Review of haploporid (Trematoda) genera with ornate muscularisation in the region of the oral sucker, including four new species and a new genus.

Species of the Haploporidae Nicoll, 1914 with elaborate muscularisation of the oral sucker belong in three trematode genera, including three new species and a new genus from the intestine of fishes in Australian waters. Spiritestis Nagaty, 1948 is resurrected and S. herveyensis n. sp. is described from the mullet Moolgarda seheli (Forsskål) collected in Hervey Bay, Queensland, Australia; the latter differs from S. arabii Nagaty, 1948 in that the position of the genital pore is pharyngeal rather than post-pharyngeal and the geographical range is off Australia rather than the Red Sea. A new genus is proposed for two new species, with a uniquely ornamented oral sucker, which infect Australian scatophagids. Members of Capitimitta n. g. are distinguished from Waretrema Srivastava, 1937, species of which have a simple oral sucker with six radially arranged anterior muscular lobes, in that their oral sucker is V-shaped with six embedded muscular finger-like structures in the anteroventral portion. The relatively small C. darwinensis n. sp., collected from Selenotoca multifasciata (Richardson) at Darwin, Northern Territory, Australia, is distinguished from C. costata n. sp., collected from Scatophagus argus (Linnaeus) in the same locality and S. multifasciata off Brisbane, Australia, and by having smaller eggs, a vitellarium commencing at a level close to the ventral sucker rather than at greater than one ovarian length posterior to the ventral sucker, and shorter tegumental body spines. Sequence data of a c.2,500 bp region of the 3' end of 18S, the entire ITS region and the 5' end of the 28S revealed that Spiritestis and Capitimitta are not as closely related as some morphological features would suggest and are probably not the closest relative of each other. What has been reported as Waretrema piscicolum Srivastava, 1937 probably consists of several species, some in different genera, and one, based on material collected by Dr Masaaki Machida, is proposed as Spiritestis machidai n. sp. from Crenimugil crenilabis (Forsskål) off Japan. Phylogenetic hypotheses, based on analysis of an alignment of partial 28S sequences with other haploporids, provide a framework for the evaluation of interrelationships within the Haploporidae. These analyses show that: (1) Spiritestis and Capitimitta are supported within the Haploporidae; (2) branches to Forticulcita Overstreet, 1982, Saccocoelioides Szidat, 1954, Spiritestis and Capitimitta create a clade that is sister to haploporines from the Mediterranean Sea; (3) the branch to Saccocoelioides, Spiritestis and Capitimitta create a polytomy; and (4) the two new species of Capitimitta, plus an immature specimen of an unnamed species, form a monophyletic clade.