Boletín de la Sociedad Zoológica del Uruguay, 2022

Vol. 31 (2): e31.2.4

ISSN 2393-6940

https://journal.szu.org.uy

DOI: https://doi.org/10.26462/31.2.4

ABSTRACT

The Serra Dona Francisca is in the subtropical region of

the Atlantic Forest and part of its area is protected by a

sustainable use conservation unit. In the present study, we

performed an anuran inventory at different elevations,

providing information on natural history of each species.

Field work was carried out from October 2012 to January

2014. We sampled the anuran by visual and auditory

surveys and used three 50-meter pitfall lines composed of

four 60L buckets with drift-fences at each elevation area.

We registered 63 species belonging to 13 families:

Brachycephalidae (4), Bufonidae (4), Centrolenidae (1),

Craugastoridae (1), Cycloramphidae (2), Hemiphractidae

(2), Hylidae (30), Hylodidae (2), Leptodactylidae (10),

Microhylidae (2), Odontophrynidae (3), Phyllomedusidae

(1), and Ranidae (1). We observed four new records for

the northeastern region of the state of Santa Catarina:

Bokermannohyla luctuosa, Scinax aromothyella,

Odontophrynus americanus, and Leptodactylus

plaumanni. We registered a higher number of species

during the hot and wet months, with subsequent drop

during the cold months. We found an increase of species

richness at higher elevations. Considering that our results

represent 50% of anuran richness of the state of Santa

Catarina, this is an important region for conservation of

biodiversity of southern Atlantic Forest.

Keywords: Subtropical, anura, conservation,

inventory

RESUMEN

La Serra Dona Francisca está ubicada en la región

subtropical de la Mata Atlántica y parte de su área está

protegida por una unidad de conservación de uso

sostenible. En el presente estudio, realizamos un

inventario de anuros a diferentes elevaciones,

proporcionando información sobre la historia natural de

cada especie. El trabajo de campo se llevó a cabo desde

octubre de 2012 hasta enero de 2014. Tomamos

muestras de los anuros mediante búsqueda activa y

utilizamos tres líneas de trampa de caída de 50 metros

compuestas por cuatro baldes de 60 litros con cercas de

deriva en cada área de elevación. Registramos 63

especies pertenecientes a 13 familias: Brachycephalidae

(4), Bufonidae (4), Centrolenidae (1), Craugastoridae (1),

Cycloramphidae (2), Hemiphractidae (2), Hylidae (30),

Hylodidae (2), Leptodactylidae (10), Microhylidae (2),

Odontophrynidae (3), Phyllomedusidae (1) y Ranidae (1).

Observamos cuatro nuevos registros para la región

nororiental del estado d e Santa Catarina:

Bokermannohyla luctuosa, Scinax aromothyella,

Odontophrynus americanus y Leptodactylus plaumanni.

Registramos un mayor número de especies durante los

meses cálidos y húmedos, con la consiguiente

disminución durante los meses fríos. Encontramos un

aumento de la riqueza de especies en elevaciones más

altas. Teniendo en cuenta nuestros resultados que

representan el 50% de la riqueza de anuros del estado de

Santa Catarina, esta es una región importante para la

conservación de la biodiversidad de la Mata Atlántica Sur.

Palabras clave: Subtropical, anura, conservación,

inventario

INTRODUCTION

The Serra Dona Francisca is a historic and natural area

that has been of great scientific value since the XIX

century, when a land glebe from the Santa Catarina

province was given as a dowry to the prince of Joinville,

François d'Orléans, with Princess Francisca Carolina,

the sister of emperor Dom Pedro II (Ficker, 1965). This

region was colonized for logging and development of

agricultural activities, which were subsistence ways for

Bol. Soc. Zool. Uruguay (2ª época). 2022. ISSN 2393-6940Vol. 31 (2): e31.2.4

ANURAN NATURAL HISTORY FROM SERRA DONA FRANCISCA, AN ATLANTIC FOREST REMNANT IN

SOUTHERN BRAZIL

Lucas Ribeiro Mariotto¹* , Sarah Mângia² and Diego José Santana²

1Instituto Blumenauense de Ensino Superior – Centro Universitário UNISOCIESC, 89010-350,

Blumenau, SC, Brazil.

2Instituto de Biociências, Universidade Federal de Mato Grosso do Sul, 79002-970, Campo Grande, MS, Brazil.

*Corresponding author: lucasrmariotto@gmail.com

Fecha de recepción: 11 de julio de 2022

Fecha de aceptación: 6 de octubre de 2022

MARIOTTO, L.R. et al.

the newly arrived European immigrants, especially

Germans and Austrians (Ficker, 1965; Rodowicz-

Oswiecimsky, 1992). The new Dona Francisca colony

also attracted many European naturalists to explore its

rich nature, which resulted in the description of new

animal species for the south of Brazil. Regarding

amphibians, George Albert Boulenger, from the British

Natural History Museum, received many specimens in

the late XIX century collected by Hr. Michäelis

(Boulenger, 1888a). Boulenger listed 21 species for the

Santa Catarina province and described four of them:

Scinax catharinae (Boulenger, 1888a), Vitreorana

parvula (Boulenger, 1894), Physalaemus nanus

(Boulenger, 1888b), and Chiasmocleis leucosticta

(Boulenger, 1888a). In the late XIX and early XX

centuries, while living in the Hansa-Humbolt colony,

adjacent to the Dona Francisca colony, Willhelm

Ehrhardt collected 43 amphibian species (Gutsche,

Kwet, Kucharzewski and Hallermann, 2007, Gutsche,

Kwet, Kucharzewski, Lingnau and Güther, 2007). In

subsequent studies made by Dr. Lorenz Müller, three

new amphibian species were described for this region:

Adenomera nana (Müller, 1922), Vitreorana

uranoscopa (Müller, 1924) and Aplastodiscus ehrhardti

(Müller, 1924). Furthermore, Müller listed 23 amphibian

species for the colony and 38 species for Santa

Catarina province (Müller, 1922).

Currently, amphibians (Lissamphibia) comprise more

than 8400 species in the world (Frost, 2021), and Brazil

harbors the highest diversity (1188 species, Segalla et

al., 2021), with 625 species occurring in the Atlantic

Forest (Rossa-Feres et al., 2017). According to Lucas

(2008), there are 122 amphibian species in the state of

Santa Catarina, 77 of which are known for the Serra

Dona Francisca region, including São Bento do Sul,

Campo Alegre, Joinville, and Corupá municipalities,

north-eastern region of the state. This number

represents 63% of all amphibian species found in the

state, which emphasizes the value of this region for

amphibian conservation. It is also worth noting that

recent species records and anuran descriptions

increased this number of species (e.g., Lucas and

Garcia, 2011, Bruschi, Lucas, Garcia and Recco-

Pimentel. 2014, Monteiro, Condez, Garcia, Comitti,

Amaral, and Haddad, 2014, Bornschein et al. 2015,

Condez et al. 2016, Monteiro, Comitti and Lingnau.

2014, Lourenço, Lingnau, Haddad and Faivovich,

2019). Although the Atlantic Forest has been greatly

reduced and fragmented (Ribeiro, Metzger,

Martensen, Ponzoni and Hirota, 2009), it harbors many

endemic species, which characterizes the biome as a

biodiversity hotspot (Myers, Mittermeier, Mittermeier,

Fonseca and Kent, 2000). Therefore, studies focusing

on tropical diversity are imperative to evaluate the

conservation status of this environment and manage

the regional species list (Verdade et al., 2012), that

adds value to a preserved patrimony.

Given the historical importance of the Serra Dona

Francisca and the lack of information for several areas

in the Atlantic Forest biome, we performed an anuran

inventory in the Serra Dona Francisca. We gathered

data on natural history for each species recorded,

describing their temporal and spatial distribution,

especially along the elevational gradient. In addition,

we compared the species composition with other

anuran communities from subtropical Atlantic Forest

Bol. Soc. Zool. Uruguay (2ª época). 2022. ISSN 2393-6940Vol. 31 (2): e31.2.4

Fig. 1. SC-301 highway

lookout located just up of

Serra Dona Francisca

showing an example of the

exuberant Atlantic Forest

regi o n ( S u b - M o n t a n e

O m b ro p h i l o u s D e n s e

Forest - Lat -26.193884º

Lon -49.035562).

3Anuran Natural History from Serra Dona Francisca

and provided comments on forest fragment

conservation and preservation efforts in this

sustainable use conservation unit.

MATERIAL AND METHODS

Study area

In the Serra Dona Francisca (Fig. 1) there is the

“Área de Proteção Ambiental Serra Dona Francisca”

(APA; Environmental Protection Area), a sustainable

use conservation unit, with 41000 hectares under the

environmental agency management FUNDEMA

(Fundação Municipal do Meio Ambiente), from Joinville

municipality, state of Santa Catarina, Brazil. The region

is characterized by geological formations of Serra

Geral and Serra do Mar mountain ranges, with the

Cubatão River as the main watershed (Scheibe, 1986,

Peluso Jr, 1991). There is no dry season in the region,

but a period of higher rainfall intensity from spring to

summer seasons was recorded (Klein, 1960). The

historical mean amplitude of temperature is

12.3–24.5ºC, and mean amplitude of precipitation is

77.7–282.3 mm (INMET, for the nearest climatic

stations, Porto União and São Francisco do Sul

municipalities, from 1931 to 1990).

Sampling

We established four sampling localities at different

elevations (100, 300, 700 and 1000 m) in the Serra

Dona Francisca, within Joinville and Campo Alegre

municipalities, with the highest elevation at 1200 m

(Table 1, Fig. 2).

Field work was carried out in 16 months, totalling

68 sampling days. The first 12 campaigns occurred

from October 2012 to March 2013, with two campaigns

each month throughout the elevation gradient

established, using the two methods described below.

The following 10 campaigns were carried out from April

2013 to January 2014, in which we performed one

campaign each month in the two categories of highest

elevation (700 and 1000 m), Araucaria Forest and

Montane Ombrophilous Forest, using only the surveys

at breeding sites method. Such change in our sampling

was due to logistic support that change through the

monitoring.

We applied two sampling methods: (1) SBS -

surveys at breeding sites, following Scott Jr and

Woodward (1994), and (2) PT - pitfall traps (Corn,

1994). The SBS method consisted of visual and

auditory surveys and counting the richness of anuran in

breeding sites either in the forest interior, streams,

swamps, or ponds. We selected at least five breeding

sites at each elevation, totalling 34 breeding sites in all

sampled area. In the forest and stream, we sampled

120 m transects, counting the anuran species that used

these habitats for reproduction through visual and

auditory surveys. Each elevation was visited for one

night in each sampling, with activities starting at dusk,

from 20:00 h to 24:00 h, totaling 272 person-hours of

sampling. The PT method consisted of three lines of

pitfall traps at each sampled elevation. The lines were

built with four 60-liter plastic buckets, all connected with

a 50 m-long and 1 m-high drift-fence. The buckets were

open for four consecutive nights in each campaign,

totalling 48 bucket-days and 4608 hours of total

sampling effort. We checked the traps every morning

while they were open, and we captured, identified,

measured, and marked all anuran found in buckets.

After these proceedings we released the animals next

to the traps where they were first captured. This method

was only applied in the first 12 campaigns of this study.

Some vouchers were collected, and the specimens

were deposited in the Zoology Scientific Collection at

the Universidade Regional de Blumenau (FURB:

22166…22952) and in the Herpetological Collection of

Ribeirão Preto (CHRP: 1387– 1390).

During fieldwork, we also obtained information on

the anurans' breeding behaviour and spatial-temporal

occupation in the environment: the habitat (forest

interior, forest edge, open area, lentic, lotic),

microhabitat (leaf litter, stream banks, shrubs, grasses,

trees, bromeliads, rivers, ponds, marshes, aquatic

vegetation, water), perch type at the reproductive

activity (bromeliads, tree, shrub, grasses, herbaceous,

soil, rocks), and activities (calling, foraging,

amplecting). The reproduction period for all the species

was categorized by the presence of ovigerous females,

amplected couples, spawning and/or tadpoles and

juveniles. We classified the species as continuous,

prolonged, or explosive breeders as proposed by

Crump (1974), Wells (1977) and Prado, Uetanabaro

and Haddad (2004). According to Prado et al. (2004),

continuous breeders are species with individuals that

reproduce throughout the year; prolonged breeders

are species with more than three months of

reproduction and asynchronous arrival of females to

breeding sites, with males showing territoriality

behaviour; and explosive breeders present

reproductive activity for only a few days or weeks with

synchronous arrival of females and males. The

reproductive modes were classified following Nunes-

de-Almeida, Haddad and Toledo (2021).

For some species, we provide bioacoustics

information to support taxonomic identification. We

registered anuran calls using a Marantz PMD 661

digital recorder (sample rate 24000) with a YOGA HT-

81 directional microphone. We analysed the recorded

calls in the Audacity software (version 2.2.1).

Ecological analyses

We performed rarefaction/extrapolation curves

based on incidence (samples in campaigns) for each

method (SBS and PT) separately. We observed the

Bol. Soc. Zool. Uruguay (2ª época). 2022. ISSN 2393-6940Vol. 31 (2): e31.2.4

Bol. Soc. Zool. Uruguay (2ª época). 2022. Vol. 31 (2): e31.2.4 ISSN 2393-6940

Anuran Natural History from Serra Dona Francisca4

Fig. 2. Sampled points (black diamonds) in Serra Dona Francisca, Joinville municipality, Brazil. The shades area repre-

sents the contour line of the sustainable use environmental protection area (Serra Dona Francisca).

5Anuran Natural History from Serra Dona Francisca

Bol. Soc. Zool. Uruguay (2ª época). 2022. ISSN 2393-6940Vol. 31 (2): e31.2.4

Table 1. Geographic coordinates (Datum: WGS 84), elevation, phytophysiognomies (Veloso et al. 2012) and relief of the four sampled

localities and their sampled breeding sites (SBS) and pitfall traps (PT) of Serra Dona Francisca, Joinville and Campo Alegre, Santa

Catarina, Brazil. Legends: DOSF - dense ombrophilous sub-montane forest, DMOF - ecotype between dense and mix ombrophilous

forest, MOFS - ecotype between mix ombrophilous forest (Araucaria Forest) and steppe vegetation.

Sampled Locality Vegetation Relief

Sampled Point Latitude Longitude elevation Method Type of Breeding Site/ Landscape

1000 MOFS upland

A15 26°12'10.56" 49°14'39.16" 970 SBS lentic permanent/forest edge

A16 26°12'11.05" 49°14'44.60" 960 SBS lentic permanent/open land

A17 26°12'8.91" 49°14'44.88" 963 SBS lentic permanent/open land

A21 26°12'57.40" 49°13'9.91" 1020 SBS lentic permanent/forest edge

A22 26°12'50.65" 49°13'13.11"1 022 SBS lentic temporary/open land

A23 26°12'47.14" 49°13'20.43" 984 SBS lentic temporary/forest interior

A24 26°13'7.05" 49°13'19.88" 917 SBS lentic temporary/open land

A25 26°12'45.81" 49°14'16.41" 1025 SBS lentic temporary/open land

A26 26°11'20.82" 49°15'3.10" 1029 SBS lentic temporary/open land

TF1000 26°12'41.19" 49°14'15.95" 992 SBS forest interior

TS1000 26°12'48.74" 49°13'18.68" 1014 SBS lotic permanent/forest interior

1000A 26°12'46.87" 49°13'14.86"1 020 PT forest interior

1000B 26°12'51.01" 49°13'11.72" 1036 PT forest interior

1000C 26°12'56.35" 49°13'9.04" 1033 PT forest interior

700 DMOF upland

B18 26°16'59.74" 49° 6'9.54" 636 SBS lentic permanent/forest edge

B19 26°16'56.81" 49° 6'9.87" 626 SBS lentic temporary/forest interior

B21 26°17'18.26" 49° 7'6.41" 613 SBS lentic temporary/open land

B22 26°17'20.82" 49° 7'4.97" 630 SBS lentic permanent/open land

B5 26°15'44.36" 49° 5'50.43" 651 SBS lentic permanent/edge forest

TF700 26°17'2.15" 49° 6'8.86" 649 SBS forest interior

TS700 26°17'0.96" 49° 6'8.28" 659 SBS lotic permanent/forest interior

700A 26°17'2.49" 49° 6'8.49" 639 PT forest interior

700B 26°17'0.21" 49° 6'9.04" 615 PT forest interior

700C 26°17'0.82" 49° 6'10.55" 626 PT forest interior

300 DOSF mountain climb

C43 26°10'13.58"48° 59'47.30" 118 SBS lentic permanent/edge forest

C48 26°10'40.78" 49° 0'22.17" 177 SBS lentic temporary/edge forest

C49 26°10'32.11" 49° 0'23.11" 165 SBS lentic permanent/open land

TF300 26°10'44.78" 49° 0'24.24" 148 SBS forest interior

TS300 26°10'48.69" 49° 0'20.24" 259 SBS lotic permanent/forest interior

300A 26°10'47.11" 49° 0'20.31" 219 PT forest interior

300B 26°10'52.50" 49° 0'20.03" 242 PT forest interior

300C 26°10'55.45" 49° 0'18.27" 262 PT forest interior

100 DOSF lowland

C1 26° 8'34.76" 49° 0'21.68" 85 SBS lentic permanent/edge forest

C24 26° 8'59.50" 48°59'33.91" 74 SBS lentic temporary/edge forest

C25 26° 8'58.92" 48°59'32.13" 72 SBS lentic permanent/open land

C28 26° 9'5.08" 48°59'27.88" 75 SBS lentic permanent/open land

C29 26° 9'2.85" 48°59'27.07" 77 SBS lentic temporary/open land

MARIOTTO, L.R. et al.

asymptote of the curves (Chao's richness index)

comparing the observed richness and estimated

richness by Hill numbers (order q=0) (the endpoint of

both methods for extrapolation was 40 campaigns)

(Hsieh, Ma and Chao, 2016). To compare the species

diversity among sampled localities (100, 300, 700 and

1000 m) per method (SBS and PT), we estimated the

richness (Hill number q=0), Shannon diversity (Hill

number q=1) and Simpson diversity (Hill number q=2)

using rarefaction/extrapolation curves based on

abundance (the standard function remained, which

was twice the sample size). Only for SBS method, we

used, as the value of abundance, the sum of the higher

abundance recorded for each species in each breeding

site during sampling campaigns (Table 2). Since the

differences of sampling efforts (number of breeding

sites) among sampled localities might influence the

species diversity comparisons, we plotted sample

completeness curves, which is a comparison of point

diversities for some sample size using the same

rarefaction/extrapolation curves described above. We

used the iNEXT package in R environment for such

analyses (Hsieh et al., 2016, R Core Team, 2020).

We compared the species composition of anuran

communities of Serra Dona Francisca with other

anuran communities from subtropical Atlantic Forest

(see Table 1 in Suppl. Info. 1) using cluster analysis.

We generated a similarity matrix of species

composition for each locality through the Jaccard

index (presence-absence) using vegan package

(Oksanen et al., 2019). To check whether the

geographical distance influenced species

composition, we search for spatial autocorrelation

between species composition (the same similarity

matrix used in the cluster analysis) and geographical

distance matrix between samples (matrix of latitudinal

and longitudinal coordinates of each sampled locality)

using the Mantel test (vegan package, Oksanen et al.,

2019). All analyses were performed using the R

language (R Core Team, 2020).

RESULTS AND DISCUSSION

1. Anuran Commented List

We found 63 anuran species sampled by both SBS

and PT methods, in all four elevational localities (Table

2, Fig. 3–8). These species belong to 13 families

(number of species in parenthesis): Brachycephalidae

(4), Bufonidae (4), Centrolenidae (1), Craugastoridae

(1), Cycloramphidae (2), Hemiphractidae (2), Hylidae

(30), Hylodidae (2), Leptodactylidae (10),

Mi c r oh y l ida e (2 ) , Od ont o p hr y n ida e (3 ) ,

Phyllomedusidae (1), and Ranidae (1).

Family Brachycephalidae

Brachycephalus sulfuratus

Condez, Monteiro, Comitti, Garcia, Amaral and

Haddad, 2016

(Fig. 3A)

Vouchers - FURB 22950, 22952

Taxonomy and distribution - This is a recently

described species endemic to the Atlantic Forest

(Condez et al., 2016). Brachycephalus sulfuratus

belongs to the B. didactylus species group and occurs

in São Paulo, Paraná, and Santa Catarina states. It is

apparently widely distributed, occurring from the sea

level up to 1000 m (Condez et al., 2016).

Ecology - We did not observe any behavior of this

species in the field, once two individuals were

captured by PT and only one individual was collected

on a shrub in the forest interior during the SBS, all at

700 m altitude. The Brachycephalus' species present

endotrophic and direct development of their eggs

(Hedges, Duellman and Heinicke, 2008). They deposit

the eggs in the leaf litter, tree barks and on the soil

(reproductive mode 27 – Nunes-de-Almeida et al.,

2021). Condez et al. (2016) observed this species

calling under the leaf-litter on rainy days with

reproductive activity throughout the year.

Bol. Soc. Zool. Uruguay (2ª época). 2022. ISSN 2393-6940Vol. 31 (2): e31.2.4

Table 1. Cont.

Sampled Locality Vegetation Relief

Sampled Point Latitude Longitude elevation Method Type of Breeding Site/ Landscape

C30 26° 9'2.59" 48°59'30.09" 73 SBS lentic temporary/edge forest

TF100 26° 9'12.26" 48°59'36.87" 99 SBS forest interior

TS100 26° 9'8.83" 48°59'36.02" 100 SBS lotic permanent/forest interior

100A 26° 9'8.93" 48°59'32.63" 97 PT forest interior

100B 26° 9'10.31" 48°59'34.13" 100 PT forest interior

100C 26° 9'10.89" 48°59'35.88" 100 PT forest interior

7Anuran Natural History from Serra Dona Francisca

Bol. Soc. Zool. Uruguay (2ª época). 2022. ISSN 2393-6940Vol. 31 (2): e31.2.4

Table 2. Amphibians' species and abundance registered in the four elevation localities of Serra Dona Francisca, Brazil. The two

sampling methods: SBS - survey at breeding sites - from October 2012 to January 2014, and PT - pitfall traps - from October 2012

to March 2013.

Taxa/Elevation localities/Methods 100 300 700 1000

SBS PT SBS PT SBS PT SBS PT

Class Amphibia - Order Anura

Brachycephalidae (2 genera, 4 species)

Brachycephalus sulfuratus Condez, Monteiro,

Comitti, Garcia, Amaral and Haddad, 2016 - - - - 1 2 - -

Ischnocnema sp. (gr. guentheri) 1 - 8 - - - - -

Ischnocnema henselii (Peters, 1872) - - - - 10 1 6 -

Ischnocnema sp. (I. lactae species series) - - 8- - - - -

Bufonidae (3 genera, 4 species)

Dendrophryniscus berthalutzae Izecksohn, 1994 - - - - 11 - - -

Rhinella ornata (Spix, 1824) 5 39 48 28 24 17 9 4

Rhinella icterica (Spix, 1824) 2 1 - - 20 1 66 18

Melanophryniscus xanthostomus Baldo, Bornschein,

Pie, Ribeiro, Firkowski and Morato, 2015 - - - - - - 4-

Centrolenidae (1 genus, 1 species)

Vitreorana uranoscopa (Müller, 1924) - - - - 4 - 15 -

Craugastoridae (1 genus, 1 species)

Haddadus binotatus (Spix, 1824) - - 2 - - - - -

Cycloramphidae (1 genera, 2 species)

Cycloramphus bolitoglossus (Werner, 1897) - - - - 1 3 1 -

Cycloramphus izecksohni Heyer, 1983 - - 9 - - - - -

Hemiphractidae (1 genus, 2 species)

Fritziana mitus Walker, Wachlevski, Nogueira-Costa,

Garcia and Haddad, 201811-45-60---Fritziana sp. - - - - - - 17 -

Hylidae (8 genera, 30 species)

Aplastodiscus sp. - - - - 29 - 41 -

Aplastodiscus ehrhardti (Müller, 1924) - - - - 9 - - -

Aplastodiscus perviridis A. Lutz in B. Lutz, 1950 - - - - - - 20 -

Boana albomarginata (Spix, 1824) 10 - - - 7 - - -

Boana albopunctata (Spix, 1824) - - - - - - 59 -

Boana bischoffi (Boulenger, 1887) - - 8 - 86 - 101 -

Boana faber (Wied-Neuwied, 1821) 9 - 3 - 23 - 58 -

Boana semilineata (Spix, 1824) 15 - 1 - - - - -

Boana semiguttata (Lutz, 1925) - - - - - - 10 -

Bokermannohyla hylax (Heyer, 1985) 6 - 7 - 12 - - -

Bokermannohyla luctuosa Pombal and Haddad, 1993 - - - - 17 - 14 -

Dendropsophus berthalutzae (Bokermann, 1962) - - 20 - - - - -

Dendropsophus elegans (Wied-Neuwied, 1824) 79 - 13 - - - - -

Dedropsophus microps (Peter, 1872) - - - - 14 - 11 -

Dendropsophus minutus (Peters, 1872) - - 1 - 16 - 465 -

Dendropsophus nahdereri (B. Lutz and Bokermann, 1963) - - - - 22 - 20 -

Dendropsophus werneri (Cochran, 1952) 66 - 19 - 29 - - -

Scinax aromothyella Faivovich, 2005 - - - - - - 30 -

Scinax catharinae (Boulenger, 1888) - - - - - - 2 -

MARIOTTO, L.R. et al.

Bol. Soc. Zool. Uruguay (2ª época). 2022. ISSN 2393-6940Vol. 31 (2): e31.2.4

Table 2. Cont.

Taxa/Elevation localities/Methods 100 300 700 1000

SBS PT SBS PT SBS PT SBS PT

Scinax aff. perpusillus - - 1 - 9 - - -

Scinax litorallis (Pombal and Gordo, 1991) 19 - 7 - - - - -

Scinax rizibilis (Bokermann, 1964) 2 - 1 - 9 - 3 -

Scinax fuscovarius (A. Lutz, 1925) 19 - - - 14 - 1 -

Scinax granulatus (Peters, 1871) - - - - - - 38 -

Scinax imbegue Nunes, Kwet and Pombal, 2012 110 - 25 - 54 - - -

Scinax perereca Pombal, Haddad and Kasahara, 1995 13 - 22 - 65 - 101 -

Scinax squalirostris (A. Lutz, 1925) - - - - - - 30 -

Scinax tymbamirim Nunes, Kwet and Pombal, 2012 53 - - - - - - -

Sphaenorhynchus surdus (Cochran, 1953) - - - - - - 112 -

Trachycephalus mesophaeus (Hensel, 1867) 27 - 4 - 9 - - -

Hylodidae (2 genera, 2 species)

Crossodactylus sp.- - - - 36 - - -

Hylodes perplicatus (Miranda-Ribeiro, 1926) - - 7 6 1 6 - -

Leptodactylidae (4 genera, 10 species)

Adenomera araucaria (Kwet and Angulo, 2002) - - - - 15 - 19 6

Adenomera nana Müller, 1922 7 12 4 25 27 23 22 -

Leptodactylus paranaru Magalhães, Lyra, Carvalho, Baldo,

Brusquetti, Burella, Colli, Gehara, Giaretta, Haddad, Langone,

López, Napoli, Santana, Sá and Garda, 2020 11 - 2 - 15 - 31 5

Leptodactylus notoaktites Heyer, 1978 21 9 8 2 16 2 7 -

Leptodactylus plaumanni Ahl, 1936 - - - - - - 26 2

Physalaemus aff. gracilis - - - - - - 50 13

Physalaemus cuvieri Fitzinger, 1826 22 4 - - 2 - 42 8

Physalaemus lateristriga (Steindachner, 1864) 1 - - 5 36 156 50 393

Physalaemus nanus (Boulenger, 1888) - - - - - - 17 4

Scythrophrys sawayae (Cochran, 1953) - - - 1 17 61 - -

Microhylidae (2 genera, 2 species)

Chiasmocleis leucosticta (Boulenger, 1888) 3 26 1 1 - 2 4 -

Elachistocleis bicolor (Valenciennes in Guérin-Menéville, 1838) 33 1 - - - - 22 1

Odontophrynidae (2 genera, 3 species)

Odontophrynus americanus (Duméril and Bibron, 1841) - - - - - - 4 -

Proceratophrys boiei (Wied-Neuwied, 1825) - 1 5 9 49 79 13 2

Proceratophrys subguttata Izecksohn, Cruz and Peixoto, 1999 - - - - - 2 - -

Phyllomedusidae (1 genus, 1 species)

Phyllomedusa disticta A. Lutz in B. Lutz, 1950 - - - - 2 - 4 -

Ranidae (1 genus, 1 species)

Lithobates catesbeianus (Shaw, 1802) - 1 - - - - - -

Total Richness 24 9 27 8 37 14 38 11

Bol. Soc. Zool. Uruguay (2ª época). 2022. Vol. 31 (2): e31.2.4 ISSN 2393-6940

MARIOTTO, L.R. et al. 9

Fig. 3. Some amphibians registered in Serra Dona Francisca: (A) Brachycephalus sulfuratus, (B) Ischnocnema henselii, (C)

Ischnocnema sp. (gr. Guentheri), (D) Ischnocnema sp. (I. lactae species series), I Dendrophryniscus berthalutzae, (F) Rhine-

lla ornata, (G) Rhinella icterica, (H) Melanophryniscus xanthostomus, (I) Vitreorana uranoscopa, (J) Haddadus binotatus, (K)

Cycloramphus bolitoglossus, and (L) Cycloramphus izecksohni.

Ischnocnema henselii

(Peters, 1870)

(Fig. 3B)

Vouchers - FURB 22375, 22390, 22719, 22730, 22738,

22739

Taxonomy and distribution - This species was

recently placed within the Ischnocnema guentheri

species group, and it is widely distributed in the south

portion of Atlantic Forest in the São Paulo, Paraná,

Santa Catarina, and Rio Grande do Sul states, and in

the northeast of Argentina (Conte et al., 2010, Gehara,

Canedo, Haddad and Vences, 2013).

Ecology - We registered males of I. henselii at the

altitude levels of 700 m and 1000 m, foraging and

calling in the leaf litter or on vegetation such as shrubs,

grasses, or trees, in the forest interior and in stream

banks. We observed males emitting advertisement

calls from October 2012 to January 2013 and

September 2013 to December 2013, and registered

ovigerous females in March 2013, December 2013 and

January 2014. Species of the genus Ischnocnema

have reproductive mode 27, with direct development of

the terrestrial eggs (Nunes-de-Almeida et al., 2021).

Ischnocnema sp. (gr. guentheri)

(Fig. 3C)

Vouchers - FURB 22283, 22374, 22380, 22795, 22812,

22913

Taxonomy and distribution - This species is part of

the I. guentheri species complex, cited as Candidate

Species 1 (CS1) in Gehara et al. (2013), which was

delimited by using molecular markers and bioacoustics

data. This putative species occurs from the states of

São Paulo to Santa Catarina (Gehara et al., 2013). The

calls (n=6 from two males) have similar acoustic

parameters if compared with Gehara et al. (2013), as

follows: dominant frequency 2325–2497 Hz (2403.833

± 63.294), call duration 1.880–2.440 s (2.118 ± 0.184)

and notes per call 29–35 (31.5 ± 1.974).

Ecology - We registered this species only at low

elevations (100 and 300 m). We observed calling males

in the leaf litter, at the stream banks or on the vegetation

in the forest interior, from October 2012 to March 2013;

we registered ovigerous females during November and

December 2012, and from January to March 2013.

Species of the genus Ischnocnema have reproductive

mode 27 (Nunes-de-Almeida et al., 2021).

Ischnocnema sp. (I. lactea species series)

(Fig. 3D)

Vouchers - FURB 22748, 22758, 22907, 22929

Taxonomy and distribution - This species was

assigned as part of the I. lactea species series based

on external morphology. There are three species

belonging to this species series occurring close of the

studied region: I. paranaensis, I. manezinho and I.

sambaqui. The collected individuals (adults, males)

differ from I. paranaensis by their larger size (SVL

21.3–21.8 mm in Ischnocnema sp.; 17.7 mm in I.

paranaensis, Langone and Segalla, 1996) and by the

presence of external tympanum (indistinct in I.

paranaensis). Ischnocnema sp. presents variegated

dorsal coloration that varies from brown to yellow;

thigh, shank and forearms with black stripes on dorsal

surfaces. The snout is round from dorsal view and

acuminate in profile. Because of the presence of the

internal tarsal tubercle, collected individuals are similar

to I. sambaqui, and differs from I. manezinho (internal

tarsal tubercle absent) (Garcia, 1996, Castanho et al.,

2000). The advertisement calls (n = 12, two males)

have duration of 0.140–0.270 s (0.207 ± 0.029), with

3.000–5.000 notes per call (3.916 ± 0.288), note

duration of 0.008–0.010 s (0.010 ± 0.001), and only one

pulse per note. The dominant frequency is

3014.000–3100.000 Hz (3071.333 ± 38.167), Fmin

1765.000–2282.000 Hz (2095.583 ± 130.311), and

Fmax 3875.000–4392.000 Hz (4083.750 ± 150.020).

The advertisement call of I. paranaensis is not

described, hampering any comparisons. However, we

observed differences in the call duration and number of

notes in comparison with I. manezinho (respectively,

0.79–0.97 s and 11–13 number of notes, Florianópolis

municipality, Santa Catarina state, approximately 160

km from Serra Dona Francisca, Castanho et al. 2000)

and I. sambaqui (respectively, 0.50–0.63 s and 6.0–8.0

number of notes, Guaraqueçaba municipality, Paraná

state, approximately 130 km from Serra Dona

Francisca, Castanho et al. 2000). Therefore, because

of the reasons explained above, we were not able to

assign the collected individuals to any of these three

species.

Ecology - We observed males of Ischnocnema sp.

calling during November 2013 in the forest interior, only

at 300 m of elevation level. Males of this species use

large rocks (more than 1 m from the soil) as calling

sites, besides trunks wrapped by mosses and

bryophytes. Species of the Ischnocnema genus have

the reproductive mode 27 (Nunes-de-Almeida et al.,

2021).

Family Bufonidae

Dendrophryniscus berthalutzae

Izecksohn, 1994

(Fig. 3E)

Vouchers - FURB 22282, 22474, 22720, 22901, 22908

Taxonomy and distribution - This species occurs in

coastal forests in the Paraná and Santa Catarina states

(Fusinatto, Cruz and Garcia, 2008) and belongs to the

D. brevipollicatus species group (Izecksohn, 1993).

Ecology - We registered D. berthalutzae only at the

elevation level of 700 m, foraging on the vegetation and

rocks in the forest interior. We observed calling males

on shrubs in the streams (1.5 m high), only in

November 2012, but we found females during all the

survey, and juveniles only in February 2013. This

species is usually found in bromeliads, where the eggs

are laid (reproductive mode 4 – Peixoto, 1995, Nunes-

de-Almeida et al., 2021).

Bol. Soc. Zool. Uruguay (2ª época). 2022. ISSN 2393-6940Vol. 31 (2): e31.2.4

MARIOTTO, L.R. et al.

Rhinella ornata

(Spix, 1824)

(Fig. 3F)

Vouchers - FURB 22166-22168, 22170-22175

Taxonomy and distribution - This species is widely

distributed from the south of Minas Gerais and Rio de

Janeiro states to the south of the state of Santa

Catarina, and from the east littoral to the west in

Argentina. It belongs to the R. crucifer species group

(Pereyra et al., 2021).

Ecology - We found individuals of R. ornata at all

elevation levels and within all sampled habitats: open

areas, forest edges and in the forest interior. We

observed calling males from October 2012 to January

2013 and from August to October 2013 in open areas

and forest edges. We captured ovigerous females

during the period of vocalization. We registered

amplected couples only at the end of October 2012.

Out of the breeding period, we captured mostly females

and juveniles in the forest interior (PT method), from

October 2012 to March 2013. This species presents the

reproductive mode 1, with eggs and exotrophic

tadpoles in lentic habitats (Nunes-de-Almeida et al.,

2021).

Rhinella icterica

(Spix, 1824)

(Fig. 3G)

Vouchers - FURB 22176, 22177, 22201, CHRP 1387,

1388

Taxonomy and distribution - The cururu toad is

widely distributed in the Atlantic Forest of the south and

southeast of Brazil and belongs to the R. marina

species group (Pereyra et al., 2021).

Ecology - We found R. icterica at all elevation levels

and observed this species in breeding activity in open

areas and in forest edges, but we also collected some

individuals in the forest interior (PT and SBS). We

observed calling males and adult females from October

2012 to January 2013, and amplected couples only at

the end of October 2012. We captured some juveniles

from October 2012 to March 2013. This species

presents the reproductive mode 1 (Nunes-de-Almeida

et al., 2021).

Melanophryniscus xanthostomus

Baldo, Bornschein, Pie, Ribeiro, Firkowski and Morato,

2015

(Fig. 3H)

Vouchers - FURB 22713, 22714, 22822, 22851

Taxonomy and distribution - Melanophryniscus

xanthostomus is a recently described species and it is

not assigned to any species group (Bornschein et al.,

2015). This species is distributed on top of mountains of

northeaster of the state of Santa Catarina (Bornschein

et al., 2015).

Ecology - We observed calling males inside

terrestrial bromeliads in the forest interior from

November 2013 to January 2014, only at the elevation

level of 1000 m. We found eggs, tadpoles and one

female near a pitfall trap fence in November 2013. As

well as reported for other species of Melanophryniscus

with this breeding behaviour, this species has the

reproductive mode 4 (Nunes-de-Almeida et al., 2021).

Family Centrolenidae

Vitreorana uranoscopa

(Müller, 1924)

(Fig. 3I)

Vouchers - FURB 22462, 22813

Taxonomy and distribution - This species inhabits

streams of the Atlantic Forest from the southeast to the

south of Brazil (Heyer, 1985a, Machado et al., 2010).

Vitreorana uranoscopa represents a species complex

with several distinct lineages (Paz et al., 2018).

Bokermann (1966) assigned Corupá municipality, state

of Santa Catarina, as the type locality of this species,

and given the proximity of the studied area (less than

15 km of airline distance from Serra Dona Francisca),

we considered this population as the nominal species

(the green dot lineage from Paz et al., 2018).

Ecology - We collected this species at the elevation

levels of 700 m and 1000 m. We observed calling males

on xaxins (Dicksonia sellowiana), palms and other

shrub leaves (up to 1 m high) in streams of the forest

interior, from November 2012 to February 2013, and

from September 2013 to January 2014; we registered

spawning on leaves pending above streams during

December 2013. This species has reproductive mode

22, with arboreal eggs hatching into exotrophic

tadpoles that drop in lotic water (Nunes-de-Almeida et

al., 2021).

Family Craugastoridae

Haddadus binotatus

(Spix, 1824)

(Fig. 3J)

Vouchers - FURB 22235, 22487, 22253

Taxonomy and distribution - This species is widely

distributed throughout the Atlantic Forest, from the

south of Bahia to Rio Grande do Sul states (Kwet,

Lingnau and Di-Bernardo, 2010, Dias, Lourenço-de-

Moraes and Solé, 2012). There is a record for Mata de

São João, state of Bahia (Brazil), that could be the

northern record for this species (Amaro, Nunes,

Canedo, Napoli and Juncá, 2013, Napoli, Menezes

and Abreu, 2017). However, individuals of H. binotatus

present some morphological differences that can

indicate a possible species complex (Amaro et al.,

2013, Napoli et al., 2017).

Ecology - We found this species only at the

elevation level of 300 m, during the SBS in the forest

interior. We have not observed individuals calling or

breeding, only in foraging activity. Haddadus binotatus

presents direct development and deposits the eggs on

the leaf litter in the forest (reproductive mode 27,

Bol. Soc. Zool. Uruguay (2ª época). 2022. ISSN 2393-6940Vol. 31 (2): e31.2.4

11 Anuran Natural History from Serra Dona Francisca

Hedges, Duellman and Heinicke, 2008, Nunes-de-

Almeida et al., 2021).

Family Cycloramphidae

Cycloramphus bolitoglossus

(Werner, 1897)

(Fig. 3K)

Vouchers - FURB 22244, 22765, 22870

Taxonomy and distribution - This species occurs in

the Atlantic Forest of Paraná and Santa Catarina states

(Heyer, 1983, Conte and Rossa-Feres, 2007), and

belongs to the C. bolitoglossus species group

(Lingnau, Solé, Dallacorte and Kwet, 2008).

Ecology - We found this species at the elevation

levels of 700 m and 1000 m. We observed several

calling males after an intense rainfall in November

2013. We also found calling males in October and

November 2012, always under the leaf litter in the

forest edge and in the forest interior. Cycloramphus

bolitoglossus breeds in the humid forest soil (Lingnau

et al. 2008). There are some controversies about the

breeding behavior of this species (see Heyer, 1983);

however, it is supposed that its reproductive mode is

similar to C. stejnegeri (Noble, 1924), which consists to

terrestrial non-froth eggs laid in constructed burrow

(reproductive mode 41, Lingnau et al. 2008, Nunes-de-

Almeida et al., 2021).

Cycloramphus izecksohni

Heyer, 1983

(Fig. 3L)

Vouchers - FURB 22646, 22712, 22882

Taxonomy and distribution - This species occurs in

Serra do Mar mountain range throughout the states of

São Paulo, Paraná and Santa Catarina (Frost, 2021),

and it is included in the C. fuliginosus species group

(Verdade, 2005).

Ecology - We found C. izecksohni in hillside

streams in the forest interior all over the region at the

elevation level of 300 m. We observed calling males on

rocks, trunks, and on the soil at the stream banks, from

November 2012 to March 2013, and juveniles from

January to March 2013; females were not recorded.

Lingnau et al. (2008) reported the presence of some

females with eggs in December 2004 and January

2006, and tadpoles in the small waterfalls at the same

locality of the present study. This species has

reproductive mode 17 (Nunes-de-Almeida et al.,

2021), which is non-froth eggs laid on rock nest in lotic

water.

Family Hemiphractidae

Fritziana mitus

Walker, Wachlevski, Nogueira-Costa, Garcia and

Haddad, 2018

(Fig. 4A)

Vouchers - FURB 22489, 22922

Taxonomy and distribution - This genus is endemic

to the Atlantic Forest, and the species are popularly

named marsupial treefrogs, because females carry

fertilized eggs in the dorsum (Duellman, Jungfer and

Blackburn, 2011). Fritziana mitus is distributed along

the Atlantic Forest, from northern of the state of São

Paulo state to northern of the state of Rio Grande do

Sul (Walker et al., 2018).

Ecology - We observed calling males calling in the

canopy and understory in the forest interior, on

bromeliads and epiphytic plants, at the elevation levels

of 100 m to 700 m. The period of vocalization

comprised the rainy season, from October 2012 to

March 2013, June 2013 and October 2013 to January

2014. We have not found female carrying eggs of this

species; however, it is known that females carry the

eggs in a dorsal pouch, and after hatching, the tadpoles

are deposited in the water-filled axils of bromeliads

(reproductive mode 67, Nunes-de-Almeida et al.,

2021).

Fritziana sp.

(Fig. 4B)

Vouchers - FURB 22749, 22881, 22926

Taxonomy and distribution - We found specimens of

Fritziana sp. only in the Araucaria Forest, which

represents the first record of the genus in this

vegetation type. We cannot relate these specimens to

any other Fritziana species because of differences in

body size and colour patterns, as follows: individuals of

Fritziana sp. are larger (SVL 23.4–25.3 mm) than

Fritziana mitus (SVL 16.2–23.7 mm, Walker et al.,

2018), presents a subovoid snout in dorsal view and

slightly acute in profile, and yellowish light orange

background with dark spots evenly distributed on

dorsum.

Ecology - We observed calling males calling from

the canopy to the ground, on bromeliads and epiphytic

plants in the forest interior, only at the elevation level of

1000 m. We highlight that there is an allopatric

distribution between F. mitus, distributed only in sub-

montane and montane ombrophilous Atlantic Forest,

and Fritziana sp., occurring only in Araucaria Forest.

This pattern may be related to climatic conditions

associated to altitudinal gradient. The period of

vocalization occurred in November and December

2013. Females of this genus carry the eggs in a dorsal

pouch and after hatching, the tadpoles are deposited in

the water-filled axils of bromeliads (reproductive mode

67, Peixoto 1995, Nunes-de-Almeida et al., 2021); we

did not find any female carrying eggs.

Family Hylidae

Aplastodiscus sp.

(Fig. 4C)

Vouchers - FURB 22306, 22461, 22463, 22478, 22818

Taxonomy and distribution - The canebrake

treefrogs are distributed mostly in the Atlantic Forest,

from São Paulo to Santa Catarina states (Berneck et

al., 2016). The collected individuals belong to the A.

albosignatus species group, and the last available

Bol. Soc. Zool. Uruguay (2ª época). 2022. ISSN 2393-6940Vol. 31 (2): e31.2.4

MARIOTTO, L.R. et al.

Bol. Soc. Zool. Uruguay (2ª época). 2022. Vol. 31 (2): e31.2.4 ISSN 2393-6940

MARIOTTO, L.R. et al. 13

Fig. 4. Some amphibians registered in Serra Dona Francisca: (A) Fritziana mitus, (B) Fritziana sp. (photo by Carlos Eduardo

Conte), (C) Aplastodiscus sp., (D) Aplastodiscus erhardti, (E) Aplastodiscus perviridis, (F) Boana albomarginata, (G) Boana

albopunctata, (H) Boana bischoffi (I) Boana faber, (J) Boana semilineata, (K) Boana semiguttata, and (L) Bokermannohyla

hylax.

phylogenetic hypothesis divided this species group

under six clades, in which the population recorded in

the present study was recognized as an undescribed

species (Aplastodiscus sp. 5, Berneck et al., 2016).

Ecology - We observed males calling from October

2012 to April 2013, and from July 2013 to January

2014, in the swamps at the forest edges and in the

forest interior, at the elevation level of 700 m. We did not

observe spawning of Aplastodiscus sp. but species of

this genus lay their eggs in underground cavities, and

after developed, tadpoles head towards a body of

water (reproductive mode 10, Nunes-de-Almeida et al.,

2021).

Aplastodiscus ehrhardti

(Müller, 1924)

(Fig. 4D)

Vouchers - FURB 22248, 22645

Taxonomy and distribution - This species occurs

from the south of the state of São Paulo, in the

municipality of Paranapiacaba, to the south coast of the

state of Santa Catarina (Conte, Lingnau and Kwet,

2005, Conte and Rossa-Feres, 2006). It belongs to the

A. albofrenatus species group (Berneck et al., 2016).

Ecology - We observed males calling from October

2012 to January 2013 and July 2013 to January 2014,

only in the forest interior, on taquaras, bromeliads and

cipós, at the elevation level of 700 m. We did not find

ovigerous females and amplected couples. As

mentioned for Aplastodiscus sp. this genus usually

presents the reproductive mode 10 (Nunes-de-

Almeida et al., 2021).

Aplastodiscus perviridis

Lutz, 1950

(Fig. 4E)

Vouchers - FURB 22236, 22285, 22307

Taxonomy and distribution - This species is widely

distributed, occurring from southeastern Brazil to the

state of Rio Grande do Sul, and north-eastern

Argentina (Frost, 2021). It is part of the A. perviridis

species group (Berneck et al., 2016).

Ecology - We recorded individuals of A. perviridis

only at the elevation level of 1000 m. We observed

males calling in open swamps and dams at the forest

edge, from October 2012 to February 2013, and from

November 2013 to January 2014. We did not find

ovigerous females or amplected couples.

Aplastodiscus perviridis reproduces in swamps or

streams and deposits the eggs in underground

cavities; after reaching advanced stages, the tadpoles

follow to adjacent water bodies (reproductive mode 10,

Nunes-de-Almeida et al., 2021).

Boana albomarginata

(Spix, 1824)

(Fig. 4G)

Voucher - FURB 22250

Taxonomy and distribution - This species occurs in

the Atlantic Forest of eastern Brazil, from state of

Paraíba to state of Santa Catarina (Frost, 2021), and

belongs to the B. faber species group (Faivovich et al.,

2005).

Ecology - We found B. albomarginata at the

elevation levels of 100 m and 700 m, occupying

herbaceous and shrubby vegetation of dams and

swamps in open areas and forest edges. We observed

males calling from October 2012 to March 2013 and

December 2013, and amplected couples in December

2012. This species deposits its eggs on the water

surface; the tadpoles are exotrophic (reproductive

mode 1, Giasson and Haddad, 2007, Nunes-de-

Almeida et al., 2021).

Boana albopunctata

(Spix, 1824)

(Fig. 4G)

Voucher - FURB 22240

Taxonomy and distribution - This species is widely

distributed in open areas in the midwest and south of

Brazil (Frost, 2021), and belongs to the B. albopunctata

species group (Faivovich et al., 2005).

Ecology - We found this species only in dams in

open areas or in the edge of the forest, at the elevation

level of 1000 m. We observed males calling perched on

grasses at the edge or inside the water body, from

October 2012 to February 2013, and from December

2013 to January 2014. We did not observe females or

spawning of B. albopunctata. This species lays eggs

directly on the water surface (reproductive mode 1 -

Muniz et al., 2008, Nunes-de-Almeida et al., 2021).

Boana bischoffi

(Boulenger, 1887)

(Fig. 4H)

Vouchers - FURB 22247, 22254, 22247,22464, 22480,

22745, 22861

Taxonomy and distribution - This species is widely

distributed in the Atlantic Forest, from the state of São

Paulo to the state of Rio Grande do Sul (Cochran,

1955, Kwet et al., 2010), and belongs to the B. pulchella

species group (Faivovich et al., 2021).

Ecology - We found individuals of B. bischoffi

foraging in both lotic and lentic environments, and

males calling only in lentic water, in dams from open

areas or on forest edges, at all elevation levels. Calling

males were observed during all the survey (specially

call type 1 - Hyla multilineata - sensu Heyer et al.,

1990), except in May 2013. Males emitted the call type

2 (sensu Heyer et al., 1990) only during the hotter

months from October 2012 to March 2013, when we

also found ovigerous females. We observed juveniles

some weeks after the beginning of the reproductive

period, at the end of November. We recorded

individuals laying their eggs on the water surface of

lentic environments. This species presents exotrophic

tadpoles (reproductive mode 1, Kwet et al., 2010,

Nunes-de-Almeida et al., 2021).

Bol. Soc. Zool. Uruguay (2ª época). 2022. ISSN 2393-6940Vol. 31 (2): e31.2.4

MARIOTTO, L.R. et al.

Boana faber

(Wied-Neuwied, 1821)

(Fig. 4I)

Vouchers - FURB 22178, 22179, 22180

Taxonomy and distribution - This species is widely

distributed in Brazil and occurs mainly in the Atlantic

Forest, from the state of Rio Grande do Norte to the

state of Rio Grande do Sul. It also occurs in adjacent

formations, such as the Cerrado, and in Paraguay and

Argentina (Lavilla, Aquino, Kwet and Baldo, 2010). The

species belongs to the Boana faber species group

(Faivovich et al., 2005).

Ecology - We found individuals reproducing in dams

and swamps in open areas and forest edges at all

sampled elevation levels. We observed males calling

from October 2012 to March 2013, and from

September to December 2013; the reproduction

events occurred in the same period as the calling, once

we observed clay nests with spawns in the breeding

sites (reproductive mode 12, Nunes-de-Almeida et al.,

2021). In December 2012 and January 2013, and in the

same period of 2013 and 2014, we observed many

females and amplected couples; juveniles were

observed some weeks later.

Boana semilineata

(Spix, 1824)

(Fig. 4J)

Vouchers - FURB 22234, 22300, 22302, 22760

Taxonomy and distribution - This species occurs in

the Atlantic Forest from the state of Pernambuco to the

state of Santa Catarina (Van Sluys and Rocha, 2010),

and belongs to the B. semilineata species group

(Faivovich et al., 2005).

Ecology - We found B. semilineata in dams and

swamps in open areas and forest edges at the

elevation levels of 100 m and 300 m. We observed

males calling from October 2012 to January 2013,

using taboa as vocalization site. We did not find

females, juveniles, or spawning. However, it is known

that this species deposits its eggs on the water surface

of lentic and lotic environments; the tadpoles are

exotrophic (reproductive mode 16, Nunes-de-Almeida

et al., 2021).

Boana semiguttata

(Lutz, 1925)

(Fig. 4K)

Vouchers - FURB 22313, 22479, 22880

Taxonomy and distribution - This species belongs to

the Boana pulchella species group. Since populations

from Joinville and São Bento do Sul municipalities were

previously assigned as B. semiguttata (Faivovich et al.,

2020), we identified the population from Campo Alegre

municipality as B. semiguttata.

Ecology - We found males of B. semiguttata calling

only in December 2012 and September 2013 to

January 2014 in streams at forest edge at the elevation

level of 1000 m; some individuals were observed

foraging during other periods. We did not observe

ovigerous or spawning. The species of the B. pulchella

group deposit their eggs on the water surface, and

present exotrophic tadpoles (reproductive mode 16,

Garcia, Faivovich and Haddad, 2007, Kwet, 2008,

Nunes-de-Almeida et al., 2021).

Bokermannohyla hylax

(Heyer, 1985)

(Fig. 4L)

Vouchers - FURB 22241, CHRP 1389, 1390

Taxonomy and distribution - The forest treefrog B.

hylax occurs from the south of the state of Rio de

Janeiro (Napoli and Pimenta, 2009) to the south of the

state of Santa Catarina (Napoli and Pimenta, 2009),

restricted to the coast region and the mountain range

(Armstrong and Conte, 2010, Lucas, 2008). This

species belongs to the B. circumdata species group

(Heyer, 1985b).

Ecology - We found B. hylax at the elevation levels

of 100 m, 300 m, and 700 m, in swamps and rivulets in

the interior of the forest. We observed males calling

from October 2012 to March 2013, and from

September 2013 to January 2014. We did not observe

any reproductive behaviour other than calling males.

This species reproduces in streams, lays eggs on the

water surface, and presents exotrophic tadpoles

(reproductive modes 12, Nunes-de-Almeida et al.,

2021).

Bokermannohyla luctuosa

Pombal and Haddad, 1993

(Fig. 5A)

Vouchers - FURB 22274, 22275, 22189, 22190

Taxonomy and distribution - Bokermannohyla

luctuosa occurs in south-eastern Brazil (Napoli, 2005).

This is the first record of this species in southern Brazil.

Despite the very similar morphology between B.

circumdata and B. luctuosa, we were able to see the

subtle difference in the tubercle of finger IV, which differ

these two species (Fig. 9) (Pombal and Haddad, 1993).

Ecology - We found males of this species on the

edge and inside the forest, at the elevation levels of 700

m and 1000 m, calling from November 2012 to January

2013, and from October 2013 to January 2014. In

December 2012 we registered one female and

observed some agonistic interactions among males

placed at a small water body on the ground.

Bokermannohyla luctuosa presents the reproductive

mode 12 (Pombal and Haddad, 1993, Nunes-de-

Almeida et al., 2021).

Dendropsophus berthalutzae

(Bokermann, 1962)

(Fig. 5B)

Vouchers - FURB 22754, 22897, 22899

Taxonomy and distribution - Dendropsophus

berthalutzae occurs in the Atlantic Forest from the state

of Espírito Santo to the state of Paraná, with a record in

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15 Anuran Natural History from Serra Dona Francisca

the state of Minas Gerais (Moura, Lacerda and Feio,

2012, Comitti, 2017). This species belongs to the D.

decipiens species group (Orrico et al., 2021).

Ecology - We found individuals of D. berthalutzae at

the elevation level of 300 m, on the herbaceous

vegetation in the forest edge. Calling males were

observed from October 2012 to March 2013. This

species laid eggs in the hanging foliage over a lentic

breeding site in February 2013 (reproductive mode 22,

Nunes-de-Almeida et al., 2021).

Dendropsophus elegans

(Wied-Neuwied, 1824)

(Fig. 5D)

Vouchers - FURB 22715, 22716, 22889, 22925

Taxonomy and distribution - This species occurs in

the Serra do Mar mountain range, from northern Bahia

(Canelas and Bertoluci, 2007; Gondim-Silva et al.,

2016) to the state of Santa Catarina (Lucas and Garcia,

2011). It belongs to the D. leucophyllatus species group

(Orrico et al., 2021).

Ecology - We found this species at elevation levels

of 100 m and 300 m, in swamps and dams in open

areas. Calling males were observed from October

2012 to February 2013, perched on herbaceous

vegetation and on the grass. We did not observe

females and spawning of D. elegans. This species lays

its eggs in the water surface (reproductive mode 1,

Bastos and Haddad, 1996, Nunes-de-Almeida et al.,

2021).

Dendropsophus microps

(Peters, 1872)

(Fig. 5E)

Vouchers - FURB 22395

Taxonomy and distribution - This species is widely

distributed in the south and southeast of Brazil (Maffei

et al., 2009), and belongs to the D. parviceps species

group (Orrico et al., 2021).

Ecology - We found D. microps only at elevation

levels of 700 m and 1000 m. Calling males were

observed from October 2012 to January 2013 and in

the same period in 2013 and 2014, perched in

herbaceous and grassy vegetation in swamps and

ponds at the edge of the forest. We did not observe

females and spawning of D. microps. This species lays

its eggs in the water surface and presents exotrophic

tadpoles (reproductive mode 1, Nunes-de-Almeida et

al., 2021).

Dendropsophus minutus

(Peters, 1872)

(Fig. 5F)

Vouchers – FURB 22237, 22381, 22860, 22879, 22898

Taxonomy and distribution – This species is widely

distributed all over the South America (Frost, 2021),

and belongs to the D. minutus species group (Orrico et

al., 2021). Despite its wide distribution, there are

several lineages within this nominal species (Gehara et

al., 2014).

Ecology - We found D. minutus at all elevation

levels, occupying swamps, dams, and streams in open

areas and in the forest edge areas. Calling males were

observed from October 2012 to March 2013, and from

August 2013 to January 2014, using herbaceous

vegetation and bushes as calling sites. We found

ovigerous females and amplected couples during

November and December 2012 and 2013. This species

lays its eggs on the water surface, and presents

exotrophic tadpoles (reproductive mode 1, Nunes-de-

Almeida et al., 2021).

Dendropsophus nahdereri

(Lutz and Bokermann, 1963)

(Fig. 5G)

Vouchers - FURB 22471, 22472

Taxonomy and distribution - This species is

restricted to the south of Brazil, mostly recorded in the

state of Santa Catarina (Conte et al., 2010), and

belongs to the D. marmoratus species group (Orrico et

al., 2021).

Ecology - We found D. nahdereri in open areas and

in the forest edge at elevation levels of 700 m and 1000

m. Calling males were observed on the ground,

perched on herbaceous vegetation or in trees near

water bodies, from October 2012 to January 2013, and

from July 2013 to January 2014. We observed

amplected couples only in November 2012. This

species lays its eggs in lentic environments, and

presents exotrophic tadpoles (reproductive mode 1,

Haddad et al., 2013, Nunes-de-Almeida et al., 2021).

Dendropsophus werneri

(Cochran, 1952)

(Fig. 5C)

Vouchers - FURB 22382, 22392, 22797, 22800, 22940

Taxonomy and distribution - This species occurs in

the Serra do Mar mountain range, from Rio de the state

of Janeiro to the state of Rio Grande do Sul (Pombal

and Bastos, 1998; Freire et al., 2016), and belongs to

the D. microcephalus species group (Orrico et al.,

2021).

Ecology - We found individuals of D. werneri in

herbaceous vegetation of dams and swamps in open

areas and on the edge of the forest at elevations of 100

m, 300 m, and 700 m, sometimes syntopically to D.

berthalutzae. We observed calling males from October

2012 to March 2013, and from November 2013 to

January 2014. Some females were sporadically

observed close to males, but we did not observe

spawning of this species. Dendropsophus werneri lays

its eggs on the water surface in lentic environments and

presents exotrophic tadpoles (reproductive mode 1,

Nunes-de-Almeida et al., 2021).

Phyllomedusa distincta

Lutz, 1950

(Fig. 8A)

Vouchers - FURB 22243, 22245

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MARIOTTO, L.R. et al.

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MARIOTTO, L.R. et al. 17

Fig. 5. Some amphibians registered in Serra Dona Francisca: (A) Bokermannohyla luctuosa, (B) Dendropsophus berthalut-

zae, (C) Dendropsophus werneri, (D) Dendropsophus elegans, (E) Dendropsophus microps, (F) Dendropsophus minutus,

(G) Dendropsophus nahdereri, (H) Scinax aromothynella, (I) Scinax catharinae, (J) Scinax littoralis, (K) Scinax rizibilis, and

(L) Scinax fuscovarius.

Taxonomy and distribution - This species occurs

from south of São Paulo to north of Rio Grande do Sul

states (Kwet et al., 2010), and belongs to the P.

burmeisteri species group (Pombal and Haddad, 1992;

Faivovich et al., 2010).

Ecology - We found individuals of P. distincta

foraging at the elevation level of 1000 m, and males

calling in a swamp at the elevation level of 700 m, all

at the edges of the forest, only in October 2012. We

did not observe females or spawning, but this species

lays its eggs in foliage hanging over lentic breeding

sites (reproductive mode 37, Nunes-de-Almeida et al.,

2021).

Scinax aromothyella

Faivovich, 2005

(Fig. 5H)

Vouchers - FURB 22259, 22314, 22647, 22772, 22723,

22891, 22944.

Taxonomy and distribution - This species occurs in

Uruguay, Argentina, and Brazil in the state of Paraná

(Pereyra et al., 2012).

Ecology - We found S. aromothyella in temporary

swamps from only in open areas, at elevation level of

1000 m. Calling males were observed in herbaceous

vegetation, in February, November and December

2013, and January 2014. We recorded ovigerous

females in the same period and in September 2013. We

did not observe the reproductive mode of this species,

which remains unknown.

Scinax catharinae

(Boulenger, 1888)

(Fig. 5I)

Vouchers - FURB 22239, 22281, 22311,22746

Taxonomy and distribution - This species occurs in

the mountain regions of the state of Santa Catarina and

Rio Grande do Sul (Lutz, 1973). There are some

records in the interior of Santa Catarina (Lucas and

Fortes, 2008) and Paraná states (Machado and

Bernarde, 2002, Crivelari et al., 2014), but in some of

these records the identification is uncertain (Scinax cf.

catharinae). The type locality of the species is

imprecise (Sierra do Catharina, Boulenger, 1888a), but

Lutz (1973) suggested that it is probably the mountains

of northern of the state of Santa Catarina, where the

German immigrants settled.

Ecology - We found individuals foraging in a stream

at the edge of the forest, only at the elevation level of

1000 m, from November 2012 to March 2013, and

October 2013. We did not observe the reproductive

behaviour, which species remains unknown.

Scinax littoralis

(Pombal and Gordo, 1991)

(Fig. 5J)

Vouchers - FURB 22278, 22280, 22711

Taxonomy and distribution - This species occurs

from São Paulo (Pombal and Gordo, 1991) to state of

Santa Catarina (Lucas and Garcia, 2011), and belongs

to the Scinax catharinae group (Faivovich et al., 2005).

Ecology - We found individuals of S. littoralis in

swamps and streams at the edge of the forest, at

elevation levels of 100 m and 300 m. Calling males

were observed on the shrubby vegetation, from

October 2012 to March 2013; only one female was

found in February 2013. This species presents the

reproductive mode 7, with eggs and tadpoles in

bromeliad axils (Toledo et al., 2012) and reproductive

modes 1 or 2, with exotrophic eggs and tadpoles in

lentic or lotic water breeding sites (Pombal and Gordo,

1991, Nunes-de-Almeida et al., 2021).

Scinax rizibilis

(Bokermann, 1964)

(Fig. 5K)

Vouchers - FURB 22371, 22372, 22391, 22473, 22869

Taxonomy and distribution - This species occurs

from the state of São Paulo (Bokermann, 1964) to the

state of Rio Grande do Sul (Kwet et al., 2010) and

belongs to the S. catharinae species group (Faivovich

et al., 2005).

Ecology - We found S. rizibilis in lentic

environments, such as swamps and dams at the forest

edge and in the forest interior, at all sampled elevation

levels. Calling males were observed on the shrubby

vegetation from October 2012 to March 2013, and we

found only one female, in October 2012. We did not

observe spawning. This species lays its eggs on the

water surface (Bokermann, 1964) and uses foam nests

to protect the eggs (reproductive mode 65, Haddad,

Pombal and Gordo, 1990, Nunes-de-Almeida et al.,

2021).

Scinax sp. (gr. perpusillus)

Vouchers - no vouchers

Taxonomy and distribution – This species is

morphologically and ecologically similar to the species

belonging to the Scinax perpusillus group, which are

distributed from the state of Espírito Santo to the state

of Paraná (Faivovich et al. 2005). Despite Bertha Lutz

(1973), in her book “Brazilian Species of Hyla” (1973)

provided the species occurrence for Guaramirin and

Joinville municipalities, state of Santa Catarina, Brazil,

we did not collect individuals, hampering further

comparisons. The advertisement calls (n = 9, one

male) have the following parameters: Fmin

3617.000–3789.000 Hz (3707.777 ± 50.166), Fmax

4608.000–4780.000 Hz (4732.222 ± 58.661), Fdom

4134.000–4177.000 Hz (4167.444 ± 18.961), call

duration 0.205–0.278 s (0.249 ± 0.026), 3–4 notes per

call (3.777 ± 0.440), note duration 0.029–0.046 s

(0.034 ± 0.003), 7–12 pulses per note (9.527 ± 1.922).

Scinax aff. perpusillus differs from S. perpusillus by the

lower dominant frequency (4134–4177 Hz), longer

duration (0.200–0.270 s), and higher number of pulses

(7–12 pulses/note) (4554–4856 Hz, 0.007–0.018 s,

3–5 pulses/note, Rio de Janeiro municipality, Rio de

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MARIOTTO, L.R. et al.

Janeiro state, 700 km from Serra Dona Francisca,

Pombal and Bastos 2003). Nevertheless, we kept the

population of the study area as S. aff. perpusillus.

Ecology - We did not observe individuals of this

species, but we heard several males calling in the

epiphytic bromeliads occurring one to two meters from

the ground, at the edge of the forest and in the forest

interior. We recorded this species at altitude levels of

300 m and 700 m, from October to December 2012,

and from December 2013 to January 2014. Is worth to

note that species belonging to the Scinax perpusillus

group are bromeligenous that lays their eggs in the

accumulated water of tank bromeliads (reproductive

mode 4, Peixoto, 1995, Nunes-de-Almeida et al.,

2021).

Scinax fuscovarius

(Lutz, 1925)

(Fig. 5L)

Vouchers - FURB 22258, 22299

Taxonomy and distribution - This species is widely

distributed in the eastern Brazil from Alagoas to Rio

Grande do Sul states, and in the northern Argentina,

northern and eastern Paraguay, and Bolivia, from 150

to 1800 m elevation (Frost, 2021), and belongs to the S.

ruber species group (Faivovich et al., 2005).

Ecology - We found individuals of S. fuscovarius at

elevation levels of 100 m, 700 m, and 1000 m, in

temporary or permanent lentic environments in open

areas and in forest edges areas. We observed males

calling in herbaceous vegetation and grasses, from

October 2012 to February 2013, and from November

2013 to January 2014. We did not find females or

spawning. This species lays its eggs on the water

surface and presents exotrophic tadpoles

(reproductive mode 1, Kwet et al. 2010, Nunes-de-

Almeida et al., 2021).

Scinax granulatus

(Peters, 1871)

(Fig. 6A)

Vouchers - FURB 22648, 22710, 22764, 22771

Taxonomy and distribution - This species is widely

distributed south of South America, occurring in north-

eastern Argentina, Uruguay, in the Brazilian states of

Paraná, Santa Catarina, and Rio Grande do Sul, and in

south-eastern Paraguay (Conte et al., 2010; Frost,

2021), and belongs to the S. ruber species group

(Faivovich et al., 2005).

Ecology - We found S. granulatus only at elevation

level of 1000 m. Calling males were observed in

temporary or permanent lentic environments, perched

on herbaceous vegetation at open areas, from

November 2012 to April 2013, and from August 2013 to

January 2014. We did not find females or spawning.

This species presents the reproductive mode 1

(Nunes-de-Almeida et al., 2021), and presents

exotrophic tadpoles (Kwet et al. 2010, Nunes-de-

Almeida et al., 2021).

Scinax imbegue

Nunes, Kwet and Pombal, 2012

(Fig. 6B)

Vouchers - FURB 22284, 22823, 22921

Taxonomy and distribution - This species is widely

distributed in the south and southeast of Brazil (Frost,

2021) and it is endemic to the Atlantic Forest (Nunes,

Kwet and Pombal, 2012). Scinax imbegue belongs to

the S. ruber species group (Nunes et al., 2012).

Ecology - We found S. imbegue in temporary and

permanent lentic environments in open areas and in

forest edges, at elevation levels from 100 m to 700

m. We observed males calling in herbaceous

vegetation and grasses, from October 2012 to March

2013, and from September 2013 to January 2014; we

found amplected couples in October and December

2012. This species has its nest absent, with non-froth

floating eggs laid in lentic water (reproductive mode

1; Nunes et al. 2012, Nunes-de-Almeida et al., 2021).

Scinax perereca

Pombal, Haddad and Kasahara, 1995

(Fig. 6C)

Vouchers - FURB 22238, 22386, 22482, 22488, 22824

Taxonomy and distribution - Scinax perereca is

known only from a few localities in the states of São

Paulo to Rio Grande do Sul states in Brazil, and

Misiones Province in north-eastern Argentina; likely in

south-eastern Paraguay (Frost, 2021). This species

belongs to the S. ruber group (Pombal, Haddad and

Kasahara, 1995; Faivovich et al., 2005).

Ecology - We found S. perereca in temporary and

permanent lentic environments in open areas and in

forest edge areas, at all sampled elevation levels. We

observed calling males on herbaceous vegetation or

in underground cavities, from October 2012 to March

2013, and from September 2013 to January 2014.

We also observed this species in forested areas,

calling on tree trunks up to 60 cm from the ground,

and did not find spawns or females. This species

presents the reproductive mode 1 (Nunes-de-

Almeida et al., 2021), with spawning laid in lentic

environments and with exotrophic tadpoles (Kwet et

al. 2010).

Scinax tymbamirim

Nunes, Kwet and Pombal, 2012

(Fig. 6D)

Vouchers - FURB 22257, 22722, 22896, 22903, 22946

Taxonomy and distribution - This species is widely

distributed in the south and southeast of Brazil (Frost,

2021), endemic to the Atlantic Forest (Nunes et al.,

2012). It belongs to the S. ruber species group (Nunes

et al., 2012).

Ecology - We found S. tymbamirim in temporary

and permanent lentic environments in open areas, only

at elevation level of 100 m, with males calling from

October 2012 to March 2013. We observed females

and amplected couples in October 2012. This species

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19 Anuran Natural History from Serra Dona Francisca

Bol. Soc. Zool. Uruguay (2ª época). 2022. Vol. 31 (2): e31.2.4 ISSN 2393-6940

Anuran Natural History from Serra Dona Francisca20

Fig. 6. Some amphibians registered in Serra Dona Francisca: (A) Scinax granulatus, (B) Scinax imbegue, (C) Scinax pere-

reca, (D) Scinax tymbamirim, (E) Scinax squalirostris, (F) Sphaenorhynchus surdus, (G) Trachycephalus mesophaeus, (H)

Crossodactylus sp., (I) Hylodes perplicatus, (J) Adenomera araucaria, (K) Adenomera nana, and (L) Leptodactylus paranaru.

occurred syntopically with S. imbegue and S. perereca.

It presents the reproductive mode 1 (Nunes-de-

Almeida et al., 2021), with spawning laid in lentic

environments and with exotrophic tadpoles (Nunes et

al., 2012).

Scinax squalirostris

(Lutz, 1925)

(Fig. 6E)

Vouchers - FURB 22369, 22729, 22755

Taxonomy and distribution - This species is widely

distributed in the south, southeast and midwest of

Brazil (Frost, 2021), and belongs to the S. ruber

species group (Faivovich et al., 2005).

Ecology - We found S. squalirostris only in

temporary swamps in open areas at elevation level of

1000 m. Several calling males were observed from

October 2012 to March 2013, and from May to

December 2013. This species occurred syntopically

with S. fuscovarius, S. perereca and S. granulatus.

Scinax squalirostris presents the reproductive mode

1 (Nunes-de-Almeida et al., 2021), with spawning laid

in lentic environments and with exotrophic tadpoles

(Kwet et al. 2010).

Sphaenorhynchus surdus

(Cochran, 1953)

(Fig. 6F)

Vouchers - FURB 22370, 22246, 22475

Taxonomy and distribution - This species is

restricted to the south of Brazil, in Araucaria Forest and

in transitional areas between other associated

physiognomies (Toledo et al., 2007).

Ecology - We found S. surdus only at elevation level

of 1000 m, in lentic environments, using herbaceous

vegetation, grasses and macrophytes as vocalization

sites in open areas and in forest edge areas. We

observed males calling from October 2012 to March

2013, and from August 2013 to January 2014. We did

not find females and spawning. This species lays its

eggs in lentic environments and presents exotrophic

tadpoles (reproductive mode 3; Nunes-de-Almeida et

al., 2021).

Trachycephalus mesophaeus

(Hensel, 1867)

(Fig. 6G)

Vouchers - FURB 22191, 22192

Taxonomy and distribution - This species occurs

along the coast from the state of Rio Grande do Sul to

the state of Pernambuco (Santana et al., 2016).

Ecology - We found T. mesophaeus in temporary

swamps in open areas, forest edges and forest

interior, at elevation levels from 100 m to 700 m. We

observed several individuals, including males and

females, after heavy rains in October, November, and

December 2012. We found amplected couples laying

the eggs on the surface of the water (reproductive

mode 1, Nunes-de-Almeida et al., 2021); five weeks

after the first reproductive event we found froglets of

this species. Lutz (1973) observed the larval

development of this species under laboratory

conditions, and the total duration until metamorphosis

was two to three months.

Family Hylodidae

Crossodactylus sp.

(Fig. 6H)

Vouchers - FURB 22725, 22747, 22763

Taxonomy and distribution - The occurrence of C.

caramaschii Bastos and Pombal, 1995 has already

been reported for the study area (Pimenta, 2008).

However, based on external morphology, such as the

presence of a gland under the tympanum, as a discreet

crest in C. caramaschii, very distinct in the vouchers,

we were not able to associate this population to C.

caramaschii or to any valid nominal species within the

genus.

Ecology - We found individuals of Crossodactylus

sp. in streams in the interior of the forest, only at

elevation level of 700 m. Calling males were

observed from October 2012 to January 2013, and in

March 2013; we did not observe females or

spawning. In January and February 2013, we found

individuals foraging in streams, and two juveniles

were captured by pitfall traps. Species of the genus

Crossodactylus present the reproductive mode 18

(Nunes-de-Almeida et al., 2021), with spawning and

tadpoles in early stages sheltered in burrows on the

banks of the stream; after development, the

exotrophic tadpoles move to water.

Hylodes perplicatus

(Miranda-Ribeiro, 1926)

(Fig. 6I)

Vouchers - FURB 22252, 22273, 22863

Taxonomy and distribution –

state of Santa Catarina, Brazil. Its type locality,

Colônia Hansa-Humboldt (Miranda-Ribeiro, 1926),

current Corupá and São Bento do Sul municipalities,

state of Santa Catarina (Frost, 2021). The

advertisement calls (n = 7 calls from one male) have

the following parameters: dominant frequency

3445.000–3617.000 Hz (3524.857 ± 62.945), call

duration 0.904–1.235 s (1.114 ± 0.114), notes per call

18–24 (21.571 ± 2.070), and note duration

0.026–0.044 (0.037 ± 0.006). These calls match with

the description of the advertisement call available for

H. perplicatus in this same locality

Ecology - We found H. perplicatus in streams in

the interior of the forest, only at elevation levels of

300 m and 700 m. Calling males were observed from

November 2012 to March 2013. We observed only

one female in February 2013, and juveniles captured,

mostly by pitfall traps, from November 2012 to

This species is

known only from few localities in the municipalities of

São Bento do Sul, Corupá and Joinville, northen of

the

(Monteiro, Comitti

and Lingnau, 2014)

Bol. Soc. Zool. Uruguay (2ª época). 2022. ISSN 2393-6940Vol. 31 (2): e31.2.4

21 Anuran Natural History from Serra Dona Francisca

February 2013. This species has reproductive modes

14 and 18 and presents exotrophic tadpoles (Nunes-

de-Almeida et al., 2021).

Family Leptodactylidae

Adenomera araucaria

Kwet and Angulo, 2002

(Fig. 6J)

Vouchers - FURB 22376, 22704, 22717, 22728, 22796,

22828, 22936

Taxonomy and distribution - This species is

distributed in the south of Brazil, occurring in western

Paraná state, and reaching the southern coast of the

state of Santa Catarina (Conte et al., 2010; Fouquet et

al., 2014).

Ecology - We found individuals of A. araucaria at

elevation levels of 700 m and 1000 m. We observed

males calling in the leaf litter at the edge and the interior

of the forest, from October 2012 to January 2013, and

from November 2013 to January 2014. From October

2012 to January 2013, we captured females with pitfall

traps, and ovigerous females during December 2012

and January 2013. In January and February 2013, we

captured two juveniles. This species lays its eggs in

underground cavities or in leaf litter, in foam nests, and

presents endotrophic tadpoles (reproductive mode 62,

Kwet et al. 2010, Nunes-de-Almeida et al., 2021).

Adenomera nana

(Müller, 1922)

(Fig. 6K)

Vouchers - FURB 22260, 22261, 22262, 22264, 22276,

22295, 22296, 22316, 22317, 22318, 22385, 22393,

22476, 22718, 22744, 22768, 22825, 22892, 22893,

22931, 22945

Taxonomy and distribution - This species occurs in

Santa Catarina and Paraná states, southern Brazil

(Kwet, 2007, Conte et al., 2010, Fouquet et al., 2014).

Ecology - We found individuals of A. nana in all

sampled elevation levels. We observed males calling in

the leaf litter at the edge and the interior of the forest,

from October 2012 to March 2013, and from

September 2013 to January 2014. We captured

females in the pitfall traps from October 2012 to

January 2013, and ovigerous females during

December 2012 and January 2013. This species lays

its eggs in underground cavities or in leaf litter, in foam

nests, and presents endotrophic tadpoles

(reproductive mode 62, Heyer, 1973, Kwet et al. 2010,

Nunes-de-Almeida et al., 2021).

Leptodactylus paranaru

Magalhães, Lyra, Carvalho, Baldo, Brusquetti, Burella,

Colli, Gehara, Giaretta, Haddad, Langone, López,

Napoli, Santana, Sá and Garda, 2020.

(Fig. 6L)

Vouchers - FURB 22187, 22188

Taxonomy and distribution - This species belongs to

the Leptodactylus latrans group (Magalhães et al.,

2020), and represents a recently described species,

which was removed from the synonymy of L. latrans

(Magalhães et al., 2020). The Southern-butter-frog is

distributed in the south-eastern coastal region of the

Atlantic Forest, from Santos municipality, in the state of

São Paulo, to the northeastern state of Rio Grande do

Sul, located east of the Serra do Mar mountain range

(Magalhães et al., 2020).

Ecology - We found this species at all sampled

elevation levels, in lentic environments (temporary or

permanent swamps and dams) in open areas and on

forest edges. Calling males were observed from

October 2012 to March 2013, and from October 2013 to

January 2014, but we also found individuals foraging in

April, August, and September 2013. We observed

females and tadpoles from November 2012 to March

2013, and juveniles were captured with pitfall traps in

January and March 2013. Tadpoles form schools that

receive adult parental care. This species lays its eggs in

foam nests in lentic environments, and presents

exotrophic tadpoles (reproductive mode 45, Nunes-de-

Almeida et al., 2021).

Leptodactylus notoaktites

Heyer, 1978

(Fig. 7A)

Vouchers - FURB 22193, 22251, 22255, 22256, 22292,

22303, 22367, 22816, 22895

Taxonomy and distribution - This species occurs

from the state of São Paulo to the state of Santa

Catarina (Heyer, 1978; Figueiredo et al., 2018) and

belongs to the L. fuscus species group (Sá et al., 2014).

Ecology - We found L. notoaktites at all elevation

levels in temporary ponds in open areas and on forest

edges, where we observed males calling from October

2012 to March 2013, and from October 2013 to January

2014. We did not find females; we captured juveniles in

pitfall traps, from December 2012 to March 2013. As

usually observed for species of the L. fuscus group, L.

notoaktites lays its eggs in foam nests in underground

cavities, and presents exotrophic tadpoles

(reproductive mode 64, Nunes-de-Almeida et al.,

2021).

Leptodactylus plaumanni

Ahl, 1936

(Fig. 7B)

Vouchers - FURB 22467, 22366, 22483

Taxonomy and distribution - This species is distributed

in north-eastern Misiones Province, Argentina, and in

the Brazilian states of Rio Grande do Sul, Paraná, and

Santa Catarina, likely into adjacent Paraguay (Kwet,

Di-Bernardo and Garcia, 2001; Frost, 2021), and

belongs to the L. fuscus species group (Sá et al., 2014).

Ecology - We found individuals of L. plaumanni at

elevation level of 1000 m, in temporary swamps in open

areas. We observed males calling from October 2012

to January 2013, and from September 2013 to January

2014; only one juvenile was captured in March 2013, by

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MARIOTTO, L.R. et al. 23

Fig. 7. Some amphibians registered in Serra Dona Francisca: (A) Leptodactylus notoaktites, (B) Leptodactylus plaumanni,

(C) Physalaemus aff. gracilis, (D) Physalaemus cuvieri, (E) Physalaemus lateristriga, (F) Physalaemus nanus, (G) Scythro-

phrys sawayae, (H) Chiasmocleis leucosticta, (I) Elachistocleis bicolor, (J) Odontophrynus americanus, (K) Proceratophrys

boiei, and (L) Proceratophrys subguttata.

pitfall trap. Leptodactylus plaumanni lays its eggs in

foam nests in underground cavities, and presents

exotrophic tadpoles (reproductive mode 64, Nunes-de-

Almeida et al., 2021).

Physalaemus aff. gracilis

(Fig. 7C)

Vouchers - FURB 22287, 22731, 22853, 22854, 22934

Taxonomy and distribution - Species of the P.

gracilis group are distributed in the south of Brazil,

Uruguay, Paraguay, and Argentina, occupying areas

above 600 m a.s.l., except P. gracilis (Nascimento,

Caramaschi and Cruz, 2005) which occurs at lower

elevations. Because we recorded individuals only at

the elevation level of 1000 m and based on acoustic

data and taxonomic remarks by Kwet et al., 2010, we

believe that specimens from Serra Dona Francisca are

not conspecific with P. gracilis. The advertisement calls

(n=32 from four males) have the following parameters:

dominant frequency 2454–2928 Hz (2823.066 ±

145.361), call duration 0.986–1.328 s (1.114 ± 0.123),

one note per call and seven harmonics.

Ecology - We registered Physalaemus aff. gracilis

at elevation level of 1000 m, in lentic environments

(permanent or temporary) in open areas and in forest

edges areas. We observed males calling from October

2012 to February 2013, and from July 2013 to January

2014. Females were captured in pitfall traps from

October 2012 to March 2013; only one juvenile was

captured in January 2013. Species of the genus

Physalaemus presents reproductive mode 45 (Nunes-

de-Almeida et al., 2021), laying their eggs in floating

foam nests, and present exotrophic tadpoles in ponds.

Physalaemus cuvieri

Fitzinger, 1826

(Fig. 7D)

Vouchers - FURB 22468, 22733, 22757, 22817, 22862,

22920, 22943

Taxonomy and distribution - This species is widely

distributed in South America (Frost, 2021) and belongs

to the P. cuvieri clade (Lourenço et al., 2015).

Ecology - We found individuals of P. cuvieri at all

sampled elevation levels, in temporary or permanent

lentic environments in open areas and in forest edges

areas. We observed males calling from October 2012

to March 2013, and from June 2013 to January 2014.

Females were captured by pitfall traps in November

2012. This species presents reproductive mode 45,

laying their eggs in foam nests in lentic environments,

and presents exotrophic tadpoles (Nunes-de-Almeida

et al., 2021).

Physalaemus lateristriga

(Steindachner, 1864)

(Fig. 7E)

Vouchers - FURB 22305, 22368, 22378, 22394, 22481,

22709, 22819, 22820, 22821, 22865, 22890

Taxonomy and distribution - This species occurs

from south-eastern state of São Paulo to north-eastern

state of Santa Catarina and belongs to the P. olfersii

species group (Cassini, Cruz and Caramaschi., 2010).

Ecology - We recorded one individual of P.

lateristriga foraging at the elevation level of 100 m and

several individuals at the elevation levels of 700 m and

1000 m, all at the edges of the forest. We observed

males calling from October 2012 to March 2013, and

from October 2013 to January 2014. During this same

period, we captured females with pitfall traps; we

captured juveniles from October 2012 to February

2013. This species presents reproductive mode 45

(Nunes-de-Almeida et al., 2021).

Physalaemus nanus

(Boulenger, 1888)

(Fig. 7F)

Vouchers - FURB 22319, 22384, 22930

Taxonomy and distribution - This species occurs

from the north of the state of Rio Grande do Sul (Kwet et

al., 2010) to the north state of Santa Catarina

(Bokermann, 1962), extending mostly through the west

of the state of Santa Catarina (Lingnau, 2009).

Physalaemus nanus belongs to the P. signifer clade

(Lourenço et al., 2015).

Ecology - We found P. nanus only at the elevation

level of 1000 m, with males calling in temporary ponds

in the forest interior, and in a swamp at the edge of the

forest, from November 2012 to January 2013, and from

September 2013 to January 2014. Only one female

was captured in November 2013, by pitfall trap. As

known for other species of the genus Physalaemus, P.

nanus presents reproductive modes 45 and 59,

producing foam nests in lentic environments, and

presents exotrophic tadpoles (Nunes-de-Almeida et

al., 2021).

Scythrophrys sawayae

(Cochran, 1953)

(Fig. 7G)

Vouchers - FURB 22727, 22815, 22827, 22872, 22888

Taxonomy and distribution - This species occurs

from the south of the state of Paraná to the south of the

state of Santa Catarina (Santos et al., 2020).

Ecology - We registered this species with pitfall

traps at the elevation level 300 m, and in high

abundance at the elevation level of 700 m (see Table 2).

During the hot and rainy season, we captured males

and females by pitfall traps, and found individuals in

calling activity from October to November 2012 and

2013. We observed some amplected couples inside

pitfall traps from October 2012 to January 2013.

Scythrophrys sawayae uses temporary ponds as

reproduction sites, where we found males calling at the

edge of the forest and in the forest interior. Some

spawning was observed in the cover of pitfall traps with

accumulated water (reproductive mode 1, Nunes-de-

Almeida et al., 2021).

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MARIOTTO, L.R. et al. 25

Fig. 9. Hands of two adult males of (A) Bokermannohyla luctuosa, from Campo Alegre municipality, state of Santa Catarina,

and (B) B. circumdata, from Doutor Pedrinho municipality, state of Santa Catarina. The red arrows indicate the differences

(bilobed in B) of the tubercles of finger I .

Fig. 8. Some amphibians

registered in Serra Dona

Francisca: Phyllomedusa

distincta.

Family Microhylidae

Chiasmocleis leucosticta

(Boulenger, 1888)

(Fig. 7H)

Vouchers - FURB 22388, 22484, 22485, 22750, 22769,

22866, 22905

Taxonomy and distribution - This species occurs

from the states of São Paulo to Rio Grande do Sul

(Cruz, Caramaschi and Izecksohn, 1997; Freire et al.,

2016) and it is not assigned to any species group (Sá et

al., 2012).

Ecology - We registered individuals of C. leucosticta

in the forest at the elevation levels from 100 m to 700 m.

We observed intense vocalization activity after heavy

rains and captured numerous ovigerous females and

amplected couples in pitfall traps, in October 2012.

Besides this explosive episode, we also captured this

species in abundance in pitfall traps during the summer

(from October 2012 to February 2013). Chiasmocleis

leucosticta builds foam nests in temporary swamps in

the forest interior, and presents exotrophic tadpoles

(reproductive mode 65, Haddad and Hödl, 1997,

Nunes-de-Almeida et al., 2021).

Elachistocleis bicolor

(Guérin-Méneville, 1838)

(Fig. 7I)

Vouchers - FURB 22387, 22887, 22928

Taxonomy and distribution - This species occurs

from the state of São Paulo to Rio Grande do Sul, with

records in Paraguay, Uruguay, and Argentina

(Caramaschi, 2010).

Ecology - We found E. bicolor only in the elevation

levels of 100 m and 1000 m, with males calling in lentic

and temporary environments at open areas. The

vocalization period lasted from October 2012 to

February 2013, and in November and December 2013.

No females and juveniles were captured by pitfall traps.

Elachistocelis bicolor lays its eggs directly on the

surface of the water (reproductive mode 1, Rodrigues,

Lopes and Uetanabaro, 2003, Nunes-de-Almeida et

al., 2021).

Family Odontophrynidae

Odontophrynus americanus

(Duméril and Bibron, 1841)

(Fig. 7J)

Vouchers - FURB 22286, 22466, 22477

Taxonomy and distribution - This species is widely

distributed, occurring from the south of the state of

Minas Gerais to the state of Rio Grande do Sul,

Paraguay, Argentina, and Uruguay, and belongs to the

O. americanus species group (Rosset et al., 2006).

Ecology - We found this species in February 2012

and November 2013, in only one reproductive site, a

temporary swamp in an open area, at the elevation

level of 1000 m. We did not find spawning of O.

americanus during fieldwork, but it is known that this

species presents reproductive mode 2, laying its eggs

in lentic environments, and presents exotrophic

tadpoles (Nunes-de-Almeida et al., 2021).

Proceratophrys boiei

(Wied-Neuwied, 1824)

(Fig. 7K)

Vouchers - FURB 22195, 22196, 22197, 22198, 22199,

22200, 22242, 22469, 22470

Taxonomy and distribution - This species is widely

distributed in Brazil, occurring from the state of Espírito

Santo to Santa Catarina, in the Atlantic Forest domain,

extending to the Cerrado domain in São Paulo and

Minas Gerais states, and belongs to the P. boiei

species group (Prado and Pombal, 2008).

Ecology - We found individuals of P. boiei at all

sampled elevation levels. Calling activity was observed

in swamps in the forest interior during the spring and

summer periods from 2012 to 2014. We captured

females and juveniles by pitfall traps, from October

2012 to January 2013, and from October 2012 to March

2013, respectively. We did not observe individuals

reproducing, but this species presents reproductive

modes 2 and 16 (Nunes-de-Almeida et al., 2021).

Proceratophrys subguttata

Izecksohn, Cruz and Peixoto, 1999

(Fig. 7L)

Vouchers - FURB 22193, 22194, 22290

Taxonomy and distribution - This species is

restricted to south of the state of Paraná and northeast

of the state of Santa Catarina and belongs to the P.

appendiculata species complex (Prado and Pombal,

2008).

Ecology - Three individuals were captured by pitfall

traps, and three juveniles were observed in the forest in

October 2012, only at the elevation level of 700 m. We

did not find males in calling activity, nor do we observe

individuals reproducing. The reproductive mode of this

species is 16 (Nunes-de-Almeida et al., 2021).

Family Ranidae

Lithobates catesbeianus

(Shaw, 1802)

Taxonomy and distribution - Widely introduced

throughout the globe, including in several regions in

Brazil (Both et al., 2011), this species is native from

eastern North America and some adjacent areas in

Central America (Frost, 2021).

Vouchers - FURB 22924

Only one juvenile was captured at the elevation level of

100 m, by pitfall trap. We did not find males in calling

activity during fieldwork. This is an exotic and invasive

species, distributed in the south, southeast, middle-

west and northeast of Brazil (Both et al. 2011), with

some records in Argentina (Akmentins and Cardozo,

2010), Uruguay (Laufer et al., 2008), Colombia,

Ecuador, and Venezuela (IUCN SSC 2015).

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MARIOTTO, L.R. et al.

2. Species Richness and Species Composition

The rarefaction/extrapolation curves for the SBS

(observed richness=62, estimated richness=62.382 ±

0.838) and PT (observed richness=23, estimated

richness=23.825 ± 1.367) methods reached the

asymptote, with no differences between the observed

and estimated species richness (Hill numbers of order

q=0) (Fig. 10). The sample completeness curves did

not show differences among the sampled localities for

each sampling method (Fig. 11). The high species

richness (63 species) found in Serra Dona Francisca is

probably related to the heterogeneous landscape of

the region. The elevation gradient presents streams,

ponds, and permanent and temporary swamps within

the phytophysiognomies of the Atlantic Forest:

lowlands, sub-montane and montane ombrophilous

dense forests, mixed ombrophilous forest (or Araucaria

Forest) and steppes (Veloso et al., 2012).

We found the greatest species richness at the

highest elevations (Fig. 12), within areas with

montane ombrophilous dense forest, mixed

ombrophilous forest and steppes. The richness is the

same for interpolated species curves between 1000 m

and 700 m, as the same for 300 m and 100 m. The

richness difference is in the interpolated species

richness curves between these groups (1000 and 700

are different from 300 and 100), and in the

extrapolated species richness curve of 300 m level

had a light overlap of confidence interval with the 700

m and 1000 m elevation levels (Fig. 12 - Hill numbers

q=0). For the SBS method, the elevation level 1000 m

registered the highest species richness

(observed=38.000, estimated=39.999 ± 3.739),

f o l l o w e d b y 7 0 0 m ( o b s e r v e d = 3 6 . 0 00 ,

estimated=38.247 ± 3.392), 300 m (observed=26.000,

estima ted=32 .228 ± 7.526) an d 10 0 m

(observed=24.000, estimated=24.998 ± 1.868). Only

the elevation level of 300 m showed differences

between observed and estimated species richness

(Table 3). For the PT method, the interpolated and

extrapolated curves had no differences among the

elevation levels for species richness (Fig. 13 - Hill

numbers q=0). The highest species richness was

registered at the 700 m elevation level

(observed=14.000, estimated=14.665 ± 1.303),

Bol. Soc. Zool. Uruguay (2ª época). 2022. ISSN 2393-6940Vol. 31 (2): e31.2.4

Fig. 10. Rarefaction curves for the Surveys at Breeding Sites (SBS) and Pitfall Traps (PT) sampling methods showing the

interpolation curve (continuum line) with the observed richness (dot) and the extrapolation curve (dashed line) representing the

estimated richness. Shaded areas represent the 95% confidence interval.

27 Anuran Natural History from Serra Dona Francisca

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Anuran Natural History from Serra Dona Francisca28

Fig. 11. Sampling Completeness curves comparing the sampling effort among the sampled localities (1000, 700, 300 and 100) for each sampling method: Surveys

at Breeding Sites (left) and Pitfall traps (right).

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MARIOTTO, L.R. et al. 29

Fig. 12. Rarefaction curves for the Surveys at Breeding Sites (SBS) sampling method showing the interpolation curve (continuum line) and the extrapolation curve

(dashed line) representing the richness (Hill number=0), Shannon diversity (Hill number=1) and Simpson diversity (Hill number=2) with twice the sample size, for

each sampled locality (1000 - green; 700 - purple; 300 - blue; 100 - red). Shaded areas represent the 95% confidence interval

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Anuran Natural History from Serra Dona Francisca30

Fig. 13. Rarefaction curves for the Pitfall Traps (PT) sampling method showing the interpolation curve (continuum line) and the extrapolation curve (dashed line)

representing the richness (Hill number=0), Shannon diversity (Hill number=1) and Simpson diversity (Hill number=2) with twice the sample size, for each sampled

locality (1000 - green; 700 - purple; 300 - blue; 100 - red). Shaded areas represent the 95% confidence interval

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MARIOTTO, L.R. et al. 31

Fig. 14. Abundance distribution of the anuran species registered for each sampled locality (100 - A, E; 300 - B, F; 700 - C, G;

1000 - D, H) and sampling method (Surveys at Breeding Sites - A-D; Pitfall traps - E-H). Grey bars – relative (SBS) and total

(PT) abundances (left axis); Red line - percentage of abundance (right axis). Aalbo - Aplastodiscus albomarginatus, Aarau

- Adenomera araucaria, Aehrh - Aplastodiscus ehrhardti, Anana - Adenomera nana, Aperv - Aplastodiscus perviridis, Balbo

- Boana albomarginata, Balpu - Boana albopunctata, Bbisc - Boana bischoffi Bfabe - Boana faber, Bhyla - Bokermanohyla

hylax, Bluct - Bokermanohyla luctuosa, Boana - Boana semiguttata, Bsemi - Boana semilineata, Bsulp - Brachycephalus

sulfuratus, Cboli - Cycloramphus bolitoglossus, Cizek - Cycloramphus izecksohni, Cleuc - Chiasmocleis leucosticta, Cross -

Crossodactylus sp., Dbert - Dendropsophus berthalutzae, Deleg - Dendropsophus elegans, Dmicr - Dendropsophus microps,

Dminu - Dendropsophus minutus, Dnahd - Dendropsophus nahdereri, Dpber - Dendrophryniscus berthalutzae, Dwern - Den-

dropsophus werneri, Ebico - Elachistocleis bicolor, Fmitu - Fritziana mitus, Fritz - Fritziana sp., Hbino - Haddadus binotatus,

Hperp - Hylodes perplicatus, Lcate - Lithobates catesbeianus, Iguen - Ischnocnema sp. (gr. guentheri), Ihens - Ischnocnema

henselii, Ischn - Ischnocnema sp. (I. lactea species series), Llatr - Leptodactylus paranaru, Lnoto - Leptodactylus notoaktites,

Lplau - Leptodactylus plaumanni, Mxant - Melanophryniscus xanthostomus, Oamer - Odontophrynus americanus, Sarom -

Scinax aromothyella, Scath - Scinax catharinae, Slitt - Scinax littoralis, Sperp - Scinax aff. perpusillus, Srizi - Scinax rizibilis,

Pboie - Proceratophrys boiei, Pcuvi - Physalaemus cuvieri, Pdist - Phyllomedusa distincta, Pgrac - Physalaemus aff. gracilis,

Plate - Physalaemus lateristriga, Pnanu - Physalaemus nanus, Psubg - Proceratophrys subguttata, Rorna - Rhinella ornata,

Ricte - Rhinella icterica, Sfusc - Scinax fuscovarius, Sgran - Scinax granulatus, Simbe - Scinax imbegue, Spere - Scinax

perereca, Ssawa - Scythophrys sawayae, Ssqua - Scinax squalirostris, Ssurd - Sphaenorhynchus surdus, Stymb - Scinax

tymbamirim, Tmeso - Trachycephalus mesophaeus, Vuran - Vitreorana uranoscopa.

followed by the 1000 m elevation level

(observed=11.000, estimated=11.249 ± 0.728) and by

the elevation levels of 100 m (observed=9.000,

estima ted=14 .936 ± 7.050) an d 30 0 m

(observed=9.000, estimated=13.442 ± 7.106). The

elevation levels of 100 m and 300 m had the same

species richness and were the only ones to show

differences between observed and estimated species

richness (Table 3).

Compared to some studies on anuran in elevation

gradients (e.g., Fauth, Crother and Slowinski, 1989,

Lescano et al., 2015), we found an inverted pattern of

species richness, with an increase in the number of

species toward higher elevations. Such pattern,

however, has already been reported for other anuran

communities (Heyer, 1967; Naniwadekar and

Vasudevan, 2007). The Shannon and Simpson

indexes are two measurements that show the

heterogeneity of diversity and the evenness of some

community (Krebs, 1999). While the Shannon index

focuses more on equitability between species,

Simpson focuses more on dominance between

common and rare species (Magurran, 2011). The

elevation levels of 300 m and 100 m showed lower

Shannon and Simpson diversity indexes for the SBS

method, with the Simpson diversity index of the 300 m

altitude level being higher than that of the 1000 m level

(Fig. 12 - Hill numbers q=1 and 2). For the SBS

method, S. imbegue was the dominant species at 100

m elevation (Fig. 14 A) and R. ornata at 300 m

elevation (Fig. 14 B). Even with lower species

richness for the PT method, the sampled elevations

100 m and 300 m had similar values of Shannon and

Simpson indexes with the elevation level 700 m (Fig.

13 - Hill numbers q=1 and 2), being R. ornata the

dominant species in both elevation levels (Fig. 14 E,

F). The elevation level 700 m had the highest

Shannon and Simpson diversity indexes, with

differences from all sampled elevation levels for SBS

and only from 1000 m for PT method (Fig. 12 and 13 -

Bol. Soc. Zool. Uruguay (2ª época). 2022. ISSN 2393-6940Vol. 31 (2): e31.2.4

Table 3. Results from asymptotic analysis comparing the Species richness (Chao Richness), Shannon diversity (Chao Shannon) and

Simpson diversity (Chao Simpson) among sampled localities: 1000, 700, 300 and 100; for each sampling method: SBS - Sampling at

Breeding Sites and PT - pitfall traps. S.E. - Standard Error, CI - confidence interval.

Method Sampled Locality Diversity index Observed Estimator S.E. Lower CI Upper CI

1000 Species richness 38,000 39,999 3,739 38,181 60,115

Shannon diversity 17,089 17,314 0,536 17,089 18,363

Simpson diversity 8,559 8,601 0,485 8,559 9,551

700 Species richness 36,000 38,247 3,392 36,266 55,016

Shannon diversity 24,322 24,972 0,706 24,322 26,356

Simpson diversity 18,857 19,325 0,932 18,857 21,152

300 Species richness 26,000 32,228 7,526 26,976 5,996

Shannon diversity 15,457 16,406 0,968 15,457 18,304

Simpson diversity 11,231 11,66 0,861 11,231 13,348

100 Species richness 24,000 24,998 1,868 24,093 5,048

Shannon diversity 13,752 14,073 0,541 13,752 15,135

Simpson diversity 9,928 10,094 0,602 9,928 11,274

1000 Species richness 9,000 14,936 7,05 9,944 46,32

Shannon diversity 4,644 5,026 0,504 4,644 6,014

Simpson diversity 3,618 3,723 0,379 3,618 4,465

700 Species richness 9,000 13,442 7,106 9,488 49,421

Shannon diversity 5,036 5,483 0,67 5,036 6,797

Simpson diversity 3,905 4,058 0,466 3,905 4,972

300 Species richness 14,000 14,665 1,303 14,057 21,789

Shannon diversity 5,734 5,845 0,302 5,734 6,437

Simpson diversity 4,099 4,132 0,249 4,099 4,62

100 Species richness 11,000 11,249 0,728 11,013 15,725

Shannon diversity 1,971 1,995 0,128 1,971 2,246

Simpson diversity 1,341 1,342 0,049 1,341 1,437

P T S B S

MARIOTTO, L.R. et al.

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MARIOTTO, L.R. et al. 33

Fig. 15. Dendrogram from cluster analysis with the Jaccard dissimilarity index (1-Jaccard) of amphibian species composition of studied areas in subtropical Atlantic

Forest. The red line represents the criterion of 40% dissimilarity.

Hill numbers q=1 and 2) due to the high uniformity of

species abundance distribution (Fig. 14 C, G). Even

with higher species richness, the sampled elevation

level 1000 m had lower Shannon and Simpson

indexes (Fig.s 12 and 13 - Hill numbers q=1 e 2) due

to the high dominance of D. minutus (SBS method)

and P. lateristriga (PT method) (Fig. 14 D, H). These

results may be related to the quality of the

environment, as the elevation level 1000 m has some

negative environmental impacts (see below).

Even if Serra Dona Francisca is characterized by

formations of Serra Geral and Serra do Mar mountain

ranges (Scheibe, 1986, Peluso, 1991), this locality

grouped (Fig. 15) with Joinville municipality (state of

Santa Catarina), Reserva Natural Salto Morato and

Morretes municipality (both in the state of Paraná), all

of which are characterized by sub-montane and

montane ombrophilous dense forest (Straube and

Urben-Filho, 2005, Armstrong and Conte, 2010,

Garey and Hartmann, 2012, Comitti, 2017). Even with

high dissimilarity in the cluster analysis (Fig. 15), the

anuran species composition of Serra Dona Francisca

is closer to north-eastern Serra do Mar mountain

range than to any other area in the south-eastern

region of the state of Santa Catarina (in Serra Geral

mountain range, such as Parque Estadual Serra do

Bol. Soc. Zool. Uruguay (2ª época). 2022. ISSN 2393-6940Vol. 31 (2): e31.2.4

Fig. 16. Regression of the similarity matrix of amphibian species compositions (Jaccard index) with the

spatial distance matrix (latitudinal and longitudinal coordinates) of communities from subtropical Atlantic

Forest. (Mantel Test - r=0.312, p=0.004).

MARIOTTO, L.R. et al.

Bol. Soc. Zool. Uruguay (2ª época). 2022. Vol. 31 (2): e31.2.4 ISSN 2393-6940

MARIOTTO, L.R. et al. 35

Fig. 17. Maps from Google Earth showing the advance of exploitation kaolin mining (red circle) in the region 1000

meters of the Serra Dona Francisca. (A) Image from 9th February 2014 of sampling site A21 and (B) image from

7th September 2018 of the same site.

Tabuleiro, Parque Estadual Serra Furada and

Siderópolis, see Table 1 of Suppl. Info. 1 for additional

information on these localities). Other nearby

localities, such as Fazenda Gralha-Azul (Fazenda Rio

Grande municipality), São José dos Pinhais, Curitiba

and Campo Largo municipalities, all in state of

Paraná, are characterized by mixed ombrophilous

forest (Araucaria Forest) and steppe vegetation, but

these localities were grouped with regions of the same

vegetation type in the state of Paraná. Finally, the

species composition of anuran communities in the

subtropical Atlantic Forest is influenced by

geographical distance (Mantel Test - r = 0.312, p =

0.004) (Fig. 16), that is, more distant locations present

more dissimilar species compositions, a pattern

evidenced in previous studies on anuran communities

(e.g., Bertoluci et al., 2007, Iop et al., 2011).

3. New Distribution Records

We provided four new records for the region:

Bokermannohyla luctuosa - distribution extended 385

km from the nearest record (São Paulo municipality,

state of São Paulo, Pombal and Haddad, 1993); Scinax

aromothyella - distribution extended 70 km from the

nearest record (Rio Negro municipality, state of

Paraná, Santos and Conte, 2014); Odontophrynus

americanus - distribution extended 65 km from the

nearest record (Fazenda Rio Grande municipality,

state of Paraná, Conte and Rossa-Feres, 2006); and

Leptodactylus plaumanni - distribution extended 200

km from the nearest record (Caçador municipality,

state of Santa Catarina, Lucas, 2008).

4. Spatial and Temporal Distribution

Out of the 14 exclusive anuran species at the

elevation level of 1000 m (Table 2), only four are usually

associated with vegetation in open areas (grasses and

shrub vegetation with few trees): S. aromothyella

(Faivovich, 2005), S. squalirostris (Gallardo, 1961), L.

plaumanni (Kwet et al., 2001) and O. americanus

(Savage and Cei, 1965); the other ten species are

usually associated with highland landscapes (Garcia,

Caramaschi and Kwet, 2001, Araújo, Bocchiglieri and

Holmes, 2007, Toledo et al. 2007, Lucas, 2008),

although Scinax granulatus and Physalaemus nanus

present a wider and more generalized distribution

(Conte et al., 2010, Kwet et al., 2010). Breeding sites in

grassland vegetation have a lower number of available

microhabitats compared to breeding sites in forest due

to the vertical stratification of the latter. Furthermore,

the highlands of Serra Dona Francisca present

different climatic conditions, with lower temperatures,

more lentic and less lotic breeding sites, compared to

the rugged terrain of the mountain climb.

We also registered exclusive species at the

remaining elevations levels (Table 2), characterized

mainly by forest and stream environments: five species

at the elevation level of 700 m - B. sulfuratus, P.

subguttata, Crossodactylus sp., A. ehrhardti and

Dendrophryniscus berthalutzae; four species at the

elevation level of 300 m elevation - Ischnocnema sp.

(gr. lactea), H. binotatus, C. izecksohni and

Dendropsophus berthalutzae; and two species at the

elevation level of 100 m elevation - S. tymbamirim and

L. catesbeianus. Furthermore, we registered five other

species exclusive to these three elevation levels, which

are not usually registered in the Araucaria Forest: F.

mitus, B. hylax, D. werneri, S. imbegue and C.

leucosticta (Conte, 2010).

Ten anuran species were registered at all elevation

levels (Table 2): R. ornata, R. icterica, B. faber, O.

rizibilis, S. perereca, A. nana, L. paranaru, L.

notoaktites, P. lateristriga and P. boiei (Table 2).

Elachistocleis bicolor was registered at the 100 m and

1000 m levels, with a distribution gap of 300 to 700 m

levels (Table 2). This result may be due to differences

in forest vegetation types, anthropic disturbance along

the entire elevation gradient, and the rugged

landscape from 300 m to 700 m elevation levels,

which is markedly different from lowlands (100 m) and

highlands (1000 m).

Out of the 74 known reproductive modes among all

amphibian species (sensu Nunes-de-Almeida et al.,

2021), we registered 21 modes for the species found in

Serra Dona Francisca, with predominance of mode 1,

which is the most common type among anuran species

(Duellman and Trueb, 1994). This result can be

explained by the fact that the frogs in the study area are

mainly inhabitants of lentic environments. We detected

46 anuran species in lentic environments, 22 species in

lotic environments and 17 species with breeding

activity in the forest.

Most hylid species were found occupying open

areas and forest edges habitats, except for

Aplastodiscus sp., A. ehrhardti and Bokermannohyla

spp., which were found in forest interior habitats. Even

with the arboreal habits, the breeding sites of the hylid

species were in lentic environments, most of them

located in forest edges or open areas. Several anuran

species registered in the forest interior present

reproductive modes associated with the forest floor,

s t r e a m s , o r e p i p h y t e v e g e t a t i o n ( e . g . ,

Br a c hy c eph a lid a e s p p ., H y l od i d ae s p p .,

Cycloramphidae spp., Hemiphractidae spp.,

Adenomera spp., Proceratophrys spp., Vitreorana

u r a n o s c o p a , C h i a s m o c l e i s l e u c o s t i c t a ,

Dendrophryniscus berthalutzae, Haddadus binotatus,

and Melanophryniscus xanthostomus). Only four

species occupied all habitats: Rhinella spp.,

Trachycephalus mesophaeus and Scinax perereca.

There are 19 forest-dependent species, 13 open-

area-dependent species, four generalist species, and

the remaining 27 species have some association with

forest or open areas. Therefore, it is relevant for the

amphibians to preserve both landscape matrixes

(Foerster and Conte, 2018), even with some anthropic

Bol. Soc. Zool. Uruguay (2ª época). 2022. ISSN 2393-6940Vol. 31 (2): e31.2.4

MARIOTTO, L.R. et al.

Bol. Soc. Zool. Uruguay (2ª época). 2022. ISSN 2393-6940Vol. 31 (2): e31.2.4

Table 4. Temporal distribution of amphibian species of Serra Dona Francisca, Joinville and Campo Alegre municipalities, state of

Santa Catarina, Brazil, between October 2012 and January 2014. Shades are the campaigns where we recorded the amphibians

foraging, calling or any other behaviour.

2012 2013 2014

Species/Campaign O O N N D D J J F F M M A M J J A S O N D J

Adenomera araucaria

Adenomera nana

Aplastodiscus sp.

Aplastodiscus ehrhardti

Aplastodiscus perviridis

Boana albomarginata

Boana albopunctata

Boana bischoffi

Boana faber

Boana semilineata

Boana sp.(gr. pulchella)

Bokermannohyla hylax

Bokermannohyla luctuosa

Brachycephalus sulphuratus

Chiasmocleis leucosticta

Crossodactylus sp.

Cycloramphus bolitoglossus

Cycloramphus izecksohni

Dendrophryniscus berthalutzae

Dendropsophus elegans

Dendropsophus microp

Dendropsophus berthalutzae

Dendropsophus minutus

Dendropsophus nahdereri

Dendropsophus werneri

Elachistocleis bicolor

Fritziana mitus

Fritziana sp.

Haddadus binotatus

Hylodes perplicatus

Ischnocnema henselii

Ischnocnema sp. (guentheri group)

Ischnocnema sp. (I. lactae species series)

Leptodactylus cf. latrans

Leptodactylus notoaktites

Leptodactylus plaumanni

37 Anuran Natural History from Serra Dona Francisca

Table 4. Cont.

2012 2013 2014

Species/Campaign O O N N D D J J F F M M A M J J A S O N D J

Richness 37 43 51 42 42 46 41 43 36 36 31 30 10 3 7 8 9 24 27 37 38 36

influence (Mendenhall et al., 2014).

Regarding the calling activity, we observed an

increase in the number of species during spring and

summer, and a decrease during autumn and winter,

an evident pattern of temporal distribution for the

subtropical Atlantic Forest (Conte and Machado,

2005, Conte and Rossa-Feres, 2007, Santos and

Conte, 2014). Nevertheless, six species showed

some activity (foraging or calling) during autumn: B.

luctuosa, Crossodactylus sp., Dendrophryniscus

berthalutzae, F. mitus, S. aromothyella, P. cuvieri;

eight species during winter: A. ehrhardti, B. faber, D.

minutus, D. nahdereri, R. icterica, S. imbegue, S.

sawayae and S. surdus; and three species only

during spring: Brachycephalus sulfuratus,

Ischnocnema sp. (gr. lactae) and P. distincta (Table

4). Usually in seasonal environments, the rainy

season is the main abiotic factor associated with

amphibian's reproduction (Arzabe, 1999, Prado et al.,

2004). Even without a dry season, the subtropical

Atlantic Forest present lower temperatures during

autumn and winter, with lower precipitation in autumn

(Conte and Machado, 2005, Conte and Rossa-Feres,

2007, Santos and Conte, 2014). This evidence

Bol. Soc. Zool. Uruguay (2ª época). 2022. ISSN 2393-6940Vol. 31 (2): e31.2.4

Melanophryniscus xanthostomus

Odontoprhynus americanus

Scinax aromothyella

Scinax catharinae

Scinax aff perpusillus

Scinax litorallis

Scinax rizibilis

Phyllomedusa disticta

Physalaemus aff. gracilis

Physalaemus cuvieri

Physalaemus lateristriga

Physalaemus nanus

Proceratophrys boiei

Proceratophrys subguttata

Rhinella ornata

Rhinella icterica

Scinax fuscovarius

Scinax granulatus

Scinax imbegue

Scinax perereca

Scinax squalirostris

Scinax tymbamirim

Scythrophrys sawayae

Sphaenorhynchus surdus

Trachycephalus mesophaeus

Vitreorana uranoscopa

MARIOTTO, L.R. et al.

reinforces the temperature as the main abiotic factor

structuring amphibian communities in subtropical

environments (Ceron et al., 2020).

Classifying the anuran species as continuous,

prolonged, or explosive breeders is not an easy task.

Many studies have used the calling behaviour of

males to identify the period of breeding activity

(Bertoluci and Rodrigues, 2002, Prado et al., 2004,

Conte and Rossa-Ferres, 2007). However, the

reproduction event, when males mate with ovigerous

females and lay their clutch eggs in the environment

or when tadpoles or juveniles are present, is more

restrict to a few days or months. Unlike seasonal

environments such as the Caatinga (Arzabe, 1999)

or the Pantanal (Prado et al., 2004) with many

explosive breeding species, in Serra Dona Francisca

we found only five explosive breeders: M.

xanthostomus, S. sawayae, S. aromothyella, T.

mesophaeus and C. leucosticta. Only B. bischoffi

presented continuous breeding behaviour

considering the male's annual calling period (except

May 2013). However, if we consider only the period

in which we observed ovigerous females, B. bischoffi

is a prolonged breeder with the reproductive period

restricted to hot and rainy months (October 2012 to

March 2013). We identified eight species as

prolonged breeders, with reproduction activity for

only four to six months during the hot and wet

months: R. ornata, B. faber, A. nana, A. araucaria,

Leptodactylus paranaru, L. notoaktites, P. lateristriga,

and P. boiei. Another 25 species presented four to six

months of calling activity, but direct evidences of

reproduction (presence of females, amplected

couples, spawning, tadpoles or juveniles) were

restricted to one or two months: Aplastodiscus sp., A.

ehrhardti, A. perviridis, B. albomarginata, B.

albopunctata, B. semilineata, B. hylax, B. luctuosa,

C. izecksohni, Dendrophryniscus berthalutzae, D.

elegans, D. microps, D. berthalutzae, D. minutus, D.

nahdereri, D. werneri, E. bicolor, F. mitus, I. henselii,

Ischnocnema sp. (gr. guentheri), L. plaumanni, S.

rizibilis, P. cuvieri, Physalaemus aff. gracilis, and R.

icterica.

5. Conservation of Serra Dona Francisca

Considering the field records and literature data,

the Serra Dona Francisca region, which covers the

Corupá, Campo Alegre, São Bento do Sul, Joinville

and Schroeder municipalities, has the highest anuran

diversity in the state of Santa Catarina, and one of

the greatest species richness of the subtropical

Atlantic Forest (total of 87 species, considering our

results and literature data; Lucas, 2008, Conte et al.,

2010), highlighting the paramount importance of this

region for anuran conservation. There are many

sustainable conservation units in the region (e.g.,

Área de Preservação Ambiental Serra Dona

Francisca, Área de Relevante Interesse Ecológico

Morro do Iriri, Parque Natural Municipal da Caieira,

and Reserva do Desenvolvimento Sustentável da

Ilha do Morro do Amaral, all in Joinville municipality,

and Área de Preservação Ambiental Estadual de

Guaratuba, in Guaratuba municipality) that contribute

to the conservation of the area, especially by the

inclusion of the local population in sustainable

economic activities, such as hotels, inns, restaurants

and recreational activities focusing on ecotourism.

We observed some illegal activities in Serra Dona

Francisca, including hunting and palm heart

exploitation, and an activity that causes great impacts

to the environment: kaolin mining. A visual impact of

the mining activity is the suppression of the

grassland, a steppe phytophysiognomy used by

anurans as breeding sites, which affects all species

that occupy open landscapes. This activity

culminates in the deposition of sediments and

suppression of breeding sites. Observing Google

images for five years (2012-2017), we notice an

increase in mining activities in the vicinity of the

breeding sites (Fig. 17). In Serra Dona Francisca,

most of the sampled environments have some

associated human activity such as farmlands or small

pastures. However, the intensity of human activities

associated with these environments should be

investigated, as it can negatively impact amphibian

communities, especially with chemical inputs on

agricultural lands (Mendenhall et al., 2014) or

chemical residuals from some types of activities (De

Lucca et al., 2017).

We believe that the creation of an integral protection

conservation unit in the region would help to stop

mining exploitation but keeping the local population in a

sustainable conservation unit and investing in

environmental education might be the best

conservation actions for the region.

ACKNOWLEDGEMENTS

We are grateful to all those contributed with this

study, especially to Carlos Eduardo Conte, Maria

Vitória Giglio, Jaqueline Reginato Koser, Guilherme

Grazzini, Daniel Son, Maria Luiza Fausto and family,

and all the residents of Serra Dona Francisca. To the

Universidade Federal do Paraná and Universidade

Regional de Blumenau by the structure support for

this research. To the Conselho Nacional de

Desenvolvimento Científico e Tecnológico (CNPq

processes 132559/2012-9, 167888/2014-5 and

311492/2017-7), Coordenação de Aperfeiçoamento

de Pessoal de Nível Superior (CAPES), and

Fundação Boticário for grants. To the SISBIO (nº

35005) and FUNDEMA (nº 013/12-GEMAP) for the

research licenses and to PLASVALE for the plastic

buckets for used as pitfall traps.

Authors' Contributions

Lucas Ribeiro Mariotto. Substantial contribution in

Bol. Soc. Zool. Uruguay (2ª época). 2022. ISSN 2393-6940Vol. 31 (2): e31.2.4

39 Anuran Natural History from Serra Dona Francisca

the concept and design of the study; Contribution to

data collection; Contribution to data analysis and

interpretation; Contribution to manuscript preparation;

Contribution to critical revision, adding intellectual

content.

Sarah Mângia. Contribution to data analysis and

interpretation; Contribution to manuscript preparation;

Contribution to critical revision, adding intellectual

content.

Diego José Santana. Contribution to data analysis

and interpretation; Contribution to manuscript

preparation; Contribution to critical revision, adding

intellectual content.

Conflicts of Interest

“The author(s) declare(s) that they have no conflict of

interest related to the publication of this manuscript”

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