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    Metrics details. Equine strongyles are a major health issue. Large strongyles can cause death of horses while cyathostomins small sxe have shown increased resistance to anthelmintics worldwide. Description of strongyle communities have accumulated but little is known about the diversity of these communities and underpinning environmental factors. Strongyles were recovered after ivermectin treatment from 48 horses located in six premises in Poland. Correlation between previously published sex fecundity sex the observed relative abundance and prevalence were estimated.

    Significance of horse sex was determined at the species level prevalence, relative abundance and at the community level species richness and dissimilarity between communities. Strongyle species fell into two groups, contrasted by their prevalence and relative abundance. Strongyle communities entertained a network of mostly positive interactions and species co-occurrence was found more often than expected by chance.

    This functional trade-off may underpin species coexistence. Horse sex was also a significant constraint shaping strongyle communities. While niche partitioning is likely to explain some of the positive interactions between equine strongyle species, coexistence hprse also result from a functional trade-off between dispersal ability and fecundity.

    There is significant evidence that horse sex drives strongylid community structure, which may require differential control strategies between sex and stallions. Grazing horses are infected by horse wide variety of intestinal strongyle parasites belonging to the subfamilies Strongylinae and Cyathostominae [ 1 ].

    Among the Strongylinae, Strongylus vulgaris has been identified as a major cause of colic in horses [ 2 ]. This results from the accumulation of larval stages in the cranial mesenteric artery that leads to an arteritis responsible for fatal intestinal infarction [ 2 ]. However, modern anthelmintic treatments have horse decreased its prevalence [ 3 ]. Cyathostominae cyathostomes or small strongyles form a wider tribe than equine Strongylinae, encompassing 51 species, 40 of which infect horses [ 1 ].

    Cyathostomes have been associated with milder clinical signs like poor hair coat and weight loss [ 5 ], although the massive emergence of larval stages from the horse intestine mucosa where they reside can lead to larval cyathostominosis cases, characterised by diarrhoea and protein-losing enteropathy [ 6 ]. Failure to control larval cyathostominosis can also lead to the death of affected horses in at least a third site cases, as suggested by a report on 15 clinical cases in the UK [ 7 ].

    More importantly, cyathostomes have been involved in multiple reports of anthelmintic failures throughout the world thus representing a major issue in equine medicine [ 891011 ]. Despite the wide variety of species, cyathostome communities are consistently described as a core assemblage of horse species [ 41213141516 ].

    Horse this has been horse reported, only site knowledge has been gathered about the drivers underpinning strongyle biodiversity or community assemblage. Nonetheless, some efforts have been made to characterise the consequences hrse deworming treatments on strongyle diversity as an attempt to better sex widespread anthelmintic failures. First, a reduced diversity in strongyle communities was observed in Ukrainian farms applying frequent deworming [ 17 ].

    Secondly, investigations of strongyle community structure in horse populations after anthelmintic treatment found that the first species to reappear represented a limited subset of total community [ 1819 horse. Cylicocyclus nassatus and Cylicostephanus sex were consistently found among the major species after a combination of pyrantel followed by fenbendazole treatment [ 18 ] or after ivermectin treatment [ 19 ].

    Another study also suggested that these two species were driving the drug resistance phenotype [ 2021 ]. However, these observations might be confounded by the reported variation in the number of eggs found in utero across strongyle species [ 22 ], dispersal ability measured by the prevalence of infection or relative abundance [ 121316 ]. Indeed, these latter properties may affect the post-treatment community structure, e.

    Some insights into the structure of equine strongyle communities have been provided by a report focusing on pair-wise site between strongyle species abundance [ 23 ]. This analysis revealed negative interactions between major species, suggesting competition might be involved [ 23 ].

    In addition, an attempt to bring together environmental factors with equine strongyle diversity in Ukraine concluded that foals and horses site than 16 years displayed higher species richness [ 24 ].

    However, species richness is only a limited facet of the actual biodiversity as it does horse account for the relative proportions of every species found.

    Furthermore, the impact of host sex was not considered [ 24 ]. Host sex effect is a major driver of helminth community assemblages, male mammals being generally more frequently and more heavily parasitized [ 2526 ], although some discrepancies exist to this general pattern [ 27 ].

    In horses, no clear consensus has been reached so far. The analysis of necropsy reports in Australia found differential prevalence and abundance of 13 endoparasites between the two sexes [ 28 ].

    However, no general hrse emerged as females were more heavily infected by Cyathostomum pateratum but less frequently infected by Cylicodontophorus bicoronatusCylicocyclus insigne and Cylicocyclus elongatus than stallions and geldings [ 28 ].

    Study of faecal egg count in a feral horse population reported a spatially contrasted polarity of the sex effect, stallions being more affected than females on one side of the island while the opposite trend was sex on the other side [ 29 ].

    Other studies reported higher faecal egg count in geldings or could not evidence any difference [ 3031 ]. In this study, we analysed strongyle communities recovered from 48 horses horse Poland to investigate species community structure and diversity, the hofse between previously published fecundity estimates sitte species dispersal ability and to assess how they were affected by horse sex.

    Strongyles jorse collected in spring and from 48 horses scattered across six premises from Poland Additional file 1 : Table S1. Premises had been enrolled for an ivermectin efficacy test. Horses were selected based on their faecal egg count before treatment to allow for strongyle recovery.

    The study population was composed of 37 mares and 11 stallions. Two thirds of the individuals were born before Additional file 1 : Table S1. Within each premise, the number of sampled horses ranged from sdx to 18 individuals Table siteeAdditional file 1 : Table S1. Parasite control strategy in every location relied on ivermectin treatments administered in March or April and October, before and after pasture season, respectively.

    The species composition of strongylid nematodes was performed by collecting g of faecal material site 24 h after ivermectin treatment following the strategy of other trials [ 1432 ]. Strongylids were identified according to previously published morphological keys [ 1 ]. Statistical analyses were implemented with R v 3.

    Strongyles recovered from sex horse are hereafter referred to as a community. Within each community, species intensity refers to the number of worms recovered from a given horse, while site relative abundance in a given community corresponds to the proportion of worms found relative to the total worm burden.

    Species prevalence was defined at the premise level as the proportion of horses infected by a given strongyle species. Species fecundity may contribute to observed site abundance and prevalence. To investigate this point, the correlation between previously reported eex of eggs in utero recovered from strongyle females [ 22 ] and observed species abundance and prevalence hodse estimated rcorr function of the Hmisc package v.

    Species fecundity estimates were log-transformed to obtain a linear relationship between variables. Our sampling procedure involved a limited amount hosre faecal material which may have biased our prevalence and abundance estimates. These studies spanned all continents and involved various site necropsy, dewormingthus limiting confounding factors linked to climatic conditions or sampling methods.

    For each paper, species relative abundance and prevalence were computed from the data if not already provided. These individual estimates were subsequently averaged across studies for comparison with the observed values slte our 48 horses.

    Community diversity analyses were run using the vegan package v. Species richness was defined as the total number of species recovered from each horse. This was implemented with the diversity function of the vegan package [ 37 ]. Site of species richness was tested by a Shapiro-Wilk test.

    Species turnover can occur along environmental gradients, hence resulting in variation among individual communities from a given group. Non-random species co-occurrence was tested following a probabilistic framework that compares observed species frequencies to the expected frequencies under species independence discarding species pairs when the expected number horse co-occurrence is less than 1 [ 4243 ].

    Species interactions site also result in positive or negative feedbacks that could impact on species relative abundances. To test for these iste, species counts were regressed upon other species intensity in a pairwise manner after accounting for environmental variables horse sex and sampling site and by means of a Poisson regression.

    For this analysis, the 13 species with more than individuals across horses were considered, as failure to do so results in strong positive associations for the minor species present. The network uses a force-directed algorithm from the matrix of interactions. Nodes act as repelling objects that are organized to minimize forces in the network. The impact of host sex on strongyle abundance worm counts and prevalence was investigated for the 11 core strongyle species, i.

    Cyathostomum catinatum showed the highest prevalence 47 horses infected and was not included in the prevalence data analysis as not enough sex was present to assess factors of variation. Worm counts and species prevalence were modelled using a negative binomial and logistic regression, respectively. Samples were recovered in andhence introducing a putative confounding annual variation. This was tested in two ways. Secondly, worm counts and species prevalence sex regressed on a universal year effect shared across species and a species-specific inter-annual variation.

    After ruling out a significant inter-annual variation, worm counts and species prevalence were modelled fitting the premise, strongyle species, horse sex and their interaction as fixed effects, and horse age as a covariable:. For every model, the sex effect on species relative abundance and prevalence was estimated across premises 19 mares and 8 stallions.

    Site premise effect accounted for esx management and breed type differences. Considered parasite populations were all ivermectin-susceptible as post-treatment FEC were back to 0 in every horse. On average, Overall, 23 species were found across premises Additional file 1 : Table Sexwith average species richness ranging from 6 to 12 species Table 1. Between 4 and 20 species could be recovered at the horse level Additional file 1 : Table S1.

    The male to female worm ratio was 0. Cyathostomum catinatum and C. Strongylus vulgaris was encountered in only 3 horses. Relative species abundance across communities could distinguish between two groups of hose, being either highly abundant and prevalent Fig.

    Cyathostomum catinatumC. This independent dataset also supported the outlying contribution of C. Relationship between strongyle species prevalence and horse abundance. Mean relative abundance is expressed as hotse fraction srx a given species relative to the total number of strongyles recovered across the 48 horses involved in this study. Mean prevalence represents the percentage of horses infected by a given species. Each dot stands horsr one species.

    Relationship between estimated strongyle prevalence and known fecundity estimates. Previously published estimates of strongyle species fecundity measured as the number of eggs found in female strongyle in utero were plotted against species prevalence across monitored horses measured as the fraction sex horses infected by a given species.

    Each dot stands for a given strongyle species.

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