REVISTA DE BIOLOGIA E CIÊNCIAS DA TERRA ISSN 1519-5228 Volume 23 - Número 2 - 2º Semestre 2023

1. REVISTA DE BIOLOGIA E CIÊNCIAS DA TERRA ISSN 1519-5228
Volume 23 - Número 2 - 2º Semestre 2023
ICHTHYOPHOENIX STOCK: DIVERSITY OF FISHES IN PUDDLES OF AN
INTERMITTENT RIVER IN THE BRAZILIAN SEMI-ARID
Hiltin Magalhães Soares1
; Francimário da Silva Feitosa²; Artur Henrique Freitas Florentino de Souza3
ABSTRACT
This study verified the stock of ichthyofauna in a stretch of the Piauí river, in the Brazilian semi-arid
region, through ecological attributes during the dry period. Sampling was carried out in July 2022
and data on length, width and depth were collected in each puddle and fish were captured using a
beach trawl. Were captured 238 individuals belonging to 4 orders, 6 families, 12 genera and 14
species. The order Characiformes was the most representative, with 2 families and 7 species,
followed by Perciformes with 1 family and 4 species. Were collected 22 individuals in puddle 1, 58
individuals in 2 and 158 individuals in 3. The data showed that there was a positive and significant
correlation between the depth of the puddles and the abundance of individuals present in the samples
(r = 0.95; p < 0.05), however the attributes richness and diversity did not show significant difference.
Considering the lack of studies on fish stocks and ecology in the Brazilian semi-arid region, the
results provide information for monitoring the assemblage in the dry season and its potential for
recolonization in the rainy season in intermittent rivers.
Keywords: Caatinga, Ichthyology, Parnaíba basin river, Temporary rivers.
ESTOQUE ICTIOFÊNIX: DIVERSIDADE DE PEIXES EM POÇAS DE UM RIO
INTERMITENTE NO SEMIÁRIDO BRASILEIRO
RESUMO
Este estudo verificou o estoque da ictiofauna num trecho do rio Piauí, região semiárida brasileira,
através de atributos ecológicos durante o período seco. Amostragem foi efetuada em julho de 2022 e
dados de comprimento, largura e profundidade foram coletados em cada poça e os peixes foram
capturados usando rede de arrasto de praia. Foram capturados 238 indivíduos pertencentes à 4 ordens,
6 famílias, 12 gêneros e 14 espécies.Aordem Characiformes foi a mais representativa, com 2 famílias
e 7 espécies, seguida de Perciformes com 1 família e 4 espécies. Foram coletados 22 indivíduos na
poça 1, 58 na 2 e 158 na 3. Os dados mostraram que houve uma positiva e significante correlação
entre a profundidade das poças e a abundância de indivíduos presentes na amostra. (r = 0.95; p <
0.05), contudo os atributos riqueza e diversidade não mostraram diferença significativa.
Considerando a falta de estudos sobre estoques de peixes e sua ecologia na região semiárida brasileira,
os resultados fornecem informações para monitoramento das assembleias na estação seca e seu
potencial para recolonização no período chuvoso em rios intermitentes.
Palavras-chave: Caatinga, Ictiologia, Bacia do rio Parnaíba, Rios temporários.
80

2. 1 INTRODUCTION
Brazil has about 43% of the freshwater
ichthyofauna of the Neotropical region, howe-
ver a good part of the assemblages that occur in
small streams and sub-basins are little known
from the point of view of diversity and geogra-
phic distribution (CAVALHEIRO; FIALHO,
2020). In the Northeast region, the problem is
even more serious due to the lack of knowledge,
as there are coastal basins and intermittent ri-
vers that have not been inventoried, compared
to the south and southeast of Brazil (BUCKUP;
MENEZES; GHAZZI, 2007) largely due to the
intermittent regime of the rivers, which histori-
cally led scientists to believe that aquatic life
was poorly composed of species common to the
Atlantic Forest and the Amazon. This only star-
ted to change from the 1990s onwards, when a
higher number of endemic aquatic organisms
began to be recognized (ROSA et al., 2003;
RODRIGUES-FILHO et al., 2020).
Ramos (2012) argues that this region
has a history of aggression to the environment
with countless effects on the freshwater ich-
thyofauna, which according to Rosa et al.
(2003), comprises about 240 species. Anthro-
pogenic alterations stand out, such as the crea-
tion of dams, deforestation, implantation of
plants and distilleries, diversion in the course
of rivers and introduction of species. In the
area of the present study, in the southeast of the
State of Piauí, these problems are also com-
mon, the main one being the introduction of
exotic species (GURGEL; OLIVEIRA, 1987).
Freshwater fish constitute an important
model for Biogeographical studies, as their
evolution is directly associated with the geolo-
gical evolution of the Earth (LUNDBERG,
1993) and more recently with the impacts cau-
sed by man. Given this scenario, it is necessary
to carry out investigations that aim at knowle-
dge about the ichthyofauna of temporary and
semi-permanent puddles in this region, with the
aim of knowing their role in the conservation of
diversity.
A conspicuous characteristic of this
ichthyofauna is the set of adaptations that allow
them to inhabit intermittent rivers [TERRA; TEI-
XEIRA; REZENDE, 2017; RODRIGUES-FI-
LHO et al., 2020), as these environments only
flow in the rainy season and then dry out in peri-
ods of high evapotranspiration. The rivers belon-
ging to the Caatinga ecosystem, represented in
this work by the Piauí river, have striking charac-
teristics such as a temporary regime of their wa-
tercourses, with the vast majority of them drying
up completely or fragmented into small puddles,
obviously related to the periods of flood and
drought (MALTCHIK; MEDEIROS, 2006; LE-
VIS; RAMOS; LIMA, 2013).
Fish and other aquatic organisms have
their life cycles altered when the dry period be-
gins. In the case of fish, semi-permanent puddles
or streams are responsible for harboring this di-
versity that will contribute to the recolonization
of lake systems in the next rainy season (LEVIS;
RAMOS; LIMA, 2013). However, during the dry
period, eutrophication is undoubtedly one of the
main problems faced by them, as well as the little
space for locomotion and foraging, which is ano-
ther great dilemma, increasing competition for
food resources (MATTHEWS, 1998). Even so,
Matthews (1998) also considers that temporary
puddles serve as important spawning sites for
some species, however it should be noted that
most neotropical freshwater fish reproduce in the
rainy season (AGOSTINHO at al., 2004) and the-
refore the puddles play a key role in the recoloni-
zation of other areas of the water system based
on their stocks.
The aim of this study was to verify the
main ecological attributes of fish assemblages in
puddles in a stretch of the Piauí River, in order to
assess the potential for recolonization in the sub-
sequent rainy season. For this, we considered (i)
inventorying the ichthyofauna in a stretch of the
Piauí River, (ii) recording the total richness,
abundance and diversity and per puddle, and (iii)
relating the structure of the puddles with the eco-
logical attributes. The specimens collected in this
study were deposited at the Laboratory of Bioar-
chaeology and Biodiversity at UNIVASF Cam-
pus Serra da Capivara for future conferences.
2 MATERIALAND METHODS
2.1. Study area
The study area comprises a stretch of the
Piauí River located in the village 'Lagoa dos
Torrões' in the municipality of São Raimundo
Nonato, State of Piauí (Figure 1), at an altitude of

3. approximately 330 meters. The climatic condi-
tions of this location present minimum tempera-
tures of 18° C and maximum of 36° C, with a
semi-arid, hot and dry climate (AGUIAR; CAR-
VALHO, 2004).
Average annual rainfall is defined by the
Continental Equatorial regime, with annual iso-
hyets around 600 mm and rainfall concentration
between December and March (AGUIAR; CAR-
VALHO, 2004).
Figure 1. Study area localization. Source: Prepared by the
authors.
Collections took place in July 2022 at
three sampling points with approximate coordi-
nate data: Puddle 1 (9°04’20”S 42°45’23”W),
Puddle 2 (9°04’11”S 42°45’49”W) and Puddle 3
(9°04’09”S 42°45’56”W).
The depth, width, and length of each pud-
dle were obtained by probing several measure-
ments along a transect using a graduated rod and
measuring tape. Data were recorded in the field
and then tabulated in Excel to calculate the ap-
proximate volume of each well. With this infor-
mation we related the structure of the puddles
with the ecological attributes. In this case, it is
expected that in more structured or deeper pud-
dles there will be more biotopes for refuge, fa-
voring the establishment of more species, conse-
quently increasing the diversity in the place.
The equipment used to capture the speci-
mens was a beach seine net due to the efficiency
of this instrument (MEDEIROS et al., 2010). Af-
ter collection, the fish were fixed in the field with
formalin (10%) after Eugenol anesthesia with a
concentration of 125 mg/L and placed in plastic
bags duly identified as to the sampling point.
They were then transported to the Bioarchaeol-
ogy and Biodiversity laboratory at UNIVASF
Campus Serra da Capivara, where they were pre-
served in 70% alcohol for identification.
2.2. Species identification
Captured fish were identified in the labor-
atory using literature from the area with identifi-
cation keys (BRITSKI; SILIMON; LOPES, 1972;
NELSON, 2006; RAMOS, 2012). They were
also classified according to their origin, being (i)
native (from the semi-arid watershed), (ii) alloch-
thonous (introduced from other watersheds in
Brazil) or (iii) exotic (originating from water-
sheds in other countries) according to Britiski;
Silimon; Lopes (1972), Nelson (1984) and Na-
katani et al. (2001).
After identification, the fish were trans-
ferred to glass vials compatible with their sizes
and submerged in a 70% alcohol solution for con-
servation. The flasks were duly identified with la-
bels with information regarding the species,
place, date and collectors.
2.3. Indexes of the assemblages and data analysis
For the description of the ichthyological
assemblage, in this study different attributes of
the community were analyzed, such as the total
number of individuals in the sample (N), species
richness (total number of species S) and species
diversity (Shannon SH diversity index) present in
the sample to verify the distribution of the ich-
thyofauna in the three puddles of the studied river
stretch. The indices were obtained from Gerhard
(2005), for the calculation of the diversity indices,
the data were tabulated in Excel and then calcu-
lated in the PAST software.
The choice of the Shannon-Weaver diver-
sity index is due to the fact that the test considers
equal weight between rare and abundant species
(MAGURRAM, 1988). Thus, the greater the va-
lue of H', the greater the diversity of the sample,
which may express richness and uniformity. The
Shannon-Weaver diversity index was obtained
through the formula: H'= -∑pi x logpi, where pi
= ni/N (pi = the relative abundance of each spe-
cies, calculated by the proportion of individuals
of a species by the total number of individuals in
the community; ni = the number of individuals in
each species; the abundance of each species; N =
the total number of all individuals; log. = nepe-
rian basis) (MUELLER-DUMBOIS; ELLEN-
BERG, 1974).

4. 3 RESULTS
The puddles had very different volumes
(Table 1), where the first puddle had almost half
the volume of the third, reflecting the abundance
and richness of species.
Table 1 - Abundance (N), richness (S), diversity index
(H’), depth (D) and volume (V) of Piauí river puddles.
Source: Research data.
In the stretch studied, 238 individuals
were collected, distributed in 4 orders, 6 families,
12 genera and 14 species (Table 2). Analysis of
the composition of the ichthyofauna showed that
the most abundant orders were Characiformes
(50%) and Perciformes (29%), with the families
Characidae (42%) and Cichlidae (29%) having
the highest number of species. However, the
most abundant species was Cyprinodontiformes
Poecilia vivipara (Bloch & Schneider, 1801)
(29%), followed by Characiformes Astyanax aff.
bimaculatus (Linnaeus 1758) (21%).
Six species were classified as native, six
as allochthonous and two as exotic (Table 2). 22
individuals were collected in Puddle 1, distrib-
uted in 6 species and H’ = 1,538. In Puddle 2, 58
individuals were collected, belonging to 8 spe-
cies and H’ = 1,841. In Puddle 3, 158 individuals
belonging to 9 species and H’ = 1,757 were col-
lected.
Table 2 – List of species present and your respectives
abundance in the puddles 1 (P1), 2 (P2) and 3 (P3). The
species were classified as natives (NAT), allochthonous
(ALL) and exotic (EXO).
Source: Research data.
The t test indicated that there was no sig-
nificant difference between the mean diversity
values between puddles (p = 0.13).
The collected data demonstrated that
there was a positive and significant correlation
between the depth of the puddles and the abun-
dance of individuals present in the samples (r =
0.95; p < 0.05) (Figure 2), corroborating the ini-
tial premise that a greater abundance would occur
in more structured places, in this case in greater
depth.
The correlation assessment was also pos-
itive when analyzing the wealth variable (r =
0.85), but the value was not significant at the 5%
level (p = 0.34) (Figure 2). The correlation anal-
ysis for the diversity variable (SH) was negative,
with no significant difference (r = 0.41; p = 0.7)
(Figure 2).

5. Figure 2. Linear correlation between the independent
(depth) and dependent variables (abundance, richness and
diversity). Source: Research data.
4 DISCUSSION
Lowe-McConnell (1999) argues that the
ichthyofauna of South America is characterized
by the dominance of Characiformes over Siluri-
formes, which was confirmed in the present study.
The most abundant species (P. vivipara and A.
bimaculatus) are small and can be classified as
opportunistic because they have a high reproduc-
tive potential, trophic plasticity and, mainly, wide
tolerance to environmental disturbances
(LOWE-MCCONNELL, 1999) which can favor
events of recolonization in the environment.
The influence of puddle depth on the eco-
logical attributes of fish assemblages (abundance,
richness, diversity) has been associated with the
volume and complexity of the habitat
(GORMAM; KARR, 1978), a similar scenario
observed in the present research, since the abun-
dance data found were positively related to depth,
with a more evident relationship in Puddle 3.
Likewise, this relationship is probably enhanced
by the greater number of refuges, contributing to
the fish being able to explore the habitat in dif-
ferent and better ways, according to their charac-
teristics biological and morphological
(WINEMILLER, 1991). Thus, fish species can
explore the environment just below the surface,
in the middle of the water column, as well as in
the sediment (LOWE-MCCONNELL, 1999;
CASATTI; CASTRO, 2006).
Súarez; Petrere-Júnior; Catella (2004)
showed that the variation in richness is positively
related to the depth of the lakes studied, which
was not found in the present study, probably due
to the smaller size of the puddles. This result was
not expected in the present study, since in deeper
puddles this characteristic would provide more
microhabitats and food resources, contributing to
increased diversity (ARAÚJO; TEREJINA-
GARRO, 2007; SÚAREZ, 2008).
The present study confirms the im-
portance of the semi-permanent puddles of the
Piauí River for the conservation of fish biodiver-
sity, as it provides refuge for forage species pre-
sent there, ensuring their survival until the arrival
of the next rainy season. Due to the high repro-
ductive capacity of many representatives of this
assemblage in terms of abundance, there must
probably be a transition between the minimum
biotic potential, in the dry period, and the maxi-
mum in the rainy season, being limited by the
great water deficit in most of the year. This bio-
logical characteristic justifies the “resurgence” of
a new assemblage in post-rain random events that
needs to be tested. However, the evidence sug-
gests that the bank of species present in the pud-
dles in the dry period can contribute to the
maintenance of other populations in that portion
of the drainage basin throughout the entire cycle.
The research confirmed a positive rela-
tionship between abundance and structure of the
puddles, since the greatest abundance occurred in
places with greater volume and depth. This was
obviously expected, but it did not occur with the
calculated diversity index. Although the 'traíra'
Hoplias malabaricus was not collected in the
present study, the most abundant species (Poe-
cilia vivipara and A. bimaculatus) are its main

6. food items and therefore such forage species are
considered to be mainly responsible for recoloni-
zation events also for serving of food; in other
parts of the basin, forage species are also food
items for the 'piranhas' Pygocentrus and Ser-
rasalmus, which are only found in large reser-
voirs in the region.
5 CONCLUSION
We conclude that in the area of this study,
forage species support the trophic web of the fish
assemblage, which in turn is greatly impacted by
the water regime as well as by the use of soil and
natural resources by humans in the drainage ba-
sin.
It is projected that in extreme droughts,
where the puddles would dry up completely, this
stock can only be replenished only from large lo-
cal reservoirs. This would have a negative impact
on the genetic heritage of this ichthyophoenix
stock, since historically the basin has been highly
fragmented and, therefore, without gene flow.
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______________________________________
1- Colegiado de Ciências da Natureza,
Universidade Federal do Vale do São Francisco
- UNIVASF, Campus Serra da Capivara, Rua
João Ferreira dos Santos s/n, Bairro Campestre,
CEP 64770-000, São Raimundo Nonato, PI,
Brasil; e-mail:
hiltin.soares@discente.univasf.edu.br
2- Colegiado de Ciências da Natureza,
Universidade Federal do Vale do São Francisco -
UNIVASF, Campus Serra da Capivara, Rua João
Ferreira dos Santos s/n, Bairro Campestre, CEP
64770-000, São Raimundo Nonato, PI, Brasil; e-
mail: francimario.feitosa@univasf.edu.br
ORCID: https://orcid.org/0000-0002-8463-3707

8. 3. Prof. do Departamento de Sistemática e
Ecologia, Universidade Federal da Paraíba -
UFPB, Cidade Universitária I, 900, CEP 58059-
900, João Pessoa, PB, Brasil; e-mail:
ahffs@ccen.ufpb.br ORCID:
https://orcid.org/0000-0002-9069-7950
87