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Notice © 1990 Institute for Systematics and Population Biology (Zoological Museum), University of Amsterdam. This manuscript is an author version with the final publication available and may be cited as: Van Soest, R. W. M., Diaz, M. C., & Pomponi, S. A. (1990). Phylogenetic classification of the halichondrids (Porifera, Demospongiae). Beaufortia, 40(2), 15-62.

BEAlJFOR TIA

INSTITUTE OF TAXONOMIC ZOOLOGY (ZOOLOGICAL MUSEUM) UNIVERSITY OF AMSTERDAM

Vol. 40, no. 2 July27, 1990

PHYLOGENETIC CLASSIFICATION OF THE HALICHONDRIDS (PORIFERA, DEMOSPONGIAE)

R. W. M. VAN SOEST*), MARIA CRISTINA DIAZ") & SHIRLEY A. POMPON!***) *) Institute of Taxonomic Zoology, University of Amsterdam, P. 0. Box 4 766-1009 AT Amsterdam, the Netherlands **) Marine Sciences, U. C. Santa Cruz, 273 Applied Sciences Building, Santa Cruz, California 95064, U.S.A. ***)Harbor Branch Oceanographic Institution Inc., 5600 N Old Dixie Highway, Fort Pierce, Florida 34946, U.S.A.

ABSTRACT

Using a multicharacter approach and numerical cladistic computer programs a phylogenetic analysis is made of a newly defined order Halichondrida (which includes all Halichondrida and parts of the Axinellida sensu L6vi, 1973), with emphasis on the newly defined family Halichondriidae (which includes the families Halichondriidae and Hymcniacidonidac sensu LCvi, 1973, and some genera formerly assigned to the Axinellidae and Desmoxyidae). The newly defined order consists of the "old" families Axinellidae/Bubaridae and Desmoxyidac, a "new" family Dic­ tyonellidae (containing Dict_yonel/a, Acanthella, Liosina and Dactylella), and the new concept of the family Halichondriidae. On the basis of numerous specimens, including many type and other original material from all parts of the world oceans, the latter group has been analyzed in depth. Next to classical morphological characters also biochemical data (secondary metabolites) were used. This resulted in the recognition of several discrete generic groups, viz. Axilu>ssa + Colloralypta, A1yrmekioderma + Didiscus, Spongosorites, TojJscntia + Epipolasis + Petromica, Ciocalypta + Amorphinopsis, and Hymcniacidon + Hafichondria. A revision comprising definitions of and remarks on all genera, including all junior synonyms, of the newly defined Halichondriidae concludes the paper.

INTRODUCTION at the subclass level and if used at lower levels are suspect as likely adaptive characters easily The Halichondrids s.s., until very recently rec­ developed independently. ognized as a separate order (cj. Levi, 1973; If reproductive strategies are abandoned as Bergquist, 1978; Hartman, 1982) are a group characters at the subclass-level, it is necessary heavily affected by the major changes of to reevaluate the contents and the position of classification (based on reproductive strategies) the Hali'thondrids. Characters of the sponges introduced by Levi (1953), which subsequently involved are discussed and evaluated, and a gained general acceptance. It has been argued new classification based on (numerical) cladistic elsewhere (e.g. Van Soest, 1987, 1990b; Diaz, techniques is presented. For a general outline of eta!., 1990a), that reproductive strategies can­ these methods see Wiley ( 1981); for sponge not be employed as synapomorphous characters examples see Van Soest (1987, 1990a and b).

15 The newly recognized valid families and genera (Axinellida) - Acanthella (?) - Dictyonella (?) - and their junior synonyms are listed, defined, Axinyssa (?) - Myrmekioderma (?) - Topsentia and illustrated, based on many type and other (Halichondrida) - Ciocalypta (Haliehondrida) - original specimens from all over the world Halichondria (Halichondrida). In fact, the oceans. definitions of the "orders" Axinellida and This study is the fourth part of a revision of Halichondrida apply only really well to the type the Halichondrids undertaken by Pomponi, genera Axinella and Halichondria, and it is only Dfaz & Van Soest. Preliminary papers on this through other characters (e.g. sinuous spicules revision have been published by Diaz et al. in Phakellia and Acanthella, and Axinella cannabina (1990a) and Pomponi et al. (1990), and a review (Esper, 1794); telescoped endings of spicules in of Central Atlantic representatives of the newly Axinella, Dictyonella, Axinyssa and Spongosorites; defined family Halichondriidae was made by size ranges of oxea in Myrmekioderma, Topsentia, Diaz et al. (in press). Ciocalypta and Halichondria), that typical genera are linked with intern1ediates. From these observations, it may be inferred that to provide AXINELLIDA-HALICHONDRIDA RELA­ definitions of two discrete groups (orders or TIONSHIPS suborders) encompassing all the associated To perform a cladistic analysis of the families genera is virtually impossible. It will not be and genera of the Halichondrids, it is necessary attempted here; instead it will be shown below to find the nearest outgroups which enables that the group of genera together comprising character polarization (Watrous & Wheeler, the presently recognized Axinellida s. s. and 1980). Van Soest (in the press) and Diaz et al. Halichondrida s. s. consists of four definable (in the press) showed that Halichondrids and genus-groups which can conveniently be given some Axinellida share many characteristics. family status. Also, pre-Levi (1953) classifiers (Topsent, Bur­ The phylogenetic position of Raspailiids ton, De Laubenfels) associated Halichondrids (including Euryponids) and Hemiasterellids with Axinellids, particularly the family has to be discussed briefly, since these have Axinellidae. In fact, both were considered been considered close to Axinellidae in the past, members of a single order Halichondrida and indeed showed up to be close in a recent Vosmaer, 1887 by Topsent and De Laubenfels, phylogenetic exercise (Van Soest, 1987), based until the Axinellida were given separate ordinal on possesswn of an axially condensed status in 1955. Halichondrida s.s., i.e. families choanosomal skeleton. Ongoing cladistic Halichondriidae and H ymcniacidonidac sensu analyses (Van Soest, 1990b), however, reveal Levi (1973), can be associated with Axinellida the probable homoplasic nature of the axially Levi (1955) (in part only: familes Axinellidae, condensed skeleton. Typical members of Bubaridae, and perhaps some Desmoxyidae, Axinellidae, Raspailiidae, Hemiasterellidae but excepting the Raspailiidae, Euryponidae, and Sigmaxinellidae share the axial condensa­ Rhabderemiidae, Trachycladidae, Sigmaxinel­ tion of the choanosomal skeleton; however, lidae ( ~ Hamacanthidae) and Hemiastercl­ each of them also shares indubitable lidae), by a convincing series of intermediate synapomorphous characters with widely dif­ genera and families. The major features of the ferent groups such as Hadromerids Axine1lida, i.e. axial condensation of the (Hemiasterellid asters) and Poecilosclerids choanosomal skeleton and indiscriminate (Raspailiid acanthostyles and Sigmaxinellid occurrence of styles and oxea, and the major sigmata). There is no solution for this distribu­ features of Halichondrida, I.e. ectosomal tion of characters other than to assume that one tangential skeleton and confused choanosomal or more homoplasic developments have occur­ skeleton, intergrade nicely in the series of red in the evolution of these groups. According genera Axinel/a (Axinellida) Plzake/lia to recent cladistic analyses (Van Soest, 1990b),

16 it is more parsimonious to assume that axial E 0"' 0 0 0 ..: ~ condensation has developed in four different ~ "' Q) 'C " a. ~ "' 0 "'Q) lines (Axinellidae, Hcmiasterelliidac, Raspailii­ Q) -f- J:"' J:"' c. J:"' ~ dae and Sigmaxincllidae), than to assume that asters, acanthostyles and sigmata have each developed independently in two separate lines (asters: Hadromerids and Hemiasterellids; acanthostyles: Raspailiids and Micro­ cionids/Myxillids; sigmata: Sigmaxincllids and Poccilosclerids). Speculations about presence of any of those characters in unknown ancestors of the groups, and subsequent independent losses in several major lines, seem to be all equally ( un-)parsin1onious and unnecessary. These results arc in support of Hooper (1990), who correctly pointed out the sitnilarities of Fig. 1. Revised classification of the Dcmospongiae based on the reevaluation of major group synapomorphies Raspailiids and Microcionids. h1oreover, axial (based on Van Socst, l990b). Character A: (sub-)radiatc condensation of a sort is found in several architecture, B: anisotropic reticulation, C: taurin syn­ Suberitid Hadrmnerids, viz. species of Rhizax­ thesis via cysteic acid, D: sigmatose microscleres. inella and the species originally described as Phakellia lobata Wilson, 1902. Comparisons of axially condensed species of all these groups show in1portant architectural differences vide strong (underlying) synapomorphies between various axial skeletons (see Van Soest, (Pomponi, el al., in press). 1990b), testifying for the probable homoplasic nature of this type of supporting skeleton. The Definition of the Halichondrida s .I.: revised classification of the Hadromerid­ DemosjJongiae with a plwnoreticulate skeletal architec­ Axinellid-Halichondrid-Poccilosclerid ture built of interchangeable styles and oxea and Demospongcs is presented in Fig. 1 (after Van intermediate spicules, of widely diverging sizes, and not Socst, 1990b). Junctional!;• localized. A problem remaining is the formal definition Remarks: The plumoreticulate skeletal of the group comprising Axinellidae s. s. and architecture maybe a synapomorphy for a Halichondrida (Levi's (1973) families much wider group of Demospongiae (see Van Axincllidae, Bubaridae, Desmoxyidae, Soest, 1990b). This character is progressively ~Halichondriidae, and Hymeniacidonidae), lost in the group formerly considered the order which may be named Halichondrida (s.l.) Halichondrida sensu Levi. However, a basic because the present contents of this group plumoreticulate or plumose architecture is largely conform to those of the similarly named found even there with few exceptions. group of Topscnt and De Laubenfels. Although Although the combination of styles and oxea as argued above a convincing series of IS also found m a few Raspailiids and intern1ediates exists linking this group closely Poecilosclerids, these groups show localization together, it seems nevertheless that easily or categorization in them. For instance, in recognizable synapomorphies are absent. Thus, M yxillids stylote choanosomal spicules are we have to rely on underlying synapomorphies combined with ectosomal diactines; in the found in many but not all members of a taxon Raspailiid Echinodictyum choanosomal oxea are and unique combinations of characters not arranged into tracts which are echinated by uniquely confined to the group. Possibly, (acantha-)styles. Indiscriminate occurrence of biochemical compounds may turn out to pro- styles and oxea is not found. Localized spicules

17 in Didiscus, Myrmekioderma and Higginsia arc sideration for their likely Raspailiid or considered microscleres. Hemiasterellid nature. The remaining 92 Sinuous longer or shorter diactinal spicules nominal genera recognized within the newly are found in Axinella cannabina, Phakellia) A1detta) defined Halichondrida could be subdivided Bubaris, Acantheila, Dactyleila, some Axinyssa and an1ong the following generic groups: some Topsentia. This distribution may be inter­ preted as retention of ancestral spicules, Family Axinellidae Ridley & Dendy, 1887 possibly related to lithistid spicules (through (emended) (Figs. 2-3, 8): Haiichondrida with Monocrepidium and fossil genera such as axially condensed and extra-axially plumoreticulate Cephalorhaphidites and Megaloraphium), and thus choanosomal skeletons. constitute an underlying synapomorphy. Isonitriles arc organic compounds found in Characteristically, oscules are star-shaped Axineila, Acanthella, Ciocalypta, Axinyssa, (Fig. 8) but in view of its (non-homologous?) '' Epipolasis' ', '' Stylotella'', ''Halichondria'', and occurrence outside the group that is a doubtful ''Hymeniacidon'' (names between quote marks synapomorphy. Next to typical Axinellidac represent doubtful generic assignments) (per­ genera like Axinella (Figs. 2, 8), Phakeilia and sonal communication Dr J. C. Brackman), and Auletta, the family Bubaridae with valid genera thus may turn out to be a good underlying Bahan's (Fig. 3), Monocrepidium, Hymerhabdia, synapomorphy, too. and Cerbarz's, are here included based on the observation that the. incrusting growth form is supported by an axially~ basally condensed RELATIONSHIPS WITHIN THE HALI­ and an extra-axially~ peripheral plumo­ CHONDRIDA (s. i.) reticulate skeleton. Although our revision will confine itself to the Halichondrida in the restricted sense (which Family Desmoxyidae Hallmann, 1917 will subsequently be named the family (emended) (Figs. 4, 9): Halichondrida with a Halichondriidae) and will not include at this reticuiatejasciculate choanosomai skeleton of spongin­ moment Axincllidae/Bubaridae/Dcsmoxyidae, enveloped tracts with a fleshy, corrugate/ridged surface. we will need to find the nearest outgroup of the group we wish to study further. So it is Genera included are Higginsia (with junior necessary to briefly discuss the contents of the synonym Desmoxya), which possesses ectosmnal Halichondrida s.l., for which purpose we con­ centrotylote acanthoxca, and Ptilocauiis (Figs. 4, sidered the descriptions of an original 148 9), which lacks this character but is very close genera, of which 56 were later left out of con- in habit. Possibly, this group is more related to

Figs. 2-7. Figs. 2. Axineila verrucosa (Esper, 1794), A. longitudinal section showing axially condensed and extra-axially plumo­ rcticulate skeleton (redrawn from Vosmaer, 1912: pl. 16 fig. 5) (scale bar (s.b.) = 1 em), B. Spicules (s.b. = 100 fJ.In). Fig. 3. Bubaris vermiculata (Rowerbank, 1866), choanosomal styles erect on a basal mass of sinuous strongyles (s.b. = 100 ~m). Fig. 4. Ptilocaulis walpersi (Duch. & N1ich., 1864), A. longitudinal section showing reticulation of spongin-enforced tracts (s.b. = 1 mm), B. spicules (s.b. = 1 oo fJ.m) (both redrawn from Zea, 1987: fig. 67). Fig. 5. Dict;·onella obtusa (Schmidt, 1868), A. dendritic spicule tracts (redrawn from Vosmaer, 1912: pl. 16 fig. 1) (s.b. = 1 mm), B. spicules (s.b. = 100 11m). Fig. 6. Scopalina lophyropoda Schmidt, 1862, A. dendritic spicule tracts (redrawn from Schmidt, 1862; pl. 7 fig. 18) (s.b. ~ 100 ~m), B. style (s.b. ~ 100 ~m). Fig. 7. Acanthella acuta Schmidt, 1862, A. dendritic spicule tracts (redrawn from Vosmaer, 1912: pl. 16 fig. 3) (s.b. =

1 mm), B. spicules (s.b. = 100 fJ.m).

18 B

3 A 2

A A 4 B 5

., "'' B

B 7 6

19 Raspailiids and Microcionids because of the families. The absence of axial condensation of converging habits found in these groups; the skeleton is here interpreted as an ancestral absence of acanthostyles in the present family character, which cannot be used as a seems to be preclude a more close association synapomorphy. The resulting picture, which is between these forms. Also there seem to be two admittedly rather weak, is graphically rep­ distinct habits in Higginsia species: one more or resented in Fig. 12. less smooth with a cover of tightly packed microscleres (e.g. Hi"gginsia petrosioides Dendy, Family Halichondriidae Vosmaer, 1887 1905), the other Ptilocaulis- like possessing long (emended): Halichondrida with a choanosomal styles. The latter may be related to the skeleton consisting of (1) a lu~~h density of spicules Euryponid Halicnemia which shares the arranged in (2) vague, ill-defined, directionless tracts, microsclere morphology. A revision of Higginsia and of (3) spicules in confusion. is necessary for evaluating the contents of the family Desmoxyidae. Remarks: As is remarked above, the new concept of the family Halichondriidae includes Family Dictyonellidae, new family (Figs. 5-7, the old families Halichondriidae and 10-11 ): Halichondrida with spongin-enforced den­ H ymeniacidonidae, as well as '' Axinellid'' dritic choanosomal spicule tracts and a fleshy co111dase genera such as Myrmekiaderma and Axilryssa; surface. moreover, also Didiscus, in the past regarded variously as Hadromerid or Poecilosclerid, but Genera included are Dictyoneila (Figs. 5, 10), properly assigned recently by Diaz et al. (1987). Scopalina (Fig. 6), Acantheila (Fig. 7), Dactylel/a, A total of 46 nominal genera (see below) are Tethyspira, and Liosina (Fig. 11). considered to belong to this group. After The presence of reticulate skeletons in examination of specimens of most type species Axinellidae and Desmoxyidae, and also in (including many holotypes), and many other many Raspailiids and Poecilosclerids, is here species assigned to these genera, we consider 12 interpreted as ancestral, be it that the particular genera as valid, with another 4 monotypical development of this reticulate architecture in genera of doubtful status. the different groups represent apomorphous states. The absence of a reticulate skeleton in Dictyonellidae and Haliehondriidae is con­ CLADISTISTIC ANALYSIS OF THE sidered a synapomorphy for the two families, HALICHONDRIIDAE thus making the Dictyonellidae the sistergroup (or functional outgroup) of the Halichon­ We selected two representative species of each driidae. Dictyonellidae and Desmoxyidae share of the twelve valid genera and included some the fleshy ectosome, which is also found in some representatives of the outgroups (Phakellia, Halichondriidae such as Axinyssa and Ptilocaulis, Dactylella and Dictyoneila). We sub­ Hymeniacidan, and this is hesitatingly considered mitted this group of 27 taxa to a formal charac­ an underlying synapomorphy of the three ter analysis. This involved drawing up a list of

Figs. 8-11. Fig. 8. Axinella donnani Bowerbank, 1875, Z:NIA specimen from Indonesia {s.b. = 1 em). Fig. 9. Ptilocaulis walpersi (Duch. & 1v1ich., 1864), ZMA specimen from Curac;ao (s.b. = 1 em) (photo L.A. van der Laan). Fig. 10. Dict)'onella obtusa Schmidt, 1868, ZMA specimen from Marseille (s.b. = 1 em) (photo L. A. van der Laan). Fig. 11. Liosina paradoxa Thiele (1903), ZlviA specimen from Indonesia (s.b. = 1 em) (photo H. van Brandwijk).

20 21 Q) C!S Q) "0 C!S "0

Q) c Halichondriidae

high spicule density vague tracts spicules in confusion

loss of reticulate skeleton

fleshy ectosome

collagenous mesohyle plumoreticulate architecture sinuous strongyles 1 non-localized interchangeable styles and oxea

Fig. 12. Phylogenetic relationships of newly defined Halichondrid families (fUrther explanations sec text).

26 characters (sec below) and tabulating the (which necessitated transformation of the distribution of these characters and their states multistate character matrix into a additive­ over the taxa, thus constructing a taxon­ binary one using the ''Factor'' option, after character matrix. Where it appeared necessary, which "Mix", and Dollo parsimony algorithn1s it was decided to use a tnulti-state coding were used) (Felsenstein, 1987). because of the obvious interdependent nature of n1any characters/states. The necessity to develop transformation-series hypotheses in the Synapomorphic characters used in the cladistic ana(ysis case of multistate characters seriously con1- of relationsluj;s in the Hahchondriidae: plicates this type of analysis; however these complications were preferred over a strict 1. Occurrence of a (()mbination of oxca, st_ylcs and mod(fiwtions binary coding, which has serious philosophical of these: this is a synamorphy of the order Hali­ chondrida. setbacks. Subsequently the taxon-character 2. Sinnon' 'picule<: these occur in some Bubaridae, matrix has been evaluated vvithout computer Axinellidae, Dictyonellidae, and Halichondriidac; assistance, and also treated with three different however, the distribution within these groups is computer programs all constructing VV agner­ restricted to a small number of species and genera, Trecs in which relationships between taxa and and within a species the proportion of sinuous spicules may be quite small (e.g. in Dacl)'ldla and Tapsmtia). their hypothetical ancestors are computed, viz. The presence of these spicules is here considered an MacC!ade 2.1 (Maddison & Maddison, 1987), underlying synapumorphy of the order Hali­ Tree Tools 2.5 (Ellis, 1987) and Phylip 3.0 chondrida.

22 3. Reticulate c!wanosomal architecture: this is an ancestral spongin (in the form of fibres or enveloping or binding character of a much larger group including the spicule tracts); this collagenous choanosome is lost in families Desmoxyidae, Axinellidae, but also probably all genera except in Axin)'Ssa, Colloca!ypta, A1;,rmekio­ the Poecilosclerida. For the Halichondriidae and Dic­ dcrma and Didiscus, and its loss (lOb) is considered a tyonellidae its loss (3b) is considered a synapomorphy, strong synapomorphy for the remaining Hali­ thus uniting both families and making the Dic­ chondrids. tyonellidae the nearest outgroup of the Halichon­ 11. iHcandering surface ltroovcs: deep surface grooves, driidae. sinuously curving or meandering, roofed over by 4. SjJongin-enforced perjJendicular tracts: these consist oflong "organic skin", leaving isolated rounded areas with somewhat wavy spicules, and often follmv a den­ ectosomal pallisadcs, form a distinct synapomorphy of dritically anastomosing course ending at the surface i\1J'rmekioderma and Didiscus. and causing a more or less pronounced conulose 12. Sinuous trichodragmata: these are a synapomorphy of exterior: a synapomorphy of the family Die- A1]nnckioderma, where there arc normally several size tyonellidae. categories. An assumed homoplasic occurrence of 5. High spicular density: the reduced spongin in the sinuous trichodragmata is found in .E'pipolasis; the Halichondriidac causes a loss of reticulation and a dragmata are definitely slimmer and much more more confused skeleton and leads to a higher spicular· curved m that genus (arranged around the density: a synapomorphy of the bmily Halichon­ mcgascleres and keeping these presumably together). driidae. Outside the Halichondrida sinuous trichodragmata 6. Vague sjJicule tracts: loss of spongin and reticulation has are found in the Poecilosclerid family Desmacellidae left only a vague organization of central confused (e.g. Dragmatel!a). clements and equally vague tracts radiating to the sur­ 13. D1'scorhabds: strong synapomorphy of Did1'scus; since face: a synapomorphy of the family Halichondriidae. they develop from thin oxea they may be homologous 7. Haphazard arrangement of spicules: also related to the with the smallest category of megascleres of other above is the confused arrangement of many single genera, but in view of their small size this is left spicules: a synapomorphy of the family Halichon­ undecided. driidae. 11. Choanosomal tracts of spicules strtlll!,lhcncd b)• some sjJOngin 8. Finely conulose surface: without ectosomal skeletal cover­ running more or less parallel with the surface: a synapomor­ ing, the type of choanosomal skeleton found in the phy for Spongosorites. Halichondriidac results in groups of spicules sticking 15. Colour-change: from yellow or orange to brown or out (often at diverging angles) beyond the ectosomc, black, when exposed to air or any other medium con­ producing a finely conulose surface: this is a taining free oxygen (acrophobic): a synapornorphy of synapomorphy of the genera Axinyssa and Collocalypta. Spongosorites. 9. Ectosomal skeletal strengthening: the remaining Halichon­ 1fi. TojJsentins: secondary metabolites found only in driidae have some sort of ectosomal skeleton (and if S'pongosorites. not they arc so obviously related to forms that possess 17. Progressive loss of spongin: presumably in two stages all it, that the subsequent loss is a likely assumption): a genera (excepting those of the genus groups Axinyssa­ strong synapomorphy of all genera except Axinyssa and Collocalypta and }vfyrmekiodmna-Didiscus and also Colloca(ypta. The strengthening- occurs in several Spongosorites) exhibit loss of spongin; most genera forms: (9b) the cctosomal skeleton may consist of a retain some binding the spicules ( 17b ), but in the mass of spicules arranged more or less vertically, genus group Topsentia-Ept'polasis-Petromica loss of mostly tightly packed, erected on a base of larger spongin is virtually total resulting in a notable brittle tangential spicules: a synapomorphy of the genera fragile consistency (17c). A1yrmekioderma and Didisws; (9c) the ectosomc may 18. Absence of tme st)'les: a "weak" synapomorphy (18b) consist of a tangential arrangement of bundles of setting Topsentia and Epipolasis apart from Petromica. spicules or single ones, a synapomorphy for the genus 19. Presence of desmata: possibly an ancestral character set­ group Ciocalypta-Amorphinopsis-Halichondria-Hymeniaci­ ting Petromica apart; maybe inadmissible, if possession don; (9d) a possible variant of this is an utterly con­ of desmata turns out to be a retained ancestral fused crust as found in the genus Topsentia; (9e) a character. recognizable variant is also the detachable ectosome of 20. Trichodragmata drajJed around the spicules: an apomor­ Hymeniacidon and Halichondria; (9f) a doubtful apomor­ phous character of bfJI'polasis. phous character state is formed by the parchment-like 21. Ectosomal small styles: a synapomorphy of the genus ectosomc that sets Epipo!asis apart: oxea arranged in a group Amorphinapsis-Cioca(ypta. tangential felt-like mass; (possibly independently 22. Bundles of longer oxeotes 'echinated' by small St)'les: a developed in the doubtful genus I)>loderma). synapomorphy of Amorphinopsis. 10. Collagenous mesohyle: most Axinellidae and Dic­ 23. Axially arranged fistule architecture: a ''vveak'' tyonellidae show a highly collagenous choanosomc synapomorphy of Ciocalypta; in Petromira the axial apart from the variable presence of recognizable strengthening skeleton is found in the wall of the 23 fistules, rather than in the centre, but in other respects Axlnyssa it is similar; in certain fistulose Toftscntia and Halichon­ dria arrangements are more or less the same as in Collocalypta Cioca(ypta. 24. Tufted larvae: a synapomorphy of Halidwndria. Myrmekloderma 25. VimjJa1:y: a synapomorphy of the genera Halichondria Dldlscus and H)'meniacidon; this is certainly an adaptive charac­ ter, easily developed anew (occurs in such widely diverging groups as Hexactinellida, Calcarea, Spongosorlles Homosclerophorida, Poecilosclerida, Haplosclerida and Keratosa). 26. St_)'les with t_ylott swellings: a synapomorphy of Topsenlia H)'meniacidon.

Epipolasis

Petro mica RESULTS Amorph!nops!s Treatment of the taxon-character matrix by Clocalypta the computer programs yielded several equally parsinwnious trees, which differed however Hymenlacldon only marginally from each other. Among them N was one tree which was identical to the one • Halichondria tnade without computer assistence and it was decided to adopt this as the most likely Fig-. 13. Phylogenetic relationships of the valid g-enera of the newly defined family Halichondriidae; numbers with representation of' phylogenetic relationships of black boxes refer to synapomorphic characters listed and the Halichondrid genera and the outgroup­ discmsed in the text. f'amilics. The inferred relationships between the genus groups which have been prelin1inary suggested already in a previous paper (sec Diaz, eta!., in dria). S'jJongosorites deviates rather strongly from press), arc here Connally presented in the the others in architecture and chemistry. cladogram of Fig. 13. In short, the Halichon­ driidae consist of three major genus groups: one DISCUSSION comprising two genera which are characterized by a finely conulose surface, and which retain Although a formal analysis of relationships still an 'Axinellid' type of collagenous within the whole order Halichondrida s.l. has choanosomc; furthermore two genera with not yet been completed it is proposed here to characteristically grooved surfaces and heavy adopt above presented results into a fonnal cctosomal protective skeleton; they, too, retain classification of the order into four families, one a collagenous mesohyl; and, finally, a group of of which (Desmoxyidae) may turn out to be eight genera which have lost the collagenous Inore related to Raspailiids. It is furthcrn1ore nature of the organic choanosomc and are char­ proposed to assign Raspailiids, Rhabcleremiids, acterized by having son1e sort of ectosomal and Euryponids for the time being as a discrete tangential crust. Relationships within the latter group to the Poecilosclerida (partly in accord­ group are further determined by details of the ance with suggestions n1adc by Hooper, 1990), ectosomal crust, which may consist of a but the relationships with this group as well as para tangential arrangement of spicules ( Topscn­ with Age/as have to be reevaluated. Trachycla­ tia, Pctromica), a reticulation of spicule bundles clids arc considered Hadromerida on account of (AmorjJhinopsis, CiocalyjJta) or feltwork (EjJipo­ similarities of their n1icroscleres with spirasters. lasis), or single spicules (Hymeniacidon, Halichon- Below, the recognized genera and their (sub-

24 jective) synonyms will be defined. In a forth­ small styles, however, is regarded as a coming monograph of the West Indian synapomorphy of Amorphinopsis. Halichondriidae (see Diaz et al., in press) the new classification will be adopted at the species level. A REVISION OF THE GENERA Characters considered to be of low value as PRESENTLY ASSIGNED TO THE synapornorphics (if used are considered 'weak') HALICHONDRIIDAE or assumed to be homoplasic are: Family Halichondriidac Vosmaer, 1887 Shape: rnost Halichondriidae tend to be 'amor­ (emended) phous', connected no doubt to the low grade of skeletal organization. Certain characteristic Halichondrida with a choanosonwl skeleton shapes, such as fistulose forrns with a n1assive consisting of a high density of spicules arranged base buried in the sediment are definitely in vague, ill-defined, directionless tracts and of hon1oplasic, as these habits do not concur with spicules in confusion. other character distributions, and tnorcover are also known in several other Demospongiae Genus group A.rinJ'SSa-Collocal:}jJta: orders (e.g. Oceanapia in Haplosclcrida and Halichondriidac with choanosomal spicule Coe!mphaera in Poecilosclerida). Striking tracts protruding slightly beyond the organic sin1ilarities between "Cl.oca(pjJ!a"-likc forms in ectosome producing a finely conulose surface different genera ( Cioca/yjJ!a, Topsentia, Halichon­ pattern. dria, Hymeniacidon, Petromica, Collocalj,pta) (sec e.g. Wells el al., 1960: Figs. 47, 49, 50, 51) are Genus Axin.yssa Von Lendcnfeld, 1897: 116 here considered the product of environn1ental Figs. 14-22 pressure on growth form and general low grade Type species (monotypy): A. tvpsenti Von skeletal organization in the Halichondrids. Lendenfcld, 1897: 116, pl. 10 figs. 134-144 Spicule morphology: the spicules in this (Holotype ZMB 2971 (Fig. 14), reexamined; group share with their outgroups an 'indif­ holotypc fi·agmcnts in BIVINH 1908:9:24:145 ference' with regard to the exact shapes of the (Fig. 17) and slide 1897:3:25:70 reexamined). spicules, especially the apices, but also the cur­ vature and axial canal width. These characters Definition (emended): Halichondriidae of the arc assumed to be partly ancestral (plesimnor­ genus group Axin)'SSa-Collocal)pta with vague phic), partly easily developed independently. choanosornal tracts ending at the surface in Etnphasis on stylote vs. oxeote nature of the regularly distributed spicule brushes producing spicules is only made occasionally (e.g. in a regularly spaced conulation. distinguishing AmorphinojJsis and Cioca()lpta, or Halichondria and lijmzeniacidon) and considered Remarks: This is a cornmon genus in tropical 'weak'. In fact, it is not so 1nuch the nature waters. The type species has smaller stylotc itself of the spicules but rather the absence of oxcotes in the surface brushes, and this has led one or more of the usual forms we are using. authors in the past to recognize a separate Spicule n1alformations and centrotylote condi­ genus (Pseudaxinyssa) for species similar to A. tion are found in several groups within and topsenti, but without clearly differentiated outside the Halichondriidae, and it does spicule size categories. In view of the cmnmon apparently not point to close relationship. The occurrence of these in various Halichondrid frequent occurrence In some, e.g. In genera, this is considered superfluous. SjJongosorites is however noteworthy. Spicule size The well known North Australian Pseudaxinyssa categories seem to occur in most genera and can stipitala Bergquist & Tizard, 1967 is not an only be succcsfully employed as apomorphy in Axinyssa but probably belongs to an Axinclid distinguishing species. A special category of g~nus close to Axinella.

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26 Synonymy: Several species assigned to the genus Leucophloeus belong to Axi1ryssa; however, the type species of that genus is Genus Pseudaxinyssa Burton, 1931:350 assigned to Ciocalypta. (Figs. 16, 18) The genus name Rhaphisia Topsent, 1892, has been used Type species (original designation): Axin;•ssa teth;·oides in connection with Axinyssa specimens by some authors Kirkpatrick, 1903: 245, pl. V fig. 12, pl. VI fig. 8 (holo· (e.g. De Laubenfels). The type species of Rhaphisia type reexamined) (monotypy), viz. R. laxa Topsent, 1892, ho\vever, is a Burton: "Axinellidae of massive form in which the clear Haliclona (see De Wcerdt & Van Soest, 1986; De skeleton is composed of vertically ascending fibres of oxea Weerdt, 1987). of one sort only, the fibres branching and anastomosing The genus name Ph)'copsis Carter, 1882, has been used for throughout their length and ending at the surface in small at least one species of Axhryssa, but the type of that genus, tufts which project to a greater or lesser extent''. P. hirsuta Carter, 1883, is a spinous Raspailiid. Remarks: The holotypc specimen of Kirkpatrick (Fig. 18) is incorporated in the collections of the British Museum (reg. no. 1902:11: 16:25), and has been reexamined. It has a dendritic reticulation of spongin-bound (?) tracts. In Species examined: Axilryssa topsenti Von many ways the specimen is quite unusual: it is black, has Lendenfcld ( 1897) (holotypc m the Berlin a coarse organic skin, large quantities of choanosomal organic (?foreign) material; the spicule tracts arc often Museum (Fig. 14), fragments in BMNH (Fig. quite discrete and seem to be unattached to the organic 17)), A. tethyoides Kirkpatrick (1903) (holotype parts. Possibly, the specimen is not a healthy represen­ in BMNH (Figs. 16, 18), A. gravieri Topsent, tative of the species as no other, similar specimens have 1893 (holotype, MNHN DT. 1873), A. aculeata ever been described. Wilson (1925) (holotype USNM. 21341 (Fig. Genus Axinomimus De Laubenfels, 1936: 163. 19) and ZMA specimens from Indonesia), A. (Fig. 22) aplysinoides (Dendy, 1922 as Halichondria (Fig. Type species (monotypy): Axinella paradoxa Ridley & 16)) (holotype BMNH.1921:11:7:31 and ZMA Dendy, 1886:482, 1887: pl. XXV frg. 4 (holotype reex· specimens from Indonesia (Fig. 20)), A. amined). De Laubenfels: "The spicules are oxcas only ambrosia (De Laubenfels, 1934 as Raphisia) and the construction is so nearly radiate that inclusion in this family (i.e. Sollasellidae) seems justified, but it (holotype USNM. 22445, holotype of deviates from typical members of the family in the direc­ Leucophloeus lewisi Van Soest & Stentoft, 1988 tion of the family Axinellidae in as much as the spicules ZMA 5401 which is now considered a junior arc more or less vaguely arranged in plumose tracts. The synonym; holotype of Dictyonellayumae Pulitzer· rubbery consistency is very noteworthy." Finali, 1986 Mus. Genova CE 47694; ZMA Remarks: The type specimen BMNH 87:5:2:68 from near Tristan da Cunha in the South Atlantic is a rubbery specimens from Cura<;ao (Fig. 21) and Puerto lobate mass with conulose surface; no cctosomal skeleton; Rico)), A. fenestra/us Ridley, 1884 (holotype, choanosomal skeleton consists of confused loose spicules BMNH 82:2:23:253; ZMA specimens from arranged mostly in vague radiating tracts; oxea on Indonesia), A. valida (Thiele, 1899 as Phycopsis) average 870 by 22 [J.m. (holotype, Mus. Basel no. 154; ZMA Several species assigned to Trach;·opsis arc here considered specimens from Indonesia), A. radiata (Levi & members of Axin)'ssa; however, the type species of that Levi, 1983 as Prostylyssa) (holotype fragment genus is considered a Topsentia. MNHN DC!. 2954 examined). Geographical

Figs. 14-16. Fig. 14. Axinyssa topsenti Von Lendenfcld, 1897, A. Habit (redrawn from Lendenfeld, 1897: pl. 10 fig. 134) (s.b. = 1 em), B. longitudinal section of one of the oscular cones to show skeletal architecture (s.b. = 1 mm), C. tangential view of ectosome showing spicule brushes and single oxea (redrawn from Lendenfe1d, 1897: pl. 10 fig. 144) (s.b. = 100fJ.m), D. spicules (s.b. = 100 fJ.m). Fig. 15. Axi?U'ssa aplysinoides (Dendy, 1922), A. Habit (redrawn after Dendy, 1922: pl. 3 fig. 3) (s.b. = 1 em), B. Cross section showing confused spicule tracts with ill-developed spicule brushes (s.b. = 1 mm), C. spicules (s.b. = 100 fJ.m). Fig. 16. Axhryssa tetlu'oides Kirkpatrick, 1903 (type of Pscudaxinyssa), A. skeletal architecture (redrawn from Kirkpatrick, 1903: pl. 6 fig. 8) (s.b. = 1 mm), B. peripheral endings of choanosomal spicule tracts enveloped in spongin coat (s.b. ~ 0.5 mm), C. oxca (s.b. ~ 100 ~m).

27 28 distribution: probably confined to the tropical Genus group Myrmekioderma-Didiscus: regions of the three oceans. Halichondriidae with sinuous surface grooves and ectosomal skeletal strengthening of Genus Coliocalypta Dendy, 1905: 199. palissade-1ikc arranged stnaller spicules erected Fig. 23 on a crust of tangential larger megascleres. Type species: Col/oca/yjJ/a digitata Dendy, 1905: 199, pl. VII fig. 6, pl. XIII figs. 1-2. (Holotype Remark: The remaining Halichondriid genera BMNH 1907:2:6:89 reexamined). arc all characterized by an ectosmnal strengthening skeleton; in most this takes the Definition: Halichondriidae of the genus group form of a tangential spicule arrange1nent Axinyssa-Coliocalypta, with Ciocalypta-like (which may occasionally be absent, e.g. in some fistulose habit and architecture but without Topsentia or Haiichondria), while in the present ectosomal skeleton and with highly collagenous genus group species have variously arranged choanosomc. crust-like ectosomes, which consist at least partly of semi-erect (non-tangential) spicules on Remarks: So far only two species have been top of a layer of tangential spicules. assigned to this genus, viz. the type species and C. aderma Levi & Vacelet, 1958. Some doubt as Genus Myrmekioderma Ehlers, 1870:32. to the generic distinctness of Colioca!ypta and Figs. 24-30 Ciocalypta seems justified. The central column of spicules with diverging spicule tracts carrying Type species (monotypy): Alcyoniwn granuiatum the ectosome is present in both. Conceivably Esper, 1794: 71, pl. XXIV (holotype not the present species are Cioca!ypta having lost the examined, possibly lost). ectoson1al skeleton (which also is assumed to have occurred in some Halichondria species). On Definition (emended): Halichondriidac of the the other hand, the fistular habit seems a clear genus group 111yrmekioderma-Didiscus with one or adaptation to life on sandy bottmn and there more categories of trichodragmata, the larger of may well be no other way to build a fistulc with which are sinuously curved. the confused spicular arrangement found in the Halichondriidae. Hence the occurrence of Remark: As a distinguishing feature of the similar fistules in unrelated Petromica, Coelo­ genus the possession of an acanthose smaller ca/yjJia ( ~ Topsentia) and Ciocalapata ( ~ category of rnegascleres has been traditionally Halichondria). used. However, recent studies of the variability of the acanthose condition of these spicules in Species examined: C. digitala (holotype and the type species and in Caribbean represen­ ZMA specimens from Indonesia). tatives has revealed that the spination may be Geographic distribution: India, Indonesia, weak or even absent in 1nany individuals. This Eastern Atlantic. led to the assignment of species of the genera

Figs. 17-21. Fig. 17. Axinyssa topsenti Von Lendenfcld, 1897, BMNH type fragment 1908:9:24:145 from Zam~ibar, photographed from outer and inner side (s.b. = 0.5 em). Fig. 18. Axit~yssa tethj•oides Kirkpatrick, 1903, holotype BMNH 1902:11:16:25 from the Natal coast (s.b. = 1 em). Fig. 19. Axinpsa aculeafa VVi!son, 1925, holotype USNM 2134-1 from the Philippine region (s.b. = 1 em). Fig. 20. Axiru•ssa aplysinoides (Dendy, 1922), ZlviA specimen from Indonesia (s.b. = 1 em) (photo H .. van Brandwijk). Fig. 21. Axinyssa ambrosia (De Laubenfels, 1934), ZMA specimen from Cura<;ao (s.b. = 1 em) (photo H. van Brandwijk).

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30 Anacanthaea and Epipolasis (several species only be abnormally developed spicules, polyaxone pro­ except the type species) to Myrmekioderma. liferations of malformed, crooked oxea. The British :Museum (Nat. Hist.) holds a spicule slide preparation Several species of A1yrmekioderma reveal the with a label in Schmidt's handwriting. The slide contains presence of quantities of spongin, which may oxeote spicules in three categories: 443-680/12-15 [lll1, cause the spicules to be arranged in irregular 196-320/4-9 (J.m, and 80-121/1-2 [J.rn; there. arc a few thick tracts. malformations, blunt or stair-stepped apices with \vide axial canals, etc. No rhaphides have been found, and coupled to the three size categories of oxea the most likely Synonyms: genus fur it would be Topscntia. However, in the absence Genus Acanthoxifer Dendy, 1905: 15 7. of a type specimen (Topsent, 19,15; Desqueyroux, pers. Type species (monotypy): A. cqloncnsis Dendy, 1905: 15 7, comm.) it is proposed here to consider the genus and the pl. IX fig. 5 (holotype not examined). species inccrtac scdis, although Tsurnamal (1969) claimed Sec for details of this synonymization: Bergquist, 1965: the rediscovery of the species in lsraelian waters. Tsur­ 177. namal's material (Fig. 26) clearly belongs to the genus ;llyrmckioderma through its possession of a confused Genus Anacanthaca Rowe, 1911: 329. skeleton of oxeote spicules of three size categories, a com­ (Fig. 25) plement uf Jlexuous rhaphides, and a palissade-like Type species (monotypy): A. nivca Rowe, 1911: 329, pl. 38 arrangement of smaller oxea in the ectosomal crust. It is fig. 17, text-fig. 14 (holotype fragment BNINH reg. no. assumed here, that Tsurnamal's material belongs to 1912:2:1:54 reexamined) ;lfpmekiodcrma spclca (Pulitzer-Finali, 1983). If Schmidt's Rowe: ''Heteroxyinae in which the oxea are not differen­ species would at any time be proven to be identical to this tiated into two forms, and without spines on them. There species, than ;\ipmckiodama Ehlers, 1870 will become a are no microscleres. '' junior synonym of Callites Schmidt, 1868. Remarks: The external surface of the upright-lamellar specimen has the characteristic grooves (Rowe's pl. 38 fig. ?Genus Hctcro.ya Topsent, 1904: 133. 17) making polygonal areas; brushes of smooth oxea form (Fig. 27) a cortical skeleton; oxea are 350-430 by 6-10 [J.m with Type species (monotypy): II. cortirata Topsent, 1904: D3, stair-stepped apices; not mentioned in the original des­ pl. I fig. 15, pl. XU Jig,. 22-23 (holotype slide MNHN cription but nevertheless present are two sizes of DT. 905 reexamined). trichodragmata (20 and 110 [lm), the larger of which is Topsent: "Tethyidae rev&tantes, sans microsclCrcs, a sinuous. choanosome a peu pres aspiculeux, a ectosome en revan­ che diffCrencie en une ecorce solide armCe d'oxes de deux ?Genus Callitcs Schmidt, 1868: 16. sortes, disposCs verticalement: les uns tres nombrcux et (Fig. 26) serres, les autres, solitaires, dCterminant le hispidation de Type species (monotypy): Cal!ites lacazii Schmidt, 1868: Ia surface''. 16, pl. III fig. 2 (micmscopic slide in BrvlNH reex­ Remark: no mention is made of surface grooves; no amined). trichodragmata have been found in the slide; the holotype Schmidt: "Eine auf dcr Grcnze zwischen den Compagi­ (Fig. 27) is described as a hispid plaque of 2 mm thick; neen und den Gummineen stehende Gattung, wo die oxca are 1600-2000 by 35, and acanthoxca are 235-420 by Sarcode-Substanz ein h6ehst unrcgelmiissiges Geriist oder 11-23 [J.Ill. Conclusion: reexamination of the holotype is Fachwerk, auch Giinge und R6hren bildet, jedoch keine necessary. die Gummineen charakterisierende Rindenschicht. Es sind Kieselgcbilde vorhanden''. Genus NcoftTOS)'fta De Laubenfels, 1954: 189. Remarks: The identity of Schmidt's species remains Type species (monotypy): N. atina De Laubenfels, 195,1: uncertain, because the description focusses on what can 190, fig. 127 (holotype USN!vl 22974 reexamined).

Fig,. 22-23. Fig. 22. Axinella ( = Axii1)1Ssa) paradoxa (Ridley & Dendy, 1886), A. habit (redrawn from Ridley & Dendy, 1887: pl. 25 fig. 4) (s.b. = 1 em), B. tangential view of ectosome (s.b. = 0.5 mm), C. cross section of peripheral region (s.b. 0.5 mm), D. spicules (s.b. = 100 [J.m) (B-D made from the BMNH type specimen). Fig. 23. Collocal)'}Jta digitata (Dendy, 1905), A. habit (redrawn from Dendy, 1905: pl. 7 fig. 6) (s.b. = 1 em), B. cross section of peripheral region (s.b. = 1 mm), with as inset a typical spicule (s.b. = 100 (.lin) (both redrawn from Dendy, 1905: pl. 13 figs. 1-2), C. longitudinal section of apex offistulc to show architecture (s.b. = 0.5 mm), D. spicule apices (,.b, ~ 50 ~m).

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32 Sec for details of this synonymization Bergquist, 1965: Type species (monotypy): D. p!acospongioides 177. Dendy, 1922: 135, pl. 7 fig. 10, pl. 18 fig. 3 (holotype BMNH 11 : 7: 112 reexamined). The genus Raspaz;~tella Schmidt, 1868 has been used for species of i\1)'Tmekioderma, e.g. by Burton (1959) and Vacelet & Vasseur (1971), but the type species (mono­ Definition (emended): Halichondriidae .of the typy), R. brunnea Schmidt, 1868, is a HaLichondria (see also Myrmekioderma-Didiscus genus-group with dis­ there). corhabds. The genus E'pipolasis De Laubenfels, 1936 has been used for species of Myrmekioderma, e.g. by Dfa:-: et all. (1987) and Van Soest & Stentoft (1988), but the type species (original Remarks: The discorhabds tend to be arranged designation), Spongosorites suiuensis Wilson (1925) has uni­ peripherally perpendicular to the surface; this que characters which make it necessary to uphold it as a arrangement is assumed to be homologous to separate genus (see also there). that of the palissade of small oxeotes of Species examined: Myrmekioderma granulata Myrmekioderma. The genus has been associated (Esper, 1794) (ZMA specimens (Fig. 28) from with Latrunculia (Hadromerida) on account of Indonesia; holotype of Neoprosypa atina), M. styx the similarity of the microscleres. This De Laubenfels, 1953 (holotype USNM 23400; similarity is non-homologous, the 'discorhabds' HBOI specimens (Fig. 24); ZMA specimens of Latrunculia are streptasters/spirasters, while including the holotype (Figs. 29-30) and those of Didiscus can be traced back to cen­ paratypes of Epipoiasis reiswigi Dfaz et ai. 1987 trotylote microxea. In some cases n1icroxea of a which is a junior synonym of M. styx), M. rea De size equal to that of the discorhabds and Laubenfels, 1934 (as Anacanthaea) (holotype ornamented with small spines are found as USNM 22301; HBOI specimens; ZMA abnormal variations of the discorhabds; these specimens), M. tulearensis (Vacelet, Vasseur & are quite similar to acanthose microxea found Levi, 1976) (holotype MNHN.D.JV.20), M. in Myrmekioderma, though much smaller. dendyi (Burton, 1959) (ZMA specimens from For a review of all species and a phylogenetic Indonesia), M. spelea Pulitzer-Fin ali, 1983 (as analysis of the genus see Hiemstra & Van Soest Raphisia) (holotype in Mus. Genova PTR D.18; (in press). Aegean specimens collected by E. Voultsiadou­ Koukoura). Synonyms: none. Geographical distribution: tropical and sub­ tropical parts of all three oceans, Mediterra­ Species examined: D. piacospongioides Dendy, nean; not known from the East Pacific. 1922 (holotype, Fig. 31), D. aceratus (Ridley & Dendy, 1886 as Latruncuiia) (ZMA specimens Genus Didiscus Dendy, 1922: 135 from Indonesia), D. oxeata Bechtel, 1983 (halo­ Figs. 31-32, 36-40 type YPM 8969 from Brazil; holotype fragment

Figs. 24-27. Fig. 21. Jvf.yrmekiodenna styx De Laubenfels (1953), A. cross section of peripheral region to show ectosomal palissade (redrawn from Dfaz et ai., 1990a) (s.b. = 1 mm), B. megasclcres (s.b. = 50 [lm), C. sinuous trichodragmata (s.b. 25 [lm) (B-C after Van Soest & Stentoft, 1988: fig. 43). Fig. 25. Anacanthaea (= 1\1yrmekioderma) nivea Row, 1911, A. habit (redrawn from Row, 1911: pl. 38 fig. 17) (s.b. 1 em), B. megascleres (redrawn from Row, 1911: fig. 14), C. sinuous trichodragmata (found in the BMNH type specimen) (s.b. = 25 [lm). Fig. 26. Ca!Lites Lacazei sensu Tsurnamal, 1969 ( = probably A1yrmekioderma speiea (Pulitzer-Finali, 1983), A. rnegascleres (s.b. = 100 [lm), B. rhaphides (s.b. = 10 11m), C. malformed spicules (s.b. = 10 [lm) (all redrawn from Tsurnarnal, 1969: fig. 26). Fig. 27. Jletero.>:J'a (? = .M)'rmekioderma) rortirata Topscnt, 1904, A. habit (s.b. = 1 em), B. cross sec Lion of peripheral region (s.b. = 0.5 mm), C. acanthoxea (s.b. = 100 f.I.In) (all redrawn from Topsent, 1904: pl. 1 fig. 15, pl. 12 figs. 22-23).

33 Figs. 28-30. Fig. 28. A1yrmekiodenna granulata (Esper, 1794), ZMA specimen from Indonesia (s.b. = 1 ern). Fig. 29. Epipo!asis reiswigi Dfaz et al., 1987 ( = 1Hyrmekioderma Sf)'X De Laubenfels, 1953), holotypc in situ in reefs of Los Roques, Vcnc:wela (photo E. VVcil). Fig. 30. 1\1pmekioderma styx De Laubenfels, 1953, SE1v1 photo of choanosomal skeletal arrangement (photo F. Hiemstra).

of D. habanensis Alcolado, 1984 (junior anisodiscus Vacelet & Vasseur, 1971 (ZMA synonym) in USNM; holotype (ZMA 4889) specimens from Indonesia) (Fig. 40), D. (Figs. 36-39) and paratypes (ZMA 4890-4892) styliferus Tsurnamal, 1968 (holotype from Tel­ of D. flavus Van Soest, 1984 (junior synonym); Aviv Museum, Mediterranean specimen of HBOI specimens from Jamaica (Fig. 32), ZMA Pulitzer-Finali, 1983 examined m Mus. specimens from Cura~ao and Colombia), D. Genova), D. verdensis Hiemstra & Van Soest, in

34 press (holotype and paratypes in ZMA from compound apparently confined to the genus. Cape Verde Islands). Geographical distribu­ Unfortunately, the specimen upon which the tion: tropical and subtropical parts of all three first report of topsentin (Bartik eta/., 1987) was oceans; not known frmn the East Pacific; occur­ based, was erroneously identified as a Topsentia. rence in North Australia (Pulitzer-Finali, 1982) Those true Topsentia species examined so far, is doubtful. did not reveal the presence of topsentin, The phylogenetic relationships of the genus arc Genus group Spongosorites- Topsentia-Epipoiasis­ uncertain; it stands rather isolated through lack Petromica- Ciocalypta- Amorphinopsis- Halichondria­ of distinct synapomorphies with other genera. Hymeniacidon: Halichondriidae which have lost the col­ Synonyms: lagenous nature of the choanosomc. Genus Aponastra Topsent, 1927: 5 (Fig. 35) Genus Spongosorites Topsent, 1896: 117. Type species (monotypy): A. dendJ•iTopscnt, 1927:5 (type Figs. 33-35, 41-44 slide reexamined). Topsent: "Spongosoritidae. a ectosome differenciC en Type species (monotypy): S. placenta Topsent, Ccorce, a charpente choanosomique grossiCrement fascicu!Ce, irrCgulihe, pourvue d'oxes et de microrhabdes 1896: 117 (type slide MNHN DT.904 reex­ plus ou mains centrotylotes, ces derniers parscmant la amined). chair et s'accumulant dans la couche externe de l'Ccorce en un feutrage tangentiel, a la fa~on de ceux de Definition (emended): Halichondriidae with a l'ectochrote des El)•!us." smooth, flaky crust of paratangentially Remarks: the type species (Fig. 35) is described as black, and has spicules 500 by 16-20 and 45-145 by 2-7 {lm. A arranged, relatively thin spicules, and a slide of the type specimen has been examined (iviNHN choanosomal utterly confused skeleton which is D.T. 2865) and it conforms completely to SjJongosorites, traversed by spongin-enforced tracts running possibly to S. placenta. more or less parallel to the surface. Most species show an acrophobic colour-change from yellow Species examined: S. placenta Topscnt, 1896 to greyish brown or black. (type slide and topotypical specimen from the North Atlantic, in MNHN) (Fig. 33), S. Remarks: The emended diagnosis is based on intricata (Topscnt, 1892) (specimens identified reexamination of the type slide in MN HN by Pulitzer-Finali, 1983; specimen in ZMA (D.T.904) and on a specimen of the MNHN from Norway), S. ruetzleri (Van Soest & Stentoft, collection identified by Prof. Levi; the type 1988) (as 'Halichondria) (type specimens in specimen (Fig. 33) itself seems to be lost. ZMA, HBOI specimens, Fig. 34)), S. siliquaria Emphasis in the past has been put to the fre­ Van Soest & Stentoft, 1988 (type specimens in quent malformations or ccntrotylotc conditions ZMA, Figs. 42-44), S. spec. (ZMA specimen in the smaller category of oxeotes, but this is not from Indonesia), S. n. spec. (HBOI specimens, a consistent character; moreover, it is also Fig. 41). infrequently found in other genera, e.g. Topsen­ Geographical distribution: tropical, subtropical tia. For that reason, Spongosorites has often been and temperate regions of all three oceans. considered a senior synonym of the latter genus (e.g. a MNHN specimen identified by Topsent Genus-group Topsentia-Epipolasis-Petromica: himself asS. placenta, D.T.1116, is a Topsentia). However, the two are clearly distinct in Halichondriidae without any spongin resulting architecture, spicule sizes and shapes, and in an utterly confused arrangement of spicules. especially in the acrophobic properties. All chemically examined species reveal the Genus Topsentia Berg, 1899: 77. presence of topsentin (Fig. 34B), a biochemical Figs. 45-55

35 31

{.

'

c

(~ l 34 0 ' 1\ N (Q NH NH

36 Type species (original designation): Anisoxya Synonyms: g/abra Topsent, 1989: 234 (type slide MNHN Genus ALlosclcria Topsent, 1927: 6. DT.1169 and topotypical specimen examined). (F;g. 47) Type species (original designation); A. tcnuisjJinosa Top­ Definition (emended): Halichondriidac of the sent, 1927: 6 (type slide 1\1NHN D.T. 1190 reexamined). 'I'ojJsentia-Ej;ipo!asis-Petromica genus-group with Topsent: "Spongosoritidac a spicules de deuX sortes, les uns monactinaux, plus grands, representant les CJCments an utterly confused ectosomal crust. principaux, d'une charpente vaguemcnt halichondroidc, ]fiche, sans sponginc, les at1tres diactinaux, de petite taille Remarks: This is a decidedly brittle-siliceous ct de rOle accessoire, l:pars et nombreux. '' type of sponge, which is also characterized by l'opsent claims there arc acanthoxea among the small oxeotes (65-120 by 1-2 [J.m), but we could not find these lacking bright colours (one bright yellow species in the type slide; the monactinal spicules are more excepted), dirty rosy white or greenish drab col­ definitely strongyloxea (350-500 by 7-10 [J.m). Later ouration being favoured. Many representatives ( 1904 ), Topscnt assigned some specimens of the type also have lost their ectosomal crust-like series of Topscntia glabra to the present' 'species"; this con­ specialization (no doubt due to absence of fusing action appears unnecessary as both "species" are probably synonymous. distinct smaller size categories of mcgascleres), in stead of which they have an utterly confused Genus Aniso.\)'a Topsent, 1898: 234 (type slide examined, peripheral skeleton (Fig. 45) with individual MNHN D.1'.1169). spicules sticking out for a short distance produc­ Type species (monotypy): A. glabra Topscnt, 1898: 234. ing an optically smooth, but rough-to-the-touch Preoccupied: this is an objective junior homonym of surface. The absence of a distinct crust is here Aniso."::ya f..'Iulsant, 1856. n1erely interpreted as independent loss. Occa­ Genus Trach)'opsis Dendy, 1905: 147. sionally, spicules maybe sinuous or twisted, (Fig. 46) showing similarities with non-Halichondriid Type species (monotypy): T. halichondrioidcs Dendy, 1905: genera as Bubaris and Dactylella. Occasionally, 147, pl. X fig. 10 (holotype reexamined). the smaller category of (peripheral) oxea is ccn­ Dendy: "Rcnierinae in which the main skeleton is com­ trotylote or malformed as is found quite com­ posed of a dense irregular network of oxea, -...vhilc the sur­ monly in Spongosorites. The present genus has face is protected by similar (or perhaps more slender) ~picule~ arranged in dense vertical brushes, which support indeed been confused or synonymized quite the pore-bearing membrane.'' often with the latter genus, but both are clearly Remarks: The type specimen (BMNH reg. no. distinct morphologically as well as bio­ 1907:2:1:44) has been reexamined and found to be like chemically (see under Spongosorites). Topscntia, but lacking a clear categ01·i:r.ation of the short TojJSentia differs fi·om the closely related genus and thick oxea, all oxea being more or less dose to 600-700 by 30-35 [J.m. The consistency is hard and compact, due Epipolasis in the absence of rhaphides, and in to the extremely dense and irregular arrangement of the the presence of unique secondary 1netabolites oxca in the choanosomc; the peripheral regions are (Pomponi cl a/., 1990). relatively \veil-organized. Several other species have been

Figs. 31-35. Fig. 31. Didiscus placospongioides (Dendy, 1922), A. habit (s.b. = 1 em), B. discorhabds in various stages of development (s.b. = 25 (llll) (all redrawn from Dendy, 1922: pl. 7 fig. 10, pl. 18 fig. 3). Fig. 32. Didiscus oxcata Hechtel, 1983, A. cross section of peripheral region to show ill-developed ectosomal palissade (s.b. = 0.5 mm), B. discorhabd (s.b. = 50 [J.m) (all redrawn from Dfaz ct a!., 1990a). Fig. 33, SjJOngosoritcs placenta Topsent, 1896, A. habit (s.b. = 1 em), B. spicules (s.b. = 100 (lm) (all redrawn from Top>ent, 190>1: pl. 10 fig. 6, pl. 12 fig. 21). Fig. 34. Spongosorites ruetzlcri (Van Soest & Stentoft, 1988), A. cross section of peripheral region to show directionlcss spicule tracts (redrawn from Dlaz et al., 1990a) (s.b. = 0.5 mm), B. topscntin (redrawn from Bartik et al., 1987). Fig. 35, Aponastra ( = Spongosorites) dendyi Topsent, 1927, A. habit (redrawn from Topsent, 1928: pl. 3 fig. 24) (s.b. = 1 em), ll-C. >pkulc> (>.b. ~ 100 ~m).

37 assigned to this genus, but only the type species conforms SjJOngosorites. However, spicules sizes arc a bit on the large to Topsentia. side: 320-400 by 5 and 400-800 by 12 jJ.m, and therefore we assume that this is a Topsentia. Genus Coeloca!ypta Topsent, 1928: 167. (Fig,. 48, 53) Genus Laminospongia Pulitzer-Finali, 1983: 546. Type species (monotypy): C. porrecta Topsent, 1928: 167, (Fig. 50) pl. II fig. 6, pl. VI fig. 4. Type species (monotypy): L. subtilis Pulitzer-Finali, 1983: Topsent: "Axinellidcs sans microscleres, composCcs 547, fig. 59, 59bis. Holotype: Genova Museum reg. no. comme les Cioca!ypta d'un corps massif, de structure 47175 (reexamined). halichondroide, d'oU s'elevent des processus digitiformes Pulitzer~Finali: "Hymeniacidonidae of laminar form, aquiferes. Ces processus sont crcux; leur paroi, faite sur~ with a halichondroid skeleton of oxeas to styles variously tout d'une couche relativement Cpaisse aU des spicules modified''. libres se placent dcbout tres pres les uns des autres, est Remarks: The consistency of the type specimen is des­ soutenue par une assise interne spiculofibreuse, plus cribed as firm, it has a Halichondria-like but not separable mince, equivalcnte a la colonne axiale de ceux des ectosomal skeleton, and the oxea-stylcs-strongyles of 350- Ciocaiypyta." (Fig. 48) 1300 by 9-19 jJ.m are in confusion; some spongin is found Remarks: The type species resembles Cioca!ypta penicillus, binding single spicules. although it is more solid and the 'fistules' are thicker (Fig. 53); the colour of topotypical specimens collected recently appeared to be strikingly yellow. The oxea of the holotype Species examined: Topsentia glabra (Topsent, are 280-1000 by 4-50 fJ.m; specimens from the Caribbean 1898) (Fig. 47) (type slide, topotypical assigned to this species (Van Soest & Stentoft, 1988) have specimen MNHN DT.1389), T. ophiraphidites a larger upper limit to spicule size; they arc not con­ (De Laubenfels, 1934) (Figs. 45, 51-52) (bolo­ specific. All known specimens have oxeotes as the only spicules. Some doubt exists over the generic differences type USNM 22334; holotype (FCLR col!. no. between this species and the genus Cioca!ypta, because the 125 from V cnczuela) and paratypes (ZMA habit and the internal fistule architecture of the two are 5839-5840 from Cura<;ao) of T. roquensis Diaz et similar. The view is taken here that both features are sub­ al., 1987, which is now considered a junior ject to environmental demands and constitute convergent synonym; holotype of Spongosorites sinuatus developments (compare also Petromica, Ciocalapata and Cioxeamastia). Pulitzer-Finali, 1986 Mus. Genova no. 47961 which is considered a junior synonym; holotypc ?Genus Oxcostilon Ferrer Hernandez, 1922: 10. YPM 8988 of Halichondria braziliensis Bechtel, (Fig. 49) 1983 frmn Brazil, which is considered a junior Type species (monotypy): 0. annandale/ Ferrer Her­ synonym), T. porrecta (Topsent, 1928 as nandez, 1922: 9, figs. 8-9, pl. I (see Fig. 49) (no material examined). Coeiocaiypta) (type in MNHN and topotypical Ferrer-Hernandez: "Axincllidas con esqueleto formado specimens in ZMA (Figs. 48, 53)), T. n.sp. por fibras cspiculosas ascendentes, que terminan en Dlaz et al., in press (Caribbean specimens iden­ pinceles de espiculas normales a la superficie y Ia cual tified as T. porrecta by Van Soest & Stentoft, atravicsan. Oxeas predominantes y dobladas en angulo y 1988 in ZMA; HBOI specimens from Bahamas estilos probablcmcnte derivados de aquellas par reduccion total o parcial de una actina o radio." (Fig. 54)), T. halichondrioides (type in BMNH Remarks: The specimen is described as a smooth plaque, (Fig. 46), specimens from Aldabra in MNHN with skeletal tracts. On that basis it is possible that it is a DC!. 361 ), T. dura (Lindgren, 1897 as Halichon-

Figs. 36-40. Fig. 36. Didiscus oxcata Hechtel (198:3), in situ (Cunu;ao reefs) photograph of surface showing grooves (s. b. = 0.5 em) (photo G. van 1\1oorscl). Fig. 37. Didiswsjlavus Van Soest, 1984 ( = D. oxeata Bechtel, 1983), holotype from Cura~ao reefs (s.b. = 1 em) (photo L. A. van der Laan). Fig. 38. Didiscus oxeata Hechtel (1983), SEM photo of choanosomal arrangement (photo F. Hiemstra). Fig. 39. Didiscus oxmta Bechtel (1983), SE1Vf photo of discorhabd (photo L. A. van der Laan). Fig. 40. Didiscus anisodiscus Vacelet & Vasseur (1971), ZMA specimen from Indonesia (s.b. = 1 em) (photo L.A. van der Laan).

39 40 dria) (ZMA specimens from Indonesia), T. trichodragmata) must be considered homo­ manaarensis (Dendy, 1905 as Spongosorites) (ZMA plasic or plesiomorphous. spccin1ens from Indonesia), T. hllbarioides (Levi The parchment-like skin resembles that of & Levi, 1983 as Spongosorites) (MNHN holotype Halichondria latrunculioides Ridley & Dendy, DC!. 2933), T. n.spec. (HBOI specimens), T. 1886, type of the monotypic genus Pyloderma subtilis (Pulitzer-Finali, 1983 as Laminospongia, Kirkpatrick, 1908 (sec Fig. 82b). However, the holotype, Fig. 50), T. aurantiaca (Schmidt, 1862 soft consistency with loose spicule tracts, and as Reniera), type specimen LMJG 15630 (Fig. absence of microsclercs, make it unlikely that 55) from Mus. Geneva. both arc congeneric. PJ'loderma is tentatively Geographical distribution: tropical and sub­ assigned to Halichondria. tropical parts of all oceans. Species examined: E. suluensis (Wilson, 1925 as Genus Epipolasis De Laubenfels, 1936: 162. Spongosorites) (type specimen, Figs. 56, 65), E. Figs. 56, 65 n.sp. (HBOI specimen from Bahamas). Geographic distribution: Philippine region, Type species (original designation): Spongosorites Bahamas. suluensis Wilson, 1925: 331, pl. 38 (fig. 8), pl. 48 (fig. 3) (holotype USNM 21297 reex­ Genus Petromica Topsent, 1898: 216. amined). Figs. 57-59, 66

Definition (emended): Halichondriidac of the Type species (monotypy): P. grimaldi Topsent, genus-group Topsentia-Epipolasis-Petromica with 1898: 216, 1904: 64, pl. 5 figs. 2-4, pl. 8 fig. 6 a parchment-like feltwork of tangential oxea, (Fig. 57) (holotype not examined). and (sinuous) trichodragmata. Definition (emended): Halichondriidae of the Remarks: So far this genus is known only by genus-group Topsentia-Epipo!asis-Petromica, with the type specimen of its type species and one sublithistid basal skeleton. West Indian specimen (Harbor Branch collec­ tion). The type is a llabellate specimen with Remarks: These are whitish massive or sides differing in ectosomal skeletal strengthen­ fistulose forms. The growth form of one of the ing: outer side with parchment-like skin, inner three species so far recorded, is similar to that ( oscule-bearing) side with palissade-like of Ciocalypta or to that of Topsentia porrecta and periphery. The West Indian specimen is there is an outside possibility that all these are massive and only shows the parchment-type closely related; hmvcver, the supporting ectosome. The genus has been misinterpreted ("axial") skeleton of the fistules of Petromica by Van Soest & Stcntoft (1988) and Dfaz el al. ciocalyptoides is found in the walls of the fistules (1987), who assigned lv!yrmekioderma species rather than in the centre. The confused spicula­ without acanthosc oxea to this genus; the tion of oxeotes, and or styles in combination similarities between the two (size range of large with desmata, however, is shared with non­ oxcotes, palissadc-like ectosmne, sinuous fistulose Petromica species.

Figs. 41-43. Fig. 41. SjJOngosorites ruetz!eri (Van Soest & Stentoft, 1988), Bahamas specimen from the Harbor Branch collection. Fig. 42. Spongosorites siliquaria (Van Soest & Stentoft, 1988), ZMA holotypc from Barbados (s.b. = 1 em) (photo L.A. van der I,aan). Fig. 43. !::i'pongosorites si!iquaria (Van Soest & Stcntoft, 1988), macroscopic SEM photo of surface (photoN. Stentoft). Fig. 44. Spongosorites siliquaria (Van Socst & Stentoft, 1988), detailed SEM photo of surface (photo N. Stentoft).

41 ' 45

[

'

46 47 ' 48

t A ~ c::::::::J <::::::::J c:::=:::J c:::=-­ 49 ~ c:=_j

42 Synonymy: Type species (monotypy): C. penicillus Bower­ Genus A1onanthus Kirkpatrick, 1903: 176. bank, 1864: 179 (holotype BMNH 1877:5:21: (Fig. 58) 1069 reexamined) (Fig. 60). Type species (monotypy): l\;f. jJlumosus Kirkpatrick, 1903: 176, pl. IV figs. 6-7 (Fig. 58) (holotype not examined). Definition (emended): Halichondriidae of the Kirkpatrick: "Desmanthidae in which the skeleton is formed of rnonocrepid dcsmas of the common type, genus-group Ciocalypta-Amorphinopsis, with separate or joined together, and of monaxon rncgasclcrcs. predominantly stylote megascleres and a Remarks: The family Dcsmanthidae shows architectural characteristic semi-transparent parchment-like, similarities with Bubaris (they are thin hairy crusts), but fistular habit. Alonanthus piumosus is described and figured (Fig. 58) as an elaborate sponge with plumose columns of oxea and styles and isolated desmata. Burton (1929) described different Ren1arks: So far, the tendency is found among growth forms of this species, one of which shmvs distinct spongologists to confine the usc of this genus tube-like fistules. name to typically fistulose forms, although many authors dealing with Indonesian sponges Species examined: Petromica ciocaiyptoides (Van (Dendy, 1889; Kieschnick, 1896; Sallas, 1902; Soest & Zea, 1986, as Monanthus) (holotype m Thiele, 1903; Hentschel, 1912) disregarded the Lcidcn Museum, paratype specimens in growth form and included Topsentia-Iike and ZMA). Amorphinopsis-like non-fistulose sponges in Geographical distribution: Azores, Caribbean Ciocalypta. Since several apparently closely and South East Africa. related fistulose, style-bearing species are in existence, it is here proposed to restrict the usc Genus group Ciocalypta-Amorphinopsis-Halichon­ of Ciocalypta to these, and to assign oxeote fistulose forms to Topsentia or Halichondria, dria-Hymeniacidon: Halichondriidae with a excepting those bearing desmata which go to distinct ectoson1al tangential reticulation of Petromica. spicule tracts, bundles or single spicules.

Synonyms: Genus group Ciocalypta-Amorphinopsis: Genus Leucophloeus Carter, 1883: :323. Halichondriidae possessing small cctosomal Type species (monotypy): Lcucoph!oeus massalis Carter, styles next to a con1plement of oxeote or stylote 1883: 323, pl. XIV fig. 15A-H (not examined). megascleres. Remarks: The true nature of this genus remains uncertain due to the fact that we have not been able to trace the original Carter specimen from West-Australia. Other Genus Ciocalypta Bowerbank, 1862: 1105. specimens incorporated in the British Museum (reg. no. Fig. 60, 67 1857:1:2:29 and 1886:12:15:82) described by Carter

Figs. 45-50. Fig. 45. Topscntia ophirhaphiditcs (De Laubenfels, 1934), A. cross section of peripheral region (redrawn from Dfaz eta!., 1990) (s.b. = 0.5 mm), B. spicules (s.b. = 100 !J.m). Fig. 46. 1'rach)·opsis ( = Topsentia) ha/ichondrioides Dendy, 1922, A. cross section of peripheral region (made from BIVINH holotype) (s.b. = 0.5 mm), B. oxca (s.b. = 100 !J.m). Fig. 47. Topsentia glabra (Topsent, 1900), spicules (redrawn from Topsent, 1904: pl. 13 fig. 1) (s.b. = 100 fJ.m). Fig. 48. Coelocalypta ( = Topsentia) porrecta Topsent, 1928, A. longitudinal section of fistule half, showing central suppor­ ting axis and brush-like ectosomal strengthening (s.b. = 0.5 mm), B. spicule (s.b. = 100 [J.m). Fig. 49. Oxeostilon (? = Topsentia) annandale/ Ferrer-Hernandez (1922), A. habit (s.b. = 1 em), B. spicules (s.b. = 100 fJ.m) (both redrawn from Ferrer-Hernandez, 1922: pl. 1 lower figure and figure on page 10). Fig. 50. Laminospongia ( = Topsentia) subtilis Pulitzer-Finali (1983), A. habit (s.b. = 2 em), B. spicules (s.b. = 50 fJ.m) (both redrawn from Pulitzer-Finali, 1983: figs. 59 and 59 bis).

43 Figs. 51-55. Figs. 51-52, Topsmtia roqucnsis Dlaz el a{. (1987) ( = T. ophirhaphidites (De Laubenfels, 1934)), type specimens in situ in the reefs of Los Roques, Venezuela (s.b. = 5 em) (photo E. Weil). Fig. 53. Topsentia porrecta (Topscnt, 1928), topotypical specimen from Boavista, Cape Verde Islands (s.b. = 1 em) (photo F. Vcrbiest). Fig. 54. Tojmntia n. spec. Dlaz ct a!. (in press), Bahama type specimens from the Harbor Branch collection. Fig. 55. Reniera ( = Topsentia) aurantiaca Schmidt (1862), Mus. Geneva holotypc from the Adriatic (s.b. = 1 em) (photo L. A. van de1· Laan).

44 (1886) and Dendy (1896) under this name from viz. New by Thomas, 1978) is not corroborated by recent Zealand and South-Australia (examined due to the findings of the species. We examined some of courtesy of Miss S.M. Stone) do not seem to conform exactly to the type description, but belong to a species of Thomas' material (MUS. Tervuren reg. no. Ciocalypta. An unregistered dry specimen from the Bower­ 1355), but the identification seems dubious; bank collection, labeled under the name Leucophloeus probably, it concerns specimens of the genus mas sa lis Australia and bearing the numbers 706 and 31, Aka ( ~ Siphonodictyon). has the characters described by Carter, viz. a thick white crust over a fibrous axially condensed choanosomal Synonyms: reticulation. The fragmentary specimen has a smooth but furrowed surface, and is hesitatingly assigned to Genus Prost)'lissa Topsent, 1925: 208. Cioralypta. A microscopic slide in the BMNH colection (Fig. 64) labeled Leucophloeus massalis SE Australia ( 1862: 12: 15: 454) Type species (monotypy): P. siamensis Topsent, 1925: 208, also seems to conform to Carter's description (thick crust figs. 1-2 (Fig. 64) (holotype reexamined). of irregularly reticulated bundles of megascleres, carried Topsent: ''Axinellidac a charpente composee d'oxcs by dendritically branched choanosomal vague tracts with inegaux. De petits styles, qui s'ajoutent a l'int6rieur en variable spongin binding the spicules); however, the proportion relativernent faible, deviennent nombreux spicules are oxea and it may be from a species similar to dans l'ectosome et constituent a cette membrane une pro­ ?'opsentia porrecta. tection speciale en se plat;ant debout, solitaires ou par groupes, le long des axes tangentielles qui la soutien­ Species examined: Ciocalypta penicillus Bower­ nent." bank, 1862 (holotypc in BMNH (Fig. 60) and Remarks: The type specimen (examined in NINHN, DT. 34; type slide: DT. 1885) is ramose, which is an unusual topotypical specimens in BMNH and ZMA growth form of the normally massive species; skeletal (Fig. 67)), C. massalis (Carter, 1886 as details· (detachable crust, confused choanosome, oxea of Leucophloeus) (topotypical specimens m 300-1050 by 40 ~m, and styles of 180-300 by 7-10 ~m) BMNH), Ciocalypta spec. (Indonesian specimen agree completely with A. exravans. in ZMA). Geographical distribution: temperate, sub­ Species examined: A. excavans Carter, 1887 tropical and tropical parts of all oceans. (type of Prostylissa siamensis (Fig. 64); ZMA specimens from Indonesia (Fig. 63)), A. Genus Amorphinopsis Carter, 1887: 77. subacerata (Ridley & Dendy, 1886 as Figs. 61-64, 68 Hymeniacidon) (holotype in BMNH 1887:5:57, Type species (monotypy): A. excavans Carter, ZMA specimens from Indonesia), A. foetida 1887: 77, pl. V figs. 12-15 (Fig. 61) (holotype (Dendy, 1887 as Hymeniacidon) (holotype in not examined). BMNH, ZMA specimens from Indonesia (fig. 68); specimen described as A. Joetida by Levi, 1961 in MNHN; holotypc of Halichondria Definition (emended): Halichondriidae of the aldabrensis Levi, 1961 MNHN no. DC!. 363 genus-group Ciocalypta-Amorphinopsis, with an which is considered a junior synonyrn); A. spec. ectosmnal skeleton of thick bundles of oxeotcs (ZMA specimens from Indonesia). "echinated" by sn1all styles. Geographical distribution: Indo-West Pacific tropical region. Remarks: The choanosomal tnegasclcres are mostly oxeotes although apices may be blunt, Genus group Haliclzondria-Hymeniacidon: occasionally stylotes. The peripheral parts include sn1aller oxeotes and styles. Unfor­ Halichondriidae with an ectosomal skeleton tunately, the type specimen of A. excavans seerns consisting of a thin, tangential, detachable to be no longer extant (pers. comm. Miss S.M. membrane charged with single spicules or Stone), so we can only rely on the excellent vague bundles, supported by choanosomal col­ redescriptions of Annandale (1919) (Fig. 62), umns traversing subdermal lacunae; larval who claims to have seen the type. The alleged strategy, where known, consists of internal fer­ excavating nature of this species (claimed also tilization and incubation of larvae.

45 E '

56

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46 Remarks: two genera Halichondria and chondria-like species (e. g. two undescribed Hymeniacidon were formerly considered the type Caribbean forms) have stylote modifications or genera of two distinct lines within the Halichon­ even dear styles in their spicule complement, driida, viz. the oxea-line and the style-line. next to oxea, so "loss of styles'' cannot be con­ Apart from the fact that ancestral forn1s quite sidered a synapomorphy; the majority of the likely already possessed both spicule-types, and species, however, possess exclusively ·sharp­ that their possession cannot be used as synapo­ pointed oxea of an intermediate to small size morphies, it is quite clear fron1 their skeletal when cmnpared to other members of the architecture that they are very close and can family. only be kept apart by assuming that the loss of Many Haiichondria-spccics have lost partly or oxea in Hymeniacidon is an apomorphous wholly their ectoson1al skeleton, and their development. Halichondria can be characterized descriptions sound very n1uch like e.g. A.i:ill)'SSa on their tufted larvae (known frmn at least two on paper, however, this loss is here interpreted members, sec e.g. Wapstra & Van Soest, 1987). as having independently occurred here and Some monotypical genera belonging in this there in the genus. genus group have doubtful status; they have tentatively been synonytnized with the two mam genera. Synonymy:

Genus Haiichondria Fleming, 1828: 520. ?Genus Ciocalajmta De Laubenfels, 1936: 134. (FiK. 73) Figs. 69-83 Type species (monotypy): Cioca(;•pta amorplwsa Ridley & Dendy, 1886:479, 1887:175, pl. XL fig. 9 (holotype reex­ Type species (original designation): Spongia amined). panicea Pallas, 1766 (type specimen probably De Laubenfels: "Halichondriidae like Ciocaf;•pta but with lost). both oxea and styles." Remarks: The type and only species is described as massive amorphous, vvith a rough tufted surface, which Definition: Halichondriidae of the genus group has a delicate dermal reticulation. Examination of a type Halichondria-Hymeniacidon with tufted larvae. fragment (Bf\'INH 1887:5:2:56) (Fig. 73) revealed two sizes of oxea (1700 by 20 and 600-750 by 8 [J.m) and one Remarks: It is assumed here, that all Halichon­ size of styles (1400/28 fJ.tn). It is very probably not a typicalllalichondria but in the absence of further speCimens dria species have larvae similar to those of H. and similar species, it is considered closely related and panicea and H. bowerbanki (see Topsent, 1911; falls probably within the definition. Possibly it is an Wapstra & Van Soest, 1987). Some Hali- atypical Biemna.

Figs. 56-59. Fig. 56. Spongosorites ( = Epipolasis) suluensis Wilson (1925), A. cross section to show overall structure (redrawn from VVilson, 1925: pl. 48 fig. 3) (s.b. = 1 em), B. cross section of "upper" peripheral region to show palissade-like brushes (s.b. = 0.5 mm), C. cross section of''lower'' peripheral region to show tangential crust(s.b. = 0.5 mm), D. tangential view of "lower" surface to show feltwork of tangential oxea (s.b. = 0.5 mm), E. megascleres (s.b. = 100 fJ.IH), F. trichodragmata (s.b. = 50 [J.m) (B-F made from the USNM type specimen). Fig. 57. Pctromica grimaldi Topsent (1898), A. habit (s.b. = 1 em), B. desma (s.b. = 50 [J.m) (both redrawn from Top­ sent, 1904: pl. 5 fig. 5, pl. 8 fig. 6). Fig. 58. A1onanthus ( = Pctromica) plumosus Kirkpatrick (1903), A. habit of the var. tubulata (s.b. = 1 em), B. section ofoscular tube (s.b. = 0.5 em), C. spicules (s.b. = 100 [J.m), D. desma (s.b. = 100 [J.m) (all redrawn from Kirkpatrick, 1903: fig. 6). Fig. 59. lvfonantlws ( = Pctromica) ciocalyptoides Van Soest & Zea (1986), A. cross section of basal mass to show subcc­ tosomal desmata (s.b. = 1 mm), B. strongylotes (s.b. = 100 [J.m), C. apices ofstrongylotes ((sb. = 10 [J.m), D. desma (s.b. = 100 fJ.In) (all redrawn from Van Socst & Zea, 1986: figs. 2-5).

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48 Genus Cioxcamastia De Laubenfels, 194-2: 265. Type species (monotypy): H. corticala Burton, 19:31: 137, (Fig. 78) fig. 2 (holotypc reexamined). Type species (monotypy): C. po(ycal;t;ta De Laubenfels, Burton: "Haploscleridae with spicules of one size only; 1942: 265 (Fig. 78) (holotype reex;;lmined). main skeleton composed of long slender fibres running De Laubenfels: "Halichondriidae with spiculation and vertically to the surface and merging into a subdermal most other characters quite typical, but differs in possess­ palissadc of oxea with scattered spicules between the ing conspicuous closed fistules, closely resembling those fibres; dermal skeleton a delicate tangential reticulation of that characterize the genus Po(ymastia of the family oxca." (Fig. 80) Suberitidae. The genus Cioca(ypta of the Halichondriidae Remarks: The type specimen (BMNH 1931:10:28:25, also has fistules, but these are coarser than those of Ciox­ reexamined) is an atypical Halichondria panicea, with camastia. The spiculation of C'ioca!ypta is not typical of its relatively feeble oxca size (due no doubt to the fact that it family.'' had been growing under artificial circumstances in the Remarks: The type specimen from 66°N 80°W (USNM Tmndheim (Norway) aquarium). 22691, reexamined) is described as being subspherical with about 50 closed fistules each 4 mm high, 1-3 nun in Genus 1\1cnandia Topscnt, 1896: 115. diameter; the ectosomc is said to he not detachable, but it (Fig. 81) is definitely tangential; oxea vary from 200-700 by 4-12 Type species (monotypy): j\,1. minchiniTopsent, 1896: 116 [J.m. It is likely that this specimen belongs to Halichondria (type slide reexamined). (Eumastia) sitiens (Schmidt, 1870), which is a circum-Arctic Topsent: "Renierinae a ectosome epais, remarquable cor­ species, iace, pourvu de spicules abondants et echevCtrCs en toutes directions sans son Cpaisseur, et forterncnt adhCrent au Genus Eumastia Schmidt, 1870: 42. choanosomc; uwites prCporales trCs reduites." (Fig. 79) Remarks: A slide of the type specimen (DT. 2099) has Type species (monotypy): E. siticns Schmidt, 1870: 42, pl. been reexamined; this conforms in many details with V fig. 12 (type material not examined). Halichondria, probably H. panicea, although the reduced Schmidt: "Unterscheidet sich von Pellina durch die kegel­ state of subdermal cavities and the heavy ectosomal und zottenfOrrnigen Ausstulpungcn der Haut, die sich skeleton arc somewhat atypical (Fig. 81). Similar sogar vcrzwe1gen k6nnen, und deren Basen em specimens have been described as Halidwndria topscnti De unregelrniissigen Labyrinth von zusarnmenhiingenden Laubenfels, 1936 from the same area; quite likely, these Riiumen bilden.'' (Fig. 79) forms are growth forms of H. j;anicea from exposed Remarks: The species is described as bearing cone-shaped habitats (sec also Vethaak et al., 1982). Van Soest (1980), papillae on a massive basis; topotypical specimens possess on the suggestion of De Laubenfels (1936: 70) doubtfully a detachable ectosornal membrane and subdermal accepted Alenanrtia as a Haplosderid (synonym of Adocia). lacunae; the skeleton consists of irregular tracts and spicules in confusion; oxea are (400-) 690 fl.lll (by 7-10 ?Genus P_yloderma Kirkpatrick, 1908: 51. ~m). (Fig. 82) Type species (monotypy): Halidzondria latnmculioidcs Genus Halichondriclla Burton, 1931: 137. Ridley & Dendy, 1886:326, 1887: pl. I figs. 5, pl. XLVI (Fig. 80) fig. 5 (holotype reexamined).

Figs. 60-64-. Fig. 60. Ciocai.J•pta pmicilhn Bowerbank (186'1), A. tangential view of surface skeleton (s.b. = 0.5 mm), B. longitudinal section through fistule showing axial support (s.b. = 05 mm), C. spicules (s.b. = 100 fJ.m) (all made from the BMNH holotype). Fig. 61. Amorphinopsis c.mwans Carter (1887), A. habit of holotype (s.b. = 1 mm), B. large mcgascleres (s.b. = 100 [J.rn), C. small style (s.b. = 100 fl.lll) (all redrawn from Carter, 1887: pl. 5 figs. 13-15). Fig. 62. Amorphinopsis cxcavans Carter ( 1887), tangential view of the surface skeleton of a ZMA specimen fmm Indonesia to show spicule tracts and "echinating" small styles (s.b. = 0.5 mm). Fig. 6:3. AmorphinojJsis exravans var. d(l!,it!fcra Annandale ( 1913), A. longitudinal section thmugh a digitiform process ( s. b. = 1 em), B. spicules (s.b. = 100 fJ.m) (both redrawn from Annandale, 1913: figs. '1-5). Fig. 64-. Prost)'lissa siamcnsisTopsent (1925) ( = AmorphinojmS exravans Carter, 1887), A. habit (s.b. = 1 em), B. tangential view of surface skeleton (s.b. = 100 fl.tn), C. spicules (s.b. = 100 fl.ill) (all redrawn from Topscnt, 1925: figs. 1-2).

49 Figs. 65-68. Fig. 65. Epipolasis su!uensis (Wilson, 1925), holotype USNM 21297 from the Philippine region (s.b. = 1 em). Fig. 66. Pelromica ciocalyptoidcs (Van Soest & Zca, 1986), photographed in situ in the reefs of the Colombian Caribbean (s.b. = 5 em) (photoS. Zea). Fig. 67. Cioca(ypta jJcnicil!us Bowerbank (1864), ZMA specimen from Lundy Island, SW Britain, collcct~.:d by Dr Vl. H. de Weerdt (s.b. = 1 em) (photo L. A. van dcr Laan). Fig. 68. AmorphinojJsisfoetida (Dendy, 1889), ZMA specimen from Indonesia (s.b. = 1 em) (photo H. van Branchvijk).

50 Kirkpatrick: ''Renierinae with a parchment-like, easily The genus Amorphina Schmidt, 11i70 has been generally separated dermal membrane in which are situated closely assumed to be a junior synonym of Haliclwndria; however, packed tangential oxea, and with distinct round or oval the type species, A. grossa, is a Jla!idona. pore areas Remarks: the type specimen (type fragment B1v1NI-I Species exatnined: Ha/ichondria panicea (Pallas, 1887:5:2:197 reexamined) is described as erect-lobosc, 1766) (ZMA specimens from all over W pear-shaped, surface uneven, soft, spongy; the dermal Europa) (Figs. 69, 74), H. bowerbanki Burton, membrane is very dense (Fig. 82B), not like typical Halic!wndria, and the choanosomal skeleton consists of 1930 (ZMA specimens from all over W Europa) irregular tracts and loose oxea of 700-1250 by 22-31 !J.m. (Figs. 70, 76), H. gibbsi (Wells et al., 1960 as If more species with similar ectosomal characters vvould be Cioca/yj;ta) (USNM holotype 23666, from North found, then this genus could be considered valid. Carolina (Fig. 72), ZMA spcomcn from

Genus Tradi)'OjJSilla Burton, 1931: 138. Bonaire), H. tyleri (Bowerbank, 1875) (as (Fig. 83) Ciocalypta) (holotype BMNH 1930:7:3:32 from Type species: T. glaberrima Burton, 1931: 138, figs. 3-4 South East Africa), H. melanodocia De (holotypc reexamined). Laubenfels, 1936 (USNM type specimen, Burton: "Haploscleridae with skeleton of oxea only, of ZMA specimens from the Caribbean); holotype variable size; main skeleton halichondroid, with loose fibres running vertically to the surface, and ending in of ilponastra modes/a Pulitzer-Finali, 1986 ti·om brushes, and with a laos~.: tang~.:ntial layer of spicules Mus. Genova no. 47688, which is considered a associated with outer ends of these spi~.:ulcs.'' junior synonyn1), H. magniconu/osa Bechtel, Remarks: The type spccinwn (Bl--iiNH reg. no. 1931:10: 1965 (USNM paratype specimen 24499, ZMA 28:77, reexamined) very probably conforms to Halichon­ specimens from the Caribbean (Fig. 75), H. dria panicea; the 'brushes' are the skeletal columns on which the tangential ectosomal skeleton rests (Fig. 83). /olea Alcolado, 1984 (USNM type fragment, Oxca arc 240-340 by '1-8 !J.m. ZMA fragment from Venezuela (Figs. 71, 85), H. n. spec. (HBOI specimens from Venezuela); Genus Raspaigcl!a Schmidt, 1868: 2:'>. H. et a/., (Fig. 77, 84) n. spec. Diaz in press (USNM holo­ Type species (monotypy): R. bnmnca Schmidt, 1868: 25 type fron1 Florida, ( = H. panicea sensu Little, (typ~.: specimens reexamined). 1963)). Schmidt: " ... erinnert an Raspailia; es stehen je doch die Geographical distribution: This seems to be a Nadel nich so hervor \vie beijener Gattung, indcm sic nur truly cosmopolitan genus, although it IS an einzelnen Stellen in unrcgclm.iissigen Biindeln bis zur definitely more common in temperate and cold OberJ1iiche und wenig tiber dicselbe herausragen. Auch lOst sich die OberJ1achenschichte stellenweise in Form waters than in tropical regions. einer Membran ah. Das innerc hat nicht die bei RasjJailia so deutliche Horngefiige. Die Nadcln licgen meist dcr Genus Hymeniacidon Bowerbank, 1862: 1112. liingsachse der Aste parallel. Ihrer sind zwei Sort en: eine Fig. 86-93 an beiden Enden allmiihlig tmgcspitztc von wechselnder Type species (original designation): Hymenia­ GrOsse herrscht vor. Die andere ist stumpf-spitz." cidon caruncula Bowerbank, 1864: 191, pl. Remarks: The alleged type specimens (LlviJG 15524 and 15350) of this species (Fig. 77) have been rediscovered in XXXV fig. 372 (holotype not examined) (Fig. the collections of the museum in Graz (Desqueyroux, per­ 87 a). This is generally considered a junior sonal communication); they arc now temporarily stored in synonym of Spongia sanguinea Grant (1829) and the MusCe d'Histoire de Geneve. The sponge has limp Spongia perlevis Montagu, 1818, so the definitive branches (reminding strongly of Ha!ichondria bowr:rbanki) name of the type species is H.ymeniacidon j;er/evis and a definite cctosomal tangential Halichondria skeleton (Fig. 84). Oxea are in two size categories: 410-600 by 7-10 (Montagu, 1818). !J.n1 and 150-260 by 3-6 !J.m. Definition: Halichondriidae of the genus group Genus Spwna P.'Iiklucho Maclay, 1870: 13. Ha/iclzondria-Hymeniacidon which have lost their oxeote spicules, retaining styles and stylotcs. Type species: S. borealis ivfiklucho l'vlaclay, 1870: 13. Remark: \'Ve have not seen the original description. Koltun (1959: 206) claims the type species is a junior Remarks: The tangential ectosomal skeleton is synonym of Ewnastia sitiens Schmidt, 1870. normally less well-developed than in Halichon-

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52 dria, and there is a decidedly fleshy consistency unlike the Jvlediterranean species in its lack of spongin, its in the type and many other spec1es. possession of vague disoriented spicule tracts, and the small size of the styles. Rhaphidosf)•ia has also been misinterpreted by Van Soest & Weinberg, 1981, who Synonymy: assigned Tethyspira spinosa Topsent, 1891, erroneously to Genus Amorphi/la Thiele, 1898: 44. Rhaphidostyia incisa. In a paper in press, Van Soest has rec­ Type species (original designation): Hymcniacidon san guinea tified this error and proposed to abandon the use of (Grant, 1829). This designation makes the genus an Rhaphidostyia in favor of Hymeniacidon (for the type species objective junior synonym, as it is also the type of R. kitchingi) and Dict;·oneila (for the other assigned species). Hymeniacidon. Genus Rhap!wxielia Burton, 1934-: 554. Genus Nailondra De Laubenfels, 1954: 182. (Fig. 90) (Fig. 88) Type species (monotypy): Batzclla corticata Thiele, 1905: Type species: N. maza De Laubenfels, 195<1: 182, fig. 121 4-J8, fig. 58 (type slide reexamined). (holotypc reexamined). Burton: "Close to R!wpho.qa Hal\mann, but differs in De Laubenfels: '' Halichondriidac with strongylcs and having a tangential dermal layer of short strongyla, not styles; structure like Halichondria." recorded by Thiele, about half the length of the principal Remarks: The specimen (USNM 2308:3, reexamined) is megascleres. '' described as amorphous, soft like "soggy bread"; it has a Remarks: The BMNH holds a slide (BJv[NH reg. no. definite tangential skeleton and subdermal spaces. The 1908.9:24:15a, reexamined) of the type specimen (which spicules are described as styles and strongyles. Although is in the Berlin Museum); this shows an ectosomal tangen­ the majority are oxeotes with notable narrowing of the tial reticulation of loose spicules, and choanosomal vague blunt apex, there are also some true styles, including ones tracts and low spicular density; spongin seems to be with a faint tylote swelling. Size of larger category of absent; spicules are styloids, oxeotes and strongyloids with spicules: 540-630/12 fJ.Il1. In the Halic!wndria-Iike architec­ variable apices. It is an atypical H;wuniacidon. ture, this species is close to Uritaia (see below). Genus S't;•lohaiina Kirk, 1909: 539. Genus Rhaphidostyia Burton, 1935: 651. Type species (monotypy): S. wnica Kirk, 1909: 539, pl. (Fig. 89) 25. Type species (original designation): R. kitchin,l!,i Burton, Remarks: The original description has not been seen, but 1935: 652 (holotype reexamined). Bergquist (1970) considers this a junior synonym of Burton: "Axinellidae with skeleton composed of HymcniaCI'don. unusually slender styli, arranged mainly in an irregular (sub-halichomlroid) manner, but frequently forming Genus Sf)•loteiia Von Lendenfeld, 1888: 185. -..vhisp-like bundles, running towards the surface; spicules (Fig. 91) characterized by a series of abrupt narrowings ending in Type species (monotypy): S. d(£tilata Von Lcndenfeld, a point, at the distal end." 1888: 185. This is generally considered a junior synonym Remarks: Burton assigned also a series of Mediterranean of H)'lncniacidon agminata Ridley, 1884: 466, holotype Dicf)'Oiu:lla species to his genus; the type specimen of R. kit­ BMNH 81:10:21:34-7 (sec Hallmann, 1914: 34-9). chingi (BMNH reg. no. 1934-:9:26:79, reexamined) is Von Lendenfeld: ''Heterorrhaphidae of very soft texture.

Figs. 69-73. Fig. 69. Halidwndria panicea (Pallas, 1776), A. tangential view of ectosom;:ll skeleton (redmwn from Hartman, 1958: fig. 9) (s.b. = 100 fJ.m), B. spicules (s.b. = 100 fJ.m). Fig. 70. Ilaiirhondria bowerbanki Burton (1930), tangential view of ectosomal skeleton (redravvn from Hartman, 1958: fig. 8) (s.b. ~ 100 ~m). Fig. 71, llalichondria iutea Alcolado (1984), A. longitudinal section of apex of one of the digitate processes (s.b. = 1 mm), B. spicules (s.b. = 100 fJ.m). Fig. 72. Cioca(ypta ( = Haiichondria) gibbsi VVells, ct al. (1960), A., habit (redrawn from Wells, eta!., 1960: fig. 4-7) (s.b. = 1 em), B. longitudinal section of fistule half (s.b. = 1 mm), C. tangential view of surface skeleton (same scale as B) (B and C made from the USNM holotype). Fig. 73. Ciocaiapata (? = Haiichondria) amorjJlwsa (Ridley & Dendy, 1886), A. tangential view of surface skeleton (s.b. = 0.5 mm), B. cross section of peripheral region (s.b. = 1 mm), C. spicules (s.b. = 100 fJ.lll) (all made from the BMNH holotype).

53 54 Megasclera styli; in bundles and scattered. No Species examined: H. perlevis (Montagu, 1818) microsdera.'' (ZMA specimens from all over Europe (Fig. 86, Remarks: The type is described as having branches and digitate processes; surface smooth; the ectosome has 87)), H. heliophila (Parker, 1910) (USNM and "horizontally disposed spicules", and the choanosome is ZMA specimens from Florida, West Indies, "occupied entirely by a dense mass of longitudinally and North Carolina), H. caerulea Pulitzer­ disposed spicules"; the only spicules arc styles of 250 by Finali, 1986 (USNM and ZMA specimens from 4 !liD· the Caribbean), H. corticata (Thiele, 1905) (type Genus Thieleia Burton, 1912: 329. slide from BMNH), H. rubigiuosa (Thiele, 1905) (Fig. 92) (type fragment from BMNH), H. kitchingi (Bur­ Type species (monotypy): 1-lymeniacidon rubiginosa Thiele, ton, 1935) (type (BMNH) and ZMA specimens 1905: 421, fig. 44 (paratype reexamined). from the British Isles). Burton: ''Main skeleton composed of delicate strands of Geographic distribution: Temperate, sub­ spicules running vertically to the surface and connected to each other by irregularly arranged masses of single tropical and tropical regions of all oceans; not in spicules. At the surface the ends of the strands diverge to (ant-)arctic regions. form dense brushes of spicules the apices of which project slightly beyond the dermis. There is no special dermal skeleton unless the dense brushes formed from the ends of the strands of the main skeleton can be regarded as such." ACKNOWLEDGEMENTS Remarks: The type is described as a flat crust, no detachable skin, with an organic skin stretched over the Drs K. Riitzler (USNM, Washington), S. M. skeletal reticulum. The lHvJNH holds a fragment Stone (BMNH, London), C. Levi (MNHN, (08:9:24:1:33a, reexamined) from a paratype specimen Paris), D. Kuhlmann (ZMB, East Berlin), V. from the Berlin Museum; this largely conforms to H)'lncniacidon, the alleged absence of a dermal skeleton is Raineri & G. Doria (Mus. Genova), and R. a matter of grade, rather than a true absence. Desqucyroux (Mus. Geneva) arc thanked for their hospitality and kindness in providing Genus Uritaia Burton, 1932: 198. access to and loans from their collections . .Nlr. (Fig. 93) H. van Brandwijk and Mr. L. A. van der Laan Type species (monotypy): Uritaia halifhondrioidcs Burton, (ITZ) made many of the photographic illustra­ 1932: 199, pl. 7 figs. 3-4, text-fig. 4 (nnt examinerl). tions of specimens frmn the ZMA collection; Burton: ''Axinellidae with skeleton composed of two lV!r F. Verbicst (National Museum of Natural categories of smooth styli; main skeleton a halichondroid History, Leiden) photographed Topsentia por­ reticulation of large styli, with a few small styli scattered between; dermal skeleton formed of brushes of smaller recta; some underwater photographs were made styli set at right angles to surface with numerous styli of by Mr. E. Wei! (F.C.L.R., Caracas), Drs G. similar size lying horizontally and scattered between van Moorsel (Alkmaar) and S. Zca them''. (INVEMAR, Santa Marta); SEM photos were Remarks: Burton compares his new genus to Amorphilla made by F. Hiemstra (ITZ) and N. Stentoft and HJ'lllClliacidon and found them to differ in the more halichondroid nature of the ectosomal skeleton. Since this (Copenhagen). is generally less well developed in H)'lncniacidon species that This is Harbor Branch Oceanographic Institu­ difference seems hardly worth erecting a separate genus. tion, Inc., Contribution number 783.

Figs. H-77. Fig. 74. Halidwndria jJanicca (Pallas, 1776), ZNIA specimen from SVV Netherlands (s.b. = 1 ern). Fig. 75. Halichondria magniconulosa Hechtel (1965), ZMA specimens from Cura~ao (s.b. = 1 em) (photo L.A. van der Laan). Fig. 76. Halidwndria bowcrbanki Burton (1930), Zl\tiA specimen from S\·V Netherlands (s.b. = 1 em) (photo L. A. van der Laan). Fig. 77. Raspaigella ( = Halidwndria) brunnca Schmidt (1868), one of the lvlus. Geneva type specimens (LMJ G 15350/o) (s.b. = 1 em) (photo L. A. van der Laan).

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'

56 REFERENCES --, 1931a. Studies on Norwegian sponges I. Det Kong. Norske Videns. Selsk. Forh., 4 (38): 136-139. ALCOLADO, P., 1984. Nuevas especics de csponjas encon­ --, 1931b. A collection of marine sponges mostly from tradas en Cuba. Poeyana, 271: 1-22. the Natal coast. Ann. Natal Mus., 6 (3): 337-358, pl. ANNANDALE, N., 1915. Some sponges parasitic on XXIII. Clionidae with further notes on that family. Rcc. --, 1932. Sponges. Disc. Reps. Cambridge Univ. Indian Museum, 11 (VI, 27): 457-478, pl. 31. Press, 6: 237-392, pis. <18-57. BARTIK, K., j.-C. BRAEKMAN, D. DALOZE & C. STOLLER, --, 1933. Report on a collection of sponges made in 1987. Topscntins, new toxic bis-indole alkaloids South Saghalin by :Nir Tomoe Urita. Sci. Rep. from the marine sponge Topsentia genitrix. Can. J. Tohoku Imp. Univ., ser. TV, 7: 195-206. Chern., 65: 2118-2121. --, 1935. Notes on British sponges, with a description BERG, C., 1899. Substituci6n de nombrcs gcncricos. III. of a new genus and species. Ann. Niag. Nat. Hist. Commun. Jvlus. Nac. Buenos Aires, 1: 77-80. (10) 15: 651-653. BERGQUIST, P. R., 1965. The sponges of iV1icronesia, part --, 1959. Sponges. In: The John Murray Expedition 1. The Palau Archipelago. Pacific Science, 19 (2): 1944-34. Scient. Rep. John Niurray Exped. London, 123-204. British i\'Iuseum (Natural History), 10 (5): 151-281. --, 1970. The marine fauna of New Zealand: Porifera, CARTF.R, H. J., 1883. Contributions to our knowledge of Dcmuspongiae, Part 2 (Axincllida and I-Ialichon­ the Spongida. Ann. :tviag. nat. Hist., (5) 12: 308- drida). New Zealand Oceanogr. lnst. fviem., 51: 329, ph. XI-XIV. 1-85. 1886. Descriptions of sponges from the --, 1978. Sponges. Univ. Calif. Press. Bcrkely: 1-268, neighbourhood of Port Philip Heads, South pk 1-12. Australia, continued. Ann. Mag. nat. Hist., (5) 17: BERGQUIST, P.R. & C. A. T!ZARD, 1967. Australian inter­ 40-43, 112-127, 431-H1, 502

Fig. 78-8·1. Fig. 78. Cioxmmastia ( = Halidumdria) polyaQ•pta De Laubenfels (1942), A. tangential view of surface skeleton (s.b. = 0.5 mm), B. longitudincll section of papilla (s.b. = 1 mm), C. oxea (s.b. = 100 [J.m). Fig. 79. Ewnastia ( = Halidwndria) sitims Schmidt (1870), A. habit (redrawn from Schmidt, 1870: pl. 5 rig. 12) (no scale given by Schmidt), B. tangential view of surface skeleton of a ZlviA specimen from the northern North Sea (s.b. = 0.5 mm). Fig. HO. llafidwndriclla ( = Halichondria) cortirata Burton (1931), A. cross section of peripheral region (s.b. = 0.5 mm), B. tangential view of surface skeleton (same scale), C. oxea (s.b. = 100 [J.m) (all redrawn from Burton, 1931: tig. 2). Fig. 81. Afcnandia ( = Halichondria) minrhini 'l'opsent (1896), A. cross section of peripheral region (s.b. = 1 mm), B. tangential view of cctosomal skeleton (s.b. = O.:J mm), C. oxea (s.b. = 100 [J.m) (all made from slide of MNHN type specimen). Fig. 82. P_y!oderma (? = Halidwndria) latrunw!ioidcs (Ridley & Dendy, 1886), A. habit (s.b. = 1 em), B. tangential view of surface skeleton showing a feltwork of oxea (s.b. "" 1 mm) (both redrawn from Ridley & Dendy, 1887: pl. 1 tig. 5 and pl. 46 fig. 5). Fig. 83. Traci!)'OjJSi!!a (= lla!irlwndria)p)abcrrima Burton (1931), A. habit (s.b. = 1 em), B. cross section of peripheral region (s.b. = 0.5 mm), C. oxea (s.b. = 100 [J.tn) (all redrawn from Burton, 1931: figs. 3-<1). Fig. 81. Rasjmigella ( = !-lalichondria) brwmca Schmidt (1868), A. tangential view of surface skeleton (s.b. = 0.5 mm), B. oxea (s.b. = 100 [J.m) (made from one of the lvius. Geneva type specimens).

57 58 by H. M.S. "Sealark" in the Indian Ocean. Trans. Linnean Soc. New South \'\'ales, 39: 263-315, 327- Linn. Soc. Lond., Zoology, 18 (1): 1-164, pis. 1-18. 376, 398·446, 10 ph, 2'1 figs. DE LAUBENFELS, see LAUBRNFF.J.S. HARTMANN, W. D., 1982. Porifera. In: S. P. Parker, eel.: DiAz, NL C., B. ALVAREZ & R. V\'. Nf. VAN SoEsT, 1987. Synopsis and classification of living organisms, vol. New species of Demospongiae (Porifera) from the 1. r..~lcGraw-Hill, New York, pp. 640-666. National Park Archipelago Los Roques, Vene7.mJa. liENTSGHEL, E., 1912. Kiesel- und Hornschwiirnme der BUd.-. Dierk., 57 (1): :ll-41. Aru- und Kei-Inseln. Abhandl. Senckenb. Naturf. DiAz, r._,r. C., S. A. PoMPONI & R. VV. r..lf. VAN SoEST, Ges., 34: 295-448, pis. 13·21. 1990a. A systematic revision of the Central-Atlantic --, 1923. Erste Unterabteilung der Nletazoa: Parazoa Halichondrida (Demspongiae, Porifera). Part I: ... Porifera= Schwamme. In: Kiikenthal, VVilly, and Evaluation of characters and diagnosis of genera. In: Krumbach, Thilo (eds.), Handbuch der Zoologic. H. Keupp & J. E. Reitner, eels.· Fossil & Recent Berlin and Leipzig", Walter de Gruyter & Co., 1: Pro­ sponges (in press). tozoa, Porifera, Coelenterata, Mesowa: 307-418. DfAz, M. C., S. A. PoMPON! & R. W. ~vl. VAN SoEST, in HooPER, J. N. A., 1990. Character stability, systematics press. A systematic revision of the Central-Atlantic and affinities between lVIicrocionidae (Poccilo­ Halichondrida (Demospongiae, Porifera). Part III. sclerida) and Axinellida. Proc. 3rd Int. Conf. Bioi. Description of valid species. Proc. Rio\. Soc. Sponges, \'\'oods Hole 1985 (in press). Washington. KIESCHNICK, 0., 1896. Kieselschwamme von Ternalc. EHLERS, E. H., 1870. Die Esperschen Spongien in der Zoo!. Anz., 19: 526-53+. Zoologischen Sammlung der K. UniversitJ.t KIRK, H. B., 1909. Two sponges from Campbell Island. Erlangen. E. Th. Jacob, Erlangcn: 1-36. In: Chilton, C., eel.: The Subantarctic islands of ELLIS, \'V. N., 1987. Tree Tools 2.3. A set of programs for New Zealand, p. 574-581, pl. XXVIL the construction of \tVagner Trees for the r._,Iacintosh KIRKPATRICK, R., 1903. Descriptions of South African computer. 1-44 (mimeographed). sponges. Part III. fviar. Invest. South Africa, 2: 233- EsPER, E. .J. C., 1791-94. Die Pflanzentieren in 264, pis. 5-G. Abbildungen nach der Natur mil Farben erleuchtet --, 1908. Porifera 2. Tetraxonida. Nation. Ant. Expeel. nebst Bcschrcibungen. NU.rnberg, Raspe, Zweyter 1901-1903, Zoo!. Botan., 4: l-56. Theil: 1-303, pls. 1-109 [sponges: 165-180, published KouuN, V. r._,r., 1959. [The cornacuspongida of the 1792, 181-220, published 1793, 221-282, published northern and fareastcrn seas of the USSR. Keys to 1799; pis. 1-Vll, VIlA, VIlE, VIIHX, lXA, X· the fauna of the USSRl, Moscow, Leningrad, xxr, xxrA, xxn-xxm, xxrrrA, XXIV· Akadcmia Nauk SSSR: 1-236, pis. 1-'l-3 (in XLVll, XLV[[A, XLV!ll-XLIXJ. Russian). --, 1798-1806. Fortsetzungen der Pflanz:enthiere LAUBENFELS, r._,f. \V. DE, 193<1. New sponges from the Niirnbcrg, Raspe, Zvveyter Teil: 1-2'1, 25-28 Puerto Rican deep. Smithson. i'vlisccl. Coli., 91 (17): [sponges: 18-24, :W-45; pls. LXII-LXIV, LXV, 1·28. LXVA, LXVI-LXV[[[, LXXJX [LXJX, --, 1936. A discussion of the sponge hmna of the Dry LXX]. Tortugas in particular and the \'\'est Indies in FELSENSTEIN, J , 1987. PIIYLIP (Phylogeny Inference g"cneral, with material for a revision of the bmilies Package) version 3.0 manual: 1-50 (on computer and orders of the Porifera. (Pub!. Carnegie Inst. disk) \'Vash., •167) Pap. Tortugas Lab., 30: 1-225. FERRER HERNANDEZ, F., 1922. i\!fas datas parae[ conoei­ --, 1942. Porifera from Greenland and Baffinland col­ miento de las esponjas de las costas espanolas. Bal. lected by Capt. Robert A. Bartlett. J. \·Vashington Pescas, 1922: 1-26, 2 pis. Acad. Sci., Zoo!., 32 (9): 263·269. FLEMING, .J., 1828. A history of British Animals. --, 195:L Sponges from the Gulf of i'vfexico. Bull. mar. Edinburgh-London: 1-565. Sci. Gulf Caribbean, 2: 511-557. GRANT, R. E., 1827. Notice of two new species of British --, 1954. The sponges of the \·Vest Central Pacific. sponges. Edinburgh new Phil. Journ., 2: 203-204. Oregon State "0.'lonogr. Zoo!., 7: i-x, 1-306. HALU.IANN, E. F., 1914. A revision of the monaxonid LENDENFELD, R. VON, 1888. Descriptive catalogue of the species described as new in Lendenfcld's ''Catalogue sponges in the Australian lvluseum, Sydney. Pub. of the sponges in the Australian tvluseum". Proc. Austral. r..~Ius., London, 16: 1-260, pis. 1-12.

Figs. 85-86. Fig. 85. Halichondria !utca Alcolado (1984), photographed in situ in the reef~ of Los Roques, Venezuela (s.b. "·" 1 em) (photo E. V/eil). Fig. 86. H;,mcniacidon pcrltvis (Montagu, 1818), ZMA specimen from S\·V Netherlands (s.b. = 1 em) (photo L.A. van der Laan).

59 87 c

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D

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c .l - ' . 90 ' I I I ! :fit\ :.\\.? I I I A l

92

60 --, 1897. Spongien von Sansibar. Ahhandl. Scnckenb. PoMPONI, S. A., A. E. WRIGHT, M. C. DIAZ & R. W. NI. Naturf. Ges., 21: 93-133, pis. IX-X. VAN SoEsT, 1990. A systematic revision of Central­ LE.vr, C., 1953. Sur une nouvelle classification des Atlantic Halichondrida (Dcmospongiae, Porifera). DCmosponges. C. R. Acad. Sci. Paris, 234: Part II. Patterns and distribution of secondary 1710-1712. metabolites, In: H. Keupp & J. E. Reitner, eels.: --, 1956. Etude des Halisarca de Roscoff. Embryologic Fossil & Recent sponges (in press). et systCmatiqucs des Demosponges. Arch. Zool. exp. PuLITZER-FINALI, G., 1983. A collection of Mediterranean gen., 93 (1): 1-181. Dcmospongiae (Porifera) with, in appendix, a list of --, 1973. Systematique de Ia classe de Demospongia the Demospongiac hitherto recorded from the (DCmosponges). TraitC de Zoologic, 3 (1): 577-631. 1v1editerranean sea. Ann Mus. Civico Storia Nat. LEvr, C. & P. LE.vr, 1983. DCmosponges bathyales Genova, 84: 445-621. rfcoltfes par le N/0 "Vauban" au sud de la --, 1986, A collection of Dcmospongiae from the West Nouvelle CalCdonie. Bull. Mus. nation, Hist. nat., Indies, with in appendix a list of the Dcmospongiae (4), 5 (A, 4): 931-997. hitherto recorded from the West Indies. Ann. Mus. LINDGREN, N. G., 1897. Beitrag zur Kenntnis dcr Civico Storia Nat. Genova, 86: 1-216. Spongienfauna des Nialaiischen Archipels und der RIDLEY, S. 0., 1884. Spongiida. Report on Zoological Chinesischen Nieere. Zool. Anz., 20: 480-487. Collections made in Indo Pacific Ocean during the --, 1898. Beitrag zur Kenntnis dcr Spongienfauna des voyage of H.M.S. "Alert" (1881-82). London; 366- Malayischen Archipcls und der chinesischcn rvieere. 482, 582-630, Pis. 39-43, 53-54. Zoo!. Jahrb. Abt. Syst., 11: 283-378, pis. 17-20. RIDLEY, S. 0. & A. DENDY, 1886. Preliminary report on NIADDISON, W. & D. MADDISON, 1987. MacClade for the the Monaxonida collected by H.M.S. 'Challenger'. Macintosh, version 2.1, 55 printed pages. Parts I and II. Ann. Mag. nat. Hist., (5) 18: 325- MrKLUcuo-MAcLAY, N., 1870. Ueber cinige Schwamme 351, 470-493. des n6rdlichen Stillen Ozeans unci des Eismeeres, --, 1887. Report on the Monaxonida collected by wclche im Zoologischen Museum der Kaiserlichcn H.M.S. "Challenger" during the years 1873-1876. Akademie dcr Wissenschaften in St. Petersburg Rep. Scient. Results Voy. Challenger, 1873-76. ausgestellt sind. Ml:m. Acad. Imp. St. Pctcrsbourg, Zoology, 20 (59): 1-275, figs. H 1, pis. 1-51. (7) 15 (3): 1-24, 2 pis. Row, R. W. H., 1911. Report on the sponges collected NloNTAGU, G., 1818. An essay on sponges, with descrip­ by Mr. Cyril Crossland in 1904-05. Part II. Non­ tions of all the species that have been discovered on Calcarea. J. Linn. Soc., 31: 287-400, pis. 35-41. the coast of Great Britain. Mcm. \tV ern, Nat. I-Iist. ScHMIDT, 0., 1868. Die Spongicn dcr Kiiste von Algier. Soc., 2 (1): 67-122, pis. 3-16. Mit Nachtr5.gen zu den Spongicn des Adriatischen PALLAS, P. S., 1766. Elenchus Zoophytorum. The Hague, Nfeeres (drittcs Supplement). Engelmann, Leipzig: P. van Cleef: 1-451. i-iv, 1-4'1, pis. 1-5. PARKER, G. H., 1910. The reactions of sponges, with a --, 1870. Grundziige einer Spongien-Fauna des Atlan­ consideration of the origin of the nervous system. J. tischen Gebietes. Engelmann, Leipzig: i-iv, 1-88, Exp. Zoo!., 8: 1-41. pis. 1-6.

Figs. 87-93. Fig. 87. H)'Jneniacidon carunwla Bower bank, 1866 ( = H. perlevis (lvlontagu, 1818)), A-B. habit and spicules (redrawn from Bowcrbank, 1874: pl. 32 fig. 1) (s.b. = 1 em; s.b. spicules = 100 fJ.tll), C. tangential view of ectosomal skeleton

(s.b. 100 fJ.m), D. blunt apices of styles to show faint tylotc swellings) (s.b. = 10 fJ.m). Fig. 88. Nailondra (? = Hymeniacidon) maza De Laubenfels (1954), A. tangential view of ectosornal skeleton (s.b. 100 fJ.m), B. cross section of peripheral region (s.b. = 1 mm), C. detail of B (s.b. = 0.5 mm), D. spicules (s.b. 100 fJ.m), E. blunt apices of stylotes (s.b. = 25 fJ.m) (all made from the USNM holotype). Fig. 89. Rhaphidostyla ( = Hymeniacidon) kitchingi Burton (1935), A. tangential view of surface skeleton (s.b. = 0,5 mm), B. cross section of peripheral region (s.b. = 1 mm), C. spicules (s.b. =50 fJ.m), D. apices of spicules (s.b. = 10 fJ.rn) (all made from the BMNH holotypc). Fig. 90. Batzella ( = Hymeniacidon) corticata Thiele (1905), spicules (redrawn from Thiele, 1905: fig. 58) (s.b. = 50 fJ.m). Fig. 91. Sty!oteila ( = Hymeniacidon) agminata Von Lcndenfeld (1888), A. habit (s.b. = 1 em), B. spicules (s.b. = 100 fJ.m), (both redrawn from Ridley, 1884: pl. 41 fig. E and pl. 53 fig. f). Fig. 92. Thie/eia ( = Hymeniacidon) rubiginosa (Thiele, 1905), spicule (s.b. = 50 fJ.m) (redrawn from Thiele, 1905: fig. 44). Fig. 93. Uritaia ( = H;•meniacidon) halichondrioides Burton (1932), A. habit (s.b. = 1 em), B. spicules (s.b. = 100 fJ.m) (both redrawn from Burton, 1932: pl. 7 fig. 3 and text-fig. 4).

61 --, 1875. Spongien. Jahresber. Cumm. wiss. Unter~ ~-, 1927. Diagnoses d'epongcs nouvelles recueillies par such. Deutsch. l\!Ieere, 2<3: 115-120. le Prince Albert Ier de f..'lonaco. Bull. Inst. SoEST, R. W. l\!I. VAN, 1980. l'vlarine sponges from Oceanogr. Monaco, 502: 1-19. Cura<;ao and other Caribbean localities. Part II. --, 1928. Spongiaires de !'Atlantique ct de Ia Mediter­ Haplosc\erida. Studies Fauna Cunu;ao Caribb. Isl., ranCc provenant des croisiCrcs de Prince Albert 1cr 62 (91): 1-173, 18 pis. de Monaco. Res. Camp. Sci. Prince f..'Ionaco, 74: 1- --, 1987. Phylogenetic exercises with monophyletic 376, ph. I-XI. groups of sponges. In: J. Vacclet & N. Boury­ TsuRNAi\fAL, M., 1969. Four new species of fvlcditena­ Esnault, eds.: Taxonomy of Porifera from the N.E. nean Demospongiae and new data on Callites lacazii Atlantic and l'vleditcrranean Sea. NATO ASI Series, Schmidt. Cah. Bioi. f..'lar., 10: 343-357. t:l: 227-2'11. VACF.T.ET' J. & p. v ASSEUR, 1971. Epunges des recifS cor­ --, 1990a. Towards a phylogenetic classification of alliens de Tulear (Madagascar). Tethys, Sup pl. 1: sponges. Proc. 3rd Int. Conf. Bioi. Sponges, \•Vuods 51-126, 4 pis. Hole, 1985 (in press). V ACELET,]., P. VASSEUR, & C. LEv1, 1976. Spongiain:s de --, 1990b. Demosponge higher taxa classification reex­ la pcnte externe des rCcifs coralliens de TulCar (sud­ amined. In: H. Keupp &]. E. Reitner, eels.: Fossil ouest de Madagascar). MCm. Mus. nation. Hist. & Recent Sponges (in press). nat. Paris, (A, Zool.) 99: 1-116, 10 pls. SoEST, R. V•/, M. VAN & D. B. SAss, 1981. 1\'Iarine sponges VAN Soi-:ST, R. \-'l. M. see SoEsT, R. W. lVL VAN. from an island cave on San Salvador Island, VETHAAK, A. D., R. J. CRONIE & R. \'V. f..11. VAN SoEST, Bahamas. Bijdr. Dierk., 51 (2): 332-344, 2 pls., 4 1982. Ecology and distribution of two sympatric, figs. closely related sponge species, Halichondria panicea SoEST, R. w. rvl. VAN & N. STENTOFT, 1988. Barbados (Pallas, 1766) and H. bmverbanki Burton, 1930 deep water sponges. Stud. Fauna Cur. Carib b. Isl., (Porifera, Dcmospongiae), with remarks on their 70: 1-175, 12 pis. speciation. Bijdr. Dicrk., 52 (2): 82-102. SoEsT, R. W. M. VAN & S. \'\'EINBERG, 1981. A note on the VosMAER, G. C. J., 1886 [1887j. Porifera. In: Bronn, H. sponges and octocorals from Sherkin Island and G., Die Klassen und Ordnungen des Thierrcichs, 2: Lough Inc, Co. Cork. Irish Naturalist's J., 20 (1): 369-49/i, pis. XXVI-XXXIV. (Title dated 1887). 1-15. VVAPSTRA, M. & R. \·V. f..if. VAN SoEST, 1987. Sexual SoEST, R. \•V. f..1. VAN & S. ZEA, 1986. A new sublithistid reproduction, larval morphology and behaviour in sponge Monanthus ciocalyptoides n. sp. (Porifera, Demospongiac from the south west of the Halichondrida) frorn the \-'lest Indian region. Bull. Netherlands. In: j. Vacelet & N. Boury-Esnault, Zool. Mus. Univ. Amsterdam, 10 ('!): 201-205. eds.: Taxonomy of Porifera fi·om the N.E. Atlantic SaLLAS, I. B.]., 1902. On the sponges collected during and f..