorsal, and B—lateral view of head; C—hand, and D—foot. Bars = 10 mm. FIGURE 7: Physalaemus olfersii (DZSJRP 13958) from Parque Nacional da Serra da Bocaina, municipality of São José do Barreiro, state of São Paulo. Photograph by Diogo B. Provete. FIGURE 8: Geographic distribution of Physalaemus olfersii , P. lateristriga comb. nov. , P. feioi sp. nov. , and P. orophilus sp. nov. FIGURE 9: Physalaemus olfersii advertisement call: A—oscilogram, B—audiospectrogram, C—power spectrum, Doscillogram of a cut of the same call evidencing pulsed structure, and E—oscillogram of a cut of the same call evidencing the subpulsed structure. Specimen MNRJ 50312, collected at the municipality of Teresópolis ( 22°27'S ; 42°58'W ), state of Rio de Janeiro, on 20/XI/2007 . Air temperature: 23°C. Advertisement call. Advertisement calls (n = 92 calls) were obtained from eight males (MNRJ 50309– 50314; MNRJ 56119–56120) from the Parque Nacional da Serra dos Órgãos, municipality of Teresópolis, state of Rio de Janeiro, Brazil . The advertisement call (Fig. 9, Table 3 ) is composed by one single multipulsed note, each pulse composed by 2 subpulses (which can be fused or not on sonogram along the call), non– harmonic structure, without frequency modulation; mean duration of 4.04 seconds (SD = 0.4, range = 2.6 to 5.1 seconds); pulse duration of 6 to 7 miliseconds, pulse rate ranges from 135 to 156 pulses per second (mean = 149, SD = 5); mean duration of time between calls 2.07 seconds (SD = 0.8, amplitude = 1.07–2.64 seconds) and the mean value of the dominant frequency 1.92 kHz (SD = 0.33, amplitude = 1.38–2.59 kHz). Remarks. Phyllobates glandulosus , described by Steindachner (1867) , was synonymized with Physalaemus olfersii by Bokermann (1966b) . According to Bokermann (1966b) , the type locality of Phyllobates glandulosus is “ Brazil , probably Serra da Estrela, Inhomirim, Rio de Janeiro.” Steindachner (1867) described the amphibians collected by the “Fregatte Novara” expedition through the years 1857 to 1859 , which ported in Brazil from August 5th to August 31 th 1857. Zoologists from “Fregatte Novara” expedition (Geroge Frauenfeld and John Zelebor) collected at the municipalities of Rio de Janeiro and Petrópolis ( Scherzer 1861 ). “Serra da Estrela” mentioned by Bokermann (1966b) and Scherzer (1861) is a chain of mountains within the Serra dos Órgãos mountain range where the municipality of Petrópolis is located. In the midway of the municipalities of Rio de Janeiro and Petrópolis, at the base of Serra da Estrela mountains, is located “Inhomirim” mentioned by Bokermann (1966b) . Scherzer (1861) didn’t specified where the zoologists of “Fregatte Novara” had collected the Phyllobates glandulosus type , and it could be anywhere in the municipalities of Rio de Janeiro, Petrópolis, Inhomirim or in the way from one to another (Figure 10). In this area, are known to occur nowadays Physalaemus cuvieri , P. maculiventris , P. marmoratus , P. olfersii , and P. signifier . Steindachner (1867) described the specimen as having a triangular head; snout very short; tongue long and rounded; evident tympanum; general color pattern of the body light brown; two turbercles well developed on metacarpus and two smaller on metatarsus and small numerous tubercles on tarsus and on the sole of the foot. Additionally, we could observe on the pictures of the type specimen (NMW 16520; Fig. 11 ) the presence of large inguinal glands associated with dark ocellus; ventral surface homogeneous light brown without blotches. TABLE 3: Acoustic parameters of species of the Physalaemus olfersii group. olf = P. olfersii ; lat = P. lateristriga ; feioi = P. feioi ; orop = P. orophilus ; agui = P. aguirrei ; soar = P. soaresi ; max = P. maximus . Values of parameters: amplitude (mean±standard deviation). Mult = multipulsed note. *What Baêta et al. (2007) “harmonics” are actually “side bands” sensu Vielliard (1993). Species olf lat feioi orop agui soar max FIGURE 10: Area travelled by zoologists of the "Fregatte Novara" expedition (Geroge Frauenfeld and John Zelebor) from August 5th to August 31 th 1857.

Locality of populations analyzed Call duration (seconds)	Teresópolis RJ 2.6–5.1 (4.04±0.4)	São José dos Pinhais PR 2.3–5.5 (3.6±0.7)	Viçosa and Araponga MG 2.5–5.6 (4.3±0.7)	Mariana MG 3.1–5.2 (3.9± 0.6)	Prado BA 0.21–0.25 (0.23±0.01)	Seropédica RJ 1.34–2.40 (1.83±?)	Ouro Preto MG 1.09–2.31 (2.01±0.26)
Structure	NH	NH	NH	NH	H	NH	NH*
Frequency modulation	No	No	ascendent along the call; descendent at the end of the call	ascendent	descedent	ascendent	weak descedent
Notes	1	1	1	1	1	1	1
Pulses Pulse duration (ms)	519–733 (603±52) 6–7	404–990 (672±235) 4–5	30-75 (45±4) 40–45	56 – 87 (78±5) 42–50	- -	mult -	mult -
Sub-pulses	2	2	14 – 19	8-11	-	-	-
Pulse rate (pulses per second) Dominant frequency	135 – 156 (149±5) 1.38–2.59 (1.92±0.33)	171–202 (183±10) 0.75–2.08 (1.68±0.23)	8–16 (12±1.36) 2.31–2.75 (2.56±0.0075)	16–19 (18±0.76) 3.0–3.39 (3.18±0.09)	- 3.10	- 3.3	- 0.732or0.775
n of calls	92	70	110	124	55	6	-
n of males	8	6	11	6	1	-	-
References	Present work	Present work	Present work	Present work	Pimenta & Cruz, 2004	Weber et al. 2005	Baêta et al. 2007

FIGURE 11: Phyllobates glandulosus , holotype (NMW 16520). A—dorsal, B—ventral, and C—lateral views (photographs conceded by Franz Tiedemman and Heinz Grillitsch). Label data: "Brasilien, Novara Expedition, 1857- 1859". The presence of inguinal glands associated with dark ocellus and evident tympanum, distinguish the type of Phyllobates glandulosus from Physalaemus marmoratus ( Nascimento et al. 2006 ) and P. olfersii (present work). Bokermann (1962) distinguished P. signifer from P. maculiventris by the presence of supranumerary tubercles on the ventral surface of the foot in P. signifer and absence on P. maculiventris . In addition, he figured the lateral view of these species and the tympanum annulus is more evident in P. signifer . Thus, from those species which are distributed within the area where Geroge Frauenfeld and John Zelebor possibly collected the type of Phyllobates glandulosus the one that better fits with the Steindachner’s descrition and photograph of the type is Physalaemus signifer . Therefore, we transfer Phyllobates glandulosus from the synonymy of Physalaemus olfersii to the synonymy of Physalaemus signifer (Girard, 1853) . Geographical distribution. Physalemus olfersii is distributed in the Atlantic Rain Forest Domain and its influence areas, from the municipality of Santa Teresa, state of Espírito Santo, southern region of the state of Minas Gerais to the municipality of São Paulo and central region of the state of São Paulo, in the municipality of Botucatu ( Fig. 8 ).