This paper reviews past, current and likely future research on the fish haemogregarine, Haemogregarina bigemina Laveran et Mesnil, 1901. Recorded from 96 species of fishes, across 70 genera and 34 families, this broad distribution for H. bigemina is questioned. In its type hosts and other fishes, the parasite undergoes intraerythrocytic binary fission, finally forming mature paired gamonts. An intraleukocytic phase is also reported, but not from the type hosts. This paper asks whether stages from the white cell series are truly H. bigemina. A future aim should be to compare the molecular constitution of so-called H. bigemina from a number of locations to determine whether all represent the same species. The transmission of H. bigemina between fishes is also considered. Past studies show that young fish acquire the haemogregarine when close to metamorphosis, but vertical and faecal-oral transmission seem unlikely. Some fish haemogregarines are leech-transmitted, but where fish populations with H. bigemina have been studied, these annelids are largely absent. However, haematophagous larval gnathiid isopods occur on such fishes and may be readily eaten by them. Sequential squashes of gnathiids from fishes with H. bigemina have demonstrated development of the haemogregarine in these isopods. Examination of histological sections through gnathiids is now underway to determine the precise development sites of the haemogregarine, particularly whether merozoites finally invade the salivary glands. To assist in this procedure and to clarify the internal anatomy of gnathiids, 3D visualisation of stacked, serial histological sections is being undertaken. Biological transmission experiments should follow these processes.
Haemogregarina bigemina Laveran et Mesnil, 1901 was examined in marine fishes and the gnathiid isopod, Gnathia africana Barnard, 1914 in South Africa. Its development in fishes was similar to that described previously for this species. Gnathiids taken from fishes with H. bigemina, and prepared sequentially over 28 days post feeding (d.p.f.), contained stages of syzygy, immature and mature oocysts, sporozoites and merozoites of at least three types. Sporozoites, often five in number, formed from each oocyst from 9 d.p.f. First-generation merozoites appeared in small numbers at 11 d.p.f., arising from small, rounded meronts. Mature, second-generation merozoites appeared in large clusters within gut tissue at 18 d.p.f. They were presumed to arise from fan-shaped meronts, first observed at 11 d.p.f. Third-generation merozoites were the shortest, and resulted from binary fission of meronts, derived from second-generation merozoites. Gnathiids taken from sponges within rock pools contained only gamonts and immature oocysts. It is concluded that the development of H. bigemina in its arthropod host illustrates an affinity with Hemolivia and one species of Hepatozoon. However, the absence of sporokinetes and sporocysts also distances it from these genera, and from Karyolysus. Furthermore, H. bigemina produces fewer sporozoites than Cyrilia and Desseria, although, as in Desseria, Haemogregarina (sensu stricto) and Babesiosoma, post-sporogonic production of merozoites occurs in the invertebrate host. The presence of intraerythrocytic binary fission in its fish host means that H. bigemina is not a Desseria. Overall it most closely resembles Haemogregarina (sensu stricto) in its development, although the match is not exact.