Metazoan Parasite Infracommunities Research Articles

Temporal and seasonal variation of metazoan parasites in Pimelodus ornatus (Siluriformes: Pimelodidae) from the Amazon River, Brazil.

This study investigated the influence of temporal and seasonal (i.e., rainy and dry seasons) in communities and infracommunities of metazoan parasites in Pimelodus ornatus, a Siluriformes from the Amazon River in the state of Amapá, in northern Brazil. A total of 144 fish were collected during 2020 and 2021. In 2020, five species of parasites were found (1 Monogenea, 3 Nematoda, and 1 Cestoda), and in 2021, seven species of parasites were found (1 Monogenea, 3 Nematoda, 1 Cestoda, 1 Digenea, and 1 Crustacea). In 2020, Spirocamallanus inopinatus was dominant, while in 2021, Contracaecum sp. was dominant. Brillouin diversity, species richness, and evenness were higher in 2021. There were differences for some parasite infracommunities between years and between seasonal periods. Brillouin diversity and species richness were higher in the rainy season, but there were no differences in evenness and Berger-Parker dominance between the seasonal periods. Temporal and seasonal variations were mainly influenced by rainfall levels, oxygen levels, and environmental temperature and, later, by the availability of infective stages of parasites in the environment and the size of the hosts. Lastly, the influence of the temporal and seasonal variations on the structure of communities and infracommunities of parasites was weaker than expected due to the low infection rates of the majority of parasites found.

Metazoan Parasite Infracommunities of the Tomtate Grunt (Haemulon aurolineatum) as Potential Bioindicators of Environmental Conditions in the Yucatan Continental Shelf, Mexico.

Metazoan parasites and their hosts experience the environmental influence of the seasonal presence of Western and Eastern subzones with different oceanographic conditions on the Yucatan continental shelf (YS). In addition, natural seeps and oil transport in the area lead to the presence of hydrocarbons. We hypothesized that the parasite infrapopulations and infracommunities of Haemulon aurolineatum will respond to environmental variability related with ongoing oceanographic subzone conditions of the YS. Spatial and multivariate statistical analyses were used to test this hypothesis. For 17 sampling sites along the YS, 55 parasite morphospecies were recovered from 146 fishes. There were significant differences in the number of parasite species between Western and Eastern subzones, but not in the number of parasite individuals. Spatial autocorrelation on environmental variables as a consequence of the Yucatan current was found. Overall, the parasite metrics suggested a region with a good ecosystem health condition naturally influenced by oceanographic processes.

Conspecific migration and environmental setting determine parasite infracommunities of non-migratory individual fish.

Parasite infracommunities tend to be stochastic in nature, although environmental characteristics such as the type of water source in streams and host traits can have an effect on the biotic assemblages and by extension the parasite fauna. We examined the effect of water source and the rate of adult fish migration on the metazoan parasite infracommunities of conspecific juvenile brown trout, Salmo trutta L. among streams flowing into Lake Lucerne (Switzerland). Juvenile (1 to 2-year old) fish harboured higher parasite species richness in groundwater-fed than in surface water-fed streams, whereas the rate of fish migration did not affect infracommunity richness. Heteroxenous species were more common in groundwater-fed streams with high and medium rates of trout migration, whereas infracommunities in surface water-fed streams and streams with low rates of fish migration were dominated by one monoxenous parasite or lacked infections. Similarity in the parasite infracommunity composition of juvenile trout across streams was explained by the interaction between type of water source and adult migration rates. Our conclusions support that similarity in the parasite composition of resident freshwater conspecifics can be predicted by the local environmental settings and host migratory behaviour, whereas parasite richness is mainly influenced by the environmental characteristics.

Community and infracommunities of metazoan parasites in Hemiodus unimaculatus (Hemiodontidae) from Jari River basin, a tributary of Amazon River (Brazil)

Abstract Parasites are an important part of biodiversity, and knowledge of species and their relationship with their hosts helps in monitoring an ecosystem over time. The aim of this study was to investigate the fauna of metazoan parasites in Hemiodus unimaculatus from the Jari River, in the eastern Amazon region, northern Brazil. Of the fish examined, 96.7% were parasitized by one or more species, and a total of 336 parasites such as Dactylogyridae gen. sp.1, Dactylogyridae gen. sp.2, Dactylogyridae gen. sp.3, Dactylogyridae gen. sp.4, Gyrodactilydae gen. sp., Urocleidoides sp.1, Urocleidoides sp.2, Urocleidoides sp.3, metacercariae of Digenea gen. sp., Procamallanus (Spirocamallanus) inopinatus, Contracaecum sp., Neoechinorhynchus sp. and Acarina gen. sp. The parasite community showed low Brillouin diversity (0.58 ± 0.29), low evenness (0.44 ± 0.21) and low species richness (7.40 ± 3.83). There was a predominance of ectoparasites, mainly monogeneans and digeneans. The parasites showed an aggregate dispersion, except for P. (S.) inopinatus, which had a random dispersion. The size of the hosts had no effect on diversity, species richness and abundance of parasites, but other factors structured the parasite community. This is the first study on the parasite community and infracommunities in H. unimaculatus.

Metazoan parasites of Plagioscion squamosissimus (Osteichthyes: Sciaenidae) of two rivers from the eastern Amazon (Brazil).

Plagioscion squamosissimus, commonly known as "white hake" or "corvina", is among the most commercially important fish in the Amazon and host to a great diversity of endoparasites. The aim of the present study was to compare the communities and infracommunities of metazoan parasites that infect P. squamosissimus in two rivers from the eastern Brazilian Amazon. A total of 75 specimens of P. squamosissimus were collected from Lago Grande do Curuái and the mouth of the Tapajós River. Morphological analysis revealed the presence of 16 parasite species, three myxozoans (Myxobolus sp., Ceratomyxa sp., Henneguya sp.), two trematodes (Austrodiplostomum compactum, Digenea gen. sp. (metacercariae), two monogeneans (Diplectanum sp., Euryhaliotrema sp.), three nematodes (Procamallanus (S.) sp., Anisakis sp., Pseudoproleptus sp. (larva), two acanthocephalans (Rhadinorhynchus plagioscionis and Neoechinorhynchus sp.), one Cestoda Ptychobothriidae gen. sp. and three crustaceans (Therodamas sp., Ergasilus sp., Dolops sp.). Six new records of parasites of P. squamosissimus were made, of which three were myxozoan species, one nematode species and two crustacean species. There were differences in the component communities of parasites of both rivers studied and these were discussed.

Metazoan Parasite Infracommunities of Rhizoprionodon acutus from Nellore Coast off Bay of Bengal Andhra Pradesh, India

Metazoan Parasite Infracommunities of Rhizoprionodon acutus from Nellore Coast off Bay of Bengal Andhra Pradesh, India

Metazoan parasite infracommunities of the dusky flounder (Syacium papillosum) as bioindicators of environmental conditions in the continental shelf of the Yucatan Peninsula, Mexico

BackgroundWe assessed metrics of the metazoan parasite infracommunities of the dusky flounder (Syacium papillosum) as indicators of aquatic environmental health of the Yucatan Shelf (YS) prior to oil extraction. We sampled the dusky flounder and its parasites along the YS, mostly during the 2015 north wind season (November–April). Our aims were: (i) to determine whether the parasite infracommunity metrics of S. papillosum exhibit significant differences among YS subregions; (ii) to determine whether the probability of the occurrence of its parasite species and individuals were affected by environmental variables, nutrients, heavy metals and hydrocarbons at the seascape level; and (iii) to determine whether there were statistical differences between the parasite infracommunity metrics of S. papillosum from YS and those of Syacium gunteri from the Campeche Sound. Multivariate statistical analyses and generalised additive models (GAMs) were used to examine the potential statistical associations between the contaminants, environmental variables and parasite community metrics, and the maximum entropy algorithm (MaxEnt) was used to characterise the habitat’s suitability for the parasite’s probability of occurrence.ResultsWe recovered 48 metazoan parasite species from 127 S. papillosum, with larval cestodes and digeneans being the most numerically-dominant. Multivariate analyses showed significant differences in parasite infracommunity metrics among Western YS, Mid YS and Caribbean subregions, with the latter being the richest in species but not in individuals. The GAM and MaxEnt results indicated a negative effect of top predators (e.g. sharks and rays) removal on parasite metrics. The parasite infracommunities of S. papillosum were twice as rich in the number of species and individuals as those reported for S. gunteri from the Campeche Sound.ConclusionsThe significant differences among subregions in parasite metrics were apparently due to the interruption of the Yucatan current during the north wind season. The fishing of top predators in combination with an influx of nutrients and hydrocarbons in low concentrations coincides with an increase in larval cestodes and digeneans in S. papillosum. The dusky flounder inhabits a region (YS) with a larger number of metazoan parasite species compared with those available for S. gunteri in the Campeche Sound, suggesting better environmental conditions for transmission in the YS.

Metazoan parasites of Plagioscion squamosissimus, an invasive species in the Tietê River, São Paulo, Brazil.

This study focused on the characterization and analysis of communities and infra-communities of metazoan parasites of Plagioscion squamosissimus caught in Promissão Reservoir in the Tietê River in Borborema (21°39'58"S, 49°8'49"W), state of São Paulo, Brazil. Fifty adult specimens caught by professional fishermen in March 2015 were necropsied. The fish presented an average standard length of 25.2 ± 2.2 cm and average weight of 328.82 ± 89.03 g. A total of 5,227 specimens of metazoan parasites were collected: 2,880 (55.1%) adult Diplectanum piscinarius (Monogenoidea: Diplectanidae) and 2,347 (44.9%) Austrodiplostomum compactum metacercariae (Digenea, Diplostomidae), both with 100% prevalence and mean abundance of 57.6 and 46.9, respectively. Parasite diversity was low (species richness = 2), with a Simpson index (D) equal to 0.505, and low values of Brillouin (HB = 0.687) and Margalef diversity (I = 0.117) indices. Berger-Parker's index of dominance (p = 0.551) indicated a slight dominance of the monogenean parasite D. piscinarius. There was a positive correlation, assessed by Pearson coefficient between parasite abundance of D. piscinarius and standard length (r = 0.43) and weight (r = 0.51) of hosts.

Ecologia da comunidade de metazoários parasitos do dourado, Coryphaena hippurus Linnaeus, 1758, (Osteichthyes: Coryphaenidae) do litoral do Estado do Rio de Janeiro, Brasil.

From April 2009 and July 2010, 68 specimens of common dolphinfish Coryphaena hippurus Linnaeus, 1758 (Osteichthyes: Coryphaenidae) collected from coast of the state of Rio de Janeiro (21-23° S, 41-45° W), were necropsied to study their infracommunities of metazoan parasites. Seventeen species of metazoan parasites were collected. All fish were parasitized by one or more metazoan. The digeneans were the majority of the specimens collected, with 90.7%. Dinurus tornatus (Rudolphi, 1819) was the dominant species with highest abundance, prevalence, frequency of dominance and mean relative dominance. The parasites species of C. hippurus showed a typical aggregate pattern of distribution. None metazoan parasites species showed correlation between total length and prevalence and parasite abundance. The metazoan parasite infracommunities C. hippurus showed dominance of endoparasites (digeneans) and no correlation between abundance and parasite species richness and the total length of the host.

Ecologia da comunidade de metazoários parasitos do dourado, Coryphaena hippurus Linnaeus, 1758, (Osteichthyes: Coryphaenidae) do litoral do Estado do Rio de Janeiro, Brasil.

From April 2009 and July 2010, 68 specimens of common dolphinfish Coryphaena hippurus Linnaeus, 1758 (Osteichthyes: Coryphaenidae) collected from coast of the state of Rio de Janeiro (21-23° S, 41-45° W), were necropsied to study their infracommunities of metazoan parasites. Seventeen species of metazoan parasites were collected. All fish were parasitized by one or more metazoan. The digeneans were the majority of the specimens collected, with 90.7%. Dinurus tornatus (Rudolphi, 1819) was the dominant species with highest abundance, prevalence, frequency of dominance and mean relative dominance. The parasites species of C. hippurus showed a typical aggregate pattern of distribution. None metazoan parasites species showed correlation between total length and prevalence and parasite abundance. The metazoan parasite infracommunities C. hippurus showed dominance of endoparasites (digeneans) and no correlation between abundance and parasite species richness and the total length of the host.

Metazoan parasites of Mandi-amarelo Pimelodus maculatus and of Jundiá Rhamdia quelen (Osteichthyes: Siluriformes) of Paraíba do Sul River, Volta Redonda, Rio de Janeiro

Forty-one specimens of mandi-amarelo Pimelodus maculatus Lacépède, 1803 (Siluriformes: Pimelodidae) and 54 specimens of jundiá Rhamdia quelen (Quoy & Gaimard, 1824) (Siluriformes: Heptapteridae) were collected from the Paraíba do Sul River, Volta Redonda, State of Rio de Janeiro, Brazil between November 2007 and October 2008. These fish underwent necropsy so their infracommunities of metazoan parasites could be studied. The same three species of parasites were collected in the two fish species studied. These were one monogenean, one nematode, and one hirudinean. Cucullanus pinnai (Travassos, Artiga, and Pereira, 1928) (Nematoda: Cucullanidae) and Aphanoblastella sp. (Monogenea: Dactylogyridae) were the dominant species with the highest prevalence in P. maculatus and R. quelen. The parasite species of P. maculatus and R. quelen showed an atypical over-dispersed pattern of distribution. No parasite species showed significant correlation between the body total length of the siluriform hosts and their prevalence and abundance. The parasite species richness showed a mean value of 0.87 ± 0.67 (0-2) and 0.57 ± 0.56 (0-2) in P. maculatus and R. quelen, respectively, and no correlation with the body total length.

Metazoan Parasite Infracommunities of the Toadfish Aphos porosus (Pisces: Batrachoidiformes) in Central Chile: How Variable Are They over Time?

The present study examines variation in parasite community characteristics of the toadfish Aphos porosus across sampling years. We analyzed and compared the species composition and numeric descriptors of the parasite infrapopulations and infracommunities in 101 fish collected during the springs of 1999, 2006, and 2007 from rocky intertidal ponds in El Tabo, central Chile. Parasites were found in 94.1% of specimens necropsied. In total 5,532 parasites were collected, representing 12 taxa, of which 7 were found in the 3 annual samples. The most prevalent and abundant parasites in the 3 sampling years were larval tetraphyllidean cestodes, followed by anisakid nematodes. The community descriptors of average richness and abundance varied significantly across the 3 yr, with the exception of parasite dominance. The parasite species composition and relative abundance showed low variability among years. A few parasite taxa (Anisakis sp., Pseudoterranova sp., tetraphyllideans, and Clestobothrium crassiceps) dominated the infracommunities of the toadfish; however, tetraphyllideans were the parasites that greatly varied over time. Consequently the parasite infracommunities of the toadfish were variable among years, with a low variation in parasite composition, but large differences in the aggregate descriptors.

Community ecology of metazoan parasites of the anchovy Anchoa tricolor (Osteichthyes: Engraulidae) from the coastal zone of the State of Rio de Janeiro, Brazil

Between October 2001 and March 2002, 103 specimens of A. tricolor from Angra dos Reis (23 degrees 01' S, 44 degrees 19' W), in the coastal zone of the State of Rio de Janeiro, Brazil, were analyzed in order to study their metazoan parasite infracommunities. Ten species of metazoan parasites were collected: 4 digeneans, 1 cestode, 1 acantocephalan, 2 nematodes, 1 copepod, and 1 hirudinean; 77.7% of the fishes were parasitized by one or more metazoan, with a mean of 3.5 +/- 6.2 parasite/fish. Digenean was the most dominant with 4 species that accounted for 53.2% of the total parasites collected; Ergasilus sp. was the most abundant species. Abundance and prevalence of Parahemiurus merus (Linton, 1910) were positively correlated with the total length of host. Relationships between total body length of fish and both total parasite abundance and mean parasite species richness were observed. Mean parasite diversity of species was correlated to host's total length, with significant differences found between male and female fishes. Two pairs of larval species showed significant positive association and covariation. The metazoan parasite infracommunities of A. tricolor presented dominance of larval endoparasites; correlation of parasite abundance, diversity, and species richness with host total length; and low number of parasite interspecific relationships. The parasite community of A. tricolor showed some similarities with the parasite community of another South American Atlantic engraulid.

Community ecology of metazoan parasites of the later juvenile common snook Centropomus undecimalis (Osteichthyes: Centropomidae) from the coastal zone of the state of Rio de Janeiro, Brazil

Between April and December 2000, seventy-nine specimens of Centropomus undecimalis from Angra dos Reis, coastal zone of the State of Rio de Janeiro (23 degrees 01'S, 44 degrees 19'W), Brazil, were necropsied to study their infracommunities of metazoan parasites. Nine species of metazoan parasites were collected: 1 digenean, 1 monogenean, 1 acantocephalan, 1 nematode, 4 copepods, and 1 isopod, and 96.2% of the fishes were parasitized by one or more metazoan, with mean of 85.3 +/- 122.9 parasite/fish. The digenean Acanthocollaritrema umbilicatum Travassos, Freitas & Bührnheim represented the majority of the parasites specimens collected, totaling, 94.7%. This species was the most abundant, prevalent, and dominant, showing positive correlation with the host's total body length and parasite abundance. The copepod species Acantholochus unisagittatus Tavares & Luque presented differences in abundance in relation to sex of host. The mean diversity in the infracommunities of C. undecimalis was H = 0.095 +/- 0.116, with no correlation with the host's total body length but correlated with the host's sex. No pair of parasite species showed positive or negative association or covariation. The dominance of digenean A. umbilicatum in the later juvenile common snook parasite community could be related with the predatory food habits of common snook and an apparent feeding transition period which might occasion great exposure to infective forms.

Community ecology of the metazoan parasites of white sea catfish, Netuma barba (Osteichthyes: Ariidae), from the coastal zone of the state of Rio De Janeiro, Brazil.

Between March 2000 and April 2001, 63 specimens of N. barba from Angra dos Reis, coastal zone of the State of Rio de Janeiro (23 degrees 0' S, 44 degrees 19' W), Brazil, were necropsied to study their infracommunities of metazoan parasites. Fifteen species of metazoan parasites were collected: 2 digeneans, 1 monogenean, 2 cestodes, 1 acantocephalan, 2 nematodes, 6 copepods, and 1 hirudinean. Ninety-six percent of the catfishes were parasitized by at least one metazoan parasite species. A total of 646 individual parasites was collected, with mean of 10.3 +/- 16.6 parasites/fish. The copepods were 37.5% of the total parasite specimens collected. Lepeophtheirus monacanthus was the most dominant species and the only species with abundance positively correlated with the host total length. Host sex did not influence parasite prevalence or mean abundance of any species. The mean diversity in the infracommunities of N. barba was H = 0.130 +/- 0.115 with no correlation with host's total length and without differences in relation to sex of the host. One pair of endoparasites (Dinosoma clupeola and Pseudoacanthostomum floridensis) showed positive association and covariation between their abundances and prevalences. The parasite community of N. barba from Rio de Janeiro can be defined as a complex of species with low prevalence and abundance and with scarcity of interspecific associations. However, because of both the presence of assemblages of sympatric ariid species as well as the spawning behavior characteristic of these fishes, additional comparative studies of the parasite component communities of ariids are necessaries to elucidate this pattern.

Ecologia da comunidade de metazoários parasitos da Maria-Luiza, Paralonchurus brasiliensis (Osteichthyes: Sciaenidae) do litoral do Estado do Rio de Janeiro, Brasil

Ninety-three specimens of banded croaker, Paralonchurus brasiliensis (Steindachner, 1875), collected from Pedra de Guaratiba (23º01’S, 43º38’W), coastal zone, State of Rio de Janeiro, Brazil, between September 2001 and March 2002, were necropsied to study their infracommunities of metazoan parasites. Fifteen species of metazoan parasites were collected. Paralonchurus brasiliensis is a new host record for 13 parasite species. The majority of fishes were parasitized by one or more metazoan. Larvae of Contracaecum sp. proved to be the dominant species, with highest prevalence and abundance values. The parasite species of P. brasiliensis showed the typical aggregated distribution pattern. Prevalence and abundance of Procamallanus (Spirocamallanus) pereirai was positively correlated with the host total length. Abundance of Contracaecum sp. was positively correlated with the host total length. The abundances of a pair of adult endoparasites (Aponurus laguncula - Procamallanus (S.) pereirai) showed negative covariation and one pair of larvae stages of endoparasites (Nybelinia sp. – Contracaecum sp.) showed positive covariations between their abundances. Paralonchurus brasiliensis showed a metazoan parasite community composed by generalist species, little ordered and with scarce quantitative evidences of interspecific associations. These patterns agree with those detected in preceding studies about the parasite communities of Neotropical sciaenid fishes.

Metazoan parasite infracommunities of Florida pompano (Trachinotus carolinus) from the coast of the Yucatan Peninsula, Mexico.

Metazoan parasite infracommunities of the Florida pompano (Trachinotus carolinus) were studied in terms of species composition, species richness, diversity, numerical dominance, and similarity. Seventy-five fishes were collected from 4 localities along the Yucatan Peninsula coast and 24 parasite species recovered. Most were digeneans (8 species) and nematodes (7). Other species were monogeneans (3). aspidogastreans (2), cestodes (1), acanthocephalans (1), and crustaceans (2). Only 4 species were common in at least I locality. Mean values for species richness, abundance, diversity, numerical dominance, and similarity in total (all species in the individual fish), gastrointestinal, and ectoparasite infracommunities were within ranges observed for most helminth infracommunities of marine fishes from temperate and tropical latitudes. These infracommunities had low species richness, abundance, diversity, and predictability (except ectoparasite infracommunities) and high dominance. Within the predictable element (common species), the specialist monogenean Pseudobicotylophora atlantica was the main reason for the increase in predictability because it was the only common species at all 4 localities. Host feeding habits, the distribution of intermediate hosts and infective stages, the local species pool, and a phylogenetic component seem to be determining the characteristics of these metazoan parasite infracommunities.

Metazoan Parasite Infracommunities of Florida Pompano (Trachinotus carolinus) from the Coast of the Yucatan Peninsula, Mexico

Metazoan Parasite Infracommunities of Florida Pompano (Trachinotus carolinus) from the Coast of the Yucatan Peninsula, Mexico

Metazoan Parasite Infracommunities of Menticirrhus (Teleostei: Sciaenidae): An Amphi-Oceanic Approximation

Metazoan infracommunities of Menticirrhus ophicephalus from the south Pacific Ocean and M. americanus from the south Atlantic Ocean were analyzed and compared. The metazoan parasite infracommunities of M. ophicephalus showed higher values of total number of parasite individuals/host and parasite species richness than that of M americanus, but M. americanus showed higher values of parasite species diversity. The infracommunities of ectoparasites showed higher values of total number of parasite individuals/host, richness, and frequency of dominance in M. ophicephalus; in contrast, endoparasites showed higher values of these infracommunity descriptors in M. americanus. Ecological analysis of the marine fish parasite infracommunities using the total number of components (ecto- and endoparasites) could hide some patterns of the community structure; thus, a separate analysis of only ecto- or endoparasites is recommended. Additional studies monitoring the influence of the "upwelling ecosystem" and the El Niño-Southern oscillation event on the fish parasite communities are necessary to determine the real influence of these ecological disturbances on the structure of fish parasite communities in South America.

Ecologia da comunidade de metazoários parasitos da anchova Pomatomus saltator (Linnaeus) (Osteichthyes, Pomatomidae) do litoral do estado do Rio de Janeiro, Brasil

Fifty-five specimens of bluefish, Pomatomus saltator (Linnaeus, 1766), collected at Cabo Frio (23oS, 42oW), State of Rio de Janeiro, Brazil, between March 1995 and November 1995, were necropsied to study their infracommunities of metazoan parasites. Sixteen species of metazoan parasites were collected. All fish were parasitized by one or more metazoan. The digeneans were the majority of the parasite specimens collected, with 44.2%, followed by the helminth larval stages (cestodes, acanthocephalans, and nematodes) with 40.6%. The Simpson index for all parasite species was 0.097, indicating lack of dominance by any species in the parasite community. Microcotyle pomatomi Goto, 1899 showed the higher frequency of dominance and mean relative dominance. The parasite species of P. saltator showed the typical overdispersed pattern of distribution. The majority of parasite species showed positive correlation between the host's total body length and prevalence or parasite abundance. Two species, Brachyphallus parvus (Manter, 1947) and Phocanema sp. have differences in their prevalences and abundances in relation to sex of the hosts. The mean diversity in the infracommunities of P. saltator was H'=1.243±0.521, with correlation with the host's total length and without difference between male and female fish. The components of the parasite community of P. saltator showed overall positive association. All pairs confonned by ectoparasites and adult endoparasites species showed signiticant positive association or covariation between their abundances. Five pairs of endoparasites larval species showed positive association or covariation, and two pairs, showed negative association and covariation. The parasite community of P. saltator was dominated by species with high prevalence values, that composed a high number of associations, thus, is considered closer to the interactive type.