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All issues Volume 32 (2025) Parasite, 32 (2025) 47 Full HTML
Open Access
Issue
Parasite
Volume 32, 2025
Article Number 47
Number of page(s) 14
DOI https://doi.org/10.1051/parasite/2025046
Published online 29 July 2025

© A. Tripathi et al., published by EDP Sciences, 2025

Licence Creative CommonsThis is an Open Access article distributed under the terms of the Creative Commons Attribution License (https://creativecommons.org/licenses/by/4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

Introduction

Cichlidogyrus Paperna, 1960 (Monogenea: Dactylogyridae) is the most species-rich African freshwater monogenean genus [37, 64]. It has 141 valid species [93] naturally parasitising primarily African cichlids (Cichlidae) and a few representatives of Cyprinodontidae Wagner, 1828 (Cyprinodontiformes) and Nandidae Bleeker, 1852 (Anabantiformes) [12, 20]. Some of these species have been identified as potentially pathogenic to fish, especially in aquaculture stocks [31, 56, 69]. Cichlidogyrus species, with few exceptions, are quite host-specific ([64], but also see [40]). An exception is Cichlidogyrus tilapiae Paperna, 1960. Since its first description from the Nile Tilapia, Oreochromis niloticus (Linnaeus 1758) (Cichliformes: Cichlidae) in Israel, C. tilapiae has been recorded in 31 different fish species in 27 countries across five continents, including Asia, Africa, North America, South America, and Australia (Table 1).

Table 1

Global distribution of Cichlidogyrus tilapiae Paperna, 1960 for 27 countries and 31 host fishes.

Pindani, Chindongo (Pseudotropheus) socolofi (Johnson, 1974) (Cichliformes, Cichlidae) is native to Lake Malawi in Africa [18, 46], and is available in two colour variants – normal (Blue pindani) and albino (White pindani) [33]. Despite the importance of C. socolofi in the ornamental fish market [74], little is known about its parasite fauna. To our knowledge, only one study on parasitic infections of C. socolofi exists [11]. These researchers studied the parasites of cichlids imported via the aquarium trade in Türkiye and recorded the protozoan parasite Trichodina pediculus Ehrenberg, 1831.

This study aimed to establish whether C. socolofi is infected by monogenean parasites and, if so, whether they were co-translocated into India via the ornamental fish trade. We demonstrate the presence of C. tilapiae in post-quarantine populations of C. socolofi collected from Indian aquarium markets. This was accomplished first by morphological characterisation (structure and measurements of the sclerotised parts of the haptor and reproductive organs) and subsequently by molecular characterisation (Sanger sequencing of 18S rRNA gene-ITS1 region and 28S rRNA genes). This paper is part of a series on exotic and/or invasive monogenean parasites imported into India via the ornamental trade [81–87].

Materials and methods

Ethics

This study was approved by the institutional ethics committee of the University of Lucknow under the protocol numbers LU/AEC/ZOO/2019 and 19/I/2024/IAEC/LU.

Sample collection and examination

Between January 2020 and December 2022, 22 specimens of C. socolofi (total weight: 3.12–6.50 g; and total length: 4.5–8.0 cm) (Fig. 1) were collected from aquarium shops in Lucknow, New Delhi, and Kolkata, India. Fish were shipped to the laboratory the same day after they were packaged in polybags containing water and pure oxygen. Individual fish were euthanised with an overdose of tricaine methanesulfonate (MS-222 @ 150 mg/L; Sigma Aldrich Co., St. Louis, MO, USA), followed by exsanguination by the removal of gill arches. Half of the gill arches were initially fixed in hot (60 °C) distilled water to relax and heat-kill the specimens before they were transferred to 4% formalin for microscopy following Kritsky [36]. The other half was preserved in 95% ethanol for genetic analysis. Some of the gill arches were examined fresh with live worms. Monogeneans were later isolated from these gills using fine dissecting needles under a stereomicroscope (Leica Microsystems, Wetzlar, Germany). Fish specimens were identified morphologically with the help of the ICAR-National Bureau of Fish Genetic Resources (ICAR-NBFGR), a premier Indian institute on fish taxonomy, biology, and genomics.

thumbnail Figure 1

Freshly dead specimens of Chindongo socolofi (Johnson, 1974) examined for the present study. A. Blue pindani, B. White pindani. Photograph by Chawan Matey.

Morphological analysis

Formalin-fixed worms were stained with either Gomori’s trichrome or Borax carmine and mounted in DPX (dibutylphthalate polystyrene xylene) for observing internal anatomy (permanent mounts); others were mounted in glycerine jelly or Hoyer’s medium for the study of sclerotised parts of the haptor and reproductive organs (temporary mounts). Additionally, some ethanol-preserved worms were treated for 20–30 min at 55 °C with 1.0 μL of digestion buffer (0.1 μl of solid tissue buffer and 0.9 μL proteinase K) (Quick DNATM Miniprep Plus Kit, ZYMO Research, Irvine, CA, USA) to digest the tissues surrounding their sclerotised parts.

The morphology of the sclerotised parts was examined under a light microscope (Leica DM4B) at a magnification of 100×, using an oil immersion lens with phase-contrast (PHA-CO) and differential interference contrast illumination. Photographs and measurements (in micrometres) were obtained using a digital camera (Leica DFC7000 T) and imaging analysis software (LAS X; Leica Microsystems Ltd.) attached to the light microscope. A composite line drawing plate was made from multiple parasite specimens using an Olympus BX-51 microscope drawing tube. Species were identified based on the morphological characters described in previous studies [14, 15, 35, 44, 57, 64]. The terminology and measurement of these characters followed Rahmouni et al. [70]. The prevalence and mean intensity of infection were calculated according to Bush et al. [8].

DNA extraction and amplification

Representative samples of ethanol-preserved specimens were morphologically identified as conspecific to the temporary and permanent mounts before being pooled in two groups (n = 3) by two collection sites (Lucknow and New Delhi) for gDNA isolation using a DNA extraction kit (Extracta DNA Prep for PCR-Tissue, Quantabio, Beverly, MA, USA), according to the manufacturer’s instructions. Partial fragments of 18S ribosomal RNA genes (18S) and internal transcribed spacer 1 (ITS1) clusters were amplified with the primers s1 [78] and ir8 [75]. Meanwhile, those of 28S ribosomal RNA genes were amplified with the primers c1 and d2 [23].

Polymerase chain reactions were performed in an automated thermal cycler (Himedia Laboratories, Thane, MH, India) with reaction mixtures (final volume 20 μL) containing 4 μL of distilled water, 10 μL of 2× PCR TaqMixture (Himedia Laboratories), 1 μL of 10 pmol/microliter of each primer, and 4 μL of DNA template. The amplification profile for the 18S rRNA gene-ITS1 region was as follows: initial denaturation at 95 °C for 3 min, then 35 cycles of denaturation at 95 °C for 30 s, annealing at 50 °C for 30 s, and extension at 72 °C for 1 min, with a final extension at 72 °C for 7 min. The amplification profile for the 28S ribosomal RNA gene followed Šimková et al. [76]. The size of the PCR products (2 μL) was analysed by electrophoresis in 1.2% agarose gel prepared in 1× TAE buffer, prestained with 0.1 μL/mL 10,000× Sybr Safe in dimethyl sulfoxide (Invitrogen, Waltham, MA, USA), at 90 V for 30 min, and visualised and documented on a Bio-Print gel documentation imaging system (Vilber Lourmat, Collégien, France).

Sequence analysis

The PCR products were purified (on 1.5% agarose using a QIAquick PCR Purification Kit; QIAGEN, Germantown, MA, USA) and Sanger sequenced (on an ABI 3730xL automated sequencer; Applied Biosystems, Foster City, CA, USA) with PCR primers by Eurofins Genomics (Bengaluru, KA, India). SnapGene version 5.3 (https://www.snapgene.com) was used to manually quality-trim the successfully sequenced amplicons. Consensus sequences (18S-ITS1, 942 bp; 28S, 660 bp and 848 bp) were generated using the BioEdit Program [22].

Sequences, together with all sequences from the same markers and species retrieved from NCBI GenBank (Tables 2 and 3) were aligned using ClustalW [24] implemented in MEGA v.7 [38]. To obtain equal lengths for sequence analysis, they were trimmed to 687 bp (18S-ITS1) and 631 bp (28S). A median-joining network [3] was inferred for each marker using PopART [39].

Table 2

Information on Cichlidogyrus tilapiae Paperna, 1960, including hosts, localities, and GenBank accession numbers of their 18S+ITS1 rRNA gene sequences (as retrieved from the NCBI database on December 04, 2024).

Table 3

Information on Cichlidogyrus tilapiae Paperna, 1960, including hosts, localities, and GenBank accession numbers of their 28S rRNA gene sequences (as retrieved from the NCBI database on December 04, 2024).

As additional assessment for species-level identification, intraspecific genetic differences (also known as genetic distances) among different geographical isolates of C. tilapiae were computed from the same dataset. This was done using the Kimura two-parameter (K2P) model [34] of nucleotide substitution in MEGA 11 [79], with gaps treated as complete deletions.

Results

Thirty individuals of a single monogenean species – namely Cichlidogyrus tilapiae Paperna, 1960 – were collected from the gills of five specimens of C. socolofi, with a low prevalence (22.72%) and low infection intensities (7–13 worms/fish). Brief morphological and molecular data of this species are presented below.

Taxonomic summary

Phylum Platyhelminthes Minot, 1876

Superclass Neodermata Ehlers, 1985

Class Monogenea van Beneden, 1858

Family Dactylogyridae Yamaguti, 1963

Genus Cichlidogyrus Paperna, 1960

Cichlidogyrus tilapiae Paperna, 1960 (Figs. 2 and 3)

Type host and locality: Nile tilapia Oreochromis niloticus (Linnaeus, 1758) (Cichliformes: Cichlidae); Israel (Paperna 1960).

thumbnail Figure 2

Photomontage of light microscopy and phase contrast (PHACO) images of Cichlidogyrus tilapiae Paperna, 1960 from Chindongo socolofi (Johnson, 1974). A. anchor-bar complex and hooks, B and C. different configurations of male copulatory organ, D. vagina. Photograph by Amit Tripathi.

thumbnail Figure 3

Line drawings of sclerotised structures of Cichlidogyrus tilapiae Paperna, 1960 from Chindongo socolofi (Johnson, 1974). A. ventral anchor, B. ventral bar, C. dorsal anchor, D. dorsal bar, E. male copulatory organ, F. hook (pairs i–vii). Scale bar = 20 μm. Figure by Amit Tripathi.

Present host: Chindongo socolofi (Cichliformes, Cichlidae).

Present material and collection date: Aquarium shops in Lucknow (26.8467° N, 80.9462° E), New Delhi (28.6139° N, 77.2090° E), and Kolkata (22.5726° N, 88.3639° E), India; January 2020–December 2022.

Site of infection: Gills.

Infection parameters: Prevalence: 22.72% (5 out of 22 C. socolofi examined); Mean infection intensity: 6.2 ± 3.86 (7–13; n = 5).

Museum material: Five voucher specimens stained with Gomori’s trichrome or Borax carmine and mounted on glass slides in DPX (Smithsonian Institution, USA; USNM 1757684-1757688).

GenBank deposition: 18S-ITS1: 942 bp (MZ266637); 28S: 660 bp (MZ265190), 848 bp (PQ675652).

Morphological data

Cichlidogyrus is distinguished by a vas deferens that does not encircle the intestinal caecum, two pairs of anchors (one dorsal and one ventral), two transversal bars (a dorsal bar with two typical auricles and a V-shaped ventral bar), seven pairs of hooks, a sclerotised or non-sclerotised vagina and a sclerotised male copulatory organ comprising a male copulatory tube and (often but not always) an accessory piece [57, 88].

Our specimens presented nearly identical morphological features of sclerotised parts (both haptoral and reproductive ones) indicated in the original description [55] and subsequent redescriptions or illustrated records [14, 15, 35, 44] of C. tilapiae (Figs. 2 and 3) (Table 4). Only two minor discrepancies were observed in the morphometry of the haptoral armaments. First, our specimens had a slightly longer and deeper outer root of the dorsal anchor (4–6 μm), compared to their conspecifics. Second, variations were observed in the ranges of measurements of the ventral bar. Paperna [57], for example, measured the length of the ventral bar to be 34–98 μm, whereas Douëllou [14] measured it to be 26–33 μm (as we did), and Kritsky and Thatcher [35], Ergens [15], and Maneepitaksanti and Nagasawa [44] measured it to be 50–65 μm.

Table 4

Comparative measurements (in μm) of reproductive organs and haptoral armaments of Cichlidogyrus tilapiae Paperna, 1960 from India (present study) and other geographical locations.

We were also able to locate the vagina in a single live specimen, which had gone unnoticed in previous studies on C. tilapiae. It resembled a short unsclerotised (muscular?) tube with a funnel-like opening at one end (Fig. 3D). We lost it quickly however, when the vitellaria burst out of the parasite body, killing it. Therefore, our identification of the vagina may not be conclusive and should be reconfirmed. Cichlidogyrus tilapiae has previously been adequately described/redescribed and, thus, does not need to be formally redescribed here.

Molecular data

The partial 18S rRNA gene-ITS1 region (942 bp) and 28S rRNA genes (660 bp and 848 bp) were sequenced from two pools of C. tilapiae specimens collected from C. socolofi in aquarium shops in India. Comparative analysis of these sequences against the NCBI GenBank non-redundant database using the “megablast” algorithm (https://blast.ncbi.nlm.nih.gov/Blast.cgi) revealed “near perfect” matches for 28S rRNA (659/660 bp; 99.85% similar identity with a query coverage of 100%) and 18S rRNA-ITS1 (935/938 bp; 99.68% similar identity with a query coverage of 99%) to C. tilapiae from Paratilapia polleni Bleeker, 1868 (Cichliformes, Cichlidae) in Madagascar deposited in GenBank under the accession numbers MH767412 (28S) and MH767400 (18S-ITS1), respectively [77]. These findings suggest their conspecificity (Tables 5 and 6).

Table 5

Intraspecific genetic distances (Kimura 2-parameter model with partial deletion option) and variations between our samples and conspecific references (most similar BLAST hits) of Cichlidogyrus tilapiae Paperna, 1960 based on 18S rRNA gene-ITS1 region.

Table 6

Intraspecific genetic distances (Kimura 2-parameter model with partial deletion option) and variations between our samples and conspecific references (most similar BLAST hits) of Cichlidogyrus tilapiae Paperna, 1960 based on 28S rRNA gene.

The intraspecific genetic distances for 28S rRNA and 18S rRNA genes between four geographic isolates of C. tilapiae from different hosts and geographical locations were determined at 0%, indicating their conspecificity (Tables 5 and 6). The genetic distance for the ITS1 sequence, another marker with higher variability, was also determined between 0 and 0.002% (Table 5).

The haplotype network indicated that Indian haplotypes, for both markers, were widespread, and shared with conspecifics from both native and introduced populations (Fig. 4).

thumbnail Figure 4

Median-joining haplotype networks based on a 687 bp fragment of small subunit rDNA and the first Internal Transcribed Spacer (left) and 631 bp of large subunit rDNA (right) from the newly sequenced individuals of Cichlidogyrus tilapiae from India, aligned with all previously published sequences from this species. Genotypes are represented by circles, with the size of the circle correlating with the number of isolates displaying the respective genotype. Colours denote the countries of sampling localities. Genotypes are connected by lines indicating the number of mutations between them.

Discussion

This paper is the first to document the presence of a member of Cichlidogyrus in India. In addition, C. socolofi has been identified as a new host species for C. tilapiae. This is also the first time a species of Cichlidogyrus is formally reported from a Malawi cichlid; members of the genus are known to occur on the lake’s cichlids but were mentioned without species-level identification [6]. Although there are a few variations in morphometrical data between different geographic isolates of C. tilapiae, we do not consider these differences to merit species-level separation. These variations may be attributable to differences in the host species [55], environmental factors [7], developmental stages [85, 90], individual variations within the species, or even the different fixation [16] or measuring methods employed thus far. For instance, just as we did, Douëllou [14] measured only one branch of the ventral bar following established norms of measurements for Cichlidogyrus [70]. Meanwhile, Kritsky and Thatcher [35] and Ergens [15] measured the total length in a “straight line extending between the two most distant parts”. Unfortunately, Paperna [57] and Maneepitaksanti and Nagasawa [44] did not specify their measurement methods.

While many publications have recorded only the occurrence of C. tilapiae without providing any morphometric data, those that have provided such data have shown a few variations in the sclerotised parts. For example, Paperna [57] found that both pairs of anchors were “about the same length”. However, all subsequent investigations have clearly shown that dorsal anchors are slightly larger than the ventral anchors. Paperna [57] also described and illustrated an accessory piece that terminated in a “bent bifurcated tip”, but this bifurcation has not been observed in any other studies. Furthermore, Ergens [15] and Douëllou [14] noted a small “groove on the base of dorsal anchor”, that no other researchers have reported. Ergens [15] also illustrated a small sliver-like structure on the outer roots of the ventral anchor, which has not been described or illustrated by other researchers.

Nonetheless, the distinctive morphology of the male copulatory organ, which lacks a heel and has a hook-shaped terminal end of the accessory piece, is consistent enough in all illustrations of C. tilapiae to be considered the most reliable diagnostic trait for identifying this species. This aligns with the notion that identification of Cichlidogyrus species is primarily based on the morphology of the reproductive hard parts [89].

Curiously, the 18S rRNA gene-ITS1 region and 28S rRNA gene sequences of C. tilapiae found in India differed from their conspecific references by only 3 bp and 1 bp, respectively (see above). Different phenotypes of C. tilapiae did not cluster monophyletically in the recent morphology-based phylogeny [51]. Therefore, we speculate that either C. tilapiae comprises a species complex of morphologically variable but closely related lineages [66] or that there are geographical variants of a single species.

The haplotype networks (Fig. 4) indicate that the haplotype of C. tilapiae found in India occurs widely throughout native and introduced host and parasite populations. Other markers than the ones used here, for example a sequence fragment of the cytochrome c oxidase subunit 1 gene, allow higher resolution distinction between populations of C. tilapiae [30] and may allow the identification of native and (co-)introduced strains of cichlid parasites [21].

Indian scenario

Since nothing is known about the monogenean fauna of C. socolofi in the wild, we cannot ascertain whether it is a natural host of C. tilapiae or whether it acquired it from other cichlids cohabiting in aquarium conditions. However, it is highly likely that C. socolofi is a regular host for C. tilapiae because the latter was consistently recovered over space (Lucknow, New Delhi, and Kolkata) and time (January 2020–December 2022). Chindongo socolofi is currently maintaining its self-sustaining populations in the country’s aquacultural ponds and has yet to be recorded in the wild. The potential negative impact of C. socolofi on India’s environment and/or economy will therefore depend on its ability to successfully establish, dominate, and expand in Indian waters. It has previously been hypothesised that the invasion success of a fish is linked to, amongst other things, favourable environmental conditions in the new habitat that are comparable to those in its native ranges (climate match theory; [1, 25]), and to the enemy release hypothesis [80]. Chindongo socolofi may be considered a potentially invasive fish species in this context because the climatic conditions in India, particularly in South India, are similar to those found in the native range of C. socolofi (southeastern Africa), including a tropical climate and a temperature range of 24–26 °C [18].

Should C. socolofi become invasive in Indian waters, the low host specificity of C. tilapiae (see above), combined with the native fish species’ lack of protective immunity against exotic parasites [72], could pose a serious biological invasion challenge. It is worth noting that C. tilapiae has already demonstrated its ability to switch from introduced cichlids to native hosts in destination environments, such as Vieja fenestrata (Günther, 1860) (Cichliformes, Cichlidae) (syn. = Paraneetroplus fenestratus) in Mexico [19], and Coptodon tholloni (Sauvage 1884) (Cichliformes, Cichlidae) in the Lower Congo Basin [29], and even non-cichlid hosts: Pachypanchax omalonotus (Duméril, 1861) (Cyprinodontiformes: Aplocheilidae) in Madagascar [77]. In fact, tilapia-infecting monogeneans have been proposed as the most ubiquitous tropical freshwater fish parasites globally, with C. tilapiae being one of the species most frequently reported as co-introduced with translocated tilapias [73]. However, to the best of our knowledge, this is the first report of this parasite from the ornamental fish trade.

The presence of C. tilapiae on C. socolofi highlights an additional challenge in India namely, illegal ornamental fish trafficking. The “Guidelines for import of ornamental fishes into India” [53] includes an “indicative list” of 92 exotic ornamental fish species that the Government of India has approved for import. Although C. socolofi is not on this list, it is widely available in Indian domestic trade ([67], this report).

Clearly, the fish were acquired illegally via international smuggling. Unfortunately, the “Guidelines” makes no clear or implicit declaration prohibiting the import of ornamental fish that are not on the “indicative list”, nor does it suggest that violators will face prosecution or even a fine. It simply states that “the import permit shall be cancelled forthwith and all the specimens imported destroyed without any notice to or permission of the importer”.

Given that the issue at hand involves not only fish trafficking but also the trafficking of accompanying (unidentified and often overlooked) parasites, merely cancelling import permits is a minor step toward protecting biodiversity and deterring traffickers. In reality, illegally imported exotic species are more likely to introduce parasites and diseases into the country because they bypass the import risk analysis and quarantine procedures of the importing country. Therefore, we recommend that the sale of a non-permitted ornamental fish species be treated as a criminal offence comparable to wildlife smuggling and implementing heavy penalties for this crime.

Acknowledgments

AT thanks Christoph Hahn (University of Graz, Austria) for a useful discussion on a previous version of the manuscript.

Funding

This work was fully supported by a project grant from the Science and Engineering Research Board (SERB), Government of India to AT (SERB–EMR/2017/003232). CM acknowledges the Ministry of Tribal Affairs, Government of India for providing her a PhD fellowship (202223-NFST-ARU-00531).

Conflicts of interest

The authors declared that they have no potential conflicts of interest with respect to the research, authorship, and/or publication of this article.

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Cite this article as: Tripathi A, Matey C, Pariselle A & Vanhove MPM. 2025. Monogeneans on exotic Indian freshwater fish. 8. Co-translocation of Cichlidogyrus tilapiae (Monogenea, Dactylogyridae) with pindani Chindongo socolofi (Cichliformes, Cichlidae): first report of this parasite genus in India within aquarium trade facilities. Parasite 32, 47. https://doi.org/10.1051/parasite/2025046.

All Tables

Table 1

Global distribution of Cichlidogyrus tilapiae Paperna, 1960 for 27 countries and 31 host fishes.

In the text
Table 2

Information on Cichlidogyrus tilapiae Paperna, 1960, including hosts, localities, and GenBank accession numbers of their 18S+ITS1 rRNA gene sequences (as retrieved from the NCBI database on December 04, 2024).

In the text
Table 3

Information on Cichlidogyrus tilapiae Paperna, 1960, including hosts, localities, and GenBank accession numbers of their 28S rRNA gene sequences (as retrieved from the NCBI database on December 04, 2024).

In the text
Table 4

Comparative measurements (in μm) of reproductive organs and haptoral armaments of Cichlidogyrus tilapiae Paperna, 1960 from India (present study) and other geographical locations.

In the text
Table 5

Intraspecific genetic distances (Kimura 2-parameter model with partial deletion option) and variations between our samples and conspecific references (most similar BLAST hits) of Cichlidogyrus tilapiae Paperna, 1960 based on 18S rRNA gene-ITS1 region.

In the text
Table 6

Intraspecific genetic distances (Kimura 2-parameter model with partial deletion option) and variations between our samples and conspecific references (most similar BLAST hits) of Cichlidogyrus tilapiae Paperna, 1960 based on 28S rRNA gene.

In the text

All Figures

thumbnail Figure 1

Freshly dead specimens of Chindongo socolofi (Johnson, 1974) examined for the present study. A. Blue pindani, B. White pindani. Photograph by Chawan Matey.
In the text
thumbnail Figure 2

Photomontage of light microscopy and phase contrast (PHACO) images of Cichlidogyrus tilapiae Paperna, 1960 from Chindongo socolofi (Johnson, 1974). A. anchor-bar complex and hooks, B and C. different configurations of male copulatory organ, D. vagina. Photograph by Amit Tripathi.
In the text
thumbnail Figure 3

Line drawings of sclerotised structures of Cichlidogyrus tilapiae Paperna, 1960 from Chindongo socolofi (Johnson, 1974). A. ventral anchor, B. ventral bar, C. dorsal anchor, D. dorsal bar, E. male copulatory organ, F. hook (pairs i–vii). Scale bar = 20 μm. Figure by Amit Tripathi.
In the text
thumbnail Figure 4

Median-joining haplotype networks based on a 687 bp fragment of small subunit rDNA and the first Internal Transcribed Spacer (left) and 631 bp of large subunit rDNA (right) from the newly sequenced individuals of Cichlidogyrus tilapiae from India, aligned with all previously published sequences from this species. Genotypes are represented by circles, with the size of the circle correlating with the number of isolates displaying the respective genotype. Colours denote the countries of sampling localities. Genotypes are connected by lines indicating the number of mutations between them.
In the text

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