Phorbas Posidoni N.Sp. (Porifera: Poecilosclerida) from the Aegean Sea, with a Discussion of the Family Anchinoidae

JOURNAL OF NATURAL HISTORY, 1991, 25, 827-836

Phorbas posidoni n.sp. (Porifera: Poecilosclerida) from the Aegean Sea, with a discussion of the family Anchinoidae

ELENI VOULTSIADOU-KOUKOLIRAt and ROB W. M. VAN SOEST~ t Department of Zoology, University of Thessaloniki, 54006 Thessaloniki, Greece Institute of Taxonomic Zoology (Zoologisch Museum), University of Amsterdam, P.O. Box 4766, 1009 AT Amsterdam, The Netherlands

(Accepted 8 March 1991)

A new species of the genus Phorbas Duchassaing and Michelotti, 1864 (junior synonym Anchinoe Gray, 1867) is described from the Aegean Sea (Eastern Mediterranean). Phorbas posidoni n.sp. is distinguished from the known Mediter- ranean species of Phorbas primarily by its elaborate repent-ramose habit combined with an extensive, spongin-enforced, plumoreticulate skeleton of oxea-like tornotes and the usual echinating acanthostyles and arcuate chelae. As in most Phorbas the surface bears characteristic areolae. The only other elaborate Mediterranean species, P. mercator (Schmidt) differs clearly in spiculation, since it lacks microscleres and has strongyles as the main skeletal elements. Phorbas fictitius (Bowerbank), P. tenacior (Topsent) and P. paupertas sensu Topsent are all incrusting and differ in spicule form and sizes as well. Likewise, Phorbas species from the neighbouring East Atlantic each show clear differences with P. posidoni n.sp. The genus Phorbas is discussed and compared with Pronax Gray (1867) sensu Lrvi (1973). It is proposed to unite the two genera because the alleged sharp difference between the two (only oxea-like tornotes in the skeletal tracts in Phorbas, only acanthostyles in the skeletal tracts in Pronax) is compromised by a range of intermediate conditions in various species. KEYWORDS: Systematics, Porifera, Poecilosclerida, Aegean Sea, Eastern Mediterranean.

Introduction During recent collecting in the Aegean Sea (Eastern Mediterranean) a species of the family Anchinoidae Topsent, 1928 (=Phorbasidae De Laubenfels, 1936) was discovered, which could not be assigned to any of the known species from the Mediterranean (for a list of these, see Pulitzer-Finali, 1983). The new species is unusual because of its elaborate repent-ramose habit, whereas most other Anchinoidae are massively incrusting. In addition to providing a description of the new species, this paper also intends to address the confused taxonomy of the family Anchinoidae, based on a critical survey of the literature and examination of the characters of the type species of most genera assigned to the family, notably Phorbas amaranthus, type of Phorbas Duchassaing and Michelotti (1864), and Pronax plumosa, type of Pronax Gray (1867). Based on the usually employed criteria in Mediterranean sponge taxonomy, see, for example, Topsent (1928), Vacelet (1969), Pulitzer-Finali (1983), the new species

002~2933/91 $5.00 © 1991 Taylor & Francis Ltd. 828 E. Voultsiadou-Koukoura and R. W. M. van Soest conforms to the genus Anchinoe Gray (1867). This genus is defined by most authors as having smooth diactinal spicules in the skeletal tracts and in the ectosomal region, with echinating acanthostyles only in echinating condition. Recently, Anchinoe was synonymized with Phorbas Duchassaing and Michelotti (1864) (Van Soest, 1984, and Discussion below) and this slowly gained wide acceptance (Bergquist, 1961; Van Soest, 1984; Bergquist and Fromont, 1988; Uriz and Rosell, 1989). Accordingly, the new Eastern Mediterranean species is assigned to Phorbas.

Systematic description Order POECILOSCLERIDA Topsent, 1928 Family ANCHINOIDAE (Topsent, 1928 (emended) (junior synonym: Phorbasidae De Laubenfels, 1936; Van Soest, 1984; even though the genus Anchinoe is now considered a junior synonym of Phorbas the family group name Anchinoidae is valid according to the ICZN rules). Poecilosclerida with plumose or plumoreticulate skeletal tracts consisting of choanosomal styles and/or smooth diactinal spicules, echinated by styles; ectosomal skeleton consisting of grouped smooth diactinal spicules lying perpendicularly, paratangentially or tangentially to the surface. Microscleres: chelae (if present) are arcuate; sigmata may frequently be absent; toxa and rhaphides are absent.

Genus Phorbas Duchassaing and Michelotti, 1864 Synonymy: Pronax Gray, 1867: 536; Anchinoe Gray, 1867; Suberotelites Schmidt, 1868; Plumohalichondria Carter, 1876; ?Clathrissa Von Lendenfeld, 1888; Stylostichon Topsent, 1892; Lissopocillon Ferrer Hernandez, 1916; Merriamium De Laubenfels, 1936. Definition (emended): Anchinoidae in which the styles composing the choanosomal tracts, if present, are heavily spined acanthostyles; echinating styles are likewise heavily spined. Occasionally, the coring acanthostyles are completely replaced by the diactinal smooth spicules. For justification of the synonymy see below.

FIG. 1. Holotype of Phorbas posidoni n.sp. Phorbas posidoni n.sp. (Porifera: Poecilosclerida) 829

Phorbas posidoni n.sp. HOLOTYPE,Zoological Museum of Amsterdam ZMA POR 8969, Gulf of Kavala, North Aegean Sea, 17 May 1977. Description. Form: the sponge is a ramose, bushy aggregate, more or less erect to sprawling, about 11 cm long, 9 cm high and 3 cm wide. The branches often anastomose and have a diameter of 0.5 to 1.5 cm and a height up to 5 cm; they are in most cases depressed and terminally thinned (Fig. 1). Colour: brown in formalin. Consistency: soft, elastic and compressible. Surface: it is smooth, with many tangential areolate pore fields. Oscules are numerous, irregularly distributed, up to 1 mm in diameter. The ectosome is fleshy and easily detachable. Skeleton: the ectosomal skeleton is made of abundant isochelae and hastate oxeas. Oxeas are arranged, at right angles to the surface, around areolate pore fields (Fig. 2). This vertical arrangement is not so strict in areas among the pore fields, where oxea-like tornotes can be found even tangentially laid in the ectosome. Isochelae are scattered without any pattern on the dermal membrane. The choanosomal skeleton is composed of well developed spongin fibres forming a more or less well organized network (Fig. 3). Oxea-like tornotes, identical to those found in the ectosome, are embedded in the spongin fibers, which are provided with echinating acanthostyles. No acanthostyles are found inside the fibres, thus having an accessory role in the skeleton. The spongin fibres have a diameter of l0 to 80 #m and they are in most areas totally cored with spicules, although in many of them only two-thirds of their width is cored. The spongin network is equally developed both in the base and near the surface of the sponge. Isochelae are present in the choanosome of the sponge but in lower abundance than in the ectosome. Spicules: megascleres. Diactinal spicules forming the main skeleton, are smooth oxea-like tornotes, measuring 265-310 x 3.5-6 #m. Acanthostyles, usually straight but sometimes slightly curved, covered with spines all along their length (Fig. 4). The spines, most of which are hooked become gradually smaller forward to the apex of the spicules. Acanthostyles measure 110-145 x 7#m. Microscleres: arcuate isochelae measuring 21.5~8.8 #m. Habitat. The sponge was dredged from a depth of about 30m. The substratum was sandy silt mixed with biogenic detritus.

FIG. 2. Ectosomal skeleton of Phorbas posidoni n.sp. 830 E. Voultsiadou-Koukoura and R. W. M. van Soest

FIG. 3. Choanosomal skeleton of Phorbas posidoni n.sp.

F~G. 4. Spicules of Phorbas posidoni n.sp. a, hastate oxea; b, acanthostyle; c, isochelae.

Etymology. The species is named after the Greek mythological God of the sea Posidon (Neptune).

Discussion Comparison with related species from the Mediterranean The new species differs from the known Mediterranean species (cf. appendix in Pulitzer-Finali, 1983) assignable to Phorbas especially in the elaborate growth form and the development of discrete spongin-enforced spicule tracts forming a reticulation. In the elaborate growth form, it is approached perhaps only by Phorbas mercator Phorbas posidoni n.sp. (Porifera: Poecilosclerida) 831

(Schmidt, 1868 as Suberotelites, see also redescription of Topsent (1938) and Pulitzer- Finali, 1983), and P. dendyi (Ferrer Hernandez, 1916 as Lissopocillon). From the former species the new Phorbas differs clearly in its possession of chelae (absent in P. mercator) and oxea-like tornotes (tylostrongylotes in P. mercator). The latter species is described as a massive sponge, but the photo (Ferrer Hernandez, 1916, Fig. 9) shows short projections and elevations. It is described in a new genus on account of the occurrence of 'pocillos', modifications of the arcuate chelae not worthy of generic separation. A further difference with our new species is the possession of two distinct sizes of acanthostyles in P. dendyi sensu Ferrer Hernandez. Note that if the synonymization of Phorbas and Pronax/Stylostichon is accepted, the combination of Phorbas dendyi would be preoccupied by that of Topsent (1892). The other species of 'Anchinoe', namely A. tenacior Topsent 1925, A. paupertas sensu Topsent (1925) and A. fictitius (Bowerbank 1866), differ in being thinly or massively incrusting. Moreover, of these only A.fictitius is similar in spiculation to our new species, differing, however, in possessing the usual two size categories of acanthostyles. Both A. tenacior and A. paupertas have smooth strongyles instead of oxea-like tornotes. The Mediterranean species assigned to 'Pronax', namely P. dives Topsent (1891), P. flbulatum Topsent (1893) and P. lieberkuehni Burton (1930), all differ in being incrusting. Moreover, P. dives and ~P.fibulatum have sigmata, and both of them as well as P. lieberkuehni have their choanosomal tracts consisting mostly of acanthostyles. 'Anchinoe' coriaceus (Fristedt, 1887) is considered a Hymedesmia (Van Soest, 1987); it lacks microscleres and the skeletal columns of strongyles are devoid of echinating megascleres. Comparison with related species from the eastern Atlantic Species referable to Phorbas reported from the neighbouring East Atlantic coasts are P. bihamiger (Waller, 1878), P. dendyi Topsent (1892), P. jecusculum (Bowerbank, 1866), P. microchelifer (Cabioch, 1968), P. microcionides (Carter, 1876), P. perarmatus (Bowerbank, 1866) (with probable synonym P. arneseni Topsent (1913), cf. Burton, 1930), P. plumosus (Moritagu, 1818), and P. repens Topsent (1904). The habit of P. dendyi may resemble that of Phorbas posidoni n.sp. in being also ramose. However, P. dendyi differs clearly in having acanthostyles dominating the choanosomal tracts, and in having smooth tylostrongylotes as ectosomal spicules. The other mentioned species are all massively incrusting. Like Phorbas posidoni both P. perarmatus and P. microcionides have their skeletal tracts dominated by oxea- like tornotes; however, P. perarmatus has isochelae twice as large as those ofP. posidoni, and P. microcionides has two distinct categories of acanthostyles. The specific identities of these two North Atlantic species remain to be confirmed, as both are poorly known. The remaining species differ from P. posidoni n.sp. in the dominance of acanthostyles in the skeletal tracts; moreover, the poorly known P. microchelifer, P. repens and P. bihamiger all have sigmata. We refrain from comparing the new species in detail with anchinoids from areas further away. In cases where the habit is comparable to that of P. posidoni n.sp. (e.g. Brasilian Anchinoe ramosa Hechtel 1983, or Namibian Anchinoe bardajii Uriz 1989)), there are invariably differences in skeletal organization of spicule categories and sizes. Discussion of the family Anchinoidae Generic content and synonymy. The original generic content of Anchinoidae included Anchinoe, Stylostichon, Echinodictyum, Lissopocillon and Hamigera. Some other genera 832 E. Voultsiadou-Koukoura and R. W. M. van Soest were added by later authors: Plumohalichondria, Clathrissa, Pronax, Suberotelites, Phorbas, and Myxodoryx. The family was erected by Topsent (1928) to receive sponges '... with diactinal ectosomal megascleres, with a skeleton consisting of more or less anastomosing polyspicular tracts echinated by megascleres, almost always with microscleres, but no toxa. Normally, the skeletal tracts contain spicules of the same type as those of the ectosome (Anchinoe Gray, 1867, Lissopocillon Ferret Hernandez, 1916, and Echinodic- tyum Ridley, 1884), but these are absent in the tracts ofStylostichon Topsent, 1892. The echinating spicules are normally acanthostyles, mostly in two sizes but in a single size in Echinodictyum, where they have a blunt point. Still, in Hamigera Gray, 1867, smooth subtylostyles echinate tracts of smooth substrongyles...' (literal translation of Topsent's text). Clathrissa Von Lendenfeld (1888) was suggested to be junior synonym of Anchinoe by Stephens (1921); this synonymization remains to be substantiated. Echinodictyum was at a later date removed from the family by L6vi (1973) and is generally assigned to the family Raspailiidae (cf. Hooper, 1991 in press). Hamigera is a well-known Mediterranean genus, with one Mediterranean and a few Australian-New Zealand representatives. Lissopocillon is known only from Ferrer Hernandez's description and is here synonymized with Phorbas. Myxodoryx Burton (1929) by virtue of its coring smooth styles is not a member of Anchinoidae (probably Myxillidae). Suberotelites Schmidt (1868) was demonstrated to be a junior synonym of Anchinoe by Topsent (1938). The history of the remaining genera Phorbas, Anchinoe, Plumohalichondria, Stylostichon and Pronax is thoroughly confused. Even though some of them (notably Anchinoe and Phorbas) have been demonstrated recently to be synonymous, the use of these names persists. After the study of many anchinoid species and an exhaustive literature survey, our conclusion is that all five genera are either objective or subjective synonyms of a single, large genus Phorbas. Because major authors (Bergquist, L6vi, Wiedenmayer) continue to use several of these names and, moreover, in an inconsistent manner, a justification to convince these authors of the correctness of our conclusion is necessary. The order in which the various genera are treated is chronological.

History of the genus Phorbas and its synonyms. Phorbas was erected by Duchassaing and Michelotti (1864) for P. amaranthus (designated type by De Laubenfels, 1936: 63) and P. vieqensis, both from the West Indies. De Laubenfels (1936) attributed P. amaranthus with the wrong characters, e.g. echinating acanthoxea, and sigmata, but Burton after examining a fragment of the type specimen in the British Museum (Natural History) decided that it conformed closely to Anchinoe (he assigned Anchinoe fictitius to Phorbas in 1957). Van Soest (1984) redescribed the type specimen (Museum Torino) and studied also several fresh specimens. From these it became clear that it shares characters with both the 'genera' Achinoe and Pronax/Stylostichon in that the skeletal columns starting from the base are composed firstly by acanthostyles which towards the periphery become progressively replaced by smooth oxea-like tornotes; these also fan out to the surface to enclose 'criMes' (areolated pore fields) known from most Anchinoe and several Pronax/Stylostichon. Gray (1867) erected Anchinoe for the Shetland species Hymeniacidon perarmatus Bowerbank (1866) (redescribed by Stephens, 1921; see also Burton, 1930), a thinly incrusting smooth sponge with numerous oscula; its ectosome is covered with abundant oxea-like tornotes and chelae, its skeletal columns consist of the same oxea- Phorbas posidoni n.sp. (Porifera: Poecilosclerida) 833 like tornotes, echinated by small acanthostyles; large acanthostyles are erect on the substrate; Bowerbank's drawing of the chelae (Bowerbank, 1874: pl. 31, Figs. 14-15) are peculiar, but presumably they are arcuate. Carter (1876) described Plumohalichondria microcionides from roughly the same locality (about 800 m): it is a smooth irregularly mamillated sponge with 'internally radiating plumose' skeleton; smooth oxea-like tornotes are confined to the tracts ('. i. the acerate skeleton-spicules are confined to the fibre of the sponge .... '); there are two sizes of acanthostyles, and arcuate (?) chelae of a large size if the measurements of Carter are correct. In most details the two above descriptions match, perhaps both concern the same species (which may have another synonym in Desmacidon neptuni Schmidt, 1875, cf. Thiele, 1903), but certainly PlumohaIichondria and Anchinoe are synonyms. Topsent (1892) discovered that in a later publication Carter (1885) emended his genus Plumohalichondria by assigning a species with acanthostyles in the skeletal tracts to it, namely Spongia plumosa Montagu (1818), and proposed to have this species, along with some related species, separated in a genus Stylostichon. Unfortunately, Topsent (1892) also included Plumohalichondria microcionides, the type of Plumohalichondria, in the species assigned to Stylostichon, thus confusing matters considerably. This confusion has persisted up till now, as Bergquist and Fromont (1988) still maintain that the type species of Plumohalichondria has plumose columns of spiny monacts, quoting Topsent (1892). Wiedenmayer (1989), rejecting Pronax for reasons given below, even reinstated Plumohalichondria as a senior synonym of Stylostichon. Lundbeck (1909) realized that Topsent (1892) had made a mistake in assigning the type of Plumohalichondria to Stylostichon and indicated S. dendyi Topsent (1892) as the type species of Stylostichon, hoping by this action to conserve the genus name. Stephens (1921) also considered PIumohalichondria a junior synonym of Achinoe and kept Stylostichon for S. dendyi. This type designation however, was not accepted by De Laubenfels (1936) and L6vi (1963, 1973), who maintained S. plumosa as the first and principal species named by Topsent in connection with Stylostichon, and the obvious type species. Unfortunately, P. plumosa was already named a type of the genus Pronax Gray (1867), a discovery made by De Laubenfels (1936), who also noted that Pronax was also used for a genus with type species Cliona lobata Hancock. De Laubenfels (1936) declared the latter Pronax to be preoccupied and gave a replacement name Grayax with the same Cliona lobata as type species. The action of De Laubenfels was not only totally unnecessary, since there is (and was) no doubt over the correct generic assignment of Cliona lobata, but it also violated the rule of page priority, since the anchinoid Pronax was named on page 536 of Gray's paper and the clionid on page 526. It seems clear that Pronax is unavailable as a genus name in the Anchinoidae, because of previous use of the name for a Cliona, of which it is a junior synonym. Lundbeck's subsequent designation of StyIostichon dendyi as the type of StyIosti- chon is valid according to the ICZN rules, because Topsent (1892) when establishing the genus, named four species as 'examples' without indicating explicitly which was the type species. A first reviser is in such cases entitled to declare any of these four as the type. Moreover, S. dendyi was the only species described fully at the erection of the genus, whereas the others were mere literature assignments. This means, that we can be certain that S. dendyi comes closer to what Topsent intended the genus to comprise than the other named species. To add to the confusion, L6vi's (1973) definition of Anchinoe allowed a mixture of smooth diactinal spicules and acanthostyles in the fibres, thus (without referring to it) 834 E. Voultsiadou-Koukoura and R. W. M. van Soest

Table 1. Speciesof Anchinoidae showing an intermediate condition of both acanthostyles and diactines in the skeletal tracts (based on specimens in the ZMA collection and descriptions in the literature).

Phorbas amaranthus Duchassaing and Michelotti, 1864 (Holotype Mus. Torino Por. 59; holotype of Merriamium tortugasense De Laubenfels, 1936, USNM 22418 which is a junior synonym; ZMA specimens from Curaqao) Phorbas plumosus (Montagu, 1818) (Bowerbank, 1866: 199, description of Montagu's type; specimens in ZMA from Ireland and Roscof0 Phorbas dendyi (Topsent, 1892) (ZMA specimens from Norway) Phorbas tenacior (Topsent, 1925) (Topsent, 1925: 668, as Achinoe, description of localization of large acanthostyle) Phorbas paupertas (sensu Topsent, 1934) (Topsent, 1934: 44, as Anchinoe, a description of condition of thinner stages) Phorbas intermedia Bergquist (1961) (Description in Bergquist & Fromont, 1988) Phorbas areolata (Bergquist & Fromont, 1988) (Description) Phorbasfuloa (Bergquist and Fromont, 1988) (Description as Pronax) Phorbas cf. tenacior (sensu Wiedenmayer, 1989) (Description) Phorbas areolata sensu Desqueyroux-Faundez (1989) (Description as Anchinoe)

admitting to the closeness to Pronax/Stylostichon. Perplexingly, Boury-Esnault and Van Beveren (1982), followed by Uriz (1989), emended the definition of Anchinoe as having skeletal columns consisting of acanthostyles and echinated by ectosomal diactines, without any discussion or comment. It is assumed here that the latter definition is a mistake by these authors. From the above-presented confusing array of whole and half mistakes, wrong emphases, and nomenclatorial actions, it appears that the major difference between Anchinoe/Phorbas and the rival genus Pronax/Stylostichon, is seriously compromised by the occurrence in the type species of Phorbas (and in many other related species) of an intermediate condition between having the skeletal tracts filled exclusively with oxea-like tornotes or with acanthostyles. Topsent's remarks (1925) on ontogenetic variability of the skeleton of anchinoids adds further evidence on the absence of clear- cut differences in two separate 'genera'. These observations effectively prevent the correct generic allocation of many specimens and species (see Table 1 for a list of examples). Accordingly, it is here proposed to unite these 'genera' under the oldest available name, namely Phorbas. It is expected that a future formal character analysis of Anchinoidae (which should include also the closely related Hymedesmiidae) will show the spicular composition of the choanosomal tracts to be part of a transformation series from completely filled with Phorbas posidoni n.sp. (Porifera: Poecilosclerida) 835 acanthostyles to completely filled with smooth ectosomal spicules. The various positions of these spicules in Hymedesmia may have considerable bearing on such a character analysis.

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