European Journal of Taxonomy 18: 1-21
http://dx.doi.org/10.5852/ejt.2012.18
BY
This work is licensed under a Creative Commons Attribution 3.0 License.
ISSN 2118-9773
www. europeanj ournaloftaxonomy. eu
2012 • Rob W.M. Van Soest et al.
Research article
Sponges of the family Esperiopsidae (Demospongiae, Poecilosclerida)
from Northwest Africa, with the descriptions of four new species
Rob W.M. Van Soest, Elly J. Beglinger & Nicole J. de Voogd
Naturalis Biodiversity Center, Darwinweg 2, 2333 CR, Leiden, The Netherlands
Email: [email protected]
Abstract. Sponges belonging to the genera Amphilectus Vosmaer, Esperiopsis Carter and Ulosa
de Laubenfels of the family Esperiopsidae were collected during 1986 and 1988 expeditions of the
Netherlands Centre for Biodiversity Naturalis (at that time the National Museum of Natural History
at Leiden and the Zoological Museum of Amsterdam) in waters off the coasts of Mauritania and the
Cape Verde Islands. Four new species, Amphilectus utriculus sp. nov., Amphilectus strepsichelifer
sp. nov., Esperiopsis cimensis sp. nov., Ulosa capblancensis sp. nov., and two already known species,
Amphilectus cf. fucorum (Esper) and Ulosa stuposa (Esper) are described and discussed.
Keywords. Sponges, new species, Esperiopsidae, Mauritania, Cape Verde Islands.
Van Soest R.W.M., Beglinger E.J. & de Voogd N.J. 2012. Sponges of the family Esperiopsidae (Demospongiae,
Poecilosclerida) from Northwest Africa, with the descriptions of four new species. European Journal of Taxonomy
18: 1-21. http://dx.doi.org/10.5852/eit.2012.18
Introduction
Knowledge of the sponge fauna of Northwest Africa is poor, with few recent publications and older
papers usually based on specimens obtained during wide-ranging expeditions with often only summary
descriptions (Lendenfeld 1907; Topsent 1918, 1928; Arnesen 1932; Burton 1956; Levi 1952, 1956,
1959, 1960; Van Soest 1993a, b). Preliminary analysis (Van Soest 1993a) suggested that the main
affinity of the sponge fauna of Northwest Africa lies with the Lusitanian and Western Mediterranean
faunas, with minor affinity with the Tropical Western Atlantic, and a considerable endemic element.
Expeditions made in the 1980’s (CANCAP 1-7; Mauritania II) yielded ample materials from the coasts
of Mauritania, the Canary and Madeira Archipelagoes and the Cape Verde Islands, incorporated in the
collections of the former Rijksmuseum van Natuurlijlce Historie at Leiden and the Zoologisch Museum
Amsterdam, recently merged into the collections of the Netherlands Centre for Biodiversity Naturalis
at Leiden. Sponges were collected by various methods (wading, snorkeling, SCUBA, Van Veen grab,
Agassiz trawls and rectangular dredges) from shallow water down to >1000 m. This material was
supplemented by specimens, previously collected by F.P. Vermeulen in Mauritanian and Senegalese
waters. The combined collections contain several sponges new to science and many interesting new
finds of ill-known sponges. A number of scientific studies containing descriptions of sponges from
these Northwest African collections preceded the present one, e.g. on Haplosclerida Topsent, 1928 (De
Weerdt & Van Soest 1986), the genera Didiscus Dendy, 1922 (Hiemstra & Van Soest 1991), Acarnus
Gray, 1867 (Van Soest et al. 1991), Hemiasterella Carter, 1879 (Voultsiadou-Koukoura & Van Soest
1
European Journal of Taxonomy 18: 1-21 (2012)
1991), and Dercitus Gray, 1867 (Van Soest et al. 2010). The present paper addresses sponges of the
poecilosclerid family Esperiopsidae Hentschel, 1923 collected off the coasts of Mauritania and the
Cape Verde Islands. The family contains four accepted genera (Van Soest & Hajdu 2002b), three of
which appear to have representatives in Northwest African waters, viz. Amphilectus Vosmaer, 1880,
Esperiopsis Carter, 1882, and Ulosa de Taubenfels, 1936. Only the genus Semisuberites Carter, 1877
has so far not been recorded. The three genera present in the material are easily distinguished, possibly
because they are ‘artificial’, but this remains unchallenged as no comprehensive phylogenetic research
has been done on members of this group. Skeletons consist of reticulated bundles of styles, there is
no special surface arrangement of spicules, microscleres combine the presence of chelae and sigmas
(Esperiopsis ), consist of chelae only ( Amphilectus ), or are absent {Ulosa). Whereas three genera tend to
have cold-water affinity (Van Soest et al. 2012), at least Ulosa appears to have also tropical or warm-
water representatives.
Fig. 1. Map of Northwest Africa, showing the approximate localities from which the sponges were
collected. 1. Mauritania, Cap Blanc. 2. Mauritania, Banc d’Arguin. 3. Mauritania, SW of Cap Timiris 4.
Cape Verde Islands, Sao Vicente 5. Cape Verde Islands, Ilheu de Cima.
2
VAN SOEST R.W.M., BEGLINGER E.J. & DE VOOGD N.J., West African Esperiopsidae
Material and methods
Specimens were collected by trawls, dredges and grabs during the CANCAP 7 Expedition (August 1986,
Cape Verde Islands, on board of HNLMS Tydemann ) and Mauritania II Expedition (June 1988, on board
of RV Tyro) (Fig. 1). All material was provisionally identified on board and subsequently preserved in
96% ethanol. Until recently, the material was incorporated in the collections of the Zoological Museum
of the University of Amsterdam (ZMA), but at present it is housed in the Netherlands Centre for
Biodiversity Naturalis at Leiden. The acronym ZMA is maintained as part of the registration numbers of
all specimens mentioned. Collection data are provided with each treatment of the species below. To study
the skeletal structure, thick sections were made by hand, air-dried on a hotplate, and mounted in Canada
Balsam. Stacked (automontage) light microscopy images of these thick sections were made using a
Leica DM5500 microscope. For measurements of the spicules and SEM examination dissoluted spicule
suspensions were made with concentrated NaOHCl, washed five times in distilled water and mounted on
light microscopic slides and SEM stubs. Spicule measurements (minimum-mean-maximum) are based
on 25 spicules of each category or type for each individual specimen. SEM photos of spicules were
combined on a black background, aligned, and cleaned using Photoshop CSS licensed to R.W.M. Van
Soest.
Results
Phylum Porifera Grant, 1836
Class Demospongiae Sollas, 1885
Order Poecilosclerida Topsent, 1928
Suborder Mycalina Hajdu, Van Soest & Hooper, 1994
Family Esperiopsidae Hentschel, 1923
Remarks
Below we assign West African species to esperiopsid genera in accordance with the account of the
family by Van Soest & Hajdu (2002b). Species with only a single chela size, and no other microscleres
are assigned to Amphilectus. The species with sigmas included is assigned to Esperiopsis. Species
without microscleres are assigned to Ulosa.
Genus Amphilectus Vosmaer, 1880
Remarks
Amphilectus differs from the closely related genus Esperiopsis in the lack of additional microscleres and
the generally smaller size of the styles. The difference in the latter character was fixed on 400 pm (Van
Soest & Hajdu 2002b), but this is to be interpreted loosely, as several species in both genera possess
styles with lengths just under or above 400 pm. The genus to date contains 17 species (Van Soest et
al. 2012), predominantly from temperate and cold water, evenly spread over the North Atlantic (five
species), North Pacific (three species), South Atlantic (six species), and South Pacific waters (three
species). The Atlantic species are summarized in Table 1.
Amphilectus utriculus sp. nov.
Figs 2-3
Etymology
utriculus (L.) = small water bag, referring to the hollow, flattened shape.
3
European Journal of Taxonomy 18: 1-21 (2012)
Table 1 . Summary of characters of Atlantic sponges belonging to genera Amphilectus Vosmaer, 1880
and Esperiopsis Carter, 1882.
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4
VAN SOEST R.W.M., BEGLINGER E.J. & DE VOOGD N.J., West African Esperiopsidae
Material examined
Holotype
ZMA Por. 22592, Mauritania, SW of Cap Timiris, depth 260-280 m, muddy bottom, coll. R.W.M. Van
Soest & J.J. Vermeulen, Mauritania II Expedition stat. 39/01, 18.8°N 16.7167°W, 3.5 m Agassiz trawl,
10 Jun.1988.
Paratypes
ZMA Por. 06636, 53 specimens, Mauritania, SW of Cap Timiris, depth 260-280 m, muddy bottom, coll.
R.W.M. Van Soest & J.J. Vermeulen, Mauritania II Expedition stat. 39/01, 18.8°N 16.7167°W, 3.5 m
Agassiz trawl, 10 Jun. 1988. ZMA Por. 06668, Mauritania, SW of Cap Timiris, depth 500 m, fossil
coral debris, coll. R.W.M. Van Soest & J.J. Vermeulen, Mauritania II Expedition stat. 40/20, 18.85°N
16°8833°W, 3.5 m Agassiz trawl, 10 Jun. 1988.
Fig. 2. Amphilectus utriculus sp. nov. A,C-E. Holotype ZMAPor. 22592. B. Paratypes ZMAPor. 06636.
C. Cross section showing skeletal structure. D. Peripheral skeleton showing protruding spicule tracts
and clusters of microscleres. E. Spicules in light microscopy.
5
European Journal of Taxonomy 18: 1-21 (2012)
Additional specimen
ZMAPor. 06627, Mauritania, SW of Cap Timiris, depth 200 m, muddy sand, coll. R.W.M. Van Soest &
J.J. Vermeulen, Mauritania II Expedition stat. 35/01, 18.75°N 16.7°W, Van Veen grab, 9 Jun. 1988.
Description
The holotype (Fig. 2A) and most other specimens (Fig. 2B) are short-stalked, with main body laterally
flattened, isodiametrical or either tapering inward near the upper end or flaring, with smaller or wider
opening. Main body hollow, like a purse. Colour reddish or greyish brown, both alive and in alcohol.
Surface irregular, shaggy. Consistency very soft, limp, easily damaged. Stalk rounded, but grading into
the main body by widening, surface slightly smoother. Size 6 cm high (holotype), up to 8 cm high
(paratypes), 0.8 cm (holotype), up to 1 cm (paratypes) in widest expansion, proportion of stalk and main
body approximately 1:3.
Skeleton. There is no special ectosomal skeleton, skeletal bundles are simply protruding beyond the
surface. Choanosomal skeleton consists of polyspicular bundles (Fig. 2C), with 3-6 spicules in cross
section, connected regularly by 1-2 spicules at right angles. Near the surface the bundles fan out to form
loose brushes. Microscleres are concentrated lining the bundles and crowding the brushed endings at
the surface (Fig. 2D). In the interior, microscleres are scattered and distinctly less numerous than at the
surface.
Fig. 3. Amphilectus utriculus sp. nov., Holotype ZMAPor. 22592, SEM images of spicules. A. Styles.
A r Detail of the apices. B. Palmate isochelae in side and ventral view.
6
VAN SOEST R.W.M., BEGLINGER E.J. & DE VOOGD N.J., West African Esperiopsidae
Spicules. (Figs 2E, 3) Styles, palmate isochelae.
Styles. (Fig. 3A, A L ) Of the mycalostyle-type, slightly constricted near the rounded end, straight, or
more commonly slightly curved, 330-377.2-414 x 11-72.7-14 pm.
Palmate isochelae. (Fig. 3B) ‘Normal’-shaped, but with the frontal alae appearing somewhat angular in
side-view, with the shaft slightly incurved, all closely similar in size, 52 -59.6-66 pm.
Distribution and ecology
Mauritania, S of Banc d’Arguin (Fig. 1, loc. 3), muddy bottom at 200-500 m depth.
Remarks
Assignment to the genus Amphilectus is based on the possession of a single microsclere type, although
technically speaking the size of the styles exceeds the 400 pm upper size given as an additional character
of Amphilectus. North Atlantic Amphilectus species with pedunculate shape were reviewed from
literature descriptions (see also Table 1). A. columnatus (Topsent, 1890) (as Esperiopsis) from deep-
sea waters around the Azores has much longer styles (750 pm) and much smaller isochelae (16 pm).
A. pedicellatus (Lundbeck, 1905) (as Esperiopsis ) from West Greenland likewise has longer styles (up
to 940 pm) and very small chelae (13-15 pm). Further species are not only different in shape, but also
show spicular differences: A. fucorum (Esper, 1794) (as Spongia) is massively-encrusting to digitate-
ramose and has smaller spicules overall (see also below). A. lobatus (Esper, 1794) (as Spongia ) is lobate
and likewise has smaller spicules, and the isochelae are verging toward an anisochelate condition. Both
A. fucorum and A. lobatus are essentially shallow-water species. A. typichela (Lundbeck, 1905) (as
Esperiopsis) is encrusting and has two distinct size classes of isochelae.
Southward along the African coasts, in Namibian and South African waters, three further species have
been recorded: A. rugosus sensu Uriz (1988) [as Esperiopsis , probably not conspecific with the Chilean
species A. rugosus (Thiele, 1905)] is a whitish massive sponge with much smaller isochelae. A. lesliei
(Uriz, 1988) (as Esperiopsis) is ramose and has strongyles as megascleres, distinctly smaller than the
styles of the new species. A. informis (Stephens, 1915) is encrusting and has much smaller spicules.
ZMA Por. 06627 from Mauritania (stat. 35/01) is a small whitish crust showing the same skeletal
structure and a spicule complement slightly smaller but essentially similar to the above described
specimens: styles 302-329.5-353 x 6-9.7-12 pm, palmate isochelae: 33-43.5-51 pm. For the time being
this is assumed to be an incipient individual of A. utriculus sp. nov., but the colour and the encrusting
habit point to a possible separate species.
Amphilectus strepsichelifer sp. nov.
Fig. 4
Etymology
The name is a combination of strepsis (L.) = twisted, and chelifer (L.) = bearing chelae, reflecting the
twisted condition of the chelae.
Material examined
Holotype
ZMA Por. 07564, Cape Verde Islands, W of Sao Vicente, Canal de Sao Vicente, depth 348-354 m, coll.
R.W.M. Van Soest, CANCAP 7 Expedition stat. 172/03, 16.8833°N 25.1167°W, rectangular dredge, 7
Sep. 1986.
7
European Journal of Taxonomy 18: 1-21 (2012)
Description
Pedunculate sponge (Fig. 4A), with long thin smooth stalk and abruptly attached small ovate main body.
Main body flattened, but solid (not hollow). Upper surface somewhat rectangular, caused by preparation
damage. Surface irregular, shaggy. Colour light brown alive, grey in alcohol. Size of main body 12 x
6 mm, stalk 35 mm long, 1.2 mm thick.
Fig. 4. Amphilectus strepsichelifer sp. nov., Holotype ZMA Por. 07564. A. Habit. B-C. Cross section
of skeleton in light microscopy. B. Peripheral skeleton. C. Detail of peripheral tracts and clustered
microscleres. D-E. SEM images of spicules. D. Style. D r Details of apices. E. Palmate isochelae, in
various stages of torsion.
8
VAN SOEST R.W.M., BEGLINGER E.J. & DE VOOGD N.J., West African Esperiopsidae
Skeleton. Of main body plumose, rather than plumoreticulate, with diffuse spicule bundles directed
at right angles to the surface (Fig. 4B), where they form loose brushes. Connecting spicules few and
arranged irregularly. Chelae in loose groups in a subectosomal layer at the base of the surface brushes
(Fig. 4C). Few chelae in the interior.
Spicules. (Fig. 4D-E) Styles, palmate isochelae.
Styles. (Fig. 4D, D t ) Thin, slightly curved, 396-430.5-462 x 2-4.6-1 pm.
Palmate isochelae. (Fig. 4E) Predominantly with ‘twisted’ shaft, causing the alae of opposite ends to
face different angles, a minority of the chelae appearing ‘normal’ but these are of the same size as the
twisted ones, and upon closer examination appear to be slightly twisted as well, 32-33.7-36 pm.
Distribution and ecology
Known only from the type locality between the islands of Sao Vicente and Santo Antao, Cape Verde
Islands (Fig. 1, loc. 4), hard bottom, at depths below 300 m.
Remarks
The species is assigned to Amphilectus on the same basis as A utriculus sp. nov. The species stands out
among stalked Amphilectus species (see above in the remarks on A. utriculus sp. nov.) by the peculiar
twisted condition of the palmate isochelae. Additionally the thin stalk carrying the main body without a
clear intermediate zone is characteristic and not found in the other North Atlantic Amphilectus species.
Amphilectus cf. fucorum (Esper, 1794)
Fig. 5
For synonymy, see Van Soest & Hajdu 2002b: 657.
Material examined
ZMAPor. 06775, Mauritania, off Banc d’Arguin, depth 100 m, bottom muddy sand, coll. R.W.M. Van
Soest & J.J. Venneulen, Mauritania II Exped. Stat. 072/14, 20.0°N 17.3°W, 2.4 m Agassiz trawl, 13
Jun. 1988. ZMAPor. 06796, Mauritania, off Banc d’Arguin, depth 48-52 m, bottom muddy sand with
some calcareous gravel, coll. R.W.M. Van Soest & J.J. Vermeulen, Mauritania II Exped. Stat. 082/19,
19.9833°N 17.5°W, 3.5 m Agassiz trawl, 14 Jun. 1988. ZMAPor. 06843, Mauritania, offBanc d’Arguin,
depth 95-100 m, bottom muddy sand with shells, coll. R.W.M. Van Soest & J.J. Vermeulen, Mauritania
II Exped. Stat. 130/09, 20.4167°N 17.6667°W, 3.5 m Agassiz trawl, 20 Jun. 1988.
Description
The material consists of several fragments of encrusting to irregularly ramose sponges (Fig. 5A).
Consistency, soft irregular surface, colour brownish alive and beige to whitish in alcohol. Size of
individual fragments 2-3 cm.
Skeleton. Irregularly plumoreticulate (Fig. 5B), with loosely defined spicule bundles connected by
individual spicules, general aspect rather confused. Spicules barely protruding beyond the surface.
Chelae clustered and singly occurring throughout the interior.
Spicules. Styles, palmate isochelae.
Styles. (Fig. 5C, C,) Straight or slightly curved, relatively short and robust, 198-237.6-276 x 9- 11.4-
14 pm.
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European Journal of Taxonomy 18: 1-21 (2012)
Palmate isochelae. (Fig. 5D) Of ‘normal’ shape, but like in A. utriculus sp. nov., the shaft is slightly
incurved. 19-23. <5-27 pm.
Distribution and ecology
Mauritania, off Banc d’Arguin (Fig. 1, loc. 2), on muddy bottom below 50 m. Elsewhere, if identification
is correct, along most of the coasts of Europe, including the Western Mediterranean. This is the
southernmost record of the species if Southern Ocean records (see below) are considered not conspecific.
Remarks
By assigning these specimens to A. fucorum , the range of this species, which was already huge, is
further extended along the East Atlantic coasts. The species is common and distinctly orange-coloured
in shallow water habitats of the British Isles and the W coast of France, but according to Van Soest et al.
Fig. 5. Amphilectus fucorum (Esper, 1794), ZMAPor. 06775. A. Habit (fragments). B. Cross section of
skeletal reticulation. C-D. SEM images of spicules. C. Style. C r Details of apices. D. Palmate isochela.
10
VAN SOEST R.W.M., BEGLINGER E.J. & DE VOOGD N.J., West African Esperiopsidae
(2000) deep-water specimens may loose their colour, and such specimens may be found down to 100 m.
Skeleton and spicule characteristics of the present material fall within the recorded variation, although
usually styles elsewhere are thinner than those of the Mauritanian specimens. Genetic comparisons may
show diversity over the range of this species and such studies are needed to decide the specific identity
of the Mauritanian populations.
From the other Amphilectus species from the area described above, A. cf. fucorum differs in habit and
chelae (A. utriculus sp. nov. has chelae twice the size, A. strepsichelifer sp. nov. has chelae with a twisted
shaft).
Amphilectus informis (Stephens, 1915) from the Atlantic coast of South Africa appears to be similar
based on published data. According to the original description its chelae have an incurved shaft but
apparently the frontal alae of the chelae are characteristic showing a ‘tubercle in front view’. However,
the illustration of this feature is indistinct. Samaai & Gibbons (2005) described the species also, but their
illustrations do not clarify these alleged differences.
Burton (1932, 1940) recorded A. fucorum from Tristan da Cunha, the Falkland Islands, South Georgia,
and off the coast of Argentina. Thiele (1905) recorded it from Chile. Goodwin et al. (2011) described
two new Amphilectus species from the Falkland Islands, which appear to cover the Burton and Thiele
records. Bergquist & Fromont (1988) recorded the European species Esperiopsis normani (Bowerbank,
1866) and Esperiopsis edwardii (Bowerbank, 1866) from New Zealand waters, but both are now
considered junior synonyms of Amphilectus fucorum. This is not to say that A. fucorum occurs in New
Zealand, but merely that these records should be compared critically.
Genus Esperiopsis Carter, 1882
Remarks
Currently, there are 33 species assigned to this genus (Van Soest et al. 2012). Possibly, some of the
species may need to be transferred to other genera because of the interpretation of the morphological
structure of the chelae (e.g. E.flava Levi, 1993 and similar species with deformed chelae, which cannot
be identified with certainty as palmate), others would have to be returned to Amphilectus if the style size
difference would be no longer a valid discriminatory character of the two genera. Future studies using
additional datasets (such as molecular sequences) are needed to clarify the status of Esperiopsis.
Esperiopsis cimensis sp. nov.
Fig. 6
Etymology
Named after the type locality, Ilheu de Cima.
Material examined
Holotype
ZMAPor. 07282, Cape Verde Islands, SW of Ilheu Rombos, SE of Ilheu de Cima, depth 165 m, hard
bottom with yellow calcareous sand, coll. R.W.M. Van Soest, CANCAP 7 Expedition stat. 030/05,
14.95°N 24.65°W, Van Veen grab, 23 Aug. 1986.
Description
Thinly encrusting on a large volcanic stone (Fig. 6A, arrow), alongside a specimen of Aplysilla Schulze,
1878. Size of crust 3.5 x 2 cm, thickness 1-2 mm. Colour: yellow alive, beige in alcohol. Surface
optically smooth. Consistency soft.
11
European Journal of Taxonomy 18: 1-21 (2012)
Fig. 6. Esperiopsis cimensis sp. nov., Holotype ZMA Por. 07282. A. Habit encrusting a volcanic rock
(indicated by arrow). B. Cross section of peripheral skeleton showing plumose spicule brushes. C-E. SEM
and light microscopy images of spicules. D. Style. D r Detail of apices. C. Sigmas. C r Large sigma. C 2 .
Small sigma. E. Palmate isochelae. E r Left: large chela in dorsal view; right: large chela in side view.
E 2 . Small chela.
12
VAN SOEST R.W.M., BEGLINGER E.J. & DE VOOGD N.J., West African Esperiopsidae
Skeleton. (Fig. 6B) Plumose, with strongly developed spicule bundles traversing the sponge at right
angles to the surface, where they fan out. Few if any connecting spicules. No special ectosomal skeleton.
There is a ‘groundmass’ of microscleres crowding the space between the spicule bundles and adhering to
it, mostly consisting of sigmas and small isochelae. Scattered larger chelae, rather rare, mostly occurring
at the surface.
Spicules. (Fig. 6C-E) Styles, sigmas, palmate isochelae.
Styles. (Fig. 6C, Cj) Long and thin, straight or slightly curved, 396-431.4-461 x 5-6.4-8 pm.
Sigmas. (Fig. 6D) In two distinct non-overlapping size categories, both thin and strongly curved: (I) 76-
85.8-96 pm (Fig. 6Dfl and (II) 3 6-45.4-56 pm (Fig. 6DJ.
Palmate isochelae. (Fig. 6E) In two distinct, non-overlapping size categories, the larger (I) ‘normal’ in
shape (Fig. 6E X ), 56-69.4-84 pm, the smaller (II) with incurved median alae (Fig. 6E 0 ), 14-76.9-21 pm .
Distribution and ecology
Only known from the type locality S of Ilheu de Cima, Cape Verde Islands (Fig. 1, loc. 5), on volcanic
bottom with yellow sand, at 165 m depth.
Remarks
The new species appears to be a member of a group of morphologically similar North Atlantic deep-water
species (see Table 1), sharing the encrusting habit, the size and shape of megascleres and two categories
of both sigmas and chelae. Probably closest is Esperiopsisflagellum Lundbeclc, 1905 from off SE Iceland.
Clear differences exist in the shape of one of the sigma categories, as these of E. flagellum are ‘flagellate’
(excessively incurved). The sigmas are also larger in size, up to 250 pm in E. flagellum, whereas they
are up to 400 pm in the similar species E. macrosigma Stephens, 1916 from waters west of Ireland. E.
praedita Topsent, 1890 from the Azores is also close, but the larger chelae category is smaller than in
our new species, and the sigmas can be as large as 200 pm. A less similar sponge is E. decora Topsent,
1904 from the Azores, sharing the overall spiculation, but having even larger (flagellate) sigmas, and in
addition possessing trichodragmas and three instead of two size categories of chelae. Other Esperiopsis
species described from the North Atlantic appear more distant: E. incognita Stephens, 1916 from Irish
waters has smaller chelae and larger sigmas. E. strongylata (Alander, 1942) from the Skagerrak and E.
strongylophora Vacelet, 1969 from the Western Mediterranean have strongylote megascleres instead
of styles. The two remaining, more elaborately shaped, North East Atlantic Esperiopsis species also
show differences in spicule sizes and categories: variably massive or ramose E. polymorpha Topsent,
1890 from the Azores has smaller chelae in a single category and smaller sigmas. Massive-erect or leaf¬
shaped E. villosa (Carter, 1874) has the size of the largest sigmas and chelae clearly in excess of those
of our new species.
Esperiopsis schmidti Arnesen, 1903 from Norwegian waters and Esperiopsis glomeris Topsent, 1904
have been assigned to the myxilline genus Echinostylinos Topsent, 1927 by Van Soest & Hajdu (2002a)
on account of their possession of tridentate isochelae. Esperiopsis typichela Lundbeck, 1905 and
Esperiopsispedicellata Lundbeck, 1905 have been transferred to Amphilectus by Van Soest et al. (2012)
as they do not possess sigmas. Esperiopsis lesliei Uriz, 1988 and Esperiopsis rugosus sensu Uriz (1988)
both from Namibia, likewise do not have sigmas and belong to Amphilectus.
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European Journal of Taxonomy 18: 1-21 (2012)
Genus Ulosa de Laubenfels, 1936
Remarks
The genus Ulosa is not well-established, with approximately 15 disjunctly occurring species (South
Australia, North East Atlantic and Mediterranean, Indonesia, Central Pacific, and Brazil, see Van
Soest et al. 2012). About half the number of species are only known from old dried types and/or are
inadequately characterized. With the simple morphological features the genus remains to be established
as a monophyletic group, and indeed as a member of the suborder Poecilosclerida, as the implied
assumption of secondarily lost chelae lacks firm evidence. In fact, a recent study of 28S gene sequences
of a large group of demosponges including poecilosclerid, hadromerid and halichondrid sponges
(Morrow et al. 2012) indicated that at least one species of the genus Ulosa could be a member of the
family Halichondriidae Gray, 1867. Here we will retain the genus as a member of Esperiopsidae until
sequences of the type species of Ulosa will be obtained.
Ulosa stuposa (Esper, 1794)
Fig. 7
For synonymy, see Van Soest 1987.
Material examined
ZMAPor. 06567, Mauritania, off Cap Blanc, depth 15 m, bottom muddy, coll. R.W.M. Van Soest & J.J.
Vermeulen, Mauritania II Exped. Stat. 006/01, 20.8167°N 17.0167°W, 1.2 m Agassiz trawl, 7 Jun. 1988.
ZMAPor. 06705, Mauritania, SW of Cap Timiris, depth 12-18 m, bottom sand overlying limestone ridge,
coll. R.W.M. Van Soest & J.J. Vermeulen, Mauritania II Exped. Stat. 049/18, 19.0833°N 16.4167°W,
rectangular dredge, 11 Jun. 1988. ZMAPor. 06714, Mauritania, SW of Cap Timiris, bottom muddy sand,
depth 30 m, coll. R.W.M. Van Soest & J.J. Vermeulen, Mauritania II Exped. Stat. 053/05, 19.0833°N
16.4667°W, 2.4 m Agassiz trawl, 11 Jun. 1988. ZMAPor. 06766, Mauritania, offBanc d’Arguin, bottom
muddy sand, depth 48-52 m, coll. R.W.M. Van Soest & J.J. Vermeulen, Mauritania II Exped. Stat.
072/06, 20.0°N 17.4°W, 3.5 m Agassiz trawl, 13 Jun. 1988. ZMAPor. 06859, Mauritania, offBanc
d’Arguin, bottom muddy sand, depth 50 m, coll. R.W.M. Van Soest & J.J. Vermeulen, Mauritania II
Exped. Stat. 137/01, 20.7167°N 17.35°W, 2.4 m Agassiz trawl, 21 Jun. 1988.
Examined for comparison
ZMA Por. 09932, Portugal, Algarve, E of Sagres, 37.0°N 8.9333°W, growing on Cystoseira alga,
coll. J.H. Stock, 14 Aug. 1976. ZMAPor. 16490, France, Bretagne, Roscoff, Le Taureau, depth 25 m,
48.6633°N 3.8833°W, coll. M.J. de Kluijver, Jul. 1992.
Ulosa digitata sensu Burton 1956, slide made from a specimen in the collections of the Zoological
Museum of Copenhagen collected by the Atlantide Expedition Stat. 147, 09.4333°N 14.9667°W, depth
45 m, bottom shells and forams, 14 Apr. 1946.
Description
Long branches (Fig. 7A) issuing from a basal mass (presumably buried in the sand), dividing sparingly,
surface irregularly conulose. Length of branches up to 10 cm, diameter 0.5-1.5 cm. Consistency soft,
easily damaged.
Colour. Variously reported as yellow, light brown, brown, blue-green or greyish.
14
VAN SOEST R.W.M., BEGLINGER E.J. & DE VOOGD N.J., West African Esperiopsidae
Skeleton. (Fig. 7B) A rectangular reticulation of thick spicule tracts, cemented by only little spongin,
which is usually not visible. Tracts divisible in slightly thicker longitudinals of 35-100 pm thickness
with a core of up to 15 spicules, and thinner interconnecting tracts of 30-60 pm thickness with a core
of 2-10 spicules. Meshes formed by the tracts of widely divergent sizes 200-750 pm. Loose spicules
are observed among the tracts, but these are absent or rare in many specimens, which appear to be
‘macerated’, possibly due to loss of tissue during alcohol preservation.
Spicules. (Fig. 7C, C t ) Styles only (no oxeote modifications were observed), sharply pointed, tending
to be thicker in the middle than at the rounded end (‘fusiform’), occasionally faintly ‘centrotylote’, of
rather variable size in the various specimens, but with limited variation within a specimen, 123-161.2-
192 x 4-7.6-11 pm.
Fig. 7. Ulosa stuposa (Esper, 1794), ZMAPor. 06766. A. Fragmented habit. B. Cross section of skeleton.
C. SEM image of style. C r Detail of apices.
15
European Journal of Taxonomy 18: 1-21 (2012)
Distribution and ecology
Mauritania, Cap Blanc, Banc d’Arguin and Cap Timiris, in sandy substrate at 12-52 m (Fig. 1, Iocs. 1-3).
Elsewhere, widespread from the British Isles southward, Mediterranean, Lusitanian region, Atlantic
islands; southernmost record Gulf of Guinea (see below).
Remarks
The West African material shows a range of growth forms including massively encrusting and branching-
erect, in accordance with the variability elsewhere. Colours in Mauritanian specimens appear duller
(mostly brownish) than in Western European material (cf. Van Soest et al. 2000) and spicule thickness
on average appears greater than in northern specimens: in two specimens from Western France (ZMA
Por. 16490) and Portugal (ZMAPor. 09932), examined for comparison, spicules measured 140-165 x
2-6 pm. Nevertheless, there is overlap in these characters and conspecificity seems likely.
Burton (1956) reported this species (as U. digitata ) from Conakry, Guinea (9.4667°N 14.9667°W). It
was described as forming a bush with branches 5-6 cm long and 0.5 cm thick. The skeleton is similar
to Western European specimens and spicules conform as well (130-145 x 3-5 pm). Apparently, the
distribution of U. stuposa extends over much of the coastal waters of the Northeast Atlantic.
Alittoral species from Faial, Azores, Ulosajullieni (Topsent, 1892) (1892: 137, pi. I, fig. 12, as Stylinos ),
has similar skeleton with slightly smaller spicules (120-130 pm), but differs from U. stuposa primarily
in growth form and surface (thickly encrusting with smooth surface and slightly elevated oscules). It is
also recorded here from the piles of the pier of Villa Baleira, Porto Santo, 33.0562°N 16.334°W (ZMA
Por. 21503, coll. R.W.M. Van Soest, 30 June 1990).
Ulosa capblancensis sp. nov.
Fig. 8
Etymology
Named after the type locality, Cap Blanc.
Material examined
Holotype
ZMAPor. 09959, Mauritania, off Cap Blanc, depth 11-35 m, coll. F.P Vermeulen, 20.7°N 17.1667°W,
1906.
Description
Erect branches issue from a basal mass (Fig. 8A), which was presumable buried in the sandy substrate.
Individual branches 8 mm diameter at the base tapering to blunt endings of 2 to 4 mm diameter. Individual
branches up to 10 cm long. Basal mass (Fig. 8A ; ) 3-6 cm thick, 4-5 cm in lateral expansion. Surface of
basal mass and branches roughly conulose and porose, consistency rather firm, but soft to the touch. No
apparent oscules. Possibly the specimen is somewhat macerated.
Colour. Brownish red in alcohol, live colour unknown.
Skeleton. (Fig. 8B) Forms a spongin-rich irregular reticulation with longitudinal meshes. Emphasis
of the skeleton is on ascending fibres, which are cored by 2-9 spicules, whereas cross connections are
predominantly cored by single spicules at widely differing angles, occasionally two or more spicules
are in the connecting spongin-fibres but then these are not aligned. Meshes average 500 pm in length.
16
VAN SOEST R.W.M., BEGLINGER E.J. & DE VOOGD N.J., West African Esperiopsidae
distance between longitudinal fibres is 200-400 pm, fibres have a thickness of 100-220 jam depending
not only on the amount of coring but also on spongin development.
Spicules. Robust styles only.
Styles (Fig. 8C, Cj) straight or slightly curved, isodiametrical over much of their length with sharply
pointed endings, 648-725.9-843 x 18-22.7-26 pm.
Fig. 8. Ulosa capblancensis sp. nov., Holotype ZMAPor. 09959. A. Habit. A r Detail of basal mass.
B. Cross section of skeleton. C. SEM image of style. C r Detail of apices.
17
European Journal of Taxonomy 18: 1-21 (2012)
Distribution and ecology
Mauritania, on sandy platform off Cap Blanc, 10-35 m (Fig. 1, loc. 1).
Remarks
The growth form of the new species is similar in most respects to Ulosa stuposa , which was found in
the same habitat. It is likely that colours differ (yellow-orange in typical U. stuposa , possibly red or red-
brown in U. capblancensis sp. nov.). However, the major distinguishing features are (1) the structure of
the skeleton, with weakly developed connecting fibres and thick irregular spongin-encased ascending
spicule tracts, and (2) the relatively long and thick styles, on average twice or thrice as long and thick as
those of U. stuposa (725 vs. 160 pm).
The present species might easily be confused with a local species of Dictyonella Schmidt, 1868 [possibly
identical or close to Dictyonella pelligera (Schmidt, 1864)], which shares possession of erect branches
and long thick styles with U. capblancensis sp. nov. Differences are pointed surface projections, the
plumose arrangement of the spicules, and the characteristic abundance of cells with reffactile granules
(see Riitlzer et al. 2003) of the Dictyonella species.
Discussion
From the present new data from an area of the oceans that is relatively well-studied, it may be concluded
that actual diversity of esperiopsid sponges is probably substantially underestimated. Many more species
from deeper water (shelf, bathyal and abyssal habitats) along the west coasts of Africa and elsewhere
may be expected to occur A problem for further discovery is the simple skeletal structure in these species,
which may prevent an unambiguous description. Cases in point are literature records of Amphilectus
rugosus , a Chilean species recorded from SW Africa, A. fucorum reported from disjunct localities all
over the globe, or potential confusion of Ulosa species with members of halichondrid genera such as
Dictyonella , Scopalina Schmidt, 1862, Stylissa Hallmann, 1914 or Hymeniacidon Bowerbank, 1858.
The distinctness of Amphilectus and Esperiopsis remains problematic, as the character differences given
by Van Soest & Hajdu (2002b) appear artificial, viz. length of styles less than 400 pm and lack of
microscleres other than palmate isochelae ( Amphilectus ), vs. longer styles and additional microscleres,
sigmas, toxas and trichodragmas ( Esperiopsis ). The differences are also eroded by several species
possessing styles with lengths in the neighbourhood of400 pm (see above). Other characters, such as the
precise shape of the microscleres, like the incurved shaft of the chelae in Amphilectus utriculus sp. nov.,
A. fucorum and A. informis, or the architecture of the skeleton, like the more reticulate architecture of
A. fucorum compared to the distinctly plumose structure observed in both Amphilectus strepsichelifer
sp. nov. and Esperiopsis cimensis sp. nov., may need to be taken into account as well to arrive at a more
‘natural’ grouping of species of both genera. Possibly this will result in a rearrangement of species
currently assigned to both genera, but support from independent datasets is needed for this.
The family Esperiopsidae, although maintained as a separate family by the latest overview of the
classification (Hooper & Van Soest 2002), was extensively emended and reduced compared to its
original content (Hentschel 1923). It is founded on the lack of an ectosomal specialization, a basically
plumoreticulate choanosomal skeleton and palmate isochelae, of which the latter may be either absent or
complemented with sigmas. The family Mycalidae Lundbeclc, 1905 is considered most closely related
and many previous authors included current genera of Esperiopsidae as members of a broader concept
of the family Mycalidae. However, most mycalids have a specialized ectosomal structure and the chelae
are anisochelate. Relationships with desmacellid and cladorhizid sponges are also evident, but most
genera of these families appear more distant. Phylogenetic relationships have not yet been seriously
18
VAN SOEST R.W.M., BEGLINGER E.J. & DE VOOGD N.J., West African Esperiopsidae
investigated and until this is addressed we refrain from proposing changes in the family content or the
integrity of the genera assigned to it.
Acknowledgements
Jan Vermeulen (formerly employed by ZMA) assisted on board of RV Tyro with securing and preserving
the sponges. The late Jaap van der Land invited the first author to participate in the CANCAP 7 and
Mauritania II cruises. Captains and crews of HNLMS Tydemann and RV Tyro are thanked for their
efforts.
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Manuscript received on: 1 May 2012
Manuscript accepted on:25 June 2012
Published on: 27 July 2012
Topic editor: Rudy Jocque
In compliance with the ICZN , printed versions of all papers are deposited in the libraries of the institutes
that are members of the EJT consortium: Museum national d’Histoire naturelle, Paris, France; National
Botanic Garden of Belgium, Meise, Belgium; Royal Museum for Central Africa, Tervuren, Belgium;
Natural History Museum, London, United Kingdom; Royal Belgian Institute of Natural Sciences,
Brussels, Belgium; Natural History Museum of Denmark, Copenhagen, Denmark.
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