Pennaria disticha Goldfuss, 1820

Calder, Dale R., 2019, On a collection of hydroids (Cnidaria, Hydrozoa) from the southwest coast of Florida, USA, Zootaxa 4689 (1), pp. 1-141 : 8

Pennaria disticha Goldfuss, 1820

Fig. 1a

Pennaria disticha Goldfuss, 1820: 89 .

Pennaria gibbosa L. Agassiz, 1860 : pl. 15, figs. 1, 2.—L. Agassiz, 1862: 278.—A. Agassiz, 1865: 186.

Pennaria symmetrica . — Jäderholm, 1896: 5.

Pennaria tiarella .— Mayer, 1910a: 25, text-fig. 2, pl. 1, fig. 5.— Fraser, 1943: 87; 1944: 84.— Menzel, 1956: 2.— Joyce, 1961: 45, pl. 7, figs. 3, 4.— Shier, 1965: 28, pl. 14.

Pennaria .— Wallace, 1909: 137.

Pennaria sp. — Joyce, 1961: 44, pl. 7, figs. 1, 2.— Shier, 1965: 30, pls. 15, 16.

Type locality. Italy: Gulf of Naples (see Calder 2013: 7) .

Material examined. Naples ( FL), Doctors Pass, north jetty, channel side, on boulder, 26°10’29.14”N, 81°48’53.45”W, ELWS, 06 December 2017, two colony fragments, up to 3.7 cm high, without gonophores, coll. D. Calder, ROMIZ B4345.— Sanibel Island , beach at Lighthouse Point, on tube of the polychaete Diopatra cuprea , ELWS, 02 January 2018, two colony fragments, up to 1.7 cm high, without gonophores, coll. D. Calder, ROMIZ B4346 .— Fort Myers Beach , 26°27’21”N, 81°57’45”W, detached at water’s edge, 16 March 2018, 18° C, 34‰, two colonies, up to 12 cm high, without gonophores, coll. D. Calder, ROMIZ B4347 .— Sanibel Island , beach at Lighthouse Point, 26°26’38”N, 82°01’36”W, on stranded Thalassia , 28 March 2018, 21° C, 35‰, one colony, 1.1 cm high, with gonophores, coll. D. Calder, ROMIZ B4348 .— Fort Myers Beach , Salty Sam’s Marina, 26°27’21.7”N, 81°56’34.6”W, on floating dock, <0.1 m, 29° C, 25‰, 27 August 2018, several colony fragments, up to 14 cm high, with gonophores, coll. D. Calder, ROMIZ B4418 .

Remarks. Hydroids identified as Pennaria disticha Goldfuss, 1820 are widespread, conspicuous, distinctive, often abundant, and easily collected in shallow, warm waters along the Atlantic coast of North America. This relatively large species was one of the earliest hydroids to be reported from the region, having been discovered in the eastern United States as early as the 1850s by Ayres (1852, as Globiceps tiarella ), Leidy (1855, as Eucoryne elegans ), and McCrady (1859, as Pennaria tiarella ). As noted in introductory remarks above, it was also the first hydroid to have been reported from the southwest coast of Florida (by L. Agassiz 1860, 1862, as P. gibbosa ). A frequent component of shallow fouling communities in tropical and temperate regions, P. disticha is held to be an invasive species with a particularly wide geographic distribution. Barcoding techniques have recently provided evidence of a complex of cryptic species within hydroids identified as P. disticha ( Miglietta et al. 2015, 2018a), but the taxonomy and nomenclature of the various lineages presently remain unresolved. While a name change may be necessary in the future, current usage of the binomen P. disticha is maintained for the species here.

The hydroid stage of this species, often identified in the western North Atlantic region as P. tiarella ( Ayres, 1852) , has been widely utilized in morphological, experimental, and ecological studies. Many of the distribution records listed below are based on specimens utilized in such work. If that hydroid proves to be a species distinct from the European P. disticha , and morphological differences in colonies from the two regions have indeed been noted ( Wallace 1909; Mayer 1910a; Brinckmann-Voss 1970), the name P. tiarella will be upheld as valid for at least one population here. The possible existence of cryptic species in Pennaria Goldfuss, 1820 on the east coast of the United States is suggested from the molecular work of Miglietta et al. (2015, 2018a) and from earlier morphological accounts such as those of Hargitt (1900) and Mayer (1910a).

More detailed accounts of this species are given elsewhere ( Calder 1988, 2010, 2013; Schuchert 2006).

Reported distribution. Gulf coast of Florida. Key West (L. Agassiz 1860: pl. 15, figs. 1, 2, as Pennaria gibbosa ; L. Agassiz 1862: 280, as Pennaria gibbosa ; A. Agassiz 1865: 186, as Pennaria gibbosa ; Jäderholm 1896: 5, as Pennaria symmetrica ; Fraser 1943: 87, as Pennaria tiarella ).—Dry Tortugas ( Wallace 1909: 137, as Pennaria ; Mayer 1910a: pl. 1, fig. 5, text-fig. 2, as Pennaria tiarella ; Fraser 1944: 86, as Pennaria tiarella ).—Florida Reefs ( Mayer 1910a: 27, as Pennaria tiarella ).—St. George Sound—Apalachee Bay region ( Menzel 1956: 2, as Pennaria tiarella ).—Seahorse Key ( Joyce 1961: 44, 45, as Pennaria sp. A and Pennaria tiarella ).—Cape San Blas area ( Shier 1965: 28, 30, as Pennaria tiarella and Pennaria sp.).

Elsewhere in western North Atlantic. USA: New York, Long Island, Sag Harbor ( Ayres 1852: 195, as Globiceps tiarella ).— USA: Rhode Island, Point Judith ( Leidy 1855: 136, as Eucoryne elegans ).— USA: New Jersey, Great Egg Harbor ( Leidy 1855: 138, as Eucoryne elegans ).— USA: South Carolina, Charleston Harbor ( McCrady 1859: 153, as Pennaria tiarella ).— Haiti: Jérémie (A. Agassiz 1865: 186, as Pennaria gibbosa ; Fraser 1944: 86, as Pennaria tiarella ).— USA: Massachusetts, Massachusetts Bay (medusa) + Suisconset (=Siasconset) (medusa) + Naushon (medusa) + Beverly (hydroid) + West Yarmouth (hydroid) + Nahant (hydroid) (A. Agassiz 1865: 189, as Pennaria tiarella ).— USA: Rhode Island, Newport (medusa) (A. Agassiz 1865: 189, as Pennaria tiarella ).— USA: Massachusetts: Woods Hole ( Verrill 1874d: 520, as Pennaria tiarella ; Bumpus 1898: 857, as Pennaria tiarella ; Smallwood 1899: 861, as Pennaria tiarella ; Hargitt 1900: 388, as Pennaria tiarella ; Hargitt 1901a: 311, as Pennaria tiarella ; Nutting 1901: 337, as Pennaria tiarella ; Weill 1934: 376, as Pennaria tiarella ; Puckett 1936: 393, as Pennaria tiarella ; Weill 1937: 1750, as Pennaria tiarella ; Petersen 1964: 18, as Pennaria tiarella ; Wyttenbach et al. 1973: 364, as Pennaria tiarella ).— USA: New Jersey, Great Egg Harbor ( Verrill 1874d: 735, as Pennaria tiarella ).— USA: Connecticut, near New Haven ( Verrill 1874d: 735, as Pennaria tiarella ).— USA: Massachusetts, Vineyard Sound, low water to 10 ftm (18 m), and floating algae ( Verrill 1874d: 735, as Pennaria tiarella ).— Cuba: Bahía Honda ( Clarke 1879: 240, as Pennaria symmetrica ).— USA: Massachusetts, Vineyard Sound ( Verrill & Rathbun 1880: 230, as Globiceps tiarella ).— USA: North Carolina, Beaufort ( Brooks 1882: 136, as Pennaria tiarella ).— USA: North Carolina, outside Fort Macon ( Brooks 1882: 144, as Pennaria inornata ).— Saint-Barthélemy ( Jäderholm 1903: 264, as Pennaria symmetrica ).— Bermuda ( Congdon 1907: 464, as Pennaria tiarella ; Weill 1937: 1750, as Pennaria tiarella ; Calder 1986: 132, as Halocordyle disticha ).— USA: Rhode Island, Newport ( Mayer 1910a: 25, pl. 1, figs. 2–4, as Pennaria tiarella ).— USA: North Carolina, Beaufort area ( Wilson 1911: 282, as Pennaria tiarella ).— USA: North Carolina, Morehead City + Beaufort ( Fraser 1912b: 355, as Pennaria tiarella ).— USA: Massachusetts, Woods Hole and vicinity + Vineyard Sound + Buzzards Bay ( Sumner et al. 1913: 561, as Pennaria tiarella ).— Virgin Islands of the United States: St. Thomas, Charlotte Amalie ( Stechow 1919: 7, as Pennaria tiarella ).— Bermuda: Hamilton Harbour + Great Sound + flats outside ( Bennitt 1922: 243, as Pennaria tiarella ).— USA: Massachusetts, Woods Hole region, on eelgrass, rocks and rockweed, pilings ( Allee 1923: 175, as Pennaria tiarella ).— USA: Florida, “Tablot” Island (= Talbot Island) ( Fraser 1933: 262, as Pennaria tiarella ).— USA: New York, Great South Bay ( Conard 1935: 449, as Pennaria tiarella ).— USA: Massachusetts, Woods Hole ( Baker 1936: 251, as Pennaria tiarella ; Ballard 1942: 329, as Pennaria tiarella ; Kepner 1943: 310, as Pennaria tiarella ).— Venezuela: Puerto Cabello ( Leloup 1937: 92, footnote, as Pennaria disticha ).— USA: North Carolina, Beaufort ( McDougall 1943: 336, as Pennaria tiarella ; Kepner 1943: 299, as Pennaria tiarella ; Maturo 1959: 123, as Pennaria tiarella ; Sutherland 1974: 861, as Pennaria tiarella ; Sutherland & Karlson 1977: 427, as Pennaria tiarella ; Martin & Thomas 1977: 198, as Pennaria tiarella ; Karlson 1978: 230, as Pennaria tiarella ; Sutherland 1978: 258, as Pennaria tiarella ; Sutherland 1981: 503, as Pennaria tiarella ; Walch et al. 1986: 353, as Pennaria tiarella ; Martin & Archer 1986a: 486, as Pennaria tiarella ; Holm et al. 1997: 192, as Pennaria tiarella ).— USA: Massachusetts, Buzzards Bay off Parkers Island, 11 ftm (20 m) + Vineyard Sound near West Chop Light, 14 ftm (26 m) ( Fraser 1944: 86, as Pennaria tiarella ).— USA: Rhode Island, Narragansett Bay off Prudence Light, 14.5 ftm (27 m) ( Fraser 1944: 86, as Pennaria tiarella ).— USA: North Carolina, off Cape Hatteras, 35°25’30”N, 75°20’30”W, 15 ftm (27 m) + 35°20’40”N, 75°18’40”W, 16 ftm (29 m) + Ocracoke ( Fraser 1944: 86, as Pennaria tiarella ).— USA: Florida, Biscayne Bay ( Weiss 1947: 57, as Pennaria tiarella ; Weiss 1948: 158, as Pennaria tiarella ; Jones 2002: 218, as Halocordyle disticha ).—Unstated location ( Woods Hole Oceanographic Institution 1952: 187, as Pennaria tiarella ).— Bermuda: vicinity of the Bermuda Biological Station ( Cowden 1965: 870, as Pennaria tiarella ; Martin & Thomas 1977: 198, as Pennaria tiarella ).— USA: Virginia, Hampton Roads, Norfolk, Norfolk Naval Base Pier 12, on fouling panels, 5 m ( Calder & Brehmer 1967: 153, as Pennaria tiarella ).— Jamaica: Kingston ( Vervoort 1968: 4, as Halocordyle disticha ).— Virgin Islands of the United States: St. Thomas, sound ( Vervoort 1968: 4, as Halocordyle disticha ).— Barbados ( Vervoort 1968: 4, as Halocordyle disticha ).— Venezuela: La Guaira + Puerto Cabello ( Vervoort 1968: 4, as Halocordyle disticha ).— Panama: Colón ( Vervoort 1968: 4, as Halocordyle disticha ).— Costa Rica: Limón ( Vervoort 1968: 4, as Halocordyle disticha ).— Curaçao: Piscadera Bay ( Vervoort 1968: 5, as Halocordyle disticha ).— Bermuda: Flatts Inlet ( Summers & Haynes 1969: 82, as Pennaria tiarella ; Summers 1970: 117, as Pennaria tiarella ; 1972: 229, as Pennaria tiarella ; Keough & Summers 1976: 507, as Pennaria tiarella ; Lesh-Laurie 1976: 366, as Pennaria tiarella ).— USA: Virginia, Perrin + Gloucester Point + Norfolk Naval Base + Cape Charles ( Calder 1971: 25, as Halocordyle disticha ).— USA: Virginia, York River, Big Mumford Island, 37°16’N, 76°31’W, on Zostera ( Marsh 1973: 93, as Halocordyle tiarella ).— USA: Gulf Stream off South Carolina, 32°00’N, 79°00’W, on Sargassum pteropleuron ( Rackley 1974: 14) .— Colombia: Santa Marta area ( Wedler 1975: 340, as Halocordyle disticha ; Bandel & Wedler 1987: 39, as Halocordyle disticha ).— Jamaica ( Mergner 1977: 122, as Halocordyle disticha ; 1987: 187, as Halocordyle disticha ).— Colombia ( Mergner 1977: 122, as Halocordyle disticha ; 1987: 187, as Halocordyle disticha ).— Costa Rica: east coast ( Mergner 1977: 122, as Halocordyle disticha ; 1987: 187, as Halocordyle disticha ).— USA: Florida, southeast coast ( Mergner 1977: 122, as Halocordyle disticha ; 1987: 187, as Halocordyle disticha ).— USA: Massachusetts, Nonamesset Island (41°31’N, 70°40’4”W) ( Osman 1978: 398, as Pennaria tiarella ).— USA: South Carolina, Murrells Inlet + Beaufort River + Calibogue Sound ( Calder & Hester 1978: 88, as Halocordyle disticha ).— USA: North Carolina, Wrightsville ( Bynum 1980: 228, as Pennaria tiarella ).— USA: North Carolina, Morehead City ( Martin & Thomas 1980: 27, as Pennaria tiarella ; Martin & Thomas 1981a: 303, as Pennaria tiarella ; Martin & Thomas 1981b: 304, as Pennaria tiarella ; Martin & Thomas 1983: 18, as Pennaria tiarella ; Martin & Archer 1986b: 116, as Pennaria tiarella ; Clark & Cook 1986: 406, as Halocordyle disticha and Pennaria tiarella ; Martin 1987: 325, as Halocordyl (sic) disticha ; Kolberg & Martin 1988: 250, as Halocordyle disticha ; Martin 1988a: 321, as Halocordyle disticha ; Martin 1988b: 67, as Halocordyle disticha ; Martin 1990: 11, as Pennaria tiarella ; Martin 1991: 76, as Pennaria tiarella ; Martin 1992: 432, as Pennaria tiarella ; Brumwell & Martin 1996: 14, as Pennaria tiarella ; Martin & Archer 1997: 42, as Pennaria tiarella ; Martin 2000: 243, as Pennaria tiarella ).— Belize: Carrie Bow Cay region (medusa) ( Larson 1983: 254, as Halocordyle disticha ).— USA: North Carolina, Wrightsville Beach, floating docks ( Hotchkiss et al. 1984: 718, as Pennaria tiarella ).— USA: Florida, Sebastian Inlet ( Winston 1982: 164, as Pennaria tiarella ).— Belize: Carrie Bow Cay ( Spracklin 1982: 240, as Halocordyle disticha ).— Colombia: Bahía de Cartagena ( Flórez González 1983: 123, as Halocordyle disticha ).— Dominican Republic: south coast ( Williams et al. 1983: 43, as Halocordyle disticha ).— USA: South Carolina, Murrells Inlet, jetties + Murrells Inlet, Capt. Dick’s marina, floating docks ( Fox & Ruppert 1985: 93, 104, as Halocordyle disticha ).— Puerto Rico: Isabela + La Parguera ( Wedler & Larson 1986: 73, as Halocordyle disticha ).— Bermuda: Coney Island ( Clark & Cook 1986: 406, as Halocordyle disticha and Pennaria tiarella ).— USA: North Carolina, coast ( Edwards et al. 1987: 381, as Halocordyle disticha ).— USA: North Carolina, Wrightsville Beach ( Thomas et al. 1987: 92, as Halocordyle disticha ).— USA: North Carolina coast ( Edwards et al. 1987: 381, as Halocordyle disticha ).— Puerto Rico: Mona Island + Desecheo Island ( Larson 1987: 514, as Halocordyle disticha ).— British Virgin Islands: Virgin Gorda ( Larson 1987: 514, as Halocordyle disticha ).— Bermuda: Castle Harbour near Tuckers Town, 7 m + Flatts Inlet, 3 m + Whalebone Bay, 1 m ( Calder 1988, as Halocordyle disticha ).— USA: South Carolina, continental shelf, fouling plates ( Van Dolah et al. 1988: 684, as Halocordyle disticha ).— USA: South Carolina, coastal areas, in stomachs of Atlantic spadefish ( Hayse 1990: 81, as Halocordyle disticha ).— Belize: Twin Cays, on Rhizophora ( Ellison & Farnsworth 1990: 96, as Halocordyle disticha ).— Belize: Big Creek, on Rhizophora + Lark Cay, on Rhizophora + Northeast Cay, on Rhizophora + Twin Cays, on Rhizophora ( Ellison & Farnsworth 1992: 90, as Halocordyle disticha ).— Colombia: Bahía de Chengue, on Rhizophora ( Reyes & Campos 1992: 108, as Halocordyle disticha ).— USA: Florida, Boca Raton, on nearshore artificial reef ( Cummings 1994: 1208, as Pennaria sp.).— USA: North Carolina, Wilmington ( Martin & Archer 1997: 42, as Pennaria tiarella ).— Cuba: Ciudad de La Habana province ( Ortiz 2001b: 68, as Halocordyle disticha ).— Costa Rica: Limón ( Kelmo & Vargas 2002: 603).— Panama: Colón, Fort Sherman dock + Colón, Isla Margareta + Galeta + Bocas del Toro area ( Calder & Kirkendale 2005: 480).—French Lesser Antilles: Guadeloupe, Basse-Terre, N of Malendure, 16°10´25.00”N, 61°46´58.00”W + Basse-Terre, Petite Anse, 16°05´47.00”N, 61°46´17.00”W + Basse- Terre, Anse à la Barque, 16°05´21”N, 61°46´00”W ( Galea 2008: 13).—French Lesser Antilles: Les Saintes, Terre-de-Haut, Pompierre Bay, 15°52´25”N, 61°34´15”W + Terre-de-Haut, Pain de Sucre, 15°51´45”N, 61°35´60”W ( Galea 2008: 13).— Cuba: Golfo de Batabanó ( Castellanos-Iglesias et al. 2011: 14, as Halochordyle (sic) disticha ).— Cuba: Golfo de Ana María ( Rodríguez-Viera et al. 2012: 33).— USA: Florida, Fort Pierce Inlet, north jetty, 0.1 m ( Calder 2013: 7).—French Lesser Antilles: Martinique ( Galea 2013: 49).— Panama: Bocas del Toro ( Miglietta et al. 2015: 5).— Honduras ( Miglietta et al. 2015: 5).— USA: Florida, Fort Pierce ( Miglietta et al. 2015: 5).— USA: North Carolina, Beaufort ( Miglietta et al. 2015: 5).—Caribbean Sea ( Wedler 2017b: 69, figs. 50—54).— Mexico: Alacranes Reef, on sponges, corals, molluscs, ascidians, rocks, artificial reefs ( Mendoza-Becerril et al. 2018b: 129).— Cuba: Havana, coral reef system west of the city (Castellanos et al. 2018: Supplementary Table S2).— USA: Florida, Fort Pierce ( Miglietta et al. 2018a: published online, no assigned pages).— USA: North Carolina, Beaufort ( Miglietta et al. 2018a: published online, no assigned pages).— Sint Eustatius ( Miglietta et al. 2018a: published online, no assigned pages).— Honduras: Caribbean coast ( Miglietta et al. 2018a: published online, no assigned pages).— USA (?): Gulf of Mexico ( Miglietta et al. 2018a: published online, no assigned pages).— Panama: Bocas del Toro area ( Miglietta et al. 2018a: published online, no assigned pages).— Panama: Bocas del Toro area, Crawl Cay + Swan’s Cay ( Miglietta et al. 2018b: 108).

Figures

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.