Margelopsis gibbesii McCrady 1859
Margelopsis gibbesii (McCrady, 1859) Figs. 1, 2 not Nemopsis gibbesii McCrady, 1859: 160, pl. 10, figs. 1–3 [medusa] [= Nemopsis bachei L. Agassiz, 1849].— Bedot, 1925: 295 [binomen N. gibbesii encompasses two species].— Stephens & Calder, 1992: 44, fig. 3, in part (reproduction of McCrady’s pl....
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Biodiversity Taxonomy Animalia Cnidaria Hydrozoa Anthoathecata Margelopsidae Margelopsis Margelopsis gibbesii |
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Biodiversity Taxonomy Animalia Cnidaria Hydrozoa Anthoathecata Margelopsidae Margelopsis Margelopsis gibbesii Calder, Dale R. Johnson, William S. Margelopsis gibbesii McCrady 1859 |
topic_facet |
Biodiversity Taxonomy Animalia Cnidaria Hydrozoa Anthoathecata Margelopsidae Margelopsis Margelopsis gibbesii |
description |
Margelopsis gibbesii (McCrady, 1859) Figs. 1, 2 not Nemopsis gibbesii McCrady, 1859: 160, pl. 10, figs. 1–3 [medusa] [= Nemopsis bachei L. Agassiz, 1849].— Bedot, 1925: 295 [binomen N. gibbesii encompasses two species].— Stephens & Calder, 1992: 44, fig. 3, in part (reproduction of McCrady’s pl. 10, figs. 1–3) [mention of medusa; historical discussion] [= Nemopsis bachei ].— Sanders & Anderson, 1999: 58 [medusa; historical discussion] [= Nemopsis bachei ]. Nemopsis gibbesii McCrady, 1859: 162 –163, pl. 10, figs. 4–7 [description of hydroid; description of medusae newly liberated from hydroid].— Allman, 1864: 370; 1872: 362 [mention of hydroid; not adult medusa].— Bedot, 1910: 333 [synonymy].— Schuchert, 2006: 361 [taxonomic discussion]. Nemopsis bachei .— L. Agassiz, 1862: 345 [in part; hydroid of Nemopsis gibbesi included in synonymy with N. bachei ] [not Nemopsis bachei L. Agassiz, 1849]. not Nemopsis bachei .— L. Agassiz, 1862: 345 [in part; medusa of Nemopsis gibbesi included in synonymy with N. bachei ].— A. Agassiz, 1865: 149 [medusa] [ Nemopsis gibbesi included in synonymy list] [incorrect subsequent spelling].— Haeckel, 1879: 93 [mention of medusa] [ Nemopsis gibbesii included in synonymy list].— Brooks, 1883: 468 [mention of medusa] [ Nemopsis gibbesii included in synonymy list]. Nemopsis gibbesi .— Frech, 1897: 565 [mention of hydroid].— Hartlaub, 1899: 221, fig. 4; 1903: 28 [mention of hydroid].— Bedot, 1918: 358 [synonymy: N. gibbesi = Margelopsis gibbesi in part] [incorrect subsequent spelling]. Margelopsis gibbesii .— Hartlaub, 1899: 224 [mention of hydroid].— Mayer, 1910, pl. 9, figs. 4–7 [illustration of medusa].— Stephens & Calder, 1992: 44, fig. 3, in part (reproduction of McCrady’s pl. 10, figs. 4–7) [mention of hydroid & young medusa; historical discussion].— Sanders & Anderson, 1999: 58 [historical discussion].— Cairns et al ., 2002: 14 [listing of hydroid]; 17 [listing of medusa].— Schuchert, 2006: 361 [taxonomic discussion].— Johnson & Allen, 2012: 94, fig. p. 95 [report of medusa]. Margelopsis gibbesi .— Hartlaub, 1903: 28 [mention of hydroid].— Hartlaub, 1907: 92 [mention of hydroid].— Mayer, 1910: 82 [report of medusa]; 83 [mention of hydroid].— Ritchie, 1915: 563 [mention of hydroid].— Bedot, 1918: 184 [literature].— Bedot, 1925: 279 [literature].— Uchida, 1927: 196 [mention of medusa].— Rees, 1941: 134 [mention of hydroid].— Fraser, 1944: 104, pl. 18, fig. 77; 1946: 156 [description of hydroid and young medusa based on McCrady 1859].— Miner, 1950: 102, figs. pl. 35 [guidebook; mention of hydroid; mention of medusa].— Werner, 1955 a: 8 [mention of medusa].— Kramp, 1959: 92, fig. 48 [account of medusa from literature].— Kramp, 1961: 49 [literature: review of medusa stage].— Allwein, 1967: 122 [report of medusa].— Gosner, 1971: 111, fig. 5.18 B [guidebook; mention of medusa].— Sandine & Swiecicki, 1975: 83 [technical report; report of medusa].— Hester, 1976: 19, pl. 1, fig. 4 [thesis; report of medusa].— Tatham et al ., 1977: 141 [technical report; report of medusa].— Calder & Hester, 1978: 88 [mention of hydroid; mention of medusa].— Harrison et al ., 1979: 37 [mention of hydroid; mention of medusa; species of “special concern”].— Cairns et al ., 1991: 16 [listing of hydroid]; 20 [listing of medusa].— Bouillon & Boero, 2000: 145 [listing of hydroid; listing of medusa].— Bouillon et al ., 2006: 245, fig. 118 B [listing of hydroid; listing of medusa; figure of medusa, from Kramp 1959 after Mayer 1910] [incorrect subsequent spelling]. not Margelopsis gibbesi. — Thiel, 1938: 294 [misidentification of medusa] [= Margelopsis australis Browne, 1910]. Margelopsis .— Werner, 1954: 144 [reference to McCrady’s 1859 hydroid]. M. (as Margelipsis ) gibbsi .— Zamponi, 1983: 178 [mention of medusa] [incorrect subsequent spelling]. Type Locality. USA: Charleston Harbor, South Carolina (McCrady 1859). Material examined. Virginia Beach, Virginia, from aquaria operated by Hampton Roads Sanitation District, May 1989, 22 ‰, several dozen hydroids, some with medusa buds, coll. Ms. Butterworth, ROMIZ B 1027.— Chincoteague, Virginia, Station 199, about 5 km offshore, depth 10 m, 24.iv. 2006, epibenthic sled, <20 cm off bottom, two male medusae, coll. W. Johnson, ROMIZ B 4077.—Assateague Island, Virginia, Station 208, just off beach near southern tip of island, surf zone, depth 1–2 m, 10.vi. 2006, epibenthic sled, <20 cm off bottom, three female medusae, some with nascent polyps, coll. W. Johnson, ROMIZ B 4078.—Assateague Island, Virginia, Station 221, ca. 200 m off beach near southern tip of island, depth 5 m, 10.vi. 2006, epibenthic sled, <20 cm off bottom, one male medusa, coll. W. Johnson, ROMIZ B 4079.—Assateague Island, Virginia, ca. 400 m offshore, depth 5 m, 10.vi. 2006, epibenthic sled, <20 cm off bottom, about 35 female medusae, some with nascent polyps, and four male medusae, coll. W. Johnson, ROMIZ B 4080. Description of material. Hydroids solitary, naked, monomorphic, free-floating. Hydranths all young, minute (<1 mm long), typically vasiform to pear-shaped, with dome-shaped hypostome at distal end and vestigial hydrocaulus at proximal end; mouth at tip of hypostome. Hydrocaulus with base hollowed out, having a sucker-like appearance. Tentacles appearing moniliform, in widely separated oral and aboral whorls; oral tentacles 5–7 in number; aboral tentacles 7–9 in number, slightly larger than oral ones. Some hydranths appearing to undergo asexual reproduction by transverse fission (Fig. 1 e) and budding (Fig. 1 f). Gonophores medusa buds arising on hydranth distal to aboral tentacular whorl; all observed medusa buds very early in development. Medusae thimble-shaped with rounded apex, up to 2.3 mm high, 1.9 mm wide; mesoglea of moderate thickness. Exumbrellar surface with scattered nematocysts, possibly subspherical heteronemes, and nematocyst patches. Apical canal present, narrow, inconspicuous. Manubrium vasiform, extending nearly to velar opening, proximal end with vacuolated gastrodermal cells. Mouth irregularly circular, without lips or oral tentacles. Radial canals four, simple. Ring canal present. Marginal bulbs four, pyriform. Ocelli lacking. Marginal tentacles 5–8 per marginal bulb, moniliform, with nematocysts arranged in rings and with an elongated terminal bulb. Velum welldeveloped. Gonad surrounding manubrium except at upper and lower extremities. Female medusae with prominent subitaneous eggs extruded from gonad, some developing as nascent polyps (actinulae). Developing polyps subspherical to discoidal at release, about 0.3 mm in diameter, with approximately 12–13 short tentacles. Cnidome of hydroid . Desmonemes (n = 10): 5.0– 5.8 µm long × 2.6–3.3 µm wide (undischarged) Haplonemes (n = 10): 6.3–7.4 µm long × 2.5–3.6 µm wide (undischarged) Microbasic heteronemes (n = 10): 7.0– 8.4 µm long × 2.8–3.6 µm wide (undischarged) Cnidome of medusa . Desmonemes (n = 10): 3.7 –4.0 µm long × 2.8–3.2 µm wide (undischarged) Haplonemes (n = 5): 5.0– 6.4 µm long × 1.6–2.5 µm wide (undischarged) Microbasic heteronemes (n = 10): 5.0– 6.1 µm long × 2.8–3.6 µm wide (undischarged) Subspherical heteronemes (n = 10): 8.4–9.5 µm long × 7.5–8.6 µm wide (undischarged) Stenoteles (n = 10): 5.8–7.3 µm long × 4.0– 5.9 µm wide (undischarged) Historical account . Under the name Nemopsis gibbesii , McCrady (1859) described and illustrated a medusa from Charleston Harbor, South Carolina, that he regarded as a new species. Included with it was an account of a hydroid, also illustrated, that he took to be its polyp stage. McCrady recognized that the medusa was referable to the same genus as Nemopsis bachei L. Agassiz, 1849 from New England, but considered it to be specifically distinct. The hydroid he associated with the species bore medusa buds, and medusae were liberated from it. Apparently seen only once by him, it was found during January as a free-floating, solitary polyp having a rudimentary hydrocaulus and two whorls of tentacles. Medusae attributed to N. gibbesii were observed during winter and spring. Without comment, but surely based on the medusa stage only, L. Agassiz (1862: 345) included Nemopsis gibbesii (spelled therein as N. gibbesi ) in the synonymy of N. bachei . Following studies on the medusa stage of N. bachei in New England, A. Agassiz (1865) also included N. gibbesii (again spelled N. gibbesi ) as a subjective synonym. The only reservation he expressed in doing so was based on seasonality of populations at the two locations, with those from Charleston occurring in winter and those from Vineyard Sound being common in September. After searching for the hydroid stage without success, the younger Agassiz concluded that McCrady’s planktonic polyp had simply been a detached hydranth of a benthic species. Brooks (1883) was first to unequivocally establish that the hydroid and medusa stages attributed to Nemopsis gibbesii by McCrady (1859) were different species. He raised medusae identical to the original account of N. gibbesii from a benthic colonial hydroid reportedly resembling species of Eudendrium Ehrenberg, 1834 or Bougainvillia Lesson, 1830, both morphologically quite unlike the polyp described by McCrady. His observations on development of the species supported the hypothesis that medusae of N. gibbesii and N. bachei were conspecific. However, that confirmation did not fully resolve the taxonomic identity of the nominal species N. gibbesii . No type specimen had been designated to objectively define it, whether hydroid or medusa, and the true affinities of McCrady’s remarkable hydroid remained unknown. As with A. Agassiz (1865) earlier, Allman (1872) questioned whether the trophosome was indeed a normally free-floating polyp. The generic identity of the hydroid McCrady (1859) had assigned to Nemopsis gibbesii was finally resolved by Hartlaub (1899) through life cycle studies of the European medusa Margelopsis haeckelii Hartlaub, 1897. He discovered that its hydroid stage was a solitary, planktonic polyp much like that described by McCrady. Hartlaub thereupon assigned McCrady’s hydroid to Margelopsis Hartlaub, 1897, as M. gibbesii . The medusa stage of the American hydroid remained unknown until Mayer (1910) discovered and described specimens from North Carolina and South Carolina, referring to them as Margelopsis gibbesi in the text (p. 82) and as M. gibbesii in captions accompanying Plate 9. While the link between McCrady’s hydroid and Mayer’s medusa is subjective, not having been based on a complete life cycle study, it seems probable that they are conspecific. For more than a century, the name M. gibbesi has commonly been applied to the two stages (see synonymy list above). Both hydroid and medusa appear endemic to the southeast and mid-Atlantic coasts of the United States. A report of the species from the South Atlantic Ocean between South Georgia and Bouvet Island by Thiel (1938) is believed to have been based instead on M. australis Browne, 1910 (Kramp 1961: 49). Nomenclature and taxonomy . As apparent from the synonymy list above, the specific name of McCrady’s species has been spelled in the literature as both gibbesii and gibbesi . Following provisions of the International Code of Zoological Nomenclature (ICZN) (International Commission on Zoological Nomenclature 1999), gibbesii is the original and correct spelling (ICZN Art. 32). The name honours noted South Carolina mathematician and naturalist Lewis Reeve Gibbes (1810–1894) (Sanders & Anderson 1999; Stephens 2000). Although the identities of the hydroid and medusa described by McCrady (1859) have long been known, nomenclatural uncertainty persists over the concept of the species to which they were assigned. The taxonomic identity of Nemopsis gibbesii remains ambiguous because of the absence of a name-bearing type for a binomen that originally included two species. McCrady did not designate a holotype of N. gibbesii , and no syntype specimens are known to exist. His collections, including types, are thought to have been destroyed by fire during the American Civil War (Stephens & Calder 1992: 44). We are also unaware of any previous lectotype or neotype selections. To stabilize the taxonomic identity of Nemopsis gibbesii , we select as lectotype the hydroid specimen illustrated by McCrady (1859) in Plate 10, Figure 7 following ICZN Arts. 74.7. 1, 74.7. 2, and 74.7. 3. Our designation is not invalidated by “…the fact that the specimen no longer exists or cannot be traced” (ICZN Article 74.4). The lectotype chosen here upholds prevailing usage of the name Margelopsis gibbesii for both hydroid and medusa stages of the species since the turn of the 20 th century. Had one of the medusae illustrated in McCrady’s work (Plate 10, Figures 1 –3) been chosen as lectotype, N. gibbesii would become a subjective synonym of N. bachei . A new specific name for the species widely known as M. gibbesii would then be required, an act we consider unnecessary and undesirable. Margelopsidae, the name of the family to which M. gibbesii belongs, is sometimes credited in error to Uchida (1927). Under the Principle of Coordination in nomenclature (ICZN Art. 36), authorship and date should be credited instead to Mayer (1910). Although Mayer established the name as Margelopsinae, for a subfamily, he is deemed under that article of the code to have simultaneously established coordinate names for all other ranks of the family group, including the name Margelopsidae. Two other genera, Pelagohydra Dendy, 1902 and Climacocodon Uchida, 1924, are usually included with Margelopsis in the family Margelopsidae (e.g. Uchida 1924; Kramp 1961; Petersen 1990; Bouillon et al . 2006; Schuchert 2006). When Pelagohydra and Margelopsis are included in the same family, a nomenclatural issue arises because the infrequently used family name Pelagohydridae Dendy, 1902 predates the more familiar name Margelopsidae Mayer, 1910. The two had earlier been recognized as distinct subfamilies by Rees (1941), with Margelopsinae (including Margelopsis and Climacocodon ) having whorls of tentacles on the hydranth and lacking a float, and Pelagohydrinae (including only Pelagohydra ) having scattered tentacles over the entire hydranth and having the proximal portion of the hydranth modified into a float. Unfortunately, the adult medusa stage of Pelagohydra is as yet unknown for comparison with that of Margelopsis . Following Schuchert (2006: 356), prevailing usage of the name Margelopsidae is maintained here pending a taxonomic or nomenclatural resolution of the problem. The family Margelopsidae has generally been classified within the superfamily Tubularioidea Fleming, 1828 (e.g. Rees 1957; Bouillon 1985; Petersen 1990). The latter group has recently been transferred from the suborder Capitata Kühn, 1913 to Aplanulata Collins, Winkelman, Hadrys & Schierwater, 2005 (Cartwright & Nawrocki 2010; Nawrocki et al . 2013). Molecular evidence supporting inclusion of Margelopsidae in Aplanulata, however, is still lacking (Peter Schuchert, personal communication, 30 June 2015). The six species commonly included in the family ( Margelopsis gibbesii , M. haeckelii , M. hartlaubii , M. australis , Pelagohydra mirabilis , and Climacocodon ikarii ) all appear to be of infrequent occurrence, and specimens have been difficult to obtain for analysis. As for the name Tubularioidea, authorship of it should be credited to Fleming (1828). While the name “Tubulariae” had been used earlier in both Goldfuss (1818) and Fischer von Waldheim (1823), it was not rendered available in either of those works (ICZN Art. 11.7.1.1). As used in them, “Tubulariae” was merely a descriptive term applied to groups that excluded the genus Tubularia Linnaeus, 1758 (see Calder 2010: 45). Purported morphological differences between M. gibbesii from the western North Atlantic and M. haeckelii from the eastern North Atlantic need re-examination. In a key to medusa stages of species of Margelopsis , Kramp (1959) highlighted three characters based on then-existing descriptions that seemed to discriminate the two. Margelopsis haeckelii was distinguished by having 3–4 tentacles on each marginal bulb, a wide axial canal above the manubrium in adults, and eggs that developed into actinula-like larval stages on the manubrium. In M. gibbesii , 5–6 tentacles were said to arise from each marginal bulb, no wide axial canal was known to exist above the manubrium in adults, and no actinula-like larval stages were thought to occur on the manubrium. The latter distinction has been shown here to be inaccurate; some female medusae of M. gibbesii studied by us were observed with developing larval polyps just as in M. haeckelii . Meanwhile, the other two characters require further evaluation. Several medusae of M. gibbesii in our material possessed an axial canal, although it was narrow in each case. As for numbers of marginal tentacles, Werner (1955 a) reported as many as 8–9 on one or more of the marginal bulbs of M. haeckelii . Up to eight were seen on the bulbs of a large male of M. gibbesii examined during this study (ROMIZ B 4077). The hydroid stage of M. gibbesii is still too poorly known to reliably assess how it may differ, if at all, from that of M. haeckelii . Overall, it seems best to maintain the two as distinct species for now, an opinion expressed earlier by Schuchert (2006). Live medusae of M. gibbesii were examined and described by Mayer (1910). The umbrella was said to be highly contractile. When relaxed, the manubrium reached little more than half-way along the length of the subumbrellar cavity but when contacted, the mouth often extended beyond the velar opening. The endoderm of the marginal bulbs was dull-yellow in colour, while that of the manubrium varied from dull-yellow to dull-green. In M. haeckelii , marginal bulbs are brown while the manubrium is gray with dark-brown pigment granules (Schuchert 2006). Life cycle . The life cycle of Margelopsis gibbesii has yet to be adequately described. Werner (1954, 1955a, b, 1956) traced the remarkable life cycle of M. haeckelii , and it seems likely that : Published as part of Calder, Dale R. & Johnson, William S., 2015, Margelopsis gibbesii (McCrady, 1859) (Cnidaria, Hydrozoa): taxonomic review, and conservation of usage by designation of a lectotype, pp. 215-225 in Zootaxa 4052 (2) on pages 216-222, DOI: 10.11646/zootaxa.4052.2.5, http://zenodo.org/record/242944 : {"references": ["McCrady, J. (1859) Gymnopthalmata of Charleston Harbor. Proceedings of the Elliott Society of Natural History, 1, 103 - 221.", "Agassiz, L. (1849) Contributions to the natural history of the Acalephae of North America. Part I. - On the naked-eyed medusae of the shores of Massachusetts, in their perfect state of development. Memoirs of the American Academy of Arts and Sciences, 4, 221 - 316. http: // dx. doi. org / 10.2307 / 25058163", "Bedot, M. (1925) Materiaux pour servir a l'histoire des hydroides. 7 e periode (1901 a 1910). 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Ichthyological Associates, Inc., Ithaca, New York, 368 pp.", "Calder, D. R. & Hester, B. S. (1978) Phylum Cnidaria. In: Zingmark, R. G. (Ed.), An annotated checklist of the biota of the coastal zone of South Carolina. University of South Carolina Press, Columbia, pp. 87 - 93.", "Harrison, F. W., Calder, D. R., Coull, B. C. & James, F. C. (1979) Status report: lower invertebrates and miscellaneous phyla. In: Forsythe, D. M. & Ezell, W. B. (Eds.), Proceedings of the First South Carolina Endangered Species Symposium, Charleston, South Carolina, pp. 34 - 40.", "Cairns, S. D., Calder, D. R., Brinckmann-Voss, A., Castro, C. B., Pugh, P. R., Cutress, C. E., Jaap, W. C., Fautin, D. G., Larson, R. J., Harbison, G. R., Arai, M. N. & Opresko, D. M. (1991) Common and scientific names of aquatic invertebrates from the United States and Canada: Cnidaria and Ctenophora. American Fisheries Society Special Publication, 22, 1 - 75.", "Bouillon, J. & Boero, F. 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(1996) The marine fauna of New Zealand: athecate hydroids and their medusae (Cnidaria: Hydrozoa). New Zealand Oceanographic Institute Memoir, 106, 1 - 159."]} |
format |
Text |
author |
Calder, Dale R. Johnson, William S. |
author_facet |
Calder, Dale R. Johnson, William S. |
author_sort |
Calder, Dale R. |
title |
Margelopsis gibbesii McCrady 1859 |
title_short |
Margelopsis gibbesii McCrady 1859 |
title_full |
Margelopsis gibbesii McCrady 1859 |
title_fullStr |
Margelopsis gibbesii McCrady 1859 |
title_full_unstemmed |
Margelopsis gibbesii McCrady 1859 |
title_sort |
margelopsis gibbesii mccrady 1859 |
publisher |
Zenodo |
publishDate |
2015 |
url |
https://dx.doi.org/10.5281/zenodo.6106039 https://zenodo.org/record/6106039 |
long_lat |
ENVELOPE(-101.250,-101.250,-71.917,-71.917) ENVELOPE(3.358,3.358,-54.422,-54.422) ENVELOPE(102.867,102.867,-65.867,-65.867) ENVELOPE(157.417,157.417,-79.633,-79.633) ENVELOPE(3.358,3.358,-54.422,-54.422) ENVELOPE(-70.100,-70.100,-69.333,-69.333) ENVELOPE(-57.976,-57.976,-63.685,-63.685) ENVELOPE(166.083,166.083,62.917,62.917) ENVELOPE(66.733,66.733,-70.700,-70.700) ENVELOPE(-110.908,-110.908,78.348,78.348) ENVELOPE(-128.387,-128.387,54.916,54.916) ENVELOPE(-57.018,-57.018,53.708,53.708) |
geographic |
Antarctic Canada Norway Indian New Zealand Petersen Bouvet Elliott Medusa Bouvet Island Hampton Chapel Hill Sever Butterworth Oyster Creek Ritchie Cartwright |
geographic_facet |
Antarctic Canada Norway Indian New Zealand Petersen Bouvet Elliott Medusa Bouvet Island Hampton Chapel Hill Sever Butterworth Oyster Creek Ritchie Cartwright |
genre |
Antarc* Antarctic Bouvet Island North Atlantic South Atlantic Ocean Spitzbergen |
genre_facet |
Antarc* Antarctic Bouvet Island North Atlantic South Atlantic Ocean Spitzbergen |
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Open Access Creative Commons Zero v1.0 Universal https://creativecommons.org/publicdomain/zero/1.0/legalcode cc0-1.0 info:eu-repo/semantics/openAccess |
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CC0 |
op_doi |
https://doi.org/10.5281/zenodo.6106039 https://doi.org/10.11646/zootaxa.4052.2.5 https://doi.org/10.5281/zenodo.242945 https://doi.org/10.5281/zenodo.242946 https://doi.org/10.5281/zenodo.6106038 |
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ftdatacite:10.5281/zenodo.6106039 2023-05-15T13:46:15+02:00 Margelopsis gibbesii McCrady 1859 Calder, Dale R. Johnson, William S. 2015 https://dx.doi.org/10.5281/zenodo.6106039 https://zenodo.org/record/6106039 unknown Zenodo http://zenodo.org/record/242944 http://publication.plazi.org/id/FFC3FFDBFFF3C115FF9E7A5EFFE2FFA9 http://zoobank.org/AF7D9D8E-4DE7-43D0-AA7A-10104E95A6E9 https://zenodo.org/communities/biosyslit https://dx.doi.org/10.11646/zootaxa.4052.2.5 http://zenodo.org/record/242944 http://publication.plazi.org/id/FFC3FFDBFFF3C115FF9E7A5EFFE2FFA9 https://dx.doi.org/10.5281/zenodo.242945 https://dx.doi.org/10.5281/zenodo.242946 http://zoobank.org/AF7D9D8E-4DE7-43D0-AA7A-10104E95A6E9 https://dx.doi.org/10.5281/zenodo.6106038 https://zenodo.org/communities/biosyslit Open Access Creative Commons Zero v1.0 Universal https://creativecommons.org/publicdomain/zero/1.0/legalcode cc0-1.0 info:eu-repo/semantics/openAccess CC0 Biodiversity Taxonomy Animalia Cnidaria Hydrozoa Anthoathecata Margelopsidae Margelopsis Margelopsis gibbesii article-journal ScholarlyArticle Taxonomic treatment Text 2015 ftdatacite https://doi.org/10.5281/zenodo.6106039 https://doi.org/10.11646/zootaxa.4052.2.5 https://doi.org/10.5281/zenodo.242945 https://doi.org/10.5281/zenodo.242946 https://doi.org/10.5281/zenodo.6106038 2022-04-01T10:40:26Z Margelopsis gibbesii (McCrady, 1859) Figs. 1, 2 not Nemopsis gibbesii McCrady, 1859: 160, pl. 10, figs. 1–3 [medusa] [= Nemopsis bachei L. Agassiz, 1849].— Bedot, 1925: 295 [binomen N. gibbesii encompasses two species].— Stephens & Calder, 1992: 44, fig. 3, in part (reproduction of McCrady’s pl. 10, figs. 1–3) [mention of medusa; historical discussion] [= Nemopsis bachei ].— Sanders & Anderson, 1999: 58 [medusa; historical discussion] [= Nemopsis bachei ]. Nemopsis gibbesii McCrady, 1859: 162 –163, pl. 10, figs. 4–7 [description of hydroid; description of medusae newly liberated from hydroid].— Allman, 1864: 370; 1872: 362 [mention of hydroid; not adult medusa].— Bedot, 1910: 333 [synonymy].— Schuchert, 2006: 361 [taxonomic discussion]. Nemopsis bachei .— L. Agassiz, 1862: 345 [in part; hydroid of Nemopsis gibbesi included in synonymy with N. bachei ] [not Nemopsis bachei L. Agassiz, 1849]. not Nemopsis bachei .— L. Agassiz, 1862: 345 [in part; medusa of Nemopsis gibbesi included in synonymy with N. bachei ].— A. Agassiz, 1865: 149 [medusa] [ Nemopsis gibbesi included in synonymy list] [incorrect subsequent spelling].— Haeckel, 1879: 93 [mention of medusa] [ Nemopsis gibbesii included in synonymy list].— Brooks, 1883: 468 [mention of medusa] [ Nemopsis gibbesii included in synonymy list]. Nemopsis gibbesi .— Frech, 1897: 565 [mention of hydroid].— Hartlaub, 1899: 221, fig. 4; 1903: 28 [mention of hydroid].— Bedot, 1918: 358 [synonymy: N. gibbesi = Margelopsis gibbesi in part] [incorrect subsequent spelling]. Margelopsis gibbesii .— Hartlaub, 1899: 224 [mention of hydroid].— Mayer, 1910, pl. 9, figs. 4–7 [illustration of medusa].— Stephens & Calder, 1992: 44, fig. 3, in part (reproduction of McCrady’s pl. 10, figs. 4–7) [mention of hydroid & young medusa; historical discussion].— Sanders & Anderson, 1999: 58 [historical discussion].— Cairns et al ., 2002: 14 [listing of hydroid]; 17 [listing of medusa].— Schuchert, 2006: 361 [taxonomic discussion].— Johnson & Allen, 2012: 94, fig. p. 95 [report of medusa]. Margelopsis gibbesi .— Hartlaub, 1903: 28 [mention of hydroid].— Hartlaub, 1907: 92 [mention of hydroid].— Mayer, 1910: 82 [report of medusa]; 83 [mention of hydroid].— Ritchie, 1915: 563 [mention of hydroid].— Bedot, 1918: 184 [literature].— Bedot, 1925: 279 [literature].— Uchida, 1927: 196 [mention of medusa].— Rees, 1941: 134 [mention of hydroid].— Fraser, 1944: 104, pl. 18, fig. 77; 1946: 156 [description of hydroid and young medusa based on McCrady 1859].— Miner, 1950: 102, figs. pl. 35 [guidebook; mention of hydroid; mention of medusa].— Werner, 1955 a: 8 [mention of medusa].— Kramp, 1959: 92, fig. 48 [account of medusa from literature].— Kramp, 1961: 49 [literature: review of medusa stage].— Allwein, 1967: 122 [report of medusa].— Gosner, 1971: 111, fig. 5.18 B [guidebook; mention of medusa].— Sandine & Swiecicki, 1975: 83 [technical report; report of medusa].— Hester, 1976: 19, pl. 1, fig. 4 [thesis; report of medusa].— Tatham et al ., 1977: 141 [technical report; report of medusa].— Calder & Hester, 1978: 88 [mention of hydroid; mention of medusa].— Harrison et al ., 1979: 37 [mention of hydroid; mention of medusa; species of “special concern”].— Cairns et al ., 1991: 16 [listing of hydroid]; 20 [listing of medusa].— Bouillon & Boero, 2000: 145 [listing of hydroid; listing of medusa].— Bouillon et al ., 2006: 245, fig. 118 B [listing of hydroid; listing of medusa; figure of medusa, from Kramp 1959 after Mayer 1910] [incorrect subsequent spelling]. not Margelopsis gibbesi. — Thiel, 1938: 294 [misidentification of medusa] [= Margelopsis australis Browne, 1910]. Margelopsis .— Werner, 1954: 144 [reference to McCrady’s 1859 hydroid]. M. (as Margelipsis ) gibbsi .— Zamponi, 1983: 178 [mention of medusa] [incorrect subsequent spelling]. Type Locality. USA: Charleston Harbor, South Carolina (McCrady 1859). Material examined. Virginia Beach, Virginia, from aquaria operated by Hampton Roads Sanitation District, May 1989, 22 ‰, several dozen hydroids, some with medusa buds, coll. Ms. Butterworth, ROMIZ B 1027.— Chincoteague, Virginia, Station 199, about 5 km offshore, depth 10 m, 24.iv. 2006, epibenthic sled, <20 cm off bottom, two male medusae, coll. W. Johnson, ROMIZ B 4077.—Assateague Island, Virginia, Station 208, just off beach near southern tip of island, surf zone, depth 1–2 m, 10.vi. 2006, epibenthic sled, <20 cm off bottom, three female medusae, some with nascent polyps, coll. W. Johnson, ROMIZ B 4078.—Assateague Island, Virginia, Station 221, ca. 200 m off beach near southern tip of island, depth 5 m, 10.vi. 2006, epibenthic sled, <20 cm off bottom, one male medusa, coll. W. Johnson, ROMIZ B 4079.—Assateague Island, Virginia, ca. 400 m offshore, depth 5 m, 10.vi. 2006, epibenthic sled, <20 cm off bottom, about 35 female medusae, some with nascent polyps, and four male medusae, coll. W. Johnson, ROMIZ B 4080. Description of material. Hydroids solitary, naked, monomorphic, free-floating. Hydranths all young, minute (<1 mm long), typically vasiform to pear-shaped, with dome-shaped hypostome at distal end and vestigial hydrocaulus at proximal end; mouth at tip of hypostome. Hydrocaulus with base hollowed out, having a sucker-like appearance. Tentacles appearing moniliform, in widely separated oral and aboral whorls; oral tentacles 5–7 in number; aboral tentacles 7–9 in number, slightly larger than oral ones. Some hydranths appearing to undergo asexual reproduction by transverse fission (Fig. 1 e) and budding (Fig. 1 f). Gonophores medusa buds arising on hydranth distal to aboral tentacular whorl; all observed medusa buds very early in development. Medusae thimble-shaped with rounded apex, up to 2.3 mm high, 1.9 mm wide; mesoglea of moderate thickness. Exumbrellar surface with scattered nematocysts, possibly subspherical heteronemes, and nematocyst patches. Apical canal present, narrow, inconspicuous. Manubrium vasiform, extending nearly to velar opening, proximal end with vacuolated gastrodermal cells. Mouth irregularly circular, without lips or oral tentacles. Radial canals four, simple. Ring canal present. Marginal bulbs four, pyriform. Ocelli lacking. Marginal tentacles 5–8 per marginal bulb, moniliform, with nematocysts arranged in rings and with an elongated terminal bulb. Velum welldeveloped. Gonad surrounding manubrium except at upper and lower extremities. Female medusae with prominent subitaneous eggs extruded from gonad, some developing as nascent polyps (actinulae). Developing polyps subspherical to discoidal at release, about 0.3 mm in diameter, with approximately 12–13 short tentacles. Cnidome of hydroid . Desmonemes (n = 10): 5.0– 5.8 µm long × 2.6–3.3 µm wide (undischarged) Haplonemes (n = 10): 6.3–7.4 µm long × 2.5–3.6 µm wide (undischarged) Microbasic heteronemes (n = 10): 7.0– 8.4 µm long × 2.8–3.6 µm wide (undischarged) Cnidome of medusa . Desmonemes (n = 10): 3.7 –4.0 µm long × 2.8–3.2 µm wide (undischarged) Haplonemes (n = 5): 5.0– 6.4 µm long × 1.6–2.5 µm wide (undischarged) Microbasic heteronemes (n = 10): 5.0– 6.1 µm long × 2.8–3.6 µm wide (undischarged) Subspherical heteronemes (n = 10): 8.4–9.5 µm long × 7.5–8.6 µm wide (undischarged) Stenoteles (n = 10): 5.8–7.3 µm long × 4.0– 5.9 µm wide (undischarged) Historical account . Under the name Nemopsis gibbesii , McCrady (1859) described and illustrated a medusa from Charleston Harbor, South Carolina, that he regarded as a new species. Included with it was an account of a hydroid, also illustrated, that he took to be its polyp stage. McCrady recognized that the medusa was referable to the same genus as Nemopsis bachei L. Agassiz, 1849 from New England, but considered it to be specifically distinct. The hydroid he associated with the species bore medusa buds, and medusae were liberated from it. Apparently seen only once by him, it was found during January as a free-floating, solitary polyp having a rudimentary hydrocaulus and two whorls of tentacles. Medusae attributed to N. gibbesii were observed during winter and spring. Without comment, but surely based on the medusa stage only, L. Agassiz (1862: 345) included Nemopsis gibbesii (spelled therein as N. gibbesi ) in the synonymy of N. bachei . Following studies on the medusa stage of N. bachei in New England, A. Agassiz (1865) also included N. gibbesii (again spelled N. gibbesi ) as a subjective synonym. The only reservation he expressed in doing so was based on seasonality of populations at the two locations, with those from Charleston occurring in winter and those from Vineyard Sound being common in September. After searching for the hydroid stage without success, the younger Agassiz concluded that McCrady’s planktonic polyp had simply been a detached hydranth of a benthic species. Brooks (1883) was first to unequivocally establish that the hydroid and medusa stages attributed to Nemopsis gibbesii by McCrady (1859) were different species. He raised medusae identical to the original account of N. gibbesii from a benthic colonial hydroid reportedly resembling species of Eudendrium Ehrenberg, 1834 or Bougainvillia Lesson, 1830, both morphologically quite unlike the polyp described by McCrady. His observations on development of the species supported the hypothesis that medusae of N. gibbesii and N. bachei were conspecific. However, that confirmation did not fully resolve the taxonomic identity of the nominal species N. gibbesii . No type specimen had been designated to objectively define it, whether hydroid or medusa, and the true affinities of McCrady’s remarkable hydroid remained unknown. As with A. Agassiz (1865) earlier, Allman (1872) questioned whether the trophosome was indeed a normally free-floating polyp. The generic identity of the hydroid McCrady (1859) had assigned to Nemopsis gibbesii was finally resolved by Hartlaub (1899) through life cycle studies of the European medusa Margelopsis haeckelii Hartlaub, 1897. He discovered that its hydroid stage was a solitary, planktonic polyp much like that described by McCrady. Hartlaub thereupon assigned McCrady’s hydroid to Margelopsis Hartlaub, 1897, as M. gibbesii . The medusa stage of the American hydroid remained unknown until Mayer (1910) discovered and described specimens from North Carolina and South Carolina, referring to them as Margelopsis gibbesi in the text (p. 82) and as M. gibbesii in captions accompanying Plate 9. While the link between McCrady’s hydroid and Mayer’s medusa is subjective, not having been based on a complete life cycle study, it seems probable that they are conspecific. For more than a century, the name M. gibbesi has commonly been applied to the two stages (see synonymy list above). Both hydroid and medusa appear endemic to the southeast and mid-Atlantic coasts of the United States. A report of the species from the South Atlantic Ocean between South Georgia and Bouvet Island by Thiel (1938) is believed to have been based instead on M. australis Browne, 1910 (Kramp 1961: 49). Nomenclature and taxonomy . As apparent from the synonymy list above, the specific name of McCrady’s species has been spelled in the literature as both gibbesii and gibbesi . Following provisions of the International Code of Zoological Nomenclature (ICZN) (International Commission on Zoological Nomenclature 1999), gibbesii is the original and correct spelling (ICZN Art. 32). The name honours noted South Carolina mathematician and naturalist Lewis Reeve Gibbes (1810–1894) (Sanders & Anderson 1999; Stephens 2000). Although the identities of the hydroid and medusa described by McCrady (1859) have long been known, nomenclatural uncertainty persists over the concept of the species to which they were assigned. The taxonomic identity of Nemopsis gibbesii remains ambiguous because of the absence of a name-bearing type for a binomen that originally included two species. McCrady did not designate a holotype of N. gibbesii , and no syntype specimens are known to exist. His collections, including types, are thought to have been destroyed by fire during the American Civil War (Stephens & Calder 1992: 44). We are also unaware of any previous lectotype or neotype selections. To stabilize the taxonomic identity of Nemopsis gibbesii , we select as lectotype the hydroid specimen illustrated by McCrady (1859) in Plate 10, Figure 7 following ICZN Arts. 74.7. 1, 74.7. 2, and 74.7. 3. Our designation is not invalidated by “…the fact that the specimen no longer exists or cannot be traced” (ICZN Article 74.4). The lectotype chosen here upholds prevailing usage of the name Margelopsis gibbesii for both hydroid and medusa stages of the species since the turn of the 20 th century. Had one of the medusae illustrated in McCrady’s work (Plate 10, Figures 1 –3) been chosen as lectotype, N. gibbesii would become a subjective synonym of N. bachei . A new specific name for the species widely known as M. gibbesii would then be required, an act we consider unnecessary and undesirable. Margelopsidae, the name of the family to which M. gibbesii belongs, is sometimes credited in error to Uchida (1927). Under the Principle of Coordination in nomenclature (ICZN Art. 36), authorship and date should be credited instead to Mayer (1910). Although Mayer established the name as Margelopsinae, for a subfamily, he is deemed under that article of the code to have simultaneously established coordinate names for all other ranks of the family group, including the name Margelopsidae. Two other genera, Pelagohydra Dendy, 1902 and Climacocodon Uchida, 1924, are usually included with Margelopsis in the family Margelopsidae (e.g. Uchida 1924; Kramp 1961; Petersen 1990; Bouillon et al . 2006; Schuchert 2006). When Pelagohydra and Margelopsis are included in the same family, a nomenclatural issue arises because the infrequently used family name Pelagohydridae Dendy, 1902 predates the more familiar name Margelopsidae Mayer, 1910. The two had earlier been recognized as distinct subfamilies by Rees (1941), with Margelopsinae (including Margelopsis and Climacocodon ) having whorls of tentacles on the hydranth and lacking a float, and Pelagohydrinae (including only Pelagohydra ) having scattered tentacles over the entire hydranth and having the proximal portion of the hydranth modified into a float. Unfortunately, the adult medusa stage of Pelagohydra is as yet unknown for comparison with that of Margelopsis . Following Schuchert (2006: 356), prevailing usage of the name Margelopsidae is maintained here pending a taxonomic or nomenclatural resolution of the problem. The family Margelopsidae has generally been classified within the superfamily Tubularioidea Fleming, 1828 (e.g. Rees 1957; Bouillon 1985; Petersen 1990). The latter group has recently been transferred from the suborder Capitata Kühn, 1913 to Aplanulata Collins, Winkelman, Hadrys & Schierwater, 2005 (Cartwright & Nawrocki 2010; Nawrocki et al . 2013). Molecular evidence supporting inclusion of Margelopsidae in Aplanulata, however, is still lacking (Peter Schuchert, personal communication, 30 June 2015). The six species commonly included in the family ( Margelopsis gibbesii , M. haeckelii , M. hartlaubii , M. australis , Pelagohydra mirabilis , and Climacocodon ikarii ) all appear to be of infrequent occurrence, and specimens have been difficult to obtain for analysis. As for the name Tubularioidea, authorship of it should be credited to Fleming (1828). While the name “Tubulariae” had been used earlier in both Goldfuss (1818) and Fischer von Waldheim (1823), it was not rendered available in either of those works (ICZN Art. 11.7.1.1). As used in them, “Tubulariae” was merely a descriptive term applied to groups that excluded the genus Tubularia Linnaeus, 1758 (see Calder 2010: 45). Purported morphological differences between M. gibbesii from the western North Atlantic and M. haeckelii from the eastern North Atlantic need re-examination. In a key to medusa stages of species of Margelopsis , Kramp (1959) highlighted three characters based on then-existing descriptions that seemed to discriminate the two. Margelopsis haeckelii was distinguished by having 3–4 tentacles on each marginal bulb, a wide axial canal above the manubrium in adults, and eggs that developed into actinula-like larval stages on the manubrium. In M. gibbesii , 5–6 tentacles were said to arise from each marginal bulb, no wide axial canal was known to exist above the manubrium in adults, and no actinula-like larval stages were thought to occur on the manubrium. The latter distinction has been shown here to be inaccurate; some female medusae of M. gibbesii studied by us were observed with developing larval polyps just as in M. haeckelii . Meanwhile, the other two characters require further evaluation. Several medusae of M. gibbesii in our material possessed an axial canal, although it was narrow in each case. As for numbers of marginal tentacles, Werner (1955 a) reported as many as 8–9 on one or more of the marginal bulbs of M. haeckelii . Up to eight were seen on the bulbs of a large male of M. gibbesii examined during this study (ROMIZ B 4077). The hydroid stage of M. gibbesii is still too poorly known to reliably assess how it may differ, if at all, from that of M. haeckelii . Overall, it seems best to maintain the two as distinct species for now, an opinion expressed earlier by Schuchert (2006). Live medusae of M. gibbesii were examined and described by Mayer (1910). The umbrella was said to be highly contractile. When relaxed, the manubrium reached little more than half-way along the length of the subumbrellar cavity but when contacted, the mouth often extended beyond the velar opening. The endoderm of the marginal bulbs was dull-yellow in colour, while that of the manubrium varied from dull-yellow to dull-green. In M. haeckelii , marginal bulbs are brown while the manubrium is gray with dark-brown pigment granules (Schuchert 2006). Life cycle . The life cycle of Margelopsis gibbesii has yet to be adequately described. Werner (1954, 1955a, b, 1956) traced the remarkable life cycle of M. haeckelii , and it seems likely that : Published as part of Calder, Dale R. & Johnson, William S., 2015, Margelopsis gibbesii (McCrady, 1859) (Cnidaria, Hydrozoa): taxonomic review, and conservation of usage by designation of a lectotype, pp. 215-225 in Zootaxa 4052 (2) on pages 216-222, DOI: 10.11646/zootaxa.4052.2.5, http://zenodo.org/record/242944 : {"references": ["McCrady, J. (1859) Gymnopthalmata of Charleston Harbor. Proceedings of the Elliott Society of Natural History, 1, 103 - 221.", "Agassiz, L. (1849) Contributions to the natural history of the Acalephae of North America. Part I. - On the naked-eyed medusae of the shores of Massachusetts, in their perfect state of development. Memoirs of the American Academy of Arts and Sciences, 4, 221 - 316. http: // dx. doi. org / 10.2307 / 25058163", "Bedot, M. (1925) Materiaux pour servir a l'histoire des hydroides. 7 e periode (1901 a 1910). 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(1996) The marine fauna of New Zealand: athecate hydroids and their medusae (Cnidaria: Hydrozoa). New Zealand Oceanographic Institute Memoir, 106, 1 - 159."]} Text Antarc* Antarctic Bouvet Island North Atlantic South Atlantic Ocean Spitzbergen DataCite Metadata Store (German National Library of Science and Technology) Antarctic Canada Norway Indian New Zealand Petersen ENVELOPE(-101.250,-101.250,-71.917,-71.917) Bouvet ENVELOPE(3.358,3.358,-54.422,-54.422) Elliott ENVELOPE(102.867,102.867,-65.867,-65.867) Medusa ENVELOPE(157.417,157.417,-79.633,-79.633) Bouvet Island ENVELOPE(3.358,3.358,-54.422,-54.422) Hampton ENVELOPE(-70.100,-70.100,-69.333,-69.333) Chapel Hill ENVELOPE(-57.976,-57.976,-63.685,-63.685) Sever ENVELOPE(166.083,166.083,62.917,62.917) Butterworth ENVELOPE(66.733,66.733,-70.700,-70.700) Oyster Creek ENVELOPE(-110.908,-110.908,78.348,78.348) Ritchie ENVELOPE(-128.387,-128.387,54.916,54.916) Cartwright ENVELOPE(-57.018,-57.018,53.708,53.708) |