Bijdragen tot de Dierkunde, 62 (1) 3-19 (1992)
SPB Academie Publishing bv, The Hague
A revision of Atlantic Asteropus Sollas, 1888 (Demospongiae), including
a description of three new species, and with a review of the family
Coppatiidae Topsent, 1898
Eduardo Hajdu & Rob W.M. van Soest
1 Laboratorio de Poriferos, Departamento de Zoologia, Instituto de Biologia, Universidade Federal do
2 Rio de Janeiro, Cep. 21941 Cidade Universitária, Rio de Janeiro, Brasil; Institute of Taxonomie
Zoology, University of Amsterdam, P.O. Box 4766, 1009 AT Amsterdam, The Netherlands
Keywords: taxonomy, Asteropus, Coppatiidae, Demospongiae
Abstract necida. A associação de Asteropus e outros génerosrelacionados
à família Coppatiidae Topsent, 1898 é discutida, levando à con-
clusão de que a família é certamente um taxon polifilético rela- Various records of A. simplex Carter, 1879 from the Atlantic cionado Van a vários grupos de Astrophorida (Hooper, 1986; are assigned to three new species of the sponge genus Asteropus Soest, 1991). viz.: brasil and Sollas, 1888, A. iensis sp. n., A. vasiformissp. n.,
A. A. niger sp. n., whereas simplex s.s. is restricted to the Indo-
Pacific. A worldwide study of Asteropus specimens resulted in
the conclusion that two species groups exist, namely “simplex ”- Introduction
like species (with true sanidasters), and “sarasinorum”- like spe-
cies (with spiny microrhabds), as previously observed by Berg- Examinationof several western Atlantic specimens quist (1965, 1968). A newly discovered microsclere complement 1879 recorded as Asteropus simplex Carter, (cf. of trichodragmata in the first group strengthens the need for
generic distinction of both lineages, and accordingly the name Boury-Esnault, 1973; Van Soest & Stentoft, 1988;
MelophlusThiele, 1899 is reinstated for the “sarasinorum” spe- Muricy et al., 1991), as well as undescribed materi-
cies group. A key to the West Atlantic species of Asteropus is al, revealed the existence of several distinct species provided. The family allocation of Asteropus and associated which all showed differences with A. simplex sensu genera in the Coppatiidae Topsent, 1898 is discussed, with the Carter (1879). It is the of this to conclusion that the family is undoubtedly a polyphyletic assem- purpose paper
related various Van describe and these blage to astrophorid groups (Hooper, 1986; compare new species.
Soest, 1991). revealed As by the present study, many Astero-
share unknown pus specimens a previously comple-
ment of trichodragmata, linking them to the genus
Resumo Holoxea Topsent, 1892 and focussing our attention
on distinction among genera in the family to which
são As citações de Asteropus simplex o Atlântico referidas is attributed.A further para Asteropus usually purpose três a novasespécies de Asteropus , viz.: A. brasiliensis sp. n., A. of this paper is to contribute to the discussion of the vasiformis sp. n. e A. niger sp. n.; A. simplex s.s. fica restrito classification of this group. A brief historical sur- ao Indo-Pacífico. O estudo de espécimens de Asteropus de todo
is to outline the mundo resultou conclusão de dois de vey necessary problem. o na que grupos espécies
estão envolvidos, espécies tipo-“simplex” (com sanidasters ver- Topsent (1898) erected the family Coppatiidae
dadeiras) e espécies tip microrabdes o-“sarasinorum” (com es- (replacing Epipolasidae Solías, 1888, which does
pinadas), como já notado Bergquist (1965, 1968). Um por fulfill for not the requirement family names; the recente descoberto complemento de tricodragmas no primeiro name Epipolasidae was never fixed by application necessidade de ambas li- grupo reforça a distinção genérica de
of a type genus name - Article 35, ICZN), for nhagense assim sendo, o nome Melophlus Thiele , 1899 é recupe-
rado para o grupo de espécies t ipo-“sarasinorum”. Uma chave sponges belonging to rather distinct lineages: Top-
identificação as de para espécies do Atlântico Ocidental é for- sentia Berg, 1899 (as Anisoxya Topsent, 1898) and
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Spongosorites Topsent, 1896 (both Halichondri- ious ancorinid genera possessing triaenes. In the
this various da); Hemiasterella Carter, 1879 and Tethya course of paper genera of Coppatiidae
various order Lamarck, 1814 (as Magog Solías, 1888) (possibly are compared to Ancorinidaein to ar-
1867 rive the allocation of Hadromerida); Jaspis Gray, (as Coppatias at proper family Asteropus.
Solías, 1888) and Asteropus Solías, 1888 (probably
Astrophorida). He stressed the possession of eu-
asters and diactinal megascleres, and absence of tri- Material and methods
aenes, as the shared derived characters for this
Hentschel added the group. Later, (1923) genera The Brazilian Asteropus specimens are deposited in the collec-
Cryptotethya Dendy, 1905 (= Tethya), Diastra tion of Porifera in the Universidade Federal do Rio de Janeiro
(UFRJ POR). They were collected during an ecological survey Row, 1911 (a possible synonym of Stelletta fauna of the sponge of Arraial do Cabo, southeastern Brazil Schmidt, 1862), and Paracordyla Hallmann, 1912 (Muricy, 1989; Muricy et al., 1991, recorded as A. simplex). Scolopes Solías, 1888). This assemblage has not (= Data about specimens quoted in "Material examined" are as
of been accepted as one monophyletic origin by complete as possible.
A ofthe ofA. brasiliensis is modern authors dueto major differences in skeletal fragment holotype sp. n. deposit-
ed in the collections of the Museum Amsterdam well distinct Zoölogisch arrangement, as as spicule types, habit, (ZMA), number POR 9761. Additional material from the collec- and chemical constituents. tions of ZMA, the Muséum National d'Histoire Naturelle, Paris there is of However, a group generacurrently ac- (MNHN), the Natural History Museum, London (BMNH), and
cepted by some authors (e.g. Bergquist, 1968) as the United States National Museum, Washington (USNM) was
studied for comparison. Section and spicule mounts for light comprising the Coppatiidae. They share a spicular
made methods described diactinal in microscopicstudy were according to by complement of megascleres arranged a Riitzler (1978). SEM preparations of spicules were made as confused/radialchoanosomal pattern and asterose described by Buizer & Van Soest (1977). "In toto" SEM prepa-
microscleres. The ectosomal skeleton is a tangential rations were treated with 20% HNOj for 3-12 h, after which
layer of megascleres (as opposed to the tangential they were coated with a thin layer of gold.
microscleres found in certain astrophorid genera
like Penares Gray, 1867 and Erylus Gray, 1867).
Systematic descriptions This group includes the genera Jaspis Gray, 1867,
Asteropus Solías, 1888, Holoxea Topsent, 1892, Genus Asteropus Solías, 1888 Melophlus Thiele, 1899, and Lamellomorpha Berg-
quist, 1968. In this group, Neamphius De Lauben- Type species: Stellettinopsis simplex Carter, 1879, fels, 1953 also seems to fit, although this is uncon- by monotypy (Article 68d, ICZN). firmed due to its possession of amphiasters. The
Coppatiidae are here considered to be close to the Diagnosis. - Ancorinidae without triaenes, with a Astrophorida, based on the observation that tangential ectosomal skeleton of megascleres; with reduced radiality, rarity/absence of triaenes, and oxyasters and sanidasters, to which trichodragmata presence of tangential microxea is also found in may be added. some Astrophorida of the family Ancorinidae. A
similar point of view, though much reduced in
scope, was expressed by Burton & Rao (1932), Asteropus brasiliensis sp. n. Bergquist (1968) and Bergquist & Hartman (1969). (Fig. 1 a-d; Pl. I A-F) The family allocation of Asteropus in the Cop-
patiidae was formally challenged by Dendy (1916a) Asteropussimplex sensu Boury-Esnault, 1973: 273, text-fig. 14;
and, more recently, also by Van Soest & Stentoft Hechtel, 1976: 249 and 252; Muricyetal., 1991: 1187, 1190(non
(1988). Bergquist (pers. comm.) also expressed Stellettinopsissimplex Carter, 1879).
doubts over it. These authors suggest that Astero- Material examined. - Holotype: UFRJ POR 3022, Pedra Ver- and also other pus possibly coppatiid are genera melha (22°59.17'S 41°59.57' W; Arraial do Cabo, southeastern Ancorinidae without simply triaenes, related to var- Brazil), 3 m depth.
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brasiliensis Fig. 1. Asteropus sp. n., spicules: a, oxeas; b, oxy-
Scales are in aster; c, sanidaster; d, trichodragma. µm.
Additional material: UFRJ POR 3371 (fragment from Paratypes: UFRJ POR 2575, Praia dos Anjos, 3 m depth,
Pernambuco State 19' S coll. E. Hajdu, 09/V/1987; UFRJ POR 2814, Praia do Forno, MNHN NBE 1025), coast (08°
and m depth, 1961/62 (Boury-Esnault, 4.5 m depth, 07/III/1988; UFRJ POR 2889, 2908, 2913, 34°39'W), 75 "Calypso"
recorded A. 2915a, all four from Pedra Vermelha, 6.5-7 m depth, 1973; as simplex).
lO/V/1987; UFRJ POR 2974, Pedra Vermelha, 6 m depth, coll.
G. UFRJ POR 3027, Pedra Vermelha, Muricy, 24/VI11/1987; - is Description. The sponge cushion-shaped to 5 3 UFRJ POR 3038, Pedra Vermelha, m depth, coll. m depth; size of 2 subglobose. Specimens attain a 25 cm by G. Muricy, 08/111/1988.
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Plate I. brasiliensis Asteropus sp. n.: A, sanidaster; B, sanidaster; C, oxyaster and sanidasters; D, oxyaster; E, trichodragmata; F, trichodragma. Scales are all 5 µm long.
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4 cm high. They are firm, hardly compressible. fusiform, tapering gradually, slightly curved at
Their colour alive is dark brown where exposed to mid-length.
the light, gradually becoming clearer towards the Microscleres: Sanidasters (Fig. 1 c; Pl. I A-B)
base of attachment of the which is fre- sponge, spined all over. Spines usually larger at end of rays.
The choanosome is white. quently dirty-white. Rays up to 15, showing no tendency for a concen-
There is an easily discernible cortex, about 1 mm tric arrangement. Irregular shape. Some rare
deep, characterized by large amounts of subspheri- reduced forms, sometimes smooth, may be present
cal pigmented cells (18 X 25 /¿m). Both the holotype (perhaps developmental stages, never microrhabd-
and show in Boury-Esnault's specimen spirit a like); length 11.3-17.5 (« = 70). Oxyasters (Fig.
brown choanosome and a whitish cortex. 1 Pl. I all sometimes b; C-D) spined over, with
The surface is minutely hispid, sometimes irregu- smooth centrum. Largest spines at distal third of
due the lar, to presence of embedded sediment. Os- rays. Rays (6-8), normally fusiform. Tapering,
cula, approximately 2 mm wide, are grouped on the gradual or sudden; length 13.8-50.6-42.5 /im
darker of the No at- = apical part sponge. firmly (s 5.8) (« = 43). Trichodragmata (Fig. 1 d; Pl. I
tached epibionts were seen at any of the specimens E-F) irregularly cylindrical (capsule-like), slightly
collected. flattened shape; length 12.6-22.7-28 /im (« = 20).
Skeleton: The cortical skeleton is characterized
- by an upper ectosomal crust of sanidasters, rein- Ecology. Relatively common at the Cabo Frio
forced by huge amounts of small trichodragmata region. The species is normally found under the
2 (up to more than 100 per 1000 /¿m ). This layer is zoanthidPalythoa sp. (which is not attached to the
adhered with usually closely to an intermediary one (ap- sponge), only the darker apical part of the
200 The proximately /im wide), consisting of tangential- sponge apparent. species has a considerable
sometimes difficult to distin- ly arranged oxeas, bathymetrical range (3 to 75 m depth).
guish from the underlying choanosomal skeleton.
These oxeas are in size - the same range as those in Distribution. Tropical southwest Atlantic.
the choanosome, as demonstratedby the unimodal
distribution in the = - The refers size-frequency holotype (n Etymology. name to the distribution
150), and by the absence of any size distinction be- of the species, which is restricted to the Brazilian
tween dissociated spicules from the ectosome and coast.
the choanosome. The inner layer is composed
mainly of radially disposed bundles of hav- Remarks. - Of all western Atlantic oxeas, Asteropus spe-
lower than the A. brasiliensis ing a spicular density choanosome, cies, sp. n. seems closest to A. sim-
and being traversed by abundant aquiferous plex (Carter, 1879) from the Indo West Pacific. A
channels. re-examination of a type slide from Carter's collec-
The choanosome consists of dense skeleton of tion a (BMNH, no registration number) revealed the
radially disposed oxeas. Large amounts of scattered occurrence of two of unexpected categories oxy-
oxeas make of the skeleton and 40—67 In other may parts very con- asters (20—32 ¿im). respects
and fused. Oxyasters are rare seem to show no it conforms precisely to spicular micrometries of A.
localization. Small trichodragmata are abundant, brasiliensis sp. n. Trichodragmata are in the same
scattered throughout the size sponge. range (17-25 /xm), although less abundant in
d Spicules (see Fig. 1 a- and Pl. I A-F; measure- A. simplex.
ments in Table = standard I; s deviation, « = num- Re-examinationof Boury-Esnault's (1973) speci-
ber of spicules measured): men, collected by the "Calypso" Expedition along
Megascleres: Oxeas (Fig. 1 a) variable in size the northeastern Brazilian coast and identified by
distribution with mode 700 her A. (unimodal at ¿im), as simplex, revealed a slight tendency for bi-
322—970.5—1984 width 6.3- length (5=325.8), modality in the size-frequency of the oxeas
= 26.8-60 n= slender or = A similar noted in /im (s 2.1) ( 150); robust, (n 150). pattern was a paratype
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Table I. Individual spicule dimensions for specimens of Asteropus brasiliensis sp. n. Means are in italics; other values are ranges; ex-
specimen was pressed as spicule length/width. The number of oxeasmeasured for both the holotype and Boury-Esnault’s (1973) 150.
100 other = if otherwise stated. Values are in In paratype UFRJ POR 2814, oxeas weremeasured. For measurements, n 20, except µm.
sanidasters oxeas oxyasters trichodragmata
UFRJ POR 3022 422-/099-1984/12.5-55.5-60 12.5-16.3 22.5-29.4-38.8 15.4-22.7-28
Holotype
UFRJ POR 2814 322-75/-1782/6.3-5/. 5-41.3 12.5-17.5 13.8-26.3 (n = 3) 18.2-25.2
Paratype
UFRJ POR 2889 510-990- 1736/15-26-35 11.3-15 not found 16.8-23.8
Paratype
UFRJ POR 2908 645-955-1314/10-54-45 11.3-16.3 30-55.5-38 18.2-22.4
Paratype
UFRJ POR 2913 570-/029-1786/11.3-24-45 12.5-15 25-30.4-35 18.2-22.4
Paratype
UFRJ POR 2915a 446-950-1686/8.8-20.9-37.5 12.5-13.8 23.8-52.Z-42.5 12.6-/6.5-21
Paratype
UFRJ POR 3027 434- 787- 1215/10-/5.9-20 11.3-15 not found 16.8-21
Paratype
Boury-Esnault (1973) 1245-1650/30-56 15-20 28-38 not given
570-1026/9-20
(remeasured) 405-925-1620/7.5-22.5-47.5 12.5-16.3 27.5-55.5-41.3 14-/7.6-19.6
of n. POR A. brasiliensis sp. (UFRJ 2814), trichodragmata (15-28 /xm) (Table II). although not as pronounced as in the previous specimen. As intermediate-sized oxeas are quite Asteropus ketostea (De Laubenfels, 1950) common in both specimens and no shape distinc- tion can be found betweensmaller and larger oxeas, Stellettinopsis ketostea De Laubenfels, 1950: 112-114;text-fig we considered this to be merely a case of in- 50. traspecific variation, and consequently Boury-
Material examined. - Two slides (thick-sections and spicules), Esnault's specimen was assigned to A. brasiliensis from the type specimen from Bermuda (USNM, no reference sp. n. It is considereddistinct from A. simplex (sen- number). De Laubenfels (1950) mentions a syntype in the su on the basis of of one Carter) possession only BMNH collection (reg. no. 1948:8:6:47), but this could not be category of oxyasters. found by Miss S.M. Stone and is assumed to be lost.
The between brasiliensis and similarity A. sp. n.
A. ketostea (De Laubenfels, 1950) (see below, Ta- Description (emended from De Laubenfels, 1950).
ble II), as revealed by the tendency towards bimo- - Shape encrusting to massive (1 cm thick), di- dality, is strongly contrasted by the smaller oxy- ameter of 2 cm. Consistency firm, hard, slightly asters and absence of trichodragmata in the latter compressible. Surface minutely verrucose. Pores species. abundant over theentire surface, each about 60 /¿m
Soest & of Asteropus syringiferus Van Stentoft, in diameter. Oscules not seen (perhaps some the
1988 (see below) is also close to the present species, pores may be exhalant apertures). Colour black ex- differing in its peculiar habit (with fistules), slightly ternally and grey in the interior. In alcohol it fades
slender larger oxyasters and more sanidasters. slightly.
Asteropus brasiliensis sp. n. is distinguished Skeleton: The ectosomal skeleton is composed of from other western Atlantic records of Asteropus large amounts of sanidasters interspersed with a
the by possession of only one variable category of skeleton of paratangential oxeas. A concentration oxeas small cells that extend (300-2000 /¿m), and rare oxyasters of pigmented (24 x 11 may
and and abundant 160-320 into the makes the ectosome (14-43 /xm) relatively large n m sponge,
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Table II. Comparative data on spicule dimensions for all western Atlantic Asteropus species. Means are in italics; other values are ranges;
expressed as spicule length/width. The number of oxeasmeasured was 150 in A. brasiliensis sp. n. and 100 in all the others. For other
spicules, n = 20. Values are in µm.
oxeas 1 oxeas 2 sanidasters oxyasters trichodragmata
Asteropus brasiliensis sp. n. 322-9//-1984/ absent 11.3-17.5 13.8-50.6-42.5 12.6-22.7-28
6.3-27-60
Asteropus ketostea 216-426-551/ 616-975-1404/ 11-14.7 17-27 absent
(De Laubenfels, 1950) 1.5-3.5 6.4-25
Asteropus niger sp. n. 182-2/5-235/ 1045-/966-2503/ 17-20 42-J7.6-98 12.6-19.6
2.2-5.6 8-25-34
Asteropus syringiferus 520-/0/0-1900/ absent 11.2-19.6 34-60 18.2-25.2
Van Soest & Stentoft, 1988 6.5-19.1-53
Asteropus vasiformis sp. n. 632-//6/-1650/ 1750-2225-2805/ 7.5-22 45-65-101 18.2-23.8
19.6-39.2 51.8-86.8
Atlantic in much darker than the rest ofthe sponge in mounted all other Asteropus having two
sections. The pigmented cells are also found 1 mm categories of oxeas, very small oxyasters, and lack-
deep in the sections, though here they are more ing trichodragmata.
sparse. Trichodragmata were not found. Re-examination of the type preparations of A.
The choanosomal skeleton has some loose ketostea revealed the presence of two categories of
ascending radial tracts of large oxeas. Many oxeas oxeas (Table II), instead of only a single category as
in the are scattered in confusion. The subectosomal reported original description (De Laubenfels,
region shows a reduced spicule content. The thinner 1950). Desqueyroux-Faúndez (1990) reported A.
to restricted to ketostea from Easter Island also with two oxeas appear be the choanosome, (Pacific),
where they are strewn at random; they are absent categories of oxeas. Nevertheless, we consider this
from both ectosomal and subectosomal regions. record to be highly dubious, because the sanidasters
her Oxyasters are restricted to the choanosome. reported by are almost three times as large (38
Table for in the her Spicules (see II measurements): ¿un) as type specimen (11-15 /xm); speci-
Megascleres: Oxeas 1 slender, fusiform, tapering men possibly represents a separate species. gradually; straight, slightly curved or sinuous; The closest western Atlantic relative of A.
width 1.5-3.5 length 216-426-551 (s=88), ketostea seems to be A. brasiliensis sp. n., which
= Oxeas 2 differs the of (« 40). fusiform, tapering gradually, by possession only a single category
616-975-1404 = of the abun- slightly curved; length (5 205), oxeas, slightly larger oxyasters, and width 6.4-25 /¿m (« = 80). dance of trichodragmata.
Microscleres: Sanidasters of variable shape; rays Asteropus kaena (De Laubenfels, 1957), from
distributed studied the basis of up to 15, bifurcating or not, irregularly Hawaii, on a holotype fragment over the shaft length; length 11-14.7 ¿im. Oxy- (USNM 23540), is similar to A. ketostea in the ab- asters slender, rays 6-10, completely microspined, sence of trichodragmata. The huge oxeas with very small centrum; length 17-27 /¿m. (2000-2400 ¿im long) reported by De Laubenfels in
the original description were not found, thus
strengthening its similarity to A. ketostea. Distribution. - Collected at 1 m depth, under a Nevertheless, both species are considered to be rock. Known only from Whalebone Bay, north valid on the basis that oxeas smaller than 800 /¿m shore of Bermuda, St. George's Parish. are rare in A. kaena, whereas oxeas only 400 /xm
long are abundant in A. ketostea. A. kaena has also
Remarks. - The species can be distinguished from slightly smaller oxyasters (16-20 /¿m).
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Plate II. A-D, Asteropus niger sp. n.: A, sanidaster; B, sanidaster; C, oxyaster; D, oxyaster. Scales are 5 µm long. E-G, Asteropus
n.: Scales 5 vasiformis sp. E, sanidaster; F, sanidaster; G, oxyaster. are µm long.
brown, almostblack. The surface is and Asteropus niger sp. n. deep hispid
2 Pl. No (Fig. a-e; II A-D) completely overgrown by haplosclerid sponges.
oscules are apparent.
Material examined. - Holotype: ZMA POR 5715, Blauwbaai, Skeleton: The ectosomal skeleton is character-
W 55 M. Curaçao (68°59' 12°08'N), m depth, coll. W. Hoppe & of ized by large numbers of scattered oxeas, some Reichert, 20/X/1984. them tangential to the surface. This arrangement of
the spicules can extend more than 5 mm deep into
Description. - One fragment, 2 cm long, 1.2 cm the sponge. The oxeas may protrude beyond the wide, 2 cm high. Consistency tough, incompressi- surface making it hispid. Masses of pigmented cells ble. Colour alive black. In ethanol the specimen is (14 X 11 íím) are visible near the surface, becoming
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Fig. 2. Asteropus niger sp. n., spicules: a, oxea 2; b, oxea 1; c,
Scales in oxyaster; d, sanidaster; e, trichodragma. are µm.
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below Some of the size - and less abundant 1 mm deep. pro- huge large oxyasters, deep-water
habitat. Both considered to be distinct truding spicules core spongin fibres of an epibiont, species are
the basis of the much smaller size of the smaller Callyspongia strongylata Hartman, 1955. on
less of in the its The choanosome has a more or radially category oxeas latter, densely rayed
with loose and its to oriented skeleton of oxeas, some very sanidasters, massive shape (as opposed
in tracts. Spicule density is high. Colour is darker in cup shape A. vasiformis sp. n.).
the choanosome (deep reddish brown after mount-
ing) than in the ectosome (dirty yellow to light
and trichodragmata are scarce brown). Oxyasters Asteropus syringiferus Van Soest & Stentoft, 1988
and seem to be restricted to the choanosome.
Sanidasters are concentrated near the surface, Asteropus syringiferus Van Soest& Stentoft, 1988: 32-34; text- though less abundant than in other species. fig. 14, Pl. V figs. 1-2.
Spicules (see Fig. 2 a-e and Pl. II A-D; mea-
surements in Table II): Material examined. - Holotype: ZMA POR 5307, off
off Megascleres: Oxeas 1 (Fig. 2 b) slender, almost Holetown, Barbados, 100 m. Paratype: ZMA POR 5308,
Holetown, Barbados, 100 m. raphidiform, slightly curved; length 182-275-235,
2 2 width 2.2-5.6 um (« = 20). Oxeas (Fig. a) slender and slightly curved, fusiform, tapering Description (emended from Van Soest & Stentoft,
- is round 2.5 in gradually; length 1045-1966-2503 (5=314.9), 1988). The holotype a mass, cm
= incrusted with dead shells, two width 8-25-34 /¿m (« 100). diameter, bearing
Microscleres: sanidasters (Fig. 2 d; PI. II A-B), conspicuous hollow fistules; the larger fistule, 2 cm
in is closed at the spines concentrated on rays, largest spines at end of in length, 4 mm diameter, apex;
the shorter one bears an apical oscule. The rays, up to 30 rays in one spicule; length 17-20 /¿m paratype
= in diameter. Both (« 20). Oxyasters (Fig. 2 c; PI. II C-D) extremely is a smaller sphere, 1 cm speci-
is hard, slender, rare, spined all over, but spines becoming mens have a rough surface; the consistency
is larger and more abundant closer to end of rays; almost incompressible. The colour in spirit grey
= the fistules. rays 6-10; length 42-57.6-98 /im (« 20). in the main body and dirty-white at
Skeleton: The ectosomal skeleton is of Trichodragmata (Fig. 2 e) of variableshape, some- composed what capsule-like; length 12.6-19.6 /im (« = 20). paratangentially oriented oxeas carrying numerous
sanidasters. The choanosomal skeleton is (para-)
- and Distribution. Collected on the reef slope at 55 m tangential to the surface in the fistule walls, depth, at Blauwbaai, Curaçao. confusedly radiate in the main body. Trichodrag-
mata (not mentioned by Van Soest & Stentoft,
- refers the black live abundant the Etymology. The name to 1988) are throughout sponge. colour reported for the specimen, but more espe- Spicules (see Table II for measurements): cially to the dark, almost black choanosome, a fea- Megascleres: Oxeas fusiform, tapering gradual- ture not usual in Asteropus. ly, slightly bent; length 520-1010-1900 (5 = 244),
width 6.5-/9.7-53 m («=100).
— of variable Remarks. Asteropus niger sp. n. is distinguished Microscleres: Sanidasters shape,
from Atlantic of or other species Asteropus by the shaft mostly thin; rays up to 15, bifurcating not,
distinct shaft possession of two categories of oxeas, one irregularly distributed over the length; small
and 11.2-19.6 very small (180-230 one very large spines concentrated on the rays; length
Its abun- = with small (1000-2500 urn). sanidasters are more /¿m (AI 20). Oxyasters slender, a very
of dantly rayed than in other Asteropus species. centrum; spines concentrated on distal half rays;
Asteropus vasiformis sp. n. (see below; Table II) rays 8-12; length 34-60 /¿m (« = 20). Trichodrag- is closest to A. niger sp. n. on account of its two mata variable in shape, slightly capsule-like; length
of - the 18.2-25.2 = categories oxeas larger one attaining a p m (« 20).
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— substrate 100 = Oxeas 2 3 slender Distribution. Rare, on shelly at m fim (« 45). (Fig. a) fusiform, or
depth, off Barbados. robust, tapering gradually; slightly curved; length
1750-2225-2805 (5=229), width 51.8-86.8 /xm
Remarks. — sizes of this The spicule species are (« = 55).
close to thoseof A. brasiliensis sp. n. However, the Microscleres: sanidasters (Fig. 3 d; Pl. II E-F)
remarkably differenthabit and habitat, the irregu- spined all over; largest spines at end of rays; rays up
lar shape of the trichodragmata, and the spines con- to 14; length 7.5-22 /xm (« = 20). Oxyasters (Fig. 3
fined to the distal half of the rays of the oxyasters c; Pl. II G) large; extremely slender and abundant;
here considered evidence for differentiat- of are strong spined all over, largest spines at end rays; rays
these ing species. 6-10; length 45-63-101 /xm (« = 20). Trichodrag-
mata (Fig. 3 e) irregularly cylindrical (capsule-like),
slightly flattened shape; length 18.2-23.8 /xm
Asteropus vasiformis sp. n. (« = 20).
(Fig. 3 a-e; Pl. II E-G)
Distribution. - Apparently quite rare at 153 m
off Barbados. Asteropus simplex sensu Van Soest & Stentoft, 1988: 31-32, depth,
text-fig. 13, PI. IV-fig. 6 (non Stellettinopsis simplex Carter,
- The refers to the 1879). Etymology. name cup shape.
Material examined. - ZMA POR off Holotype: 5302, Payne's Remarks. - Important differences between A.
Bay, Barbados (13°10'N 59°40'W), 153 m depth;coll. N. Sten- vasiformis sp. n. and A. simplex (examined from a toft, 1978/80. of type slide) are the occurrence two oxyaster
categories (20-32 and 40-67 /xm) and only one Description (emended from Van Soest & Stentoft, category of oxeas (780-1500 /xm long) in the latter. 1988). - Cup-shaped, with one side not quite Asteropus vasiformis sp. n. can be distinguished closed; 10 cm high, 7 cm in diameter at the top, with from other Atlantic species of Asteropus by its two steep walls. Six oscules, one facing four others on categories of oxeas, both with means above 1000 sides opposing inner of the break in the cup wall, urn, large oxyasters (up to 100 abundance of one outside the cup wall. Rim and inner surface microscleres, its cup shape, and deep-water habitat. smooth, outer side hispid. Consistency tough, in- Its closest relative seems to be A. niger sp. n., which compressible, crumbly. Colour in the dry state is differs in much having a smaller category of small light brown at the rim and inner side, and dark pur- oxeas, massive habit, and less abundantly rayed ple at the outer side but this is probably not the sanidasters. proper live colouration.
Skeleton: The ectosomal skeleton is made up of a loose layer of tangential oxeas carrying abundant Key to the western Atlantic Asteropus sanidasters and some trichodragmata. These oxeas are smaller than the choanosomal as 1. ones, demon- Oxyasters 50-100 jim 2
- strated by a bimodal size frequency distribution. Oxyasters < 50 /JL m 3
2. Two distinct of 500 categories oxeas (one < um long), The choanosomalskeleton has a radial architecture
sanidasters with up to 30 rays, shape massive. A. niger■ sp. n. of huge oxeas which often protrude beyond the sur-
- Two categories of oxeas with meanlengths both above 1000 face. become abundant 3 mm Oxyasters abruptly sanidasters with 15 urn, at most rays, cup-shaped beneath the surface. A. vasiformis sp. n.
3. Trichodragmata abundant Spicules (see Fig. 3 a-e and Pl. II E-G; mea- 4
- No A. ketostea trichodragmata .... (De Laubenfels, 1950) surements in Table II): 4. Shape massive-sprawling, oxyasters 14-43 (im Megascleres: Oxeas 1 (Fig. 3 b) fusiform, slender A. brasiliensis sp. n. or robust, tapering gradually; slightly curved; - Shape globular, with fistules, oxyasters 34-60 /¿m length 612-1161-1650 (s= 258), width 19.6-39.2 A. syringiferus Van Soest & Stentoft, 1988
Downloaded from Brill.com10/04/2021 04:39:43PM via free access 14 E. Hajdu & R. W.M. van Soest - Revision of Atlantic Asteropus Sollas, 1888
Fig. 3. Asteropus vasiformis sp. n., spicules: a, oxea 2; b, oxea
in 1; c, oxyaster; d, sanidaster; e, trichodragma. Scales are µm.
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Discussion (1909), Dendy & Frederick's (1924), Wilson's
(1925), Lévi's (1967), Bergquist's (1968, 1969),
Vacelet al.'s Indo-Pacific records of Asteropus Tanita's (1970), et (1976),
Desqueyroux-Faúndez's (1990), and Tanita &
A study of Carter's type slide of A. simplex Hoshino's (1990) specimens is needed in order to
revealed the ascertain what the real in A. is. (BMNH, no reference number) occur- variability simplex
of which renders It is that this will reveal the occurrence of rence of two categories oxyasters, predicted the specimen unique among other records of the several discrete Indo-Pacific species. species. No Atlantic Asteropus is known to have
which is consid- two categories of oxyasters, here
evidence restrict the of the ered as strong to use Phylogenetic affinities of Asteropus
name A. simplex to Indo-Pacific records.
that of Data from the literature on Asteropus are barely Bergquist (1965) observed two groups spe-
informative authors cies couldbe within The first as most give only representa- recognized Asteropus.
and/or had sanidasters and included A. tive measurements, without ranges means. one true simplex,
found in both the A. and A. kaena The larger oxeas presently type ketostea, (the “simplex” group).
add A. specimen and in Hong Kong specimens (the latter To this assemblage we can A. syringiferus,
brasiliensis A. and A. reported by Van Soest, 1981) were in the sp. n., niger sp. n., vasifor-
which is much smaller than mis n. The second was characterized by 1400-1600 /xm range, sp. group
Dendy's (1924) 2100 /xm, Wilson's (1925) 3100 /xm having spiny microrhabds and included A.
and A. cherbonnieri and Bergquist's (1968) 2900 /xm. sarasinorum (Thiele, 1899)
" " The live colour of A. simplex has been reported (Levi, 1961) (the sarasinorum group). Re-
in the literature only by Bergquist (1968: light yel- examination of Lévi's (1965) Stellettinopsis solida,
low; 1969: white), and by Desqueyroux-Faúndez also associated with the latter group, revealed that
(1990: greyish to black). The latter record of A. triaenes were relatively common and a proper tan-
ectosomal skeleton that the simplex is considered highly dubious on account of gential was absent, so
in is here to Ecionemia the possession of much smaller oxyasters (13 /¿m; species assigned Bowerbank,
Hawaii 1862. the same size range as in A. kaena from the
that the difference Islands) and a second, also much smaller category Bergquist (1965) considered
between the above-mentioned of oxeas (130 /xm). Many specimens were reported two Asteropus spe-
cies was similar to that found between An- as being dark-coloured in spirit (e.g. Dendy, 1916a; groups
Lévi, 1967) but it is not possible to ascertain corina (with sanidasters) and Ecionemia (with spiny
whether or not this is the live colour of the microrhabds). This had also previously been point-
specimen. ed out by Solías (1888) and Topsent (1928). As both
studied, considered Remarkably, all the Asteropus species genus groups were synonyms by most
decided to the except for A. ketostea and A. kaena, possess more authors, Bergquist (1965) keep
These under the or less abundant trichodragmata. structures Asteropus lineages same generic as-
have not previously been reported in the genus, and semblage.
will The of abundant small this discovery has a bearing on systematics, as present discovery
of in of be established below. This recent discovery trichodragmata many species Asteropus as-
" " but in trichodragmata widespread in Asteropus speci- signed to the simplex group, not A.
well of in sarasinorum ZMA POR 8049, from Indone- mens, as as a second category oxyasters (e.g.
shows how identified the second is here consid- the type specimen of A. simplex, easily sia, by author)
spicule categories have probably been overlooked ered strong evidence for generic distinctionbetween
in previous descriptions of Asteropus. It is clear both lineages. Lévi (1961) did not mention
Carter's for A. cherbonnieri either. The that a thorough revision of (1879, 1886), trichodragmata
available for Dendy's (1905, 1916a, 1916b, 1924), Hentschel's name Melophlus Thiele (1899) is the
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“sarasinorum” and is considered The second could be if group, presently synapomorphy genuine
valid. A. sarasinorum is considered distinct from the various tangential conditions were homolo-
true Ancorina/Ecionemia on the basis of its posses- gous. However, the condition in Jaspis s.s. - i.e.
of skeleton of sion a very thick tangential oxeas, conforming to that of the type species, J. johnstoni
be defined follows: - and can as (Schmidt, 1862) seems to be widely different
from that found in Asteropus-Melophlus-Holoxea
Melophlus Thiele, 1899 s.S.: Jaspis s.s. has small oxeas forming a tightly ar-
Type species: Melophlus sarasinorum Thiele, 1899, ranged fine-grained surface, leaving characteristic
by monotypy. round pore fields free of spicules, while the other
taxa have an almost paratangential, confused ar-
- Ancorinidae with of Diagnosis. without triaenes, a rangement oxeas of intermediate or large size.
thick ectosomal skeleton of Holoxea tangential megascleres, violacea Boury-Esnault, 1973 has no tan-
and microsclere of this is a complement oxyasters and gential skeleton, but interpreted as a subse-
microrhabds. spiny quent loss.
included: M. sarasinorum 1899 On the other Species Thiele, hand, Asteropus microscleres are
and M. (type species) cherbonnieri (Lévi, 1961). closely similar to those of Stryphnus Gray, 1867, as
already suggested by Solías (1888) and Van Soest &
Stentoft (1988), and the somewhat irregular, con-
allocation of fused/radial Family Asteropus arrangement of choanosomal mega-
scleres is also is comparable. It not difficult to con-
Various authors (Solías, 1888; Topsent, 1900; Berg- sider both genera as closely related, differing in the quist, 1968) have pointed out that generic differ- absence of triaenes and possession of trichodrag-
within the ences Coppatiidae paralleled those of mata in Asteropus. Holoxea Topsent, 1892, may be certain of but this associated with genera Ancorinidae, apparently this group on account of its did motivate these authors abandon the first not to trichodragmata. Assigned species are H. furtiva
if characters family. Still, such as the possession of Topsent, 1892 (specimen studied: ZMA POR 9760, euasters, and/or microrhabds, and/or trichodrag- Cape Verde, identified by the second author), H.
considered mata are primitive (and there can be lit- collectrix Thiele, 1903 (specimen studied: ZMA tle doubt of this in view of the wide distributionof POR 3053, Siboga Expedition, det. M. Burton; the these characters the there over tetractinomorphs), type specimen could not be found in the Sencken- are only two coppatiid synapomorphies of dubious berg Museum, Frankfurt), and H. violacea Boury- value remaining, viz. the absence of triaenes and Esnault, 1973 (specimen studied: fragment from the most of the Brazilian The is occurrence (in coppatiids) a comple- type specimen, coast). genus ment of tangential megascleres. considered valid on the basis of the loss of oxy-
The first synapomorphy, the absence of triaenes, asters. Melophlus shows important similarities with is since a suspect character, "lipotriaene" Stelletta Ecionemia: the acanthomicrorhabds of both are and Geodia species have been reported frequently, similar in shape (centrotylote) and size.
S. distincta erected e.g. (Thiele, 1900, as Rhabdastrella),S. Bergquist (1968) the genus Lamellomor- sterrastrea (Row, 1911, as Diastra), and S. pha for L. strongylata, a species having sanidasters cribriporosa (Dendy, 1916a, as Aurora); quite a in combination with a loose skeleton of flexuous
' ' few are disguised as “Jaspis” or “Dorypleres” Sol- strongyles. The affinities of this species remain ob-
1888. of Stelletta las, Relationships Schmidt, 1862, scure. She also assigned Coppatias baculifer Kirk-
Rhabdastrella Thiele, 1900, and Diastra Row, patrick, 1903, to Lamellomorpha, but it lacks the
still need be re-examined. In fact there 1911, to is "streptaster" category present in the type species, little for the that loss of triaenes ground assumption L. strongylata Bergquist, 1968. Its megascleres are did not and happen frequently independently in also true oxeas, which makes it closer to true Anco- various ancorinid groups. rina/Ecionemia, and especially to Hezekia De
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1888 and Ancorina Laubenfels, 1934, which was erected for an Stryphnus Solías, are compara-
Ancorina-like ( H lacking euasters. ble to Asteropus (sanidasters), and Ecionemia is species . demera)
al- to and Holoxea (spiny Jaspis is similarly problematic, where many comparable Melophlus
leged Jaspis do not conform to J. johnstoni but are microrhabds).
which follows like Stelletta, although they lack triaenes. For the Based on the above rationale, the
is collective ideas of Row De latter group the name Dorypleres Solías, 1888 (1911), Dendy (1916a),
available; if the loss of triaenes is considered to be Laubenfels (1936), Bergquist (1968 and pers.
of Stelletta-like and Van Soest & Stentoft (1988), the poly- a synapomorphy of a related group comm.),
be de- species, then re-erecting Dorypleres could be con- phyletic origin of the Coppatiidae cannot
sidered. The phylogenetic relationships of Jaspis nied. We postulate that the loss of triaenes occurred
s.s. remain obscure. If the ectosomal small oxeas in several ancorinid lineages and that the presence
of are homologous to the microxeas of Penares Gray, a complementary, variously developed, tangen-
skeleton be used 1867 (Burton & Rao, 1932; Bergquist, 1968), these tial ectosomal cannot as a synapo-
character for such a two could be related. However, certain Jaspis such morphic a group bearing
in as J. serpentina Wilson, 1925 share similarities diverse asterose microsclere complement. The oc-
ectosomal skeleton in their microscleres with Hemiasterella. currence of a tangential
of revision. Holoxea, Melophlus, Neam- The genus Jaspis is badly in need a Jaspis, Asteropus,
efforts the and is here interpreted as an Previous to split genus (i.e. Lindgren, phius, Lamellomorpha
of skele- 1898; Thiele, 1900, 1903; Topsent, 1904) were all adaptive answer, as a way strengthening a
considered impractical by Dendy (1916a), who was ton that has lost the "umbrella" formed by the cla-
of ectosomal triaenes. followed by Wilson (1925). The question that arises domes the
is all form These conclusions show that the whether or not Jaspis a monophyletic as- family Coppatii-
Its J. is The semblage. type species, johnstoni (Schmidt, dae, as presently understood, polyphyletic.
- and Holoxea 1862) hypothesized to be of pachastrellid origin genera Asteropus, Melophlus, are
by Dendy (1916a) and Bergquist (1968) - has much transferred to the Ancorinidae. The family Cop-
is retained accommodate smaller oxeas than those normally found in As- patiidae provisionally to
and until trophorida and the remaining coppatiids. Boury- Jaspis, Neamphius, Lamellomorpha a
revision of Pachas- Esnault (1973) mentioned that two species groups more complete Ancorinidae,
within the with and related "lithistid" were recognizable Jaspis: first, trellidae, Coppatiidae,
microxeas (e.g. J. johnstoni and J. salvadori forms has been undertaken. The affinities of the
Boury-Esnault, 1973) and the second, without restricted Coppatiidae must remainundecided until
then. microxeas (e.g. J. reptans [Dendy, 1905], J. topsen-
tiThiele, 1900 and J. novaezelandiaeDendy, 1924).
Boury-Esnault (1973) mistakenly assigned J. coria-
cea (Carter, 1886) to the second group. Carter Acknowledgements
(1886) clearly mentions the presence of a small category of oxeas (100 /xm long), restricted to the The authors wish to thank Dr. N. Boury-Esnault (Marseille),
Dr. Klaus Rützler (Washington), and Miss S.M. Stone (London) surface of his specimen. Jaspis reptans was
for sending specimens on loan. Mrs. Chantal Bezac (Marseille) described without a tangential skeleton of oxeas, is thanked for technical assistance in obtaining the SEM photos which makes it like Burton & Rao's (1932) "lipotri- A-D of Plate 1. Fieldwork in Arraial do Cabo (Brazil) was sup- aenose Stelletta” (or "reduced stellettids" [= an- ported by grants from Conselho de Ensino para Graduados corinids] sensu Bergquist, 1968). The relationships from the Universidade Federal do Rio de Janeiro (CEPG-UFRJ)
and from Fundação de Amparo à Pesquisa do Estado do Rio de between reduced ancorinids and "coppatiids" seem
Janeiro (FAPERJ). The first author received a fellowship from particularly close when certain genera (character- Conselho Nacional de Desenvolvimento Científico e Tecnológi- ized by a slightly reduced radiality) are considered. co from Brazil (CNPq) to undertake Ph.D. studies at the Univer-
In this Penares 1867 is to Gray, comparable A was way, sity of Amsterdam. first draft of this paper greatly im-
Jaspis (at least those with small ectosomal oxeas); proved after valuable comments from two anonymous referees.
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