On a collection of hydroids (Cnidaria, Hydrozoa) from the southwest coast of Florida, USA Author Calder, Dale R. text Zootaxa 2019 2019-10-25 4689 1 1 141 journal article 25128 10.11646/zootaxa.4689.1.1 a63613b4-e611-42e8-89cd-2d3aae126759 1175-5326 3519047 41BFBBDF-41AD-4329-B6B9-CF38D64815A6 Turritopsis nutricula McCrady, 1857 Fig. 1e Oceania ( Turritopsis ) nutricula McCrady, 1857: 55 , pl. 4, figs. 1–10, 12–15, 28a, pl. 5, figs. 11, 16 18, 28b [medusa stage]. Turritopsis nutricula McCrady, 1857: 58 [medusa stage].— Mayer, 1900b: 39 [medusa stage]; 1910a: 143, text-figs. 75, 76, pl. 15, figs. 12, 13 [hydroid & medusa stages]. Turritopsis nutricola .— Vanhöffen, 1916: 418 [medusa stage] [incorrect subsequent spelling]. Type locality. USA : South Carolina , Charleston Harbor area ( McCrady 1857: 55 ) . Material examined. Fort Myers Beach, 26°27’27”N , 81°57’51”W , on detached barnacle cluster at water’s edge, 22 March 2018 , 21° C, 33.5‰, one colony, 2 mm high, without gonophores, coll. D. Calder, ROMIZ B4327. Remarks. The original account of Turritopsis nutricula by McCrady (1857) was based solely on the medusa stage. A previously unknown hydroid was linked to it in life cycle studies by Brooks (1883) . While he provided no illustration of either hydroid or medusa of the species in that work, an excellent one appeared in another publication three years later ( Brooks 1886 : pl. 37). The same drawing was included later in the monograph of Mayer (1910a : fig. 76). McCrady’s (1857) description of T. nutricula made no direct mention of the collection locale. However, his pioneering research on hydrozoans of the southeastern United States was carried out in Charleston Harbor and vicinity, South Carolina ( Stephens & Calder 1992 ). The type locality can therefore be taken to be the Charleston Harbor area. The species was described again in a classic paper on Hydrozoa of Charleston Harbor ( McCrady 1859 ), and specimens collected from Charleston, by McCrady, were included in a catalogue of the Museum of Comparative Zoology by A. Agassiz (1865) . In a report on anthoathecate hydroids of Bermuda ( Calder 1988 ), I wrote that the generic name Turritopsis Mc- Crady was threatened by a virtually unused subjective synonym ( Clavula Wright, 1859 ). McCrady’s paper, in which Turritopsis was first established, is now known to have been published in 1857 ( Calder et al. 1992 ) and not in 1859 as generally believed at the time. Turritopsis therefore has priority over Clavula , a name proposed by Wright (1959) in the July 1859 issue of Edinburgh New Philosophical Journal . For the same reason, the binomen Clavula gossii Wright, 1859 is not a nomenclatural threat to Turritopsis nutricula . Indeed, the two species names are no longer held to be synonyms ( Schuchert 2004 ). An account of the medusa of T. nutricula by A. Agassiz (1865) from Naushon, Massachusetts , was based on a misidentification ( Brooks 1882 ; Mayer 1910a ). Reports of the species (misspelled as T. nutricola ) by L. Agassiz (1862) from the same location, in collections by A. Agassiz, are likewise excluded from distribution records below as likely misidentifications. So too is a report of it from Massachusetts , based on the records of L. and A. Agassiz, by Verrill (1874d: 454 , 734). Recent fanciful accounts of the species as “the immortal jellyfish” have largely been based on misidentifications of the European T. dohrnii ( Weismann, 1883 ) . By the mid- and late 20 th century, both hydroid and medusa stages of T. nutricula were widely thought to be almost circumglobal in distribution ( Kramp 1961 ; Calder 1988 ). Evidence from recent reproductive and molecular studies ( Schuchert 2004 ; Miglietta et al . 2007 ; Miglietta & Lessios 2009 ; Miglietta 2016 ; Kubota & Nagai 2018 ) indicate that the species is far more restricted in distribution, occurring largely or exclusively in the warm-temperate western Atlantic. As noted by Schuchert (2016) , however, 16S sequences of specimens attributed to T. nutricula thus far are inconclusive in having been based on specimens collected at considerable distances from the type locality in South Carolina . Analyses of topotypic material are needed to confirm results of those sequences. Medusae of T. nutricula are dioecious and oviparous ( Schuchert 2004 ), thereby differing from those of the European T. polycirrha ( Keferstein, 1862 ) . Morphological characters distinguishing T. nutricula from T. polycirrha , the Mediterranean T. dohrnii , and the Pacific T. rubra ( Farquhar, 1895 ) are outlined in Schuchert (2004) , with differences being most apparent in medusae of the species. The hydroid colony examined here from Fort Myers Beach lacked medusa buds, with identification of it as T. nutricula being based largely on its occurrence within a biogeographic region related to that of the type locality of the species elsewhere in the southern United States . Moreover, medusae of T. nutricula have been reported nearby from the Dry Tortugas in southwest Florida ( Mayer 1900b , 1910a ; Vanhöffen 1916 ). Hydroids identified as the relatively deep water T. fascicularis Fraser, 1943 also occur in the region ( Fraser 1944 ; Calder 2013 ; Miglietta 2016 ), but they differ in having much larger, polysiphonic colonies. As for T. fascicularis , it has now been linked through DNA sequence data to Oceania armata Kölliker, 1853 ( Schuchert 2016 ) . Turritopsis nutricula has been reported infrequently in the Gulf of Mexico . As noted above, medusae of the species have been recorded at the Tortugas . Medusae were also collected in the Bahía de Campeche , Mexico , by Martell-Hernández et al . (2014). Earlier, Segura-Puertas (1992) reported the medusa stage from the Yucatan Shelf and the Mexican Caribbean, but it is unclear whether the species was collected in the Gulf of Mexico . The hydroid stage has been identified from coastal petroleum platforms in Texas ( Fotheringham 1981 ) and Louisiana ( Lewbel et al . 1987 ). Its reported range elsewhere in the western North Atlantic extends from southern Massachusetts to the southern Caribbean Sea (see records below). Records of T. nutricula from South America are summarized in Oliveira et al . (2016) . FIGURE 1. a, Pennaria disticha : part of branch, with pedicel and hydranth, Fort Myers Beach, ROMIZ B4347. Scale equals 0.5 mm. b, Corydendrium parasiticum : part of colony with a reduced hydranth, Southwest Florida Shelf, ROMIZ B449. Scale equals 0.5 mm. c, Rhizogeton sterreri : hydranth and stolon, Fort Myers Beach, ROMIZ B4326. Scale equals 0.2 mm. d, Rhizogeton sterreri : male gonophore and stolon, Fort Myers Beach, ROMIZ B4326. Scale equals 0.2 mm. e, Turritopsis nutricula : colony with two hydranths, Fort Myers Beach, ROMIZ B4327. Scale equals 0.2 mm. f, Cordylophora caspia : part of colony with hydranth and female gonophore, Caloosahatchee River at Fort Myers, ROMIZ B4329. Scale equals 0.2 mm. Reported distribution. Gulf coast of Florida. Tortugas (medusa) ( Mayer 1900b: 39 ; 1910a: 144 , captions to pl. 15, figs. 12, 13).— Tortugas (medusa) ( Vanhöffen 1916: 418 , as Turritopsis nutricola ). Elsewhere in western North Atlantic. USA : South Carolina , Charleston Harbor (medusa) ( McCrady 1857: 58 ; 1859: 128 ).— USA : Massachusetts , Buzzards Bay, Naushon (medusa) ( Fewkes 1881b: 149 , as Modeeria multitentaculata ).— USA : North Carolina , Beaufort, the most common medusa (medusa) ( Brooks 1882: 143 ).— USA : North Carolina , Beaufort (medusa) + Morehead City (hydroid) ( Brooks 1883: 465 ; 1886: 390 ).— Bermuda : Castle Harbor ( Fewkes 1883: 79 , 80, as Modeeria multitentaculata and Modeeria ( Turritopsis ) nutricula ).— USA : Virginia , Hampton Roads (medusa) ( Brooks 1886: 389 ).— USA : Rhode Island , Newport (medusa) ( Mayer 1900b: 39 ; 1910a : figure caption to plate 15).— Cuba : (medusa) ( Mayer 1900b: 39 ; 1910a: 144 ).— USA : South Carolina , Charleston Harbor (medusa) ( Mayer 1900b: 39 ; 1910a: 144 ).— USA : Massachusetts , Woods Hole (medusa) ( Nutting, 1901: 375 ).— Bahamas : (medusa) ( Mayer 1904: 10 ; 1910a: 144 ).— USA : Massachusetts , Woods Hole (medusa) ( Hargitt 1904: 37 ).— USA : North Carolina , Beaufort (medusa, juvenile hydroid reared from medusa) ( Brooks & Rittenhouse 1907: 433 , 437).— USA : North Carolina , on pilings of bridge between Beaufort and Morehead City, low water + Bogue Sound, 10 feet ( 3 m ) + Cape Lookout, on boathouse pilings ( Fraser 1912b: 345 ).— Virgin Islands of the United States : St. Thomas, Charlotte Amalie, surface, on algae on an old barge ( Stechow 1919: 12 , as Turritopsis nutricola ).— USA : Massachusetts , Woods Hole region (medusa) ( Fish 1925: 124 ).— Virgin Islands of the United States : St. Thomas, Charlotte Amalie, surface, on algae ( Fraser 1944: 38 ).—Northwest Atlantic Ocean: 26°56’N , 53°09’W , 1000 m (medusa, identification uncertain) ( Kramp 1959: 9 ).— USA : North Carolina , Beaufort region, inshore waters (medusa) ( Allwein 1967: 122 ).— Curaçao : Port + Schottegat, on algae ( Vervoort 1968: 6 ).— Panama : Colón , on algae on experimental plates ( Vervoort 1968: 6 ).— USA : Virginia , York River (Tue Marsh Light + Gloucester Point + Page’s Rock) + James River (Hampton Roads Middle Ground) + southern Chesapeake Bay ( Calder 1971: 30 ).— Colombia : Santa Marta area ( Wedler 1975: 333 ).— USA : South Carolina , estuaries ( Calder 1976: 169 ).— USA : South Carolina , Price Creek + Charleston Harbor + North Edisto River + St. Helena Sound + Colleton River ( Calder & Hester 1978: 89 ).— USA : South Carolina , Bull Bay + Price Creek + Inlet Creek + Charleston Harbor + North Edisto River (medusa) ( Calder & Hester 1978: 89 ).— USA : Texas , Buccaneer oil field, on oil platform ( Fotheringham 1981: 194 ).— Belize : Carrie Bow Cay, on reefs, mangroves, sand troughs ( Spracklin 1982: 248 ).— Colombia : Bahía de Cartagena ( Flórez González 1983: 123 ).— USA : South Carolina and Georgia , inner, middle and outer continental shelf ( Wenner et al . 1984: 20 , 39).— USA : South Carolina , North Inlet area, Town Creek and tributaries + Murrells Inlet, Capt. Dick’s Marina, floating docks + Charleston area + Folly River area, Oak Island , oyster reefs + Isle of Palms, marina, floating docks ( Fox & Ruppert 1985: 61 , 104, 141, 152, 177).— Bermuda : common inshore and on offshore buoy chains ( Calder 1986: 134 ).— Puerto Rico , La Parguera, on mangrove roots and sponges, 1–5 m ( Wedler & Larson 1986: 86 ).— USA : Louisiana , on coastal petroleum platforms ( Lewbel et al . 1987: 214 ).— Colombia : Santa Marta area ( Bandel & Wedler 1987: 39 ).— Bermuda : Whalebone Bay + Flatts Inlet ( Calder 1988: 8 ).— USA : South Carolina , continental shelf, fouling plates ( Van Dolah et al . 1988: 684 ).— USA : South Carolina , inner continental shelf, on artificial reef ( Wendt et al . 1989: 1116 ).— Belize : Twin Cays ( Calder 1991b: 223 ).— Mexico : Campeche Bank + Mexican Caribbean (medusa) (Segura-Puertas & Ordóñez- López 1994: 108).— Bermuda : Argus (=Plantagenet) Bank ( Calder 2000: 1134 ).— USA : North Carolina , Beaufort Inlet (medusa) ( Schuchert 2004: 323 ).— Panama : Mole Buoy, Atlantic entrance to canal + Colón , Fort Sherman dock, wood, 09°22’12”N , 79°56’59”W , 0-2 m + Colón , bridge near Fort Sherman, 09°17’33”N , 79°55’22”W , 0-1 m + Colón , Fort Sherman dock, marina, 09°20’57”N , 79°54’10”W , 0-2 m + Colón , Club Nautico, steel pilings, 09°21’51”N , 79°53’39”W , 0-1 m + Colón , Isla Margareta, Fort Randolph, shore, 09°23’15”N , 79°53’11”W , 0-1 m + Bocas del Toro area, Hospital Point, 09°20’00.7”N , 82°13’06.8”W , 0-2 m + Bocas del Toro area, Mangrove Inn, 09°19.52.6”N, 82°15’17.7’W, 2-3 m + Bocas del Toro area, Almirante pilings, 09°16.218’N , 82°23.382’W , 1-10 m + Bocas del Toro area, Hospital Point, 09°20’01.9”N , 82°13’07.7”W , 2-13 m + Bocas del Toro area, Crawl Cay, 09°15.261’N , 82°07.787’W , 2-4 m + Bocas del Toro area, Boca del Drago, no coordinates, 0-3 m ( Calder & Kirkendale 2005: 479 ).— USA : Massachusetts , Woods Hole ( Miglietta et al . 2007: 13 ).— USA : South Carolina , North Inlet estuary (medusa) (Marshalonis & Pinckney 2007: 1032).—French Lesser Antilles: Guadeloupe , Basse-Terre, Petite Anse, 16°05’47.00”N , 61°46’17.00”W , rocky shore ( Galea 2008: 9 , identification provisional, as Turritopsis cf. nutricula ).— USA : Massachusetts , Woods Hole ( Miglietta & Lessios 2009: 833 ).— Cuba : Bahía de Cochinos, 0.5 m , on Pecten sp. (Varela et al . 2010: 30, as Turritopsis nutricola ).—French Lesser Antilles: Martinique , Le Prêcheur, Les Jardins des Abîmes, 14.809044 , -61.228853 , 10–15 m , on sponge and worm tubes + Le Prêcheur, Pointe Lamare, 14.780461°, -61.211935°, 15–18 m , on Pennaria disticha ( Galea 2013: 6 ) .— Mexico : southern Gulf of Mexico (medusa) (Martell-Hernández et al . 2014: 23).— USA : Massachusetts , Woods Hole ( Devarapalli et al . 2014: 590 ).— USA : east coast ( Miglietta 2016: 431 ).—Caribbean Sea ( Wedler 2017b: 43 , figs. 38, 39A, B).— USA : Massachusetts , Woods Hole ( Kubota & Nagai 2018: 3 ).— USA : New Jersey , Barnegat Bay (medusa) ( Bologna et al . 2018: 222 ).— Cuba : Havana , coral reef system west of the city (Castellanos et al . 2018: Supplementary Table S2).