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Manta rays (Mobula alfredi and M. birostris) are poorly understood in South Africa, despite their ecological importance and charismatic appeal. This study analyzed a 41-year dataset from the KwaZulu-Natal bather protection program to investigate catch per unit effort between 1981-2021. We used Generalized Additive Models and the probability of encounter to assess annual and seasonal trends, as well as the effect of location and moon phases on catch rates. We also evaluated the size composition and demographics of caught manta rays using the same dataset. Our analysis revealed a significant decline in overall manta ray catches since the late 1990s (p

Subfamily BERNDTIINAE Utinomii, 1950 Berndtiidae Utinomi, 1950: 457. Weltneriinae Kolbasov & Newman, 2005: 40. Diagnosis. Females with five pairs of terminal cirri. Caudal appendages present in Weltneria, but not in Berndtia. Remarks. Currently there are two genera described from this monophyletic subfamily, Berndtia Utinomi, 1950 and Weltneria. No species of Berndtia are reported from Africa (Kolbasov, 2009), however further investigation of coral species could potentially reveal their presence, as corals are their preferred substrate (Chan, Hsieh & Kolbasov, 2014; Chan et al., 2014)., Published as part of Botha, Thomas P. A. & Griffiths, Charles L., 2021, South African Acrothoracica (Crustacea: Cirripedia), pp. 45-78 in Zootaxa 4949 (1) on page 55, DOI: 10.11646/zootaxa.4949.1.3, http://zenodo.org/record/4635765, {"references":["Utinomi, H. (1950) General account of Acrothoracica. In: Nakamura, K. (Ed.), Some Problems in Modern Biology. Zoshindo, Osaka, pp. 428 - 461. [in Japanense]","Kolbasov, G. A. & Newman, W. A. (2005) Revision of the Lithoglyptidae sensu Tomlinson, 1969 and Lithoglyptes Aurivillius, 1892 (Cirripedia, Acrothoracica), including a new species from Bermuda. Zootaxa, 1013 (1), 35 - 64. https: // doi. org / 10.11646 / zootaxa. 1013.1.3","Kolbasov, G. A. (2009) Acrothoracica, burrowing crustaceans. KMK Scientific Press Ltd, Moscow, 452 pp.","Chan, B. K. K., Hsieh, W-P. & Kolbasov, G. A. (2014) Crustacean Fauna of Taiwan: Barnacles. Vol. III. Cirripedia: Acrothoracica. Biodiversity Research Center, Academia Sinica, Taipei, 107 pp."]}

Australophialus utinomii Tomlinson, 1969 Figs 5, 6 Australophialus utinomii Tomlinson, 1969: 119–121.— Kolbasov 2009: 354, fig. 127. Deposited material. SAMC-A091096, near East London, Eastern Cape, April 2017, 79 specimens in 7 Burnupena cincta shells (4 not deposited, used for SEM and genetic analyses). SAMC-A091097, near East London, Eastern Cape, April 2017, 29 specimens in 5 Burnupena cincta shells (11 not deposited, used for SEM and genetic analyses). SAMC-A091099, Gonubie, Eastern Cape, 27 April 2017, 6 specimens in Mancinella capensis shell. SAMC-A091098, Chintsa West, Eastern Cape, 26 April 2017, 6 specimens in 2 Dinoplax gigas shell plates. SAMC- A091100, near East London, Eastern Cape, April 2017, 7 specimens in 3 Turbo sarmaticus shells. SAMC-A091101, near East London, Eastern Cape, April 2017, 81 specimens from 4 Turbo sarmaticus shells (4 not deposited, used for dissections). SAMC-A091102, Gonubie, Eastern Cape, 27 April 2017, 4 specimens in Ranella gemmifera shell. SAMC-A091093, Isipingo, KwaZulu-Natal, 21 September 2017, 44 specimens in Purpura panama shell (34 deposited, 10 used for dissections). SAMC-A091091, Banana Beach, Port Shepstone, KwaZulu-Natal, 20 September 2017, 35 specimens from 2 Purpura panama shells. SAMC-A091092, Banana Beach, Port Shepstone, KwaZulu-Natal, 20 September 2017, 5 specimens in Purpura panama shell. SAMC-A091094, Banana Beach, Port Shepstone, KwaZulu-Natal, 20 September 2017, 7 specimens in Purpura bufo shell. SAMC-A091095, Port Shepstone, KwaZulu-Natal, 19 September 2017, 1 specimen from Dinoplax validifossus shell plate. Other records. PP08, Banana Beach, Port Shepstone, KwaZulu-Natal, 20 September 2017, 2 specimens from Purpura panama shell. Diagnosis. Similar to A. turbonis, but without the distinctive notch in opercular margin, rather has an entire margin in shape of small shield, no prominent hooks or spines, fringed with small, numerous bifid teeth. Description. Female, body shape characteristic of cryptophialids, with mantle bottle-like and neck elongated (fig. 5A). Length 0.65–2 mm (mean = 1.18 mm), width 0.4–0.95 mm (mean = 0.68 mm). Opercular bars with average length 0.35 mm, with plenty of setae and simple, bifid and rarely trifid teeth on margin (fig. 6D, E). Large teeth and setae present in posterior part of opercular bar (fig. 6B). Anterior end of opercular bar elongated with several teeth and setae (fig. 6E). Comb collar consisting of small projections fused at their bases (fig. 6A). Surface of opercular area striated (as is whole of mantle), with small multifid scales (fig. 6F). Pair of lateral bars support either side of mantle, these bars pointed and associated with 3–4 rows of small teeth. Reinforcing bar found within apertural region at anterior end. Colour light brown/white when fresh, becoming slightly paler when preserved in alcohol. Burrow opening small, oval-shaped (fig. 5B). Four pairs of biramous, multi-annulated terminal cirri present (fig. 5C). Pair of rudimentary uniramous, unsegmented mouth cirri present on anterio-ventral thorax area, posterior to mouth. Labrum pronounced, tongue-shaped and extends far beyond opercular area; large in comparison to others of this genus (fig. 6D). Mouthparts typical of genus, a pair of mandibles with palps present, along with a pair of maxillae and maxillules. One dorsal body process present. Females with maximum of four dwarf males recorded. Dwarf males 0.29 x 0.16 mm. Cypris larvae 0.25 x 0.15 mm (fig. 5D) Hosts. Previously known only from Dinoplax gigas. Additional hosts in the Eastern Cape include Burnupena cincta, Mancinella capensis, Ranella gemmifera and Turbo sarmaticus. Additional hosts in KwaZulu-Natal include Dinoplax validifossus, Purpura bufo and P. panama. Distribution. Type locality Qolora (3241’50.5”S, 2822’25.1”E), Eastern Cape, roughly 50 km east of East London. Ranges from Gonubie (3256’33.8”S, 2802’00.9”E) to Isipingo (2959’44.1”S, 3057’00.8”E). Endemic to Eastern Cape and KwaZulu-Natal. Remarks. Common, often in association with Kochlorine bocqueti Turquier, 1977, Kochlorine sp. A and Weltneria spinosa Berndt, 1907. Due to its small size, attempts to dissect this species were not as successful as others., Published as part of Botha, Thomas P. A. & Griffiths, Charles L., 2021, South African Acrothoracica (Crustacea: Cirripedia), pp. 45-78 in Zootaxa 4949 (1) on pages 52-55, DOI: 10.11646/zootaxa.4949.1.3, http://zenodo.org/record/4635765, {"references":["Tomlinson, J. T. (1969) The burrowing barnacles (Cirripedia: Order Acrothoracica). Bulletin of the United States National Museum, 296, 1 - 162. https: // doi. org / 10.5479 / si. 03629236.296.1","Kolbasov, G. A. (2009) Acrothoracica, burrowing crustaceans. KMK Scientific Press Ltd, Moscow, 452 pp.","Turquier, Y. (1977) Etude de quelques Cirripdes Acrothoraciques de Madagascar. III. Kochlorine bocquette n. sp. Archives de Zoologie Experimentale et Generale, 118 (2), 133 - 154.","Berndt, W. (1907) Uber das System der Acrothoracica. Archiv fur Naturgeschichte, 73 (1), 287 - 289."]}

Genus Australophialus, Tomlinson, 1969 Australophialus Tomlinson, 1969: 117. Diagnosis. Characterised by having four pairs of terminal cirri and one whip-like dorsal body process (Chan, Hsieh & Kolbasov, 2014). Remarks. Globally, there are currently five species in this genus, two of which are found in South Africa (both endemic)., Published as part of Botha, Thomas P. A. & Griffiths, Charles L., 2021, South African Acrothoracica (Crustacea: Cirripedia), pp. 45-78 in Zootaxa 4949 (1) on page 49, DOI: 10.11646/zootaxa.4949.1.3, http://zenodo.org/record/4635765, {"references":["Tomlinson, J. T. (1969) The burrowing barnacles (Cirripedia: Order Acrothoracica). Bulletin of the United States National Museum, 296, 1 - 162. https: // doi. org / 10.5479 / si. 03629236.296.1","Chan, B. K. K., Hsieh, W-P. & Kolbasov, G. A. (2014) Crustacean Fauna of Taiwan: Barnacles. Vol. III. Cirripedia: Acrothoracica. Biodiversity Research Center, Academia Sinica, Taipei, 107 pp."]}

Superorder ACROTHORACICA Gruvel, 1905 Cryptosomata Hancock, 1849: 313. Abdominalia Darwin, 1854: 583. Acrothoracica Gruvel, 1905: 310. Diagnosis. Small, soft-bodied barnacles. Females burrowing into calcareous substrates (molluscs, corals, limestone, thoracicans, etc.). Minute dwarf males attached either to the female, or to her burrow. Lacking calcareous shell plates, but possessing a single pair of chitinous plates, also known as opercular bars, guarding the aperture. 3–5 pairs of terminal cirri, a single pair of mouth cirri developed or reduced, adults with reduced abdomen. Caudal appendages present (some lithoglyptids) or absent (all cryptophialids). Remarks. Currently the Acrothoracica consists of two orders, the Cryptophialida and the Lithoglyptida (Kolbasov, 2009)., Published as part of Botha, Thomas P. A. & Griffiths, Charles L., 2021, South African Acrothoracica (Crustacea: Cirripedia), pp. 45-78 in Zootaxa 4949 (1) on page 48, DOI: 10.11646/zootaxa.4949.1.3, http://zenodo.org/record/4635765, {"references":["Gruvel, A. (1905) Monographie des cirrhipedes ou thecostraces. Masson et Cie, Paris, 472 pp.","Hancock, A. (1849) Notice of the occurrence on the British coast of a burrowing barnacle belonging to a new order of the class Cirripedia. Annals and Magazine of Natural History, Series 2, 4 (23), 305 - 314. https: // doi. org / 10.1080 / 03745486009494840","Darwin, C. (1854) A monograph on the sub-class Cirripedia. Vol. II. The Balanidae. In: Barrett, P. & Freeman, R. B. (Eds.), The Works of Charles Darwin. Vol. 12. New York University Press, Washington Square, New York, pp. vii - viii + 1 - 684, figs. 1 - 7.","Kolbasov, G. A. (2009) Acrothoracica, burrowing crustaceans. KMK Scientific Press Ltd, Moscow, 452 pp."]}

Kochlorine bocqueti Turquier, 1977 Figs 18–20 Kochlorine bocqueti Turquier, 1977: 134.—1978: 107.— Kolbasov 1999: 139.— 2002: 536, 540, fig. 7 (u).—2009: 222 fig. 10 (в), fig. 13 (З), fig. 15 (е), fig. 109.. Deposited material. ELMC 0420, Kwelera Bay, Eastern Cape, 7 January 1986, 21 specimens in Charonia lampas shell (7 specimens not deposited, used for SEM, dissections and genetic analyses). SAMC-A091087, Gonubie, Eastern Cape, 27 April 2017, 4 specimens in Ranella gemmifera shell (1 not deposited, used for genetic analyses). SAMC-A091086, Gonubie, Eastern Cape, 27 April 2017, 2 specimens in 2 Turbo sarmaticus shells. SAMC- A091089, Gonubie, Eastern Cape, 27 April 2017, 2 specimens in 2 Haliotis midae shells. SAMC-A091088, Chintsa West, Eastern Cape, 26 April 2017, 1 specimen in Dinoplax gigas. Other records. Gonubie, Eastern Cape, 27 April 2017, 1 specimen on Mancinella capensis shell (specimen used for genetic analyses). Diagnosis. Opercular bars with a pair of small posterior processes, and with bifid and 8–13 long, lance-shaped teeth. Description. Female, length 1. 7–3.4 mm (mean = 2.69 mm) and width 0.85–1.85 mm (mean = 1.56 mm). Mantle sac oval-shaped (fig. 18C; 20A). Opercular bars average 0.98 mm long, armed by medial row of ‘arrowhead-shaped’ teeth and setae, also lined with lateral row of bifid teeth and more setae (fig. 19D, E). Two posterior processes on opercular bar with several bifid and simple teeth (fig. 19A). Comb collar feather-like, with small projections along entire edge (fig. 19B). Orificial knob well developed (fig. 20B). Surface below opercular bars smooth and ‘plate-like’, in contrast to surface of rest of animal, which is smooth (fig. 19A). Developed lateral bars running from opercular bar downwards, reinforcement bars running from anterior part of opercular bar downwards. Female bright red below opercular area, rest of body brownish-red (fig. 18A), becoming dark brown when preserved (fig. 18C). Burrow opening slit-like, elongated (fig. 18B). Three pairs of terminal cirri and caudal appendages two-segmented (fig. 20C). Mouth cirri with four-segmented posterior ramus shorter than six-segmented anterior ramus, both rami with long, plumose setae on each segment; rami extending off two-segmented protopod (fig. 20H). Mandible with three large teeth, first separated from remainder by notch; inferior angle with small teeth and setae (fig. 20E). Maxillule with two cuspidate setae above notch, with several setae, lower two-thirds of cutting edge with several sharp setae (fig. 20F). Maxilla triangular, with long dense setae at tip (fig. 20D). Mandibular palp trapezoid with long dense setae at tip (fig. 20G). No males observed. Hosts. Previously described from Charonia tritonis and Conus terebra. Collected in South Africa in the areas surrounding East London from Charonia lampas, Dinoplax gigas, Haliotis midae, Mancinella capensis, Ranella gemmifera and Turbo sarmaticus. Distribution. Previously known from Madagascar and Socotra Island (Kolbasov, 2002). In South Africa found intertidally, and subtidal to 30 m depth, in the Eastern Cape, ranging from Kwelera Bay (3250’08.0”S, 2807’09.5”E) to Gonubie (3256’33.8”S, 2802’00.9”E). In South Africa found only in Eastern Cape Province. Remarks. Differences were observed in the number of segments of each ramus of the mouth cirri; the posterior ramus with 6 segments, the anterior with 4. There are currently 7 species in this genus, with the removal of K. bihamata (now considered a nomen nudum, see above), the addition of Kochlorine sp. A. (below), and not including two undescribed species reported by Chan, Hsieh & Kolbasov (2014). Our identification is based partly on similarity in overall body shape and length to Kochlorine bocqueti, which has a length of 2.1 mm and maximum width of 1.2 mm (Kolbasov, 2002 a), both being within the ranges of the South African specimens (1.7–3.4 mm and 0.85–1.85 mm, respectively). Similarly, the reported opercular bar size (0.75 mm) is within the range found in South Africa (0.6–1.35 mm). More importantly, the South African specimens fit the diagnosis reported for K. bocqueti, which is (translated from Russian) “Opercular bars with a pair of small posterior processes armed with bifid and 8–13 long, sharp lance-shaped teeth”. Finally, K. bocqueti is already known from sites in the western Indian Ocean (Madagascar and Socotra Island) and thus its discovery in the East London area represents a plausible range extension down the east coast of Africa. This discovery not only represents a considerable range extension but adds several new hosts. The species was also found in association with A. utinomii, W. cf. hirsuta and W. spinosa., Published as part of Botha, Thomas P. A. & Griffiths, Charles L., 2021, South African Acrothoracica (Crustacea: Cirripedia), pp. 45-78 in Zootaxa 4949 (1) on pages 70-73, DOI: 10.11646/zootaxa.4949.1.3, http://zenodo.org/record/4635765, {"references":["Turquier, Y. (1977) Etude de quelques Cirripdes Acrothoraciques de Madagascar. III. Kochlorine bocquette n. sp. Archives de Zoologie Experimentale et Generale, 118 (2), 133 - 154.","Kolbasov, G. A. (1999 a) The external mantle morphology of burrowing barnacles of the families Lithoglyptidae and Cryptophialidae (Cirripedia, Acrothoracica). Crustaceans and the Biodiversity Crisis. Proceedings of the Fourth International Crustacean Congress, 1998, 1, 139 - 149.","Kolbasov, G. A. (2002) A new species of burrowing barnacles, Kochlorine grebelnii, from the Cape Verde Islands. Description of external ultrastructure of the genus Kochlorine (Thecostrata, Cirripedia, Acrothoracica). Zoological Journal, 81 (5), 529 - 546.","Chan, B. K. K., Hsieh, W-P. & Kolbasov, G. A. (2014) Crustacean Fauna of Taiwan: Barnacles. Vol. III. Cirripedia: Acrothoracica. Biodiversity Research Center, Academia Sinica, Taipei, 107 pp."]}

Australophialus turbonis (Barnard, 1925) Figs 2–4 Cryptophialus turbonis Barnard, 1925: 5 (type material lost).— Newman & Ross, 1971: 26. Australophialus turbonis Tomlinson, 1969: 121–125, fig. 33.—1987: 66, fig. 3.— Kolbasov & Høeg 2007: 129, fig. 3 (F), fig. 4 (K), fig. 10 (A).— Kolbasov 2009: 352, fig. 3 (a), fig. 7 (в), fig. 19 (б), fig. 20 (б, e, ж), fig. 21 (u), fig. 22 (б), fig. 24 (б), fig. 28 (б, e), fig. 38 (ж), fig. 50 (a), fig. 52 (г), fig. 55 (ж), fig. 59 (a), fig. 126. Deposited material. SAMC-A091103, False Bay, Western Cape, February 2017, 30 specimens in 3 Turbo cidaris shells. SAMC-A091104, Wooley’s Pool, False Bay, Western Cape, 30 January 2017, 3 specimens in 2 Turbo sarmaticus shells. SAMC-A091105, False Bay, Western Cape, January and February 2017, 308 specimens in 4 Turbo sarmaticus shells (19 not deposited, used for dissections, SEM, light microscopy and genetic analyses). SAMC-A091106, Melkbosstrand, Western Cape, 28 February 2017, 1 specimen in Fusinus ocelliferus shell. SAMC- A091107, Melkbosstrand, Western Cape, 28 February 2017, 2 specimens in Haliotis midae shell. SAMC-A091108, Bloubergstrand, Western Cape, April 2017, 2 specimens from 2 Argobuccinum pustulosum shells. SAMC-A091109, False Bay, Western Cape, 15 March 2017, 1 specimen in Burnupena cincta shell. SAMC-A091110, False Bay, Western Cape, 15 March 2017, 1 specimen in Burnupena cincta shell. SAMC-A091111, False Bay, Western Cape, 27 September 2018, 17 specimens from 3 Crepidula porcellana shells. SAMC-A091112, False Bay, Western Cape, 15 March 2017, 1 specimen in Burnupena cincta shell. Diagnosis. Australophialid with a notch one-third distance from anterior end on apertural margin, otherwise similar to other South African australophialid, Australophialus utinomii. Description. Female, mantle bottle-shaped, with characteristic elongated neck of all cryptophialids (Figs 2C; 4A). Length 1–3 mm (mean 1.95 mm), width 0.6–1.6 mm (mean 1.19 mm). Opercular bars ~ 300 m long and wellarmed on margins by long, simple teeth and setae, with nodules on flat surface of opercular bar (fig. 3A, D, F). Distinct notch on opercular bars about one-third from anterior end (fig. 3D, F; 4B). An infolded velum with lamellae extends into comb collar, which is composed of long, fine projections, ending in bristles (fig. 3E). Lateral surfaces of operculum with dense rows of multifid scales, with setae (fig. 3B). Mantle surface striated, with no obvious projections other than small teeth that are mostly bifid and uniformly distributed. Large reinforcing bars present, with one extending dorsally toward attachment area on dorsal end of aperture (fig. 4B). Pair of lateral bars extends from apertural notch downward along mantle on either side (fig. 4B), with fewer teeth on their surface, as normal for this genus. Attachment area masked by layers of previous moults (cemented exuviae) that form a hard disk onto which males attach (Tomlinson, 1969). Colour similar to W. spinosa, light reddish and white colour when fresh turning light brown/white when preserved in ethanol (fig. 2C). Burrow oval-shaped (fig. 2B), averaging 0.3 x 0.25 mm. Terminal cirri with four pairs, without caudal appendages (fig. 4C). One whip-like dorsal body process (fig. 4D). Mouth cirrus (paired) reduced, consisting only of small processes with several bristles (Tomlinson, 1969). Characteristic of the genus, labrum elongated and tongue-shaped, extending out of aperture, with setae at tip (fig. 4F). Mouthparts typical of genus, with ovate maxilla with distal setae (fig. 4E). Mandible with three teeth on cutting edge. Maxillule with no notch, no dense and long setae (fig. 4E). Mandibular palp elongated, with setae at distal end. Mature attached males averaging 0.45 x 0.25 mm. As many as 17 males attached to single female. Hosts. Type host Turbo sarmaticus. Previously known to bore into shells of Burnupena cincta and Burnupena limbosa. New hosts reported here are Argobuccinum pustulosum, Crepidula porcellana, Fusinus ocelliferus, Haliotis midae and Turbo cidaris. Distribution. Type locality False Bay (3407’57.8”S, 1826’44.1”E), South Africa. Ranges from Paternoster (3248’28.90”S, 1753’29.05”E) to Hermanus (3425’1.62”S, 1914’55.46”E). Endemic to the Western Cape. Remarks: Australophialus turbonis was previously reported from the Western Cape in various hosts (Barnard, 1925) and at Umpangazi in KwaZulu-Natal (Tomlinson, 1969) from Purpura persica (previously Thais rudolphi). However, this was probably a misidentification and likely was a specimen of A. utinomii, which is morphologically similar to A. turbonis (see below). Australophialus turbonis is only known from the Western Cape, whereas A. utinomii occurs in the Eastern Cape around East London and in KwaZulu-Natal. Moreover, A. utinomii is also known from two Purpura species (see below), while this is not a known host for A. turbonis. Commonly found in the same hosts in association with Weltneria spinosa., Published as part of Botha, Thomas P. A. & Griffiths, Charles L., 2021, South African Acrothoracica (Crustacea: Cirripedia), pp. 45-78 in Zootaxa 4949 (1) on pages 49-52, DOI: 10.11646/zootaxa.4949.1.3, http://zenodo.org/record/4635765, {"references":["Barnard, K. H. (1925) Report on a collection of Cirripedia (barnacles) from South African waters. Fisheries and Marine Biological Survey, Union of South Africa, Report, 4 (6), 1 - 5.","Newman, W. A. & Ross, A. (1971) Antarctic Cirripedia: Monographic Account Based on Specimens Collected Chiefly Under the United States Antarctic Research Program, 1962 - 1965. Vol. 14. American Geophysical Union, Washington, D. C., 257 pp.","Tomlinson, J. T. (1969) The burrowing barnacles (Cirripedia: Order Acrothoracica). Bulletin of the United States National Museum, 296, 1 - 162. https: // doi. org / 10.5479 / si. 03629236.296.1","Kolbasov, G. A. & Hoeg, J. T. (2007) Cypris larvae of acrothoracican barnacles (Thecostraca: Cirripedia: Acrothoracica). Zoologischer Anzeiger-A Journal of Comparative Zoology, 246 (2), 127 - 151. https: // doi. org / 10.1016 / j. jcz. 2007.03.001","Kolbasov, G. A. (2009) Acrothoracica, burrowing crustaceans. KMK Scientific Press Ltd, Moscow, 452 pp."]}

Genus Kochlorine Noll, 1872 Kochlorine.— Tomlinson, 1969: 65. Diagnosis. Females with three pairs of terminal cirri, caudal appendages present (two-segmented), developed lateral bars and orificial knob. Dwarf males with elongated attachment stalk and lateral projections (Chan, Hsieh & Kolbasov, 2014). Remarks. There are currently seven described species in this genus, of which K. bihamata, was previously thought to occur in South Africa. Below we suggest that report be discounted. However, two other species belonging to this genus are reported for the first time from the region, one of which is considered new to science., Published as part of Botha, Thomas P. A. & Griffiths, Charles L., 2021, South African Acrothoracica (Crustacea: Cirripedia), pp. 45-78 in Zootaxa 4949 (1) on page 69, DOI: 10.11646/zootaxa.4949.1.3, http://zenodo.org/record/4635765, {"references":["Noll, F. C. (1872) Kochlorine hamata N., ein bohrende Cirripede. Bericht uber die Senckenbergische naturf. Gesellsch 1871 - 1872, 4, 50 - 58","Tomlinson, J. T. (1969) The burrowing barnacles (Cirripedia: Order Acrothoracica). Bulletin of the United States National Museum, 296, 1 - 162. https: // doi. org / 10.5479 / si. 03629236.296.1","Chan, B. K. K., Hsieh, W-P. & Kolbasov, G. A. (2014) Crustacean Fauna of Taiwan: Barnacles. Vol. III. Cirripedia: Acrothoracica. Biodiversity Research Center, Academia Sinica, Taipei, 107 pp."]}

Subfamily KOCHLORININAE Gruvel, 1905 Gruvel, 1905: 335. Diagnosis. Females with three pairs of biramous terminal cirri, two-segmented caudal appendages present. Remarks. Consists of two genera; Kochlorine Noll, 1872 and Kochlorinopsis Stubbings, 1967, with only the former present in South Africa. Globally these genera contain 7 and 1 species, respectively., Published as part of Botha, Thomas P. A. & Griffiths, Charles L., 2021, South African Acrothoracica (Crustacea: Cirripedia), pp. 45-78 in Zootaxa 4949 (1) on page 69, DOI: 10.11646/zootaxa.4949.1.3, http://zenodo.org/record/4635765, {"references":["Gruvel, A. (1905) Monographie des cirrhipedes ou thecostraces. Masson et Cie, Paris, 472 pp.","Noll, F. C. (1872) Kochlorine hamata N., ein bohrende Cirripede. Bericht uber die Senckenbergische naturf. Gesellsch 1871 - 1872, 4, 50 - 58","Stubbings, H. G. (1967) The cirriped fauna of tropical West Africa. Bulletin of the British Museum (Natural History), Zoology, 15 (6), 229 - 319. https: // doi. org / 10.5962 / bhl. part. 27518"]}

Genus Weltneria Berndt, 1907 Weltneria Berndt, 1907: 289. Diagnosis. Females with five pairs of biramous terminal cirri, two-segmented caudal appendages. This is considered plesiomorphic, as this represents the ground pattern for all thecostracans (Kolbasov & Newman, 2005). Lateral bars weak or absent, orificial knob absent. Dwarf males pear-shaped, tapering towards the top, no lateral projections, attachment antennules with stalk absent (Chan, Hsieh & Kolbasov, 2014). Remarks. At present, there are 12 Weltneria species globally (Kolbasov, Chan & Cheng, 2017). Two of these occur in South Africa, of which W. spinosa is endemic., Published as part of Botha, Thomas P. A. & Griffiths, Charles L., 2021, South African Acrothoracica (Crustacea: Cirripedia), pp. 45-78 in Zootaxa 4949 (1) on page 55, DOI: 10.11646/zootaxa.4949.1.3, http://zenodo.org/record/4635765, {"references":["Berndt, W. (1907) Uber das System der Acrothoracica. Archiv fur Naturgeschichte, 73 (1), 287 - 289.","Kolbasov, G. A. & Newman, W. A. (2005) Revision of the Lithoglyptidae sensu Tomlinson, 1969 and Lithoglyptes Aurivillius, 1892 (Cirripedia, Acrothoracica), including a new species from Bermuda. Zootaxa, 1013 (1), 35 - 64. https: // doi. org / 10.11646 / zootaxa. 1013.1.3","Chan, B. K. K., Hsieh, W-P. & Kolbasov, G. A. (2014) Crustacean Fauna of Taiwan: Barnacles. Vol. III. Cirripedia: Acrothoracica. Biodiversity Research Center, Academia Sinica, Taipei, 107 pp.","Kolbasov, G. A., Chan, B. K. K. & Cheng, Y. R. (2017) Weltneria acanthostoma sp. nov., a burrowing barnacle (Cirripedia: Acrothoracica) from deep-waters of the South China Sea. Zootaxa, 4290 (3), 591 - 599. https: // doi. org / 10.11646 / zootaxa. 4290.3.12"]}

Order CRYPTOPHIALIDA Kolbasov, Newman & Høeg (Cited in Kolbasov, 2009) Cryptophialida Kolbasov, Newman & Høeg in Kolbasov, 2009: 241.— Chan, Hsieh & Kolbasov 2014: 2. Diagnosis. Mainly characterised by females having long, narrow-necked operculum, with small circular aperture and developed lateral and reinforcing bars, tongue-shaped labrum, no caudal appendages and special gastric mill at end of stomach. Rudimentary mouth cirri, thorax with one or two whip-like dorsal processes (Chan, Hsieh & Kolbasov, 2014). Posterior end of dwarf males with circular, cuticular ribs and frequently with mantle teeth; simple attachment antennules without stalk. Cypris larvae posses rudimentary thorax and thoracopods, carapace covers body incompletely. Burrow opening rounded and small. Remarks. Cryptophialida consists of the monotypic family Cryptophialidae., Published as part of Botha, Thomas P. A. & Griffiths, Charles L., 2021, South African Acrothoracica (Crustacea: Cirripedia), pp. 45-78 in Zootaxa 4949 (1) on page 48, DOI: 10.11646/zootaxa.4949.1.3, http://zenodo.org/record/4635765, {"references":["Kolbasov, G. A. (2009) Acrothoracica, burrowing crustaceans. KMK Scientific Press Ltd, Moscow, 452 pp.","Chan, B. K. K., Hsieh, W-P. & Kolbasov, G. A. (2014) Crustacean Fauna of Taiwan: Barnacles. Vol. III. Cirripedia: Acrothoracica. Biodiversity Research Center, Academia Sinica, Taipei, 107 pp."]}

Family LITHOGLYPTIDAE Aurivillius, 1892 Lithoglyptidae Aurivillius, 1892: 133.— Kolbasov 2009: 239.— Chan, Hsieh & Kolbasov 2014: 13. Diagnosis. Females with well-developed opercular bars, comb collar, thoracic lappets and mouth appendages; twosegmented protopod mouth cirri, biramous multi-annulated terminal cirri, caudal appendages present in some species, intestine with anus. Dwarf males pear-shaped, with pair of lobes at base of attachment antennules (Chan, Hsieh & Kolbasov, 2014). Remarks: Consist of three subfamilies: Berndtiinae, Lithoglyptinae and Kochlorininae, all present in South Africa., Published as part of Botha, Thomas P. A. & Griffiths, Charles L., 2021, South African Acrothoracica (Crustacea: Cirripedia), pp. 45-78 in Zootaxa 4949 (1) on page 55, DOI: 10.11646/zootaxa.4949.1.3, http://zenodo.org/record/4635765, {"references":["Aurivillius, C. W. S. (1892) Neue Cirripeden aus dem Atlantischen, Indischen und Stillen Ocean. Koniglich Vetenskapakademiens Forhandlingar, 3, 123 - 134.","Kolbasov, G. A. (2009) Acrothoracica, burrowing crustaceans. KMK Scientific Press Ltd, Moscow, 452 pp.","Chan, B. K. K., Hsieh, W-P. & Kolbasov, G. A. (2014) Crustacean Fauna of Taiwan: Barnacles. Vol. III. Cirripedia: Acrothoracica. Biodiversity Research Center, Academia Sinica, Taipei, 107 pp."]}

Weltneria spinosa Berndt, 1907 Figs 7–9 Weltneria spinosa Berndt, 1907: 289.— Tomlinson 1969: 32–36, fig. 1.— Tomlinson 1987: 65, fig. 2.— Kolbasov & Høeg 2007: 129, fig. 3 (A), fig. 5 (A, F), fig. 7 (A), fig. 8 (C), 1 fig. 9 (E), fig. 11 (E).— Kolbasov 2009: 284 (in Russian), fig. 2 (a), fig. 5 (ж), fig. 14 (е, ж), fig. 25 (г), fig. 28 (б), fig. 37 (a), fig. 39 (e), fig. 49 (a), fig. 51 (a), fig. 53 (a, e, ж, З, u), fig. 55 (в), fig. 56 (б, u), fig. 58 (б), fig. 60 (б), fig. 85. Deposited material. SAMC-A091113, False Bay, Western Cape, January and February 2017, 34 specimens from 6 Turbo sarmaticus shells (5 not deposited, used for genetic analyses). SAMC-A091114, False Bay, Western Cape, January and February 2017, 376 specimens from 8 Turbo sarmaticus shells (10 not deposited, used for dissections and light microscopy). SAMC-A091115, Chintsa West, Eastern Cape, April 2017, 25 specimens from 4 Turbo sarmaticus shells (2 not deposited, used for SEM). SAMC-A091116, Gonubie, Eastern Cape, April 2017, 105 specimens from 5 Turbo sarmaticus shells. SAMC-A091117, Gonubie, Eastern Cape, 27 April 2017, 12 specimens from Haliotis midae shell. SAMC-A091118, Gonubie, Eastern Cape, 27 April 2017, 223 specimens from Haliotis midae shell (3 not deposited, used for genetic analyses). SAMC-A091119, Melkbosstrand, Western Cape, 28 February 2017, 13 specimens from Haliotis midae shell. SAMC-A091120, Gonubie, Eastern Cape, 27 April 2017, 30 specimens from Ranella gemmifera shell. SAMC-A091121, Gonubie, Eastern Cape, 27 April 2017, 6 specimens from Ranella gemmifera shell. SAMC-A091122, False Bay, Western Cape, 19 February 2017, 1 specimen from Argobuccinum pustulosum shell (2 not deposited, used for SEM). SAMC-A091123, False Bay, Western Cape, 17 February 2017, 9 specimens from Argobuccinum pustulosum shell. SAMC-A091124, False Bay, Western Cape, 14 March 2017, 11 specimens from Argobuccinum pustulosum shell. SAMC-A091125, False Bay, Western Cape, 14 March 2017, 3 specimens from Argobuccinum pustulosum shell. SAMC-A091126, Melkbosstrand, Western Cape, 28 February 2017, 5 specimens from Turbo cidaris shell. SAMC-A091127, Melkbosstrand, Western Cape, February 2017, 30 specimens from 2 Turbo cidaris shells. SAMC-A091128, Chintsa West, Eastern Cape, 26 April 2017, 3 specimens from Dinoplax gigas. SAMC-A091129, False Bay, Western Cape, 15 March 2017, 2 specimens from Burnupena cincta shell. SAMC-A091130, East London surrounds (Gonubie, Chintsa West), Eastern Cape, April 2017, 8 specimens from 2 Mancinella capensis shells (2 not deposited, used for genetic analyses). SAMC-A091131, Gonubie, Eastern Cape, 27 April 2017, 51 specimens from Mancinella capensis shell. SAMC-A091132, Summerstrand, Eastern Cape, 4 July 2017, 6 specimens from Mancinella capensis shell. SAMC-A091133, L’Agulhas, Western Cape, 15 June 2010, 1 specimen in Heydrichia woelkerlingii. SAMC-A091134, Summerstrand, Algoa Bay, Eastern Cape, 4 July 2017, 30 specimens in Lithophyllum sp. SAMC-A091135, Morgan Bay, Eastern Cape, 13 July 2010, 12 specimens from Lithophyllum neoatalayense Masaki. SAMC-A091136, False Bay, Western Cape, 4 August 2016, 4 specimens from Scutellastra tabularis shell. Diagnosis. Females with pair of large hooks forming posterior processes of operculum, inner and outer margins of operculum with simple or bifid teeth. Also characterised by presence of pair of weak and irregular lateral bars. Description. Female, body oval-shaped (fig. 7B; 9A). Length 2.1–4.7 mm (mean = 2.98 mm), width 1.05–2.85 mm (mean = 1.69 mm). Opercular bars on average 0.87 mm long, with pair of large, curved posterior processes at top of operculum armed with several simple teeth and setae (fig. 8B, D; 9B). Both inner and outer margins of operculum lined with bifid or simple teeth and setae (fig. 8C, D, E). Rostral end of opercular bar with simple spines and setae (fig. 8E). Lateral surface of operculum lined with several teeth, setae and large multifid scales (fig. 8B, F). Comb collar consisting of long feather-like projections (fig. 8A). Developed orificial knob absent. Feeble lateral bars present running down mantle near operculum (fig. 9A). Row of small, mostly bifid teeth running down ventral surface on mantle. Females bright red when fresh, cirri also bright red (fig. 7A, C). Aperture resembles typed apostrophe, with one end having more pointed tapered slit (fig. 7C). Five pairs of terminal cirri, with pair of two-segmented caudal appendages (fig. 9C, D), two conical processes present (fig. 9C). Mouth cirri with four-segmented posterior ramus shorter than five-segmented anterior ramus, both with long setae (fig. 9G). Labrum saddle-shaped, mandibular palp trapezoid, with dense setae (fig. 9H). Mandible with three teeth, excluding inferior angle (fig. 9E), first tooth separated from second and third by large notch, inferior angle with several small spines and setae. Maxillule with single notch in middle of outer margin, two long cuspidate setae below notch, with three short, sharp setae above notch (fig. 9F). Male much smaller than female, average of 0.66 x 0.32 mm, with up to four on a single female. Hosts. The type-host is Haliotis midae. Also previously known from Argobuccinum pustulosum and Turbo sarmaticus (Tomlinson, 1969). New hosts in the Western Cape include Burnupena cincta, Dinoplax gigas, Scutellastra tabularis and Turbo cidaris. Collected in the Eastern Cape from two new hosts, Ranella gemmifera and Mancinella capensis, as well as from T. sarmaticus and H. midae. Also found inhabiting coralline red algae (Botha, Griffiths & Maneveldt, 2020) in the Eastern Cape in a rhodolith (Lithophyllum sp.), a discoid species, Lithophyllum neoatalayense, also from Heydrichia woelkerlingii in the Western Cape. Distribution. Type locality Hermanus (3425’1.62”S, 1914’55.46”E) in the Western Cape. Ranges from Melkbosstrand (3344’08.1”S, 1826’15.2”E) to Qolorha (3250’08.0”S, 2807’09.5”E). Endemic to Western and Eastern Cape provinces. All specimens were collected in water Remarks: Weltneria spinosa Berndt, 1907 is the type species of the genus and is the most common acrothoracican in South Africa, with up to 235 individuals found in a single Haliotis midae shell. Often found in the same host with Australophialus turbonis in the Western Cape and Australophialus utinomii and Kochlorine bocqueti in the Eastern Cape. In the Western Cape, empty burrows of this barnacle are often occupied by the unique amphipod Ampelisca excavata, which appears to be restricted to, and anatomically adapted, to live in this specialised niche (Gray & Barnard, 1970). We recorded this amphipod in W. spinosa holes on both T. sarmaticus and, in great abundance, on the coralline algae H. woelkerlingii., Published as part of Botha, Thomas P. A. & Griffiths, Charles L., 2021, South African Acrothoracica (Crustacea: Cirripedia), pp. 45-78 in Zootaxa 4949 (1) on pages 56-57, DOI: 10.11646/zootaxa.4949.1.3, http://zenodo.org/record/4635765, {"references":["Berndt, W. (1907) Uber das System der Acrothoracica. Archiv fur Naturgeschichte, 73 (1), 287 - 289.","Tomlinson, J. T. (1969) The burrowing barnacles (Cirripedia: Order Acrothoracica). Bulletin of the United States National Museum, 296, 1 - 162. https: // doi. org / 10.5479 / si. 03629236.296.1","Tomlinson, J. T. (1987) The burrowing barnacles (Acrothoracica). In: Southward, A. J. (Ed.), Barnacle Biology. Crustacean Issues. Vol. 5. Balkema, Rotterdam, pp. 63 - 71. https: // doi. org / 10.1201 / 9781315138053 - 3","Kolbasov, G. A. & Hoeg, J. T. (2007) Cypris larvae of acrothoracican barnacles (Thecostraca: Cirripedia: Acrothoracica). Zoologischer Anzeiger-A Journal of Comparative Zoology, 246 (2), 127 - 151. https: // doi. org / 10.1016 / j. jcz. 2007.03.001","Kolbasov, G. A. (2009) Acrothoracica, burrowing crustaceans. KMK Scientific Press Ltd, Moscow, 452 pp.","Botha, T. P., Griffiths, C. L. & Maneveldt, G. W. (2020) Coralline red algae - a new host taxon for burrowing barnacles (Cirripedia, Acrothoracica). Marine Biodiversity, 50 (1), 1 - 5. https: // doi. org / 10.1007 / s 12526 - 019 - 01038 - 7","Gray, W. S. & Barnard, J. L. (1970) South African Ampelisca excavata KH Barnard (Amphipoda, Gammaridea), a redescription with notes on the domicile. Crustaceana, 19 (1), 67 - 83. https: // doi. org / 10.1163 / 156854070 X 00644"]}

Kochlorine bihamata Noll, 1883 (deletion from fauna list) Kochlorine bihamata.— Barnard 1924: 99.— Tomlinson, 1969: 72.— Kolbasov, 2009: 320. Diagnosis. Females with no anterior conical processes, mantle aperture with two posterior hooks and spiny teeth on rim; size 5 mm (Tomlinson, 1969). Hosts. Collected as dried specimens observed in an Haliotis shell. Although not explicitly stated which Haliotis species is referred to, it can be assumed this is the abalone, Haliotis midae, which is known to host other acrothoracicans. Remarks. This species has not been observed since its description from Cape Town by Noll (1883) and there remains grave doubt regarding the validity of the species. Noll (1883) mentioned that: “Should new specimens of Kochlorine arise from Cape Town, one can well label it Kochlorine bihamata ”. Therefore, it is safe to assume that the genus Kochlorine occurs from Cadiz, Spain to Cape Town, South Africa and burrows into shells of Haliotis spp.. Although Tomlinson (1969) calls into question the validity of the species, he later states that K. bihamata should be treated as a valid species. However, following our survey no trace of this species was found, despite the type host being collected at the type locality and several adjacent localities. Thus, we consider that it is safe to assume that this is a nomen nudum., Published as part of Botha, Thomas P. A. & Griffiths, Charles L., 2021, South African Acrothoracica (Crustacea: Cirripedia), pp. 45-78 in Zootaxa 4949 (1) on page 69, DOI: 10.11646/zootaxa.4949.1.3, http://zenodo.org/record/4635765, {"references":["Barnard K. H. (1924) Contributions to the crustacean fauna of South Africa, 7. Cirripedia. Annals of the South African Museum, 20, 1 - 103.","Tomlinson, J. T. (1969) The burrowing barnacles (Cirripedia: Order Acrothoracica). Bulletin of the United States National Museum, 296, 1 - 162. https: // doi. org / 10.5479 / si. 03629236.296.1","Kolbasov, G. A. (2009) Acrothoracica, burrowing crustaceans. KMK Scientific Press Ltd, Moscow, 452 pp."]}

Weltneria cf. hirsuta (Tomlinson, 1963) Lithoglyptes hirsutus Tomlinson, 1963b: 299, figs 1–7. Utinomia newmani Tomlinson, 1963a: 265: 265, figs 1–5. Weltneria hirsuta Kolbasov, 2009: 280 (in Russian), fig. 8 (a), fig. 12 (a), fig. 14 (б), fig. 15 (a), fig. 17 (в), fig. 18 (e), fig. 26 (г), fig. 37 (б, в), fig. 39 (a, в), fig. 82.— Chan, Hsieh & Kolbasov 2014: 49, fig. 42, fig. 43. Deposited material: ELMC 0421, Kwelera Bay, Eastern Cape, 7 January 1986, 14 specimens on Charonia lampas shell (7 specimens not deposited, used for SEM, dissections and genetic analyses). Diagnosis. Weltneria with opercular bars that have numerous sharp, simple teeth and long setae, opercular bars terminating in tapered, conspicuous, spine-shaped posterior process. Description. Female, length 1.6–2.5 mm (mean = 2.05 mm) and width 1–1.7 mm (mean = 1.31 mm). Body oval-shaped, except opercular area flat surface, not rounded (fig. 10A; 12A). Opercular bars broad, concave on outer surface, average of 1.05 mm, with large upper surface and pair of posterior processes (fig. 11C, D), simple teeth along edges and down lateral surface of opercular area (fig. 11C, A, F). Occludent margins of opercular bars forming small, narrow aperture with almost no internal structures visible (fig. 11C). Row of simple and bifid teeth on either side of opercular bars, with several others randomly distributed on dorsal surface (Fig. 11A). Comb collar of long, feather-like projections (fig. 11B). Several bifid and trifid teeth on surface of mantle. Lateral surface below opercular bars with spines, setae and broad multifid scales, similar to those of W. spinosa (fig. 11E, F). Opercular knob and lateral bars absent. Mantle surface with several, randomly distributed, bifid teeth. Live colour unknown; dark brown/orange when preserved in ethanol (fig. 10A). Burrow oval-shaped, opercular bars well-fitted to burrow (fig. 10B). Terminal cirri five pairs with two-segmented caudal appendages, with setae at distal segment (fig. 12B, C). Mouth cirri with three-segmented posterior ramus shorter than four-segmented anterior ramus (fig. 12D, E). Both rami with setulated setae. Labrum saddle-shaped, bullate, upper edge convex, armed with developed dorsal process, while anterior edge horseshoe-shaped, smooth (fig. 12D). End of mandibular palp trapezoid, with dense simple setae as well as setae with small sparse setules on tip (Chan, Hsieh & Kolbasov, 2014). Mandible with three major teeth becoming smaller towards inferior angle, first upper tooth separated from the rest. Inferior angle with several denticles, inferior angle ending in two large denticles (Chan, Hsieh & Kolbasov, 2014). Maxillule with single notch, two large cuspidate and small setae above notch, more than five sharp setae at margin below notch (fig. 12F). Lateral surfaces with dense serrate setae. Maxilla sub-triangular with many simple setae on outer margin and apex (fig. 12E). Hosts. Originally found in two corals, Psammocora and Porites, in Hawaii, and shells of Bursa in Japan and Murex in Taiwan (Chan, Hsieh & Kolbasov, 2014). Found in South Africa in Charonia lampas from Kwelera Bay, collected in 1986 and stored in the East London Museum (ELMW0952). This is a new record for South Africa. Distribution. Type locality is Hawaii (Chan, Hsieh & Kolbasov, 2014). Also Japan, Taiwan and a single sample from South Africa at Kwelera Bay (3250’08.0”S, 2807’09.5”E), near East London, Eastern Cape. In South Africa only occuring in the Eastern Cape. Remarks. Live colour unknown. Some mouthparts could not be dissected out. Only 14 individuals were available from a single sample, collected in 1986 and held at the East London Museum. The individuals were in bad condition. Given that only a single, poorly-preserved South African sample, collected more than 30 years ago, was available for examination, and that this record is so geographically distant from the type locality, this identification should be confirmed using genetic methods and is recorded here as ‘cf’. It is possible that this is a cryptic new species., Published as part of Botha, Thomas P. A. & Griffiths, Charles L., 2021, South African Acrothoracica (Crustacea: Cirripedia), pp. 45-78 in Zootaxa 4949 (1) on pages 60-62, DOI: 10.11646/zootaxa.4949.1.3, http://zenodo.org/record/4635765, {"references":["Tomlinson, J. T. (1963 b) Lithoglyptes hirsutus (Cirripedia: Acrothoracica), a new burrowing barnacle from Hawaii. Pacific Science, 17, 299 - 301.","Tomlinson, J. T. (1963 a) Two new acrothoracican cirripeds from Japan. Publications of the Seto Marine Biology Laboratory, 11, 263 - 280. https: // doi. org / 10.5134 / 175342","Kolbasov, G. A. (2009) Acrothoracica, burrowing crustaceans. KMK Scientific Press Ltd, Moscow, 452 pp.","Chan, B. K. K., Hsieh, W-P. & Kolbasov, G. A. (2014) Crustacean Fauna of Taiwan: Barnacles. Vol. III. Cirripedia: Acrothoracica. Biodiversity Research Center, Academia Sinica, Taipei, 107 pp."]}

Family CRYPTOPHIALIDAE Gerstaecker, 1866 Cryptophialidae Gerstaecker, 1866: 534. Diagnosis. As for Cryptophialida. Remarks. Contains the genera Australophialus Tomlinson, 1969 and Cryptophialus Darwin, 1854 with five and 16 species, respectively. In South Africa, only Australophialus observed., Published as part of Botha, Thomas P. A. & Griffiths, Charles L., 2021, South African Acrothoracica (Crustacea: Cirripedia), pp. 45-78 in Zootaxa 4949 (1) on page 48, DOI: 10.11646/zootaxa.4949.1.3, http://zenodo.org/record/4635765, {"references":["Gerstaecker, A. (1866 - 1879) Arthropoda. In: Winter, C. F. (Ed.), Bronn's Klassen und Ordnungen des Tierreichs. Vol. 5. C. F. Winter, Leipzig und Heidelberg, pp. 406 - 589.","Tomlinson, J. T. (1969) The burrowing barnacles (Cirripedia: Order Acrothoracica). Bulletin of the United States National Museum, 296, 1 - 162. https: // doi. org / 10.5479 / si. 03629236.296.1","Darwin, C. (1854) A monograph on the sub-class Cirripedia. Vol. II. The Balanidae. In: Barrett, P. & Freeman, R. B. (Eds.), The Works of Charles Darwin. Vol. 12. New York University Press, Washington Square, New York, pp. vii - viii + 1 - 684, figs. 1 - 7."]}

Order LITHOGLYPTIDA Kolbasov, Newman & Høeg (Cited in Kolbasov, 2009) Lithoglyptida Kolbasov, Newman & Høeg in Kolbasov, 2009: 239.— Chan, Hsieh & Kolbasov 2014: 12. Diagnosis. Females with wide aperture, sac-like mantle, operculum without a neck, with opercular bar less than aperture length, well-developed mouth cirri, large, saddle-like labrum and lack special gastric mill at end of stomach (Kolbasov, 2009; Chan, Kolbasov & Cheang, 2012; Chan, Hsieh & Kolbasov, 2014). Thorax without long dorsal processes. Burrow shape elongated. Males have surfaces with several cuticular denticles and lack mantle teeth. Cypris larvae with developed thorax and thoracopods with an unperforated carapace with fronto-lateral pores. Remarks. Consists of two families, the Lithoglyptidae and Trypetesidae. No members of the Trypetesidae known from South Africa (although reported from south-west of Madagascar)., Published as part of Botha, Thomas P. A. & Griffiths, Charles L., 2021, South African Acrothoracica (Crustacea: Cirripedia), pp. 45-78 in Zootaxa 4949 (1) on page 55, DOI: 10.11646/zootaxa.4949.1.3, http://zenodo.org/record/4635765, {"references":["Kolbasov, G. A. (2009) Acrothoracica, burrowing crustaceans. KMK Scientific Press Ltd, Moscow, 452 pp.","Chan, B. K. K., Hsieh, W-P. & Kolbasov, G. A. (2014) Crustacean Fauna of Taiwan: Barnacles. Vol. III. Cirripedia: Acrothoracica. Biodiversity Research Center, Academia Sinica, Taipei, 107 pp.","Chan, B. K. K., Kolbasov, G. A. & Cheang, C. C. (2012) Cryptic diversity of the acrothoracican barnacle Armatoglyptes taiwanus in the Indo-Pacific waters, with description of a new species from the Mozambique Channel collected from the MAIN- BAZA cruise. Zoosystema, 34 (1), 5 - 20. https: // doi. org / 10.5252 / z 2012 n 1 a 1"]}

Balanodytes flexuosus (Chan, Kolbasov & Cheang, 2012) Figs 13, 14 Material examined. CLP14-4, Port Shepstone, Eastern Cape, August 2017, 1 specimen in Charonia lampas shell (specimen used for dissection). Diagnosis. Balanodytes with opercular bars with bifid teeth, posterior processes L-shaped, strongly bent/recurved at tip. Description. Female, mantle sac oval-shaped, narrowing below to opercular bars (figs 13, 14A). Length 1.65 mm, width 1.05 mm, opercular bars 500 µm in length, with row of bifid teeth and setae (fig. 14B). Posterior processes hook-like (anteriorly bent/recurved), L-shaped, with bifid teeth and setae along length (Figs 13, 14B). Comb collar with long, feather-like projections, fused at bases. Orificial knob distinct, with setae and simple or bifid teeth (fig. 14B). Lateral area below opercular bar covered with sparse, simple setae and rows of large, multifid scales. Feeble lateral bars. Burrow opening narrow, oval-shaped. Four pairs of terminal cirri and pair of two-segmented caudal appendages (fig. 14C, D). Mouth cirri with both rami three-segmented, with anterior ramus shorter than posterior ramus, both rami with plumose setae (fig. 14H). Labrum saddle-like, with small, blunt teeth on anterior margin (fig. 14E). Mandible with three large teeth, two smaller teeth close to lower margin, large notch between first and second teeth. Inferior angle with two sharp denticles and setae (fig. 14F). Maxillule with two long, sharp upper and single, smaller lower cuspidate setae. Notch with short, sharp seta, lower 2/3 of cutting edge with numerous short, sharp setae (fig. 14G). Maxilla triangular with setae on both exterior and interior margins. Mandibular palp trapezoidal, with simple setae along exterior margin. Hosts. Previously found in the coral Pavona sp. from Mozambique Channel (Chan, Kolbasov & Cheang, 2012). Collected in South Africa from Charonia lampas shell. Distribution. Originally described from Mozambique Channel (2611’00”S, 3501’00”E). Collected in South Africa only in Port Shepstone (3044’38.8”S, 3027’28.7”E). This is the first record of this species from South Africa. Remarks. Only one individual collected, found in association with Balanodytes sp. A, Published as part of Botha, Thomas P. A. & Griffiths, Charles L., 2021, South African Acrothoracica (Crustacea: Cirripedia), pp. 45-78 in Zootaxa 4949 (1) on pages 63-64, DOI: 10.11646/zootaxa.4949.1.3, http://zenodo.org/record/4635765, {"references":["Chan, B. K. K., Kolbasov, G. A. & Cheang, C. C. (2012) Cryptic diversity of the acrothoracican barnacle Armatoglyptes taiwanus in the Indo-Pacific waters, with description of a new species from the Mozambique Channel collected from the MAIN- BAZA cruise. Zoosystema, 34 (1), 5 - 20. https: // doi. org / 10.5252 / z 2012 n 1 a 1"]}

The beaked clam Eumarcia paupercula (Holten 1802) (Bivalvia: Veneridae) is an important fishery resource for local artisanal fishers in Maputo Bay. Its annual reproductive cycle was described by following seasonal fluctuations in the condition index of the population occupying a tidal flat, and by analysing fresh gonad smears to confirm that changes in condition were, in fact, due to gonad state. Macroscopic gonad observations and changes in body condition confirmed that E. paupercula is a year-round breeder with three spawning peaks, with the major spawning periods occurring during summer. The spawning pattern found in this study is similar to other clams inhabiting similar tropical ecosystems.

Genus Calvadosia Clark, 1863 Remarks. Calvadosia was originally proposed by Clark (1863) to accommodate a species described by Lamouroux (1815), ��� Lucernaire campanul��e ��� (or Lucernaria campanulata), from Calvados, France, therefore proposing the name Calvadosia campanulata (Lamouroux, 1815). Its main difference from other Lucernaria is the ���four pilasters [���] not muscular, as are the pilasters in the pedicel of Lucernaria quadricornis ��� (Clark 1863: 556), i.e., absence of interradial longitudinal muscles associated with the septa of the peduncle. However, Clark���s (1863) proposal was overlooked for many years. Later, Uchida (1929) proposed a new genus, Lucernariopsis, for the same ��� Lucernaria campanulata ���, including species with one-chambered peduncle without muscles, overlooking the availability of the older name Calvadosia Clark, 1863. Apparently, Gwilliam (1956: 10) was the only author to notice this nomenclatural issue, concluding that according to the ���law of priority, the proper generic name of Lucernariopsis Uchida, 1929 is Calvadosia Clark, 1863 ���, but he never published his PhD Dissertation on the taxonomy of the Stauromedusae. More recently, Lucernariopsis Uchida, 1929 was officially recognized as a synonym of Calvadosia Clark, 1863 (Miranda et al. 2016b). In addition, based on molecular and morphological evidence, the former genera Kishinouyea Mayer, 1910 and Sasakiella Okubo, 1917 were also incorporated into Calvadosia (Miranda et al. 2016b). Therefore, Calvadosia is currently one of the most diverse genera in Staurozoa, with 11 species: Calvadosia campanulata (Lamouroux, 1815), Calvadosia nagatensis (Oka, 1897), Calvadosia vanhoeffeni (Browne, 1910), Calvadosia cruciformis (Okubo, 1917), Calvadosia hawaiiensis (Edmondson, 1930), Calvadosia tsingtaoensis (Ling, 1937), Calvadosia capensis (Carlgren, 1938), Calvadosia cruxmelitensis (Corbin, 1978), Calvadosia corbini (Larson, 1980), Calvadosia tasmaniensis (Zagal, Hirano, Mills, Edgar & Barrett, 2011), and Calvadosia lewisi sp. nov. described in this study., Published as part of Miranda, Luc��lia S., Branch, George M., Collins, Allen G., Hirano, Yayoi M., Marques, Antonio C. & Griffiths, Charles L., 2017, Stalked jellyfishes (Cnidaria: Staurozoa) of South Africa, with the description of Calvadosia lewisi sp. nov., pp. 369-389 in Zootaxa 4227 (3) on pages 371-372, DOI: 10.11646/zootaxa.4227.3.5, http://zenodo.org/record/268342, {"references":["Clark, H. J. (1863) Prodromus of the history, structure, and physiology of the order Lucernariae. Journal of the Boston Society of Natural History, 7, 531 - 567.","Lamouroux, J. V. F. (1815) Memoire sur la Lucernaire campanulee. Memoirs du Museum d' Histoire Naturelle, II, 460 - 473.","Uchida, T. (1929) Studies on the Stauromedusae and Cubomedusae, with special reference to their metamorphosis. Japanese Journal of Zoology, 2, 103 - 193.","Gwilliam, G. F. (1956) Studies on West Coast Stauromedusae. PhD. Dissertation. University of California, Berkeley, 192 pp.","Miranda, L. S., Hirano, Y. M., Mills, C. E., Falconer, A., Fenwick, D., Marques, A. C. & Collins, A. G. (2016 b) Systematics of stalked jellyfishes (Cnidaria: Staurozoa). PeerJ, 4, e 1951.","Mayer, A. G. (1910) Medusae of the world. Volume III. Scyphomedusae. Carnegie Institution Publishing, Washington, Publication 109, pp. 499 - 735.","Okubo, T. (1917) Preliminary note on a new genus of Stauromedusae from Hokkaido. Zoological Magazine, 29, 317 - 322.","Oka, A. (1897) Sur une nouvelle espece Japonaise du genre Lucernaria. Annotationes Zoologicae Japonenses, 1, 141 - 145.","Browne, E. T. (1910) Reports of the natural history results of the voyage of the SS Discovery in the Antarctic regions in 1901, under Captain R F Scott RN. Coelentera. V. Medusae. British Museum (Natural History), London, 62 pp.","Edmondson, C. H. (1930) New Hawaiian medusae. Bernice P. Bishop Museum Occasional Papers, 9, 1 - 16.","Ling, S. W. (1937) Studies on Chinese Stauromedusae. I. Stauromedusae from Tsingtao. Amoy Marine Biological Bulletin, 3, 1 - 35.","Carlgren, O. (1938) Eine neue sudafrikanische Lucernariidae, Lucernariopsis capensis. Kungliga Fysiografiska Sallskapets i Lund Forhandlingar, 8, 1 - 6.","Corbin, P. G. (1978) A new species of the stauromedusan genus Lucernariopsis (Coelenterata: Scyphomedusae). Journal of the Marine Biological Association of the United Kingdom, 58, 285 - 290.","Larson, R. J. (1980) A new stauromedusa, Kishinouyea corbini (Scyphozoa, Stauromedusae) from the tropical western Atlantic. Bulletin of Marine Science, 30, 102 - 107.","Zagal, C. J., Hirano, Y. M., Mills, C. E., Edgar, G. J. & Barrett, N. S. (2011) New records of Staurozoa from Australian coastal waters, with a description of a new species of Lucernariopsis Uchida, 1929 (Cnidaria, Staurozoa, Stauromedusae) and a key to Australian Stauromedusae. Marine Biology Research, 7, 651 - 666."]}

Lipkea stephensoni Carlgren, 1933 (Fig. 11) Lipkea stephensoni Carlgren, 1933: 1 ���15, figures 1���14; Carlgren 1935: 19, 23; Kramp 1961: 299; Grohmann et al. 1999: 386; Pisani et al. 2007: 7; Zagal et al. 2011: 652, 663, 664; Miranda et al. 2016b: 20, 26, 32, 38. Material examined. Iziko South African Museum MB-A083794, 2 specimens, Onrust River, Overberg, Western Cape, South Africa (-34.4196, 19.1801), January 1992, intertidal pools, under rock, ethanol 70%, col. & det. C. Griffiths. Not kept, 2 specimens, Smitswinkel Bay, False Bay, Cape Town, Western Cape, South Africa (-34.2743, 18.4728), between 1990���2003, depth 3���4 m, observed by G. Spiby. Not kept, 3 specimens, False Bay (between Partridge Point and Castle Rocks), Cape Town, Western Cape, South Africa (-34.2457, 18.4795), April 2012, depth 23 m, observed and photographed by G. Jones. Information on type material. The holotype is probably S185, from Still Bay, Eden, Western Cape, South Africa (type locality), found in pool under stones. The material is listed in the ���University of Cape Town Ecological Survey��� but could not be located in the Iziko South African Museum���s collection. Description. (complemented with Carlgren 1933). Calyx elongated, funnel-shaped (Fig. 11). Peduncle singlechambered, with four interradial longitudinal muscles. Four interradial septa visible in calyx (Fig. 11 D, F, G). Variable number of marginal lappets (arms?), with reduced (rudimentary) tentacles in one row along their margin (Fig. 11 A���C), although some specimens with smooth marginal lappets (Fig. 11 D���G). Gastrovascular cavity not divided by claustrum. Manubrium with four perradial lips (Fig. 11 B, D, G). Numerous gastric filaments in gastrovascular cavity. Gonads embedded in gastrovascular cavity of subumbrella, not extending into marginal lappets (confined to basal part of calyx) (Fig. 11 B, D, G). Perradial and interradial anchors absent. Pad-like adhesive structures absent. Coronal muscle entire (Fig. 11 D, E). White spots of nematocysts on subumbrellar surface, margin of calyx, and marginal lappets (Fig. 11). General color of body pinkish white (Fig. 11). Total body length about 8.0 to 16.0 mm. Distribution and habitat. The type locality for L. stephensoni is Still Bay, Eden, Western Cape, South Africa (Carlgren 1933). These new records are also in the Western Cape: Onrust River, Overberg; between Partridge Point and Castle Rocks, False Bay, Cape Town; Smitswinkel Bay, False Bay, Cape Town (Fig. 5 F). The species was recorded from intertidal pools to 23 m deep, attached to rocks. Remarks. There are three valid species in the genus Lipkea: Lipkea ruspoliana Vogt, 1886, Lipkea sturdzii (Antipa, 1893), and Lipkea stephensoni Carlgren, 1933. Lipkea ruspoliana was originally recorded from a single specimen from Alghero, on the Sardinian coast of the Mediterranean Sea (Vogt 1886, 1887). The Sardinian specimen was defined by its peculiar morphology, with eight marginal lappets in the perradii and interradii with mucous glands, but without tentacles (Vogt 1886, 1887). New specimens recently found in aquaria of the Oceanographic Museum of Monaco have 8���12 adradial lappets (Pisani et al. 2007), raising questions about intraspecific variation and the homology of these structures with arms and primary tentacles/anchors (Miranda et al. 2016b). Capria sturdzii Antipa, 1893 was described based on one specimen from Capri Island, Gulf of Naples, Italy, being the only species of the genus Capria and the family Capriidae (Antipa 1893). Carlgren (1933) synonymized Lipkea and Capria, proposing the name Lipkea sturdzii (see also Kramp 1961). Lipkea sturdzii has not been observed since its original description, but it differs from L. ruspoliana by the presence of a row of 16���20 toothlike or short finger-shaped rudimentary tentacles, which are fused one to another by a web (Antipa 1893; Mayer 1910). Finally, Lipkea stephensoni is the only formally described Lipkea species outside of Europe. The species was also described based on a single specimen from Still Bay, Eden, South Africa (Carlgren 1933), characterized by eight adradial short lappets with 30���40 reduced tentacles in one row along their margin (Fig. 11 A���C). However, the additional specimens that we observed show that the number of marginal lappets is variable in L. stephensoni (Fig. 11; as seems to be common in Lipkea species, Pisani et al. 2007; Zagal et al. 2011), as is the presence of the rudimentary marginal tentacles (Fig 11 C, E). The original description of L. stephensoni was based on a specimen found in the intertidal zone (Carlgren 1933), but photographs of not-kept L. stephensoni stauromedusae revealed that specimens from deeper water (about 23 m deep) have rudimentary tentacles in the lappets (Fig. 11 A���C), whereas intertidal and subtidal specimens (Fig. 11 D���G) have smooth marginal lappets, similar to those of L. ruspoliana, indicating either an intraspecific variation in L. stephensoni or the existence of an additional (new) species, a hypothesis that cannot be tested until more specimens become available. Indeed, the identification of Lipkea species is difficult (Zagal et al. 2011), based on subtle differences (presence or absence, and number and distribution of reduced tentacle-like structures along the margin of each lappet, Zagal et al. 2011) that have never been tested in a phylogenetic context. In addition, there is little information on intraspecific variation (Vogt 1886, 1887; Pisani et al. 2007) and no information on morphological changes during development. As a consequence, there are numerous unidentified records around the world, including specimens from Australia (Zagal et al. 2011), New Zealand (Cairns et al. 2009), and Japan (Miranda et al. 2016b). Australian specimens seem to have a similar morphology to L. stephensoni, with a single row of 18���27 reduced tentacles along the margin of the lappets (Carlgren 1933; Zagal et al. 2011), although, as discussed, it is equivocal whether the presence of rudimentary tentacles should be considered a diagnostic character (Uchida 1929; Zagal et al. 2011). Therefore, the validity of described and possibly new Lipkea species remains to be vigorously tested in further investigations., Published as part of Miranda, Luc��lia S., Branch, George M., Collins, Allen G., Hirano, Yayoi M., Marques, Antonio C. & Griffiths, Charles L., 2017, Stalked jellyfishes (Cnidaria: Staurozoa) of South Africa, with the description of Calvadosia lewisi sp. nov., pp. 369-389 in Zootaxa 4227 (3) on pages 384-386, DOI: 10.11646/zootaxa.4227.3.5, http://zenodo.org/record/268342, {"references":["Carlgren, O. (1933) Zur Kenntnis der Lucernariiden Lipkea, Capria und Brochiella. Kungliga Fysiografiska Sallskapets Handlingar, 44, 1 - 19.","Carlgren, O. (1935) Uber eine neue Sudafrikanische Lucernariidae, Depastromorpha africana n. gen., n. sp., nebst Bemerkungen uber den Bau und die Systematik dieser Tiergruppe. Kungliga Svenska Vetenskapsakademiens Handlingar, 15, 1 - 24.","Kramp, P. L. (1961) Synopsis of the medusae of the world. Journal of the Marine Biological Association of the United Kingdom, 40, 292 - 303.","Grohmann, P. A., Magalhaes, M. P. & Hirano, Y. M. (1999) First record of the order Stauromedusae (Cnidaria, Scyphozoa) from the tropical southwestern Atlantic, with a review of the distribution of Stauromedusae in the southern hemisphere. Species Diversity, 4, 381 - 388.","Pisani, V., Otero-Ferrer, F., Lotto, S., Maurel, P. & Goy, J. (2007) Lipkea ruspoliana Vogt, 1887 (Stauromedusae, Scyphozoa, Cnidaria) dans les aquariums du Musee Oceanographique de Monaco. Bulletin de la Societe Zoologique de France, 132, 183 - 190.","Zagal, C. J., Hirano, Y. M., Mills, C. E., Edgar, G. J. & Barrett, N. S. (2011) New records of Staurozoa from Australian coastal waters, with a description of a new species of Lucernariopsis Uchida, 1929 (Cnidaria, Staurozoa, Stauromedusae) and a key to Australian Stauromedusae. Marine Biology Research, 7, 651 - 666.","Miranda, L. S., Hirano, Y. M., Mills, C. E., Falconer, A., Fenwick, D., Marques, A. C. & Collins, A. G. (2016 b) Systematics of stalked jellyfishes (Cnidaria: Staurozoa). PeerJ, 4, e 1951.","Vogt, C. (1886) Sur une medusaire sessile, Lipkea ruspoliana. Archives des Sciences, 16, 356 - 362.","Antipa, G. (1893) Eine neue Stauromeduse (Capria n. sturdzii n.). Mittheilungen aus der Zoologischen Station zu Neapel, 10, 618 - 632.","Vogt, C. (1887) Sur un nouveau genre de medusaire sessile Lipkea ruspoliana c. v. Imprimerie Centrale Genevoise, Geneve, 53 pp.","Mayer, A. G. (1910) Medusae of the world. Volume III. Scyphomedusae. Carnegie Institution Publishing, Washington, Publication 109, pp. 499 - 735.","Cairns, S. D., Gershwin, L. - A., Brook, F. J., Pugh, P., Dawson, E. W., Ocana, V. O., Vervoort, W., Williams, G., Watson, J. E., Opresko, D. M., Schuchert, P., Hine, P. M., Gordon, D. P., Campbell, H. J., Wright, A. J., Sanchez, J. A. & Fautin, D. G. (2009) Phylum Cnidaria: corals, medusae, hydroids, myxozoans. In: Gordon, D. P. (Ed.), New Zealand Inventory of Biodiversity, Volume 1. Canterbury University Press, Christchurch, pp. 59 - 101.","Uchida, T. (1929) Studies on the Stauromedusae and Cubomedusae, with special reference to their metamorphosis. Japanese Journal of Zoology, 2, 103 - 193."]}

Calvadosia lewisi sp. nov. (Figs 1���4) Kishinouyea sp. South Africa ��� Miranda et al. 2016b: 8, 12���15, figures 3���5. Calvadosia sp. 4 South Africa ��� Miranda et al. 2016b: 17, 34, 36, figures 7, 16. Material examined. Holotype: MZUSP 3415, 1 specimen, Simon���s Town, Cape Town, Western Cape, South Africa (���34.2100, 18.4626), 0 1 December 2014, depth 3 m, on Sargassum, formaldehyde solution 4%, col. C. Foster. Paratype: MZUSP 3416, 1 specimen, Simon���s Town, Cape Town, Western Cape, South Africa (���34.2100, 18.4626), 0 1 December 2014, depth 3 m, on Sargassum, formaldehyde solution 4%, col. C. Foster. Additional material: MZUSP 2731, 1 specimen, Miller���s Point, Cape Town, Western Cape, South Africa (���34.2320, 18.4745), February 2013, subtidal, on Brassicophycus brassicaeformis and Anthophycus longifolius, ethanol 90%, col. D. Robertson-Anderson. MZUSP 2732, 1 specimen, Miller���s Point, Cape Town, Western Cape, South Africa (��� 34.2320, 18.4745), February 2013, subtidal, on Brassicophycus brassicaeformis and Anthophycus longifolius, ethanol 90%, col. D. Robertson-Anderson. MZUSP 3417, 1 specimen, Miller���s Point, Cape Town, Western Cape, South Africa (���34.2320, 18.4745), February 2013, subtidal, on Brassicophycus brassicaeformis and Anthophycus longifolius, ethanol 90%, col. D. Robertson-Anderson. CMNH-ZG7819, 1 specimen, A-Frame, False Bay, Cape Town, Western Cape, South Africa (���34.2159, 18.4650), 19 January 2003, shallow subtidal (depth 2���3 m), on kelp, formaldehyde solution 5% and transferred to ethanol 70%, col. Y. Hirano. Iziko South African Museum MB- A083793, 1 specimen, Miller���s Point, Cape Town, Western Cape, South Africa (���34.2320, 18.4745), February 2013, subtidal, on Brassicophycus brassicaeformis and Anthophycus longifolius, ethanol 80%, col. D. Robertson- Anderson. Iziko South African Museum MB-A084062, 1 specimen, Betty���s Bay, Overberg, Western Cape, South Africa (���34.3723, 18.8875), May 2013, depth 1 m, on kelp Ecklonia maxima, formaldehyde solution 4%, col. E. Firl & C. Pickering. Not kept, 1 specimen, Castle Rocks, False Bay, Cape Town, Western Cape, South Africa (��� 34.2385, 18.4766), February 2010, depth about 5 m; on wrack growing on flat rock, observed by G. Zsilavecz. Not kept, 1 specimen, A-Frame, False Bay, Cape Town, Western Cape, South Africa (���34.2159, 18.4650), April 2014, depth 5 m, on Caulerpa filiformis growing on sand; observed by G. Zsilavecz. Diagnosis. Gonadal curved nodular lobes, regular (symmetric) and smooth in shape, facing interradii, arranged in a ���zigzag��� row on subumbrella. Description. Body divided into two clearly demarcated regions: calyx and peduncle (Fig. 1 D). Calyx wider than high, cruciform (Fig. 1), height 3.11���6.15 mm (mean 4.64 mm, number of individuals measured n = 5), width 5.88���16.48 mm (mean 12.03 mm, n = 5). Peduncle short (Figs 1 D, 2C), about 1/4 of the total height, 1.20���1.32 mm tall (mean 1.26 mm, n = 5), width 1.39���2.03 mm (mean 1.85 mm, n = 5). Broad, swollen adhesive circular pedal disk at base of peduncle (Fig. 2 C, E), width 2.48���4.31 mm (mean 3.38 mm, n = 5). Small central pore in the pedal disk of some specimens (Fig. 2 E). Peduncle without interradial longitudinal muscle bands, with a single cruciform chamber (in cross-section) that becomes four-chambered basally within the pedal disk (Fig. 2 D). Calyx without anchors (rhopalioids) or primary tentacles (Fig. 1). Manubrium with four perradial pleated lips (Figs 1 E; 2B, F). Many gastric filaments in gastrovascular cavity. Gastrovascular cavity not divided by vertical tissue composed of double layer of gastrodermis with internal mesoglea, known as claustrum. Eight arms (width 1.00��� 1.55 mm, mean 1.27 mm, n = 5), organized in four interradial pairs, resembling a cross (Figs 1, 2), hence the common name ���cross of pearls���. Four U-shaped perradial notches about four times as deep as the U- or V-shaped interradial notches (Fig. 2 A). Eight gonads not embedded in gastrovascular cavity, but contained within evaginations from the gastrovascular cavity. Each gonad consisting of several nodular lobes, relatively regular and smooth in shape, and arranged in a single row on subumbrella (Figs 1���3). However, in mature specimens nodules are tightly packed, giving the impression of two rows of nodular lobes (but actually in a ���zigzag��� row; see Fig. 3 D). Regular nodular lobes curved, facing interradii (Fig. 3 A, E). Each arm with 8���12 nodular lobes (n = 5). Gonads extending from manubrium to tips of arms (Figs 1, 2 B). Each arm with a terminal cluster of secondary tentacles, each cluster with 12���27 (n = 5) hollow, knobbed tentacles, similar in shape (Fig. 2 J). Broad pad-like adhesive structures forming a glandular cushion (height 0.46���1.02 mm, mean 0.69 mm, n = 5; width 1.37���3.53 mm, mean 2.39 mm, n = 5) on tips of arms, on exumbrella, surrounding outermost tentacles near their base (Fig. 2 I). Four interradial longitudinal muscles at calyx base, each divided in pyloric region into two bands toward adradial arms (Fig. 2 G, H). Coronal muscle divided into eight segments by arms (Fig. 2 H). Exumbrella finely granulated with nematocyst (euryteles) warts (Fig. 4 D). Conspicuous white nematocyst spots on subumbrella, distributed along calyx margin, at interradii and perradii, to tips of arms (Figs 1 A, E; 3B, C). Higher concentration of white spots at perradii, organized in 1���4 rows. White spots also associated with gonads. General color of body reddish to greenish brown (Fig. 1). Cnidome. Secondary tentacles with two types of nematocysts: isorhiza (abundant), length 18.02���19.65 ��m (mean 18.72 ��m, number of capsules measured n = 10), diameter 2.12���3.16 ��m (mean 2.61 ��m, n = 10); and eurytele (scarce), length 14.18���15.96 ��m (mean 15.27 ��m, n = 10), diameter 5.28���8.04 ��m (mean 6.85 ��m, n = 10) (Fig. 4 A, B). White nematocyst spots with one type of nematocysts: rhopaloids (abundant), length 12.77���15.40 ��m (mean 14.29 ��m, n = 10), diameter 10.20���11.96 ��m (mean 11.34 ��m, n = 10) (Fig. 4 C). Etymology. Named after Lewis Jason, a legendary volunteer at the Two Oceans Aquarium (Cape Town, South Africa) who first brought our attention to this animal and made passionate and lengthy notes on the species. Lewis Jason passed away on December 17th, 2014, at the age of 90. Type locality. Simon���s Town, Cape Town, Western Cape, South Africa. Distribution and habitat. Calvadosia lewisi was found at various localities in the Western Cape, South Africa: A-Frame, False Bay, Cape Town; Betty���s Bay, Overberg; Castle Rocks, False Bay, Cape Town; Miller���s Point, Cape Town; and Simon���s Town, Cape Town (Fig. 5 C). The species is found from intertidal to shallow subtidal regions, up to 5 m deep, attached to different species of algae. Remarks. Molecular phylogenetic analyses of Staurozoa (Miranda et al. 2016b) revealed a putative new species in South Africa, referred to Calvadosia sp. 4 in that study, and herein properly described as Calvadosia lewisi sp. nov. This species is closely related to C. tasmaniensis and C. corbini, in a clade whose possible synapomorphy is a broad pad-like adhesive structure on the tip of each arm (Miranda et al. 2016b). This feature is also present in C. hawaiiensis (Edmondson 1930; Grohmann et al. 1999) and in C. capensis (Carlgren 1938; Miranda et al. 2012b), suggesting that they too may belong to this clade (Miranda et al. 2016b). Calvadosia cruxmelitensis has a slightly different pad-like adhesive structure on the tip of the arm, in which the secondary tentacles arise directly from this structure (Corbin 1978; Miranda et al. 2016b). The morphology of C. lewisi is similar to C. tasmaniensis and C. corbini, in accordance with molecular results (Miranda et al. 2016b). Calvadosia tasmaniensis has been recorded only from Australia (Zagal et al. 2011) and C. corbini was originally described from Puerto Rico (Larson 1980), subsequently recorded from Brazil (Grohmann et al. 1999) and Mexico (Lechuga & Alamo 2005). These species have a wide ���open��� calyx, cruciform in C. lewisi and in C. corbini (Figs 1, 6) (Larson 1980; Grohmann et al. 1999), with interradial pairing of arms, a feature not evident in C. tasmaniensis (Zagal et al. 2011). These three species also share a relatively short peduncle (Larson 1980; Grohmann et al. 1999; Zagal et al. 2011) and broad, pad-like adhesive structures on the tips of the arms (Figs 2 I; 6E; 7D). In addition, the gonads of these species are not embedded in the gastrovascular cavity, but are nodular evaginations from the gastrovascular cavity resting on the subumbrella (Figs 1���3, 6, 7) (Larson 1980; Grohmann et al. 1999; Zagal et al. 2011). The generally smooth shape of the nodular gonads is the hypothetical main distinguishing feature of C. lewisi. Analyzed specimens of C. corbini and C. tasmaniensis (Table 2) have nodular erect gonads, with irregular shape (described by Larson (1980) for C. corbini as looking like ���small raisins���), which can have many evaginations, and are distributed in a single, relatively straight row (Figs 6, 7). However, the nodular gonads in C. lewisi have a smooth, regular shape, curved toward the interradii, and are tightly arranged in a ���zigzag��� row (Figs 1���3). Our observations include specimens of different sizes, making it unlikely that there is ontogenetic variation of this feature., Published as part of Miranda, Luc��lia S., Branch, George M., Collins, Allen G., Hirano, Yayoi M., Marques, Antonio C. & Griffiths, Charles L., 2017, Stalked jellyfishes (Cnidaria: Staurozoa) of South Africa, with the description of Calvadosia lewisi sp. nov., pp. 369-389 in Zootaxa 4227 (3) on pages 372-376, DOI: 10.11646/zootaxa.4227.3.5, http://zenodo.org/record/268342, {"references":["Miranda, L. S., Hirano, Y. M., Mills, C. E., Falconer, A., Fenwick, D., Marques, A. C. & Collins, A. G. (2016 b) Systematics of stalked jellyfishes (Cnidaria: Staurozoa). PeerJ, 4, e 1951.","Edmondson, C. H. (1930) New Hawaiian medusae. Bernice P. Bishop Museum Occasional Papers, 9, 1 - 16.","Grohmann, P. A., Magalhaes, M. P. & Hirano, Y. M. (1999) First record of the order Stauromedusae (Cnidaria, Scyphozoa) from the tropical southwestern Atlantic, with a review of the distribution of Stauromedusae in the southern hemisphere. Species Diversity, 4, 381 - 388.","Carlgren, O. (1938) Eine neue sudafrikanische Lucernariidae, Lucernariopsis capensis. Kungliga Fysiografiska Sallskapets i Lund Forhandlingar, 8, 1 - 6.","Miranda, L. S., Haddad, M. A., Mills, C. E. & Marques, A. C. (2012 b) Lucernariopsis capensis Carlgren, 1938 (Cnidaria, Staurozoa) in Brazil: first record outside its type locality in South Africa. Zootaxa, 3158, 60 - 64.","Corbin, P. G. (1978) A new species of the stauromedusan genus Lucernariopsis (Coelenterata: Scyphomedusae). Journal of the Marine Biological Association of the United Kingdom, 58, 285 - 290.","Zagal, C. J., Hirano, Y. M., Mills, C. E., Edgar, G. J. & Barrett, N. S. (2011) New records of Staurozoa from Australian coastal waters, with a description of a new species of Lucernariopsis Uchida, 1929 (Cnidaria, Staurozoa, Stauromedusae) and a key to Australian Stauromedusae. Marine Biology Research, 7, 651 - 666.","Larson, R. J. (1980) A new stauromedusa, Kishinouyea corbini (Scyphozoa, Stauromedusae) from the tropical western Atlantic. Bulletin of Marine Science, 30, 102 - 107.","Lechuga, G. R. & Alamo, M. A. F. (2005) Primer registro de Kishinouyea corbini Larson, 1980 (Cnidaria: Scyphozoa, Stauromedusae) para Mexico. Revista de la Sociedad Mexicana de Historia Natural, 7, 107 - 110."]}

Calvadosia capensis (Carlgren, 1938) (Fig. 8) Lucernariopsis capensis Carlgren, 1938: 1 ���6, figures 1���5; Corbin 1978: 285, 289; Grohmann et al. 1999: 386; Zagal et al. 2011: 652, 660���664; Miranda et al. 2012b: 60 ���64, figures 1, 2; Miranda et al. 2016b: 16. Calvadosia capensis ��� Miranda et al. 2016b: 19, 34, 36. Source of data. The original description of C. capensis (Carlgren, 1938) was based on one specimen collected from intertidal rocks in East London, on the southeast coast of South Africa. The holotype is probably L393, from Shelly Beach, East London, Eastern Cape, South Africa (type locality). The material is listed in the ���University of Cape Town Ecological Survey��� (UCT). In the mid-1980s, UCT donated its museum collections to Iziko South African Museum. However UCT lent samples to specialists without loans being recorded. Iziko South African Museum���s curator could not locate the material. In addition, Carlgren (1938) prepared histological sections from this material and one slide is deposited in the Invertebrates Collection of the Swedish Museum of Natural History, catalog number NRM:EVmain:115053 (GBIF ID 1099397008). A second individual was recorded from Brazil and is deposited in the Museum of Zoology of the University of S��o Paulo, Brazil, catalog number MZUSP 1566 (Miranda et al. 2012b). To date the species is known only from these preserved specimens and it has never been seen or photographed alive, despite attempts to collect the species on April 7���8th, 2016, in tidal pools on the rock shore in East London (type locality) and Gunube (about 10km east of East London). Description. (after Carlgren 1938: figures 1���5 and Miranda et al. 2012b, figure 1). Body divided into two clearly demarcated regions: calyx and peduncle (Fig. 8 A, C���E). Calyx higher than wide, pyramidal, narrowing basally (Fig. 8 A, C���E). Peduncle long, about same length as calyx (Fig. 8 A, C���E). Peduncle without interradial longitudinal muscle, with single chamber at median region (no histological details at base of peduncle). Broad, swollen adhesive circular pedal disk at base of peduncle (Fig. 8 A, E). Calyx without anchors or primary tentacles (Fig. 8 A, C���E). Gastrovascular cavity not divided by claustrum. Manubrium with four perradial lips. Numerous gastric filaments in gastrovascular cavity. Eight arms, short, organized in four interradial pairs (Fig. 8 A, C���E). Eight gonads not embedded in gastrovascular cavity (but contained within evaginations from the gastrovascular cavity), extending from manubrium to distal end of arms, consisting of several nodular lobes of irregular shape. Each arm with a cluster of secondary hollow knobbed tentacles, similar in shape (Fig. 8 B, F). Coronal muscle divided into eight segments by arms. Secondary tentacles with numerous nematocysts of two types: isorhiza and eurytele. Thin, pad-like adhesive structures (abaxial cushion) on tips of arms, on exumbrella, at base of tentacular cluster (Fig. 8 B, F). White spots of nematocysts on subumbrella. Preserved specimens green to yellowish brown (only information on preserved specimens available). Total body length about 6.70 to 13.80 mm. Distribution and habitat. Although known from only two records, this species is widely distributed, having been recorded in Shelly Beach, East London, Eastern Cape, South Africa, in the Indian Ocean (Carlgren 1938); and in Itanha��m, S��o Paulo, Brazil, in the western Atlantic Ocean (Miranda et al. 2012b) (Fig. 5 A, D). South African and Brazilian specimens were collected attached to Sargassum sp. in the intertidal zone (���University of Cape Town Ecological Survey���; Miranda et al. 2012b). Remarks. Calvadosia capensis was described from South Africa based on a single specimen (Carlgren 1938), and its general morphology (Fig. 8) is similar to C. hawaiiensis, from Hawaii (Edmondson 1930), with a narrowlyopened (pyramidal) calyx, paired arms, and thin pad-like adhesive structures on the tips of the arms (although overlooked in C. hawaiiensis by Larson 1980; see Grohmann et al. 1999). However, based on the literature, C. capensis has shorter arms and longer peduncle compared to C. hawaiiensis (Edmondson 1930, figure 6; Carlgren 1938, figure 1, Miranda et al. 2012b, figure 1)., Published as part of Miranda, Luc��lia S., Branch, George M., Collins, Allen G., Hirano, Yayoi M., Marques, Antonio C. & Griffiths, Charles L., 2017, Stalked jellyfishes (Cnidaria: Staurozoa) of South Africa, with the description of Calvadosia lewisi sp. nov., pp. 369-389 in Zootaxa 4227 (3) on pages 378-381, DOI: 10.11646/zootaxa.4227.3.5, http://zenodo.org/record/268342, {"references":["Carlgren, O. (1938) Eine neue sudafrikanische Lucernariidae, Lucernariopsis capensis. Kungliga Fysiografiska Sallskapets i Lund Forhandlingar, 8, 1 - 6.","Corbin, P. G. (1978) A new species of the stauromedusan genus Lucernariopsis (Coelenterata: Scyphomedusae). Journal of the Marine Biological Association of the United Kingdom, 58, 285 - 290.","Grohmann, P. A., Magalhaes, M. P. & Hirano, Y. M. (1999) First record of the order Stauromedusae (Cnidaria, Scyphozoa) from the tropical southwestern Atlantic, with a review of the distribution of Stauromedusae in the southern hemisphere. Species Diversity, 4, 381 - 388.","Zagal, C. J., Hirano, Y. M., Mills, C. E., Edgar, G. J. & Barrett, N. S. (2011) New records of Staurozoa from Australian coastal waters, with a description of a new species of Lucernariopsis Uchida, 1929 (Cnidaria, Staurozoa, Stauromedusae) and a key to Australian Stauromedusae. Marine Biology Research, 7, 651 - 666.","Miranda, L. S., Haddad, M. A., Mills, C. E. & Marques, A. C. (2012 b) Lucernariopsis capensis Carlgren, 1938 (Cnidaria, Staurozoa) in Brazil: first record outside its type locality in South Africa. Zootaxa, 3158, 60 - 64.","Miranda, L. S., Hirano, Y. M., Mills, C. E., Falconer, A., Fenwick, D., Marques, A. C. & Collins, A. G. (2016 b) Systematics of stalked jellyfishes (Cnidaria: Staurozoa). PeerJ, 4, e 1951.","Edmondson, C. H. (1930) New Hawaiian medusae. Bernice P. Bishop Museum Occasional Papers, 9, 1 - 16.","Larson, R. J. (1980) A new stauromedusa, Kishinouyea corbini (Scyphozoa, Stauromedusae) from the tropical western Atlantic. Bulletin of Marine Science, 30, 102 - 107."]}

Depastromorpha africana Carlgren, 1935 (Figs 9, 10) Depastromorpha africana Carlgren, 1935: 1 ���24, figures 1���12; Kramp 1961: 300; Grohmann et al. 1999: 386, 387; Dawson 2004: 252, figure 1; Collins & Daly 2005: 222, 226, figures 5���7; Cairns et al. 2009: 70; Zagal et al. 2011: 651 ���666, figures 2, 4, 8; Miranda et al. 2016b: 2, 6, 12���15, 17, 19, 25, 26, 36, figures 1, 3���5, 7, 12, 13. Material examined. MZUSP 3418, 8 specimens, Sea Point, Cape Town, Western Cape, South Africa (-33.9130, 18.3874), 22 December 2014, intertidal pools, on Gigartina polycarpa, formaldehyde solution 5%, col. & det. C. Griffiths. MZUSP 3419, 1 specimen, Kalk Bay, Cape Town, Western Cape, South Africa (-34.1266, 18.4498), 28 March 2013, intertidal pools, on Caulerpa filiformis, formaldehyde solution 5%, col. & det. C. Griffiths. MZUSP 2733, 1 specimen, Kalk Bay, Cape Town, Western Cape, South Africa (-34.1266, 18.4498), 25 April 2013, intertidal pools, on Caulerpa filiformis, ethanol 90%, col. & det. C. Griffiths. MZUSP 2734, 1 specimen, Kalk Bay, Cape Town, Western Cape, South Africa (-34.1266, 18.4498), 25 April 2013, intertidal pools, on Caulerpa filiformis, ethanol 90%, col. & det. C. Griffiths. Iziko South African Museum MB-A083795, 1 specimen, Kalk Bay, Cape Town, Western Cape, South Africa (-34.1266, 18.4498), 28 March 2013, intertidal pools, on Caulerpa filiformis, formaldehyde solution 5%, col. & det. C. Griffiths. Iziko South African Museum MB-A083796, 1 specimen, Kalk Bay, Cape Town, Western Cape, South Africa (-34.1266, 18.4498), 25 April 2013, intertidal, on Caulerpa filiformis, ethanol 80%, col. & det. C. Griffiths. Iziko South African Museum MB-A083797 (CP 718A), 1 specimen, Kalk Bay, Cape Town, Western Cape, South Africa (-34.1266, 18.4498), 9 April 1947, intertidal, on Caulerpa filiformis, ethanol 80%, col. University of Cape Town Ecological Survey, det. C. Griffiths. Iziko South African Museum MB-A084063, 5 specimens, Camps Bay, Cape Town, Western Cape, South Africa (-33.9566, 18.3755), 12 January 2016, intertidal, on various large algae in rock pools, ethanol 80%, col. & det. C. Griffiths. Iziko South African Museum H 5108, 5 specimens, Castle Rocks, False Bay, Cape Town, Western Cape, South Africa (-34.2385, 18.4766), 19 January 2003, depth 1 m, ethanol 80%, col. Y. Hirano, det. C. Griffiths. Not kept, 3 specimens, Scarborough, Cape Town, Western Cape, South Africa (���34.2017, 18.3702), 19 August 2013, from rockpools, on Sargassum longifolium, observed by G. Jones. Not kept, several specimens, Green Point, Cape Town, Western Cape, South Africa (���33.8990, 18.4078), 17 March 2014, on Brassicophycus brassicaeformis, observed by C. Griffiths. Not kept, 4 specimens, Moullie Point, Cape Town, Western Cape, South Africa (��� 33.8996, 18.4046), 22 December 2014, intertidal, on Champia, observed by C. Griffiths. Not kept; 1 specimen, Hermanus, Overberg, Western Cape, South Africa (���34.4210, 19.2437), March 1991, on Codium, observed by C. Griffiths. Not kept, 5 specimens; Dalebrook, Cape Town, Western Cape, South Africa (���34.1196, 18.4401), 2 April 1966, shallow subtidal, on Caulerpa filiformis, observed by G. Branch. USNM 1233748, 5 specimens, Gerloff Bay, Victoria, South Australia, Australia, 13 January 1998, intertidal, formaldehyde solution 5%, col. & det. Y. Hirano Information on type material. The holotype is probably A194, from Oudekraal, Cape Town, Western Cape, South Africa (type locality). As mentioned for C. capensis, the material is listed in the ���University of Cape Town Ecological Survey��� but could not be located in the Iziko South African Museum���s collection. Description. (complemented with Carlgren 1935). Calyx higher than wide, cylindrical (Fig. 9). Peduncle with four perradial chambers and four interradial longitudinal muscles. Gastrovascular cavity divided by claustrum. Manubrium with four perradial lips (Fig. 10 D, E). Numerous gastric filaments in gastrovascular cavity. Gonads extending from pyloric region to perradial margin of calyx, embedded in gastrovascular cavity (Figs 9 B, C; 10E). Gonadal vesicles organized in rows (Fig. 9 B, C). Eight short (rudimentary) adradial arms (Figs 9, 10). Each arm with a cluster of about 25 secondary hollow, knobbed tentacles (Figs 9, 10). Individual, pad-like adhesive structures in outermost secondary tentacles (Fig. 10 A���C). Perradial and interradial anchors between arms, with remnant of primary tentacles (Fig. 10 A���C). Coronal muscle entire, internal to anchors. White spots of nematocysts associated with perradial gonads (Fig. 10 D, E). General color of body very variable: orange, green, yellow, white (Fig. 9). Total body length about 10.0 to 15.0 mm. Distribution and habitat. The species was recorded for the first time at Oudekraal, Cape Town, Western Cape, South Africa (Carlgren 1935) (Fig. 5 B, E). Zagal et al. (2011) extended the occurrence of D. africana to Victoria and Tasmania, in Australia, and Otago, in New Zealand (Fig. 5 B), but molecular data indicated that Depastromorpha in Australia might be a distinct species (see Depastromorpha sp. AUS in Miranda et al. 2016b and remarks below). In this study we provide several new records for the species in the Western Cape, South Africa: Camps Bay, Cape Town; Castle Rocks, False Bay, Cape Town; Dalebrook, Cape Town; Green Point, Cape Town; Hermanus, Overberg; Kalk Bay, Cape Town; Moullie Point, Cape Town; Scarborough, Cape Town; Sea Point, Cape Town (Fig. 5 B, E). Depastromorpha africana is generally found from the intertidal to shallow subtidal depths, up to 14 m deep (Zagal et al. 2011), attached to various species of algae (Zagal et al. 2011; this study). Remarks. Depastromorpha was erected by Carlgren (1935) and comprises the single species, D. africana, whose original description was based on a single specimen from Oudekraal, Cape Town, South Africa. There are unpublished observations of the species in Australia and New Zealand (Grohmann et al. 1999), subsequently confirmed in Victoria and Tasmania (Australia), and Otago (New Zealand) (Zagal et al. 2011; specimens deposited in the Tasmanian Museum and Art Gallery, TMAG K3857���K3862). According to Zagal et al. (2011), individuals from Australia and New Zealand matched Carlgren���s (1935) description, although they have slightly smaller dimensions. However, molecular markers from D. africana of South Australia differ substantially from those from South African specimens, indicating the existence of a second species of the genus (Miranda et al. 2016b). If confirmed, D. africana would be endemic to South Africa. Detailed morphological and molecular studies, comparing the populations from South Africa, Australia, and New Zealand are necessary to assess this hypothesis and the possible existence of cryptic species (see examples in Knowlton 2000; Dawson & Jacobs 2001). Whatever the outcome, the South African species will retain its name by priority., Published as part of Miranda, Luc��lia S., Branch, George M., Collins, Allen G., Hirano, Yayoi M., Marques, Antonio C. & Griffiths, Charles L., 2017, Stalked jellyfishes (Cnidaria: Staurozoa) of South Africa, with the description of Calvadosia lewisi sp. nov., pp. 369-389 in Zootaxa 4227 (3) on pages 381-384, DOI: 10.11646/zootaxa.4227.3.5, http://zenodo.org/record/268342, {"references":["Carlgren, O. (1935) Uber eine neue Sudafrikanische Lucernariidae, Depastromorpha africana n. gen., n. sp., nebst Bemerkungen uber den Bau und die Systematik dieser Tiergruppe. Kungliga Svenska Vetenskapsakademiens Handlingar, 15, 1 - 24.","Kramp, P. L. (1961) Synopsis of the medusae of the world. Journal of the Marine Biological Association of the United Kingdom, 40, 292 - 303.","Grohmann, P. A., Magalhaes, M. P. & Hirano, Y. M. (1999) First record of the order Stauromedusae (Cnidaria, Scyphozoa) from the tropical southwestern Atlantic, with a review of the distribution of Stauromedusae in the southern hemisphere. Species Diversity, 4, 381 - 388.","Dawson, M. N. (2004) Some implications of molecular phylogenetics for understanding biodiversity in jellyfishes, with emphasis on Scyphozoa. Hydrobiologia, 530 / 531, 249 - 260.","Collins, A. G. & Daly, M. (2005) A new deepwater species of Stauromedusae, Lucernaria janetae (Cnidaria, Staurozoa, Lucernariidae), and a preliminary investigation of stauromedusan phylogeny based on nuclear and mitochondrial rDNA data. Biological Bulletin, 208, 221 - 230.","Cairns, S. D., Gershwin, L. - A., Brook, F. J., Pugh, P., Dawson, E. W., Ocana, V. O., Vervoort, W., Williams, G., Watson, J. E., Opresko, D. M., Schuchert, P., Hine, P. M., Gordon, D. P., Campbell, H. J., Wright, A. J., Sanchez, J. A. & Fautin, D. G. (2009) Phylum Cnidaria: corals, medusae, hydroids, myxozoans. In: Gordon, D. P. (Ed.), New Zealand Inventory of Biodiversity, Volume 1. Canterbury University Press, Christchurch, pp. 59 - 101.","Zagal, C. J., Hirano, Y. M., Mills, C. E., Edgar, G. J. & Barrett, N. S. (2011) New records of Staurozoa from Australian coastal waters, with a description of a new species of Lucernariopsis Uchida, 1929 (Cnidaria, Staurozoa, Stauromedusae) and a key to Australian Stauromedusae. Marine Biology Research, 7, 651 - 666.","Miranda, L. S., Hirano, Y. M., Mills, C. E., Falconer, A., Fenwick, D., Marques, A. C. & Collins, A. G. (2016 b) Systematics of stalked jellyfishes (Cnidaria: Staurozoa). PeerJ, 4, e 1951.","Knowlton, N. (2000) Molecular genetic analyses of species boundaries in the sea. Hydrobiologia, 420, 73 - 90. https: // doi. org / 10.1023 / A: 1003933603879","Dawson, M. N. & Jacobs, D. K. (2001) Molecular evidence for cryptic species of Aurelia aurita (Cnidaria, Scyphozoa). Biological Bulletin, 200, 92 - 96."]}