(stratoudakis et al., 2000). infected eggs have been observed with a maximum of three parasites. most studies that have followed the progression of the host embryo after hatching have reported the death of all, or nearly all, hatched larvae following the rupturing of the yolk sac as the parasites are released into the water. gestal et al., 2006; shadrin et al., 2010) identification infected host larvae will develop an opaque coloration of the yolk sac due to being full of parasites. the yolk sac of infected larvae becomes increasingly more filled with dark parasite cells as development progresses (skovgaard et al., 2009). most eggs will be infected within two hours after spawning. one to two hours post-spawning, the parasite genome may be detected in host eggs by pcr analysis. after eight hours post-spawning, the parasites will start to become detectable by light microscopy. at this stage, parasites are generally 13-15 μm in diameter. two hours after being released from the host, parasites begin to swim with their two flagella. the transverse flagellum is about twice as long as the longitudinal flagellum, but both are around 300 nm in thickness (skovgaard et al., 2009). at this stage, they are between 20 and 24 μm in diameter. taking a cross section of the flagella reveals both flagellar cross-sections as being smooth and spherical, unlike typical dinokaryote dinoflagellate cells, which have flagellar dimorphism (skovgaard et al., 2009). in typical dinokaryote dinoflagellate cells, the transverse flagellum is ribbon-like and the axoneme runs along the longer, outer edge of the flagella, and a contractile fiber runs along the shorter, inner edge, while the other flagellum is spherical and smooth in cross section (skovgaard et al., 2009). the cingulum is visible in both light microscopy and scanning electron microscopy (sem) (skovgaard et al., 2009). when looking at the dorsal side of the cell, the cingulum divides the cell into two equal parts, the hyposome and the episome (skovgaard et al., 2009). however, when looking at the left lateral view of the cell, the hyposome appears twice as large as the episome (skovgaard et al., 2009). the sulcus can be viewed as a shallow depression in sem images of some cells (skovgaard et al., 2009). it extends ventrally from the site of insertion of the longitudinal flagellum on the apical end of the cell to the posterior end of the cell (skovgaard et al., 2009). the weak appearance of the cingulum and sulcus also make the morphology of ichthyodinium sp. distinct from typical dinoflagellates, which tend to have a more pronounced cingulum and sulcus (skovgaard et al., 2009). in a 2009 study, no mastigonemes were found on the flagella of the asymmetric i. chabelardi zoospores (skovgaard et al., 2009). this contradicts observations from 2006, in which mastigonemes were reported on the posterior flagellum of spherical zoospores (gestal et al., 2006). it is possible that this discrepancy is the result of the two different cell types having flagella of different morphology, or the result of the flagella in the 2009 study being damaged during fixation (skovgaard et al., 2009). economic significance ichthyodinium sp. has been reported in a large number of fin fish species of europe and asia. some of these, such as bluefin tuna, yellowfin tuna, mackerel, and cod, are of large economic significance (pedersen and koie, 1994; skovgaard et al., 2009). the annual catch of the japanese pilchard (sardinops melanostictus) has been rapidly dropping since 1989, and ichthyodinium sp. may be playing a role in this. in 2000, it was estimated that if sardine eggs infected with i. chabelardi experience 100% mortality, then this would roughly match the reported rates for the average daily mortality of sardine eggs and early larvae (stratoudakis et al., 2000). this suggests that ichthyodinium sp. may play a significant role in controlling populations of some fin fish species in the wild. ichthyodinium sp. has also been documented as a problem in some fish farms (mori et al., 2007;. in 2005, a semi-enclosed system tank of yellowfin tuna in bali, indonesia experienced substantial losses due to larval deaths. the yolk sacs of the larvae were observed to be completely filled with parasites, which caused them to rupture and kill the newly hatched larvae. a phylogenetic analysis of the ssu rrna sequence of the parasite revealed it to be ichthyodinium sp. a multi-year study conducted in nha trang bay, vietnam found a worrying trend of rapidly increasing prevalence of infected eggs and larvae (shadrin et al., 2010). the prevalence of infection in 1993 was estimated to be no more than 1% of eggs and larvae collected (shadrin et al., 2010). by the late 1990s, however, it was reported that a large proportion of the eggs and larvae collected in most samples were infected (shadrin et al., 2010). in 2006, it was recorded that 98.6% of eggs and larvae were infected in nearby cam ranh bay (shadrin et al., 2010). samples collected in nearby van phuong bay and phy quoc island in 2006 and 2008 also showed high rates of infection, between 29.3 and 59.4% (shadrin et al., 2010). it has been hypothesized that as global climate change leads to rising ocean temperatures, the ranges of many generalist species, such as i. chabelardi, may expand. the rising ocean temperatures could also increase the rates of infection of host species. together, these factors could lead to drastic increases in losses in the commercial fishing industry if fish tanks and surrounding seawater are not properly monitored, and appropriate steps taken to prevent infection of spawned eggs by ichthyodinium sp. ecology while ichthyodinium sp. was the suspected culprit of parasitized cod and turbot larvae found off the coast of denmark, (pedersen et al., 1994; it has been identified with certainty only in warm temperate and tropical waters (skovgaard et al., 2009). in a study conducted in 2003, samples of mackerel eggs were taken from fifty-nine locations stretching from cape finisterra, spain (43on) to the outer hebrides, uk (57on). eggs containing i. chabelardi were found from 43.4on to 53.8on, with a negative latitudinal trend in the proportion of eggs at each station that were parasitized. in the southernmost quartile of the samples, there was a mean infection rate of 54%, while in the northernmost quartile, the mean infection rate was only 9%. this suggests that i. chabelardi is most prevalent in warmer waters, closer to the equator. samples of fish eggs and larvae around vietnam between 2004 and 2008 showed high rates of infection (29.3 - 98.6%) in the southern samples, but no infected eggs at all were found in the northern samples (shadrin et al., 2010). a year-long survey of sardines from the adriatic sea in 1990 found i. chabelardi present in every month that had sardine egg spawning (dulcic, 1998). the infection rates varied considerably from month to month, with the colder months having higher infection rates and the warmer months having lower infection rates (dulcic, 1998). december (50%) and january (48.2%) had the highest rates of infection, while may (6.8%) and september (2.6%) had the lowest rates of infection (dulcic, 1998). no sardine eggs were spawned in june, july, or august, so no infection rates could be taken (dulcic, 1998). reported host species bluefin tuna (thunnus thynnus): identified in bluefin tuna in japan in 2004. bogue (boops boops): identified in bogue eggs off the coast of portugal (skovgaard et al., 2009). bristlemouth (mourolicus pennant): (hollande and cachon, 1952; cod (gadus morhua) *suspected: cod eggs collected off the coast of denmark were found to be infected with a protistan parasite that was hypothesized to be i. chabelardi with rates ranging between 14 and 88% (pedersen and koie, 1994). parasites were found in at least some eggs of all eight batches collected (pedersen and koie, 1994). european pilchard (sardina pilchardus): i. chabelardi was first identified in sardines from the mediterranean by hollande and cachon in 1952.(hollande & cachon, 1952; in 1997 and 1999, sardine eggs were sampled off the coast of portugal, and approximately 30% of them were found to be infected with i. chabelardi (stratoudakis et al., 2000). gilthead seabream (sparus aurata): (marinaro, 1971; leopard coral grouper (plectropomus leopardus): mass mortality events of leopard coral grouper around okinawa, japan caused by an unknown parasite have been reported repeatedly since 1990 (mori et al., 2007). based on an analysis of the parasite's mode of infection, development, and ssu rdna sequence, the unknown parasite was identified as ichthyodinium sp. (mori et al., 2007). mackerel (scomber scombrus): in 1998, fifty-nine samples of mackerel eggs, totaling 2,409 eggs, were collected in the north-east atlantic and checked for parasitism by i. chabelardi. of these, 1,099 eggs (46%) were found to be infected. thirty-nine (66%) of the collection stations had at least one parasitized egg. turbot (scopththalmus maximus) *suspected: turbot eggs collected off the coast of denmark were found to be infected with a protistan parasite that was hypothesized to be i. chabelardi (pedersen and koie, 1994; yellowfin tuna (thunnas albacares): in 2005, larval deaths of yellowfin tuna in a semi-enclosed system tank in bali, indonesia were attributed to ichthyodinium sp. geographical distribution the complete global distribution of ichthyodinium sp. is currently unknown (gleasen et al., 2019). among general surveys conducted to sample dinoflagellate species, no data for ichthyodinium sp. has been collected at most sites (gleasen et al., 2019). more surveys of ichthyodinium sp., taken at broader ranges, are needed to gain a better grasp of its true distribution (gleasen et al., 2019). ichthyodinium sp. has so far been recorded in the waters of algeria (holland and cachon, 1952), croatia (dulcic, 1998), denmark (pedersen and koie, 1994), indonesia, japan(mori et al., 2007), portugal (stratoudakis et al., 2000; skovgaard et al., 2009), spain, the united kingdom, and vietnam (shadrin et al., 2010).