Record of aquatic invertebrates associated with the bromeliads Aechmea ornata (Baker) and Aechmea recurvata (Klotzsch) L. B. Sm in two Atlantic Rainforest fragments of south Brazil

Abstract

Bromeliads are numerous both in diversity and abundance in the Atlantic Rainforest, one of the most threatened biomes on Earth. They are also an important habitat for several animal species that live in the water retained by its rosette structure. In this study, we aimed to characterize and compare the macroinvertebrate community of two bromeliad species from Atlantic forest fragments. One is characteristic of Dense Ombrophilous Forests, Aechmea ornata, and the other occurs in Mixed Ombrophilous Forests, Aechmea recurvata. We sampled and identified macroinvertebrates form 24 bromeliads, 12 of each species. A total of 19 taxa and 656 specimens were sampled. In both bromeliad species, Diptera was the most abundant insect order (n=417), represented mostly by mosquitoes (n=227) and non-biting midges (n=107), followed by Scirtidae marsh beetles (n=216). Aechmea ornata presented higher abundance and lower invertebrate richness compared to Aechmea recurvata. Seven taxa were exclusive of A. recurvata, eight were common to both, and four, including a sponge, occurred only in A. ornata. The most abundant invertebrate taxa were common to both bromeliads and are regarded as typical of lentic environments. On the other hand, the high number of exclusive species indicates that conservation efforts should include bromeliads from these two phytophysiognomies to secure protecting the diversity of phytotelmata communities.

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Acta Limnologica Brasiliensia, 2021, vol. 33, e25
https://doi.org/10.1590/S2179-975X3621
ISSN 2179-975X on-line version
is is an Open Access article distributed under the terms of the Creative Commons Attribution License, which permits
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Record of aquatic invertebrates associated with the bromeliads
Aechmea ornata (Baker) and Aechmea recurvata (Klotzsch) L. B.
Sm in two Atlantic Rainforest fragments of south Brazil
Registro de invertebrados aquáticos associados às bromélias Aechmea ornata (Baker) e
Aechmea recurvata (Klotzsch) L. B. Sm em dois fragmentos de Mata Atlântica do sul
do Brasil
Yuri Kataoka Silva1 , Anderson de Souza Moser2 , Mônica Ceneviva-Bastos1  and
Ana Lucia Suriani Aonso1* 
1
Universidade Estadual do Centro-Oeste – Unicentro, Alameda Élio Antonio Dalla Vecchia, 838,
CEP 85040-167, Guarapuava, PR, Brasil
2
Universidade Federal do Paraná – UFPR, Rua XV de Novembro, 1299, CEP 80060-000, Curitiba,
PR, Brasil
*e-mail: analuciabio@gmail.com
Cite as: Silva, Y.K.etal. Record of aquatic invertebrates associated with the bromeliads Aechmea
ornata (Baker) and Aechmea recurvata (Klotzsch) L. B. Sm in two Atlantic Rainforest fragments of
south Brazil. Acta Limnologica Brasiliensia, 2021, vol. 33, e25.
Abstract: Bromeliads are numerous both in diversity and abundance in the Atlantic Rainforest,
one of the most threatened biomes on Earth. ey are also an important habitat for several animal
species that live in the water retained by its rosette structure. In this study, we aimed to characterize
and compare the macroinvertebrate community of two bromeliad species from Atlantic forest
fragments. One is characteristic of Dense Ombrophilous Forests, Aechmea ornata, and the other occurs
in Mixed Ombrophilous Forests, Aechmea recurvata. We sampled and identied macroinvertebrates
form 24 bromeliads, 12 of each species. A total of 19 taxa and 656 specimens were sampled. In
both bromeliad species, Diptera was the most abundant insect order (n=417), represented mostly by
mosquitoes (n=227) and non-biting midges (n=107), followed by Scirtidae marsh beetles (n=216).
Aechmea ornata presented higher abundance and lower invertebrate richness compared to Aechmea
recurvata. Seven taxa were exclusive of A. recurvata, eight were common to both, and four, including
a sponge, occurred only in A. ornata. e most abundant invertebrate taxa were common to both
bromeliads and are regarded as typical of lentic environments. On the other hand, the high number
of exclusive species indicates that conservation eorts should include bromeliads from these two
phytophysiognomies to secure protecting the diversity of phytotelmata communities.
Keywords: phytotelmata; aquatic insects; Ombrophilous Forests; diversity.
Resumo: As bromélias são numerosas em diversidade e abundância na Mata Atlântica, um dos
biomas mais ameaçados do planeta. Elas também são um habitat importante para várias espécies de
animais que vivem na água retida por sua estrutura em roseta. Neste estudo, objetivamos caracterizar
e comparar a comunidade de macroinvertebrados de duas espécies de bromélias de fragmentos de
Mata Atlântica. Uma é característica de Floresta Ombróla Densa, Aechmea ornata, e a outra ocorre

2 Silva, Y.K.etal.
Acta Limnologica Brasiliensia, 2021, vol. 33, e25
Ombrophilous Forest (MOF), sampled at the
Parque Municipal das Araucárias, Guarapuava,
Paraná (25°21’35” S; 51°28’16” W), within an area
of approximately 30 m2 (with an average distance of
3 m among individuals). e two sampling sites are
preserved areas within Conservation Units, though
the surroundings of the Parque Municipal das
Araucárias are under stronger anthropic pressure.
e A. ornata bromeliads were in more opened
canopy sites; their tanks held on average 142 ml of
water (± 104.8), with an average pH of 6.2 (± 1.2)
and 8.5 mg.L-1 (± 0.7) of dissolved oxygen. Aechmea
recurvata were in closed canopy and held 55 ml (±
12.2) of water on average, with an average pH of
4.5 (± 0) and 5.5 mg.L-1 (± 3.5) of dissolved oxygen.
The sampled macroinvertebrates were fixed
in 10% formalin solution, sorted, identied, and
preserved in ethanol 70%. To characterize bromeliad
communities, we calculated macroinvertebrate
relative abundance and manually built a descriptive
network with species occurrence in each bromeliad
species. We also did a similarity analysis (ANOSIM)
in the Primer v6 software (Clarke & Gorley, 2006),
with log (X+1)-transformed abundance data and
the Bray-Curtis similarity coefficient, to check
whether invertebrate communities diered between
bromeliads.
A total of 656 specimens of 19 taxa were
sampled, being 12 taxa and 413 specimens sampled
in A. ornata and 15 taxa and 243 specimens in A.
recurvata (Table1). Seven taxa were exclusive of A.
recurvata, eight were common to both, and four,
including a sponge, occurred only in A. ornata
(Figure1). Overall, the macroinvertebrate samples
from A. ornata and A. recurvata were separated,
though with a large overlap (ANOSIM: R = 0.337;
P = 0.001).
e epiphytic habit of A. recurvata bromeliads
in the MOF, a site with dense canopy cover and
high precipitation, probably contributed to the
large accumulation of leaf litter and organic matter
e Atlantic Rainforest is a biodiversity hotspot
that harbors several endemic species (Myersetal.,
2000; Fundação SOS Mata Atlântica, 2017).
For instance, of the 803 species of bromeliads that
occur in the Atlantic Forest, 653 are endemic from
this biome (Martinellietal., 2008). e bromeliads
are rosette-forming plants classied as phytotelmata
(see Freitasetal., 2016 for examples). e typical
bromeliad morphological structure originates
from the overlapping of leaves and formation of a
tank that accumulates water (i.e., the phytotelma)
and decomposing organic matter. is creates an
ecologically important microcosm for diverse animal
communities, which use the bromeliads as a site for
foraging, refuge, and reproduction (Richardson,
1999; Sodréetal., 2010). Aquatic invertebrates
such as annelids, atworms, gastropods, bivalves,
crustaceans, mites, and mostly insects comprise
the major animal groups that occupy bromeliad
habitats (Hauer and Resh, 1996). In this study,
we aimed to characterize the macroinvertebrate
community of two bromeliad species from distinct
Atlantic Forest phytophysiognomies. The two
species, Aechmea ornata and Aechmea recurvata,
are botanically known and well-studied, though we
did not nd studies that investigated the associated
macroinvertebrate fauna.
We sampled aquatic macroinvertebrates
in November/2015, September/2016, and
December/2016 from 24 bromeliads by aspiration
of the tank water, according to the methodology
described by Sodréetal. (2010), selected to prevent
plant damage. Twelve of these were the rupicolous
bromeliad Aechmea ornata, which is a common
endemic species in the Dense Ombrophilous Forest
(DOF) (Martinellietal., 2008), sampled at the Ilha
do Mel, Paranaguá, Paraná (25°33’44” S; 48°19’07”
W) within an area of approximately 40 m2 (with
an average distance of 2 m among individuals).
e other 12 were the epiphytic Aechmea recurvata,
a common and widespread species in the Mixed
em Floresta Ombróla Mista, Aechmea recurvata. Amostramos e identicamos macroinvertebrados
de 24 bromélias, 12 de cada espécie. No total foram registrados 19 táxons e 656 espécimes. No geral,
Diptera foi a ordem de inseto mais abundante (n = 417), representada principalmente por culicídeos
(n = 227) e quironomídeos (n = 107), seguidos por besouros Scirtidae (n = 216). Comparativamente
a A. recurvata, A. ornata apresentou maior abundância e menor riqueza de invertebrados. Sete táxons
foram exclusivos de A. recurvata, oito foram comuns a ambas e quatro, incluindo uma esponja,
ocorreram apenas em A. ornata. Até onde sabemos, este é o primeiro registro de esponjas em bromélias.
Observamos que os táxons de invertebrados mais abundantes eram comuns a ambas as bromélias e
são considerados típicos de ambientes lênticos. Por outro lado, o elevado número de táxons exclusivos
indica que os esforços de conservação devem incluir bromélias dessas duas tosionomias para garantir
a diversidade das comunidades de totelmatas.
Palavras-chave: totelmata; insetos aquáticos; Floresta Ombróla; diversidade.

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Acta Limnologica Brasiliensia, 2021, vol. 33, e25
Figure 1. Network representing the occurrence (lines) of macroinvertebrate taxa (white circles) in the bromeliads
Aechmea ornata (AO) and Aechmea recurvata (AR) (black circles) in Atlantic Rainforest fragment of the Paraná
state. e circle size is proportional to species abundance categories (intervals of 50 specimens), with smallest circles
representing taxa with up to ten specimens and largest circles representing taxa with 150-200 specimens. Taxa
abbreviations are listed in Table1.
Table 1. List of taxa, name abbreviations (used in Figure1), abundance, functional feeding groups and richness of
macroinvertebrates sampled in Aechmea ornata and Aechmea recurvata bromeliads from Dense Ombrophilous Forest
and Mixed Ombrophilous Forest fragments, respectively, Paraná state, Brazil.
Animal taxa Abbreviation
Bromeliad species
Functional feeding group
Aechmea
ornata
Aechmea
recurvata
Porifera Porif 1 0 Filterer
Naididae
Pristina Prist 0 13 Collector gatherer
Araneae Aran 2 0 Predator
Hydracarina Hydr 0 1 Predator
Collembola Coll 0 1 Collector gatherer
Formicidae Form 1 3 Shredder
Culicidae Cul 145 82 Collector gatherer
Ceratopogonidae (adult/pupae) Cerato 2 11 Collector gatherer; predator
Atrichopogon Atric 0 6 Collector gatherer
Bezzia Bezz 0 6 Predator
Ceratopogonidae morphotype I* D_morph 34 23 –
Chironomidae
Ablabesmyia Ablab 8 0 Predator
Limnophyes Limno 14 66 Collector gatherer
Polypedilum Polyp 13 0 Shredder; collector gatherer
Empididae Empid 2 1 Predator
Psychodidae Psychod 1 1 Collector gatherer, scraper
Tipulidae Tipul 0 2 Predator
Noteridae Noter 0 1 Predator
Scirtidae Scirt 190 26 Scraper
Total abundance – 413 243
Total taxa richness – 12 15
*Ceratopogonidae morphotype I – unidentied Ceratopogonidae genus (D_morph = Diptera morphotype).

4 Silva, Y.K.etal.
Acta Limnologica Brasiliensia, 2021, vol. 33, e25
in their tanks. is nding may be related to the
exclusive presence of Pristina (Naididae) earthworms
in these bromeliads, since they feed on plant detritus
and play an important role in organic matter
processing (Gorni & Alves, 2015). On the other
hand, the rupicolous habit of A. ornata bromeliads,
present in more opened areas of DOF near the
beach, probably contributed to clearer water and
lower organic matter concentration (pers. obs.).
is nding may in turn be related to the exclusive
presence of sponges in these bromeliads. Sponges
can inhabit several freshwater environments, both
permanent and temporary, being more frequent in
clear waters, free from contaminants (Mugnaietal.,
2010), and in silicate-rich waters due to their
skeleton constitution (Volkmer-Ribeiro & Parolin,
2010). ere are numerous records of sponges in
Atlantic Forest streams, such as Oncosclera jewelli
(Volkmer, 1963), registered in streams of the
Planalto das Araucárias and Campos de Altitude
regions (Macieletal., 2001), though their presence
in bromeliad plants is quite uncommon – to our
knowledge, this is the rst record. However, it was
one single record and since it was not possible
to identify the sponge, it is possible that this
occurrence may have been accidental. us, further
studies are needed to investigate sponge occurrence
in tank plants and their relationships with other
bromeliad animals.
e most abundant insect order was Diptera,
especially the mosquito larvae (Culicidae, the
most abundant family) and non-biting midges,
followed by Coleoptera, represented mostly by
scirtid beetles. Mosquito larvae are cosmopolitan
and quite common in lentic waters, occupying
from large lakes to rainfall ponds and phytotelmata.
ey play an important role in bromeliad food
webs and can occupy dierent trophic levels, form
primary consumers, as microbial lter-feeders, to
top predators (Kitching, 2001). Representatives of
many mosquito genera, such as Aedes, reproduce
almost exclusively in phytotelmata (Kitching,
2001). Similarly, many beetles of the Scirtidae
family, which also presented a high abundance
in our study, are also closely associated with
phytotelmata (Mestreetal., 2001). They are
detritivorous species (i.e., feed on dead organic
matter) that break-up large detritus into finer
particles, making them available for other animals;
they can even act as facilitators to other species,
such as midges (Kitching, 2001). Conversely,
non-biting midges of the Chironomidae family
are associated not only to lentic habitats, being
widespread, rich, and abundant in virtually all other
freshwater environments - and some salty ones as
well (Ferrington-Junior, 2008). ey were the third
most abundant family in this study. Limnophyes
chironomid midges had already been sampled in
Atlantic Forest and mountain streams, though they
are rarely found in bromeliads (Trivinho-Strixino,
2011); Polypedilum, however, are known to occur
in both epiphytic and rupicolous tank plants
(Saetheretal., 2010). Collector-gatherers, which
feed on ne particulate organic matter deposited
on the submerged substrate, and scrapers, which
scrape periphyton from the surface of the leaves
(Berg, 1995), were the predominant functional
feeding groups in both bromeliads.
e phytotelmata present in A. ornata and A.
recurvata harbored an aquatic invertebrate richness
comparable to another Aechmea species (Aechmea
distichantha Lem.), which found 16 morphospecies
of four insect orders (Amadeoetal., 2017). ere
are several factors that can inuence the diversity
of bromeliad invertebrate communities, from
uctuations of tank water volume and limnological
parameters (Nevesetal., 2019) to owering events
(Amadeoetal., 2017). When it comes to rupicolous
bromeliads near the sea, however, variations in
salinity can also occur due to sea breeze. Although
it was not possible to measure tank water salinity
in our study, the lower taxa richness and higher
abundance registered in A. ornata may have resulted
from it. For instance, Polypedilum and Ablabesmyia
are more commonly found in freshwaters, though
they can tolerate some salinity (Williamsetal.,
1991), what may explain their occurrence in A.
ornata.
In summary, the most abundant
macroinvertebrate taxa were common to both
bromeliad species and are regarded as typical of
lentic environments. Notwithstanding, over half
of the sampled taxa were exclusive, being 21%
exclusive of A. ornata and 37% exclusive of A.
recurvata.
us, our results indicate that conservation
efforts should include bromeliads from these
two phytophysiognomies to ensure the diversity
of phytotelmata communities in the Atlantic
Rainforest. Further studies that add up to a broader
sampling eort to inventory macroinvertebrates
of bromeliad phytotelmata in the Atlantic
Forest would be welcome and could include the
investigation of the occurrence of sponges in these
tank plants.

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Record of aquatic invertebrates associated with the bromeliads...
Acta Limnologica Brasiliensia, 2021, vol. 33, e25
Acknowledgements
We are thankful to Dr. Rosângela C. Tardivo
from the Universidade Estadual de Ponta Grossa
and to Dr. Adriano Silvério from the Universidade
Estadual do Centro-Oeste for the identication of
the two bromeliad species.
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Received: 10 May 2021
Accepted: 14 September 2021
Associate Editor: Victor Satoru Saito.