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INTRODUCTION
Bryozoans are sessile, aquatic, filter feed ing invertebrates that form colonies of different shapes, colors, and size. Their distribution can be influenced by substrate complexity, temperature, depth, currents, and salinity (Ryland, 1970). Bryozoans contribute to the food web of different marine ecosystems, they are important bioconstructors of habitats (Ryland, 1970; Bastos et al., 2018; Ramalho et al., 2018; Ramalho et al., 2021), and some species are being studied for the development of new drugs (Figuerola & Avila, 2019). The calcareous structures produced by most bryozoans generate rich fossil records, thus being also useful as palaeoindicators (e.g. Smith, 1995; Taylor, 2005; Berning, 2007; Ramalho et al., 2017; Bastos et al., 2018). Furthermore, bryo zoans are an important group of fouling organ isms encrusting ship hulls, pipes, cables, buoys, nets and almost any submerged hard substrate; fouling causes financial loss and is a potential mechanism of invasive species dispersal (Gordon et al., 2006; Bressy & Lejars, 2014).
The bryozoan fauna of Uruguay (Southwest Atlantic) has been poorly studied (López-Gappa & Lichtschein, 1988). The existing information is scattered among taxonomic and ecological publi cations, particularly scarce and fragmentary for coastal ecosystems and shallow sublittoral. The first records of bryozoans in Uruguayan waters were made by Busk (1881b, 1884, 1886), who described the species collected during the HMS Challenger expedition (February 1876). Almost one century after that pioneering contribution, the material collected by the RV Atlantis II cruise 60 (March 1971) provided the basis for the new information published by d’Hondt (1981, 1982, 1983), d’Hondt & Hayward (1981), Harmelin & d’Hondt (1982) and d’Hondt & Schopf (1984). Several ecological or general faunistic stud ies also mentioned bryozoans in shallow and coastal environments (Barattini & Ureta, 1961; Juanicó & Rodríguez-Moyano, 1976; Milstein et al., 1976; Riestra et al., 1992; Obenat et al., 2001; Giménez et al., 2005). In a compilation of the bryozoan fauna of the continental shelf and slope off Argentina, López-Gappa (2000) mapped most of the deep-water bryozoans from Uruguay. Another bibliographic compilation by Scarabino (2006) mentioned 13 taxa from marine and es tuarine Uruguayan waters (inner shelf). More recently López-Gappa et al. (2020) described a new species, Hippomonavella charrua, from Argentina and the continental shelf off Uruguay.
The aim of this study is to compile an updat ed checklist, bringing together all the published information on the Uruguayan marine fauna of bryozoans.
MATERIAL AND METHODS
Study area
The area considered is the Uruguayan coast line influenced by marine waters (Fig. 1), the ad jacent territorial waters in the Río de la Plata and inner continental shelf, as well as the Uruguayan Economic Exclusive Zone (URY EEZ), which extends 200 nautical miles off the Uruguayan coastline. Only one sampling site (station 242 of the cruise 60 of the RV Atlantis II) falling some miles off this zone is here included considering the forthcoming definitive enlargement of the URY EEZ.
Criteria for the construction of this checklist
The included species have been recorded from the area specified above. Scientific papers and book chapters taken into account are in dicated as “References for the area”. Full data of the stations of the HMS Challenger and RV Atlantis II cruises are provided in Appendix 1.
The bathymetric zonation regarding the dis tribution of species was defined as follows: shelf (0-200 m), upper slope (200-1500 m), lower slope (1500-3000 m) and abyssal plain (3000-5500 m).
The synonymy includes the original descrip tion and, when it existed: a) references which re-describe the type material from Uruguay (e.g. Turritigera stellata Busk, 1884), and b) species described from Uruguay that have been later syn onymized to other species (e.g. Formosocellaria abyssicola d’Hondt, 1981). For species having a long or relatively complex taxonomic history (which we do not fully detail here) we also list at least one reference detailing its synonymy.
References to the geographic distribution of littoral species include: a) general/major re visions containing comprehensive treatment of these, or b) works extending considerably (i.e. thousands of kilometers) the range of a given species. In the case of deep-sea species we have considered all available references.
It must be noted that the station 320 of the HMS Challenger falls exactly on the maritime limit between Argentina and Uruguay and there fore the species recorded there are considered as part of the faunistic inventory of both countries.
RESULTS
Class GYMNOLAEMATA Allman, 1856
Order CHEILOSTOMATIDA Busk, 1852a
Suborder MEMBRANIPORINA Ortmann, 1890
Superfamily MEMBRANIPOROIDEA Busk, 1854
Family ELECTRIDAE Stach, 1937
Genus Electra Lamouroux, 1816
Electra monostachys (Busk, 1854)
Membranipora monostachys Busk, 1854: 61.
For synonymy see: Hayward & Ryland (1998); Occhipinti Ambrogi & d’Hondt (1981).
Geographic distribution: Britain (Busk, 1854); Netherlands, North Sea, Wadden Sea (De Blauwe, 2009), North Atlantic Ocean (Hayward, 2001). South Atlantic: Brazil (Rio de Janeiro, São Paulo) (Marcus, 1938; Ramalho, 2006); Argentina (Calvet, 1904; López-Gappa, 2000); Uruguay: 35°31’S, 55°36’ - 55°48’W (Obenat et al., 2001).
Bathymetric distribution in Uruguayan waters: Intertidal-shallow subtidal.
References for the area: Obenat et al. (2001); Scarabino (2006).
Remarks: Encrusting species occurring on stones or shells in estuarine (under high and fluctuating salinity conditions) and lagoonal coast or offshore environments (Marcus, 1938; Hayward & Ryland, 1998). In Uruguay it was found at outer Río de la Plata with salinity varying 26.7 - 29.9 ppt. This species, identified by one of us (JLG), was found growing on the tubes of Phyllochaetopterus socialis (Polychaeta) (Obenat et al., 2001). Considered as cryptogenic in Uruguay-Argentina by Schwindt et al. (2020).
Genus Conopeum Gray, 1848
Conopeum reticulum (Linnaeus, 1767)
Millepora reticulum Linnaeus, 1767: 1284.
For synonymy see: Harmer (1926: 211); Osburn (1940: 350); Osburn (1950: 31); Marcus (1941: 15).
Geographic distribution: widely distribut ed. Pacific: from Alaska to Southern California (Osburn, 1950). Atlantic: Brazil (Espírito Santo, São Paulo, Paraná, and Santa Catarina States) (Ridley, 1881; Marcus, 1938, 1939, 1941); Argentina (López-Gappa, 2000; Orensanz et al., 2002; Figuerola et al., 2014); Uruguay (Obenat et al., 2001; Scarabino, 2006).
Bathymetric distribution in Uruguayan waters: Intertidal-shallow subtidal.
References for the area: Riestra et al. (1992), Obenat et al. (2001), Orensanz et al. (2002), Giménez et al. (2005), and Scarabino (2006), also see below.
Remarks: This species was initially listed by Obenat et al., 2001 on the basis of mate rial collected alive on tubes of the polychaete Phyllochaetopterus socialis and identified by one of us (JLG). Frequently found in brackish envi ronments, in the lower intertidal zone, and up to shallow subtidal waters; its most common sub strates are oyster valves, stones, and other hard substrates (Hayward & Ryland, 1998). Orensanz et al. (2002) and Schwindt et al. (2020) consid ered this species as cryptogenic, but its presence in the area since the mid-Holocene has been recently confirmed (López-Gappa & Pereyra, 2020). Recent observations made by us indicate that the references made by Barattini & Ureta (1961) (as Membranipora tehuelcha, partim), Riestra et al. (1992) and Giménez et al. (2005) (all as Membranipora sp.) belong to this species, which is very abundant and widely distributed in the Uruguayan coast. Since we have found anoth er yet unidentified species of Membraniporidae in Uruguayan ports, it is not possible to assign with certainty the record by Calvo (1984) as Membranipora sp. to C. reticulum.
Family MEMBRANIPORIDAE Busk, 1852b
Genus Jellyella Taylor & Monks, 1997
Jellyella tuberculata (Bosc, 1802)
Flustra tuberculata Bosc, 1802: 118.
For synonymy see: Tilbrook et al. (2001); Florence et al. (2007).
Geographic distribution: Widespread, sub tropical (Taylor & Monks, 1997; Tilbrook et al., 2001; Florence et al., 2007). Atlantic: West coast of South Africa (Florence et al., 2007); Brazil (Pernambuco, Rocas Atoll, Espírito Santo, Rio de Janeiro, São Paulo and Rio Grande do Sul - d’Or bigny, 1842; Gliesch, 1925; Marcus, 1937, 1939 and 1955); Uruguay (see below).
Bathymetric distribution in Uruguayan waters: Intertidal-shallow subtidal.
References for the area: Barattini & Ureta (1961) as Membranipora tehuelcha, partim; Scarabino (2006).
Remarks: This species mainly encrusts fronds of algae, crustaceans and hydroids (Marcus, 1937; Tilbrook et al., 2001; Florence et al., 2007).
The Membraniporidae studied by Barattini & Ureta (1961) include both materials collected on shells (herein referred to C. reticulum) and on algae, referred by Scarabino (2006) and also here to J. tuberculata.
Suborder AETEINA Gordon & Bock in Cook et al., 2018
Superfamily AETEOIDEA Smitt, 1868
Family AETEIDAE Smitt, 1868
Genus Aetea Lamouroux, 1812
Aetea anguina (Linnaeus, 1758)
Sertularia anguina Linnaeus, 1758: 816.
For synonymy see: Osburn (1950); Marcus (1955); Hayward & Ryland (1998).
Geographic distribution: Widely distributed, including records from Brazil (Marcus, 1937, 1955; Braga, 1968; Ramalho, 2006), Uruguay (Obenat et al., 2001; Scarabino, 2006) and Argentina (e.g. Hastings, 1943; Amor & Pallares, 1965; López-Gappa & Lichtschein, 1988; López- Gappa, 2000; Figuerola et al., 2014) (also see Remarks below).
Bathymetric distribution in Uruguayan waters: 11-12 m.
References for the area: Obenat et al. (2001); Scarabino (2006).
Remarks: This species was listed by Obenat et al. (2001) based on material identified by one of us (JLG). The Southwest Atlantic materi al identified as A. anguina should be critically re-examined, as Vieira et al. (2016) stated that the Brazilian records of A. anguina by Vieira et al. (2008) and Almeida et al. (2015) belonged in stead to A. arcuata Winston & Hayward, 2012.
This bryozoan grows on tubes of the polychaete Phyllochaetopterus socialis (Obenat et al., 2001; Scarabino, 2006). It was considered as crypto genic in Uruguay-Argentina by Schwindt et al. (2020).
Suborder FLUSTRINA Smitt, 1868
Superfamily CALLOPOROIDEA Norman, 1903
Family CHAPERIIDAE Jullien, 1888
Genus Chaperiopsis Uttley, 1949
Chaperiopsis erecta (Busk, 1884)
Membranipora galeata var. erecta Busk, 1884: 65.
For synonymy see: Hayward & Thorpe (1988).
Geographic distribution: Only known from Argentine Basin, Argentina-Uruguay (Busk, 1884; Hayward & Thorpe, 1988; López-Gappa, 2000; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 1097 m (=600 ft, Busk, 1884).
References for the area: Busk (1884); López- Gappa (2000).
Remarks: Colonies collected during the HMS Challenger expedition at station (stn.) 320. The bathymetric range of this species is striking, as it was also found by the Discovery expedition at a depth of 199 m (stn. 474, South Georgia).
Family FARCIMINARIIDAE Busk, 1852b
Genus Farciminaria Busk, 1852b
Farciminaria cribraria Busk, 1884
Farciminaria cribraria Busk, 1884: 49, pl. 5, fig. 2.
Geographic distribution: Abyssal plain of Argentine Basin (Uruguay) (Busk, 1884). Also recorded for the North Atlantic (39°19.5’N, 33°47’W, 3360 m) by d’Hondt (1975, 1982).
Bathymetric distribution in Uruguayan waters: 3475 m.
References for the area: Busk (1884); López- Gappa (2000, as Columnella cribraria).
Remarks: Collected at HMS Challenger stn. 323, which is the type-locality of this species (Busk, 1884). Recorded as Columnella cribraria by d’Hondt (1975, 1982).
Genus Columnella Levinsen, 1914
Columnella magna (Busk, 1884)
Farciminaria magna Busk, 1884: 49, pl. 5, fig. 1.
Farciminaria magna var. armata Busk, 1884: 50, pl. 31, fig. 1.
For synonymy see: Hayward (1981).
Geographic distribution: This species was recorded from the North and South Atlantic, western Indian Ocean (Hayward & Cook, 1979; Hayward, 1981; d’Hondt, 1982; Figuerola et al., 2014) and Argentine Basin: 35°39’S, 50°47’W (Busk, 1884; Hayward, 1981; d’Hondt, 1983; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 3475 m.
References for the area: Busk (1884); López- Gappa (2000, as Columnella magna var. armata).
Remarks: collected at HMS Challenger stn. 323, which is its type-locality (Busk, 1884). This species is recorded from deep waters.
Columnella gracilis (Busk, 1884)
Farciminaria gracilis Busk, 1884: 50, pl. V, fig. 3.
For synonymy see: d’Hondt (1981: 13).
Geographic distribution: Brazil (Alagoas) (Busk, 1884); Argentine Basin, Argentina- Uruguay (d’Hondt, 1981, 1983; López-Gappa, 2000; Figuerola et al., 2014); North Atlantic (d’Hondt, 1983).
Bathymetric distribution in Uruguayan waters: 3815-3822 m.
References for the area: d’Hondt (1981); López-Gappa (2000).
Remarks: Collected at RV Atlantis II, cruise 60, stn. 243 (d’Hondt, 1981; López-Gappa, 2000). This is a deep-water species.
Family FOVEOLARIIDAE Gordon & Winston in Winston, 2005
Genus Foveolaria Busk, 1884
Foveolaria elliptica Busk, 1884
Foveolaria elliptica Busk, 1884: 68, pl. 23, fig. 5.
For synonymy see: Gordon (1986) and López- Gappa & Lichtschein (1990).
Geographic distribution: Australia, New Zealand, Southern Chile (Moyano, 1974; Gordon, 1986; Figuerola et al., 2014). Atlantic: Uruguay and Argentina (Busk, 1884; López-Gappa, 2000; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 1097 m.
References for the area: Busk (1884); López- Gappa (2000).
Remarks: Collected by the HMS Challenger at Bass Strait (Australia) and the continen tal slope off Argentina and Uruguay (stn. 320) (Busk, 1884). This genus is recorded only for the Southern Hemisphere (South America to New Zealand) (Winston, 2005). Busk (1884) identi fied these specimens (from Australia and the Argentine basin) as belonging to the same spe cies, without designating a holotype. The pres ent identification therefore depends on the con firmation that all specimens examined by Busk (1884) belong to the same taxon. A lectotype was not designated by subsequent authors who studied or mentioned this species (Canu, 1900; Gordon, 1986).
Foveolaria terrifica (Hincks, 1881)
Membranipora terrifica Hincks, 1881: 147, pl. 8, fig. 5.
Foveolaria falcifera, Busk, 1884: 69, pl. 15, fig. 6.
For synonymy see: López-Gappa & Lichtschein (1990).
Geographic distribution: Chile, Magellan Strait, Burdwood Bank, Patagonian shelf; Uruguay (Busk, 1884; López-Gappa & Lichtschein, 1990; López-Gappa, 2000; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 1097 m.
References for the area: Busk (1884); López- Gappa (2000).
Remarks: collected at HMS Challenger stn. 320, which is the type-locality of F. falcifera Busk, 1884, a species synonymized with F. terrifica by López-Gappa & Lichtschein (1990). Frequently found encrusting bilaminar erect bryozoans (López-Gappa & Lichtschein, 1990).
Superfamily FLUSTROIDEA Fleming, 1828
Family FLUSTRIDAE Fleming, 1828
Genus Securiflustra Silén, 1941
Securiflustra bifoliata (d’Hondt, 1981)
Nematoflustra bifoliata d’Hondt, 1981: 11, pl. III, 5-6, Text fig. 2.
For synonymy see: López-Gappa (1982).
Geographic distribution: Argentine Basin; Burdwood Bank; Malvinas Islands (d’Hondt, 1981; López-Gappa, 2000; Moyano, 2005; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 2707 m.
References for the area: d’Hondt (1981); López-Gappa (2000).
Remarks: Collected at RV Atlantis II, cruise 60, stn. 245, which is the type-locality of this spe cies (d’Hondt, 1981). López-Gappa (1982) trans ferred it to the genus Securiflustra.
Family SINOFLUSTRIDAE Gordon, 2009
Genus Membraniporopsis Liu in Liu et al., 1999
Membraniporopsis tubigera (Osburn, 1940)
Conopeum tubigerum Osburn, 1940: 352, pl. 2, figs 12, 13.
Membraniporopsis tubigera, Gordon et al., 2006: 336, figs 1, 3-6.
Geographic distribution: Japan Sea, Australia, New Zealand, Puerto Rico, Texas, Florida, Brazil (Espírito Santo, São Paulo, Paraná and Santa Catarina), Uruguay (33°54’S, 53°30’W) (Osburn, 1940; Allen, 1953; Kubanin, 1977; Gordon et al., 2006; López-Gappa et al., 2010).
Bathymetric distribution in Uruguayan waters: Inner shelf (recorded during stranding events).
References for the area: López-Gappa et al. (2010).
Remarks: This invasive species develops small and foliaceous colonies that spread from an encrusting base. In Brazil, Uruguay and New Zealand the colonies accumulate as beach-drift during short periods (Gordon et al., 2006; López- Gappa et al., 2010).
Superfamily BUGULOIDEA Gray, 1848
Family BUGULIDAE Gray, 1848
Genus Camptoplites Harmer, 1923
Camptoplites reticulatus (Busk, 1881a)
Bugula reticulata Busk, 1881a: 12, pl. 1, fig. 7 (partim).
For synonymy see: d’Hondt (1981).
Geographic distribution: Southern Indian and Pacific Oceans; South West Atlantic (off Argentina and Uruguay) (Busk, 1881a; d’Hondt, 1981; López-Gappa, 2000; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 1097-2480 m.
References for the area: Busk (1881a); d’Hondt (1981); López-Gappa (2000).
Remarks: collected by HMS Challenger near Crozet Island, off Chile and in stn. 320 (Busk, 1884) and by RV Atlantis II, cruise 60, stns. 242, 245, 259 and 262 (d’Hondt, 1981). Species re corded from deep waters.
Camptoplites asymmetricus Hastings, 1943
Camptoplites asymmetricus Hastings, 1943: 466, figs 53D, 54C, D, 55C-F.
Geographic distribution: New Zealand, Chile, South Georgia, Patagonian shelf, Argentine Basin (Hastings, 1943; López-Gappa, 2000; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 1097 m.
References for the area: Hastings (1943); López-Gappa (2000).
Remarks: Colonies collected by the HMS Challenger at stn. 320 (holotype and paratypes). Hastings (1943: 466) stated that a part of the material assigned by Busk (1881a, 1884) to Bugula reticulata Busk, 1884 actually belongs to C. asymmetricus.
Camptoplites unicornis (Busk, 1884)
Bugula reticulata var. unicornis Busk, 1884: 40, pl. 9, fig. 2.
For synonymy see: d’Hondt (1981); Gordon (1986).
Geographic distribution: Pacific Ocean (Gordon, 1986); Atlantic Ocean: European continental slope (Hayward, 1978), Uruguay- Argentina, Argentine Basin (Busk, 1884).
Bathymetric distribution in Uruguayan waters: 1097-3917 m.
References for the area: Busk (1884); d’Hondt (1981).
Remarks: Collected at HMS Challenger, stn. 320 and RV Atlantis II, cruise 60, stns. 256 and 264 (d’Hondt, 1981).
Camptoplites lutaudae d’Hondt, 1975
Camptoplites lutaudae d’Hondt, 1975: 572.
For synonymy see: d’Hondt (1981).
Geographic distribution: North and South Atlantic, off Uruguay (d’Hondt, 1981).
Bathymetric distribution in Uruguayan waters: 3906-3917 m.115
References for the area: d’Hondt (1981).
Remarks: Collected by RV Atlantis II, cruise 60, stn. 256, off Uruguay.
Camptoplites bicornis (Busk, 1884)
Bugula bicornis Busk, 1884: 40, pl 9, fig. 1.
For synonymy see: d’Hondt (1981).
Geographic distribution: New Zealand, Uruguay, Argentina, and Antarctica (Busk, 1884; Moyano, 1974; d’Hondt, 1981; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 3305-3822 m.
References for the area: d’Hondt (1981, 1983); López-Gappa (2000).
Remarks: Colonies collected by RV Atlantis II, cruise 60, stns. 243 and 259 (d’Hondt, 1981, 1983).
Genus Kinetoskias Danielssen, 1868
Kinetoskias smitti Danielssen, 1868
Kinetoskias smitti Danielssen, 1868: 24.
Geographic distribution: Arctic Ocean (Kluge, 1975); North Atlantic (Ryland & Hayward, 1991); Argentine Basin, Uruguay (d’Hondt, 1981, 1983; López-Gappa, 2000).
Bathymetric distribution in Uruguayan waters: 2195-3343 m.
References for the area: d’Hondt (1981, 1983).
Remarks: Collected by the RV Atlantis II, cruise 60, stns. 240, 245, and 246 (d’Hondt, 1981, 1983; López-Gappa, 2000).
Genus Himantozoum Harmer, 1923
Himantozoum (Himantozoum) margaritiferum (Busk, 1884)
Bugula margaritifera Busk, 1884: 41, pl. 8, fig. 4.
Geographic distribution: Indian, North and South Atlantic oceans; Argentine Basin (Busk, 1884; d’Hondt & Schopf, 1984; d’Hondt, 1985b; David & Pouyet, 1986; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 3475 m.
References for the area: Busk (1884); López- Gappa (2000).
Remarks: collected by HMS Challenger at stn. 323 (Busk, 1884).
Himantozoum (Himantozoum) variabile (Kluge, 1914)
Dendrobeania variabilis Kluge, 1914: 632, pl. 28, fig. 6, text-fig. 19b.
Geographic distribution: North Atlantic (d’Hondt, 1983; d’Hondt & Schopf, 1984); Southern Ocean (Kluge, 1914), Argentine Basin (Uruguay) (d’Hondt, 1983; López-Gappa, 2000).
Bathymetric distribution in Uruguayan waters: 3343-3822 m.
References for the area: d’Hondt (1983).
Remarks: Colonies collected by RV Atlantis II, cruise 60, at stns. 243 and 246 (d’Hondt, 1983).
Family CANDIDAE d’Orbigny, 1851
Genus Amastigia Busk, 1852b
Amastigia crassimarginata (Busk, 1884)
Caberea crassimarginata Busk, 1884: 28, pl. 11, fig. 1.
Geographic distribution: Argentine Basin (Busk, 1884; Hastings, 1943; d’Hondt, 1981; López-Gappa, 2000; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 1097-2707 m.
References for the area: Busk (1884); d’Hondt (1981); López-Gappa (2000).
Remarks: Collected by the HMS Challenger at stn. 320, which is its type-locality (Busk, 1884). Referred with doubts by d’Hondt (1981), due to poor state of his material, for the RV Atlantis II, cruise 60, at stn. 245.
Genus Notoplites Harmer, 1923
Notoplites atlanticus d’Hondt, 1981
Notoplites atlanticus d’Hondt, 1981: 25, pl. II, figs 5-6.
Geographic distribution: Argentine Basin off Uruguay (d’Hondt, 1981).
Bathymetric distribution in Uruguayan waters: 3906-3917 m.
References for the area: d’Hondt (1981).
Remarks: Colonies collected by the RV Atlantis II expedition, cruise 60, stn. 256 (d’Hondt, 1981).
Notoplites crateriformis (Busk, 1884)
Cellularia crateriformis Busk, 1884: 16, pl. 3, fig. 1.
Geographic distribution: South Indian Ocean; Argentine Basin (Busk, 1884; d’Hondt, 1984; López-Gappa, 2000; Figuerola et al., 2014).116
Bathymetric distribution in Uruguayan waters: 3475 m.
References for the area: Busk (1884); López- Gappa (2000).
Remarks: Collected by the HMS Challenger at stn. 323 (Busk, 1884).
Notoplites paradoxus d’Hondt, 1981
Notoplites paradoxus d’Hondt, 1981: 25, pl. 4, figs 5-6.
Geographic distribution: Argentine Basin off Uruguay (d’Hondt, 1981).
Bathymetric distribution in Uruguayan waters: 2707 m.
References for the area: d’Hondt (1981).
Remarks: Lower bathyal species, collected during the RV Atlantis II cruise 60 at stn. 245, off Uruguay, which is its type-locality (d’Hondt, 1981).
Notoplites cymbalicus d’Hondt, 1981
Notoplites cymbalicus d’Hondt, 1981: 26, pl. 4, figs 3-4.
Geographic distribution: Argentine Basin off Uruguay (d’Hondt, 1981, 1983).
Bathymetric distribution in Uruguayan waters: 3815-4402 m.
References for the area: d’Hondt (1981, 1983).
Remarks: Colonies collected during the RV Atlantis II, cruise 60 at stns. 242 and 243 off Uruguay (d’Hondt, 1981, 1983).
Superfamily CELLARIOIDEA Fleming, 1828
Family MEMBRANICELLARIIDAE Levinsen, 1909
Genus Membranicellaria Levinsen, 1909
Membranicellaria dubia (Busk, 1884)
Melicerita dubia Busk, 1884: 97, pl. 23, fig. 10.
Geographic distribution: Argentine basin (Busk, 1884; López-Gappa, 2000; Figuerola et al., 2014) and Antarctica (Ross Sea) (Hayward & Winston, 2011).
Bathymetric distribution in Uruguayan waters: 1097 m.
References for the area: Busk (1884); López- Gappa (2000).
Remarks: Collected during the HMS Challenger expedition at stn. 320, which is the type-locality of this species (Busk, 1884).
Genus Cookinella d’Hondt, 1981
Cookinella flustroides d’Hondt, 1981
Cookinella flustroides d’Hondt, 1981: 14, pl. 3, fig. 1; text fig. 3.
Geographic distribution: Argentine Basin (d’Hondt, 1981, 1983; López-Gappa, 2000; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 4382-4402 m.
References for the area: d’Hondt (1981).
Remarks: Abyssal species collected by the RV Atlantis II, cruise 60, at stn. 242, which is its type locality (d’Hondt, 1981). It was also re corded at greater depths (5208-5223 m) at stn. 247 during the same cruise, i.e. off Patagonia (d’Hondt, 1983).
Family CELLARIIDAE Fleming, 1828
Genus Cellaria Ellis & Solander, 1786
Cellaria dubia (Busk, 1884)
Salicornaria dubia Busk, 1884: 91, pl. 12, fig. 2, text-fig. 10.
Geographic distribution: Argentine basin (Busk, 1884; López-Gappa, 2000).
Bathymetric distribution in Uruguayan waters: 1097 m.
References for the area: Busk (1884), López- Gappa (2000).
Remarks: Collected during the HMS Challenger expedition at stn. 320. Colonies of another species were collected at 13-40 m depth near Uruguay (Parcel do Carpinteiro, off Rio Grande and São José cities and Santa Vitória do Palmar city - Rio Grande do Sul State, Brazil) and identified as Cellaria riograndensis Ramalho & Calliari, 2015.
Genus Euginoma Jullien, 1882
Euginoma biseriata d’Hondt, 1981
Euginoma biseriata d’Hondt, 1981: 16, pl. 1, figs 5-6.
Geographic distribution: North Atlantic; South West Atlantic (Brazilian and Argentine basins) (d’Hondt, 1981, 1983; d’Hondt & Schopf, 1984; López-Gappa, 2000; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 2707 m.
References for the area: d’Hondt (1981); 117
López-Gappa (2000).
Remarks: Abyssal species. Colonies collected by the RV Atlantis II cruise 60 at stn. 245, which is its type-locality (d’Hondt, 1981).
Euginoma crispa d’Hondt, 1981
Euginoma crispa d’Hondt, 1981: 17, pl. 1, figs 1-2.
Geographic distribution: Argentine Basin (off Uruguay) (d’Hondt, 1981).
Bathymetric distribution in Uruguayan waters: 2707 m.
References for the area: d’Hondt (1981).
Remarks: Collected during the RV Atlantis II cruise 60 at stn. 245, which is its type-locality (d’Hondt, 1981).
Euginoma cylindrica d’Hondt, 1981
Euginoma cylindrica d’Hondt, 1981: 18, fig. 4.
Geographic distribution: South Atlantic, Argentine Basin: Uruguay (d’Hondt, 1981).
Bathymetric distribution in Uruguayan waters: 2707 m.
References for the area: d’Hondt (1981).
Remarks: Collected during the RV Atlantis II cruise 60 at stn. 245, which is its type-locality.
Euginoma cavalieri Lagaaij, 1963
Euginoma cavalieri Lagaaij, 1963: 179, pl. 2, fig. 6.
For synonymy see: d’Hondt (1981).
Geographic distribution: North Atlantic; Brazil (off Pernambuco) and Argentine Basin (off Uruguay) (Lagaaij, 1963; d’Hondt, 1981, 1983; d’Hondt & Schopf, 1984; López-Gappa, 2000; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 2041-2048 m.
References for the area: d’Hondt (1981), López-Gappa (2000).
Remarks: Species from deep water (365-2480 m), originally described from the Gulf of Mexico (Lagaaij, 1963).
Genus Formosocellaria d’Hondt, 1981
Formosocellaria magnifica (Busk, 1884)
Salicornaria magnifica Busk, 1884: 93, pl. 12, figs 4 and 6.
Formosocellaria abyssicola d’Hondt, 1981: 20, pl. 2, figs 3-4; pl. 3, fig. 2; text-fig. 5.
Geographic distribution: North Atlantic; South Indian Ocean; South Western Pacific; Argentine Basin (off Uruguay and Argentina); Brazil Basin (off Pernambuco and Alagoas) (Busk, 1884; d’Hondt, 1981, 1983, 1984; d’Hondt & Schopf, 1984; López-Gappa, 2000; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 3305-3917 m.
References for the area: Busk (1884); López- Gappa (2000).
Remarks: Colonies collected by the HMS Challenger at stn. 323, and by the RV Atlantis II cruise 60, at stns. 243, 246, 256 (type locality of F. abyssicola) and 259. d’Hondt & Schopf (1984) synonymized F. abyssicola with F. magnifica.
Genus Cryptostomaria Canu & Bassler, 1927
Cryptostomaria cylindrica (Harmer, 1926)
Aspidostoma cylindricum Harmer, 1926: 323, pl. 22, figs 1-4.
Geographic distribution: Indonesia (Harmer, 1926), Brazil and Argentine basins (off Uruguay) (d’Hondt, 1983; d’Hondt & Schopf, 1984; López- Gappa, 2000).
Bathymetric distribution in Uruguayan waters: 3815-3822 m.
References for the area: d’Hondt (1983); López-Gappa (2000).
Remarks: Colonies collected by the RV Atlantis II cruise 60 at stn. 243 off Uruguay.
Genus Melicerita Milne-Edwards, 1836
Melicerita atlantica Busk, 1884
Melicerita atlantica Busk, 1884: 96, pl. 14, fig. 1.
Geographic distribution: Argentine Basin, Argentina-Uruguay (Busk, 1884; López-Gappa, 2000; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 1097 m.
References for the area: Busk (1884); López- Gappa (2000).
Remarks: Collected during the HMS Challenger expedition at stn. 320, which is the type and only known locality for this species.
Superfamily CATENICELLOIDEA Busk, 1852b
Family CATENICELLIDAE Busk, 1852b118
Genus Talivittaticella Gordon & d’Hondt, 1985
Talivittaticella problematica (d’Hondt, 1981)
Orthoscuticella (?) problematicum d’Hondt, 1981: 42, pl. VII, fig. 5; text fig. 11.
Geographic distribution: New Zealand; Brazilian (off Pernambuco) and Argentine ba sins (Uruguay) (d’Hondt, 1981, 1983; d’Hondt & Schopf, 1984; Gordon & d’Hondt, 1985; López- Gappa, 2000; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 2707-3822 m.
References for the area: d’Hondt (1981, 1983); López-Gappa (2000).
Remarks: Collected by the RV Atlantis II, cruise 60, at stns. 243 and 245, the latter being the type locality of this species (d’Hondt, 1981, 1983).
Superfamily CRIBRILINOIDEA Hincks, 1879
Family CRIBRILINIDAE Hincks, 1879
Genus Jolietina Jullien, 1886
Jolietina latimarginata (Busk, 1884)
Cribrilina latimarginata Busk, 1884: 131, pl. 22, fig. 10.
For synonymy see: López-Gappa et al., 2021: 3, figs 2.1, 2.2.
Geographic distribution: Southeast Pacific (Chile); Southwest Atlantic (Argentine Basin and Patagonian shelf, including Malvinas Islands) (Busk, 1884; Moyano, 1984; López-Gappa & Lichtschein, 1988; López-Gappa, 2000; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 1097 m.
References for the area: Busk (1884); López- Gappa (2000).
Remarks: Collected during the HMS Challenger expedition at stn. 320, which is its type-locality (Busk, 1884). At the Patagonian shelf these col onies are usually found on the cyclostome bryo zoan Hornera (López-Gappa et al., 2021). At the type-locality it was reported as growing on dead coral (Busk, 1884).
Superfamily BIFAXARIOIDEA Busk, 1884
Family BIFAXARIIDAE Busk, 1884
Genus Domosclerus Gordon, 1988
Domosclerus corrugatus (Busk, 1884)
Bifaxaria corrugata Busk, 1884: 80, pl. 13, fig. 3; pl. 24, fig. 6.
Sclerodomus sp. aff. D. corrugatus d’Hondt, 1981: 46.
Geographic distribution: D. corrugatus has been recorded for the Indo-Pacific, New Zealand, South Africa, Brazil (off Pernambuco), Uruguay and Argentina (Busk, 1884; Harmer, 1957; d’Hondt, 1981, 1983; López-Gappa, 2000; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 3815-4402 m.
References for the area: d’Hondt (1981, 1983); López-Gappa (2000).
Remarks: Four fragments of Sclerodomus sp. and colonies of Domosclerus corrugatus were collected during the RV Atlantis II expedition, at stns. 242 and 243, respectively, off Uruguay. Sclerodomus sp. specimens do not have oral avic ularia, a fact that did not allow to confirm its identification as D. corrugatus.
Superfamily HIPPOTHOOIDEA Busk, 1859
Family HIPPOTHOIDAE Busk, 1859
Genus Celleporella Gray, 1848
Celleporella hyalina (Linnaeus, 1767)
Cellepora hyalina Linnaeus, 1767: 1286.
For synonymy see: Soule et al., 1995: 183, fig. 66A-D.
Geographic distribution: this species com plex (Waeschenbach et al., 2012) is distributed worldwide, including Brazil, Argentina, and Uruguay (Marcus, 1937, 1955; López-Gappa & Lichtschein, 1988; Obenat et al., 2001; Scarabino, 2006; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 11-12 m.
References for the area: Obenat et al. (2001); Scarabino (2006).
Remarks: At Uruguay the colonies were fixed on tubes of the polychaete Phyllochaetopterus so cialis (Obenat et al., 2001).
Superfamily LEPRALIELLOIDEA Vigneaux, 1949
Family LEPRALIELLIDAE Vigneaux, 1949
Genus Celleporaria Lamouroux, 1821
Celleporaria (?) sp.
Celleporaria sp. d’Hondt, 1981: 40.
Geographic distribution: Argentine Basin: Uruguay (d’Hondt, 1981).
Bathymetric distribution in Uruguayan waters: 1661-1679 m.119
References for the area: d’Hondt (1981).
Remarks: Colonies were collected during the RV Atlantis II cruise 60 at stn. 239. These spec imens lacked ovicell, an important character to identify the colonies at the species level.
Family SCLERODOMIDAE Levinsen, 1909
Genus Sclerodomus Levinsen, 1909
Sclerodomus denticulatus (Busk, 1884)
Bifaxaria denticulata Busk, 1884: 82, pl. 24, fig. 3.
Geographic distribution: Magellanic region and Argentine Basin (Busk, 1884; López-Gappa, 2000; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 1097 m.
References for the area: Busk (1884); López- Gappa (2000).
Remarks: Colonies were collected during the HMS Challenger expedition at stn. 320, which is its type-locality.
Superfamily SMITTINOIDEA Levinsen, 1909
Family SMITTINIDAE Levinsen, 1909
Genus Smittina Norman, 1903
Smittina uruguayensis d’Hondt, 1981
Smittina uruguayensis d’Hondt, 1981: 37, text fig. 8.
Geographic distribution: only known from the lower slope of the Argentine Basin (Uruguay) (d’Hondt, 1981; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 1661-1679 m.
References for the area: d’Hondt (1981); López-Gappa (2000).
Remarks: Colonies were collected during the RV Atlantis II, cruise 60, at stn. 239, off Uruguay, which is its type and sole locality at which it has been recorded (d’Hondt, 1981).
Smittina smittiana (Busk, 1884)
Smittia smittiana Busk, 1884: 151, pl. 17, fig. 3.
Geographic distribution: Magellanic region and Argentine Basin (Busk, 1884; López-Gappa, 2000; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 1097 m.
References for the area: Busk (1884); López- Gappa (2000).
Remarks: Colonies were collected during the HMS Challenger expedition at stn. 320, which is its type-locality (Busk, 1884).
Smittina sp. (aff. S. exclusa Harmer, 1957)
Smittina sp. d’Hondt, 1981: 39, text fig. 10.
Geographic distribution: Uruguay (d’Hondt, 1981).
Bathymetric distribution in Uruguayan waters: 2707 m.
References for the area: d’Hondt (1981).
Remarks: Colonies white, erect, dichotomous ly branched; collected at stn. 245 during the RV Atlantis II cruise 60.
Genus Pseudoflustra Bidenkap, 1897
Pseudoflustra sp. (aff. P. aviculata (Calvet, 1906))
Pseudoflustra sp.: d’Hondt, 1981: 44.
Geographic distribution: Argentine Basin: Uruguay (d’Hondt, 1981).
Bathymetric distribution in Uruguayan waters: 2707 m.
References for the area: d’Hondt (1981).
Remarks: Two tiny fragments were collected during RV Atlantis II cruise 60 at stn. 245.
Family BITECTIPORIDAE MacGillivray, 1895
Genus Hippomonavella Canu & Bassler in Bassler, 1934
Hippomonavella charrua López-Gappa et al., 2020
Hippomonavella charrua López-Gappa et al., 2020: 145, fig. 2; table 1.
Geographic distribution: Continental shelf off Uruguay and off Buenos Aires Province, and Canal de las Escobas Formation (mid-Holocene deposits) - Argentina (López-Gappa et al., 2020).
Bathymetric distribution in Uruguayan waters: 119-128 m.
References for the area: López-Gappa et al. (2020).
Remarks: Bilaminar colonies on hydrozoan stems were collected by the vessel Undine off the Uruguayan coast, at 34°38’ S, 52°15’ W, 119-128 m depth, which is the type-locality of this species. Another species, H. brasiliensis Ramalho et al., 2008 was identified near Uruguay: Rio Grande do Sul state, Brazil (Parcel do Carpinteiro, off Rio Grande and São José cities). The loose bil aminar fragments of the Brazilian species were collected at 24-25 m depth (Ramalho & Calliari, 2015).
Superfamily SCHIZOPORELLOIDEA Jullien, 1883
Family MARGARETTIDAEHarmer, 1957
Species incertae sedis aff. Margaretta Gray, 1843
Species incertae sedis aff. g. Margaretta d’Hondt, 1983: 87, pl. 4, figs 5-6; text fig. 2.
Geographic distribution: Argentine Basin (d’Hondt, 1983).
Bathymetric distribution in Uruguayan waters: 2707 m.
References for the area: d’Hondt (1983).
Remarks: Two small fragments were collected during the RV Atlantis II cruise 60 at stn. 245 (d’Hondt, 1983).
Family CRYPTOSULIDAE Vigneaux, 1949
Genus Cryptosula Canu & Bassler, 1925
Cryptosula pallasiana (Moll, 1803)
Eschara pallasiana Moll, 1803: 64, pl. 3, fig. 13.
For synonymy see: Hayward & Ryland, 1999.
Geographic distribution: This species has a worldwide distribution, including Argentina, Uruguay and Brazil (Marcus, 1942; Lichtschein de Bastida & Bastida, 1980; Hayward & Ryland, 1999; López-Gappa, 2000; Scarabino, 2006; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: Shallow subtidal.
References for the area: Calvo (1984) as Cryptosula sp.; Scarabino (2006).
Remarks: This species is a frequent member of the fouling community (Orensanz et al., 2002). It is typically and commonly found in intertidal habitats, but it can also be found in the sublit toral, growing on stones, shells, and other hard or soft substrates such as polychaete tubes, al gae and hydroids. It can be found from the up per littoral down to 200 m depth (Marcus, 1942), and it is considered as introduced in Uruguay- Argentina (Schwindt et al., 2020).
Family CALWELLIIDAE MacGillivray, 1887
Genus Ichthyaria Busk, 1884
Ichthyaria profunda d’Hondt, 1981
Ichthyaria profunda d’Hondt, 1981: 49, fig. 13.
Geographic distribution: Argentine Basin off Uruguay (d’Hondt, 1981).
Bathymetric distribution in Uruguayan waters: 2707 m.
References for the area: d’Hondt (1981); López-Gappa (2000).
Remarks: Flexible and delicate colonies of this species were collected off Uruguay by the RV Atlantis II cruise 60 at stn. 245 (type-locality).
Ichthyaria oculata Busk, 1884
Ichthyaria oculata Busk, 1884: 46, pl. 13, fig. 7.
Geographic distribution: South Indian Ocean (Crozet shelf and Prince Edward Islands); Magellanic region, Patagonian shelf and Argentine Basin (Busk, 1884; d’Hondt, 1984; López-Gappa & Lichtschein, 1988; López-Gappa, 2000; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 1097 m.
References for the area: Busk (1884); López- Gappa (2000).
Remarks: Colonies were collected during the HMS Challenger expedition at stn. 320 (Busk, 1884).
Family MYRIAPORIDAE Gray, 1841
Genus Myriapora de Blainville, 1830
Myriapora simplex (Busk, 1884)
Myriozoum simplex Busk, 1884: 170, pl. 25, fig. 1.
Geographic distribution: Argentine Basin, Argentina-Uruguay (Busk, 1884; López-Gappa, 2000).
Bathymetric distribution in Uruguayan waters: 1097 m.
References for the area: Busk (1884); López- Gappa (2000).
Remarks: Collected during the HMS Challenger expedition at stn. 320, which is its type and unique known locality for this species.
Superfamily CELLEPOROIDEA Johnston, 1838
Family PHIDOLOPORIDAE Gabb & Horn, 1862
Genus Reteporellina Harmer, 1933
Reteporellina moyanoi d’Hondt, 1981
Reteporellina moyanoi d’Hondt, 1981: 45, fig. 12.
Geographic distribution: Only known from the lower slope of Argentine Basin (Uruguay) (d’Hondt, 1981; López-Gappa, 2000; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 2707 m.
References for the area: d’Hondt (1981); López-Gappa (2000).
Remarks: Colonies erect, dichotomously branched, collected by the RV Atlantis II cruise 60 at stn. 245, which is its type-locality and the unique site in which this species has been record ed (d’Hondt, 1981).
Reteporellina sp. 2 d’Hondt, 1981
Reteporellina sp. 2 d’Hondt, 1981: 46.
Geographic distribution: Only known from the lower slope of the Argentine Basin (Uruguay) (d’Hondt, 1981).
Bathymetric distribution in Uruguayan waters: 4382-4402 m.
References for the area: d’Hondt (1981).
Remarks: Colonies erect, dichotomously branched, collected by the RV Atlantis II expedi tion, at stn. 242 (d’Hondt, 1981).
Genus Reteporella Busk, 1884
Reteporella magellensis (Busk, 1884)
Retepora magellensis Busk, 1884: 126, pl. 26, fig. 20.
Reteporella magellensis: Hayward & Winston, 2011: 2320, fig. 29A.
For synonymy see: López-Gappa & Lichtschein (1990).
Geographic distribution: Southern Chile, Magellanic region, Patagonian shelf, Argentine Basin (Busk, 1884; Waters, 1905; López-Gappa & Lichtschein, 1990; Hayward & Winston, 2011; Figuerola et al., 2014) and Antarctic Peninsula (Hayward & Winston, 2011).
Bathymetric distribution in Uruguayan waters: 1097 m.
References for the area: Busk (1884).
Remarks: Colonies infundibuliform or cupped, rising from a short and thick peduncle. They were collected during the HMS Challenger ex pedition at stn. 320, which is its type-locality (Busk, 1884; Hayward & Winston, 2011).
Family CELLEPORIDAE Johnston, 1838
Genus Buskea Heller, 1867
Buskea sp.
Buskea sp. d’Hondt, 1983: 89, pl. 2, fig. 6.
Geographic distribution: Only known from the lower slope of the Argentine Basin (Uruguay) (d’Hondt, 1983).
Bathymetric distribution in Uruguayan waters: 2195-2323 m.
References for the area: d’Hondt (1983).
Remarks: Erect single fragment collected by the RV Atlantis II cruise 60, at stn. 240 (d’Hondt, 1983).
Genus Orthroporidroides Moyano, 1974
Orthroporidroides erectus (Waters, 1888)
Cellepora armata var. erecta Waters, 1888: 36, pl. 3, figs 4, 41, 43
For synonymy see: Cook & Hayward (1983)
Geographic distribution: Uruguay (Cook & Hayward, 1983); Magellan Strait up to Madre de Dios Archipelago (Chile - Pacific Ocean) (Moyano, 1974).
Bathymetric distribution in Uruguayan waters: 1097 m.
References for the area: Cook & Hayward (1983).
Remarks: Colony erect, branching, arising from encrusting bases. Colonies were collected during the HMS Challenger expedition at stn. 320, which is the type-locality (Cook & Hayward, 1983).
Genus Osthimosia Jullien, 1888
Osthimosia rudis (Busk, 1881b)
Cellepora rudis Busk, 1884: 199, pl. 28, fig. 7, pl. 36, fig. 7.
Geographic distribution: Argentine Basin, Argentina-Uruguay (Busk, 1881b, 1884; Hayward, 1992; López-Gappa, 2000).
Bathymetric distribution in Uruguayan waters: 1097 m.
References for the area: Busk (1881b, 1884); Hayward (1992); López-Gappa (2000).
Remarks: Colony with a short, cylindrical, thick stem rising from a broad base, and divid ing into two rounded lobes, collected during the HMS Challenger expedition at stn. 320, which is the type and only known locality (Busk, 1884; Hayward, 1992).
Superfamily CONESCHARELLINOIDEA Levinsen, 1909
Family ORBITULIPORIDAE Canu & Bassler, 1923
Genus Sphaerulobryozoon d’Hondt, 1981
Sphaerulobryozoon pedunculatum d’Hondt, 1981
Sphaerulobryozoon pedunculatum d’Hondt, 1981: 47, pl. VI, figs 5-6, pl. VII, fig. 1-4, pl. VIII.
Geographic distribution: North Atlantic (d’Hondt, 1981), Brazil (off Pernambuco), and Argentine Basin (d’Hondt, 1981, 1983; d’Hondt & Schopf, 1984; López-Gappa, 2000; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 2041-2480 m.
References for the area: d’Hondt (1981, 1983); López-Gappa (2000).
Remarks: Conescharelliniform colonies collect ed at lower bathyal depths by the RV Atlantis II cruise 60, at stns. 240, 262 and 264. Station 262 is the type-locality of this species (d’Hondt, 1981, 1983).
Family LEKYTHOPORIDAE Levinsen, 1909
Genus Turritigera Busk, 1884
Turritigera stellata Busk, 1884
Turritigera stellata Busk, 1884: 130, pl. 24, fig. 1.
For synonymy see: Cook & Hayward (1983); Hayward & Winston (2011).
Geographic distribution: Southeast Pacific (Chile), Cape Horn, Magellanic region, Argentine Basin (Busk, 1884; Moyano, 1974; Cook & Hayward, 1983; López-Gappa & Lichtschein, 1988; López-Gappa, 2000; Ramalho et al., 2011; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 1097 m. Turritigera spectabilis was col lected at 1661−4402 m (see Remarks below).
References for the area: Busk (1884); López- Gappa (2000).
Remarks: Erect and well calcified colonies were collected by the HMS Challenger expedition at stn. 320 (Busk, 1884). The original description was also based on material from South Africa. A colony from stn. 320 was selected by Hayward & Winston (2011) as the lectotype, that stn. thus becoming its type locality.
Turritigera spectabilis d’Hondt, 1981 was de scribed from material obtained at similar depths at very close (stns. 239, 242 and 245 of the RV Atlantis II cruise 60) to the type locality of T. stellata. As a formal revision of type materials is needed to conclude that T. spectabilis is a junior synonym of T. stellata, in this compilation we have preferred to cite T. spectabilis as a separate species.
Turritigera spectabilis d’Hondt, 1981
Turritigera spectabilis d’Hondt, 1981: 41, pl. 5, figs 1-3.
Geographic distribution: Argentine Basin off Uruguay (d’Hondt, 1981).
Bathymetric distribution in Uruguayan waters: 1661-4402 m.
References for the area: d’Hondt (1981).
Remarks: bathyal-abyssal specimens collected by the RV Atlantis II, cruise 60, at stns. 239, 242, and 245, the former being the type locality of this species (d’Hondt, 1981).
Genus Catadysis Canu and Bassler, 1927
Catadysis immersum (Busk, 1884)
Myriozoum immersum Busk, 1884: 170, pl. 25, fig. 4.
Geographic distribution: Magellanic re gion and Argentine basin (Busk, 1884; Cook & Hayward, 1983; López-Gappa, 2000; Figuerola et al., 2014).
Bathymetric distribution in Uruguayan waters: 1097 m.
References for the area: Busk (1884), Cook & Hayward (1983); López-Gappa (2000).
Remarks: Erect colonies with divergent and bifurcating branches were collected during the HMS Challenger expedition at stn. 320.
CHEILOSTOMATIDA INCERTAE SEDIS
Genus Acanthodesiomorpha d’Hondt, 1981
Acanthodesiomorpha problematica d’Hondt, 1981
Acanthodesiomorpha problematica d’Hondt, 1981: 9, pl. 4, figs 1-2, text fig. 1.
Geographic distribution: Argentine Basin (d’Hondt, 1981), New Zealand (Gordon, 1987).
Bathymetric distribution in Uruguayan waters: 2707 m.
References for the area: d’Hondt (1981).
Remarks: Collected by the RV Atlantis II cruise 60 at stn. 245 (d’Hondt, 1981). Gordon (1987) recorded this species for New Zealand, at 3999 m depth.
Genus Ogivalia Jullien, 1882
Ogivalia elegans (d’Orbigny, 1842)
Vincularia elegans d’Orbigny, 1842: 21, pl. 9, figs 25-28.
Vincularia labiata Busk, 1884: 73, text fig. 3.
For synonymy see: Moyano (1974).
Geographic distribution: Magellanic region and Argentine Basin (d’Orbigny, 1847; Moyano, 1974; López-Gappa & Lichtschein, 1988; López- Gappa, 2000).
Bathymetric distribution in Uruguayan waters: 1097 m.
References for the area: Busk (1884) as Vincularia labiata; Moyano (1974); López-Gappa (2000).
Remarks: Colonies were collected during the HMS Challenger expedition at stn. 320, de scribed as Vincularia labiata. Waters (1905) synonymized Vincularia labiata with Vincularia elegans.
Order CTENOSTOMATIDA (Busk, 1852a)
Suborder ALCYONIDIINA d’Hondt, 1985a
Superfamily ALCYONIDIOIDEA Johnston, 1838
Family ALCYONIDIIDAE Johnston, 1837
Genus Alcyonidium Lamouroux, 1813
Alcyonidium sp.
Geographic distribution: Uruguay: 35°23’ - 35°31’S, 55°36’ - 55°48’W.
Bathymetric distribution in Uruguayan waters: 11-12 m.
References for the area: Obenat et al. (2001); Scarabino (2006).
Remarks: Colonies were found on polychaete tubes (Phyllochaetopterus socialis) near the mouth of Río de la Plata estuary (Obenat et al., 2001).
Superfamily HAYWARDOZOONOIDEA d’Hondt, 1983
Family HAYWARDOZOIDAE d’Hondt, 1983
Genus Haywardozoon d’Hondt, 1983
Haywardozoon atlantae d’Hondt in d’Hondt & Hayward, 1981
Haywardozoon atlantae d’Hondt & Hayward, 1981: 279, fig. 4.
Geographic distribution: Argentine Basin off Uruguay (d’Hondt & Hayward, 1981).
Bathymetric distribution in Uruguayan waters: 2440-2480 m.
References for the area: d’Hondt and Hayward (1981); López-Gappa (2000).
Remarks: Lower bathyal uniserial colonies collected during the RV Atlantis II cruise 60 at stn. 262, which is its type-locality (d’Hondt and Hayward, 1981).
Suborder VESICULARIINA Johnston, 1838
Superfamily VESICULARIOIDEA Hincks, 1880
Family VESICULARIIDAE Hincks, 1880
Genus Amathia Lamouroux, 1812
Amathia sp.
Geographic distribution: Uruguay (Calvo, 1984; Scarabino, 2006).
Bathymetric distribution in Uruguayan waters: Shallow subtidal.
References for the area: Calvo (1984).
Remarks: Small and transparent colonies were found on wood (experimental panels) in Uruguayan ports (Calvo, 1984). This species was identified as Bowerbankia sp. by Calvo (1984), but this genus was later regarded as a junior synonym of Amathia (see Waeschenbach et al., 2015).
Class STENOLAEMATA
Order CYCLOSTOMATIDA Busk, 1852a
Suborder ARTICULINA Busk, 1859
Family CRISIIDAE Johnston, 1838
Genus Crisia Lamouroux, 1812
Crisia acuminata Busk, 1886
Crisia acuminata Busk, 1886: 5, pl. 3, fig. 1.
Geographic distribution: Argentine Basin, Argentina-Uruguay (Busk, 1886; López-Gappa, 2000).
Bathymetric distribution in Uruguayan waters: 1097 m.
References for the area: Busk (1886); López- Gappa (2000).
Remarks: Erect and delicate white colonies were collected during the HMS Challenger ex pedition at stn. 320, the type and only locality known for this species.
Suborder TUBULIPORINA Milne-Edwards, 1838
Remarks: An Oncousoeciidae, Filisparsa calyp so Buge, 1979 has its type and only locality close to the area here considered, i.e. 37°36´S, 54°56´W, 740 m depth (stn. 171 of the RV Calypso, cruise 1961-1962 to South America) (Buge, 1979). Moreover, Buge (1979) stated that the record of Idmonea marionensis Busk, 1875 (a species cur rently considered a taxon inquirendum) collect ed at stn. 320 of the HMS Challenger expedition may be the same as F. calypso. However, no fur ther research has been done on this subject.
Family STOMATOPORIDAE Pergens & Meunier, 1886
Genus Stomatopora Bronn, 1825
Remarks: Stomatopora eburnea (d’Orbig ny, 1847), known from the Magellanic region and Argentine Basin (d’Orbigny, 1847; Buge, 1979; López-Gappa, 2000), has been cited from a locality close to the area considered here, i.e. 37°36´S, 54°56´W, 740 m depth (stn. 171 of the RV Calypso, cruise 1961-1962 to South America) (Buge, 1979).
Stomatopora sp.
Stomatopora sp. Harmelin & d’Hondt, 1982: 6.
Geographic distribution: Argentine Basin off Uruguay (Harmelin & d’Hondt, 1982).
Bathymetric distribution in Uruguayan waters: 2707 m.
References for the area: Harmelin & d’Hondt (1982).
Remarks: Two very damaged fragments fixed on rocks were collected during the RV Atlantis II cruise at stn. 245 off Uruguay.
Family DIASTOPORIDAE, Gregory, 1899
Diastoporidae n.d., Harmelin & d’Hondt, 1982: 10, pl. 3, fig. 5.
Geographic distribution: Argentine Basin off Uruguay (Harmelin & d’Hondt, 1982).
Bathymetric distribution in Uruguayan waters: 2707 m.
References for the area: Harmelin & d’Hondt (1982).
Remarks: Only one specimen was collected by the RV Atlantis II expedition, cruise 60, stn. 245 (Harmelin & d’Hondt, 1982).
Suborder CANCELLATA Gregory, 1896
Family HORNERIDAE Smitt, 1867
Genus Hornera Lamouroux, 1821
Hornera sp.
Hornera lichenoides: Busk, 1886: 15
Geographic distribution: Argentine basin, Argentina-Uruguay (Busk, 1884).
Bathymetric distribution in Uruguayan waters: 1097 m.
References for the area: Busk (1884), Buge (1979).
Remarks: Collected during the HMS Challenger expedition at stn. 320. According to Buge (1979), the specimen found in the Southwest Atlantic should not be regarded as conspecific with H. lichenoides (Linnaeus, 1758), a boreal-arctic spe cies.
Family PSEUDIDMONEIDAE Borg, 1944
Genus Pseudidmonea Borg, 1944
Pseudidmonea fissurata (Busk, 1886)
Idmonea fissurata Busk, 1886: 14, pl. 3, fig. 5.
For synonymy see: Buge (1979).
Geographic distribution: Argentine Basin, Magellanic region (Busk, 1886; Borg, 1944; Androsova, 1968; Buge, 1979).
Bathymetric distribution in Uruguayan waters: 1097 m.
References for the area: Busk (1886); Buge (1979); López-Gappa (2000).
Remarks: Some colonies and other fragments were collected during the HMS Challenger expe dition, at stn. 320. Buge (1979) cited this species from 37°36´S, 54°56´W, 740 m depth (stn. 171 of the RV Calypso, cruise 1961-1962 to South America), i.e. from a locality close to the area considered here.
Cyclostomatida incertae sedis n° 2
Incertae sedis n° 2 Harmelin & d’Hondt, 1982: 12: pl. IV, fig. 6.
Geographic distribution: Argentine Basin off Uruguay (Harmelin & d’Hondt, 1982).
Bathymetric distribution in Uruguayan waters: 2707 m.
References for the area: Harmelin & d’Hondt (1982).
Remarks: One colony of an inverted umbrella shape and a fixation peduncle was collected by the RV Atlantis II expedition, cruise 60, stn. 245 (Harmelin & d’Hondt, 1982).
Cyclostomatida incertae sedis n° 3
Incertae sedis n° 3 Harmelin & d’Hondt, 1982: 13, pl. IV, fig. 3.125
Geographic distribution: Argentine Basin off Uruguay (Harmelin & d’Hondt, 1982).
Bathymetric distribution in Uruguayan waters: 2707 m.
References for the area: Harmelin & d’Hondt (1982).
Remarks: Eleven small fragments of erect col onies were collected by the RV Atlantis II expe dition, cruise 60, stn. 245 (Harmelin & d’Hondt, 1982).
DISCUSSION AND CONCLUSION
This study compiled 73 taxa recorded in Uruguayan waters, from all three orders of extant marine bryozoans (Cheilostomatida, Ctenostomatida and Cyclostomatida). The most diverse order was Cheilostomatida with 63 taxa, followed by Cyclostomatida (7 taxa) and Ctenostomatida (3 taxa).
With an estimated diversity of more than 4,900 species in 2013 (Bock & Gordon, 2013), the Cheilostomatida is the most successful group among the Recent bryozoans. Its high species-di versity seems to be related to their variability in the use of substrates and the acquisition of spe cial features such as zooidal polymorphism and complex reproductive patterns (Ryland, 1970).
On the other hand, the Cyclostomatida usual ly show modest levels of biodiversity when com pared with the Cheilostomatida, accounting glob ally for up to 11% of the species in modern bryo zoan faunas (Rosso, 2003; Ramalho et al., 2009). Three out of the seven taxa of Cyclostomatida herein mentioned were found at between 200 and 1500 m depth, while another four between 1500 and 3000 m. Hayward & Ryland (1985) mentioned that Cyclostomatida species are usu ally found to occur in shallow waters less than 1000 m depth, but this statement is undergoing changes from the new studies that are being car ried out in deep waters. Grischenko et al. (2018) studying the bryozoan fauna associated with nodules, collected in the Russian deep sea (4510 and 5280 m), showed that cyclostomes are much more diverse than previously thought, compris ing 34% of the total bryozoan fauna identified.
Table 1 Summary of the species of Bryozoa recorded at Uruguayan waters. Taxa are arranged follow ing their taxonomic order in the text. The depths detailed only refers to the records from Uruguay.
The Ctenostomatida commonly exhibit low biodiversity levels, but usually are very common and abundant in shallow waters. Three ctenos tomes were recorded in the present study, two of them from depths up to 200 m and one at deeper waters.
Thirty out of the 73 taxa (41%) are known only from deep waters off the Río de la Plata, with an additional species also known from a sta tion in Patagonian deep waters. Even consider ing there are some undetermined species, this re sult shows a high degree of endemism, as it was already shown for several benthic groups such as Bivalvia, Ascidiacea and Pycnogonida (Scarabino et al., 2016, 2018, 2019, and references therein). Furthermore, also molecular analysis could iden tify even more endemic species amongst some of the more seemingly widely distributed species. The remaining recorded species could be classi fied in three groups: 1) deep-sea species widely distributed over other basins (15), 2) shelf and deep-sea species of Antarctic and Subantarctic distribution (17), and 3) shelf species, including shallow water cryptogenic or introduced taxa (10). A similar pattern exists concerning the pro portion of species for Ascidiacea and Pycnogonida from Uruguayan waters (Scarabino et al., 2018, 2019) and it represents a bias. In fact, most of the species compiled in this study were gath ered during two oceanographic cruises that sur veyed the continental slope and abyssal plain off Uruguay. This fact, together with the absence of local bryozoan taxonomists, has resulted in the unprecedented situation that the deep-sea bryozoan fauna of Uruguay is better known than the coastal and shelf representatives of the phylum. Therefore, future efforts should be di rected to coordinate the sampling and taxonomic study of shallow and shelf specimens in order to bridge the present gap in the knowledge of the Uruguayan bryozoan fauna. The present study is designed to kick-start more detailed empirical taxonomic studies to fill the gap in the knowl edge of bryozoan distribution patterns along the Atlantic coast of South America.
