Reports of Pennellidae Burmeister, 1835 (Copepoda: Siphonostomatoida) post metamorphosed females off the coast of southern Africa.

Seven species belonging to Pennellidae are reported from marine teleosts caught off southern Africa. Additionally, complete re-descriptions are provided for Propeniculus stromatei and Sarcotretes scopeli. Examination of Lernaeenicus gonostomae, deposited in the Iziko South African Museum, indicated that it has the morphological features of Sarcotretes rather than Lernaeenicus and thus should be moved to Sarcotretes i.e. S. gonostomae n. comb. for which a re-description is also provided. Reports of new host records include those of Pennella instructa from Seriola lalandi; Propeniculus stromatei from Rhabdosargus holubi and Pomadasys commersonnii; Sarcotretes scopeli from Nansenia tenera, and Sarcotretes longirostris from Centrolophus niger. New geographical records include those of P. instructa, P. stromatei, S. scopeli, S. longirostris, and L. longiventris off southern Africa. Additionally, an attempt to estimate the evolutionary relationships amongst some genera is done from partial COI sequences deposited in Genbank.

Twenty years later: Biodiversity of marine symbiotic Siphonostomatoida (Copepoda) off Southern Africa.

Siphonostomatoida (Copepoda) consists of 40 families of symbionts infecting vertebrates (17 families) and invertebrates (23 families) found mostly in marine habitats. In 2004, a list was compiled of all the reported families, genera and species symbiotic with marine fish in Southern African waters. Since this was done 20 years ago, it is necessary to re-evaluate the progress made in 20 years regarding our knowledge of the diversity of marine siphonostomatoids. To assess the current knowledge, the 2004 list was updated with reports published since 2004 as well as with new host and locality records including species with changes in taxonomy. Additionally, species collected but unreported as well as species previously reported but with new hosts and/or localities were also added. Currently reports include 16 families, 75 genera and 234 species. However, amongst these are reports of only two families (3 species) infecting invertebrates. Even though the report includes 71 additional species it still compares poorly with the about 2 274 accepted species, especially regarding species infecting invertebrates. Considering South Africas wealth in possible marine host species, examination of more hosts (especially marine teleosts and invertebrates) will definitely result in an increase in the current knowledge about the biodiversity of marine siphonostomatoids off Southern Africa.

Lophoura Kölliker in Gegenbaur, Kölliker & Müller, 1853 species (Copepoda: Siphonostomatoida: Sphyriidae) off South Africa with a key to all valid species.

Lophoura Kölliker in Gegenbaur, Kölliker & Müller, 1853 is one of the eight genera of Sphyriidae which currently consists of 19 accepted species. Lophoura species are mesoparasites of various teleosts occurring worldwide. Post metamorphic females are highly transformed with longitudinally elongated cephalothoraces and elongated necks bearing holdfast organs anteriorly. The shape and structure of the holdfast organ is mostly the main character used for the identification of species. Additionally, the structure of the posterior processes attached laterally to the perianal swelling on the posterior margin of the trunk is also used to distinguish different species. The posterior processes consist of a central porous peduncle with stalks of varying shapes and sizes. The morphology of Lophoura males bear resemblance to that of lernaeopodid males. Re-descriptions of the habitus of the post metamorphic females of L. tetraloba Ho & Kim I.H., 1989, L. cf. edwardsi Kölliker, 1853, L. caparti (Nunes-Ruivo, 1962) and L. cornuta (Wilson C.B., 1919), including notes on the immature females of L. tetraloba and L. cf. edwardsi, are provided. Additionally, information regarding the structure and position of some of the appendages of L. tetraloba, L. cf. edwardsi and L. caparti is provided. Furthermore, the study provides the first illustrated descriptions of the male of L. tetraloba. A description is also done of an incomplete unidentied Lophoura species which differs from other accepted species. A morphological identification key to post metamorphic females of all accepted Lophoura species is provided, as well as new host and geographical records of Lophoura species.

Symbiotic Siphonostomatoida (Copepoda) of the hammerhead shark species Sphyrna zygaena (Carcharhiniformes: Sphyrnidae) and stingray Dasyatis pastinaca (Myliobatiformes: Dasyatidae) off the Canary Islands, with a re-description of Pseudocharopinus pillaii Kabata, 1979.

The biology, including accompanying ectoparasitic fauna, remains unstudied for several elasmobranchs off the Canary Islands in spite of concerns over their conservation. Within this region, this is the first report of copepod associations with these fishes. Twenty smooth hammerhead sharks, Sphyrna zygaena, and one common stingray, Dasyatis pastinaca, were inspected for symbionts on their body surface. Symbiotic siphonostomatoids were collected from 18 S. zygaena hosts and represent two different species, i.e., Alebion crassus (Caligidae) and Pandarus cranchii (Pandaridae). A high prevalence of 90% and a mean infection of 8.3 individuals per infected host were exhibited by A. crassus, while only a single P. cranchii was collected. Both these findings represent new geographical records. Additionally, one free-ranging common stingray Dasyatis pastinaca was opportunistically inspected and 11 Pseudocharopinus pillaii (Lernaeopodidae) collected which constitutes a new host record and a new geographical record. A re-description with detail regarding the morphology of the appendages is provided for the first time. Due to the morphological similarity between P. pillaii and P. malleus, a molecular analysis of the partial mitochondrial DNA cytochrome oxidase I gene was done in an attempt to determine if there is molecular divergence between the two species. This provided an estimate of the phylogenetic relationships amongst four Pseudocharopinus species (P. bicaudatus, P. pteromylaei, P. malleus, and P. pillaii) with Charopinus dubius as an outgroup as an alternative hypothesis to compare with the previous morphological topology estimated for all Pseudocharopinus species.

Late postnaupliar development of the freshwater copepods Lovenula falicifera and Metadiaptomus colonialis (Calanoida: Diaptomidae) from South Africa.

The life cycle of calanoid copepods consists of eggs hatching into nauplii (6 stages) which then moult into copepodids (5 stages), followed by the final moult into the adult female and male. The family Diaptomidae contains two subfamilies, Diaptominae and Paradiaptominae, with paradiaptomids almost exclusively consisting of African taxa. The copepodid stages III, IV and V were described for some freshwater diaptomine genera (i.e., Eudiaptomus Kiefer, 1932, Aglaodiaptomus Light, 1938, Skistodiaptomus Light, 1939, Leptodiaptomus Light, 1938, Megadiaptomus Kiefer, 1936 and Diaptomus Westwood, 1836). Copepods collected from Turfloop Dam, South Africa, with a plankton net were fixed and preserved in 70% ethanol. Calanoid copepods were studied under stereo- and light microscopes, using the wooden slide technique and features drawn. Examined specimens were identified as the copepodid stages of two African species, Lovenula falcifera (Lovén, 1845) and Metadiaptomus colonialis (van Douwe, 1914). Copepodids of the two species can be distinguished by their body size and the structure and size of the maxillipeds. The description and illustrations of three postnaupliar stages (CoIII, CoIV and CoV) are provided for both species. The identification of different stages is based on the number of urosomites, antennule development, the segmentation of legs 1-4, and the development of the fifth leg. These copepodids are compared with those of other described diaptomid genera.

Neoalbionella izawai n. sp. from the smallfin gulper shark Centrophorus moluccensis Bleeker and additional host records for N. etmopteri (Yamaguti, 1939) off South Africa.

Neoalbionella Özdikmen, 2008 comprises 10 accepted species, which all infect sharks of Squaliformes and Carcharhiniformes. Adult females belonging to species of Neoalbionella, based on the maxillule palp armed with three setae and the maxilliped subchela claw with only one secondary denticle, were collected from sharks off the coast of South Africa. Neoalbionella izawai n. sp. collected from the anterodorsal part of the spiracle opening of Centrophorus moluccensis Bleeker differs from its congeners by having maxillae that are separated except at the tapering tips where they are fused and that are longer than the trunk, uropods originating from the pointed posterior margin of the trunk and that are well developed, and maxillipeds without an additional spine at the base of the subchela barb. Neoalbionella etmopteri (Yamaguti, 1939) is herein reported from two new hosts (Etmopterus spp.) off South Africa.

A redescription of the female of Lernaeopoda bivia Leigh-Sharpe, 1930 (Lernaeopodidae: Siphonostomatoida) and a first description of the male.

Within the family Lernaeopodidae Milne Edwards, 1840 the females of Lernaeopoda spp. are very similar to those of Neoalbionella spp. Contrarily, the male morphology is different enough to allow distinction between species of the two genera. Unfortunately, males are not always attached to collected females and therefore often not described. Hence the validity of some of the Lernaeopoda spp. are still questionable due to incomplete descriptions of the female and no description of the male. Lernaeopoda bivia Leigh-Sharpe, 1930 is an example of an accepted species due to a redescription of the female done in 1986 but without a description of the male. Lernaeopoda bivia has only been reported twice, collected from Schroederichthys bivius (Müller & Henle). Recently, females with attached males were collected from Triakis semifaciata Girard from Inner Cabrillo Beach in southern California. The female morphology of L. bivia is similar to the previous redescription with some additional features and illustrations while the male conforms to the morphology of males of Lernaeopoda spp. The dental formula of males and females of the species of the genus differ with males having one P1, S1 less than females. The structure and armature of the appendages of the three described species of males (L. galei Krøyer, 1837, L. bidiscalis Kane, 1892 and L. bivia) is mostly similar with the tympanal membrane of the three species the most distinctive feature. A key to identify the females of the valid species of Lernaeopoda is provided.

Cladistic analysis of the morphological characters of Pseudocharopinus Kabata, 1964 and keys to the species of Pseudocharopinus and Charopinus Krøyer, 1863 based on the morphology of adult females.

Kabata separated species of Pseudocharopinus Kabata, 1964 and Charopinus Krøyer, 1863 in 1964 based mainly on four characteristics. Currently Pseudocharopinus has 11 species while Charopinus consists of three species considered valid. Pseudocharopinus malleus (Rudolphi in Nordmann, 1832) and Charopinus dubius T. Scott, 1900 were collected and studied from hosts off South Africa. Additional to previous reports from South African waters, P. malleus is reported from Torpedo sinuspersici Olfers and T. fuscomaculata Peters while C. dubius is reported from Leucoraja wallacei (Hulley) and Rajella dissimilis (Hulley), all new host records. In an attempt to estimate the evolutionary relationships among Pseudocharopinus spp. a cladistic analysis was performed by means of parsimony using described and illustrated features of the adult females. Due to the mostly unresolved 50% majority rule tree, a key for the identification of the adult females of Pseudocharopinus spp. is compiled. Additionally, a key for the identification of adult females of Charopinus spp. is provided.

Resurrection of Alebion difficile (van Beneden, 1892) (Copepoda: Siphonostomatoida: Caligidae) with notes on other species of Alebion Krøyer, 1863 collected from elasmobranchs off South Africa and an updated key of the adult females.

Alebion Krøyer, 1863 is one of the genera of the Caligidae Burmeister, 1835 and can easily be distinguished from other genera by the presence of modified exopodal spines on legs 1-3. Currently there are eight recognised species infecting a variety of elasmobranch hosts. Alebion difficile (van Beneden, 1892) is resurrected based on the difference in the structure of the spermatophores on the genital complex of the female but also the difference in the relative lengths of the posterior processes of the genital complex, the lateral processes on the first abdominal somite and the caudal rami. Additionally, the adult male of A. difficile is described and illustrated and notes provided about the morphology of the sub-adult females and males. Other Alebion species and their hosts collected off South Africa are reported and an updated key to the adult females of the species of Alebion is provided.

Description of four new species and a revision of the genus Tripaphylus Richiardi in Anonymous, 1878 (Copepoda: Siphonostomatoida: Sphyriidae).

Tripaphylus Richiardi in Anonymous, 1878 is one of the genera of the Sphyriidae Wilson, 1919 and it currently accommodates ten species. Only scanty descriptions are available for the females and some males of the species. Tripaphylus specimens, described and illustrated in this paper, were collected from elasmobranch hosts off the South African coast or came from the private collection of the late Dr G. Benz. Collected specimens were fixed and preserved in 70% ethanol, examined by light - and scanning electron microscopy, and compared with the known species. Descriptions and illustrations are provided for the transformed adult females and males of T. elongatus (Wilson, 1932) and T. versicolor (Wilson, 1919) while illustrated descriptions of the transformed adult females are also provided for T. ferox (Wilson, 1919) and T. vaissierei (Delamare Deboutteville & Nuñes-Ruivo, 1954). Additionally, the material previously reported as T. ferox is considered to represent T. elongatus and that reported as T. lobatus (Kirtisinghe, 1964) is considered a junior synonym of T. vaissierei. Based on the collected material the transformed adult female and male of T. lewisi n. sp. are described as well as the adult transformed females of T. benzi n. sp., T. hoi n. sp., and T. beatricae n. sp. A summarised review of the main features is provided for each of the previously described T. musteli (van Beneden, 1851), T. hemigalei Kirtisinghe, 1964, T. australis (Kabata, 1993), T. triakis (Castro Romero, 2001) and T. asymboli (Turner, Kyne & Bennett, 2003). Furthermore, reasons are provided for the recommendation that T. hemigalei be considered a species inquirenda. An identification key to the transformed adult females of all species of Tripaphylus is provided and the host records are also updated.

Trebius benzi n. sp. (Siphonostomatoida: Trebiidae) infecting Squalus acutipinnis Regan off South Africa.

Trebius Krøyer, 1838 currently consists of 15 accepted species all infecting elasmobranchs. Apart from two species, i.e. T. caudatus Krøyer, 1838 and T. latifurcatus Wilson, 1921, that have been reported from ten and eight host species, respectively, the other 13 species have each been reported from only one or two host species. Trebius benzi n. sp., collected from Squalus acutipinnis Regan, is described and illustrated after examination through stereo- and compound microscopes. This species can be distinguished from the other known species by a combination of characters including an abdomen that is shorter than the genital complex, a maxillule with an endite that consists of a single-tined dentiform process, sternal furca tines that are blunt and as long as the base, and the innermost spine of the last exopodal segment of leg 1 the shortest.

Schistobrachia kabata sp. nov. (Siphonostomatoida: Lernaeopodidae) from rajiform hosts off South Africa.

The genus Schistobrachia Kabata, 1964 (Lernaeopodidae: Siphonostomatoida) currently accommodates five species of which two infect holocephalans and three utilize elasmobranchs. Kensley & Grindley (1973) reported three females from "Dipturus batis" (Linnaeus, 1758) collected in Table Bay, South Africa, which they assigned to S. ramosa (Krøyer, 1863), a species previously known only from the North Atlantic. Re-examination of these specimens (labelled Charopinus ramosus Krøyer, 1863 and deposited in the Iziko South African Museum) and additional Schistobrachia material newly collected from the gills of various Rajiformes off the South African west and south coasts, showed that Kensley & Grindley's (1973) earlier record of S. ramosa was misidentified and in reality represents a distinct species, S. kabata sp. nov. Both sexes of the new species are described using light and/or scanning electron microscopy.

Barnard's Brachiella sp., Parabrachiella supplicans (Barnard, 1955) and Eubrachiella sublobulata (Barnard, 1955) (Copepoda: Siphonostomatoida: Lernaeopodidae) deposited in the Iziko South African Museum.

The family Lernaeopodidae Milne Edwards, is one of the largest families in Siphonostomatoida Thorell, with more than 260 valid species. Brachiella sp., Parabrachiella supplicans (Barnard) and Eubrachiella sublobulata (Barnard) collected from Table Bay by Barnard in 1955 off Liza ramada (Risso), Genypterus capensis (Smith) and Congiopodus torvus (Gronov) respectively, are re-examined. Brachiella sp. is identified as Parabrachiella mugilis (Kabata, Raibaut & Ben Hassine) and is synonymized with Parabrachiella exilis (Shiino) based on the general morphology of the female habitus, the type host and the attachment site on the host. Parabrachiella supplicans is redescribed and synonymized with Parabrachiella genypteri (Capart) also collected off G. capensis from Fort Rock Point, Namibia. Additionally, Eubrachiella sublobulata is redescribed and represents a different geographical range and host record for the genus Eubrachiella as species were previously reported from teleosts of Antarctic waters.

Does Alella Leigh-Sharpe, 1925 (Siphonostomatoida: Lernaeopodidae) really consist of seven species?

Alella Leigh-Sharpe, 1925 (Lernaeopodidae) consists of seven species, namely A. pagelli (Krøyer, 1863), A. canthari (Heller, 1865), A. macrotrachelus (Brian, 1906), A. ditrematis (Yamaguti, 1939), A. pterobrachiata (Kabata, 1968), A. tarakihi Hewitt & Blackwell, 1987 and A. gibbosa van Niekerk & Olivier, 1995. Specimens of Alella collected from Rhabdosargus sarba (Forsskål), R. holubi (Steindachner), R. globiceps (Valenciennes) and Acanthopagrus berda (Forsskål) (Sparidae) from South African waters were examined. The specimens from R. sarba are from the same host individuals that A. gibbosa were described. However, comparing the armature of the appendages, no consistent and conclusive differences were found between the examined specimens and the other congeners. Additionally, a comparison of the morphology of the appendages and their armature of the examined specimens and those described and illustrated for the known species also revealed limited differences which may be due to regional or intraspecific variation and inconsistent observations rather than distinguishing characteristics of different species. Therefore it is proposed that Alella only has one valid species namely A. pagelli.

How females of Achtheinus spp. (Pandaridae: Siphonostomatoida) attach to their elasmobranch hosts with notes on their effects on the hosts' fins.

Copepods of the genus Achtheinus Wilson, 1908 (Pandaridae) are parasites of elasmobranchs that attach to their fins, gill slits and around the nostrils. Specimens of Achtheinus pinguis Wilson, 1912 were collected and examined using histology and scanning electron microscopy to determine their way of attachment to the host and the possible effect on the host. They insert their antennae deep into the dermis of the shark's skin, which causes the most damage due to possible tissue compression and/or fibrosis as well as rupture of the connective tissue. Additionally, the presence of the copepod on the skin causes cell erosion of the epidermal cells and thus reduces the number of epidermal layers. The maxillipeds are used to attach to the placoid scales that cover the shark's skin and probably serve to keep the copepod and inserted antennae in position. This is accomplished by the insertion of the placoid scales into the flaccid corpus of the maxillipeds. Observed damage seems to be negligible to the shark apart from the possibility of secondary infection.

Two new species of Pupulina van Beneden, 1892 (Copepoda: Siphonostomatoida: Caligidae) from mobulid rays off South Africa.

The caligid genus Pupulina van Beneden, 1892 currently has three accepted species. Two new species, Pupulina cliffi n. sp. and P. merira n. sp., are described from Mobula kuhlii (Müller & Henle) and M. eregoodootenkee (Bleeker) (Mobulidae) caught along the east coast of South Africa. Pupulina cliffi can be distinguished from all the other species by the absence of posterolateral processes on the genital complex, whereas P. merira has very short, rounded posterolateral processes on the genital complex compared to the three previously known species. Additionally, P. merira is the only species with the abdomen only about two-thirds the length of the genital complex and the caudal rami about the same length as the abdomen. A dichotomous key to distinguish the five species of Pupulina is provided.

A redescription of Pseudocharopinus pteromylaei Raibaut et Essafi, 1979 (Siphonostomatoida: Lernaeopodidae) collected from the South African east coast.

Pseudocharopinus Kabata, 1964 is one of the 48 genera comprising the Lernaeopodidae (Copepoda, Siphonostomatoida). Currently there are 11 accepted species contained in Pseudocharopinus. Pseudocharopinus pteromylaei Raibaut et Essafi, 1979 infects the spiracles and gill filaments of Pteromylaeus bovinus (Geoffroy St. Hilaire) and is most similar in general appearance to P. pteroplateae (Yamaguti et Yamasu, 1959) from which it clearly differs in the size and dimensions of the dorsal shield and the posterior processes. Additional features of the antenna and the maxilliped ofP. pteromylaei, not previously illustrated, are discussed and illustrated while more detailed illustrations of other appendages are provided. This is the first report of the occurrence of a Pseudocharopinus species from the west Indian Ocean and the first report of P. pteromylaei off the east coast of South Africa.

Cytochrome oxidase I sequences reveal possible cryptic diversity in the cosmopolitan symbiotic copepod Nesippus orientalis Heller, 1868 (Pandaridae: Siphonostomatoida) on elasmobranch hosts from the KwaZulu-Natal coast of South Africa.

Over the past decade, numerous molecular phylogenetic studies uncovered cryptic diversity within the Copepoda, yet very few investigations focused on symbiotic copepods. Here we report mitochondrial DNA cytochrome oxidase I diversity in the cosmopolitan elasmobranch symbiont Nesippus orientalis off the KwaZulu-Natal coast of South Africa. Analysis of partial COI sequences of copepods sampled from a diversity of shark hosts, revealed the presence of two divergent clades. Diversity within the clades does not appear to be structured based on host species, host individual, geographic locality or time of sampling. However, divergence between the two clades seems to be related to host species. Phylogenetic analyses of representatives from the two clades, along with Nesippus spp., Caligus spp. and Lepeophtheirus spp. outgroups, further supports the distinction between the two clades. Future molecular phylogenetic investigations of widespread copepod symbionts most likely will reveal far greater levels of biodiversity than currently recognized.

Description of the adult female and male of Naobranchia kabatana n. sp. (Copepoda: Lernaeopodidae) from Muraenesox bagio (Hamilton) (Muraenesocidae) caught in the Indian Ocean off South Africa.

Naobranchia Hesse, 1863 is a genus of the family Lernaeopodidae. This report presents an illustrated description of the adult female and male of a new species of this genus collected from the gill filaments of Muraenesox bagio (Hamilton) (pike conger) caught off the coast of Richards Bay, KwaZulu-Natal, South Africa. Its morphological features were examined using light and scanning electron microscopy. Characteristic features of the female Naobranchia kabatana n. sp. include egg-sacs situated along most the length of the trunk, cephalothorax and trunk of about equal lengths, and egg-sacs, lacking distinct sclerotised bands, which meet at the posterior end of the body and usually obscure the abdomen and caudal rami in ovigerous specimens. The male has no division into the cephalothorax and trunk, resulting in an unsegmented body with a prominent genito-abdominal papilla in a subterminal position.

New host and geographical records of siphonostomatoid copepods associated with elasmobranchs off the KwaZulu-Natal coast, South Africa.

The Siphonostomatoida (Copepoda) consists of 40 families comprised mainly of symbiotic members. Representatives of 12 families occur in symbiosis with elasmobranchs. Only eight families, 19 genera and 35 species have been reported from 37 elasmobranchs off the coast of South Africa. Elasmobranchs caught in the nets of the Natal Sharks Board and by fishermen of the Meer-en-See Boat Club, KwaZulu-Natal Province were examined for symbiotic siphonostomatoids. Additional to previous families reported, members of Cecropidae and Sphyriidae are being reported. Genera for the first time reported from South African waters include Entepherus, Carnifossorius, Eudactylina, Eudactylinodes, Kroeyerina, Pannosus, Phyllothyreus and Paeon. Therefore, currently ten siphonostomatoid families are reported from 49 species of elasmobranchs including 27 genera and 58 species.