Description and Shaping Factors of Diet and Feeding Ecology of Neotropical Tadpoles: A Case Study and A Comprehensive Review

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Description and shaping factors of diet and feeding ecology of neotropical

tadpoles: A case study and a comprehensive review
Article in Austral Ecology · March 2023

DOI: 10.1111/aec.13302
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Received: 17 August 2022 Revised: 17 January 2023 Accepted: 30 January 2023
DOI: 10.1111/aec.13302
RESEARCH ARTICLE
Description and shaping factors of diet and feeding

ecology of neotropical tadpoles: A case study and a
comprehensive review
Iara Matias Gonçalves1 | Jéssica Stephanie Kloh2 | Katharina Ruthsatz3 |

Cleber Cunha Figueredo2 | Paula Cabral Eterovick1,3
1
Programa de Pós-Graduação em Abstract
Biologia de Vertebrados, Pontifícia
Universidade Católica de Minas Gerais,
Tadpoles are important components of many aquatic food webs as they influ-
Belo Horizonte, Brazil ence populations at other trophic levels and, ultimately, energy transfer between
2
Programa de Pós-Graduação em aquatic and terrestrial ecosystems. What tadpoles consume is a key factor to
Ecologia, Conservação e Manejo da Vida their ecological functions. Neotropical anuran assemblages include many spe-
Silvestre da Universidade Federal de
Minas Gerais, Belo Horizonte, Brazil
cies with a larval stage, however, the diets of neotropical tadpoles are relatively
3
Zoologisches Institut –Technische
poorly known. We performed a systematic review on the available literature on
Universität Braunschweig, Braunschweig, in situ studies of neotropical tadpole diets and found out that most studies are
Germany descriptive and temporally restricted, failing to consider seasonal variations.
Correspondence
Studies frequently also fail to consider possible ontogenetic dietary variations
Paula Cabral Eterovick, Zoologisches or tadpoles' capacity to select food based on what is available to them in their
Institut –Technische Universität habitats. We contributed to this topic by investigating seasonal variations and
Braunschweig, Braunschweig, Germany.
Email: [email protected]
comparing the diets of two developmental stages of Scinax curicica tadpoles.
Tadpole diet varied between dry and rainy seasons, but not between the studied
Funding information developmental stages. These results, together with published studies, suggest
Conselho Nacional de Desenvolvimento
Científico e Tecnológico, Grant/Award
that multiple ecological/physiological factors are likely to influence tadpole food
Number: 309513/2020-0 intake and should not be overlooked. Studies on neotropical tadpole feeding
ecology, digestion capacity, physiological effects of different diets and microbi-
ome versus diet interactions are highly desirable but largely unexplored. Such
studies are likely to add valuable information to understand tadpole roles in
aquatic food webs, their growth and development performance resulting from
food acquisition and assimilation, and consequently their effects on other spe-
cies in aquatic and terrestrial ecosystems.
KEYWORDS
diet, food selectivity, neotropical region, ontogenetic variation, seasonality, tadpole
I N TROD UC T I O N (Costa & Vonesh, 2013; Hite et al., 2018). As tad-

poles show trophic plasticity in response to biotic
Tadpoles play important roles in nutrient cycling, en- (Arribas et al., 2015; Caut et al., 2013) and abiotic fac-
ergy flow and bioturbation (Montaña et al., 2019). tors (Carreira et al., 2016), it is also important to take
They are important components of aquatic ecosys- these into account to understand their trophic roles
tems (Iwai & Kagaya, 2005) and influence nutrient (Montaña et al., 2019). Tadpoles can be herbivores,
transfer between aquatic and terrestrial food webs omnivores, carnivores or detritivores (Altig et al., 2007;
This is an open access article under the terms of the Creative Commons Attribution-NonCommercial-NoDerivs License, which permits use and distribution in
any medium, provided the original work is properly cited, the use is non-commercial and no modifications or adaptations are made.
© 2023 The Authors. Austral Ecology published by John Wiley & Sons Australia, Ltd on behalf of Ecological Society of Australia.
Austral Ecology. 2023;00:1–13. |

wileyonlinelibrary.com/journal/aec 1
|
2 GONÇALVES et al.
Schiesari, 2006), but their diets usually include both autotrophic and het-
erotrophic organisms (Vera Candioti, 2007).
Temperature changes, for instance, may make herbivory more advan-
tageous than omnivory (Carreira et al., 2016) and alter food availability,
leading to seasonal diet variations, which are known to exist for some tad-
pole species (Kloh et al., 2018). Food availability also varies among micro-
habitats and tadpoles occupying specific microhabitats that are favoured
by their ecomorphotypes will be exposed to the food available there (Kloh
et al., 2019).
Like the food ingested by tadpoles (Kupferberg, 1997), the microbial
communities in animal guts have important implications to animal physiol-
ogy and development (Tuddenham & Sears, 2015). These microbial com-
munities are affected by the host's genetic background, immune system,
age and diet (Tuddenham & Sears, 2015). This association allowed animals
to successfully explore a variety of food sources that would not be avail-
able otherwise, and to amplify their metabolic potential intertwining their
genomes with their microbiome (McFall-Ngai et al., 2013). Besides, animal
microbiomes are associated with the habitats occupied by their hosts and
may influence tadpole development (Vences et al., 2016).
Considering the great diversity of anuran species with a tadpole stage
in the neotropics (Provete et al., 2012), studies on tadpole natural history
and ecology are still scarce, despite their ecological importance (Andrade
et al., 2007; Rossa-Feres et al., 2015). Here, we summarize knowledge
on neotropical tadpole diets and describe the diet of tadpoles of Scinax
curicica (Hylidae) addressing two knowledge gaps: seasonal variability
and developmental-stage-related variability in food ingestion. We chose
Scinax curicica as a model because it has a nektonic tadpole, and nek-
tonic tadpoles have a diversified diet compared to other ecomorphotypes
(e.g. Antoniazzi et al., 2020). Besides, S. curicica tadpoles can be found at
different seasons and developmental stages.
MATER IAL S AN D M E T H ODS
Literature review
We conducted a non-automated systematic literature review applying the

PRISMA methodology (Page et al., 2021). We first used Clarivate Web of
Science, (© Clarivate 2022) to search for publications containing “tadpole”
and “diet” (considering all fields) and selected within the results only those
studies conducted in South and Central America countries, as well as in
Mexico, without timespan restrictions. This yielded 87 papers. After check-
ing their titles and abstracts, we selected 26 papers that were performed
with wild, native tadpoles, excluding experiments aimed at testing artificial
diets for raising Lithobates catesbeianus (Ranidae) for commercial pur-
poses or a few others that used artificial, manipulated diets to study their
effects on tadpoles. After reading the full texts, we excluded one more
paper that included tadpoles from South America but did not analyse their
diet. We performed complementary searches in Scopus (© Elsevier 2022)
using the same criteria, and retrieved 86 papers, from which we screened
24, all of them but one already obtained from Web of Science. We also
searched SciELO (Scientific Electronic Library Online, Brazil) using “tad-
pole” and “diet”, which resulted in five papers, two of which met our selec-
tion criteria and one of which was not previously found in Web of Science
or Scopus (Figure 1). The searches were conducted during March 2022
and re-done on December 6, 2022, to check for eventual new publications.
The literature review was led by P. C. Eterovick; the other authors also
DIET OF NEOTROPICAL TADPOLES
3|
Identification of studies via databases
Records identified from
Identification
databases (n = 178)
Web of Science: 87
Scopus: 86
Scielo: 5
Criteria for exclusion: no

information on tadpole diet in
natural habitats
Records excluded based on title

Web of Science: 87 and abstract (n = 60)
Record excluded after reading
the whole text (n=1)
Screening

Scopus: 86 and abstract (n = 62)
Duplicate records removed after
screening (n = 23)

Scielo: 5 and abstract (n = 2)
Duplicate records removed after
screening (n = 2)
Included
Studies included in review

(n = 28)
F I G U R E 1 PRISMA diagram (sensu Page et al., 2021) showing the steps of the systematic literature review performed on diets of
neotropical tadpoles in their native habitats.
searched additional sources (i.e. Google Scholar, reference lists, personal

communications) and checked whether selected papers were included in
the results obtained from the systematic review. No additional papers fulfill-
ing our criteria were found in these other sources.
Diet of Scinax curicica tadpoles
Scinax curicica is a treefrog endemic to the Espinhaço Mountain Range

in southeastern Brazil (Eterovick et al., 2020). It breeds usually in puddles
and ponds in high altitude areas from September to February. Tadpoles are
diurnal, nektonic and can be found year-round (Eterovick et al., 2020). We
collected tadpoles from a backwater of a temporary stream (19°17′28.7″ S
43°34′39.8″ W; 1374 m a.s.l.) at the surroundings of the National Park Serra
do Cipó on January 14 (rainy season) and September 21 (dry season)
2020. The rainy season in the area extends from October to March and
the dry season from April to September. Rainfall varies between 1460 and
2490 mm and mean temperatures range from 13 to 29°C (Viveros, 2010).
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4 GONÇALVES et al.
We collected 20 tadpoles in each period, 10 of them in Gosner (1960)

stage 25 and the other 10 in stage 30. The tadpoles were immediately
euthanized in xylocaine 10% (Pharlab Industria Farmaceutica SA) and
then preserved in 10% formalin (Kloh et al., 2019). We confirmed tadpole
stages under stereomicroscope at the laboratory. Next we removed the
tadpole's guts and used material extracted from the anterior third of the gut
to quantify ingested items. The material was diluted in 1 mL distilled water
and placed in a Sedgewick Rafter counting chamber and examined under
optic microscope (Olympus cx40) under 400× magnification (following Kloh
et al., 2019). We identified ingested items based on specialized literature
(Bicudo & Menezes, 2005; Silva et al., 2020).
We compared the gut contents of S. curicica tadpoles between the
two developmental stages (25 and 30) and two seasons (rainy and dry;
i.e., January and September). We adjusted the data on individual tadpole
diets (numbers of items consumed for each food category) to two axes
representing latent variables and built a model to test for effects of the
studied developmental stages and seasons on the food items consumed
by tadpoles. For the modelling we used the packages mvabund (Wang
et al., 2022) and ecoCopula (Popovic et al., 2019) in R version 4.2.2. (R
Core Team, 2021).
RESULT S
Literature review
The search for publications on neotropical tadpole diets resulted in 28 pa-

pers, 14 of which reporting on studies conducted in Brazil, 13 in Argentina
(one of them also including species from Panama) and one in Mexico
(Figure 2a, Appendix S1). Tadpoles of 63 species had their diets studied
at least on one occasion/habitat (Figure 2b). Most studies focused only on
the visual analyses of gut contents, just two included also stable isotope
analyses. Only four studies considered temporal variability in tadpole diet
composition, four searched for possible differences among tadpole devel-
opmental stages and seven evaluated food preferences based on food
availability quantification. Just one study approached tadpole microbiomes
but associated gut content analyses were not performed for the neotropical
tadpoles included.
Scinax curicica tadpoles ate unicellular and filamentous Bacillariophyceae

and Zygnematophyceae, Euglenophyceae, Cyanophyceae, Fungi,
Spermatophyta (pollen) and Testate Amoeba (Figure 3, Appendix S1).
Some amorphous material could also be seen in tadpole guts and may
have resulted from items that are quickly digested or most likely from
ingestion of amorphous material itself. Tadpole diets differed between
seasons (dev = 29.666, p = 0.001). Developmental stages did not ex-
plain diet variation (dev = 8.099, p = 0.201), nor interacted with sea-
sons (dev = 11.963, p = 0.076). According to univariate tests, the food
items that contributed to variability among tadpoles at different seasons
were testate amoeba (dev = 13.276, p = 0.001) and Euglenophyceae
(dev = 11.047, p = 0.001; Figures 3), both consumed at higher amounts
in the rainy season.
DIET OF NEOTROPICAL TADPOLES
5|
F I G U R E 2 Countries where studies on tadpole diets were conducted (and from where the tadpoles are native; a), and taxonomic
distribution of studied tadpoles (b) based on a systematic review of the literature with no time restriction. Tadpoles of 63 species had
their diets studied at least on one occasion/habitat, most of them in Brazil and Argentina. Taxa included six species in one genus
from Bufonidae (Rhinella arenarum, R. diptycha, R. fernandezae, R. jimi, R. rubescens and R. spinulosa), four species in three
genera from Ceratophryidae (Chacophrys pierottii, Ceratophrys cranwelli, Lepidobatrachus laevis, and L. llanensis), 34 species in 12
genera from Hylidae (Aplastodiscus cavicola, A. sibilatus, Boana cordobae, B. pulchella, B. rosenbergi, Bokermannohyla alvarengai,
B. capra, B. nanuzae, B. saxicola, Dendropsophus sp., D. microcephalus, D. nanus, Lysapsus limellum, Pithecopus hypochondrialis,
P. megacephalus, P. nordestinus, Phasmahyla jandaia, Phyllomedusa sauvagii, Pseudis minuta, P. paradoxa, Scinax angrensis,
S. boulengeri, S. fuscovarius, S. fuscomarginatus, S. granulatus, S. littoreus, S. machadoi, S. nasicus, S. perpusillus, S. x-signatus,
S. similis, Trachycephalus cunauaru, T. typhonius and Triprion petasatus), 13 species in three genera from Leptodactylidae (Alsodes
gargola, Leptodactylus ocellatus macrosternum, L. bolivianus, L. fuscus, L. natalensis, L. podicipinus, Physalaemus albonotatus,
P. biligonigerus, P. centralis, P. cuvieri, P. fuscomaculatus, P. nattereri and P. santafecinus), three species in two genera from Microhylidae
(Dermatonotus muelleri, Elaschistocleis bicolor and E. ovalis), two species in two genera from Odontophrynidae (Odontophrynus
americanus, Macrogenioglotus alipioi) and one species from Telmatobiidae (Telmatobius cf. atacamensis). Most studies were based in
visual inspection of gut contents and did not approach seasonal or developmental stage variability in diets.
F I G U R E 3 Multivariate comparison of the diets of S. curicica tadpoles according to types and amounts of consumed items in two
developmental stages (25 and 30, sensu Gosner, 1960) and in two seasons (rainy and dry). Testate amoeba and Euglenophyceae were
more consumed in the rainy season than in the dry season, leading to seasonal variation in diet composition. Diet did not vary between
developmental stages.
D I SCUS S I O N
Neotropical tadpole diet composition
Most of the studies on neotropical tadpole diet analysed gut contents

visually, as already noticed by Montaña et al. (2019). Only two studies
employed stable isotope analyses (Antoniazzi et al., 2020; Huckembeck
et al., 2014), a technique that gives a more realistic picture of what is
|
6 GONÇALVES et al.
actually assimilated by tadpoles. Based on available studies, diatoms

(Bacillariophyta, Bacillariophyceae) were usually the numerically pre-
dominant items in the diets of most neotropical tadpoles (e.g. Baffico &
Úbeda, 2006; Kloh et al., 2019; Protázio et al., 2020; Santos et al., 2016).
The most remarkable exceptions were the predaceous tadpole of
Ceratophryidae, which ate insects, crustaceans and tadpoles (Fabrezi
& Cruz, 2020; Vera Candioti, 2005), the bromeliad-dwelling tadpoles of
Scinax littoreus and S. perpusillus, which ate mainly Cyanophyceae and
Zygnematophyceae respectively (Sabagh et al., 2012) and the tree hole
breeder Trachycephalus cunauaru, which ate mainly conspecific eggs
(Schiesari et al., 2003). Tadpoles also had other items besides algae in
their guts, however, usually in small proportions, such as fungi (Protázio
et al., 2020), testate amoeba (Kloh et al., 2018, 2019; Pollo et al., 2019;
Sabagh et al., 2012; Vera Candioti, 2005; Vera Candioti, 2007), insect
remains/eggs/larvae (Dutra & Callisto, 2005; Pollo et al., 2019; Protázio
et al., 2020; Sabagh et al., 2012), Rotifera, Cilliophora, Nematoda,
Crustacea, Oligochaeta (Verburg et al., 2007), pollen (Kloh et al., 2019;
Protázio et al., 2020; Sabagh et al., 2012). The proportion of diatoms in
tadpole diets may be overestimated because many studies analysed the
whole digestive tract and the capsules of diatoms can be identified even
after digestion, whereas other items are digested and not seen in poste-
rior portions of the gut (Kloh et al., 2018). However, even in studies that
considered just the first third of the gut (e.g. Babini et al., 2017; Baffico &
Úbeda, 2006; Bionda et al., 2012, 2013; Kloh et al., 2018), where the pro-
portions of ingested items can be observed with the least digestion (Kloh
et al., 2018), diatoms were the main item in the diets of several species.
Bacillariophyceae are periphytic algae and their consumption by benthic
tadpoles is expected, as these tadpoles supposedly feed on the bottom
(Altig & Johnston, 1989), although nektonic tadpoles have also been re-
ported to consume large amounts of these algae (e.g. Pithecopus mega-
cephalus; Kloh et al., 2019).
The role of tadpole ecomorphotypes and microbiome

on diet
Tadpole ecomorphotypes are likely to have an important influence on

what tadpoles consume (Antoniazzi et al., 2020), as they relate to the use
of specific microhabitats which vary in food availability (Kloh et al., 2019).
Microhabitats have different surfaces for periphyton attachment (e.g.
Kloh et al., 2019), and substrate orientation can also influence food ac-
quisition by scraping tadpoles (Annibale et al., 2019). Consequently, the
diets of tadpoles reflect in a large extent the food availability and distri-
bution in the microhabitats they use (Kloh et al., 2019). For instance, the
neustonic tadpole of Phasmahyla jandaia has a diet largely differentiated
from those of other sympatric and syntopic benthic/nektonic stream tad-
poles (Kloh et al., 2019, 2021; Appendix S1). Isotopic signatures also dif-
fer among microbial feeders, herbivores and neuston feeders (Verburg
et al., 2007).
Interestingly, the composition of diets of tadpoles around the world
do not change considerably regarding the main taxonomic groups con-
sumed (e.g. Chen et al., 2008; Ocock et al., 2018; Pryor, 2014). The gut-
associated microbiota also showed similarities between tadpoles from
Brazil and Madagascar, corroborating microbiome and diet association
(Vences et al., 2016). Although Vences et al. (2016) did not analyse the diet
of Brazilian tadpoles, the tadpoles from Madagascar consumed mainly di-
atoms, with smaller proportions of algae, plant fragments, fungi and insect
DIET OF NEOTROPICAL TADPOLES |
7
fragments (Vences et al., 2016), a diet with similar composition to those of

many neotropical tadpoles (Appendix S1).
Temporal and ontogenetic variation in diet composition
Most studies did not test nor consider variations among developmental
stages, except for the study on the diets of Scinax angrensis (Sousa-Filho
et al., 2007), Bokermannohyla saxicola (Kloh et al., 2018) and Phasmahyla
jandaia tadpoles (Kloh et al., 2021). The diet of S. angrensis tadpoles var-
ied between developmental stages (Sousa-Filho et al., 2007) as also did
the diet of B. saxicola but, in this case, just considering number of ingested
items and not biovolume (Kloh et al., 2018). However, P. jandaia tadpoles
showed subtle variations in food preference among developmental stages,
but they could not be precisely assigned to developmental stage based on
their diets. This diet similarity throughout development may be related to
the more specialist habits of these tadpoles (Kloh et al., 2021). Variation
throughout tadpole developmental stages has also been recorded for tem-
perate species (Schriever & Williams, 2013). Regarding seasonal varia-
tion in food consumption, only the tadpoles of B. saxicola were compared
within the same developmental stage throughout the year, and showed
different diets, what may be related to food availability (Kloh et al., 2018).
Bromeliad-dwelling tadpoles of Scinax littoreus and S. perpusillus did not
show diet variation between dry and rainy seasons, however, a broad range
of developmental stages was pooled for the analyses (Sabagh et al., 2012).
Food selectivity and other knowledge gaps
Food selectivity has rarely been tested for neotropical tadpoles. Tadpoles
of Phasmahyla jandaia showed preference for pollen, testate amoeba and
Euglena (Kloh et al., 2021), tadpoles of Scinax littoreus preferred algae
and fungi, whereas S. perpusillus, in the same bromeliad phytotelmata
habitat, showed no preferences, eating food items in the same propor-
tions available (Sabagh et al., 2012). The diets of Brazilian stream tadpoles
of five species reflected the availability of food items in the microhabitats
they used, however, some items were consumed in larger proportions,
whereas others were apparently avoided, the discrepancy between avail-
able and consumed proportions being larger for some species than others
(e.g. Bokermannohyla saxicola; Kloh et al., 2019). Although tadpoles are
broadly assumed to be non-selective feeders (e.g. Dutra & Callisto, 2005;
Protázio et al., 2020), the available studies that accessed food availability
and compared to consumption by tadpoles indicate that this should not be
considered a general rule (e.g. Antoniazzi et al., 2020).
The data on diet composition of neotropical tadpoles are scarce con-
sidering their species diversity but relatively large compared to other
regions (Montaña et al., 2019). However, there are still important gaps
regarding feeding ecology, as previously noticed by Altig et al. (2007).
Tadpole flexibility in food acquisition based on the exploitation of avail-
able resources and in response to varying nutritional needs throughout
development is an interesting topic that needs to be further explored in
the neotropics, as well as in other regions (Montaña et al., 2019). Also,
digestibility of different food items was not accessed in any of the pub-
lished articles reporting natural diets of neotropical tadpoles, indicating
a huge knowledge gap. Some studies addressed assimilation efficiency
under different temperatures and diets (Benavides et al., 2005) or ef-
fects of different diets on growth, development (Toledo et al., 2014) and
|
8 GONÇALVES et al.
morphological plasticity (Castañeda et al., 2006; Ruthsatz et al., 2019,

2022), but they all used manipulated/artificial diets that did not corre-
spond to what tadpoles eat in situ. The investigation of natural diets is
important because even if nutritional composition of artificial diets mirror
natural ones, digestibility is likely to vary among food items (Pryor, 2014)
and play a role on the benefits of each diet, as assimilation of consumed
items vary (Schmidt et al., 2017).
Environmental impacts and tadpole diet composition
Besides natural sources of variation, environmental impacts may also influ-

ence tadpole food acquisition. A comparison of diets of Rhinella arenarum
and Physalaemus biligonigerus tadpoles from preserved areas versus
areas impacted by agriculture and livestock showed reduced food con-
sumption and growth in impacted areas with increased representation of
resistant taxa, suggesting that diet evaluation can also be an early indicator
on how habitat quality may affect tadpole growth and development (Babini
et al., 2017; Bionda et al., 2012, 2013). Testate amoebae are common in
low-quality aquatic habitats (Sousa, 2008) and were found in the diets of
many tadpoles (Babini et al., 2017; Kloh et al., 2018, 2021; Pollo et al., 2019;
Sabagh et al., 2012; Vera Candioti, 2005; Vera Candioti, 2007). This sug-
gests that, as a topic worth further exploration, increased consumption of
testate amoeba could be an indicator of decreasing habitat quality.
The diet of Scinax curicica tadpoles varied between rainy and dry sea-
sons. This result differs from results obtained for Bokermannohyla saxi-
cola, whose tadpoles had greater than expected niche overlap among
seasons (Kloh et al., 2018). However, stage-specific nutritional needs did
not seem to define diet composition for neither S. curicica nor B. saxicola
as tadpoles from both stages 25 and 30 had similar diets within seasons
(Kloh et al., 2018; this study). Scinax angrensis tadpoles also had similar
diets in two studied developmental stages (Sousa-Filho et al., 2007).
Phasmahyla jandaia tadpoles had very similar diets at three develop-
mental stages, however, analyses of food availability showed that they
have strong food selectivity, and their diet reflects ecomorphotype related
microhabitats (Kloh et al., 2021). Thus, it cannot be said that the seasonal
diet variation for S. curicica is simply due to the combination of indiscrimi-
nate consumption and expected seasonal variations in available food items
(e.g. Ferragut et al., 2010). The information available so far on tadpole feed-
ing ecology, although limited, indicates that both differences in microhabi-
tat use (Kloh et al., 2019) and food selectivity (e.g. Antoniazzi et al., 2020)
are likely to be important. The diets of tadpoles are thus likely influenced
by a complex combination of variables, which have frequently not been well
controlled in previous studies.
Regarding the algae found in the guts of S. curicica, there is little specific
information in the literature about their relevance for tadpoles. According
to Lee (2008), the green algae (Chlorophyta and Charophyta) are relevant
sources of carotenoids for animals that cannot synthesize these import-
ant antioxidant agents. Cyanobacteria is used as human and animal food
supplements, and diatoms (Bacillariophyta) are preferred by some inver-
tebrates (Lee, 2008), being highly nutritive due to their fatty acid content
(Lavens & Sorgeloos, 1996). Although there are specific differences in the
biochemical composition of different microalgal taxa, the protein fraction is
9
always the major organic constituent, commonly followed by lipid, and then
by carbohydrate (Lavens & Sorgeloos, 1996).
Testate amoeba and Euglenophyceae were the food items that contrib-
uted the most to the seasonal variation in the composition of S. curicica
tadpoles. The tests of testate amoeba are rich in iron and manganese
(Sousa, 2008), which are important for vertebrate immunity and metab-
olism (Pinto-Coelho, 2009). Their consumption by S. curicica tadpoles
was higher in the summer (rainy season). The cause of this increased con-
sumption remains to be investigated, as it could include higher availability
of this item in the habitat and/or higher selectivity by tadpoles, maybe due
to an increase in metabolism in the warmer season.
The presence of some amorphous material in the guts may indicate
other food sources with rapid digestion or organic material with no deter-
mined shape. For example, tadpoles can scrape the bodies of other dead
tadpoles (PCE, pers. obs.) and eat their own faeces (Pryor, 2014). The
proportion of amorphous material in relation to identifiable items increases
through the end of the gut (Kloh et al., 2018, JSK pers. obs.). Thus, we
optimized diet characterization by restricting our analyses to the first third
of the gut.
Concluding remarks
Studies on the diet of neotropical tadpoles (Appendix S1) approached in

just a few instances the effects of seasonality, developmental stage and
food selectivity as natural sources of variability in diet composition, but
the available data indicate none of these sources of variation should be
overlooked. The dynamic of food acquisition by tadpoles is likely to reflect
not only in their health but also in aquatic (Costa & Vonesh, 2013; Hite
et al., 2018; Schmidt et al., 2017) and terrestrial food webs, as diet influ-
ences populations of aquatic prey (Costa & Vonesh, 2013) as well as time to
metamorphosis (Díaz–Páez & Canales–Arévalo, 2018; Kupferberg, 1997).
In summary, studies on neotropical tadpole feeding ecology, digestion
capacity, physiological effects of different diets, and microbiome versus
diet interactions are highly desirable and still practically unexplored. Such
studies are likely to add valuable information to understand tadpole roles in
aquatic food webs, their growth and development performance. All these
parameters result from food acquisition and assimilation and have conse-
quent effects on other species in aquatic and terrestrial ecosystems.
AU T H O R C O N T R I B U T I O N S
Iara Matias Gonçalves: Conceptualization (equal); data curation (equal);
investigation (equal); writing –original draft (equal); writing –review and
editing (supporting). Jéssica S. Kloh: Conceptualization (equal); data
curation (equal); methodology (equal); project administration (equal); su-
pervision (equal); writing –original draft (supporting); writing –review and
editing (supporting). Katharina Ruthsatz: Data curation (supporting); vali-
dation (equal); writing –review and editing (supporting). Cleber Cunha
Figueredo: Data curation (supporting); investigation (equal); methodology
(equal); resources (equal); supervision (equal); writing –review and edit-
ing (supporting). Paula Eterovick: Conceptualization (equal); data curation
(equal); formal analysis (lead); methodology (equal); project administration
(equal); supervision (equal); writing –original draft (lead); writing –review
and editing (equal).
|
10 GONÇALVES et al.
AC K N OW L ED G EM E N T S

We are thankful to Yasmim Mossioli and Felipe S. F. Leite for suggestions in
a previous version of this manuscript, the Parque Nacional da Serra do Cipó
for support during the field trip, Conselho Nacional de Desenvolvimento
Científico e Tecnológico (CNPq) for a Scholarship for JSK and a Research
Productivity grant for PCE. Collection permits were provided by Sisbio/
ICMBio (64500) according to ethical guidelines. Open Access funding ena-
bled and organized by Projekt DEAL.
F U N D I N G I N FO R M AT I O N
This study was financed by Conselho Nacional de Desenvolvimento
Científico e Tecnológico (CNPq) through a Scholarship to JSK and a
Research Productivity grant to PCE. Collection permits were provided by
Sisbio/ICMBio (64500) according to ethical guidelines.
C O N F L I C T O F I N T E R E S T S TAT E M E N T
The authors declare no conflicts of interest.
DATA AVA I L A B I L I T Y S TAT E M E N T

The data sets generated and/or analysed during the current study are avail-
able from the corresponding author on reasonable request.
ORCID
Paula Cabral Eterovick https://orcid.org/0000-0002-1316-1255
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13
S U PP O R T I N G I N FO R M AT I O N
Additional supporting information can be found online in the Supporting How to cite this article:
Information section at the end of this article. Gonçalves, I.M., Kloh, J.S.,
Ruthsatz, K., Figueredo, C.C.
& Eterovick, P.C. (2023)
Description and shaping
factors of diet and feeding
ecology of neotropical
tadpoles: A case study and a
comprehensive review.
Austral Ecology, 00, 1–13.
Available from: https://doi.
org/10.1111/aec.13302
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Description and shaping factors of diet and feeding ecology of neotropical

Article in Austral Ecology · March 2023

Jessica Stephanie Kloh Katharina Ruthsatz

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Paula Cabral Eterovick

Amphibians in a changing world: An ecophysiological perspective on amphibian metamorphosis View project

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Description and shaping factors of diet and feeding

Iara Matias Gonçalves1 | Jéssica Stephanie Kloh2 | Katharina Ruthsatz3 |

I N TROD UC T I O N (Costa & Vonesh, 2013; Hite et al., 2018). As tad-

Austral Ecology. 2023;00:1–13. ﻿ |

MATER IAL S AN D M E T H ODS

We conducted a non-­automated systematic literature review applying the

Identification of studies via databases

Records identified from

Criteria for exclusion: no

Records excluded based on title

Records excluded based on title

Records excluded based on title

Studies included in review

searched additional sources (i.e. Google Scholar, reference lists, personal

Diet of Scinax curicica tadpoles

Scinax curicica is a treefrog endemic to the Espinhaço Mountain Range

We collected 20 tadpoles in each period, 10 of them in Gosner (1960)

The search for publications on neotropical tadpole diets resulted in 28 pa-

Diet of Scinax curicica tadpoles

Scinax curicica tadpoles ate unicellular and filamentous Bacillariophyceae

Neotropical tadpole diet composition

Most of the studies on neotropical tadpole diet analysed gut contents

actually assimilated by tadpoles. Based on available studies, diatoms

The role of tadpole ecomorphotypes and microbiome

Tadpole ecomorphotypes are likely to have an important influence on

fragments (Vences et al., 2016), a diet with similar composition to those of

Temporal and ontogenetic variation in diet composition

Food selectivity and other knowledge gaps

morphological plasticity (Castañeda et al., 2006; Ruthsatz et al., 2019,

Environmental impacts and tadpole diet composition

Besides natural sources of variation, environmental impacts may also influ-

Diet of Scinax curicica tadpoles

Studies on the diet of neotropical tadpoles (Appendix S1) approached in

AC K N O​W L E​D G E​M E N T S

DATA AVA I L A B I L I T Y S TAT E M E N T

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Austral Ecology. 2023;00:1–13. |

We conducted a non-automated systematic literature review applying the

AC K N OW L ED G EM E N T S