Marine Ecology.

ISSN 0173-9565

ORIGINAL ARTICLE

Hydroid assemblages from the Southwestern Atlantic

Ocean (34–42 S)
Gabriel N. Genzano1,2, Diego Giberto2,3, Laura Schejter2,3, Claudia Bremec2,3 & Pablo Meretta1
1 Estación Costera Nágera, Departamento de Ciencias Marinas, Facultad de Ciencias Exactas y Naturales, Universidad Nacional de Mar del Plata,
Mar del Plata, Argentina
2 Instituto Nacional de Investigación y Desarrollo Pesquero (INIDEP), Mar del Plata, Argentina
3 Consejo Nacional de Investigaciones Cientı́ficas y Técnicas (CONICET), Buenos Aires, Argentina

Keywords
Assemblages; biogeography; Hydrozoa;
richness; South Western Atlantic;
Sub-Antarctic.
Correspondence
G. N. Genzano, Estación Costera Nágera,
Departamento de Ciencias Marinas, Facultad
de Ciencias Exactas y Naturales, Universidad
Nacional de Mar del Plata, Funes 3350,
Mar del Plata, Argentina.
E-mail: genzano@mdp.edu.ar
Accepted: 25 April 2008
doi:10.1111/j.1439-0485.2008.00247.x
Abstract
This paper provides updated taxonomic knowledge about hydrozoan species
and provides ecological information including geographical and bathymetric
distributions and biological substrata for the various hydroid assemblages from
the Sub-Antarctic Biogeographical Region, mainly Buenos Aires and the Uru-
guayan coasts. Five of the 41 species found are new records for the study
region. Thirty-one species (75.6%), all found at depths of less than 80 m, have
cosmopolitan distributions. Biodiversity decreased markedly below 80 m depth,
where nine species (22%) distributed in the Southern hemisphere and one
endemic species (2.4%) were found. Most species were non-specific epizoites,
occurring on diverse substrata. A non-parametric multivariate similarity analy-
sis revealed nine species groups that were correlated with large-scale and local
oceanographic patterns and with the availability of suitable substrata. The main
hydroid substrata for eight of the groups were cnidarians, molluscs (mainly
blue mussels), quartzite rocks and sponges. In a single group found in Patago-
nian scallop beds, the main biological substrata were polychaete tubes, other
hydroids and scallops.

continuous work in this area has only begun in the last

Problem
In the Southwestern Atlantic Ocean, species richness, dis-
tributional patterns and the population biology of benthic
hydroids have been studied in tropical and subtropical
Brazilian waters (Grohmann et al. 1997; Rosso & Marques
1997; Kelmo & Attrill 2002; Migotto et al. 2002; Kelmo
et al. 2003), but hydroids of the temperate and sub-Ant-
arctic regions of Uruguay and Argentina remain scarcely
known despite their widespread distribution and high
species richness.
The first taxonomic reports for sub-Antarctic waters
were based on samples from Southern Patagonia at the
end of 19th and the beginning of the 20th centuries.
Hydroids from the Northern Argentine Sea (i.e. Buenos
Aires coast) remained unknown until the 1960s and
two decades (Blanco 1994; Genzano & Zamponi 1997,
2003 and information therein). However, with the excep-
tion of some ecological analyses on colonies from the
rocky intertidal of Mar del Plata (e.g. Genzano 2001,
2002, 2005; Genzano & San Martı́n 2002), the studies
from these vast areas have been largely limited to
taxonomic records; even rudimentary information on the
geographic and bathymetric distributions, habitats and
biological substrata of the recorded species is scarce.
In this paper we provide information about 41 species
of hydroids (508 colonies) of the temperate zone of the
Sub-Antarctic Biogeographical Region, including many
unexplored localities of the Buenos Aires and Uruguayan
coasts. The data include 269 samples obtained from vari-
ous bottom types and biological substrata over more than

Marine Ecology 30 (2009) 33–46 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd 33
Hydroid from SW Atlantic: richness and assemblages
18 years of collecting. The goals of this work are to
update taxonomic (species richness) knowledge, provide
ecological information including geographical and bathy-
metric distributions, biological substrata and colonization
strategies. By characterizing the different hydroid assem-
blages from this study area, we provide a useful starting
point for future investigations.
Material and Methods
Study area
The study area comprises Uruguayan and Argentinean
waters, from 35–42 S and 54–62 W. In general, soft
sediments (sand with minor quantities of shell and mud)
dominate the continental shelf and finer sediments pre-
dominate from the 200-m isobath (170 km from the
coast) to deeper waters (Parker et al. 1997). Harder bot-
toms composed of loose gravel and rocky outcrops are
also present. The region is influenced by the fluvial
Fig. 1. Study area, with the distribution of scallop and mytilid banks and other bottoms type sampled: loess rocks, quartzite rocks, muddy
bottoms and sandy beaches.
34 Marine Ecology 30 (2009) 33–46 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd
Genzano, Giberto, Schejter, Bremec & Meretta
discharge of Rı́o de la Plata and shows a muddy fringe
almost parallel to the northern coast in Uruguay, whereas
southern areas are sandy with some gravel and shells
(Urien & Martins 1979).
Many distinct subtidal faunal assemblages are found
along the study area (see Fig. 1) including the mixohaline
muddy assemblages of the Rı́o de la Plata estuary, marine
sandy assemblages off the Rı́o de la Plata estuary (Roux &
Bremec 1994, 1996; Giberto et al. 2004), mytilid grounds
distributed in middle shelf areas both in Uruguay and
Argentina and along Uruguayan rocky islands (Maytı́a &
Scarabino 1979; Penchaszadeh 1979; Riestra et al. 1992;
Bremec & Roux 1997), assemblages dominated by poly-
chaetes at Bahı́a Blanca (Elı́as et al. 2004), and scallop
(Zygochlamys patagonica) assemblages along the 100-m
isobath (Bremec et al. 2000; Bremec & Lasta 2002).
The intertidal zone is characterized by extensive sandy
beaches (see Escofet et al. 1979; Bastida et al. 1991). There
are a few hard bottoms of consolidated sedimentary rocks
Genzano, Giberto, Schejter, Bremec & Meretta Hydroid from SW Atlantic: richness and assemblages

(loess), which sometimes reach sublittoral areas, present

a
along the Buenos Aires coast. The Mar del Plata area is the
only exception to this general pattern, its coast being char-
acterized by the presence of quartzite rocks that originate
in the Tandilia system (SE Buenos Aires). The large rocky
blocks that characterize the intertidal zone in this area
extend several miles seaward to a depth of 20–25 m (Rı́os
& Alvarez 1989; Rı́os et al. 1990). The Uruguayan littoral
has large rocky promontories of variable composition that
separate sandy coastal beaches.

Data collection
Hydroid colonies were obtained from 168 of the 269 sam-
ples analysed (Fig. 2a). Study material was collected from
various habitats over a period of 18 years. Numerous
samples came from the intertidal fringe, collected since
1989. Others were collected by snorkelling and SCUBA
diving in Mar del Plata harbour and neighbouring sublit-
toral rocky outcrops (to a depth of 33 m) from 1993 to b
2005. Hydroids from other harbours were collected from
panels used for fouling studies. Samples from greater
depths, collected with dredges, came from explorations
off the coasts of Northern Argentina and Uruguay carried
out by the National Institute of Fisheries Research and
Development (INIDEP) since 1987. Sampling was
conducted throughout the year, but a few zones were
sampled only during the summer (Table 1).
Species were classified following the taxonomic system
proposed by Cornelius (1992). Colonies were preserved,
together with their substrata when possible, in 5% forma-
lin solution and deposited in the collections of Departa-
mento de Ciencias Marinas, Mar del Plata University.

Data analysis

The data used in this study were collected over nearly

two decades and come from sampling programmes asso-
ciated with various research activities including fisheries
stock assessment, environmental monitoring, and general
benthic studies. The resulting database is unique for the
study area and contains all the available biological and
ecological information on hydroid species.
The sampling effort was highly variable during the
study period and a number of different sampling methods
were required for the many different bottom types and
depths (Fig. 1). Consequently, it was not feasible to
obtain densities in some cases, or to standardize density
values across habitats. Instead, we converted species pres-
ence–absence data to frequencies using a grid of 1 lati-
tude · 1 longitude (31 quadrats in total) to obtain a
large-scale synoptic view of species occurrence along the
study area (Fig. 2b). Similarity of species composition
Fig. 2. Sampling coverage. a: Positive (filled circles) and negative
(empty circles) sites. b: Frequency grid of 1 · 1º (quadrants 1–31)
indicating total sampling sites (upper number) and positive sites
(lower number) for each quadrant.
among those quadrats was determined by non-parametric
multivariate analysis (Field et al. 1982; Clarke 1993;
Clarke & Warwick 2001). Classification methods (group
average sorting of the Bray–Curtis similarity measures
based on fourth root transformed frequency data) were
carried out using PRIMER software (Clarke & Warwick
2001). SIMPER analysis (similarity percentages) was used
to identify those species which contributed most to
(dis)similarities among and within groups. This analysis
calculates the ‘average similarity’ (contribution of the ith
species to the overall dissimilarity between the groups

Marine Ecology 30 (2009) 33–46 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd 35
Hydroid from SW Atlantic: richness and assemblages Genzano, Giberto, Schejter, Bremec & Meretta

Table 1. Code, depth (m), dates and method of sampling for the hydroid collection.

expedition latitude (S) longitude (W) depth (m) date collection method

Uruguay, intertidal 3445¢ 5450¢–5510 0–1 December, 2000–September, 2001 manual collecting
Uruguay sublittoral 3455¢ 5500¢ S 18 January, 1999 Picard dredge
H 09–93 3405¢–4159¢ 5505¢–6150¢ 17–50 July, 1993 Picard dredge
EH 09-99 3408¢–4110¢ 5306¢–6152 17–50 November, 1999 Picard dredge
H 09-01 3500¢–3720¢ 5210¢–5618¢ – November, 2001 Picard dredge
Banco Limón 3500¢–3750¢ 5450¢–5610¢ 20 February, 2002 modified dredge
H 06 ⁄ 93 3658¢–3756¢ 5553¢–5837¢ 65–75 July 20, 1992 modified oyster dredge
CC 02-02 3736¢ 5636¢ 25–38 February, 2002 by catch
Rocky intertidal, Santa Clara 3750¢ 5730¢ 0–1 October–November, 1990, February, manual collecting
1992. April, 2000–2004
Rocky intertidal, Mar del Plata 3805¢ 5732¢ 0–1 continuously monitored since 1989 manual collecting
Mar del Plata harbour 3806¢ 5727¢ 2–5 continuously monitored since, 1988 snorkeling and SCUBA
diving
Fouling Mar del Plata 3806¢ 5827¢ 0.5 monthly, during 1993 fouling panels
Sublittoral rocky outcrops 3806¢–3808¢ 5728¢–5728¢ 4–14 May, 1997, 1998, August, 1998, SCUBA diving
(Restingas) June, 2000 and continuously
monitored since, 2005
Sublittoral rocky outcrops 3807¢–3810¢ 5724¢–5727¢ 19–23 April, June, 2000 and monthly from SCUBA diving
(Bancos) November, 2000 to December, 2001.
January and April, 2002, February,
2005. May, 2006
OB 02, 04, 06, 10 ⁄ 87. OB 04 3810¢–3920¢ 5511¢–5712¢ 35–507 bimonthly during 1987; November Picard dredge
and 06 ⁄ 88. OB 01 ⁄ 89 1988 and February, 1989
EPEA 3850¢ 5700¢ 50 February, 2003 Picard dredge
Fouling Quequen fouling panels
Fouling Ing. White 3845¢ 6212¢ 0.5 monthly during 1992 fouling panels
CC 06 01 3805¢–3905¢ 5534¢–5550¢ February and March, 2001 modified oyster dredge
CC 10 06 3805¢–3905¢ 5534¢–5550¢ 84–100 October, 2006 modified oyster dredge
Reta, intertidal 3900¢ 6117¢ 0–1 January and April, 1999 manual collecting
Orense, intertidal 3841¢ 5947¢ 0–1 January, 1995, January 1999, manual collecting
April, 2001
El Rincón 3917¢ 5721¢ 69 December, 1997 by catch
El Rincón 3946¢ 6139¢ 72 March, 1998 by catch
El Rincón 4012¢ 5558¢ 500 March, 2003 by catch
El Rincón 4118¢ 6045¢ 70 December, 1997 by catch

considered) and the ‘internal similarity’ (contribution
each species makes to the average similarity within each
group considered) (see Clarke & Warwick 2001 for
details).
Results
A total of 41 hydroid species was found, five of them new
for the area. Only six species previously reported in the
literature were not found in our samples. The anthoathec-
atae colonies were less represented than leptothecate ones
(8 versus 33 species, respectively). Most species (71%) do
not have medusa stage and the larva (planula or actinula)
is the only free swimming planktonic stage; the other spe-
cies (29%) release medusae.
The complete list of all benthic hydroids found is listed
in Table 2. Zoogeographical notes and biological remarks
were included when referring to unknown or scarcely
reported species in the region. Depth distributions are
summarized in Fig. 3.
A total of 502 colonies were found colonizing 58 differ-
ent substrata (Table 3). The species–substratum analysis
shows that cnidarian colonies, especially stems of other
hydroids, supported the greatest number of epizoic spe-
cies, followed by sponges, mollusks, algae, polychaete
tubes and rocks. The remaining groups were occasionally
found with epizoic hydroids. However, this general
scheme changes in different habitats (see below).
Halecium beanii and Plumularia setacea were the spe-
cies most frequently collected (52% of the analysed sam-
ples each) in the study area, followed by Amphisbetia
operculata (39%), Sertularella striata (32%), Obelia
longissima and Symplectoscyphus subdichotomus (29%
each). The highest species richness was found in quadrat
14 (29 species), followed by quadrats 15, 17 (11 species
each) and 16 (10 species). The remaining quadrats had

36 Marine Ecology 30 (2009) 33–46 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd
Genzano, Giberto, Schejter, Bremec & Meretta Hydroid from SW Atlantic: richness and assemblages

Table 2. Benthic hydroids found in the study area. Species code used in this table and Table 3, and the quadrat where each species was found
are indicated. Asterisks indicate species previously reported from the area but not found in this study.

species code quadrat remarks

subclass Anthoathecata order Filifera

Family Clavidae
Rhizogeton nudus (Broch, 1909) Rn (s) 14 –
Family Bougainvilliidae
Bimeria vestita Wright, 1859 Bv (s) 14 –
Bougainvillia macloviana (Lesson 1830) Bma (m) 14 –
Bougainvillia muscus Allman, 1863 Bm (m) 6, 22 –
Family Eudendriidae
Eudendrium ramosum (Linnaeus 1758) Er (s) 4, 14, 15, 16, 28 –
Order Capitata Family Corynidae
Coryne eximia Allman, 1859 Ce (m) 14, 20 Species previously reported in
the region as Syncoryne sarsii or
Sarsia sarsii (see Genzano &
Zamponi 1997, 2003)
Family Corymorphidae Corymorpha –
januarii Steenstrup, 1854 Cor (m) 23
Family Tubulariidae
Ectopleura crocea (Agassiz, 1862) Ec (s) 2, 14, 20 –
*Hybocodon unicus (Browne 1902) The record of H. unicus reported
by Zamponi & Facal (1987) is a
missidentification. The same specimen
was revised in this study and identified
as Corymorpha januarii (see above)
subclass Leptothecata order Conica
family Lafoeidae
Hebella scandens (Bale, 1888) Hs (m) 14 –
Filellum antarcticum (Hartlaub, 1904) Fa (s) 15, 17 –
Filellum serratum (Clarke, 1879) Fs (s) 7, 8, 11, 14, 15, 17, 19 –
Lafoea fruticosa (Sars, 1851) Lf (s) 9, 12, 13, 14, 15, 16, 17 –
Grammaria magellanica Allman, 1888 Gm (s) 13, 16, 17 Species previously reported from Tierra
del Fuego and Malvinas Islands to
Penı́nsula de Valdes (see Blanco et al.
1994 and information therein). This
record in Buenos Aires coast extends
its geographical distribution northwards
Stegopoma irregularis (Totton, 1930) Sirr (s) 30 Species only reported in deep waters
around Malvinas Islands (Blanco et al.
1994). This record on Buenos Aires
coast extends its geographical
distribution northwards
Family Haleciidae
Halecium beanii (Johnston, 1838) Hb (s) 7, 8, 9, 10, 11, 12, 14, 15, 16, –
17, 18, 19, 20, 21, 23, 26
Halecium delicatulum Coughtrey, 1876 Hd (s) 14 –
Family Campanulinidae
Opercularella belgicae (Hartlaub, 1904) Ob (s) 14 The genus Opercularella includes species
that produce fixed sporosarcs and also
some species with unknown gonophores.
The analysed colonies have scarce and
immature gonothecae. The generic name
is thus still provisional pending a better
knowledge of the gonophore
Phiallela chilensis Hartlaub, 1905 Pch (m) 1, 6, 9, 17 –
Family Syntheciidae
Synthecium robustum Nutting, 1904 Sr (s) 7, 9, 12, 13, 14, 15, 16, 17 –

Marine Ecology 30 (2009) 33–46 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd 37
Hydroid from SW Atlantic: richness and assemblages Genzano, Giberto, Schejter, Bremec & Meretta

Table 2. Continued

species code quadrat remarks

Family Sertulariidae
Dynamena cornicina McCrady, 1858 Dc (s) 14 –
Amphisbetia operculata Ao (r) 2, 3, 8, 11, 12, 14, 15, –
(Linnaeus, 1758) 24, 25, 27, 28, 29
Sertularella conica Allman, 1877 Sc (s) 14, 15 –
Sertularella mediterranea Hartlaub, 1901 Sm (s) 9, 14, 16, 17 –
Sertularella striata Stechow, 1923 Sst (s) 7, 8, 9, 11, 12, 14, 17, –
19, 28, 29
*Sertularella uruguayensis Mañé-Garzon – – This species was described from Uruguayan coasts
& Milstein 1973 by Mañe-Garzon & Milstein (1973), based on scarce
fragments and one immature gonotheca, which
raises concerns about its validity. The original
description constitutes the only record
Symplectoscyphus subdichotomus Sys (s) 9, 10, 12, 13, 14, 15, –
(Kirchenpauer, 1884) 16, 17, 25
Family Halopterididae
Monastaechas quadridens Mq (s) 14 –
(McCrady, 1858)
Family Plumulariidae
Plumularia insignis Allman, 1883 Pi (s) 3, 7, 16 –
Plumularia pulchella Bale, 1888 Pp (s) 14 –
Plumularia setacea (Linnaeus 1758) Ps (s) 2, 5, 7, 8, 9, 10, 11, –
14, 15, 19,
20, 21, 22, 23, 24, 25
Family Aglaopheniidae
Aglaophenia acacia Allman, 1883 Aa (s) 5, 8, 11, 12, 14, –
19, 26, 28
Thecocarpus canepa Blanco and Tc (s) 16, 17, 30, 31 –
Miralles, 1971
Order Proboscoida Family Campanulariidae
Campanularia agas Cornelius,1982 Ca (s) 4, 11, 14, 24, 25, –
26, 28, 29
Campanularia hincksii Alder, 1856 Ch (s) 14
*Campanularia subantartica – – This species was frequently reported from Patagonia,
Millard, 1971 Tierra del Fuego and Malvinas as C. everta,
C. raridentata, C. tincta and C. tincta euricaulix,
(see sinonimies in Blanco 1994). Recently, matures
colonies were found in Ushuahia, Tierra del Fuego
(Genzano, unpublished data). However, records of
small colonies found in Buenos Aires coast and
assigned to C. subantartica (Genzano 1990) are
considered doubtful. Its presence in this area needs
to be confirmed
Clytia gracilis (M. Sars, 1851) Clg (m) 11, 14, 15, 24, 27, 28 –
Clytia laevis Hartlaub, 1905 Clae (m) 19 The unique record from South Western Atlantic Ocean
was near Tierra del Fuego (Vervoort 1972). Our specimens
constitute the first record on the Buenos Aires coasts,
extending northwards its geographical distribution
Clytia hemispherica Linnaeus, 1767 Clh (m) 25 –
*Gonothyraea inornata Nutting, 1901 – – This species was not found in our samples, but it was
previously reported in the fouling community from
Mar del Plata harbour. However, that identification is
in doubt because specimens were scarce and poorly
preserved (Blanco 1968); presumably that record was referring to
G. loveni colonies
Gonothyraea loveni (Allman, 1859) Gl (s) 14 –

38 Marine Ecology 30 (2009) 33–46 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd
Genzano, Giberto, Schejter, Bremec & Meretta Hydroid from SW Atlantic: richness and assemblages

Table 2. Continued

species code quadrat remarks

Hartlaubella gelatinosa Hg (s) 21, 24, 25, 27, 28 This cosmopolitan species, was previously reported in
(Pallas, 1766) Argentinean waters from south Patagonia
(Jäderholm 1903). This first record in Buenos Aires
coast extends its geographical distribution
*Obelia angulosa Bale, – – This species, not observed in our collections, was
1888 reported as part of the fouling community in Mar
del Plata harbour by Blanco (1968), who remarked
its similarity with O. australis, which is considered a
synonym of O. dichotoma. Further studies will be
necessary to determine the taxonomic status of
O. angulosa
Obelia bidentata Obi (m) 14, 21, 22, 26 –
(Clarke, 1875)
*Obelia dichotoma – – Zamponi (1987) reported this species in Mar del
(Linnaeus, 1758) Plata coast, located in the same area where we
have found abundant colonies of O. longissima
during more than 16 years of continuous sampling.
In this study, the specimens of O. dichotoma
reported by Zamponi (op cit.) are considered
a missidentification and therefore conspecific
with O. longissima
Obelia longa Stechow, Ol (m) 16 –
1921
Obelia longissima Ols (m) 7, 8, 12, 14, 19, –
(Pallas, 1766) 20, 22, 24, 25

Life cycle of species is indicated as: (s), sporosarc or fixed medusoid; (r), reduced medusae or medusoid release; (m), fully functional medusae, real
metagenetic life cycle. Rn, Rhizogeton nudus; Bv, Bimeria vestita; Bma, Bougainvilla macloviana; Bm, Bougainvillia muscus; Er, Eudendrium
ramosum; Ce, Coryne eximia; Cor, Corymorpha januarii; Ec, Ectopleura crocea; Hs, Hebella scandens; Fa, Filellum antarcticum; Fs, Filellum
serratum; Lf, Lafoea fruticosa; Gm, Grammaria magellanica; Sirr, Stegopoma irregularis; Hb, Halecium beanie; Hd, Halecium delicatulum;
Ob, Opercularella belgicae; Pch, Phiallela chilensis; Sr, Synthecium robustum; Dc, Dynamena cornicina; Ao , Amphisbetia operculata; Sc, Sertular-
ella conica; Sm , Sertularella mediterranea; Sst , Sertularella striata; Sys , Symplectoscyphus subdichotomus; Mq , Monastaechas quadridens;
Pi, Plumularia insignis; Pp, Plumularia pulchella; Ps, Plumularia setacea; Aa, Aglaophenia acacia; Tc, Thecocarpus canepa; Ca, Campanularia agas;
Ch, Campanularia hincksii; Clg, Clytia gracilis; CIae, Clytia laevis; CIh, Clytia hemispherica; GI, Gonothyraea loveni; Hg, Hartlaubella gelatinosa;
Obi, Obelia bidentata; Ol, Obelia longa; Ols ,Obelia Iongissima

lower species richness, and in quadrats 1, 18 and 31 only
one species was collected.
Species composition and richness varied considerably
in the different zones defined by the cluster analysis
(Fig. 4), as follows:
Group 1 (63% internal similarity SIMPER analysis). This
group clusters samples from two sectors from deep cold
waters (>500 m depth), where the anchoring species
Thecocarpus canepa (100% average similarity) and Stegophoma
irregularis were exclusively collected from soft bottoms.
Group 2 (63% internal similarity). Clusters sampled from
the mixohaline muddy bottoms of the Rı́o de la Plata. The
species found in the area were Phialella chilensis (100%),
attached to living specimens of the clam Mactra isabelleana
(< 15 m depth), and Bougainvillia muscus (2 m depth).
Group 3 (35.5% internal similarity). This heterogeneous
group (five species) clusters samples from two distant
sectors, where Campanularia agas (100%) was the only
shared species.
Group 4 corresponds to sector 3 (off Punta del Este)
where two species, A. operculata and Plumularia insignis,
were collected.
Group 5 (49.5% internal similarity). This area (‘El
Rincón’ and off Rı́o Negro) is represented by 11 species
assemblage and characterized by A. operculata (43%),
Hartlaubella gelatinosa (20%) and C. agas (18%). Hydroid
colonies and polychaete tubes (36.4% each) were the
main substrata, followed by fish egg capsules and Mytilus
edulis platensis.
Group 6 (36% internal similarity). This group (nine
species assemblage) clusters samples collected in hard bot-
toms, both from rocky intertidal and subtidal Mytilus
beds. The most characteristic species were P. setacea
(78%), Ectopleura crocea (12%) and O. longissima (11%).
The main substrata were polychaete tubes (43%),
sponges, hydroids, mytilids and rocks (14.3% each).
Group 7 (49% internal similarity). This hetero-
geneous group (six species assemblage) clusters samples

Marine Ecology 30 (2009) 33–46 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd 39
Hydroid from SW Atlantic: richness and assemblages Genzano, Giberto, Schejter, Bremec & Meretta

Table 3. List of substrata colonized by hydroid species in the study area (see species codes in Table 2).

SUBSTRATA ⁄ HYDROIDS Rn Bv Er Ce EC Hs Fa Fs Lf Gm Hb Hd Ob Pch Sr DC Ao Sc Sm Sst Sys Mq Pi Pp Ps Aa Ca Ch Clg Hg Gl Obi Ols Total

Algae 13
Ulva sp. X 1
Corallina officinalis X X X X X 5
Bossiella orbignyana X X X 3
Rhodymenia sp. X X X X X X X X X 9
Polysiphonia sp. X 1
Phaeophyta X X 2
Dyctiota sp. X X X X X 5
unidentified red algae X 1
Porifera 14
Tedania sp. X X X X X X 6
Esperiopsis sp. X 1
Hymeniacidon X X X X X 5
sanguinea
Halichondria sp. X X X 3
lophon sp. X X X 3
Haplosclerida X X 2
unidentified sponge X X X X X X X 7
Cnidaria 18
Bimeria vestita X 1
Ectopleura crocea X X X X X X 6
Eudendrium ramosum X X X 3
Halecium beanii X X X X X 5
Dynamena cornicina X 1
Amphisbetia operculata X X X X X 5
Sertularella striata X X X X 4
Sertularella mediterranea X X X X X X X 7
Symplectoscyphus X X X 3
subdichotomus
Plumularia setacea X X X X X X 6
Monastaechas quadridens X X 2
Aglaophenia acacia X 1
Obelia longissima X 1
Plumulariidae X 1
Tubulariidae X X 2
Campanulariidae X 1
Trypalea clavaria X X X 3
Gorgonacea X X X 3
Thesea sp. X X X 3
unidentified Anthozoa X X 2
Anthipatharia X X 2
Bryozoa 7
Bicelariella sp. X X X X X X X 7
Membranipora sp. X 1
unidentified bryozoan X 1
Polychaeta 18
Sabellidae X X X X 4
unidentified Polychaeta X X X X X X X X X X X X X X X X 16
Mollusca 17
Mactra isabelleana X 1
Brachydontes rodriguezi X X X X 4
Mytilus edulis platensis X X X X X X X X X X X 11
Zygochlamys patagonica X X X X X 5
Ostrea puelchana X X X 3
Crepidula sp. X X X X X X 6
shells X 1

40 Marine Ecology 30 (2009) 33–46 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd
Genzano, Giberto, Schejter, Bremec & Meretta Hydroid from SW Atlantic: richness and assemblages

Table 3. Continued

SUBSTRATA ⁄ HYDROIDS Rn Bv Er Ce EC Hs Fa Fs Lf Gm Hb Hd Ob Pch Sr DC Ao Sc Sm Sst Sys Mq Pi Pp Ps Aa Ca Ch Clg Hg Gl Obi Ols Total

Crustacea 2
Balanus sp. X 1
Majiidae X 1
Tunicata 9
Ciona sp. X 1
Ascidiacea 1 X X X X X X X X 8
Ascidiacea 2 X 1
Sycozoa umbellata X X 2
Pisces 2
egg capsules (Rajidae) X X 2
others 17
rocks X X X X X X X X X X X X X X X 15
wood X 1
panels X X X X X 5
total 2 8 6 15 9 1 3 9 5 1 26 3 1 3 3 2 4 1 11 12 6 2 1 2 17 1 17 1 22 1 1 4 13 213

Fig. 3. Depth and geographical distribution

of the hydroid species reported in this study.

characterized by H. beanii (72%) and P. setacea (28%).
Hydroid colonies were mainly found on polychaete tubes
and mytilid valves (66% and 33% of the found substrata,
respectively).
Group 8 (55% internal similarity). This group clusters
samples with highest species richness (31 species), charac-
terized by H. beanii (33%), S. striata (19%), O. longissima
and P. setacea (13% each), Aglaophenia acacia (12%),
Filellum serratum (11%) and A. operculata (7%). The
samples were collected between the intertidal zone and
50 m depth, most of them in mytilid beds. The main
substrata were cnidarians (hydroids and gorgonaceans;
30%), molluscs (17%, mainly intertidal Brachydontes
rodriguezii and subtidal M. edulis platensis); quartzite
rocks (14%) and sponges (12%).
Group 9 (54% internal similarity). This group consists
mainly of samples from Patagonian scallop beds and
includes 18 hydroid species. The most characteristic spe-
cies are of magellanic origin, such as S. subdichotomus
(26%), Lafoea fruticosa (22%), Synthecium robustum
(17%), which are distributed between 50 and 250 m
depth, and H. beanii (12%). The main biological sub-
strata were polychaete tubes (46%) and hydroid colonies
(31%).

Marine Ecology 30 (2009) 33–46 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd 41
Hydroid from SW Atlantic: richness and assemblages
Discussion
A total of 41 species were found in the study area, five of
which resulted in new records. Six other species previ-
ously reported for the region were not present in our
samples and the validity of some species or misidentifica-
tion of others is discussed (see Table 2). This number of
species is quite high considering that the reported species
richness does not exceed 120 for the entire Argentinean
continental shelf (Genzano & Zamponi 1997). Most of
the habitats herein studied (intertidal and sublittoral
rocky outcrops, scallop beds, mytilid beds, etc., see
Table 1) have been frequently monitored during the last
decade, showing a similar hydroid species composition
over time.
The largest number of species was found at depths
shallower than 70–80 m, with biodiversity decreasing
markedly beyond a depth of 80 m. This relatively high
number of species in shallow water could be due to the
diversity of substrata and microhabitats. Information
about other invertebrate groups in the study area shows
that, in general, the species richness reflects the intensity
of sampling effort expended on an area (López Gappa
2000; Giberto 2003; López Gappa & Landoni 2005). How-
ever, the 41 hydroid species recorded at present seem to
fit closely to total hydroid richness, as the sampling effort
has increased dramatically over time, mainly during the
last years in coastal and shelf areas (from 8 expeditions in
the early 1990s to 39 in recent years). Hydroids from
42 Marine Ecology 30 (2009) 33–46 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd
Genzano, Giberto, Schejter, Bremec & Meretta
Fig. 4. Study area with spatial distribution of
species assemblages (A) clustered in the
multivariate analysis (B).
previously unexplored areas are included (e.g. Rı́o de la
Plata estuary, Patagonian scallops beds). However, new
findings are expected as deeper bottoms (> 200 m) still
remain almost unstudied.
Many hydroid species undergo sharp seasonal cycles
(Calder 1990; Bavestrello et al. 2005). The use of records
from different periods of the year could cause biodiversity
to be underestimated. However, seasonality is more likely
to be a factor in small colonies living in shallower waters
(Genzano 1994; Genzano et al. 2002) and shallow sam-
pling was conducted all year around, or at least during
the warmest season, when most hydroid species were
present. On the Buenos Aires coast, few species seem to
thrive in winter, but they are present in the benthic com-
munity throughout the year (Genzano 1994; Genzano
et al. 2002).
The study area spans two different biogeographic prov-
inces included in the Sub-Antarctic Biogeographic Region
(Balech 1964), which adds complexity, caused by a large
number of possible niches, to the spatial patterns of both
subtidal and intertidal benthic assemblages. The
Magellanic Province (from Chiloé island, eastern Pacific
Ocean, to 35 S, southwest Atlantic) is dominated by
cold-temperate waters, and the Argentinean Province
(from 43 ⁄ 44 S to Cabo Frio, 23 S) is characterized by
warm temperate waters. According to the cluster analysis,
the 41 species collected can be ascribed to five major
groups. Two groups of hydroid species are related to the
cold waters of sub-Antarctic origin: Group 1, which
Genzano, Giberto, Schejter, Bremec & Meretta
includes soft bottom species collected in the two deepest
sampling stations available, and Group 9, which has spe-
cies from scallop (Z. patagonica) bed assemblages mainly
along the 100-m isobath. Group 5 has colonies from ‘El
Rincón’ and off Rı́o Negro, with species mainly settled on
other hydroid colonies and polychaete tubes. Groups 6
and 8 cluster many species that mainly inhabit mytilid
banks from intertidal to middle shelf warm temperate
waters of the Argentine province. Group 2 has the two
species collected in mixohaline muddy bottoms of the
Rı́o de la Plata. Finally, Groups 3, 4, and 7 cluster differ-
ent hydroid species collected from different shallow areas
both along both the Uruguayan and Buenos Aires coasts.
Most of the hydroid species found in our study are
cosmopolitan (31 species, 75.6%). Some species were
exclusive to the Southern hemisphere (nine species,
22%), whereas only T. canepa (one species, 2.4%) is
endemic to the Southwestern Atlantic. The shallow-water
hydroid fauna comprises a large number of widely
distributed species (87%), whereas in deeper waters,
cosmopolitan species decrease (50%) and the number of
species having sub-Antarctic or southern distributions is
higher. A few species showed a wider bathymetric range,
from shallow to deep waters, with H. beanii being
the only species found at all analysed depths. Analysis of
hydroid distribution patterns confirms that marked
changes in faunal composition occur between 60 and
80 m. This is a transitional zone between the mentioned
biogeographical provinces, as previously reported for
cnidarians, bryozoans, molluscs and echinoderms (López
Gappa & Lichtschein 1988; Roux et al. 1988; Bastida et al.
1992; Zamponi et al. 1998).
Hydroids tend to be substratum generalists. Most
species found in this study were non-specific epizoites,
occurring on diverse substrata. Few species exhibited
apparent substratum preferences; colonies of Filellum
serratum and F. antarcticum were more frequent on other
hydroid stems, a common characteristic of many Lafoeidae
species (see Boero et al. 1997). Hebella scandens was
found exclusively on stems of Dynamena cornicina; a close
species-specific relationship has already been reported in
the area (Genzano 1992).
Hydroids are generally more diverse and abundant on
firm or hard substrata (Calder 1991). The colonization on
different substrata is related to the colony morphology.
Species with bushy forms or grouped in dense clumps,
such as large colonies of the tubularid E. crocea, the sertu-
larids A. operculata, Sertularella mediterranea and the plu-
mularids P. setacea and A. acacia, attach directly to rocks
or, more frequently, to the shells of mytilid bivalves.
Species usually found on rocky outcrops can also
invade soft bottoms by settling on small ‘islands’ of hard
substratum dispersed across the sedimentary environment
Marine Ecology 30 (2009) 33–46 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd
Hydroid from SW Atlantic: richness and assemblages
(Morri et al. 1991). Plumularia setacea and A. operculata
were frequently found on small rocks or pebbles sur-
rounded by sandy bottoms. These colonies were smaller
and less abundant than those on rocky outcrops.
Clumps of large colonies provided substrata for other
hydroid species, such as Bimeria vestita, F. serratum, C. agas
and Clytia gracilis. These epizoic colonies showed a ‘guer-
rilla strategy’ with an extensive hydrorhizal growth; the
asexual reproduction provides new hydrocaulus which can
be colonized quickly by epizoic hydroids from neighbour-
ing hydrocauli by means of stoloniferous growth. This
colonization strategy of substrata reduces inter-specific
competition for space and avoids the negative effects of sed-
iment deposition (Hughes 1975; Boero 1984; Garcı́a Rubies
1987; Llobet et al. 1991; Genzano & Rodrı́guez 1998;
Genzano 2002). The use of this mode of asexual reproduc-
tion permits a rapid occupation of the substrata, and it is
remarkable that these epizoic colonies (in species both with
metagenetic and holobenthonic life cycles, see Table 1)
rarely develop gonothecae. Gonangia were never found in
any of the analysed Lafoeidae of which, populations seem
to be maintained by means of asexual propagation.
Gonothyraea loveni was only species found attached to
experimental submerged plates for fouling studies con-
ducted in Mar del Plata harbour. Although this is one of
the best studied areas, this species has never been found
in the natural community. It is a hydroid widely dis-
tributed in both hemispheres, frequently reported in
harbours and probably spread by ships (Millard 1975).
In the same way, Obelia bidentata, another campanulariid
having a disjunct world wide distribution, seems to be
associated with human dispersal mechanisms and it was
possibly introduced by nautical activities. This species was
found on submerged plates both in Mar del Plata and In-
geniero White harbours, and occasionally found near the
latter location in the natural community, living on poly-
chaete tubes and bivalves (see Table 3). Monastaechas
quadridens is widely distributed in tropical waters but
mature colonies have been reported in recent years in
temperate waters of Buenos Aires (Mar del Plata sublit-
toral rocky outcrops; Genzano et al. 2002, and this study).
Further studies will be necessary to assess its successful
colonization southwards.
Only three species are able to live directly on soft sub-
strata: Corymorpha januarii as a ‘digging’ species, living
partially buried in the sediments, and T. canepa and
P. insignis as ‘rooting’ species which settle initially on a
small firm substrate (e.g. pebbles, shell debris), grow in size,
and then stay erect by an anchoring system. Many other
species found in soft sediment areas were attached to poly-
chaete tubes, sponges and scallops. The importance of
worm tubes as hydroid substrata on soft bottoms was noted
by Less (1986). In areas where hard substrata are scarce,
43
Hydroid from SW Atlantic: richness and assemblages
tubes provide a stable surface on which colonies can settle,
grow and extend at least a few centimetres above the
abrasive sand–water interface. Additionally, the empty
tubes are sufficiently resilient to remain on the sea floor
for a long period of time, supporting the attached hydroid
epibionts.
Puce et al. (2005) pointed out that the associated
hydroid fauna thriving on or inside sponges shows a wide
range of relationships with its hosts. Hydroids may be
simply epibiotic on sponges. Alternatively, the stolons
may grow inside the sponge body, the polyps emerging
from the sponge surface or, in other cases, stolons and
branches may develop deep inside the sponge body, pro-
ducing a skeletal network for sponge growth. In our sam-
ples, most colonies growing on sponges showed their
stolons running on the host surface, not embedded in the
sponge tissues, and might be considered occasional epizo-
ites. By contrast, the sponges Tedania sp. and Halichon-
dria sp. can overgrow the hydrorhizae and basal portion
of hydrocauli of P. setacea, being occasional epizoites of
the hydroid colonies.
Scallops are common substrata for hydroids on deep
soft bottoms (e.g. Henry & Kenchington 2004). In our
study area, scallops not only provide substrata for sev-
eral colonies (Schejter & Bremec 2007 and Table 3),
but in addition the hydroids Grammaria magellanica
and S. subdichotomus constitute the main primary set-
tlement substrate for Z. patagonica spat (Bremec et al.
2008). The availability of filamentous substrata is an
important factor influencing bivalve population recruit-
ment. On the coasts of Buenos Aires, hydroid colonies
of A. operculata, P. setacea and O. longissima frequently
serve as primary substrata for mytilid plantigrades, as
other filamentous substrata, as algae, are very small and
scarce (Genzano et al. 2003). The authors suggested that
big hydroid colonies provide refuge from predators
such as echinoderms during the first benthic stages and
could ensure a higher probability of survival to scallop
spat by reducing predation from gastropods, sea urchins
and sea stars (Botto et al. 2006), which are very
abundant in the scallop communities (Bremec et al.
2000).
This study contains baseline information that may be
useful in future analyses, such as changes in biodiversity
attributable to natural and anthropogenic impact (i.e. glo-
bal warming, overfishing). Our present study presents an
updated inventory of hydrozoan species, their biological
substrata and bathymetric ranges for the temperate zone
of Buenos Aires and Uruguayan waters (Sub-Antarctic
Biogeographical Region). Distinct hydroid assemblages
are defined, at a large scale, by the oceanographic patterns
and, locally, by the availability of suitable substrate for
the development of the colonies.
44 Marine Ecology 30 (2009) 33–46 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd
Genzano, Giberto, Schejter, Bremec & Meretta
Acknowledgements
This paper was partially funded by EXA 372 ⁄ 07, PICT
1-15080, PICT 2006 No.1553, PIP 5009, Antorchas
13900-13. This is INIDEP Contribution No. 1518.
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