Elmoor-Loureiro et al 2022

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Limnologica xxx (xxxx) xxx
Please cite this article as: L.M.A. Elmoor-Loureiro, Limnologica, https://doi.org/10.1016/j.limno.2022.126008
Available online 4 August 2022
0075-9511/© 2022 Elsevier GmbH. All rights reserved.
Towards a synthesis of the biodiversity of freshwater Protozoa, Rotifera, 
Cladocera, and Copepoda in Brazil 
L.M.A. Elmoor-Loureiro a,b,1, F.D.R. Sousa b,c,*,1, F.R. Oliveira d,1, C.Y. Joko e,1, 
G. Perbiche-Neves f,1, A.C.S. da Silva g, A.J. Silva h, A.R. Ghidini i, B.R. Meira d, C.E.G. Aggio j, 
C.S. Morais-Junior h, E.M. Eskinazi-Sant’Anna k, F.M. Lansac-Tôha d, G.S. Cabral l, 
J.L. Portinho m, J.R. Nascimento n, J.V.F. Silva g, L. Veado a, L.J. Chiarelli o, L.O. Santana d, 
L.P. Diniz k, L.S.M. Braghin d, L.T.F. Schwind p, M. Melo Júnior q, M. Progênio d, M.A. Rocha r, 
M.L.C. Silva a, M.S.M. Castilho-Noll o, N.J. Silva d, N.G. dos Santos o, P.H.R. Morari o, 
P.M. Maia-Barbosa s, P.M. Oliveira t, R.L. Arrieira u, R.L. Macêdo h, S. Deosti d, T. Mantovano j, 
V. Gazulha a, C.C. Bonecker d,g,2, F.A. Lansac-Tôha d,2, P.H.C. Corgosinho t,2, L.F.M. Velho d,2, 
N.R. Simões n,2 
a Independent researcher, Brazil 
b Laboratory of Animal Taxonomy, Federal University of Jataí, Goiás, Brazil 
c PPZoo/University of Brasília, Distrito Federal, Brazil 
d Nupelia/PEA/CCB/Maringá State University, Paraná, Brazil 
e Faculty of Biological Sciences/Distrito Federal University Center, Distrito Federal, Brazil 
f Department of Hydrobiology/CCBS/São Carlos Federal University, São Paulo, Brazil 
g Nupelia/PGB/CCB/Maringá State University, Paraná, Brazil 
h Department of Biology and Evolutionary Ecology/CCBS/São Carlos Federal University, São Paulo, Brazil 
i CCBN/Acre Federal University, Acre, Brazil 
j Norte do Paraná State University, Paraná, Brazil 
k Department of Biodiversity, Evolution and Environment/Ouro Preto Federal University, Minas Gerais, Brazil 
l Federal University of Pará, Brazil 
m Department of Biodiversity/Paulista State University, São Paulo, Brazil 
n CAF/Sul da Bahia Federal University, Bahia, Brazil 
o Department of Biological Sciences/IBILCE/Paulista State University, São Paulo, Brazil 
p DCI/Universidade Estadual de Maringá, Paraná, Brazil 
q Department of Biology/Pernambuco Rural Federal University, Pernambuco, Brazil 
r Institute of Biology/Bahia Federal University, Bahia, Brazil 
s Federal University of Minas Gerais, Minas Gerais, Brazil 
t PPGBURN/State University of Montes Claros, Minas Gerais, Brazil 
u Paranaense University, Paraná, Brazil 
A R T I C L E I N F O 
Keywords: 
South America 
Neotropical 
Aquatic biology 
Taxonomy 
Ecology 
A B S T R A C T 
Although Brazil is considered a megadiverse country, its rich freshwater biodiversity is still poorly known. A 
general overview of to-date knowledge on Protozoa, Rotifera, Cladocera, and Copepoda species and distribution 
in Brazilian Hydrographic Regions is presented here, based on literature data since the 1890s. Ecological studies 
provided most of the occurrence records. The results show high richness for all studied biological groups and 
unequal distribution of the occurrence records, which are substantially influenced by research groups’ location. 
* Correspondence to: Laboratório de Taxonomia Animal, UAE Ciências Biológicas, Universidade Federal de Jatai—UFJ, BR 364 km 195 no. 3800, CEP 75801-615 
Jataí, GO, Brazil. 
E-mail address: fdiogo.rs@gmail.com (F.D.R. Sousa). 
1 L.M.A Elmoor-Loureiro, F.D.R. Sousa, F.R. Oliveira, C.Y. Joko, G. Perbiche-Neves are joint first author 
2 C.C. Bonecker, F.A. Lansac-Tôha, P.H.C. Corgosinho, L.F.M. Velho, N.R. Simões are joint senior author 
Contents lists available at ScienceDirect 
Limnologica 
journal homepage: www.elsevier.com/locate/limno 
https://doi.org/10.1016/j.limno.2022.126008 
Received 28 April 2022; Received in revised form 4 July 2022; Accepted 12 July 2022 
mailto:fdiogo.rs@gmail.com
www.sciencedirect.com/science/journal/00759511
https://www.elsevier.com/locate/limno
https://doi.org/10.1016/j.limno.2022.126008
https://doi.org/10.1016/j.limno.2022.126008
https://doi.org/10.1016/j.limno.2022.126008
Limnologica xxx (xxxx) xxx
2
Center of studies 
Trends 
The data also revealed that Brazilian zooplankton biodiversity still needs to be better studied, taxonomically, 
although from the beginning of the last century until 1980 these kinds of studies were predominant. 
1. Introduction 
Brazil is considered a megadiverse country (Myers et al., 2000), but 
its rich freshwater biodiversity is still poorly known. The knowledge 
about zooplankton biodiversity in Brazil has been constructed since the 
19th century, when the first reports were published on Cladocera 
(Richard, 1897), Copepoda (Poppe, 1889 in De Guerne and Richard, 
1889), Rotifera (Zelinka, 1891), and Protozoa (Daday, 1905). The rapid 
development of Brazilian Limnology after the 1970s (Esteves, 2011) 
brought a consequent increment in the occurrence records of 
zooplankton organisms. That, in turn, led to attempts in cataloging this 
biodiversity (e.g., Velho et al., 1996; Velho and Lansac-Tôha, 1996; 
Elmoor-Loureiro, 1998; Rocha and Coelho-Botelho, 1998; Santos-Silva, 
1998; Garraffoni and Lourenço, 2012), also evidencing the gaps in 
existing knowledge. 
The incompleteness of our comprehension of zooplankton biodiver-
sity is more worrying when contrasted with its importance in the func-
tioning of freshwater ecosystems (Litchman et al., 2013) and with 
Anthropocene threats that reduce biodiversity, such as biological in-
vasions, pollution, overexploitation of natural resources, land-use 
change, and climate change (Dudgeon, 2019). Thus, it is not prema-
ture to suggest that freshwaters are the most endangered ecosystems in 
the world, despite providing valuable goods and services for human 
societies (Dudgeon et al., 2006). 
The knowledge on Brazilian zooplankton biodiversity is limited in 
diverse aspects – species taxonomy, distribution, abundance, evolu-
tionary patterns, abiotic tolerance, species traits, and biotic interactions. 
Among these seven shortfalls, the most critical are the Wallacean and 
Linnean: the former has a significant impact on the sampling bias, dis-
tribution, and geographical pattern of biodiversity (Hughes et al., 2021). 
Regarding Linnean shortfall, the lack of information on species taxon-
omy prevents the development of knowledge of all other aspects (Hortal 
et al., 2015). 
Therefore, there is an urgent need to improve the taxonomic infor-
mation on Brazilian zooplankton organisms. It is true that there has been 
some recent activity in this area (e.g., Garraffoni and Lourenço, 2012; 
Narciso et al., 2020; Sousa et al., 2021; Meira et al., 2021), but efforts 
should be more planned and coordinated. In this sense, the creation of 
the Neotropical Zooplankton Network is welcome. The present paper is 
one of the Brazilian group’s first contributions, and it aims to present a 
general overview on to-date knowledge on Protozoa, Rotifera, Clado-
cera, and Copepoda species and distribution in Brazil. It is expected that 
this synthesis serves as a basis and drives future investigations. 
2. Material and methods 
The database was composed of scientific papers on zooplankton in 
Brazil, published from January 1900 to August 2021, and indexed in 
Scielo (https://www.scielo.org/), Clarivate (www.isiwebofknowledge. 
com), and SciVerse Scopus (www.scopus.com). To search for studies 
from each analyzed group, a combination of keywords was used. For 
cladocerans, it was zooplankton OR cladocer* OR ctenopoda OR 
anomopoda AND Brazil; for rotifers, it was zooplankton OR Rotifer* OR 
rotifer* AND Brazil; for Protists, it was zooplankton OR protist OR 
Protozoan* OR testate amoebae OR flagellate OR Ciliate* OR Amoebae 
AND Brazil; and, for copepods, it was zooplankton OR copepod* OR 
Copepoda AND Brazil. Additionally, the database contains records found 
in classical papers published prior to 1900 and in book chapters.The list 
of considered publications is presented in Supplementary File 1. 
Each analyzed paper provided the geographical distribution records 
of cited taxa, types of ecosystems sampled (lotic, lentic natural, lentic 
artificial, and miscellaneous), and the Hydrographic Regions associated 
with each one (Fig. 1). These papers were also separated into four cat-
egories related to their main subject: Ecology (population or commu-
nity), Functional Diversity, Molecular Diversity, and Taxonomy. Then, 
these data were analyzed from a temporal point of view. 
The richness of Protozoa, Rotifera, Cladocera, and Copepoda was 
quantified for each type of ecosystem and Hydrographic Region. Next, a 
heatmap was built using a raster file dividing the map of Brazil into a 
grid with 1◦ × 1◦ cell, c.a. 110 km. For each cell in that grid, we summed 
occurrence records totaling the species richness. However, some cells 
did not presented records in the literature. Thus, to estimate the species 
richness in the empty cells and maintain the suavized map, we applied 
the geostatistical interpolation technique of ordinary krigingbased on 
inverse distance power defined in Gräler et al. (2016). This approach has 
been very used in ecology and biogeography (Alves et al., 2020) because 
interpolates between sampled quadrats exclusively according to the 
spatial dependence of the response variable and ignores underlying 
environmental gradients but has the advantage of an exact interpolation 
method at sampled locations (Kreft and Jetz, 2007). Finally, an accu-
mulation curve for each zooplankton group was built, using the rare-
faction method based on the incidence data for Hydrographic Regions 
(Gotelli and Colwell, 2001). 
3. Results and discussion 
3.1. Protozoa 
One hundred and thirty-nine published studies on zooplankton 
protists were conducted in Brazil, with testate amoebae and ciliates 
being the best documented groups. Taxonomic and ecological studies 
involving communities of heterotrophic flagellates and naked amoebae 
are still extremely scarce, probably due to their relatively small size and/ 
or absence of notable morphological characters (Regali-Seleghim et al., 
2011), arousing little interest and greatly limiting the training of tax-
onomists in the study of these protists. 
In the inspected publications, we have recorded 181 families, 320 
genera and 804 species of protists, of which the ciliates present the most 
species, with 435 species, followed by testate amoebae, with 240 spe-
cies. Flagellates and naked amoebae had 113 and 16 species, respec-
tively (Supplementary File 3). 
The most diverse families were Difflugiidae (101), Arcellidae (27), 
Hyalospheniidae (27) and Centropyxidae (22) among testate amoebae; 
Oxytrichidae (28) and Vorticellidae (18) for the ciliates; Entosiphonidae 
(23) and Astasiidae (22) for the flagellates; and Amoebidae (7) among 
the naked amoebae. 
Considering the Brazilian hydrographic regions, the highest values of 
protozooplankton species richness were recorded for the Upper Paraná 
River region (520 species), followed by the Southeast Atlantic (377 
species, Tocantins-Araguaia (174 species) and the Amazon (159 species) 
(Table 1). These considerable geographic differences in the diversity of 
protozooplankton seem to be determined by the sampling effort and are 
less related to biogeographic patterns. Thus, high species diversity has 
been recorded in regions with a greater number of studies. This fact is 
especially evident in the Upper Paraná River Floodplain, where long- 
term ecological studies have been conducted for over 30 years 
(Fig. 2). On the other hand, the lack of studies in the Uruguay River basin 
means that the diversity of these organisms in this region is totally 
unknown. 
Considering the types of aquatic environments, the natural lentic had 
the highest number of species (646 species), followed by lotic (387 
species), artificial lentic (334 species) and miscellaneous (271 species). 
L.M.A. Elmoor-Loureiro et al. 
Limnologica xxx (xxxx) xxx
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Lentic environments are those with greater hydrodynamic stability 
(lower water flow); that is, they are the preferred habitat of typically 
planktonic species. Natural lentic environments usually have a high 
number of submerged or free-floating macrophytes and consequently 
support greater species richness, considering that they provide food 
availability and predator refuge (Buosi et al., 2011). The greater di-
versity of species recorded in natural lentic environments, compared to 
artificial lentics, may be associated with greater environmental hetero-
geneity, observed for natural lentics, which results in a greater exchange 
of species between the compartments of the environments. Similarly, the 
higher diversity observed for lotic environments, when compared to 
artificial lentic ones, is probably due to the contribution of organisms 
from other compartments. 
Another aspect that must be considered to explain the differences in 
diversity is the number of studies carried out in each type of environ-
ment. For protozooplankton, about 36% of the papers were carried out 
in natural lentic, while lotic and artificial lentic environments corre-
spond to 25%, each. 
Ecological studies have markedly predominated in research carried 
out with zooplankton protists, except at the beginning of the last cen-
tury, when taxonomic studies predominated (Fig. 2). Exponential 
growth has been observed since then in ecological studies, especially 
from the 1990s onwards. These studies initially involved aspects related 
to the characterization of protozooplankton community attributes 
(Gomes and Godinho, 2003), progressively including new approaches 
(e.g. Paiva and Silva-Neto, 2004; Segovia et al., 2016;) until testing 
complex ecological theories (e.g. Lansac-Tôha et al., 2021). 
Taxonomic studies on protozooplankton began in the early 20th-cen-
tury (Prowazek, 1910; Cunha, 1913, 1916). After a gap of a few decades, 
studies on the taxonomy and occurrence of testate amoebae species were 
carried out in different aquatic environments in the Upper Paraná River 
floodplain (Velho and Lansac-Tôha, 1996; Velho et al., 1996). Since the 
beginning of the 21st century, there has been an increase in taxonomic 
and review studies (Lansac-Tôha et al., 2000; Alves et al., 2007; 
Regali-Seleghim et al., 2011; Miranda et al., 2020). 
Few endemic protozooplankton species have been recorded in Brazil. 
Currently, there is a debate about the geographic distribution of these 
microorganisms, whether all species have a cosmopolitan distribution, 
and are selected by environmental conditions (Finlay, 2002; Fenchel and 
Finlay, 2004) or if at least a considerable number of the species have 
certain degree of endemism (“moderate endemicity model”, Foissner 
et al., 2007). Furthermore, it should be highlighted that the lack of 
taxonomists, and even of studies in different parts of the country, 
strongly limits knowledge about the real diversity and distribution of 
protozooplankton in our country. 
Despite that, some records of species are restricted to Brazil, such as, 
among the testate amoebae, Suiadifflugia multipora Green, 1975, which 
was described in the Suiá Missu river, in the Amazon basin. Currently, 
this species occurs beyond the Amazon region, in the Paraguay, Paraná, 
Tocantins-Araguaia and São Francisco river basins (Table S3.1 - Sup-
plementary File 3). Furthermore, Arcella brasiliensis Cunha (1913) has 
been recorded in plankton samples from the North to South of the 
country (Lansac-Tôha et al., 2000; Velho and Lansac-Tôha, 1996; Rocha 
et al., 2021). 
Among the ciliates, there was the description of Oxytricha marcili 
Paiva and Silva-Neto, 2004, which was reframed into a new genusby 
Shao et al. (2011), Urosomoida marcili. The species was recorded in 
Cabiúnas lagoon, in the Southeast Atlantic region (Paiva and Silva-Neto, 
2004). 
3.2. Rotifera 
A total of 220 studies were analyzed, and 13,649 occurrences of 630 
species were recorded in different freshwater aquatic environments 
(lotic, lakes, natural and artificial). Of the 43 species considered by 
Segers (2007) as endemic to the Neotropical region, 30 have so far only 
been recorded in Brazil, mainly Lecanidae (8 species) and Brachionidae 
(7 species) (Table S3.2 - Supplementary File 3). 
Among the 30 families registered in the studies, Lecanidae was the 
most representative (115 species), followed by Notommatidae (83 spe-
cies), Brachionidae (62 species), Trichocercidae (59 species) and Lep-
adellidae (58 species) (Table S3.2, Supplementary File 3). The results 
were similar, as shown in the last checklist of Rotifera in Brazil (625 
species), although the present study considered only valid species and 
Fig. 1. Brazilian Hydrographic Regions according to Resolution 32 of Conselho Nacional de Recursos Hídricos. 
L.M.A. Elmoor-Loureiro et al. 
Limnologica xxx (xxxx) xxx
4
sub-species and taxonomic forms or varieties were not included, as they 
were in the checklist. Notommatidae, Brachionidae, Trichocercidae and 
Lepadellidae were also important in the last checklist, but in the present 
study Notommatidae was represented by more species (62 taxa in Gar-
raffoni and Lourenço, and 82 species in the present study). The differ-
ence could have arisen from the existence of more studies in their 
identification in the last few years, due to their representativeness in 
rotifer communities. The majority of the studies focused on ecological 
data (181 studies), followed by taxonomic data (22 studies) and func-
tional diversity data (16 studies). 
The rotifer species occurred mainly in more than one type of envi-
ronment in the same study analyzed (496 species), followed by lakes 
(449 species), rivers and streams (361 species) and artificial environ-
ments (reservoirs and aquaculture ponds) (265 species). Most of the 
occurrence of the species in lentic environments was related to the 
importance of the reduced flow for the establishment and development 
of populations. Most species are non-planktonic (Lecanidae and 
Notommatidae); however, they are commonly found in plankton of the 
Neotropical region due to the lesser/reduced depth of most lakes and the 
occupation of banks of aquatic macrophytes, where this species may be 
associated. 
Kellicottia bostoniensis (Rousselet, 1908) is an invasive species that 
comes from the Nearctic region and showed 101 occurrences in the 
different aquatic environments, mainly in artificial lentic environments 
(53 occurrences), such as reservoirs. Artificial environments facilitate 
biological invasions (Havel et al., 2005) by altering the environmental 
conditions of natural ecosystems (rivers) and changes in the availability 
of food resources due to anthropogenic modifications to ecosystems. 
This species showed a wide distribution in the country, occurring in 10 
hydrographic regions, from the North to the South, and mainly in Paraná 
(70 occurrences). In contrast, Kellicottia longispina (Kellicott, 1879), also 
from the Nearctic region, showed a low occurrence (five records) in the 
environments, mainly in the natural lakes, including the Amazonas 
Lowlands. 
Still considering the hydrographic regions, the rotifer species 
occurred mainly in Paraná (484 occurrences) (Table 1). This greater 
occurrence could be related to the higher number of taxonomists 
studying the freshwater environments in this region (taxonomic and 
ecological studies) and consolidated research groups in the zooplankton 
community. Brachionus dolabratus Harring, 1914, Brachinous falcatus 
Zacharias, 1898, Brachionus mirus Daday (1905), Keratella cochlearis 
(Gosse, 1851) and Testudinella patina (Hermann, 1783) were the only 
species that occurred in all hydrographic regions. 
Considering the rotifer studies in Brazil, the biodiversity data 
increased, as shown by the higher number of registers observed by 
Garraffoni and Lourenço (2012) and the present study. Moreover, some 
regional reviews were also carried out after Garraffoni and Lourenço 
(2012), but some hydrographic regions should be explored more, such as 
the as Northeast Atlantic Oriental Region (20 species) and Parnaíba (29 
species). To increase this knowledge, it is also necessary to encourage 
taxonomic studies with these rotifers as the number of taxonomists is 
very small. There are groups of researchers that study these organisms 
throughout the country, but most of these researchers carry out 
ecological studies and use taxonomy only as a tool to support their 
studies. 
3.3. Cladocera 
The distribution of overall cladoceran taxa richness throughout the 
Brazilian Hydrographic region is summarized in Fig. 2. The highest taxa 
richness was recorded in the Paraná Hydrographic Region (131 taxa), 
followed by the Amazon (107 taxa), Tocantins-Araguaia (104 taxa) and 
Paraguay (100 taxa). As we expected, taxon richness is locally concen-
trated in regions with higher sampling efforts (Table 1), considering the 
presence of more consolidated research groups dedicated to limnology 
and community ecology of zooplankton, including Cladocera. These 
findings follow the same pattern throughout the world (Forró et al., 
2008). Specifically, the heat map revealed high richness in the Paraná, 
part of the São Francisco, and the Paraguay Hydrographic Regions 
(Fig. 2). Tocantins/Araguaia also presented high richness due to recent 
efforts to integrate sampling floodplains in Brazil (e.g., Gomes et al., 
2020). However, it is important to highlight that this pattern was not 
observed for the Amazon basin, which presented a low sampling effort 
compared to other regions, but with the second highest species richness 
among the studied basins (Table 1). 
From 471 studies a total of 169 taxa were reported (Table S3.3 - 
Supplementary File 3), of which 50 should be considered a priority in 
future taxonomic studies because reports remain doubtful or represent 
Table 1 
Number of studies, occurrence records and taxa of Protozoa, Rotifera, Cladocera and Copepoda in Brazilian Hydrographic Regions. AM = Amazon; EA = East Atlantic; 
NAOC = Northeast Atlantic Occidental; NAOR = Northeast Atlantic Oriental; SEA = Southeast Atlantic; SA = South Atlantic; PA = Parnaíba; PG = Paraguay; PR 
= Paraná; SF = São Francisco; TA = Tocantins-Araguaia; UR = Uruguay. 
AM EA NAOC NAOR SEA SA PA PG PR SF TA UR Total 
Protozoa 
Studies 
(%) 
12 
(7.45) 
2 
(1.24) 
2 
(1.24) 
3 
(1.86) 
20 
(12.42) 
6 
(3.73) 
3 
(1.86) 
10 
(6.21) 
78 
(48.45) 
12 
(7.45) 
13 
(8.07) 
0 
(0.00) 
161 
Records 
(%) 
9 
(5.17) 
2 
(1.15) 
2 
(1.15) 
3 
(1.72) 
32 
(18.39) 
5 
(2.87) 
3 
(1.72) 
6 
(3.45) 
89 
(51.15) 
14 
(8.05) 
9 
(5.17) 
0 
(0.00) 
174 
Richness 159 32 21 47 377 99 125 54 520 133 174 0 804 
Rotifera 
Studies 
(%) 
34 
(16.11) 
9 
(4.26)) 
1 
(0.47) 
32 
(15.17) 
19 (9.00) 4 
(1.9) 
2 
(0.95) 
7 
(3.32) 
82 
(38.86) 
9 
(4.26) 
12 
(5.69) 
2 
(0.95) 
211 
Records 
(%) 
116 
(20.35) 
36 
(6.32) 
1 
(0.18) 
50 
(8.77) 
74 
(12.98) 
5 
(0.88) 
6 
(1.05) 
13 
(2.28) 
232 
(40.7) 
15 
(2.63) 
22 
(3.86) 
8 
(1.40) 
570 
Richness 397 149 184 20 191 72 29 200 484 183 134 49 630 
Cladocera 
Studies 
(%) 
98 
(18.01) 
21 
(3.86) 
20 
(3.67) 
67 
(12.31) 
72 
(13.23) 
39 
(7.16) 
6 (1.10) 40 
(7.35) 
231 
(42.46) 
57 
(10.47) 
33 
(6.06) 
5 
(0.91) 
474 
Records (%) 153 
(10.75) 
82 
(5.76) 
23 
(1.61) 
120 
(8.43) 
171 
(12.01) 
47 
(3.30) 
23 
(1.61)51 
(3.58) 
550 
(38.65) 
139 
(9.76) 
58 
(4.07) 
6 
(0.42) 
1423 
Richness 107 79 41 85 95 89 38 100 132 78 104 27 169 
Copepoda 
Studies 
(%) 
46 
(12.81) 
10 
(2.78) 
1 
(0.27) 
29 
(8.07) 
35 
(9.74) 
16 
(4.45) 
2 
(0.55) 
14 
(3.89) 
156 
(43.45) 
29 
(8.07) 
14 
(3.89) 
7 
(1.94) 
359 
Records (%) 332 
(13.02) 
125 
(4.09) 
1 
(0.04) 
124 
(4.86) 
162 
(6.36) 
79 
(3.10) 
21 
(0.82) 
220 
(8.63) 
1262 
(49.51) 
158 
(6.20) 
55 
(2.16) 
10 
(0.39) 
2549 
Richness 88 36 1 53 52 44 7 50 81 36 37 5 194 
L.M.A. Elmoor-Loureiro et al. 
Limnologica xxx (xxxx) xxx
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poorly described taxa. The identified taxa are distributed in 52 genera, 
subordinate to nine families. Most of the taxa reported for Brazil do not 
inhabit plankton, but live associated with several kinds of substrates. 
The Chydoridae was the family with most species in both lotic and lentic 
ecosystems (90 taxa), followed by Daphniidae (24), Macrothricidae 
(15), Sididae (13), Moinidae and Bosminidae (8), Ilyocryptidae (7), and 
Eurycercidae (3). Holopedidae was represented only by Amazonian 
endemic Holopedium amazonicum Stingelin, 1904 (Table S3.3 - Supple-
mentary File 3). About 17% of the species, so far, have been reported 
only in Brazil, such as Moina rostrata McNair 1980, Macrothrix sioli 
Smirnov 1982, and Kisakiellus aweti Sousa & Elmoor-Loureiro, 2018 
(Table S3.3 - Supplementary File 3). Some species might represent truly 
endemic species, but others is likely to occur outside Brazilian territory. 
Brazil should be considered a megadiverse country in terms of 
cladoceran fauna due to harboring approximately 70% of valid species 
observed in the Neotropics. In contrast, there are relatively continuous 
species concentrations in a region extending from Paraná to the State of 
São Paulo. However, data published from other parts in Brazil have also 
helped to improve the knowledge about local endemism (e.g., Kotov and 
Elmoor-Loureiro, 2008; Sousa et al., 2016), the presence of non-native 
species (e.g., Zanata et al., 2003; Elmoor-Loureiro et al., 2010) 
(Tables 1–2), and kinds of ecosystems sampled and their dynamics. 
Herein, we found that richness was quite similar when comparing 
different types of water bodies, with highlights to lentic natural eco-
systems (Fig. 2). 
Over time, ecological studies have been the main focus of the Cla-
docera research in Brazil (Fig. 2), and they are the most important 
source of richness and distribution data shown in our analysis. The first 
data on the biodiversity of Cladocera in Brazil dates from the late 19th to 
the early 20th century (e.g., Richard, 1897; Sars, 1901; Daday, 1905). 
For about a century after that, new information was sporadic, with the 
exception of the work of Francisco Bergamin during the 1930s. The final 
years of the 1990s brought the first synthesis on Brazilian Cladocera 
diversity (Elmoor-Loureiro, 1998). 
The taxonomic work on this group began to increase in the first years 
of the 21st century (Fig. 2), when studies by non-Brazilian researchers 
initially predominated (e.g., Van Damme et al., 2005), followed by 
intense activity by Brazilian taxonomists (e.g., Elmoor-Loureiro, 2014; 
Sousa and Elmoor-Loureiro, 2017; Sousa et al., 2021). 
This current situation has provided a broad and integrated perspec-
tive on biodiversity aspects of Brazilian cladocerans. However, it does 
not seem enough to face the huge challenges that lie ahead. As said 
before, about 50 of the taxa inventoried here need taxonomic revision, 
including widespread and typically planktonic species, such as Moina 
micrura, Ceriodaphnia cornuta, Bosmina longirostris, and Bosmina freyi. At 
the same time, an improvement in the sampling coverage of Brazilian 
freshwaters brings the perspective of new taxa to be described and the 
detection of initial stages in invasion processes by non-native taxa. At 
the risk of sounding repetitive, these important tasks can only be ach-
ieved with investment in resources and taxonomic training. 
3.4. Copepoda 
Approximately 190 species of free-living freshwater copepods were 
found for Brazil in our study, and two exclusively freshwater species are 
invasive (Mesocyclops ogunnus Onabamiro, 1957 from Africa and 
Fig. 2. Synthesis of overall data obtained from the literature to Protozoa, Rotifera, Cladocera, and Copepoda. Each box is composed of a map showing the richness of 
taxa along Brazilian Hydrographic Regions, a donut chart presenting the contribution of lotic and lentic ecosystems to the richness, an accumulation curve built by 
rarefaction method, a bar chart with temporal distribution of studies within Ecology (population or community), Functional Diversity, Molecular Diversity, and 
Taxonomy categories. 
Table 2 
Non-native species of Protozoa, Rotifera, Cladocera, and Copepoda occurring in 
Brazil. 
Taxa Zoogeographic Native Range 
Protozoa – 
Rotifera 
Kellicottia bostoniensis (Rousselet, 1908) Nearctic 
Kellicottia longispina (Kellicott, 1879) Nearctic 
Cladocera 
Eurycercus lamellatus (O.F. Muller, 1785) Palearctic 
Daphnia lumholtzi Sars, 1885 Afrotropic, Australasian, Oriental 
Moina macrocopa (Straus, 1820) Palearctic 
Copepoda 
Mesocyclops ogunnus Onabamiro, 1957 Afrotropic 
Thermocyclops crassus (Fischer, 1853) Palearctic 
L.M.A. Elmoor-Loureiro et al. 
Limnologica xxx (xxxx) xxx
6
Thermocyclops crassus (Fischer, 1853) from Europe). The richest families 
are Cyclopidae (74 species) and Diaptomidae (71 species), followed by 
Parastenocarididae (11 species) (Table S3.4 - Supplementary File 3). 
Considering 12 hydrographic regions in Brazil, the Copepoda rich-
ness is clearly high in the Amazon (87 species) and Paraná (81 species) 
hydrographic regions, which are the largest hydrographic basins in 
Brazil in terms of area (Fig. 2). This result agrees with Perbiche-Neves 
et al. (2014), in a study which pointed to a positive relationship between 
copepod richness and area in freshwater ecoregions of the neotropics. 
Additionally, these basins have the oldest center of studies in limnology, 
and have carried out several studies over time, similar to other studies 
(Silva and Perbiche-Neves, 2017) (Table 1). 
Seven hydrographic regions have between 36 and 53 species, with a 
mean of 44 ± 7 species (East Atlantic-36, Northeast Atlantic Oriental 
Region-53, South Atlantic-44, Southeast Atlantic-52, Paraguay-50, São 
Francisco-36 and Tocantins-Araguaia-37). The hydrographic regions 
with the lower number of species are Parnaíba (7 species), Uruguay (5 
species), and Northeast Atlantic Occidental Region (1 species) (Table 1). 
Most of the copepod species analyzed here occur in lentic environ-
ments (natural and artificial), followed by lotic and miscellaneous 
(Fig. 2), and this finding agrees with Silva and Perbiche-Neves (2017) 
for microcrustacean trends in Brazil between 1990 and 2014. 
The richness curve for Copepoda is far from being stabilized, ac-
cording to our data, considering all hydrographic regions (Fig. 2). 
Among the types of the studies on copepods, taxonomy was predominant 
until 1979. Between 1980 and 1989, ecology was still equal to taxonomy 
and, after 1990, the number of ecology studies increased greatly, rep-
resenting more than 80% after 2000. Molecular diversity is poorly 
studied, and functional ecology, although there are still few studies, 
looks set to increase in the next few years (Fig. 2). 
The oldest valid species described for Brazil was Notodiaptomus gibber 
(Poppe, 1889 in De Guerne and Richard, 1889); however, ProfessorCarlos E. F. da Rocha (pers. comm.) pointed out that the first but not 
validated was Cyclops brasiliensis described by Dana in 1848, in Rio de 
Janeiro. He considers the species as inquirenda in the “Catálogo de 
Crustacea do Brasil” (Rocha and Coelho-Botelho, 1998), because of the 
very simple description of details and drawings. Professor Rocha 
searched for Dana’s materials and discovered that they were lost in the 
sinking of the ship that transported the Dana collection, near San 
Francisco Bay (USA). 
Cyclopoida do not show a clear endemism pattern across the country, 
but for the Diaptomidae family of the Calanoida order, several clades of 
freshwater ecoregions separated by parsimony analysis were formed in 
the Neotropical region and within Brazil (Perbiche-Neves et al., 2014), 
agreeing with other studies, such as Brandorff (1976) and 
Suárez-Morales et al. (2005). For Harpacticoida, the data did not allow 
the detection of endemism patterns. 
3.5. Functional and molecular diversity 
Studies on functional diversity of zooplankton organisms in Brazil 
are recent, having started in the 2010s, especially for Protozoa (e.g., 
Segovia et al., 2016) and Rotifera (e.g., Moreira et al., 2016). All the 
changes in the zooplankton functional diversity are related to the fea-
tures of the environment (abiotic and biotic factors), such as presence or 
absence of predator, competitor and habitat availability, which change 
the structure and trophic dynamics in the environment. The functional 
richness, for example, rose according to the increase in the macrophytes 
and fish richness due to the increase in food and habitat availability and 
selective predation, as well as rising functional beta diversity and fish 
abundance. Considering functional traits, the body size was negatively 
influenced by macrophyte and fish predation, and feeding type was 
positively influenced by macrophytes (Deosti et al., 2021). Eutrophica-
tion and cyanobacterial dominance change the composition of traits, 
which favored the specialized species and reduced functional dispersion, 
leading to zooplankton niche overlap (Josué et al., 2019; Duré et al., 
2021). However, the most difficult decision related to methodology in 
the studies is choosing the functional traits of the species that can really 
translate the functional role of the species in the environment. 
Research on organisms recorded in planktonic samples with a focus 
on molecular diversity has appeared on the Brazilian scientific scene 
only in the last decade (e.g., Abreu et al., 2010; Castilho et al., 2015; 
Lentendu et al., 2019; Fernandes et al., 2021). Despite being a promising 
research field, the number of papers is still low (Fig. 2). 
These results reveal the need and urgency for more research on 
functional and molecular diversity of all studied groups and in different 
continental aquatic environments in the country. 
4. Conclusions 
From the data analyzed herein, similar patterns of richness were 
observed for all zooplankton groups investigated. Portions of Hydro-
graphic Regions with high richness seem to be under long-term effort of 
sampling or closer to consolidated research groups dedicated to Ecology, 
Limnology or Taxonomy. The most critical data correspond to Protozoa, 
which presented large richness concentrated only in areas of the Paraná 
Hydrographic Region. In all cases, such findings demonstrate that 
geographical distribution and richness patterns are extremely dependent 
on spatial proximity to the centers of study, increasing the effect of 
shortfalls on the knowledge of Protozoa, Rotifera, Cladocera, and 
Copepoda in Brazil. Perhaps for this reason, the accumulation curves for 
all zooplankton groups investigated did not present asymptote. Thus, 
there are many areas still to be sampled in Brazil. Cooperation projects 
dealing with freshwater biodiversity inventories in as yet unstudied 
areas might have an effect on the data of geographical distribution of 
known Neotropical species, including non-native species, and on the 
description of new taxa. 
Another similar pattern was found when observing the types of 
ecosystems, with high contribution to richness observed in lotic and 
lentic natural water bodies. At the same time, lentic artificial ecosystems 
are very important to freshwater biodiversity conservation in Brazil 
because they are associated with several events of biological invasion, 
especially with microcrustaceans and Rotifera. On the other hand, an 
increase in studies focusing on other environments, such as ponds and 
semi-terrestrial, going beyond the strict focus on plankton, are needed. 
These kinds of environments frequently presented endemic fauna. 
While taxonomic studies were abundant until the end of the 1980s, 
after 1990 ecological studies have taken on great importance, and new 
areas such as functional ecology are promising. However, much work is 
still needed on taxonomy and molecular ecology, which are areas with 
several gaps. 
Credit authorship contribution statement 
Conceptualization - C.C. Bonecker; P.H.C. Corgosinho; L.M.A. 
Elmoor-Loureiro; C.Y. Joko; F.A. Lansac-Tôha; G. Perbiche-Neves; F.D. 
R. Sousa; L.F.M. Velho. Methodology - C.E.G. Aggio; R.L. Arrieira; L.S.M. 
Braghin; C.C. Bonecker; M.S.M. Castilho-Noll; P.H.C. Corgosinho; S. 
Deosti; L.P. Diniz; N.G. Dos Santos; L.M.A. Elmoor-Loureiro; E.M. Eski-
nazi-Sant’Anna; V. Gazulha; A.R. Ghidini; C.Y. Joko; F.A. Lansac-Tôha.; 
F.M. Lansac-Tôha; R.L. Macêdo; P.M. Maia-Barbosa; T. Mantovano; B.R. 
Meira; M. Melo Júnior; C.S. Morais-Junior; J.R. Nascimento; F.R. Oli-
veira; P.M. Oliveira; G. Perbiche-Neves; J.L. Portinho; M. Progênio; M.A. 
Rocha; L.O. Santana; L.T.F. Schwind; A.J. Silva; J.V.F. Silva; M.L.C. 
Silva; N.J. Silva; N.R. Simões; F.D.R. Sousa; L. Veado; L.F.M. Velho. 
Formal analysis - A.R. Ghidini; J.L. Portinho; N.R. Simões; F.D.R. Sousa. 
Investigation - C.E.G. Aggio; R.L. Arrieira; L.S.M. Braghin; C.C. Bone-
cker; G.S. Cabral; M.S.M. Castilho-Noll; L.J. Chiarelli; P.H.C. Corgo-
sinho; S. Deosti; L.P. Diniz; N.G. Dos Santos; L.M.A. Elmoor-Loureiro; E. 
M. Eskinazi-Sant’Anna; V. Gazulha; A.R. Ghidini; C.Y. Joko; F.A. Lansac- 
Tôha.; F.M. Lansac-Tôha; R.L. Macêdo; P.M. Maia-Barbosa; T. Man-
tovano; B.R. Meira; M. Melo Júnior; C.S. Morais-Junior; P.H. Morari; J. 
L.M.A. Elmoor-Loureiro et al. 
Limnologica xxx (xxxx) xxx
7
R. Nascimento; F.R. Oliveira; P.M. Oliveira; G. Perbiche-Neves; J.L. 
Portinho; M. Progênio; M.A. Rocha; L.O. Santana; L.T.F. Schwind; Silva, 
A.C.S. da Silva; A.J. Silva; J.V.F. Silva; M.L.C.M. Silva; N.J. Silva; N.R. 
Simões; F.D.R. Sousa; L. Veado; L.F.M. Velho. Data Curation - R.L. 
Arrieira; C.C. Bonecker; L.S.M. Braghin; P.H.C. Corgosinho; L.P. Diniz; 
N.G. Dos Santos; L.M.A. Elmoor-Loureiro; A.R. Ghidini; C.Y. Joko; F.M. 
Lansac-Tôha; M. Melo Júnior; C.S. Morais-Junior; Nascimento, J.R; F.R. 
Oliveira; G. Perbiche-Neves; J.L. Portinho; M.A. Rocha; L.T.F. Schwind; 
J.V.F. Silva; M.L.C. Silva; N.R. Simões; F.D.R. Sousa; L.F.M. Velho. 
Writing - Original Draft - C.E.G. Aggio; R.L. Arrieira; C.C. Bonecker; L.S. 
M. Braghin; P.H.C. Corgosinho; S. Deosti; L.P. Diniz; L.M.A. Elmoor- 
Loureiro; E.M. Eskinazi-Sant’Anna; C.Y. Joko; F.A. Lansac-Tôha.; F.M. 
Lansac-Tôha; R.L. Macêdo; T. Mantovano; B.R. Meira; M. Melo Júnior; C. 
S. Morais-Junior; F.R. Oliveira; G. Perbiche-Neves; J.L. Portinho; M. 
Progênio; M.A. Rocha; L.O. Santana; L.T.F. Schwind; M.L.C. Silva; N.J. 
Silva; N.R. Simões; F.D.R. Sousa; L.F.M. Velho. Writing - Review & 
Editing- C.E.G. Aggio; R.L. Arrieira; L.S.M. Braghin; C.C. Bonecker; P.H. 
C. Corgosinho; S. Deosti; L.P. Diniz; N.G. Dos Santos; L.M.A. Elmoor- 
Loureiro; E.M. Eskinazi-Sant’Anna; A.R. Ghidini; C.Y. Joko; F.A. Lan-
sac-Tôha.; F.M. Lansac-Tôha;R.L. Macêdo; P.M. Maia-Barbosa; T. 
Mantovano; B.R. Meira; M. Melo Júnior; C.S. Morais-Junior; F.R. Oli-
veira; G. Perbiche-Neves; J.L. Portinho; M. Progênio; M.A. Rocha; L.O. 
Santana; L.T.F. Schwind; A.J. Silva; J.V.F. Silva; M.L.C. Silva; N.R. 
Simões; F.D.R. Sousa; L.F.M. Velho. Visualization - C.E.G. Aggio; R.L. 
Arrieira; L.S.M. Braghin; C.C. Bonecker; M.S.M. Castilho-Noll; P.H.C. 
Corgosinho; S. Deosti; L.P. Diniz; N.G. Dos Santos; L.M.A. Elmoor- 
Loureiro; E.M. Eskinazi-Sant’Anna; V. Gazulha; A.R. Ghidini; C.Y. 
Joko; F.A. Lansac-Tôha.; F.M. Lansac-Tôha; R.L. Macêdo; P.M. Maia- 
Barbosa; T. Mantovano; B.R. Meira; M. Melo Júnior; C.S. Morais- 
Junior; J.R. Nascimento; F.R. Oliveira; P.M. Oliveira; G. Perbiche- 
Neves; J.L. Portinho; M. Progênio; M.A. Rocha; L.O. Santana; L.T.F. 
Schwind; A.C.S. da Silva; A.J. Silva; J.V.F. Silva; M.L.C. Silva; N.R. 
Simões; F.D.R. Sousa; L. Veado; L.F.M. Velho. Supervision - C.C. Bone-
cker; P.H.C. Corgosinho; L.M.A. Elmoor-Loureiro; C.Y. Joko; F.A. Lan-
sac-Tôha.; F.R. Oliveira; G. Perbiche-Neves; F.D.R. Sousa; L.F.M. Velho. 
Project administration - L.M.A. Elmoor-Loureiro; F.D.R. Sousa. 
CRediT authorship contribution statement 
This manuscript has not been published previously it is not under 
consideration for publication elsewhere. The final version was approved 
by all authors and tacitly or explicitly by the responsible authorities 
where the work was carried out, and that, if accepted, it will not be 
published elsewhere in the same form, in English or in any other lan-
guage, including electronically without the written consent of the 
copyright-holder. 
Declaration of Competing Interest 
The authors declare that they have no known competing financial 
interests or personal relationships that could have appeared to influence 
the work reported in this paper. 
Appendix A. Supporting information 
Supplementary data associated with this article can be found in the 
online version at doi:10.1016/j.limno.2022.126008. 
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	Towards a synthesis of the biodiversity of freshwater Protozoa, Rotifera, Cladocera, and Copepoda in Brazil
	1 Introduction
	2 Material and methods
	3 Results and discussion
	3.1 Protozoa
	3.2 Rotifera
	3.3 Cladocera
	3.4 Copepoda
	3.5 Functional and molecular diversity
	4 Conclusions
	Credit authorship contribution statement
	CRediT authorship contribution statement
	Declaration of Competing Interest
	Appendix A Supporting information
	References