Systematics of the Euretidae (Porifera: Hexactnellida: Hexactinosa)

SYSTEMATICS OF THE EURETIDAE (PORIFERA: HEXACTNELLIDA: HEXACTINOSA)

Benjamin Wheeler

Redpath Museum and the Department of Biology

McGi 11 University, Montreal February , 1 999

"A rhesis szrbmilted to the Facuhy of Graduute S~udiesand Reseurch in parliai fui$llmen( cf rhe reqtrirernents of the degree of Master of Science"

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TABLEOF CONTENTS ...... ,7 ABSTRACT...... 3 RESUME...... 3 ACKNOWLEDGMENTS...... 4 INTRODUCTION ...... 5 LITERATUREREVIEW ...... ,...... 7 METHODS...... 25 SYSTEMATICS...... 28 KEYTO GENERA...... 34 Subfamily Euretinae ZIITEL ...... 35 EURETE SEMPER, 1868 ...... 35 CALY PTORETE OKADA, 1 925 ...... 42 CONOFtETE IJIMA, 1927 ...... 47 BATHYXIPHUS SCHULZE, 1899 ...... 54 PARARETE IJIMA, 1 927 ...... -58 PLEUROCHORIUM SCHRAMMEN, 191 2 ...... 64 VERRUCOCOELOIDEA REID, 1969 ...... 68 -. Subfamily Iphitinae (n. sf.)...... 12 IPHITEON BO WERBANK, 1869A ...... 72 CHONELASMA SCHULZE. 1887~...... 79 LEFROYELLA THOMSON. 1 877 ...... 85 MYLIUSIA GRAY . 1 859 ...... -92 PERIPHRAGELLA MARSHALL. 1875 ...... 100 PTYCHODESIA SCHRAMMEN, 19 12 ...... 105 .i Heterorete-group "...... 111 HETERORETE DENDY . 19 1 6 ...... 111 GYMNORETE IJIMA, 1927 ...... iI5 ENDORETE TOPSENT, 1928...... 120 Drscljssro~...... 124 GEOGRAPHICDISTRIBUTION ...... 134 CONCLUSIONS...... 134 REFERENCES...... 136 APPENDIXA ...... 144 Abstract A long overdue taxonomic revision is performed on the family Euretidae (Hexactineltida). World Iiterature is compiled, world type material is gathered, prepared, re-examined, measured. described and illustrated, tauonomic histories and world species distributions are provided. A generic key to the Euretidae is also provided. Of the 19 generic names associated with the Euretidae at one time or another, 16 are recognised here (Buthpiphus. Calyprorere. Chonciusma. Cortorere. Endarete, Eurece, Gymnorere. Heterorete, iphiteon, Myliwia. Pararele. Periphrugelfa, P~ezrrochorirrm,P~chodesia. Lefioyella and Verrucocoeloidea). Syringidium is put in to sy nonymy with Lefioyellq Joanella and MarguritelZa are synonym ized w itli /phireon: two subspecies and one form are elevated to species sbtus; three new species combinations are presented and two new species are named. Phylogenetic analysis suggests the majority of morphological euretid characters are pi-one to homoplasy and are not useful for phylogenetic classitication. Subfamilies based on channelization and the presence or absence of key spicules are designated.

Résumé Une révision taxonomique longuement attendue a été esécutée sur la famille Euretidae (Hesactinellida). Mondialement. la littérature est compilée. le matériel de type du monde est recueil1 i. préparé, esam iné de nouveau. mesuré, décrit, et illustrées; les histoires ta,,onomiques et les distributions des espèces du monde sont fournies. Une clée générique de la famille I'Euretidae est également fournie. Des 19 noms génériques associés à I'Euretidae en même temps ou encore. 16 sont reconnus ici (Bathyxiphlrs, Culyprorere, Chonelasma. Conorete. Endorele. Ezrt-ere, Gynrnorefe.Heterorete, @hiteon, Mvliusia. Pararete. Periphragella, Plcurochorirrrn. Piychodesia. Lefioyella et Vernrcocoeloidea).Syrirlgidizm est m is dans la synon y m ie avec L~'fioyclZu;Joanella et Margaritella sont mis dans synonymie avec /phireon; deux sous-espèces et une forme sont élevées au status d'espèce; trois nouvelles combinaisons d'espèce sont présentées et deus nouvelles espèces sont nommées. L'analyse phylogénétique suggère que la majorité des caractères morphologiques des euretids soient sujets a I'homoplasy et ne sont donc pas utile pour la c!assification phylogénétique. Sous-familles basées sur le channelization et la présence ou absence des spicules principaux sont indiqués. Ackno wledgments

Henry Reiswig provided me not only with access to his extensive specimen and iiterature collections, but also to his valuable time and patience (especially with the synonomies!). Without his positive attitude, encouragement and support this study would not have been possible. I'd like to acknowledge Dr. G. Bell and Dr. D. Green for their participation in this project and the "Biodiversity lab" for the use of their advanced technology and positive atmosphere. Ttianks also goes to Jason for his helpful cladistic input. 1 would also Iike to acknowledge the BMNH. the Leiden museum. Musée de Monaco and MC2 for their valuable specimen loans. For financial support 1 would like to recognize NSERC and McGill University.

Ingrid, if I could only express how much your help, happiness and reassurances have contnbuted to this project! Thanks for making my Master's a wonderful experience. I'd also like to extend my gratitude to the Anand family for their reassuring support and nurnerous nourishing "take-home" meals! ! Harry. your bright positivity canied me throughout this project. What more cm 1 say but THANKS!! Finally, I'd Iike to thank Chris and Susan, for without whom. I'd no longer have my best friend (Hi Bau-xi!) and would still be somewhere in India!!

I'd like to dedicate this thesis to my parents whose example and support defies imagination and great distances. ln troduction

Hexactinellids, or glass-sponges, are perhaps one of the more poorly understood and unrecognised marine invertebrate groups. Due to thsir remote abyssal or bathyal habitat, little is known about modes of reproduction. development, diversity, geographic distri bution and evolutionary relationships. Perhaps more practically, hexactinellids have remained obscure due to their unstable taxonomy. which, in turn. has complicated the identification of newly collected material. The majority (approx. 50%) of hexactinellid genera are based upon single. rarely complete. specimens crudely collected and described ofien over one hundred years ago. In fact. nearly 70% of hexactinellid specimens recorded in the literature were descnbed before 1930. To fùrther aggravate matters. historicall y, hexactinellids have not been widely known and many earl y researchers mistakenly identified hexactinellids as "zoophytes", plants or sea-cucumbers. Needless to Say that. of those researchers that did recognise the sponge affinities of hesactineilids. few followed a single. well-defined. morphological terminology. Also. at the time wlien the majority of the hexactinellid descriptions were published. forma1 international zoological taxonomic codes were just forming and. for the most part. were not in practice and this unfortunately added yet another level of instability to hexactinellid taxonomy. Discovery of hexactinellid-like forms in the famous Ediacaran fauna has established hexactinelfids as the oldest known members of the Animalia and lends evidence to their role as ancestors of the Metazoans. Equally important has been the confirmation of the syncytial. or multi-nucleate. organisation possessed by hexactinellids which lends fùrther evidence to their roles in early Metazoan diversification and also nominates them as candidates for an, or the, origin of multicellularity. In order to further our understanding of these unique organisms a systematic revision of their tavonomy is

pararnount, for, as Simpson (1 945) astutely remarked ". . . animals cannot be discussed or treated in a scientific way until some taxonomy has been achieved.. .". The aim of this study is therefore to aid in the taxonomic revision of the liexactinellids by thoroughly reviewing one large farnily of hexactinellids. the Euretidae. Historicaily, the Euretidae has been differentiated from the five other families within the order Hexactinosa by the presence of 'broom-fork'. or scopule. spicules and unchannelized three-dimensional skeletal frameworks, however, this family has become a convenient place for any sponge with a three-dimensional skeletai framework, with or without skeletal channelization or scopule spicules. Since its conception in 1877, eleven international authors have erected and placed genera within the family Euretidae and over eighteen authors have descnbed euretid species. The Euretidae had attained a maximum of sixteen genera, many of which were based upon single. ofien fiagmentary specirnens, and several generic type species were based upon more than one specimen. The objectives of this revision are to stabilise the taxonomy of the Euretidae by locating and identiwing important type specimens from world museum collections. reviewing the taxonomie literature of each genus, and applying modem andytical techniques (filtration and image analysis) to a group of sponges known primarily frorn the 1 9'h century. This snidy will be the first to analyse al1 euretid genera together at one time, and, as a result. to provide the opportunity to apply modem systematic techniques in liopes of suggesting a more 'phylogenetic' classification and to analyse character homoplasy, a factor which cannot be ignored. Also, geographical distribution maps and an artificial key will be produced to aid in identification. Hopefully this study will provide some insight into the diversity of one family of hexactinellids and succeed in stirnulating awareness and interest into their unique and important biology. Literature review

Current ta.xonomic designation (Bergquist, 1978; Vacelet, 1994; Levi, 1 997; Reiswig, 1994; Hooper & Weidenmayer, 1994) recognises the glass sponges. or Hexactinellida, as one of three classes within the phylum Porifera (Bergquist, 1978; Vacelet. 1985; Hooper & Weidenmayer, 1994). Hexactinellids differ considerably from the classes Calcarea and Demospongiae in basic organisation both at the structural and cellular level. Structurally, hexactinellids possess hexactine, or six-rayed. siliceous skeletal elements not found in any other group of sponges. At the cellular level, researchers (Schulze, 1887; Ijima, 1903; Okada, 1928; Mackie & Singla. 1983; Boury- Esnault & Vacelet, 1994) have discovered that hexactinellids are syncytial in organisation; a striking contrast to both the cellular Demospongiae and Calcarea. As a result. some authors have suggested the Hexactinellida deserve Subphylum (Reiswig. 1979) or Phylum (Bergquist, 1985) status.

Basic Biology Unfortunately, basic hexactinellid biology is not well understood. Although several early authors (Schulze. 1880; 1887; Ijima, 190 1, 1903) attempted soft-tissue examinations, it wasn't until the application of the electron microscope that hexactinellid ~Itrastructurewas confidently revealed. The first authors (Reiswig. 1979; 1991 : Mackie & Singla. 1983) to undertake this new investigation not only confimed suspicions of Schulze and Ijima of syncytiality but also discovered several unique morphoIogica1 features within the lyssacine Rhabdocalyprus dawsoni Lambe: 1 ) collar-bearing tissue significantly different from that found in Demosponges and Calcareous sponges; 2) cells and tissues interconnected by cytoplasmic bridges containing plugs; 3) spicules secreted intracellularly in multinucleate giant cells and, 4) absence of pinacocytes found in other sponges. These features, and one other significant feature. the presence of a secondary reticulum. have since been reconfirrned in Aulorossella vanhoeffeni (Salomon & Barthel. 1990), Furrea occa (Reiswig & Melil, 199 1 ), Ducrylocalyx pzrrnicezis (Reiswig. 199 1 : although no secondary reticulum was observed in D. ptrmicetrs) and Oopsucus minuta (Boury-Esnault & Vacelet 1994). Unfortunately, there have been no investigations into the soft tissues of members from the family Euretidae, upon which this project is focused. Although reproduction of hexactinellids has never been observed, an exciting report on Oopsacus minuta, a lyssacifie hexactinellid, corn the famous "deep-sea like" Mediterranean cave (Boury-Esnault & Vacelet, 1994) has significantly built upon Okada's (1 928) initial studies on the reproduction and deveIopment of Farrea solfasii. a dictyonine hexactinellid. Okada ( 1 928) observed that breeding occurred al1 year in F. soffusiiand descrïbed archaeocyte-like oocytes, spermatozoa, total and regular cleavage. a planula-like blastula. and a larval form. the younger stages of which were without flagellated chambers. Boury-Esnault & Vacelet ( 1994) reported similar findings in Oopsuczw minuta and found a "trichimet la" larvae similar in forrn to that of F. solim-ii but di ffering in the notable possession of multi flagellated cells not found in any other sponges. Obviously, more studies must be performed in order to establish modes of reproduction and larval dispersal of hexactinellids. Like most sponges, hexactinellids are filter feeders. Water is sucked through the dermal ostia into incurrent canals or spaces which lead through openings (prosopyles) to flagellated chambers (Wyeth et al.. 1996). As water flows through the sponge. particulate organic material and dissolved organic material are filtered out. Very little is known regarding the naturai food of hexactinellids. however. Mackie & Singla (1983) found evidence of phagocytosis of bacteria in Rhabdocalyptus dawsoni Lambe while Reiswig (1 990) suggested that R. dcrwsoni retains only dissolved organic carbon and Aphrocaflistes vustus fi lters both coIloidal particles and dissolved organic carbon. An elegant study by Wyeth et al. ( 1996) showed, by means of a special "Sandwich Culture". that Escherichia coli and Isochrysis galbana are rernoved from the water primarily in the region of the flagellated chambers. Leys & Mackie ( 1 994) have also suggested that vigorous cytoplasmic streaming seen in regenerating hexactinellid tissues may be used for nutrient translocation. making amoebocytic transport redundant. Perhaps the most significant findings of the past two decades regarding hexactinellid biology involves the SCUBA-accessible Rhabdocalyprus dmvsoni Lambe from the West coast of British Columbia. Mackie and Lawn (Mackie, 1979; Mackie et al., 1983; Lawn et al., 198 1) have provided physiological evidence showing that the syncytial network of Rhabdocafyptus dawsoni Lambe is capable of propagating electrical impulses. Certainly, more emphasis must be placed on obtaining more biological information on these enigmatic sponges. Although several authors (Schulze, 1887, 1899. 1904; Reid, 1968; Levi. 1964; Koltun, 1970) have provided depth (bathyrnetric) or geographical information of some. or all. of the known hexactinellid collections, it wasn't until 1994 that global hexactinellid distribution was discussed in detail (Tabachnick. 1994). Using material from over three hundred publications and unpublished data, Tabachnick (1 994) performed a biogeographical analysis and concluded that hexactinellid distribution may be partitioned into five latitudinal bathyal zones. As well as confirming Levi's (1964) earlier observations that hexactinellids are prirnarily bathyal (200 - 2000rn below sea level (Gage & Tyler, 1991 )) and not abyssal (> 2000 m depth) organisms, Tabachnick (1 994) found that many taxonomie units of the order Hexactinosa (in which the Euretidae is placed) are distributed within low latitudes whereas they are absent in al1 other regions. Although not conclusive. Tabachnick (1 994) also performed a geographical cluster analysis of 12 regional hexactinellid faunas. Unfortunately. Tabachnick (1 994) did not discuss the artifact of collection nor possible factors affecting hexactinellid distribution. However. Reid (1 968) did show that geographical aspects of recent hexactinellid distribution did not show correlation wi th low temperatures and offered several other limi ting factors: (a). light: (b), physical or physico-chernical disturbance; (c), high oxygen levels: and (d) two or more of these factors together.

Spicules The class Hexactinellida, like the lithistid demosponges, contain members where fusion of siliceous skeletal elements, spicules, forms a permanent framework. Within the Hexactinellida, the order Hexactinosa encompasses those sponges with rigid and permanent frameworks. Members of the family Euretidae, one of five farnilies within the Hexactinosa and the focus of this project, typically show cup, fume1 or tube shapes. however other shapes are observed (Bathyxiphus subtilis Schulze, Pterrrochorium ~tnnundulciKirkpatrick). It is upon the body shape, spicules and frameworks that traditional (Zittel 1877. 1878; Schulze 1885, 1886, 1 887, 1904; Schrarnmen 1910, 1924, 1936; Ijima 1 90 1-3. 1 937; Reid 1964) systematic classifications among hexactinellids rely.

Spicule Formation Almost al1 siliceous sponge spicules contain a definite organic axial filament system; when hexactinellid spicules are viewed in cross section. the axial canal (fomed by the axial filament) is regularly square (Reiswig, 1 97 1 ). Although the role of axial canai shape in spicule formation is not known, it must be an important charactenstic for al1 siliceous demosponge spicules have triangular (Hooper & Weidenrnayer, 1994) axial callds- Although it is believed (Reiswig, 1971; Bergquist, 1978: Levi. 1989: etc.) that spicules are formed by deposition of silica onto an organic filament. there is some confusion as to where this deposition takes place. Recently, BOUN-Esnauit& Vacelet ( 1994) confirmed that the early stauractin spicules of Oopsacza minuta first appear as a pseudocrystailine axial filament surrounded by a silicalemrna in a process not unlike that observed in the demosponges, thereby indicating a scleroblast (cells reponsible for spicule formation) depositional origin. However, according to several studies (Ijirna. 190 1 ; Schulze. 1904; Okada, 1928; Reiswig. 199 1; Reiswig & Mehl. 1994) that have analysed the soft tissues of hexactinellids. spicule secretion takes place intrasyncytially . Regarding the order of spicule type development, Meis known and there are only a few early studies on this subject (Schulze, 1887; Ijima, 190 1 ; Okada, 1928). Smooth stauractins appear to be the first spicules secreted in larval forms, as noted by Okada ( 1928) in F. sollasii and confirmed by Boury-Esnault & Vacelet ( 1994) in Oopsaczrs rninzrtu. however, the order of appearance of other types of spicules rernains vague. as Okada (1 928) based his "Order of Formation" from subjectively detemined larval stages. Nonetheless. Okada (1 928) did offer the following order of spicule formation of Farrea sollasii: a) stauractins; b) hexasters (not clear if oxyhexasters or discohexasters or both);

C) dermal pentactins; d) hexasters (?); e) uncinates; f) dermal clavules: g) gastral pentactins and h) gastral clavules. Likewise, the process of spicule development has seldom been studied (Schulze. 1 887; Ij ima, 190 1; Okada, 1928), and therefore is not well understood. Perhaps the application of new techniques, such as the isolation of axial filaments develojxd by Schmizu (1 998), to hexactinellids will shed light on spicule development-

Spicule Size Generally, spicules form two groups, the distinction based mainly upon size. The Iarger spicules are the main supporting elements and are called skeletal spicules or megascleres (Figure 1. A); they can range fiom several hundred micrometers to several tens of centimeters long. The smaller spicule forms. which occur between the megascleres. are known as microscleres (Figure 1. B) and range from tens to several hundred micrometers in diarnetre. Although data on hexactinellid spicule sizes have not been consistently exarnined, it is believed (Bergquist, 1978; Reiswig, personal communication) that in some species, distinct size classes exist within individual spicule types.

Spicule Shape The geometry of megascleres and microscleres in Hexactinellida is very diverse. In general. spicules are named primarily according to (i) their symmetry. as judged from the arrangement of the axial filaments, and (ii) the number of rays present (Reid, 1964). As the narne hexactinellid implies, the common megasclere spicule is six-rayed and is termed a hexactin. It is widely believed (Ijima, 1927) that this six-ray spicule is the template on which al1 other hexactinellid types are based. A pentactin, for example. is simply a hexactin spicule in which one ray is reduced. Spicules Vary from the central hexactin type (Figure 1) in one or more of three ways: (a) through reduced development of one or more rays. (b) through reduction of rays accompanied by development of special terminal spines, and (c) through development of lateral spines, without associated reduction of rays (Reid, 1964). The basic type of microsclere found in the family Euretidae is the hexaster. however, the many forms of hexasters (Figure 1) are not believed to have been derived through reduction of rays (as in the megascleres) but by formation of terminal ray appendages (Ijima, 1927). uncinates

Firrire 1. Diagrammatic rcpresentations ofliexactinellid (A) nlegascleres; (B) n~icroscleres;(C) sceptriiles (Taken from Hooper Br Weidenmayer. 1 994).

IHe?ractinellid microscleres are always triaxial in origin. and they are apparently sccreted in the same way as rnegascleres (this observation taken from Scimrclinnia crrcricir Scl~iilze,1900 and Okada, 1928). Common eiiretid microscleres include discoliesasters, osyhexastcrs, onycliohexasters and tyloliexasters. Ar?c!lier spiciile type is the nionactin, in which only one of the original rays reniain and the axial cross is foiind at the end of the ray (Reid, 1964). As axial filaments have not been found in the appendages of monactines it remains unclear whether the appendages are homologous to the rays of the hexactine or bexaster. Cornmon monactine spicules are the sceptrules of which there are four types (Figure 1, C): sarules. scopules. lonchioles and clavules (Ijima, 1927). Sceptrule presence is an important feature for higher classitkation of hexactinosan sponges: for example. those sponges possessing scopules are known as the suborder Scopularia. in which the family Euretidae is placed, and those possessing clavules are known as the suborder Clavularia. MorphologicalIy. the scopule cmbe divided into three regions (Figure 2): (a), the tines. which. like the secondary rays of the hexasten. have not been found with axial canals; (b). the capitulurn, where the axial cross (the junction of remnant axial canals) is ofien observed and as a result is ofien swollen or knobbed. and (c). the sliafi; which may be straight, curved, rnicrospined, spined or smooth and proximally pointed or swollen. Tines are generally straight. geniculate (bent). microspined or smooth and may Vary from two to fifieen in number. Within the Euretidae five different scopule types have been observed (Figure 2). The most common scopule is the tyloscopule (Figure 2, 'T'). so narned for the noticeable 'tyle'. or club-like swelIing on the distal end of the tines. Ofien this 'tyle' is adomed with distinct, prosimally directed. recurved thoms. but the most distal portion is left smooth resulting in a 'bald' appearance. Strongylscopules (Figure 2. OS')are characterised by their straight. undivergent. and often thick microspined tines. Oxyscopules (Figure 2. '0') have straight. undivergent and distally pointed tines and may resemble another type of sceptrule. the sarule, but are distinguished by the location of the axial cross. Although not common. the subtyloscopule (Figure 2, 'St') is found in some euretid species. This scopule has a small but noticeable 'ball' on distal tine tips. The discoscopule (Figure 2, 'Da) is distinguished by the serrated caps on the distal tips of the tines. Closely resembling the monactines are the uncinate spicules (Figure 1 ) The presence of an axial filament has not been established in the hexactinosan uncinates and consequently their relations to other spicule types are unknown. The presence or absence of uncinates has been used by Schulze (1887) in his classification of the Hexactinosa: those with uncinates, the Uncinataria, and those without. the Inermia; however. this scheme is no longer accepted. Figure 2. Diagrmatic representation of scopule morplioloçy and ewetid scopule types (ti, tines; c, capitulum; sh, shaft; T, tyl~scopule:S. strongyloscopule: 0. o~yscopule;DI discoscopule; St, subtyloscopule).

Another feature, which varies greatly, is the degree of oriianicniation on a spiculc. This is usually seen as the presence or absence of spines. and tlic quantity and size of spines (Bergquist? 1978). The terminology associated witli tliesc spines (wlicn found on bcanis and spicules only), from srnall to large. is 'microspines' ( 1-2 prn in leiiptli). 'spines' (2-5 pm) and 'tliorns' (>5 pi).

Spicule Location Individual spicules, or spicules fused into networks, as is the case in the family Euretidae, dictate the form of the sponge. Particular spicule types occur in specific locations within the skeleton and these locations are given specific terms as sliown in the

t accompanying diagram (Figure 3). Generally, tliree zones are diffcrentiated: the first lying on or just below the dermal membrane (demal); the second lying in the interior of ihe trabeculae and flagellated chamber layer (parenchymal); and the tiiird lying bclow the riicinbrane lining the atrial cavity (gastral).

Fieure 3. Diagrammatic representation of a cross-section througli a wali ot'a typicd èiiretid (Di. dictyoiialia; D, dermalia; G, gastralia; P. parciicliyriialia).

Skeletal Framework Althougli not proven, it is thought tliat al1 hexactinellid spicules are separate frorn one anotlier when first formed and in many cases rernain separate throughout life. Tiiose sponges wliose megascleres do not regularly fuse by direct connection are termed lyssacinc (Ordcr Lyssacinosa, Ijima = Order Lyssacina, Zittcl). Sonic Iicsactiricllid spiculcs (typically megascleres) regularly become connected rigidly in three dimensions, thus forming a framework. The order Hexactinosa Schrarnrnen (iDictyonina Zittel). in which the Euretidae is placed. contains those sponges that form rigid. three-dimensionai 'dictyonal' frameworks - the 'dictyonine' sponges. As the process of fossilisation favours ngid structures, the majority of the hexactinellid fossil record is of the dictyonine sponges and paiaeontologists (Reid. Reitner, Mehl) have endeavoured to describe and characterise these complex three-dimensional structures. Indeed, in his important work "The Upper Crebceous Hexactinellida" Reid (1 964) perceived, defined and illustrated many significant framework features and offered hypotheses on the development of these features. Unfortunately, the complexity and tenninology of hexactinosan frameworks restricts an adequate review of these features, however. several key features have been selected in order to enable the reader to differentiate between euretids. Conceming the method of spicule fusion, Reid (1 963, 1964) suggested three possibilities: (i) enclosure of two parallel rays in a cornmon silica envelope, (ii) attachment of the tips of rays to other dictyonal beams or (iii) simple fusion of rays at arbitrary points (Reid, 1964). In the first case. the meshwork (Figure 4. 'm.) resulting from the fusion of the dictyonalia rnegascleres is ofien square or rectangular. Reid (1964) also pointed out that the common enclosure of parallel rays (by means of a silica envelope) of neighbouring megascleres forms a strand-like structure or dictyonal strand (Figure 4. 'ds'). Reid (1964) considered al1 dictyonalia belonging to the senes from which the dictyonal strands are forrned as primary dictyonalia. In contrast, al1 dictyonaiia which do not belong to the dictyonal strands (e.g. adcied on later) are considered as secondary dictyonalia. This is an important concept. for several euretid frameworks possess both primary and secondary dictyonalia. Upon the general architecture of the hexactinosan dictyonaI frameworks Reid (1 964) then recognised tliree principal types: (1) Farreoid; (2),Euretoid and (3). Aulocalycoid skeIetons. In Reid's 'euretoid' skeletons. the primary meshwork is itself three- dimensional. with one to many meshes between its derrnal and gastral surfaces. and has dictyonal strands distributed through its whole depth. without layered arrangement. and with their direction of growth obliquely longitudinal to radial (Reid. 1964). Note should be made here that this author is in disagreement with Reid (1964) and feels that dictyonal frameworks may be layered. Ficure 4. Diagrammatic 'cut-away' of a typical euretoid skeleton sliowing irregular ('D'. 'G') franiework surfaces surrounding regular ('1') interna1 framework (ac, asid cross; b. beam; D, irregular derma1 framework; dc, example of a dictyonal cortex; ds, dictyonal strand; G, irregular gastral framework; 1, regular intemal franiework (shaded): m, mesh; 11, node).

Reid's definition of the 'curetoid' skeleton was later (Reid, 1964) cspanded to include thosc dictyonalia which occur in laterally corresponding positions so tliat tlieir transvcrsely aligned rays unite to form successive transverse skeletal Imieilae. Anotlier important concept is tliat of the 'dictyonal cortex' (Figure 4. 'dc'). upon wliich Reid (1 964) Iiad this to Say: " A cortex may generally be said to be present when the outermost 9 iiieshwork of the framework is different in appearance from ihat seen bcncath, and wlien this is not due simply to differences in orientation." (Reid, 1964: 1-xxxix). Dictyonal cortices are often observed in members of the farnily Euretidae (Neterorete Dendy) and appear to be caused by the secondary fusion of small oxyhexactins, thereby foming triangular or irregularly shaped meshwork (Figure 4, 'D'. 'G'). As neigbouring dictyonaiia are joined together by secondary siIica deposition it is often difficult to discern the original dictyonaiia. or nodes (Figure 4, 'n') of the framework. Generally, nodes may be considered as either tme, or false, nodes. Tme nodes are those that have the original (six) number of rays (where the axial cross (Figure 4. 'ac') is present) and false nodes are those having greater or less than sis bearns (Figure 4. 'b') ernanating frorn it (axial cross is absent). Those dictyonal rays that occur at the extremities of the dictyonal Framework, and are not fused to other dictyonalia. are ofien ' finger-like' and are termed spurs (Figure 4, -s'). Obviously, this short review grossly oversimplifies the nature of these very complex structures. The complexity of these structures significantly increcises when the body walls of the sponge fold together, as appears to be the case in several euretids, or when channelization occurs.

Channelization In sponges within the order Hexactinosa water passes through normal skeletal meshes via channels that are ofien, but not always, larger in diameter than that of the skeletal mesh. If the channels are greater in diameter than the skeletal mesh. they are preserved in skeletal remains and can be used as classification characters. According to Reid (1 964), channelization may be of two main types: (1) intradictyonal. occuring in the primary framework; and, (2) extradictyonal, occuring only in the secondary framework. The following definitions of types of channelization follow primarily from Reid (1 964) but also draw upon the work of Ijima (1927). Epirhyses (Figure 5) are channels that penetrate the dermal (or external) surface perpendicularly. their blind end being situated close to the gastral (or intemal) surface (Ijima, 1927). Aporhyses (Figure 5) are similar channels that penetrate the gastral (or intemal) surface perpendicularly, their blind end situated close to the dermal surface (Ijima, 1927). Diplorhysis (Figure 5) is the condition in which the primary fiamework shows two sets of deeply penetrating intradictyonal cavities. which open on opposite sides of the framework and are inhalant and exhalant respectively (Reid, 1964). Diarhysis (Figure 5) is the condition where both dermal and gastral surfaces are direcd y comected by means of a channel. Arnararliy ses (Figure 5) is tlic terin given to tunnel-like canals of varying length wliich mn longitudinally witIi iiiten& of an indefiiiite widt11 between theni; are open on tlie gastral surfacc by dit-likc apertures; and where tliere exist ledge-like or papilla-likc proniineiices on the derttial side of tlie wall, send out radial branch canals into these to open accessory oscula on their top (Ijirna, 1927; see Plate 13 for diagrammatic representation of arnararliysis). The term 'cavaedial space' is used for those spaces enclosed by folding of the body waIl.

sis

Firiure 5: Diagrammatic "peel-away" representation of corninon cliannelization in Iicxact inosan spoiiges.

Hexactinellid Relationships Due to the bathyal and abyssal habitat of llesactinellids fresli specimens are diflicult to obtain and, tlierefore, to study. For tliis reasoii, relationsliips anlong estant Iiexactinel lids have remained large1y unexamined. However, using Ij ima7s(1927) and Schulze's (1 904) classification scliemes of extant niaterial, Salomon (1988) outlined some phylogenetic problenis witliin tlie 1-icxactiiiellida (~i~uré6): (a), the use of the presence of ampliidisc and Iiexaster spicules to divide the wliole class into two groups is problematic since they are not believed to be lioniologous (Reid, 1958); (b), the observation (Ijima? 1927) of ioose lyclmisc spicules in yoiing forms ofrlulocysris riflefi suggests that fusion 01-spiculcs (Iy~liriiscsaiid hesactines) into dictyonal frarneworks occurred twice. not a very parsimonious situation; (c) the division of the Hexactinosa into two groups (Scopularia and Clavularia) based on sceptrule presence fails in the light of Claviscop~diuinfernlediu, wliicii possesses both clavulcs and scopules, (d), the presence or absence of uncinate spicules docs not suggcst distinct groups, (e), nor does the presence or absence of microscleres suggest distinct groups.

HEXACTiNOSA

TAKEN FROM SALOMON, 1988

Figure 6. Pliylogenetic tree depicting hexactinellid relationships (AU, Aulocalycidae; AP. AphrocaIIistidaé; TR, Tretodictyidae; CR, Craticulariidae; EU. Euretidae; FA, Farreidae; LK. Lycliniscosa; LS, Lyssacinosa; l-iY, Hyalonematidae; MO, Moiiorhapliidae; PH, Plieronematidae; DS, dictyonal skeleton - nunibered cliaracters as follows: 1, stauractins; 2, uncinates; 3, reduction of parenchymalia to rliabdodiactines 3a. sceptres; 3b, tauactines; 4, loss of uncinates; 5, sarules; 6, loncliiole; 7, dictyorial skeleton witli more than one layer; 8, triangular dictyonal meslies; 9, cliannels in dictyonal skeleton; 10, "rnultiradial" dictyonal meslies; 1 1. epirhysis and aporhysis; 12. schizorliysis; 13, diarhysis). In a further attempt to analyse hexactinellid phylogeny, Mehl (1992) combined fossil and extant evidence and reconfirmed the two groups Clamlaria and Scopularia within the Sceptnilophora (Salomon, 1988). Using scopule type as a character, Mehl's ( 1992) phylogenetic analysis grouped several rnembers of the Euretidae with genera belonging to other families (e.g. A4phrocallistesGray with Chonelasma Schulze. Periphrugella Marshall, Pararete Ijima, Pleurochorium Ki rkpatric k and Eurere Semper). Mehl (1992) also reviewed the taxonomy of several ewetid genera, however, as her decisions did not incorporate original type material, her results must be considered inconclusive. Although littie attention has been paid to hexactinellid phylogeny. hexactinellids have been used in recent studies (Van Soest, 1987; West & Powers, 1993; Cavalier-Smith et al.. 1 996; Reitner & Mehl. 1 996; Koziol et al.. 1997: Borchiellini et al.. 1 998: Kruse et al.. 1998) analysing poriferan phylogeny. In a reassessment of Hartrnan's (1 982) classification, Van Soest (1 987) used hexactinellids as an outgroup on the basis of their syncytial nature. Cavalier-Smith et al. (1 996). using West and Power's (1 993) molecular information on Farrea occu (Farreidae), suggested that sponges as a whole are monophyletic with the demosponges and hexactinellids forming a monophyletic group of siliceous sponges. Using thirty-one biochemical and morphological characters ranging from the presence of aggregating factors to the "parenchymella larvae" of Farrea sollasii. Reitner & Mehl (1 996) established hexactinellids as the sister taon of the Pinacophora (Calcarea + Hornosclerornorpha + Demospongiae) and established the Porifera as the adelphotmon of the Eumetazoa/Placozoa. Afier cornparisons of genetic material for the Hsp70 (Heat Shock Protein) protein, Koziol et al. (1997) were unable to resolve the phylogenetic relationships between the three classes of sponges. BorchieIIini et al. (1 998)- using the same Hsp70 protein. confirmed Koziol's (1997) result and the monophyly of the Metazoa. Using the recently isolated protein Kinase C from Rhubdo~~lypt~sdaiosoni Larnbe, Kruse et al. (1 998) suggested that within the phy 1um Porifera, the class Hexactinellida diverged from a common ancestor to the Calcarea and Demospongiae, which both appeared later. Surprisingly, this study (Kruse et al., 1998) also suggested that the higher invertebrates (Drosophila melanoguster and Lytechinus pictus) are more closely related to the calcareous sponges. Fossil Information

Additional support for an early ongin of the HexactineIlida comes from paleontological evidence, which shows that hexactinellid-like fonns, probably in possession of stauractin spicules (Reitner & Mehl, 1995)?were present in the Ediacaran (Gelhing & Rigby, 1996; Steiner et al.. 1993) ages, before the well-known Burgess Shale deposits (ProfospongiaSalter and E~ffeeliiuWalcott) and "Cambrian Explosion". Use of paleontological evidence for the study of extant hexactinefiid diversification is problematic. As previously rnentioned. the process of fossilisation is biased towards ngid and permanent structures (i-e. the dictyonine sponges) and not those forms which are loosely held together (lyssacine sponges). This may well explain the observation that a cornparison of fossil and recent hexactinellid faunas shows a decrease in the dominance of forms with very rigid. fused skeletons and predominance of forms with non-fùsed skeletons today (Barthel & Tendal. 1994). The discovery of both amphidisc and hexaster spicules in sarnples from the Paleozoic time period (Mostler dL Mehl. 1990) suggests an early hexactinellid radiation. however. information concerning the further diversification of these rwo subclasses (Amphidiscophora and Hexasterophora) is limited. However, based upon his interpretation of the development of a spicule known only from the fossil record, the paraclavule, Finks (1 970) felt that the Hexasterophora and Amphidiscophora could have diverged from a common stock during the Paleozoic and that they could be the sole survivors of a number of early lineages in some of which both hexasters and amphidiscs conceivably could have occwred together. According to the "Fossil Record" (Benton, 1993). the oldest sponge considered as a niember of the family Euretidae, Plectospyris eleguns Sollas. comes from the mid- Jurassic. approximately 80 million years before the oldest known member of the family Farreidae, a sister group to the Euretidae. This evidence is not in accordance with Reid's (1 964) views that single mesh frameworks comrnon to members of the Farreidae are ancestral to the three-dimensional euretoid frameworks. S ystematics

Hexactinellid classification

The class Hexactinellida has been classified in many ways, however, rnost of these classifications are interrelated. The first widely accepted classification of hexactinellids was by Zittel(1877). In his scheme Zittel considered the fusion. or lack of same. between skeletal spicules as distinctive and developed the terms Dictyonina (hwd spicules) and Lyssacina (spicules not Fused). Although this system is generally thought unsatisfactory (Reid. 1964) some researchers (Bergquist, 1978) continue to use Dictyonina and Lyssacina at the ordinal level. Following Zittel. Schulze (1 885) initially incorporated the terms Dictyonina and Lyssacina but used them as subdivisions based on rnicrosclere type. Schulze later (1 887) decided to replace Lyssacina with the new names Amphidiscophora and Hexasterophora which refer to the amphidisc and hesaster microscleres. In short, the names Arnphidiscophora and Hexasterophora are now used to su bdi vide the class and the narne Dictyonina has changed to Hexactinosida (Schrarnmen), however. the term dictyonine is still used descriptively to refer to the regular fusion of spicules into a fiamework. The classification of the subclass Hexasterophora was formalized in Ijima's

( 1927) widely recognized (Reid. 1964: Koltun, 1967; Reiswig. 199 1 ; Barthel & Tendai. 1994) treatise of the "Siboga" expedition through division into three orders: Hexactinosida, Lychniscosida and Lyssacinosida. Sponges in the order Hexactinosida. in which the farnily Euretidae is found. are distinguished as having hexactine dictyonine niegascleres, the Lychniscosida having dictyonine lychniscs and the Lyssacinosida having free (or rarely fused) rhabdodiactin megascleres. Familial division within the order Hexactinosida relies prîmarily on the presence of scopute or clavule sceptrule spicules: suborder Scopularia and Clavularia respectively. Channelization has also been used as a key character in the classification 9f the Hexactinosida. Please refer to "Taxonomic Remarks" in the "Systematics" section for a systematic review of the Euretidae.

Phyfogenetic systematics The field of comparative biology has progressed considerably since the collection and description of most hexactinellids. As Willi Hennig. the founder of modem phylogenetic systematics, or cladistics, defined it. phylogenetic systematics rests on the assumption that descent with modification. or evolution, has taken place (Hennig. 1966: Janvier. 1984). The recognition of homologies (characters inherited from a comrnon ancestor) is a crucial issue in systematics because it is through homologies that relationships among taxa are inferred (Wiley, 198 1; Forey et al, 1992). However. to bridge the gap between homology. as descent from a common ancestor. and hornolopy as observable similarity, Hennig (1 966) proposed using only those evolutionarily novel cliaracters that are shared between taxa (synapomorphies) to discover phylogenetic relationships.

"Birr the 'problem o/homology' is broken by simpiy realking fhar homologies con be rreared as hypo!lwses which ure resred by orher bporhmes o/homology and rheir associaredp@logeneric lypoihcses. " Takcn from Wiley. 198 1. Pg. 138- 139

Phylogenetic reconstmctions are vital. for they not only shed light on both the genus and its rnember species. but they also yield hypotheses concerning evolution and history of diversification. Although cladistics as a tool for classification is gaining wide acceptance (McLennan, 1996) it has yet to be applied to the hexactinellids. However. numerous cladistic studies have been published on Demospongiae groups [eg.(Hajdu, 1994,1996): (Hiemstra, 1991 ); (Hooper, 1991, 1994); (Matdonado, 1994); (Sara, 1 994); (Soest. 1 990. 1 99 1a. 199 1 b. 1993)]. These studies are sources of information regarding characters and cliaracter States, as well as reliability or strength of characters. and are thus directly pertinent to the study of hexactinellids. Hajdu (1 996) examined al1 seventeen published demosponge data matrices and found that sixteen character classes predominate. The performance of each class of characters was calculated in terms of their consistency index in order to assess character reliability. However. due to the small sarnple size and the procedure involved in establishing character reliability, the results of this analysis are not conclusive. Generally, the numerous phytogenetic studies performed on demosponges follow closely the methodology outlined by Hooper ( 1991 ). Hooper performed a cladistic analysis on the family (seventeen taxa) Raspailiidae (Demospongiae) using PAUP Version 2.4 on eleven characters: structure of the choanosomal skeleton, differentiation of the axial and exa-axial skeletons. architecture of the axial skeleton, megascleres in the axial skeleton. architecture of the extra-axial skeleton. megascleres of the extra-axial skeleton, structure of the ectosomal skeleton, geometry of echinating megascleres. geometry of spines on acanthostyles. distribution of echinating megascleres. and presencehbsence, type and form of microscleres. Data used in this "Wagner" analysis were derived fkom an unordered multistate character set, and apomorphy was judged by outgroup comparisons.

Methods Specimens, or fragments ofspecimens. used in this study were predominantly from the extensive hexactinellid coliections of the Redpath Museum. Montreal, Canada. or. loaned directly fiom world museums. The majority of this material was collected by dredçe or trawl on historic nautical expeditions ('Challenger ', 'Siboga', 'Albatross '.

' BZrkc. '. 'Princess Alice '. etc.. .)- Spicule preparations followed Reiswig and Browman (1 987). Where appropriate. specimens were examined under a Wild dissecting microscope to determine spicule presence and location. Fragments of specimens were then subjected to concentrated nitric acid to dissolve al1 organic material. Undigested siliceous spicules and dictyonal frarneworks were then washed with distilled water and filtered under vacuum ont0 a 0.22 Fm Millipore GS filter. Filters were bnefly dried and mounted to glass slides using Canada balsam. Spicules were measured and counted using a 'canera lucida' (cornputer digitising tablet and microscope). Number of spicules analysed varied upon abundance. Hesactine and pentactine ray lengths were taken from the centre (axial cross) of the spicule to distal t i ps; measurements were reported as ray lengths. Scopule measurements included tine lengths, capitulum width, and overall scopule length; tine length was taken from the axial centre (or axial cross when visible) of the capitulum to the distal most tine tip; capitulum widths were taken closest to the centre (or axial cross); overall lengths were taken fiom the proximal tip of the shaft to the distal rnost tine tip. Microsclere measurements included pnmary ray length, secondary ray length and overall spicule radius; primary ray lengths were taken from the centre of the spicule to the converging point. or capitulum. of secondary rays; secondary ray length was taken from the above capitulum to the distal most tip of the secondary ray. Microsclere measurements were reported as diarnetres (radii multiplied by two). Uncinate lengths were taken fiom proximal to distal tip. Measurements of dictyonal frarnework included mesh width and length, beam width and channel diarnetre. Mesh widths and lengths were taken from the inside beam surfaces (i-e. not the centre of the beam); one channel diametre was taken per opening and was taken through the centre of the spherical or oval opening. Statistical analyses (means. standard deviation. modes, frequency distributions) were performed on Excel for Windows 95. Line-drawings were made directly fiom calibrated video grabs of spicules using Adobe Illustrator for Windows 95. Photographs of dictyonal frameworks were taken using the Zeiss Photomicroscope with phase contrat optics. Scales were detemined by using a micrometre. Whole specimen photographs were taken using a Wild dissection canera. Scales were determined using physical scale bars. Film negatives were scanned directly into Adobe Photoshop and placed within Adobe Illustrator. Scanner magnification was established using negatives of a photographed micrometre. Invalid famiiy names are indicated in the synonomy of Euretidae by a comma inserted after the name. Generic synonyms follow standard zoological ta~onomicformat. Cladistic analyses were performed on PAUP v. 3.1 and MacClade v. 3.0 for the Macintosh. Only unordered (Wagner Parsimony) binary character states were used. Information for data matrix construction was taken pnmarily fiom examination of type material, however. original descriptions and illustrations were also used when information was lacking in the type specimen or type fkagment. Constant, symplesiomorphic and autapomorphic characters were not included in the data matrix. Characten used in this analysis were detemined, and hence weighted. apriori through a preliminq "total evidence" (Kluge, 1989) analysis (including 192 characters and 17 taxa) not presented here. Abbreviations of rnuseums fiom which loans were procured are BMNH (British Museum of Natural History), MC2 (Museum of Comparative Zoology, Harvard). MNHN (Muséum National d'Histoire Naturelle, Paris). Musée de Monaco and the Leiden Museum. Sysfema tics

Class HexactineIIida Schmidt, 1870 Subclass Hexasterophora Schulze, 1 886 Order Hexactinosa Schrammen. 19 12 Suborder Scopularia Schulze, 1885 Family Euretidae ZitteI, 1877

MacAndrewiadae Gray, 1859: 440 (in part). Hesactinellidae Schmidt. 1870: 13 (in part); Carter, 1873: 349 (in part). Dactylocalycidae Gray (in part) 1872: 505: Schulze, 1885: 450 (in part); sensu Ijima, 1903: 25; Schulze, 1904: 179 (in part). Hyalothaumadae Gray, 1872: 45 1 (in part). Vitreohexactinellida, Carter, 1875: 134 (in part). Pleionakidae, Marshall, 1876: 123 (in part); sensu Zittel 1877. Monakidae, Marshall, 1876: 123 (in part); sensu Zittel 1877. Euretidae Zittel, 1877: 35; Schulze, 1885:450; Schulze. 1887: 265: Schulze 1899: 99: Schulze, 1 904: 1 72; Schrammen, 19 1 2: 1 90; Ij ima, 1 927: 1 64; de Laubenfels. 1 936: 186: Levi, 1964: 104: Reid ( 1963): 22 1 ; Van Soest. 1988: 1 1 : Barthel & Tendal. 1994: 58; Hooper & Weidenmayer: 1994: 5 1 5. Maeancirospongidae, Zittel. 1877: 38 (in part); Schulze, 1887: 265. Coscinoporidae Zittel 1 877: 36; Schulze, 1 887: 265; Schulze. 1 889: 1 05: Schulze. 1904: 1 72: Barthel & Tendall, 1994: 60 Myliusidae Schulze, 1885: 45 1. Chonelasmatidae Schrammen, 19 1 2: 190. Euritidae Carter, 1885: 393. Euryplegmatidae, de Laubenfels, 1 955: E78.

Diagnosis Growth form erect. ranging fiom simple fùmel-shaped tubes. usually with lateral oscular branching. to cup-shaped forms of intricately arranged tubes: attached to substrate by means of a basal plate; central atrium present in most forms; extemal or atrial surfaces smooth, pock-marked, with horizontal ridges or longitudinal ridges andor grooves; dictyonal fiamework of secondarily fûsed hexactin megascleres; fiamework three- dimensional and greater than one meshwork thick, peripheral areas of skeletal formation may be one meshwork thick; dictyonal cortices may be present; meshwork generaily rectangular, triangular and/or rotdate; ofien regular, rectangular-meshed, interna1 framework is enclosed by irregulru, triangular-meshed external frarnework; external and basal portions of framework ofen with smaller. more irregular meshwork; small oxyhexactins commonly fûsed to extemal or basal portions of fiamework; dictyonal strands and laminae present or wanting; smooth, microspined or tubercuiate dictyonal nodes tme or false; dermal and gastral spurs present but may be absent: beams generally Formed by arnalgamation of two rays of adjacent hexactines or synapticulation: beams smooth. microspined or spined; intradictyonal and extradictyonal framework channelized or unchannelized; channelization as epirhyses, aporhyses, diplorhyses, diahryses. or arnararhyses; cavaedial spaces rnay be present; dermalia and gastrdia commonly present but may be absent; dermalia and gastralia as pentactine (with or without vestigial knob of distal ray), hexactine and pimulated hexactine megascleres; sceptrules. if present. as scopuies only: parenchymal microscieres as oxy-. disco-. tylo- or onychohexasters: uncinates present or absent.

Taxonornic Remarks This farnily is the largest and most diverse of Hexactinosa Schrammen. It is distinguished from the five other hexactinosan farnilies (Farreidae Schulze. Tretodictyidae Schuize. Aulocalycidae Ijima, CraticuIariidae Rauff and Aphrocallistidae Gray) primarily by type of skeletal framework (euretoid), channelization (al1 types of channelization excepting schizorhyses) and presence of scopule sceptrules. The farnily Euretidae was created by the paleontologist. Zittel (1 877). to incorporate those extinct and extant genera having cup-shaped hm,three-dimensional skeletal 'latticework', unswollen nodes, and dermal 'ostia'. Zittel(1877) also divided the Euretidae into two groups: (a) those forms with a canal system and, (b), those forms without, or scarcely developed, canal systems, and, although he correctly used Eurefe Semper as type genus, he wrongly cited Marshall as author of Eztrere. Within the same classification, Zittel (1 877) erected the farnily Mæandrospongidae. for those forrns (Peripl~rugelluMarshall. Mylitcsiu Gray) constructed of meandering and intenvoven tubes. According to forma1 taxonomie procedure, the name Maeandrospongidae was not based upon an existing genus. and therefore can not be accepted as valid. Although he did recognise the importance of Zittel's (1 877) work. Schmidt (1 880) did not offer any significant contributions to existing familial classification and felt that a scheme based primarily on skeletal remains is inadequate. In an attempt to reduce the importance of skeletal fiameworks in classification, Carter (1 875, 1885, 1886) created a classification centred upon methods of spicule fùsion and placed the family Euretidae (misspelled as Euritidae) in his Vitreohexactinellida. Schulze (1 885). however, ignored Carter's (1875) classification scheme and divided Zittel's (1 887) order Dictyonina into two groups based on the presence or absence of uncinate spicules: (a) the Uncinataria, into which Schulze (1 885) placed the farnilies Chonelasmatidae (for Chonelasma Schulze), Euretidae (for Eurete Semper, Periphragellu Marshall and Lefroyella Thomson) and four other families. and (b). the Inermia, into which he placed four families including Myliusidae (for Mylizrsici Gray). Dactylocalycidae (for Ductyloccrlyx Stutchbury. Scleroplegmu Schmidt and ~MurgurireiZa Schmidt) and Euryplegmatidae (for Eurypkgma Schulze and Joanella Schmidt). Schulze (1 885) erected the tribes Scopularia and Clavularia for those Dictyonina (Zittel. 1877) with scopule and cIavule sceptrules respectively and diagnosed the family Euretidae simply as those forrns of branched or anastomosing tubes or of a goblet with lateral outgrowths. Schulze then (1887) modified his (1 885) classification by changing: (a) his Chonelasmatidae to Zittel's (1 887) Coscinoporidae. and (b). his Myliusidae to Zittel's (1 887) Maeandrospongidae (for Dacryloculys Stutchbury, Scleroplegmu Schmidt. Arilocystis (= hreoatrlocystis Volgodin) ,and Myliusia Gray). In this later (1 887) scheme. Schulze considered himself as the authonty for the family Euretidae and did not include Murgaritella Schmidt or Joanella Schmidt. Schulze (1 899) later: (1) placed the new genus Bathyxiphus Schulze into Zittel's (1 877) Coscinoporidae; (2), followed Ijima's (1 903) advice and accepted Dactylocalycidae in place of Maeandrospongidae for ~CfylitrsictGray, Murguritellu Schmidt and three other genera. and (3). placed (1 904) the genus Furrea Bowerbank into the Euretidae. The paleontologist Rauff (1893) synonomyzed Zittel's (1877) concept (based on Etrr-erc simpiicissimzrm Semper, not in possession of spicules) of Euretidae with his Craticularidae and, contary to current zoological procedure. considered Schulze's concept of Euretidae (based upon the extant and Eureie farreopsis Carter, which was in possession of spicules) as valid (Euretidae Schulze). Schrammen (1 912) closely followed Schulze's (1 885. 1887. 1889. 1904) classification and included Farrcu Bowerbank, Eurere Semper, Periphragella Marshall and LL.foyeila within the Euretidae. Schrammen (1 9 12) rejected Schulze's (1 887) concept of Zittel's (1 877) Coscinoporidae and cited himself as author of the family C honelasmatidae, which Schulze ( 1 885) had previously ( 1 885) erected and Iater ( 1 887) rejected. Schrarnrnen (1 9 12) also followed Ijima's (1 903) classification and included Dactylocalycidae as a family within Schulze's (1885) subtribe Inennia. In short, Schulze ( 1885. 1887, 1899. 1904) had little influence upon the wider treatment of Euretidae and focused primarily on providing descriptions and illustrations of newly collected euretid material. Unfortunately. Schulze (1 885, 1887, 1 899- 1904) made no mention of Iphiieon Bowerbank and was noticeably silent with regard to channelization and its role in classification. Although initiaily (Ijima, 1903) supporting Schulze's (1 887) classification. Ijima ( 1927). reporting on the material collected from the 'Siboga' expedition. completely restructured the Euretidae and is largely responsible for its current concept: "Scopularia with wall usually not canalized. cxceptionally developing either arnararhyses or epirhytic depressions but unaccompagnied by aporhysis. Skeletal framework may be essentially similarly structured as in the Farreidae. but frequently deviates from this in showing meshes of a triangular or irregular shape and certain nodes which give off more than six internodal bearns. these being singly ruming hexactin rays and in part amaigamations of two rays. belonging to different adjacent hexactins. Scopule generally present. but may sonietimes be wanting." (Ijima. 1927: 163-4). With this diagnosis Ijima (1 927) expanded the number of genera within the Euretidae, from the original (Schulze, 1887) three (Eztrefe Semper. Peripphragella Marshall and Lefroyeila Thomson) to a total of fifteen, by: (a) including two genera (Pleztrochoriztm Schrammen, 19 12; Piychodesia Schrammen, 191 2) known primaril y from the fossil record: (b), importing three genera, (Dactylocal' Stutchbury. Iphileon Bowerbank and Myliusia Gray) previously described before the inception of the Euretidae and never previously considered as euretids: (c). transferring three "post-Zittel" (1 877) non-euretid genera (Margurifella Schmidt, Chonclusma Schulze and Bufhyiphzts Schulze) into the Euretidae; (d), including Dendy 's Heterorefe from the famil y Dactylocalycidae and, (e) crecting three new genera (Conorete. Gyrnnorete and Pararete) based upon three significantly different members of Ezrreie Semper. Ijima (1 927) also removed Farrea Bowerbank from the Euretidae and reinstated it as type genus for the family Farreidae. Mernbership within the Euretidae was then altered by de Laubenfels (1936). de LaubenfeIs (1936, 1955) brought the total number of euretid genera to sixteen by including Okada's (1 925) Calyptorete, Topsent's Pityrete and Scleropiegma Schmidt , and rejecting, with no explanation, Gymnorete Ijima, Heterorete Dendy and Endoretc Topsent. Controversially placing increased importance on frameworks and channeiization and Iess on spicule composition. Reid (1964) reorganised the farnily Euretidae to include nine genera and five subgenera. Reid (1964): (1) designated several of Ijima's genera (Prrrcrrele, Gymorete, Conorete), Heterorete Dendy and Endorerc Topsent as subgenera of Ezrrere Semper; (2). reinstated Schmidt's Syringidiztm; (3). replaced Ijima's concept of Pt@miesia Schratnmen with a new genus. Tretachone Reid; and (4)rejected Pityrete Topsent, Dactyiocalyx Stutchbury, MargariteZla Schmidt and Scieropkgma Sclimidt. Levi ( 1 964) rejected Ijima's Purarete and Reid ( 1969) added the last new genus. ~~crr~~cocoeloideu,to the Euretidae. Al though not having ci ted references. Hartman (1 982) noted fifieen euretid genera , some species of which may possess sarule sceptniles. Subsequent authors (1982 to present) have made relatively few changes to the farnily: (a) Van Soest (1 988) returned DacfyZocaZyx Stutchbury to the Euretidae; (b) Barthel & Tendall (1 994) reinstated Zittel's (1 877) Coscinoporidae and therefore removed C'lmnel(isrna Schulze and Bathyxiphus Schulze from the Euretidae and. (c) in their review of Australian hexactinell ids, Hooper & Wiedenmayer ( 1 994) mentioned on1y fourteen valid euretid genera.

Review As stated in the 'Introduction' there are several problems associated with the taxonomy of the Euretidae. al1 of which contribute to instability in its classification. Thrse problems need to be addressed in order to recognise valid genera and species and to produce a phylogenetic classification. (a) The primary literature is antiquated and descriptions are often too brief for modem analytical procedures. Of the sixteen genera, considered as valid in this publication. over half (56%) were described before 1910 and the bulk (93%) were described before 1930. The present work deals primdy (when possible) with type material thereby reducing reliance on the literature. However, the literature was used when type material proved inadequate or incomplete. (b) The implementation of formal taxonomie procedure was just starting when many euretid genera were described, causing a lack of consistency and scientifk method among descriptions. For example. holotypes often were not designated and descriptions may have been based upon more than one specimen, and in several cases. more than one species. To overcome this problem. in-depth tavonomic reviews of each genus were conducted and where appropnate. proper type specimens (holotypes. lectotypes etc.. .) were designated. (c) Crude methods of collection resulted in damaged or incomplete type specimens. Also, several specimens may have been collected in the same sample, increasing the risk of spicule contamination. For the purposes of this review. supporting evidence from incomplete or inadequate type material was sought fiom available collections. (d) Genera and species within the Euretidae were described in different languages (primarily German, French and English). This necessitates translation (which ofien does not convey the true thoughts of the original author) and also makes obtaining original pubIications difficult. Fortunately, this author had available to him the world's largest assembly of hexactinellid Iiterature compiled and managed by Dr. H. M. Reiswig. (e) I'here are no single 'modern' (post 1900) generic revisions of al1 known euretid genera with no one system of accepted morphological terminology. Lack of consistency in morphological terminology greatly harnpers any attempt at systematic analyses. Although the author of this work does not have a complete understanding of available hexactinellid morphological terminology, effort was made to remain consistent throughout al1 generic revisions. Explanation of features and concepts may be found in the literature review.

SUBFAMILY EURETINAE zr-rrm

Cup, or funnel-shaped, ewetids with lateral branching; dermalia and gastralia as pentactins or pinnulated hexactins; scopules always present and are generally tyloscopules; oxy- and strongyloscopules common; uncinates present; framework generally with smooth dictyonal beams; without epirhyses and aporhyses: microscleres generally as discohexasters and/or oxyhexasters.

EURETE SEMPER, 1868

Eurefe Semper, 1868: 372; Marshall, 1875: 18 1; Carter, 1877: 124; Zittel. 1877: 344: Schmidt, 1880: 34; Carter, 1885: 393; Schulze, 1887: 289; Schulze. 1899: 72; Topsent. 190 1 a: 463; Topsent, 190 1b: 38; Topsent, 1904: 45; Schulze. 1904: 143; WiIson. 1904: G3: Kirkpatrick, 1908: 2 1; Lendenfeld. 1915: 126; Ij ima. 1927: 165: Topsent. 1928a: 3: Topsent. l928b: 92; Okada, 1932: 43; Reid, l958b: 17; Reid. 1963: 234; Koltun. 1967: 49: Levi & Levi. 1982: 298; Tabachnik. 1988: 173: Reiswig, 1990: 735.

Not Eureta Vosmaer, 1 887: 242.

TYPE SPECIES Etrr.ete simplicissirnu Semper. 1868 (holotype by original designation, Semper. 1868)

BASIS OF DIAGNOSIS Diaçnosis based upon Schulze's (1 887) original description of Elu-etc hocr.c.rhcankii Schulze. 1887 (reasons given in 'Remarks').

DIAGNOSIS Body fom tubular, branching and anastomosing; accessory oscula present. Three- dimensional unchannelized primary dictyonal framework formed of fused hexactins; no peripheral secondary dictyonal framework. Meshes commonly triangular or rectangular. Beams slightly microspined; framework nodes slightly swollen or unswollen; niicrospined spurs as free rays of primary peripheral dictyonal ia. Derrnal ia and gastralia as pentactins; loose spicules as scopules and uncinates; oxyhexasters are the only microscleres.

REMARKS The taxonomie history of the genus Eurete is long and confusing. Semper (1 868a.b) initially erected this genus for a single, macerated specimen collected from Cebu. by Dr. 2. Legaspi. This specimen was named Eurete simplicissimum Semper for its complete lack of loose spicules and the genus was described primarily upon body form and skeletal architecture: "a rather compact net of fine siliceous tubes, which sometimes blend irregularly, but sometimes cross each other very regularly, thus forming a network includinç rectangular meshes." Semper (1868) was unable to determine if the complete lack of spicules was a valid feature of the genus or the result of maceration or bleaching. For his 'Review of the Hexactinellida' Marshall ( 1875) obtained Semper's type specimen and. afier a thorough search. contirmed the complete lack of loose spicules. However. unlike Semper, Marshall (1875) postulated that the lack of spicules was not the result of physical maceration but of a fault in the spicule formation process. Marshall (1 875) also included an illustration of the conl-like body form of E. simplicissimtcm and clearly figured the prominent axial canals of the dictyonal framework. In his review of known sponçes. Claus (1 876) committed a typographical error and mistakenly cited a previously unknown combination. E. schtilzei Semper. and made no mention of E. simplicissimzrrn. Carter (1877) regarded the lack of spicules in E. simplicissimum as a result of macention. as evidenced by the prominent framework axial canals, and, using the dictyonai framework as a basis. narned a new sponge, with loose spicules, from the Philippines as Ezrrele farreopsis. In doing so, Carter indirectly redefined the genus Eurere as having not only a dictyonal framework greater than one layer thick, but also dermal and gastral pentactins. scopules, uncinates, discohexasters and oxyhexactin microscleres. Based upon the conspicuous absence of spicules within E. sirnplicissin~rrrn.Zittel (1877) provisionally synonomyzed Bowerbank's Farreajisircla~awith E. simplicissimtrrn and mistzkenly cited Marshall. and not Semper, as the authority of Eurete. Like Zittel. Schmidt (1 880) noted strong similarities between Semper's Ezrrete and Bowerbank's Fm-rea and questioned the validity of the genus Eurete. In his report on the hexactinellids from the 'Challenger' expedition. Schulze (1 887) thoroughly reviewed the genus Eicrete Semper and described four new species found off Little Kei Island and one fiom the Philippines (E. carreri. E. marshalli. E. schrnidtii, E. semperi and E. bowerbankii). Although Schulze ( 1 887) included illustrations with his descriptions, he failed to cite important measurements such as spicule sizes and heworkdimensions. Although Schulze (1 887) obtained Semper's type specimen and discovered 'several' oxyhexasten. he considered E. farreopsis as the generic type species. Schulze (1 887) also included E. simplicissirnurn Semper and the erroneous combination. E. schttfzei Claus. as valid members within the genus Eurere and mentioned several unidentified specimens of Eurere. Using matenal collected by the 'Albatross' expedition from the Galapagos and Guadeloupe Islands. Schulze (1 899) described a new species, Eureie erectirm. and mentioned several unidentified specimens of Eirrefe. Schulze (1 899) also changed the authorship of Eirrc're back to Semper from (Schulze. 1887) Carter. however. he cited no reason for this move. Two years later ( 1 90 1), Topsent described E. alicei (190 1a) from the Azores and E. gerlrrchei ( 1 90 l b) from the Arctic and later (1904) reported an unidentifiable specimen of Eurete fiom the Azores. In his last publication on Eurete, Schulze (1904) correctly cited Semper as the original author of Eurefe, described an unknown specimen of Eurete collected by the 'Valdivia' expedition off St. Paul, and included discohexasters and oxyhexasters as euretid microscleres. Using 'Albatross' material fiom the Pacific. Wilson (1904) defined three subspecies of E. erecium Schulze and Kirkpatrick ( 1908) described a distinct specimen. E. unnandalci from the Indian Ocean. Lendenfetd ( 19 15) later described the new E. spinostrm from northern Peru. redefined Wilson's (1904) subspecies as forms of 15.erecrttrn. and reported several unknown specimens of Errrefe. Ijima (1 927) completely revised the genus Eurerc. and made numerous important tasonomic decisions, to include several new species collected by the 'Siboga' expedition from the Sagami Sea (E. schrnidti kampeni. E. schmidii treubi. E. frcelandi. and E rr~~~hycloctrs).Ij ima ( 1 927) chose the next oldest member of the genus Eirreie. Schulze 's E. hoiverbankii, as the genenc type species and redefined Eirrete as "Tubular. branching. and anastamosing. Dermalia and gastralia pentactins. With oxyhexasters onIy." (Ijima. 1927: 165). He (1 927) then split most of the remaining species into three new genera: (1) Pararefe received E. furrcopsis Carter, E. carreri Schuize, E. gcrlachei Topsent and E. semperi Schulze; (2) Conorete was erected for E. erectzrm and its various forms; and (3) Gyrnnorek for Topsent's E. alicei. Ijima (1927) transferred the remaining E. crrtnandalei Kirkpatrick to the genus Plezrrochorizrm Schrammen. Okada later (1 932) described three new species. E. nipponica. E. irregirlaris and E. succul~ormis.collected from. or near. Sagami Sea by the 'Albatross' expedition. On describing the 'Upper Cretaceous Hexactinellida' Reid (1958b) rsjected E. borvwbunkii Schulze as the generic type species and redefined Eurete upon Semper's original 'spicule-less' E. simplicissimum. Reid (1958b) remarked that zoologists offen based the genus Eurete upon loose spicules. and their variations among specimens. and concluded that such groupings may be convenient only at the subgeneric level. Reid later ( 1963~)recognised the absence of skeletal canalisation (= channelization) as an important distinguishing feature of Eurefe Semper and. using E. simplicissimum Semper as a basis. formally relegated Eurcte (sensu Ijima). Pararete Ijima. Endorefe Topsent. Gymnorete Ijima. Fieferorete Dendy, and Conorere Ijima as subgenera of Eurefe Semper. In addition to collecting a specimen of E. irrcguh-is Okada from the Bering Sea. Koltun (1 967) redefined the genus Ewete Semper to include oxyhexasters and onychasters (= onychohexasters) and considered E. bowerhunkii Schulze as the generic type species. Levi and Levi (1982) did not accept Ijima's (1927) generic differentiation based upon microscleres and synonomyzed Pururefe Ijima to Eurere Semper. The last author to describe a new species of Eurete was Tabachnick (1 988). who. using the research vesse1 'Akademik Mstislav Kyeldish', coIIected and described E. lameIlina from the western Pacific. Finally. Reiswig (1990) clarified several of Ijima's taxonomic moves in his correction of ijima's (1927) 'Final List'. Although a search of world museum collections for the type specimens of E. simplicissimwn Semper and E. bowerbankii Schulze has proven unsuccessfùl, a recent review of the literature has aided considerably in understanding the genus Ewm Semper. Carter's (1877) initial inclusion of E. furreopsis Carter into Eurefe. on the basis of skeletal simiiarity, may be regarded as the primary source of taxonomic confusion within the genus Eurcte. Clearly, Carter (1877) was in error to associate a specimen void of tasonomically important spicules (E. simplicissimum) with one that does possess spicules (E. fart-eopsis). Claus's ( 1 876) citation of E. schulzei Semper for E. .simplici~wintzim Semper is clearly an editoria1 error and the name E. schulzci may be considered a nomen nzrdurn. The specimen of Farreafitulnra Bowerbank. which Zittel (1 877) provisionally synonomyzed with E. simpIicissimum Semper, is a long dead and unrecognisable fragment (Schulze 1887: 282) and therefore can not be considered synonymous to E. simplicissimurn Semper. As pointed out by Levi & Levi (1 982), Ijima's redefinition of Elrrete. based solely upon the presence of oxyhexasters. and the formation of Pararcte is controversial. However, Ijima's actions are consistent with the discovery (Schulze. 1 887) of oxyhexasters wi thin the generic type specimen, E. sirnplicissimtrrn. Althouçh Reid ( 195 8b) correctly recognised Semper's ( 1 868) taxonomie priority, his subgeneric chssification (1963a) for an extant group of sponges, based upon a single, macerated and 'spicule-free' specimen, is not acceptable. Schulze's E. borver~ankiiis therefore considered the generic type species of the genus Ezlrete Semper (sensu Ijima. 1927).

DISTRIBUTION Generic twe Ezrrête borverbankii Schulze. 1887 collected from Sagami Sea, Japan.

Generic species (Type specimens) E. simplicissimurn Semper, 1868 collected near Cebu, Philippines. E. schmidtii Schulze, 1877 collected fiom the Philippines. E. marshalli Schulze, 1877 collected from Little Kei Islands, Banda Sea. E. spinoszrrn Lendenfeld, 1915 collected fiom N.Peru,South West of Aguja E. schrnidfi kumpeni Ijima, 1927 collected off Kei Islands. E. schrnidri treubi Ijima, 1927 collected off Kei Islands. E. frmlundi Ijirna, 1927 collected from the Sulu Archipelago. E- frcrchydocus Ijima. 1927 collected from Kei Island, Banda Sea. E. rtipponicu Okada 1933 collected south West of the Goto Islands. E. il-rêgzrfuris Okada, 1932 collected near Sagkalin Island. E. sacczrliforrnis Okada, 1932 collected south west of the Goto Islands E. lumellina Tabachnick, 1988 coltected north of New Guinea. * see Plate 1, Fig. H for geographicai distribution map.

Eurete bowerbankii, Schulze (Plate 1. Figs. A-H)

Eurete bowerbankii Schulze, 1887: 297, PI. LXXIX. Figs. 9-1 3; Schulze, 1899: 1 17: Topsent. 190 1 : 466; Schulze, 1901: 143; Ijima, 1927: 165; Reid, 1964: civ; Reiswig, 1990: 735.

TYPE MATERIAL Eimre bowerbankii Schülze, 1887; holotype collected from Sagami Bay by Dr. Doderlein; preserved in alcohol; present location unknown.

DESCRIPTION Descriptions of the body form, skeletal fiamework and spicules are based upon Schulze's (1 877: 297, Pl. LXXIX, Figs. 9-1 3) original description and illustrations. Al1 rneasurements are approximations taken from Schulze's (1 887) illustrations.

Body Form (Plate 1, Fig. A): The illustrated holotype is incomptete. erect. "tree-like" and tubular; total sponge height 30 mm; width 3-1 1 mm. Body consists of a pnmq tube. which terminates distally in a funnel-shaped oscula. Laterd tubes, presiimed as accessory oscula (3 mm in diametre), with circular or nearly circular apertures. appear to originate from the primary tube and are directed distally. Method of attachnîent to substrate unknown. Schulze (1 887) functionally determined the intemal surface as gastral and the external surface as dermal.

Skeletal Framework (Plate 1. Fig. G): The dictyonal fiamework is a regular and irregular three-dimensional network of fused (ankylosis) hexactin spicules; meshes rectangular or triangular (0.3 mm width; 0.6 mm length) elongate or 'stump-like' microspined spurs as free rays of external primary dictyonalia; dictyonal strands presurnably oriented parallel to growth axis; nodes slightly swollen; beams (50 - 80 pm thick) finely microspined throughout.

Loose spicules (Plate 1. Fig. B-F): Dermalia and gastralia are smooth pentactins (Plate 1, Fig. B): sixth ray absent or vestigial; tangential rays (Iength unknown) slightly bowed. smooth and terminally blunt; proximal ray (length unknown) straight and proximal1y rounded. Scopules of two types (Plate 1. Fig. D. E). Shorter (total length unknown)(Plate 1, Fig. E) tyloscopules are cornmon: four microspined and barely divergent tines (70 prn length) end distally with a swelling on which are numerous, prominent, proximally oriented, spines; unswollen capitulum gives rise to a microspined, straight and terrninally rounded shaft. The second tyloscopule (total length unknown; Plate 1, Fig. D) is nearly identical to the first tyloscopule except that the four tines are significantly divergent and the shah is noticeably longer. The oxyhexaster (90 pm diametre ?) (Plate 1. Fig. C) is the only microsclere present. Long (30 pm ?) primary rays (twice as long as the secondary rays) end in four sharply pointed, and greatly divergent, secondary rays. Barbed uncinates (Plate 1. Fig. F) of varying thickness and length are common.

DISTRIBUTION E. simplicissimum Semper, 1 868 collected fiom Cebu, Philippines. E. bowerbunkii Schulze, 1877 collected from Sagarni Bay. E. nzcirshalli Schulze, 1877 collection location: 5" 8' S, 132" 2' E., depth 236 m. E. schmidii Schulze, 1877 collection location: 35" 1 ' N1138" 8' E.. depth 240 m. E. spinosztm Lendenfeld, 191 5 collection location: 6" 9' S, 83" 6' E., depth 4062 m. E. schmidti kampeni Ijima, 1927 collection location: 5" 5' S. 132" E.. depth 90 m. E. schmidi trwbi Ij ima. 1927 collection location: 10" 7' S. 123" 7- E.. depth 520 m. E. freekundi Ijima. 1927 collection location: 6" 1 ' N. 12 1" 3' E., depth 275 m. E. fr.crchydocus Ijima, 1927 collection location: 5" 5' S, 132" E., depth 204 m. E. nipponim Okada, 1932 collection location: 32" 5' N, 128" 5' E., depth 194 m. E. irr~'gzrlarisOkada. 1932 collection location: 46" 5' N, 145" 8' E., depth 3293 m E. sacczrliforrnis Okada, 1932 collection location: 32" 5' N, 128" 6' E., depth 247 m. E. lumellina Tabachnick. 1988; collection location (sta. 1074) 2" 12' N. 149" 3' E.. depth 1570m. Etrrete sp. Schulze. 1887; collection location (Chal. Stn. 192): 5" 8' S, 132" 2' E., depth 236 m. Errrtilc sp. Schulze. 1899; four specimens collected off the West coast of Mexico (Albatross stas. 2808,2980; 2986; 2992); 33" N - O" S; 119" - 89" W.. depth 84-1251 m. Plate 1 (Euretc borverbankii Schulze): Figure A: the holotype (modified from Schulze, 1877: PI. LXXIX. Fig. 9); B, diagrammatic representation of derrnal and gastral pentactin; C. oxyhexaster; D. larger tyloscopule; E. smaller tyloscopule; F. diagrammatic representation of uncinate; G, diagrammatic cross-section of dictyonal framework; H, geographical distribution (solid dot denotes practical type locality, open dot denotes locality generic members, stars denote unconfirmed or unidentified specimens).

CALYPTORETE OKADA. 1925

Calyptorete Okada. 1925: 285; de Laubenfels. 1936: 187; Reid. 1964:~~~~.

TYPE SPECIES Cu~~ptomteijimai Okada, 1925, (lectotype designated by Wheeler, this publication)

BASIS FOR DIAGNOSIS Diagnosis based solely upon Okada's (1 925) original description and reinterpretation of his description.

DIAGNOSrS Body form erect, cup-shaped; composed of branching and radiating tubes; supported by a tube-like stalk. Apertures of tubes ofien spanned by a peripheral membrane which also extends across cavaedial interspaces between tubes; this membrane is supported by a network of paratangentially orientated pentactins. which may undergo rigid union. Dictyonal framework is regular and irregular; meshes rectangular or triangular: beams

*~enerally microspined; regular dictyonal nodes not swollen, spurs present. iMegascleres as pentactins. scopules and uncinates microscleres as discohexactins and discohexasters.

REMARKS Okada (1 925) erected this poorly known genus for the distinct body form of five (A-E) specimens coilected fiom several regions near Sagami Sea between the years 1894 and 19 13. Okada (1 925) briefly described al1 five specimens of Calyprorete ijimui as having a stalked cup-shaped form "made up of nwnerous tubes almost definitely arranged" (Okada, 1935: 285) and an unusually wide variety of pentactins, scopules, uncinates. discohexasters and discohexactins- Okada (1 925: Figs. 1.2) based his description of body fom on the largest and best-preserved specirnen (specimen A, no. 570 at the University of Tokyo Museum: designated here as lectotype), which he illustrated; he used specimcn C as the primary source for spiculation. Reid (1 964) did not accept Okada's ( 1925) definition of the genus Cafyptoreie and offered another. which concentratcd primarily on the complex channelization and body form of Culyprorefc. Reid (1964: cxxx) also rejected Okada's (1 925) daims of pentactin hypodermalia and hypogastralia, and concluded that true dermalia and gastralia were lacking in Calyptorete. Recent revision of Okada's (1925) original description has revealed further ambiguity within the genus Calyptorete . By not designating a single specimen as holotype upon which to base the descnption of Calyporere yimai. Okada (1925) greatly confused rnatters. Okada (1 925) also lacked consistency between his description. figure Ieçend and illustration, ofien referring to a spicule by one name in the descnption and later refemng to it in the figure iegend by another. It is possible that the unusual variation within spicule type descnbed by Okada (1 925) is due to the presence of several (5?) different species within the type series. In the opinion of the author. Okada's (1 925) description must therefore be viewed with distrust. As the identity and collection location of the material Reid (1964) used for his analyses is unknown, his description of C. i/'imai Okada must also be viewed with uncertainty. Until Okada's (1925) type series is reexamined, and new material is collected, the genus Culyptoretê must be considered incompletely understood.

DISTRIBUTION Culyprorere #mai Okada 1925; lectotype (Specimen A, no. 570) collected outside of Iwadogake. Sagarni Sea. The other four specimens of the type series were coIlected either inside or just outside Sagami Sea (Plate 2. Fig. Q).

Calyptorete ijimai Okada, 1925 (Plate 2, Figs. A-Q)

Calyptorete rjimai Okada, 1925: 285. 1 Pl.; Reid. 1964:cxxx.

TYPE MATERIAL Giyprorete i/'imai Okada. 1925, type series thought to be stored at the University of Tokyo Museum; collected in or near Sagami Sea. exact location and depth of collection unknown.

DESCWPTION The following description is an interpretation of the information presented and itlustrated by Okada (1925). Shape and skeletal framework description is based upon the lectotype (here designated as specimen A) (Plate II. Fig. A); the spicule descriptions are based upon Specimen C of Okada's type series.

Body shape and size (Plate II. Fig. A, E): Body caliculate, or cup shaped; 13 - 14 cm in height and with a stalk 1-4 cm thick. The anterior cup portion of the body (6-8 cm in height and 13.3 cm in diametre) is a tightly packed radiating cluster of branching and anastomosing tubes. Tubes originate from the proximaI stalk and open laterally to the exterior (PlateII, E) or distally to the paragastral surface (Plate 2. Fig. A). Tubes open freely or are spanned by transverse membranes. which extend across. and enclose. the inter-tube spaces in cavaedial fashion. Unable to bctionally determine gastral or dermal surfaces. Basal plate present.

Skeletal Framework (Plate II. Fig. N. O, P): Dictyonal framework components distinguished on the basis of location. Intemal layered (2- 3 layers) primary dictyonal frarnework with regular meshes 1 mm in width; prominent dictyonal strands oriented parallel to growth axis joined together by heavily microspined transverse beams (3.5 - G pm thick); regular nodes not swollen. Framework of the exterior Iateral surfaces regularly quadrate meshed ( 1 mm width) (Plate II. Fig. O) or irregularly meshed. the beams of which are thin (4.5 - 6 pm) and sparsely or thickly microspined. Slender (0.7 mm long; 0.04 mm thick) "finger-like" and heavily microspined spurs curve strongly from unswollen nodes of the primq framework. The framework of the paragastral sudace with smaller (0.5 - 0.7 mm width) and more irregular meshes than that of the lateral surfaces and is formed by a network of igidly comected, paratangentially oriented. pentactins (Plate II? Fig. P, N). The smooth. or weakly microspined. beams (1 0 - 15 pm thick) of these meshes are comected to unswollen nodes from which small(15 - 35 pm thick) weakly microspined, curved, spurs with rounded tips emanate. Vertically oriented dictyonal strands of this region are 0.5 - 0.75 mm apart and course towards the lateral margins. Bearns (O. 15 - 0.2 mm thick) of the tube frarnework are strongly and densely spined; meshes (1 - 1.2 mm width) quadrate or irregular; heavily microspined spurs (0.21 - 0.48 mm length) emanate from unswollen regular nodes. Loose spicules (Plate II. Figs. B-D,F-K, L, M): Dermalia as pentactin megascleres of three size classes. Large microspined pentactins (Plate II, Fig. B) with equal length (0.90- 0.1 15 mm), straight, distally pointed tangential rays; straight proximal ray sarne Iength as tangential rays; distal ray as a vestigial knob. Pentactins (not figured) of medium size are common and rnay fuse into a reticular lanice. Tangentid rays straight, distally pointed and of equal length (0.45 - 0.80 mm); the proximal ray is reportedly shorter than that of the larger pentactins and the spicule is lightly microspined except at the node and ray bases: distal ray reduced to a knob-like swelling. Information concerning the rare. smaller lightly microspined pentactins (not figured) is sparse: tangential rays (0.30 - 0.45 mm in length) terminate distally in a point. distal ray is reduced but prominent. Of scopules there are four types. The first. (scopule 1) is a strongyloscopule (Plate II. Fips. M). with four straight undivergent tines (0.08 mm length). Tines are completely microspined except distally resulting in a "bald" appearance; the unswollen capitulum attaches to a microspined shaft that rnay or rnay not be straight; the terminus of the shafi is also unknown. Overall Iength of this scopule is reported as 0.35-0.37 mm. The second scopule (scopule 2) (Plate II. Fig. L) is a tyloscopule (0.20 - 0.25 mm in length) commonly found in the "choanosome"; with six, or eight (?), slightly divergent tines 33 - 35 pm in length (Okada erroneously reports this measurement as 0.33-0.35 mm); tines are completely microspined except proximally, where they connect to a swollen capitulum. and distally, resulting in a disk-like head: the smooth shah rnay be proximally microspined. Distinct tyloscopules (scopule 3) (Plate II. Figs. F) with 12-14 slightly divergent tines (22 - 27 pm length) are also found in the choanosome. Tines originate from a capitulum having four "cruciately disposed" knobs. each showing avial canals; tines are comptetely microspined except distally. resulting in a bald appearance. The shafi rnay or rnay not be straight, is smooth or microspined and the terminus is unknown. Information regarding the fourth scopule (scopule 4) (Plate II. Fig. G) is sparse (no measurements) and has been compiled from Okada's illustration. This oxyscopule appears to have six, nearly parallei, tines emanating from a strongly swollen capitulum which appears to also have the four "cruciately" disposed knobs previously observed in scopule 3. Okada erroneously reports this scopule as a younger stage of scopule 3 (this is the only scopule that has six well developed tines and one would expect a younger spicule to have the sarne number of fine, or weakly developed. tines as other scopules present in the sponge). Uncinates are of four types, Long uncinates (uncinate 1 ; 2.5 mm length; 15 - 20 pm thick) similar to those figured in Plate II. Fig. J have prominent barbs which are strongly angled from the shaft. These are abundant in peripheral lateral regions either perpendicular to the lateral surface or scattered in the choanosome. Smaller. "spindle-

shaped" uncinates (uncinate 2; 0.2 - 0.35 mm; 12 Pm thick) (Plate II. Fig. H) are abundant in the paragastral region and have grooves or brackets with no barbs. A nodular swelling approximately 5 pm in width is observed 0.13 - 0.15 mm from the thicker end of this uncinate. Located together with uncinate 2 and perpendicular to the paragastral surface is uncinate 3 (Plate II. Fig. 1). Uncinate 3 is 0.3 mm in length and 0.01 5 mm thick with minute dense barbs that are closely pressed against the shaft. The diagnostic vdue of the fourth type of uncinate (uncinate 4, Plate II, Fig. K) is dubious as it was only observed in one specimen of the type series. This uncinate is irregularly curved and slender (0.5 - 0.7 mm long; 0.002 mm thick) and bears poorly developed barbs and well developed brackets. Uncinate 4 is rarely found in the "subdermal" or peripheral Iateral region. MicroscIeres are of two types. Common parenchymal discohexasters (Plate II, Fig. D) with principal rays bearing two to three slender. slightly bent and smooth secondary rays. range from 0.06 - 0.08 mm in diarnetre and may be larger in the "dermal" regions. Secondary rays terminate distally in a convex disc with four to sis "minute recurved teeth". Rare discohexactin spicules (Plate II, Fig. C) with smooth and slender axial rays (0.1 mm diametre) terminate distally with toothed discs which resemble those of tlie discohexasters. Discohexactins are located primarily in peripheral lateral regions (Le. Okada's "dermal").

DISTRIBUTION Culyprorefe ijinai Okada, 1925; lectotype (specimen A, no. 570*) collected outside of 1wadogake, Sagam i Sea. C. ijimni Okada, 1 925 ; paratype B (no. 224*) col lected near Hom ba South of 1 wadogake. Sagam i Sea. CI ijintai Okada. 1925; paratype C (no. 57 1 *) collected near Homba. South of Iwadogake. Sagami Sea. C. ijituai Okada, 1925; paratype D (no. 572*) collected in the Sagami Sea. C: ijirnai Okada, 1925; paratype E (no. 573 *) collected outside of Iwadogake. * thought to be catalogue numbers at the University of Tokyo Museum. Plate 2 (Cuiypforefe vimai Okada): Figure A. top view of Specimen A (rnodified fiom Okada. 1925, PI. 1); B, pentactin; C,discohexactin; D, discohexaster; E, lateral view of Specimen A: FI tyloscopule (scopule 3); G, oxyscopules (scopule 4); Htuncinate (uncinate 2); 1, uncinate (uncinate 3); J, uncinate (uncinate 1); K, uncinate 4); L, tyloscopule (scopule 2); M, stongylscopule (scopule 1); N, diagrammatic representation of "aragastral' surface; 0, diagrarnmatic representation of lateral surface of dictyonal franiework; P. diagrarnmatic representation of 'paragastrai' surface; Q, geographical distribution (solid dot denotes type locality).

CONORETE IJIMA, 1927

Corrorete Ijima, 1927: 165; Topsent 1928: 337; Reid, 1958: 14; Reid, l964:cli; Reiswig, 1990: 736.

Not Pityrete Topsent, 1928: 92.

TAXONOMIC DECISION FOR SYNONYMY Ijima, 1927: 165

TYPE SPECIES Ewete erectum Schulze, 1899 (lectotype designated by Wheeler. this publication)

BASIS FOR DIAGNOSIS Diagnosis is based upon Schulze's (1 899) original description of Eurefe erecttm and reexamination of a fragment of the lectotype.

DIAGNOSIS Body form erect, tubular, branched or unbranched; tubes end distaily in a central osculum and are ornamented with severai accessory oscula. Basal plate and stalk present. Regular and irregular dictyonal framework several layers thick in interna1 and proximal portions but reduced to one layer distally and marginally; dictyonal beams generally smooth: primary dictyonal strands prominent; spws present on nodes of pnrnary frarnework. Loose spicules include prominent smooth dermal pinnular hexactins, heavily spined gastral pentactins, scopules, uncinates, discohexasters, oxyhexasters and onychohexasters.

REMARKS The çenus Conorete was erected by Ijima (1927) to include those euretids having distinct pinular hexactinedermalia and pentactine or pinular hexactine gastralia. Ijima (1 927) based this genus upon Schulze's Eureté erectum. Several specimens of E. erectum were collected near the Galapagos Islands by the 'Albatross' expedition at sta. 28 19 but Schulze (1 899) figured only one whole specimen (lectotype) and one fragment (the origin of which Schulze failed to include) and gave no information regarding the other specimens. Schulze (1 899) described this new species as having a dichotomously branched, tubular. body form, smooth pinular hexactinehypodermalia. heavily spined pentactine or pimular hexactine hypogastralia, scopules with four to six tines; uncinates and discohexasters. Schulze (1 899) also included a diagrammatic illustration of the spicule location within "'an average body wall" of E. erectum but rnistakenly depicted the dermal pinnular hexactins with spiny tangential and radial rays. From a total of nineteen specimens of E. erecrum collected by the 'Albatross' off the south coast of Panama. Wilson (1904) descnbed three new subspecies. He narned E. erectzrm tubuliferrtrn (3 specimens fiom sta. 3358; 7 specirnens from sta. 3359) to includr: those unbranched body forms having onychohexasters instead of discohexasters and distally smooth scopule tines. E. erectirrn rnwronatum (4 specimens from sta. 3358: 1 specimen from 3359) for those specimens with oxyhexaster microscleres. and E. erecfurn gracile (4 specimens from sta. 3380) for those elongated and spiral body forms with onychohexasters. Having obtained more material trawled off the southem coast of Western Panama by the 'Albatross' (two specimens from sta. 3622) Lendenfeld reviewed Schulze's ( 1899) and Wilson's ( 1904) material. Lendenfeld (1 9 15) rejected Wilson's ( 1904) subspecies designation and felt the minor differences in 'superficial spiculation' were best denoted as forms. of which he established four. E, erectrrm foma A was established to accommodate Schulze's ( 1899) dichotomously branched type specimens: E. erecfrtrn forma B for Wilson's unbranched E. erecturn rzrbuliferum; E. er-ectrrm forma C for Wilson's E. erectrrrn gracile and finally E. crecrrcm forma D. for those unbranched specirnens with minor differences in scopule forrn and tine number. Lendenfeld (1 9 15) felt that the possession of osyhexasters instead of discohesasters in Wilson's E. crccrurn muer-onatum was of sufficient systematic importance to merit distinction from E. erecttrm. however. he formalized no replacement narne. As previously mentioned, Ijima (1927) used SchuIze's E. erectztm as the type species for his new genus Conorete to incorporate those euretids with pinnular hexactine dernialia and pentactine or pinnu!ar hexactine gastralia. Although Ijima (1 927) did not personally examine the 'Albatross' E. erectum material. he was responsible for some important taxonomie moves involving E. erecturn Schulze. Edi tord errors in his monograph considerably coniùsed matters. Under his list of valid euretid genera (1 927: 165) Ijima did include Conorete, but. in his "Final List" (1927: 368) Conorete was not included. However, in the "Final List" Ijima renamed E. erectum mucronatum Wilson as Conorete mucronatum Wilson and included E. erecturn erectum Schulze. E. erectum tubulr~erumWilson, E. erectum gracile Wilzon and E. erecrum forma D Lendenfeld. The next year (1 928) Topsent erected the genus Piryrere for several specimens collected near the Azores by the 'Prince Albert' expedition. The type specimen for this genus was named Pityrete azoricum Topsent and was reported as having dermal pinular hexactins. However, later in the same publication Topsent (1928) recognized Ijima's (1 927) Conorete and synonomized Pityrete to Conorete, including Conorete azoricrtm Topsent as a valid member of Conorete. Reid (1 958, 1964, 1964) was the most recent to review the taxonomie status of C'onorere and it was clear that he viewed the validity of the genus Conorete with uncertainty. refemng it to a subgenus within Eurete. Several unconfirmed specimens of Eurere erectum have been reported: Talmadge (1 973) reported several fragments possibly attributable to E. erecturn from northern Caiifomia and Foell & Pawson (1986) recently reported a photograph of a specirnen possibly belonging to E. erectum from the abyssal depths of the north Pacific ocean. Reiswig (1990) corrected several errors in Ijima's (1 927) rnonograph by indicating that symbols in his "Final List'. are interpreted to reflect Ijirna's uansfer of E. erectum forma D Lendenfeld and E. mrrcronatum Wilson to the genus Conorete ljima and that text statements clearly assigned E. erectum erecttcm Schulze. E. crectirm tubull#èrzcm Wilson, E. erectzrm gracile Wilson and E. erectrrm forma D Lendenfeld to the genus Conorete Ijima. A recent search at the British National Museum found only one specimen (the lectotype) of Schulze's type material of E. erectum and the other specimens are here considered lost. Reexamination of a fiagment of the lectotype generally confirmed Scl~ulze's(1 899) description but found: (1 ) subtle variation in scopule tine number and form (2) small differences in reported spicule dimensions and (3) the presence of rare hemidiscohexasters. A review of Wilson's (1904) description of E. erectzrm suggests that Wilson's specimens are not in possession of discohexasters. as Schulze (1 899) had ot-iginally described in the type specimen. Similarly, Lendenfeld (1 91 5) found significant differences in (1 ) body forrn; (2) the number and form of scopule tines and (3) overall spicule sizes among his four fonns of E. erectum. Thetefore, Wilson's (1 904) three remaining subspecies (Ijima ( 1927) formalized E. erecrum rnzrcronatzrrn Wilson as Conorete mucronaium Wilson) and Lendenfeld's (1 9 15) three remaining forms should be viewed as separate species within the genus Conorefe. Topsent (1928) originally based Piiyrete azoricum Topsent on AphrocaZZistes uzoricus Topsent (1 904), which, in tum, had been based upon a single specimcn misidentified earlier (Topsent. 1892) as A. ramostrs. It is clear from the description and figures of Aphrocaifistes ramosus Topsent. 1892 that it does not possess the characteristic regular framework of Conorere and has mesohexactins which are not associated with known members of Conorete. Ijirna ( 1927) informally synonomized Aphrocallistes azoricus Topsent to Aphrocallistes beutr&rramostrs Gray, 1858. Conorete Ijima is then recognized as a valid genus within the farnily Euretidae and valid members of Conorete include: C. ereciurn (Schulze, 1899)[= E. erecfum Schulze, (1899) = E. erectum forma A Lendenfeld (1 9 15)J. C. mucronaturn (Wilson.

1904)[= E. erecium mucronaturn Wilson (1904). = E. mttcronatun~(?j Lendenfeld ( 19 1 31, C. tubuliferurn (Wilson. 1904)[= E. erecirrm irrbulijétrrn Wilson (1905), = E. crectzrrn forma B Lendenfeld (1 9191. C. gracile (Wilson. 1904)[= E. erecirrm gracile Wilson (1904). = E. erectum forma C Lendenfeld (1 9 15)] and C. lendenfeldi n. sp. [= E. et-ecfztmforma D Lendenfeld ( 19 1 31.

DISTRIBUTION The Albatross Expeditions collected numerous sarnples of Conorete erectrtm Ijima between 1888 and 1904. The Albatross log was used to detennine location, depth and substrate.

Generic lectotv~e C. erectum (Schulze), several specirnens including the lectotype were collected in 1888 near tlie Galapagos Islands, Generic species C. tzthrtf~erzrm(Wilson). several specimens collected off the south Coast of Panama. C. tnucronaturn (Wilson), five specimens collected off the south Coast of Panama. C. gracile (Wilson), four specimens from the Gulf of Panama. C. fendenfildi n. sp., two specimens from the Columbian Pacific.

See PIzte 3, Fig. O for geographic distribution. Conorete erectum (Schulze, 1899) (Plate 3, Figs. A-O)

Eurete erectum Schulze, 1 899: 72, PLXVII, Fig. 1-3; Schulze. 1 904: 1 43 ; Wilson, 1 904: 62; Lendenfeld, 1915: 126. PI.XXX, Fig. 1- 17, Pl-XXXI, Fig. 1 -28; Ijima. 1927: 165; Topsent. 1928: 92; Reid. 1958: 14; Reid, 1964: cli; Talmadge. 1973: 57; Foell & Pawson. 1986: 63; Reiswig, 1990: 741.

Cortorete erec!um Ijima, 1927: 165; Topsent, 1928: 337; Reid, 1958: 14; Reid, 1964: cli; Reiswig, 1990: 736

Not Pityrete erectum Topsent, 1928: 92.

TYPE MATERIAL Conorere etvcfrrrn (Schulze. 1899) = Emre erecrum Schulze. lectotype collected by the '.4lbatross. expedition near the Galapagos Islands: currently stored in alcohol at BMNH 1908.09.24.39.

MATEFUAL EXAMINED TYPE MATERIAL: Conor-ere erecrum (Schulze) Ijima. 1927 = Eurere erectrrm Schulze (details as above)

DESCRIPTION The size and shape description is based upon Schulze's original description; the skeletal framework and spicule descriptions are based upon reexamination of a small(1cm') fragment of the type material (preserved in alcohol). Data in micrometres (pm) f

Size and shape: The illustrated (Schulze, 1899) lectotype (Plate 3. Fie. A) is erect. "Y- shaped" and tubular. The stalk, or stem (2-3 cm long: 8- 10 mm in diarnetre). is fixed to the substratum by a basal plate and is folded transversely (Reid. 1964) into two l~ollow branches (6-8 cm in height; 12 mm in diametre; walls 1 -1.5 mm thick) which. presumably, open distally as central oscules. The branches are perforated by severai (5-7) circular apertures (presumabiy accessory oscula), with short protruding rims, and are oriented in a clockwise or counter-clockwise spiral. From this, the branches may be then considered to have both gastral (interior of branch) and dennal (exterior) surfaces. Skeletal Framework (Plate 3, Figs. KM): unchannelized. three-dimensionai network of hexactinemegascleres: generally in 2 - 3 layers but may be reduced to one layer at distal (growth) margins. Primary dictyonal framework (Reid, 1964) regular (Plate X, Fig. Mj; meshes rectangular (1 65fj833pm width; 3 1 1+4434 pm length); dictyonal strands (1 65+3833pm apart) appear nearly parallel (Plate 3, Figs. K, M) but may fan-out to lateral surfaces; spurs formed fiom fiee rays of hexactins of the primary dictyonalia are straight. or slightly curved, pointed and densely microspined. Dermal framework sudaces (Plate 3, Figs. K. L) irregular; oxyhexactins (rarely spiny pentactins) commonly intercallated to primary frarnework: meshes irregular and uiangular: beams, believed to be newly fomed, are thin and fragile (Plate 3, Fig. K); false (greater than six rays) nodes formed by incorporation of oxyhexactins common. Gastral surfaces of fnmework slightly more regular than dermal surface; meshes rectangular to triangular; oxyhexactins rarely intercallated to primary framework. Bearns (6OkI pn thick) generally sinooth throughout framework: nodes unswollen.

Loose spicules (Plate 3. Figs. B-J): The dermal location and the continuous layer formed by distinct pimular hexactins (Plate 3, Fig. D) are the notable characteristics of the genus Conorere. Their densely thorned, bushy distal rays (89+33;0 length) project externaliy: the smooth and pointed proximal ray (1 14k4019length) extends into fi-amework; smooth. pointed and straight tangential rays (98k2S;0 length) are aligned paratangentially on dermal surface. Gastrally. pentactins. or pinnule derivatives (Lendenfeld. 19 15). (Plate 3.

Fig. C) with vestigial (approximately 52 - 76 pm length). sometimes weakly thorned or knob-like distal rays, are common and are manged in a similar, but less regular. network as the dermal pimular hexactins. The long (275+171; length), heavily spined, slightly bowcd tangential rays are btuntly tipped; pointed proximal rays are straight and lack spines. Microscleres (Plate 3, Figs. H-1) are of two types. Hemidiscohexasters (Plate 3. Fig. 1) (601fr852diametre) with slightl y swollen nodes are abundant; prirnary rays (6-t1 length) give rise to two or three (mode is two) curved secondary rays which terminate distally in a small disk. Similarily sized discohexasters (Plate 3. Fig. H) with weakly microspined primaq rays give rise to four curved secondary rays which terminate distally in a small disc. Scopules appear to be of two types and a size-frequency distribution was bimodal (overall length 258+1 IOs2). Sma11 (1 80-265 pm length) dermal strongyloscopules (Plate 3, Figs. F, J) with three to five straight and undivergent microspined tines (tines plus capitulurn 14-45 pm) are common; tines aise from a swollen. or unswollen (4- 7 Fm width) capitulum which is connected to a straight or slightly cunred. pointed. and weakly microspined shafi; tines are distally rounded. The four. or five. divergent and geniculate tines (tines plus capitulurn 87-1 01 pm) of the gastral tylo-scopules (Plate 3. Figs. E) (360-877 pm length) are microspined and the terminal tyle is adorned with distinct, proximally oriented, curved spines; capitulurn (7.5-9.6 pm width) and shafi are microspined and the shah terminates proximally with a sharp point. Straight, or slightfy curved uncinates (Plate 3. Fig. G) (overall length 67951 Vi8)with long curved barbs. highIy angled from the shaft. are common.

DISTRIBUTION C. erecturn (Schutze), several specimens including the lectotype were collected near the Galapagos Islands; sta. 28 19,0° 8's. 90" 06' W. depth 671 fims.. 1328 m: substrate sand (Schulze (1 899) cites O" 24'S, 89" 06' W. depth 7 17 m; substrate saiid). C ~rïhrtlz~erurn(Wilson). several specimens collected off the south Coast of Panama at sta. 3358 (6" 30'N. 81" 44' W. depth: 555 fims.. 1015 m: substrate: green sand) and sta. 3359 (6" 22' 20W, 81 " 52' W. depth 465 ftms.. 85 I m; substrate rocky). C'. mrtcronutum (Wilson). four specimens from sta. 3358: one specimen from 3359 (see above for station coordinates. C. gracile (Wilson), four specimens from the Gulf of Panama at sta. 3380 (4" 3'N. 8 1" 3 1 ' W. depth: 899 ftms.. 1645 m; substrate: rock) C. lendenfeldi n.sp., two specimens collected in the Colurnbian Pacific at sta. 4622 (6" 52' N. 81" 73' W., depth 1063; substrate sandrock). Lendenfeld (1915) cites the locality as the southern Coast of western Panama, (6" 3 1 ' N., 8 1 O 44' W., depth: 1067m: substrate: green sand and rock) ? Corxwetc. sp. cotlected by Foe11 & Pawson (1986) from the eastern Pacific (14" 40' N.. 125" 49' W), depth 445 1 m. ? Conorete sp. collected by Talmadge (1973) from northern California Plate 3 (Conorete erectum (Schulze)): Figure A, lectotype (modified from Schulze. 1899: Pl. XVII, Fig. 1); B, spiny pentactine; C, pentactine or pinnule hexactine derivative: D. pinnular hexactin; E, gastral tyloscopule; F, dermal strongyloscopu~e;G. uncinate; H. discohexaster; 1, hemidiscohexaster; Jt dermal strongyloscopule: K. primary dictyonal framework (dermal and gastral layen dissected off) of lectotype: L, low magnification of irregular dermal surface (arrow indicates thin. newly formed, bearn); M, higher niagnification of dermal surface (arrow indicates false node); N, geographic distribution (solid dot denotes type locality; open dots represent generic species; solid star represents unconfirmed specimens).

BATHYXIPHUS SCHULZE, 1899

Bathyxiphm Schulze, 1899:82, PIXVII, Fig.4-5, PI-XVIII, Fig. 1-2.; Schulze & Kirkpatrick. 19 10:v; Ijima, 1927: 165; delaubefels. 1936: 187; Reid, 1964: liii: Talmadge. 1973: 57; Reiswig. 1990:736: Mehl, 199254.

TYPE SPECIES Barhyxiphzt~.subrilis Schulze. 1899; (holotype by monotypy. Schulze, 1899: 82)

BASIS FOR DIAGNOSIS Diagnosis is based upon reexarnination of holotype material and Schulze's original ( 1 899) description.

DIAGNOSIS Body form blade-like; long, thin and lenticular in cross section: no tubes present; gastral and dermal sufaces not distinguishable; dictyonal framework regular and irregular: meshes rectangular and triangular; bearns smooth; spurs present on externai dictyonalia; nodes regular or polynodal; megascleres as pentactins (?). pinnular hexactins: tylo- and strongyloscopules; uncinates and possibly (?) pileate clavules. Microscleres as oxyhexasters, hemioxyhexasters and oxyhexactins.

REMARKS Schutze (1 899) erected this monospecific genus for the distinct blade-like body form of a single macerated specimen collected by the historic Albatros Expedition (station 2986) near Guadeloupe Isl., off Lower California. Schulze (1 899) correctly remarhd on the "farreid-like" framework of Barlzyxiphus subrilis. however. regarding its spicule component he was quite vague. Schulze (1 899) described, in some detaii. pinnular hexactins, oxyhexactins, pentactins, and uncinates. casually mentioned oxyhexasters and speculated on the presence of several scopules and clavules. However. in his illustration (Schulze. 1899: PI. XVII* Fig. 5) of the "Spicule order of an average wall" Schulze failed to include pentactins and clavules. Nowhere in this early description (Schulze. 1899) did Schulze explain how he differentiated the gastral and demal surfaces. Later (1 9 1O), while describing material collected fiom the Antarctic Schulze mentioned and illustrated several fragments he attributed to an unknown species of Bathpiphus. Although he had no spicules to report, Schulze (1910) did note the "farreid-like" framework as well as the presence of several "white. dense oval masses" found within the wall of several of these fragments. In reviewing Schulze's (1899) descriptions. Ijima (1927) simply reiterated Sc hulze md noted dennal pinnular hexactins and gastral pentactins and included Barhyxiphus Schulze as a valid genus within the Euretidae. Presumably using Schulze's original ( 1 899) description Reid (1 963) remarked on the absence of "dictyonal cortices" (distinct extemal dictyonal layers) in Burhyxiphus suhrilis Schulze and concluded that epirhyses and aporhyses may occur. Talmadge (1973) unofficially reported a new specimen from Northern California as Barhyxiphus subrilis Schulze and remarked on the similarity in spicules between his new specimen and those figured by Schulze ( 1899). In their list of known hexactinellids Van Soest and Stentoff (1 988) confused the Guadeloupe location as Schulze's holotype as the West Indies. Finally, based upon Schulze's (1 899) casual mention of clavules within the holotype, Mehl (1 992) transferred Buthyxiphus from the Euretidae to the Farreidae. Records show a Eure~esp. and Furrea occu Bowerbank were collected together with B. strbtilis at the Afbatross sta. 2986. and may have been the cause of contamination. Reexarnination of fragments from the holotype has not resoIved the true spicule complement of Barhjmphus sztbtilis Schulze. Although pinnules, oxyhexactins, uncinates. scopules. clavules and oxyhesasters were observed in spicule preparations none of Schulze's ( 1899) complete "gastral-like" pentactins were found. No channelization was observed in the dictyonal framework contrary to Reid's (1963) suggestion. Until new material becomes available it remains unlikely that the genus Buhyxiphrcs Schulze will be properly understood. Also. the identity of Schulze's (1 9 10) Antarctic fragments and Talmadge's northern Californian specimen will remain uncertain until this rnaterial can be examined. The genus Barhyxiphzrs Schulze is thus retained within the farnily Euretidae.

DISTRIBUTION Burhy,riphus subrilis Schulze, holotype collected south of Guadeloupe Isl.. off Lower California.

Bathyxiphus subtilis, Schulze (Plate 4. Figs. A-O) Bathyxiphus subtilis Schulze, l899:9O, PI.XVII, Fig.4-5. Pl.XVZI1, Fig. 1-2.; Schulze & Kirkpatrick, 191 O:7; Ijima, 1927:165; Talmadge, 197357; Reiswig, l990:736; Mehi, 1992: 56.

TYPE MATERIAL: Barhyxiphus subtilis Schulze, holotype currently stored wet at USNM 7528: several fragments of a single specimen collected by the U.S.S. Albatross Expedition station 2986.

MATERIAL EXAMINED TYPE MATERIAL: Buthyxiphzis srtbt ilis Schulze (see above for details)

DESCRIPTION Size and shape based upon Schulze's (1 899) original description. Skeletal and spicule descriptions are based upon re-examination of two (A and B) small (A=6 mm X 3 mm; B= 1.3 cm X 7 mm) fragments of the holotype (preserved in alcohol) together with the original literature description (Schulze, 1899). Data in micrometres (pm)+ st-dev.~.

Size and shape: The illustrated holotype (Phte 2, Figs. A. G, N) is approximatley 30 cm in Iength and is comprised of three pieces. Generally, the sponge tapers from 5-1 0 mm in width and 2-5 mm in diickness from tip to base. The body fom is elongate, blade-like lenticulur in cross-section with two sharpened and slightly wavy sides. with a basal plate 2-3 n~rnthick and 2 cm' in area. Unable to detennine functional gastral and demal surfaces.

Skeletal Framework (Plate 4. Figs. B. H. L): The framework can be divided into interna1 and exterior regions. lntemally, the primary framework is comprised of smooth beams

(46.329.3 50 pm thick), connected by regular. unswollen, six-rayed nodes. in a roughly regular cubic meshwork (36211 1 17 pm width) (Plate 4. Fig. L). Distal and extreme edges of both margins consist of one layer of the primary skeleton. These layers increase to 4-5 in number towards the centre of the body. Primary dictyonal (Reid. 1964) strands (289+3911 Pm apart) are oriented parallel to the growth avis and eventually fm out to lateral margins. On external surfaces microscleres and uncinates are ofien soldered to bearns tl~erebyforming a more dense and irregularly shaped secondary frarnework. Free rays of extemal dictyonalia project as spurs. Extemal surface (Plate 4, Fig. H) nodes not swollen and oflen false (greater than six rays). Basal sections of frarnework are thicker and denser than dista1 portions due to increased intercallated oxyhexactins. No osculum, tubes or channels are present.

Loose spicules (Plate 4, Figs. C-F. 1-K, M): Pentactins (Plate 4, Fig. C) are reported (unconfirmed in re-examination) to display four terminally bowed. slightly tubercled. tmgential rays. a smooth proximal ray and an occasional knob of the vestigial distal ray. Similarly, hexactins (Plate 4, Fig. 1) with pinnular distal rays are present on both surfaces. Scopules appear to be of two types although they were too rare for a size-fiequency distri bution to be performed (252+ 144~(5 - 403) overdl length). Small. rare strongyloscopules (Plate 4, Figs. E) with two to four straight and undivergent microspined tines (tines plus capitulum 57%) arise from an unswollen (9.8k2.47 width) capitulum, which is co~ectedto a straight, pointed, and weakly microspined shafi. The five, slightly divergent and weakly bent tines of the tylo-scopules (Plate 4, Fig. D) are microspined except for the top-most region, giving a "bdd" appearance; tines terrninate distally with small proximally oriented curved thorns; the capitulum is not swollen and the shafi is weakly microspined. Pileate clavules (Plate 4. Fig. J: 574~3801j (1 5 - 1283) overall length) with distinctly serrated caps and weakly microspined shafts are common (possibly of contaminant origin). Microscleres (Plate 4. Fig. K) are comrnonly oxyhexaxsters (76k2O2?( 16 - 106) diarnetre). Long primary rays ( 19f 6.gzj) terminate distally in four sharply pointed and slightly divergent secondary rays. Small, smooth oxyhexactins (Plate 4. Fig . M) with slightly (6.1 + 1 .gS0)swollen nodes are abundant and have equal length

(jO.2f 3.1 100 (6 - 100) length), straight, and pointed rays. Hemioxyhexasters are rare. Uncinates (Plate 4, Fig. F; 1025S3449(49- 1426) overall length) with moderately long barbs, significantly spreading from shaft. are common.

DISTRIBUTION Ba~hyxiphussubtifis Schulze, holotype collected south of Guadeloupe Isl., off Lower California: 28"57N, 1 18' I4'3O"W. Depth: 125 1 m; substratum: mud. Barhyxiphus sp. Schulze, collected in the Antarctic (66"2'9"S, 89'38'E). Burhyxiphrrs sp. Talmadge, collected off Northem California. Plate 4 (Bafhyxiphus subtilis Schulze): Fig. A, distal fragment of holotype (modified fiom Schulze, 1899: 90, Pl. XVII, Fig. 4); B, diagrammatic cross-section through wall showing spicule location (Schulze, 1899; PI. XVII, Fig. 5); C, diagrammatic representation of pentactin according to Schulze (1 899);D, tyloscopule; E, strongyloscopule; F. uncinate: G, rniddle fragment of hoIotype; H. diagrammatic representation of external dictyonal framework: 1. diagrammatic representation of a pinnule according to Schulze (1 899); J, pikate clavule (possibly of foreign origin); K, oxyhexaster; L, diagrammatic representation of interna1 dictyonal framework; M, oxyhexactin; N, basal portion of holotype; O, geographical distribution (solid dot denotes type locality, open dots are localities of unconfirmed specimens).

PARARETE IJIMA, 1927

Eurete Semper (in part) Levi & Levi, 1982: 298; Tabachnick, 1990: 173.

Pararete Ijima, 1927: 178; Okada, 1925: 285; Reid. 1963: 224; Levi, 1964: 104; Mehl. 1992: 68; Reiswig & Mehl, 1994: 153; Barthel & Tendal, 1994: 58.

TAXONOMIC DECISION FOR SYNONYMY Ijima. 1927: 178

TYPE SPECIES Ezrr-cfefamopsis Carter, 1877: 12 1 (neotype designated by Wheeler. this publication).

BASIS OF DIAGNOSIS Diagnosis based upon reexarnination of neotype (see Remarks) together with Schulze's ( 1887) description and Ijima's (1 927) generic diagnosis of Pnrarere.

DIAGNOSIS Body form bush-like: network of branching and anastomosing tubes: dictyonal fiamework regular and irregular; meshes triangular or rectangular; nodes commonly swollen and tubercled; loose spicules as pentactins, scopules. uncinates, and discohexasters; no oxyhexasters.

REMARKS The genus Purclrete was erected by Ijima (1 927) for those euretids having discohexasters as the only microscleres. Ijima (1 927) based this new genus upon Carter's (1 877) Errr-etc fut-1-eopsis. the second oldest species of Errrete Semper. Carter ( 1877) illustrated and described a large (10 cm height) bush-like specimen of E. farreopsis (fiom Dr. Millar) from the Philippines as having distinctive four-tined scopules, barbed uncinates. oxyhexactins and small "fleur-de-lis" discohexasters. Carter (1 877) also briefly mentioned a second smaller specimen of E. farreopsis collected by Dr. Meyer. from the Philippine Islands. In his report on material collected by the 'Challenger', Schulze (1 887) reviewed Carter's ( 1877) description and redescribed E. farreopsis based upon material from sta. 192 near Little Ki Island. Although Schulze (1 887: PI. LXXIX, Fig. 5) figured and initially referred to a single specimen. his list of specimens coilectcd by the 'Challenger' ( 1 887: 477) indicates that two specimens were fond. Schulze (1 887) described the 'Challenger' specimen(s?) with a tubular feltwork body fom, square dictyonal meshes with slightly tubercled beams and thickened nodes, osyhexactins. discohexasters with "perianth-like", curved secondary rays. pentactins. uncinates and geniculate-tined scopules. As previously stated, Ijima (1927) erected Parurete for those euretids having discohexasters instead of oxyhexasters and based the genus on Carter's Eurete furreopsis. however, he (1927) did not mention if he had examined Carter's original specimens or simply described Pararete fiom Carter's original description or from five specimens identi fied as P. farreopsis coilected by the 'Siboga' . Ij ima ( 1927) transferred several other species fiom the genus Eurefe to Pururete (E. gerlachi Topsent. 190 1b; E. semperi Schulze, 1887 and E. curferi Schulze 1887) and described three new species (P. baliense. P. hngeungunrtnr and P. freeri) and three subspecies of Pararete (P. farreopsis subglobosrtm, P. farreopsis jakosalemi and P. furreopsis fiagjénim). Okada (1 932) later described two new specimen of E. farreopsis but did not mention why he did recognize Ij ima's Pururete. As with many other genera within the Euretidae. Reid (1963) recognized Purarete as a subgenus within Eurete but made no significant obsemations or tasonomic decisions. Levi (1 964) reported one specimen of E. furreopciis from the collections of the 'Galathea' and later (Levi & Levi, 1982) reported another specimen from New Caledonia. Levi (1 964) initially recognized Ijima's Pararete but. in reviewing (Levi & Levi, 1982) the taxonomie history of Eurete Semper. decided that E. farreopsis (the next oldest species described under Eurefe Semper) be considered the generic type species of Eztrete Semper. and there fore synonornyzed Pararete with Eurere. From material collected by the SRV 'Professor Shtokman' from the Nasca or Sala-y-Gomez range, Tabachnick ( 1990) described a specimen of E. furreopsis with oxyliexasters and graphiocomes and in her dissertation on hexactinellids of the Mesozoic, Mehl (1 992) retumed E. farreopsis to Ettrefe and redefined the type species of Pararete as P. semperi Schulze. Barthel & Tendall ( 1994) reviewed P. gerluchei (Topsent) and included it in their review on hexactinellids fiom the Antarctic. A recent search for Carter's (1 877) original specimens of E. farreopsis was not successfûl. however. a preliminary examination of four slides labeled as Carter's Type slides at the British Museum suggests that oxyhexasters and pentactine megascleres are common. Carter (1 877), however. made no mention of either spicule type in his original description, and for this reason, the origin and value of these slides is dubious. A search for Schulze's 'Challenger' (1 887) specimen(s) was successful and the location of collection of one specimen (Little Ki Island) is close to Carter's (1 877) "Philippine" locality. Although macerated, this specimen. (BMNH 1887.10.20.122) is therefore designated as the neotype for E. farreopsis. Reexarnination of a fragment of the neotype confirmed Schulze's (1 887) description and oxyhexasters were not found, however, contrary to Schulze (1 887) small bracketed uncinates without barbs were observed together with larger. barbed uncinates. A preliminary examination of Okada's (1 936) specimens of E. furreopsis revealed substantial differences from Carter's (1 877) and Schutze's (1 887) descriptions and neotype material, and until these specimens can be fùrther analyzed they are considered as unidentified species of Purarete. Ijima's (1927) erection of Put.ar-etc was controversial. as pointed out by Levi & Levi (1982). however, for reasons outlined in the "Remarks" section of Ewere Semper (see page 38) E. farreopsis is still considered as the type species of Pararete. As Tabachnick's (1990) specimen from the Nasca or Gomez range does contain oxyhexasters and graphiocomes it cannot be considered as E. frrrreopsis; its taxonomic status remains unresolved. Mehl's (1 992) actions are not clear and she included no reasons for the replacement of E. farreopsis as the generic type species of Parurete. As she did not review the taxonomic status of Pararete, or declare speci fi c type specimens, her actions cm not be considered authoritative. E. fnrreopsis Carter is thus here retained as the valid type species for Pararefe.

DISTRIBUTION Generic Tv~e: Et,rc?~cfin-reos Carter. neotype col lected near Little Ki Island.

Other S~ecies: P. farr-eopsis farreopsis. Ijirna, collected south of Kur 1.. Kei Island. P. furreopsis subglobosirrn Ijima. collected in the China Sea. P. furreopsis jakosulemi Ij ima, obtained near Pearl Bank, Sulu Island. P. farreopsis fragi3rtrm Ijima, collected from the Halmaheira Sea. P. carteri (Schulze), collected near Little Ki Island. P. gerlachi (Topsent), coilected from the Antarctic. P. semperi (Schulze), collected south of Kur I., Kei Island. P. baliense Ijima, obtained south of Kangean Island in Bali Sea. P. kangeanganum Ijirna, obtained south of Kangean Island in Bali Sea. P. freeri Ijima obtained near Pearl Bank, Sulu Island. Puramte sp. Okada, coltected west of Goto Island. Pcrrur-ete sp. Okada, collected off Kagoshima Gulf.

Pararete farreopsis, (Carter) (Plate 5, Figs. A-3)

Eurete farreopsis Carter, 1877: 12 1, PI.IX, Fig. 1-7; Schulze, 1887: 295, PI.LXXIX, Fig. 5-8: ijirna 1927: 165; Okada, 1932: 49; Levi, 1964:104; Mehl, 1992: 70.

Not Eurere farreopsis Carter sensu Levi & Levi. 1982: 298; Tabachnick. 1990: 172.

PurtrrereJarreopsis (Carter). Ijima. 1927: 1 78: Levi. 1964: 1 (W.

TYPE MATERIAL Ettrefefut-reopsis Carter, neotype. designated here, collected by the 'Challenger' off Little Ki Island; currently stored in alcohol at BMNH l887.lO,2O.l22.

MATERIAL EXAMINED TYPE MATERIAL: Ere-ere.furreopiS Carter. neotype (details above).

OTHER MATERIAL: Pururere sp. (= E-furreopsis sensu Okada, 1 932): USNM 22047 collected from the Eastern Sea, Ose Saki Lt.. West of Goto Iles., Albatross Stn. # 4890: Depth 247 m. Pcrrurefc sp. (= E.farreopsis sensu Okada, 1932): USNM 22048 collected from the Eastern Sea, Sata Misaki Lt, off Kagoshima Gulf. Albatros Stn. # 4934: Depth 280m.

DESCRIPTION The following description of body form is based upon Schulze's (1 887) original description and illustration. Description of the skeletal frarnework and spicules is based upon examination of a small (km') fragment of the neotype material (preserved in alcohol) which is incomplete and washed-out. Data in micrometres (pm)It ~t.dev.~. (range).

Body fonn (Plate 5. Fig. A): The illustrated fragment of the neotype is washed-out and macerated; bush-like; height 3.5 cm; width 2.5 cm; complex network of branching and anastamosing tubes; tube apertures circular or fiuinel-shaped, 5-8 mm diametre; tube wall 0.5 - 1 .O mm thick; walls thicker at base than distal margins; attachent to substrate unknown.

Skeletal Frarnework (Plate 5, Figs. H-1): Dictyonal hework is a regular. and irregular, three-dimensional network of hexactin megascleres; mesh rectangular to triangular: wall thickness 0.5- 1 .O mm, one dictyonal layer thick at margins increasing at basal and interna1 reg ions; dictyonal strands evident, orientation unknown; true dictyonal nodes swollen and beset with small(1-2 pn) blunt tubercles (Plate 5, Fig. 1); false nodes (axial cross absent) rare1y swollen: 1-3 finger-li ke, spined. spurs commonly present on nodes; bearns (52 1 lds um), formed by ankylosis and synapticulation, are smooth or sparsely microspined; round gaps (Plate 5, Fig. H; 500-700 um in diametre) in dictyonal framework as diarhyses-li ke channelization.

Loose Spicules (Plate 5' Figs. B-G): Small. smooth oxyhexactins (Plate 5. Fig. D) with equal length straight, terminally pointed. rays (38&249 pm length) are abundant in the parenchyma or are often fused. by one ray. to the dictyonal framework. Hypoderrnalia

and gastralia as pentactins (Plate 5, Fig. B). Tangential rays (2OO+3 6 1 1 1 pm len-~) slightly bowed, terminally rounded and microspined; proximal ray (169k489 pm length) slightly shorter than the terminally rounded and microspined tangentials. Parenchymal

microscleres as discohexasters (36+ 3 100 pm diametre) (Plate 5, Fig. G). Four "s-shaped" secondary rays with terminai serrated discs emanate from short (4f 1100 pm length) primary rays. Tyloscopules (Plate 5, Fig. E) (209f32jo pm length) regularly with four, rarely with five or six. geniculate tines (49+gs3 pm Iength) present on dermal and gastral surfaces. Tines terminate distally in a club, which is ornarnented with proximally oriented recurved thorns. Straight, or curved, uncinates are found in the parenchyma. Long uncinates (Plate 5. Fig. C) (1 150-2200 pm length) with prominent barbs held closely to the shafi, and smaller (250-1 100 pm length; Plate 5. Fig. F). bracketed uncinates (barbs visible with oil immersion light microscopy) are straight or curved and found in the parenchyma (uncinate size distribution reveals a bi-modal distribution: overall length: 954t 5 1 7100 pm).

DISTRIBUTION Distribution of tvDe swcies Author # Expedition Location Depth Specimens

Carter 1877 l? Dr. J. Millar Philippine Islands 9 Schulze 1887 l? Challenger Stn-ff 1 92 5O49'S. 132O14-E 234 m Ijima 1927 Frags. Siboga Stn. #97 5"48'S, 119O49'E 564 m Ijima 1927 +3 Siboga Stn. #25 1 5"28'S, 1 3S00'E 204 m Ijima 1927 2 Siboga Stn. #254 5040°S, 132O36.E 3 i O m Levi 1964 1 Galathea Stn.#423 1 0°27'S. 124" 18'E 8 10 ni LeviaLevi 1982 1 MNHN HCL 54 22"49'S, 167" 12.E 395 m

Distribution of other species P. furreopsis farreopsis (Carter). 'Siboga' sta. 25 1 (5O28'S. 132"O'E). depth 204 m. P. fo-reopsis subglobosum Ijima, ( 17O44'N. 1 10°26'E). depth 220m. P. furreopsis jakosdemi Ijima, 'Siboga' sta. 97 (5O48'N. 1 19"49'E), depth 564m. P. furreopsis fragzyerum Ijima, 'Siboga' sta. 156 (0°29'S, 130°5'E), depth 469 m. P. carreri (Schufze), 'Challenger' sta. 192 (5'493, 132" l4'E). depth 244m.

P. gerlachi (Topsent), (70"-7 1O 1 8'S1 8 1"92"W), depth between 400-569m. P. semperi (Schulze), 'Challenger' sta. 25 1 (5'28's: 13Z0O'E). depth 204 m. P. buliense Ijima, 'Siboga' sta. 15 (7O2'S. 1 1S023'E), depth IO0 m. P. kmgeungunum Ijima 'Siboga' sta. 15 (7OS'S. 1 1 S0S3'E). depth 100 m. P. fi-eer-i ljima. 'Siboga' sta. 97 (S048'N, 1 19O49'E). depth 564m. Pr,)-u~-c.resp. (Okada). 'Albatross' sta. 4890 (32'26'N. 1X036'E), depth 247m. Purcrrerc sp. (Okada). 'Albatross' sta. 4934 (30°55'N, 1 30°32'E), depth 190 m. Plate 5 (Pamrêrefarreopsis (Carter)): Figure A, diagrammatic illustration of neotype (taken from Schulze, 1887: Pl. IX, Fig. 1); B. pentactine megasclere; C. barbed uncinate; D. oxyhexactin; E, tyloscopule; F. bracketed uncinate; G. discohexaster; H. photograph of dictyonal skeleton (g = gaps); 1, higher rnagnification of dictyonal skeleton showing swollen. tubercled nodes; J. geographic distribution (solid dot denotes Carter's type IocaIity. open star denotes neotype iocality, open dots denote other specimens of E. firrr-eopsis. open squares denote locality of other Pararete species. solid square denotes unidenti fied specimens).

PLEUROCHORIUM SCHRAMMEN, 1912

Pleurochorium Schrammen, 1912: 25 1; Zittel, 191 5: 74; Ijima. 1927: 165; de Laubenfels, 1955: E86; Reid, 1958a: 265; Reid, 1958b: 14; Burton. 1959: 153; Reid, 1964: Ivi (Fig.); Mehi, 1992: 69.

Ewete Semper, 1868: 372: (in part) Ijima. 1927: 196.

TAXONOMIC DECISION FOR SYNONYMY Ijima 1927: 195

TYPE SPECIES PZettrochorium schuizei T Schrammen, 1912: 25 1 (by original designation).

TYPE SPECIES FOR MODERN REPRESENTATIVES Ple~crochoriirrnannandaki (Kirkpatrick), originally Eurete annandalci Kirkpatrick. 1908.

BASIS OF DIAGNOSIS Diagnosis based upon reexamination of lectotype material of Eztrere crnncrndulei Kirkpatrick together with Kirkpatrick's (1 908) description and Ijima's (1 927) generic diagnosis of Pleurochoriurn.

DIAGNOSIS Body form tubular, erect, with conspicuous plate-like external appendages: dictyonal framework regular or irregular; rnesh rectangular or triangular; dictyonal strands prominent; beams smooth or sparsely microspined: tnie and faIse nodes: pentactin or hexactin rnegascleres, scopules, discohexasters and uncinates as loose spicules.

REMARKS Schrammen (1912) erected this poorly known genus. for a single fossiiized sponge framework (P. schulzei i-) fiom the Cretaceous, to include sponges with conspicuous plate4 ke external appendages. Ijima (1 927) later recognized the distinct body form of the extant Ezrrere annandulei Kirkpatrick, 1908 as a rnember of Pierrl-ochor-iron. Kirkpatrick (1 908) originally descebed Eurerc unnundalc.i liom the Indim Ocean as having mesohexactins. scopules. discohexasters and uncinates. Of the seven specimens that Kirkpatrick collected ( 1 908), only one, the specimen here selected as lectotype (that Kirkpatrick illustrated (1 908, PI. 1, Fig. 2); see below), was not macerated and still contained spicules for Kirkpatrick to descnbe (Kirkpatrick did not descnbe any dermalia or gastralia). Ijima (1 927) also descnbed a new species- P. cornufurn Ijima, with similar body form and spiculation. De Laubenfels (1955) erected the farnily Wapkiosidae fûr Schrarnmen's Pfeurochot-ium and wrongly cited the type specimen as P. ferchuizei. Burton ( 1959) later descnbed several specimens of P. annundalei fiom the Maldives and Reid ( 1964: IV)only cornrnented on the development and origin of the lateral plates of E. unnondalei. Mehl (1 992) reexamined Burton's material and mistakenly identified one scopule type as a sarule. A recent reexamination of the lectotype material of E. unnundalei Kirkpatrick found no spicules and a preliminary examination of Burton's material (Burton, 1959) suggests that it merit separation from P. unnundalri as P. bttrroni n. sp.. The genus Pleurochoriurn is considered a valid member of the Euretidae.

DISTRIBUTION Generic Tv~e PZerrrochoriurn schulzei t Schrammen, 1 9 1S.

Generic Tvpe for modem re~resentatives Eurete annandalei Kirkpatrick. 1908. lectotype (BMNH: 1907.08.09.002) collected by the 'Investigator' (station Z.E.V. 1422) from the Indian Ocean. Eurete annandaki Kirkpatrick, six paralectotypes (currently only one found at BMNH: 1907.08.09.00 1) were coIlected by the 'Investigator' (station Z.E.V. 2 145).

Other Species Plezcrochorium cornttrum Ijima, two specimens and one fragment collected by the Siboga from Station 297 near Timor. P. hwroni n-sp. collected from the Maldives (originally described as P. annanduki by Burton, 1959).

Pleurochorium annandalei, Kirkpatrick (Plate 6. Figs. A-L)

Eurere annandalei Kirkpatrick, 1908: 2 1, PI.1. Fig. 1 - 13; Ijima, 1927: 166; Burton, 1959: 178; Reiswig, 1990: 744; Mehl, 1992: 79.

PIcuroc/~oriumannondalei Ijima. 1937: 196; de Laubenfels, 1955: E86; Reid. 1964: iv (Figd); Tabachnick, 1989: 50; Reiswig. 1990: 735; Mehl, 1992: 79.

Not P/et~roc/roriumanncmdafei Burton. 1959: 1 53. TYPE MATERIAL Eurete annondalei Kirkpatrick, 1908, one partial specimen, the lectotype (lectotype by subsequent designation) collected by the expedition 'Investigator' (station Z.E.V. 1422) from the Indian Ocean. (currently stored dry BMNH 1907. 08.09.002)

MATERIAL EXAMiNED TYPE MATERIAL: Eurerc mnandaki Kirkpatrick, 1908 (details above).

OTHER MATERIAL: Pletu.ochoriitm bur~oninsp. (= P. annundalei sensu Burton ( 1959) and Mehl ( 1992)). collected (currently stored at BMNH 1 936.03 .O4.OO6) by the Murray Expedition near the Maldives; 4'49'N, 72O46'E to 4O48'N. 7Z040'E; depth 878; substrate green sand.

DESCRIPTION Size and shape description based primdy upon Kirkpatrick's (1908) original description combined with reexamination of the complete lectotype. The skeletal description is based upon a small (2 cm') washed-out fragment of the lectotype. Spicule information is takrn primarily from Kirkpatrick's (1908) original description and several fragments of non- type material (Burton. 1959) were examined for supporting evidence.

Size and shape: The dry-preserved fragment (Plate 6. Fig. A) is an erect cylindrical. hollow tube 9 cm in height. diametre 7- 10 mm, with fourteen vertical lateral larnellar branc h-li ke appendages (width of sponge including lateral appendages 3 -5- 4.5 cm). The lateral appendages (Plate 6, Figs. 1, J, K) are at right angles to the centml tube and are arranged in four vertical longitudinal series and in opposite pairs with each pair forming one axis of a cross (2)(Plate 6. Fig. J). Saddle-shaped Iateral appendages (2-5 mm thick; 1.5 - 2.0 cm width and length) with lateral edges curving downwards have a well-defined round orifice (3-4 mm diametre) on both the upper and lower surfaces, presumably as oscules (Plate 6, Figs. 1, K). The appendages are connected to the central tube by means of a short, hollow, laterall y compressed stem. Extemal and internal surfaces deterrnined as dermal and gastral respectively.

Skeletal Framework (Plate 6. Figs. G. H): Dictyonal frarnework is an irregular three- dimensional network of hexactin megascleres; mesh triangular; wall thickness 0.5-0.7 mm: layering not apparent; dictyonal strands prominent (Plate 6, Fig. G), orientation unknown. True dictyonal nodes on external surfaces with one microspined and curved spur; false dictyonal nodes (without axial cross) without spurs: bearns (49f um). formed by ankylosis and synapticulation, are smooth (Plate 6, Fig. H).

Loose spicules (Plate 6, Figs. B-F): Dermalia and gastralia unknown. Location of hexactins (Plate 6, Fig. B) not confirmed. Rays (400450 um Length) are uniformly rnicrospined, straight or curved and terminal1y rounded. Scopules of two types: ( 1 ) large (436 um length) tylo-scopules (Plate 6, Fig. C) with four or five tines are found on dermal surfaces. Tines (62 um length) are geniculate and microspined: distal end swollen and beset with nurnerous recurved thoms. Capitulurn microspined and slightly swoilen. The smooth shafi may be straight or curved and is terminally microspined and bluntly pointed. (2) Iarger (694 um length) oxy-scopules (Plate 6. Fig. D) with five smooth. lanceolate and distally sharpened rays (82 um length) are also found on the demal surface. Capitulurn microspined and slightly swollen. Presence of both scopules on gastral surface is not confirmed. Parenchymal spicules as discohexasters (diametre 45 um)(Plate 6. Fig . E). Four 's-shaped', or curved, secondary rays (16 um Iength) distally tipped with minute disks arise from long (6.3 um length) prirnary rays. Curved uncinates (Plate 6. Fig. F) (1 300 pm length) with prominent barbs held closely to the shafi are present.

DISTRIBUTION

Author Date ff Expedition Location Specimen Kirkpatrick 1 908 1(icctotype) Investigator Stn- # 1422 7"55'N, 8 1"47-E * Kirkpatrick 1 908 6(syntypes) Investigator Stn. # 2 145 j04'N. 80°22'E *

Other species of Pleurochoritrrn Ij ima 1927 2+ Siboga Stn. #297 1 0°39'S, 123'40'E Burton 1959 5 Murray Stn. # 152 4O49'N. 72O46'E to 4O48'N. 72O40'E Plate 6 (Pleurochoriurn annandalei Kirkpatrick): Figure -4, lateral view of lectotype (modified from Kirkpatrick, 1908: PI. 1, Fig. 1); B, microspined hexactin; C. tyloscopule; D. oxyscopule; E, discohexaster; F, curved uncinate; G, photograph of external framework (dictyonal strands follow direction of mw);H, higher magnification of framework showing smooth beams and synapticulation; 1, diagrammatic representation of upper suface of lateral plate; J, diagrmatic representation of lateral plate alignment to central axis (large black circle represents central axis, smaller black circles represent oscula on lateral plates); K, diagrammatic representation of underside of lateral plate: L. geographic distribution (solid dot denotes lectotype locality, open dot denotes syntype locality; open square represents other species locality).

VERRUCOCOELOIDEA REID, 1969

Kerrucocoeloidea Reid, 1969: 485; Rigby & Mohanti, 1993: 919

TYPE SPECIES lferrucocoeioiàea burtoni Reid, 1 969 (holotype by original designation, Reid. 1 969:485)

BASIS OF DIAGNOSIS Diagnosis based upon Reid's (I 969) original description and reexamination of holotype materiai.

DIAGNOSIS Body form funnel-like; tubular lateral outgrowths on extemal surface; surface of central atrium with longitudinal series of prominences. Regular dictyonal frame meshwork basically elongate-rectangular; beams smooth, rarely microspined: nodes smooth; spurs present. Loose spicules as pentactins, hexactins. tyloscopules, uncinates, oxyhexasters and discohexasters. Irregular scopuies common and hexaster variants rare.

REMARKS Reid erected (1969) this most recent genus within the Euretidae to encompass the distinct funnel-like body fom and spiculation possessed by two specimens collected near Bomeo. Reid designated one specimen as the holotype (BMNH 1967.5.23.1) and the other as paratype (BMNH 1967.5.23 -2)of V'errricocoeioidea brrrtoni Reid. 1969. The name Verrzrcocoeioidea was used by Reid to denote the strong similarity of body fonn to Veri-ucocoeiiaEtallon, a fossil genus known only by body form and framework. Reid (1 969) also recognized a strong similarity in spiculation and body form between Vcrrrrcocoeloideu burtoni Reid and two members of the genus Periphrageliu Mars ha1 1. 1 875. P. chcdicngeri Ij ima and P. purva Ijima, and considered making the two genera synonymous. However. since the type P. elisae Marshall did not resemble the body form of L! hzirtoni, he recommended that the two genera. Verrz~cocoeloideaReid and Periphrugella Marshall, remain separated. Not until the type specimens of P. challengeri Ijima and P. parva Ijima are reexamined cm Reid's suspicions be verified. Reexamination of holotype material confirms Reid's original description and suggests that the spicules that Reid referred to as "irregular variants approaching diasters" are Iess common than intirnated. Material fiom the paratype was found to be washed-out and without spicules. The genus Verrtrcocoeloideu Reid is retained here as a valid genus within the farnily Euretidae.

DISTRIBUTION Verrzrcocoeloidea burioni Reid, 1969; holotype and paratype collected off Brunei, Bomeo. Depth 200 m. (see Plate 7, Fig. J)

Verrucocoeloidea burtoni, Reid (Plate 7, Figs. A-J)

Verrucocoeloidea burtoni Reid, 1 969: 485, Fig. 1 -8.

TYPE MATERIAL Verrzrcocoeloidea brrrtoni Reid. 1969: holotype part of the Dampier Collection coliected off Brunei, Bomeo currently stored wet as BMNH 1967.5.23.1-

MATERIAL EXAMINED TYPE MATERIAL: I/Lrr-zrcocoeloidea btrrtoni Reid (details above).

OTHER MATERIAL: Ve~.rzrcocoeloideaburtoni Reid. 1969; paratype (same location as holotype); stored wet as BMNH 1967.5.23.2.

DESCRIPTION The size and shape description is based upon Reid's (1 969) original description of hrrrrcocoeloidea burioni. Skeletal and spicule descriptions are based upon examination of a small fragment (2 cm2) of the holotype. Data in micrometres (pm)t sr-dev.~.

Size and shape: The nearly complete specimen (Plate 7. Fig. A) is an erect fuiinel 1 1 cm taIl, and 2.9 cm wide at the distal margin. The specimen was secured to the substrate by a basal plate from which a nearly smooth and conical stalk extends. Reid (1 969) describes the inside surface of the central atrium with longitudinal series of prominences 4-5 mm wide extending (5 mm) into the atrium. External surface with distinct lateral outgrowths presumed as accessory oscula (= parietal oscula. Reid, 1969). arranged in nearly longitudinal series. Lateral outgrowths are rudimentary and hood-like just above the stalk and tubular (5-9 mm diametre) towards the distal margin. Wall thickness unknown. Skeletal Framework: Dictyonai framework is a regular three-dimensional network of hrxactin megascleres. Regular, layered (2-3 layers), intemal meshwork basically elongate rectangular (936+289 pm length; 3 18L-1029 pm width; Plate 7, Fig. 1) but rnay be locally obscured by intercalated small hexactins. Extemai surfaces less regular than interna1 meshwork: synapticula present: meshes triangular. Dictyonal strands prominent. orientation presumably parallel to axis of gmwth, ofien with slight lateral spreading beams connecting dictyonal strands are arranged to forrn transverse lamellae (Reid. 1969: 386; Plate 7, Fig. 1). Dictyonal nodes are smooth and finger-like microspined spurs are present on external fiamework nodes. Dictyonal beams (65+1 7s7pm thick) smooth. rarely microspined, and are commonly formed by ankylosis. Reid (1 969) reports skeletal caps (1 mm wide) in the external ("demai") surface fiamework as ostia. C

Loose spicules: Dermalia and gastralia as pentactins (Plate 7. Fig. C). Unequal length (266+43108 pm) tangentid rays are slightly bowed, microspined and terminally blunt; proximaI ray smooth or microspined, 0.1 5 - 0.25 mm length (Reid. 1969). and tenninolly blunt; sixth ray often as vestigial knob. Rays of freely occuring hexactins (Plate 7. Fig. B) of unequal length (6O+-2Og7 pm), straight or curved, distally microspined and terrninally blunt. Tyloscopules (429+62103pm overal length: unimodal size distribution) reçularly on both surfaces (Plate 7. Fig. D). Six to eleven divergent. straight. microspined

tines (9621 411 1 pm length) tenninate distally as a club, which is heavil y beset with numerous. proximaliy directed, curved thoms; shaft is commonly smooth, straight. and

proximally blunt. Tines mise from a slightly swollen. smooth capitulum (14+31 11 pm width) which may be significantly enlarged in irregular tyloscopules (Plate 7. Fig. F).

Microscleres as oxyhexasters and discohexasters. Oxyhexasters (Plate 7, Fig. E; 58+38 101 pm diarnetre; unimodal size distribution) with 2, 3 or 4 sharply pointed. cuwed. secondary rays and short (5+1 Iol pm length) primary rays are abundant. Discohexasters

(48+ 1 SZC)pm diarnetre; Plate 7. Fig. H) have short (6k 1 28 pm) primary rays which give rise to 2. 3 or 4 slightly cuwed secondary rays. Secondary rays are tipped with minute four-pronged disks that appear spherical at iow magnifications. Discohexasters common. The number of secondary rays per primary ray may Vary within individual oxy- and discohexasters. Irregular forms of oxy- and discohexasters (= diasters Reid. 1969: 488) are rare. Straight or curved uncinates (3.6k0.7 mm length: Plate 7. Fig. G) may occur freely or fused to the dictyonal frarnework; barbs are outwardly curved and divergent from the shaft.

DISTRIBUTION Vcrrz~ocoeloideaburtoni Reid, 1969: holotype and paratype collected off Brunei. Borneo (5" 5' N; 1 14" 6'W); depth 200m. PIate 7 (Verrucocoeloidea brrrtoni Reid): Figure A, lateral view of holotype (taken from Reid. 1965): Fig. 1)); B, hexactine (scale bar same as 'C'); C. pentactine; D. tyloscopule (enlarged tyie showing thorns): E. oxyhexaster; F. irregular tyloscopule; G. uncinate: H. discohexaster (scale bar same as 'E'): 1, photograph of dictyonal framework (arrows indicate lamellae): J. geographic distribution (solid dot denotes holotype and paratype locality).

SUBFAMILY IPHITiNAE (hl. SF.)

Body form varied; commonly with wall-folding and lateral branching; dermalia and çastralia, if present are pentactins: scopules. if present. may be disco-, strongylo- or tyloscopules; uncinates present or absent; dictyonal framework commonly with epirhyses. aporhyses, microspined or spined beams: microscleres as discohexasters andor. oxyhexasters.

IPHITEON BOVVERBANK, 1869A

Iplriteon Bowerbank, 1869a :76; Bowerbank, 1869b: 324; Gray. 1872: 453; Bowerbank. 1875~:274; Schulze, 1887: 306; Ijima, 1927: 166.

Joarrella Schmidt, 1880: 55; Schulze, 1885: 45 1 ; Schulze, 1887: 16; Vosmaer, 1887: 2 16; Delage & Herouard. 1899: 129; Ijirna 1927: 23 1; Van Soest et al.. 1988: 1 1.

Margaritella Schmidt, 1 880: 54; Marshall. 188 1: 1 80; Zittel. 1883 : 1 87; Schulze. 1 885: 45 1 : Schulze. 1887: 15; Schulze, 1904: 180; Schramrnen. 19 12: 358: Ijima. 1927: 1 65; Reiswig. 1 990: 736; Desqueroux-Faundez & Stone, 1 992: 1 58.

TAXONOMIC DECISION FOR SYNONYMY Wheekr. this publication

TYPE SPECIES /phireon punicea Bowerbank. 1869b (holotype by monotypy Bowerbank. 1869b: 323)

BASIS FOR DIAGNOSIS Diagnosis based upon reexamination of hoIotype of I. punicea and examination of non- type material of J. compressa Schmidt and M. coeloptychioides Schmidt.

DIAGNOSIS Body form hel-shaped, erect. External surface with large and small pores. as accessory oscula and epirhyses respectivety. intemal surfaces with altemating longitudinal grooves and ridges; srnall pores on ridges presumably aporhyses. Amararhysis-like channelization of body wall; wall appears pieated or folded. Distinct rotulate, or 'spoke-like'. dictyonal framework; meshes triangular. Loose spicules include spined pentactine and hexactine megascleres, regular and irregular discohesasters. small tylohexasters and oxy- and hemioxyhexasters. Sceptrules and uncinates absent.

REMARKS The genus Iphireon is the second oldest genus of the family Euretidae. Bowerbank ( 1 858) first published the generic name in association with Stutchbury's (1 84 1 ) Ductylocalyxpumicea and made reference to a specimen at the Museum of the Jardin des Plantes at Paris. Bowerbank then (1 862b: 8 17) cited I. panicea as the generic type specimen and, in diagnosing DactyIocalyx Stutchbury, synonomyzed ( 1862~:1096) i'phiteon with D~crylod~~.Bowerbank (1 864: Pl. XXV. Fig. 340) later figured "gemmuies" within the skeletal framework of /. panicea. In his great 'Sponge System'. Gray ( 1867) recognized i'hitcor~ and placed it in his farnily Dactylocalycidae. The first generic description of iphiteon came when. on the basis of framework mesh shape. Bowerbank (1 869a: 76) separated lphiteon from Ductylocalyx: "Skeleton silico-fibrous. Fibres solid. cylindrical. Reticulations symmetrical. Areas rotulate. confluent." Bowerbank (1869a) also cited Valenciemes as the author of Iphiteon. declared the *Porto Rican' (1 799) specimen. at the Museum of the Jardin des Plantes, as the type specimen and re-determined two specimens, labelled as I. paniceu collected fiom La Martinique, as Drrc~yloculyxpztrniceu. I. panicea was first described in Bowerbank's (1 869b) second installment of his monograph on "Siliceo-fibrous sponges' . Here. Bowerbank mentioned the stalked, cup-shaped body form, symmetrical framework, triangular meshes. gemmules and two types of hexasters but did not include information regarding dermalia. gastralia. sceptrules or uncinates. Surprisingly, Bowerbank later (1 869b: 324) stated that he 'helieved that the fragment he used to describe I. panicea. given to him 'some years earIier' by Prof. Melville (who procured it from Prof. Valenciennes). came from the specimen labeiled I. panicea Valenciennes (Porto Rico. 1799) at the Museum des Plantes in Paris. in the same publication, Bowerbank erected four new species: I. beatrix (formerly Aphrocallistes beatrix Gray), 1. subglobosa (formerly Dacrylacalyx subglohosa Gray ). i. inga!li (former1 y Dactyfocalyx ingulli Gray) and 1. callocyuthus (formerl y Mylirrsia callocyathtcs Gray). In Iiis report on the Hexactinellida collected by the 'Challenger', Schulze ( 1 8 87) reviewed the historical status of lphitcon but added no new species to the genus and offered no suggestions as to its taxonomie placement. Schulze (1 887) accepted I. panicea as a valid species within Bowerbank's lphiteon (Vaienciennes could not be considered the author as he did not describe the genus), however, he returned Iphireon bearrix Bowerbank back to AphrocaiIistes beati-ix Gray, L subglobosa Bowerbank back to Dactylocaiyx subgiobosu Gray, l. ingalli Bowerbank to Dactylocal' pumicea Gray and 1. callocyarhus Bowerbank back to ibfylitcsiu callocyuthus Gray. ljima (1927) was the most recent to review iphiteon and is responsible for its placement within the farnily Euretidae. He cited Bowerbank (1 869) as the correct author of the genus and described lphifeon as: "An incompletely known genus. Irregularly cup-shaped. large. Dictyonal frarnework with triangular meshes forming confluent rotulate areas: beams weakly tubercled. Only species: I. panicea (Val.) Bwbk.. .. ." Ijirna (1 927) also indicated that the two specimens labeled I. panicea. from La Martinique. were identifiable with D~~tyIocuI~pzrrniceusStutchbury. In Ijima's (1 927) 'Final List'. Iphifeon as misspelled as Iphiton and 'Martinique' was wrongly included as a locality for 1. pcrnicea. Reesamination oFa fragment of the macerated holotype confirmed the presence of the distinct "rotulate" framework and the absence of sceptrules and uncintes within I. pmziceu. however. contrary to Bowerbanks's (1 869a) original description. loose spined pentactines and hexactines were observed. Also. numerous irregular discohexasters were found together with regular discohexasters. oxy- and hemiosyhesasters. A reccnt review of type. and non-type, material of Schmidt's (1 880) JoaneIlu compressa and it4urguriteIlu coeloptychioides. has uncovered strong simi lari ties to [phiteon punicea. Al1 three species have been found to possess: (1 ) fumel-shaped body form: (2) complex amararhysis-like channeIization of the body wall and "pleating" of the wall; (3)rotulate frarneworks; (4) spined pentactine megascleres; (5) oxyhexasters with crooked secondary rays: (6) similar discohexasters and (7) sirnilar irregular discohexasters. The type. and non-type, material of these three genera also differ in several regards: (A) Margari~ella coeloprychioides lacks loose spined hexactine megascleres possessed by both i. panicea and J. compressa; (B) 1. panicea lacks small tylohexasters possessed by J. compressa and 1bL coeiopfychioides:(C) J. compressa possesses numerous spined diactins with four central knobs (perhaps incompietely formed pentactins) not found in M. corioptychioides and I. panicea and (D)small differences in spicule sizes. Due to the incomplete and fragmentary nature of Schmidt's (1 880) type material of J. compressa and M. coeloptychioides, the menature of their body form and external surface remained obscure, and it is pnmarily for this reason that the three genera (Iphiteon, Margariteh and Joaneila) have remained ta~onornicallyseparate. However. exarnination of a complete specimen (Yale Peabody 005 19) of J. compressa suggests that body form and external surfaces may vary significantly within individual specimens. Based upon this evidence. and the above seven points (1-7). it is then reasonable to consider Jouneila and A4argarifella as junior synonyms of Iphireon. Although it is quite possible that the lack of loose hexactine megascleres in M. coeiop~ychioidesis the result of age of the specimen, and the lack of small tylohexasters within Iphiteon is the result of the macerated state of the holotype. there is not adequate evidence to merit specific synonymy. Membership of Iphileon is therefore considered to be 1. punicca Bowerbank, I. coekoptychioides (Schmidt) and I. compressu (Schmidt). On the basis of absence of sceptniles. the current (Ijima. 1927) inclusion of Iphi~eonwithin the order Scopularia and family Euretidae is not well supported. However. until a major systematic revision of the Hexactinosa is conducted, Iphiteon will remain a valid genus within the Euretidae.

DISTRIBUTION Generic Type /phiteou puniceu Bowerbank: collected near Porto Rico.

Other species I. coeZoptychioides (Schmidt); holotype (designated by Schmidt. 1880: 54) collected off Cuba (currently stored dry at MCZ fC6342). I. compressu (Schmidt); lectotype (designated here) collected off Santiago de Cuba (currently stored dry at MC2 6 181 n-1).

See Plate 8, Fig. J for geographic distribution.

Iphiteon panicea Bowerbank (Plate 8. Figs. A-J) Ip/riteorz panicea Bowerbank 1869b: 324, P1.XXI-II; Marshall, 1876: 123; Schulze, 1887: 344; ljima, 1927: 166; Soest & Stentofi, 1988: 1 1 ; Reiswig, 1990: 736. TYPE MATERIAL lphiteon panicea Bowerbank collected at Porto Rico, currently stored in alcohol at MNHN Paris L.B.I.M. No. H.X.6; P3763.

MATERIAL EXAMINED TYPE MATERiAL: /phiteon panicea Bowerbank (see details above)

OTHER MATERIAL: Jourrellu compressa Schmidt, 1880 MCZ 618In-/ J. conpressa Schmidt, 1 880 MCZ 6824 J. conpressa Schmidt, 1 880 Yale Peabody Museum 005 19 A4~u-gurife/lucodoptychioides Schmidt, 1 880 MCZ 6342 hl. codoptychioides Sclim idt, 1 880 USNM 2332 1 M. coelop~chioidesSchmidt, 1880 USNM 0765 1 M. coe/op~ychioidesSchmidt, 1 880 MC2 U-15-21 iii

* al1 the above specimens are considered as I'hireon Bowerbank.

DESCRIPTION Observations regarding size and shape are based upon Bowerbank's (1 869b) original description and examination of a photograph of the holotype. The skeletal framework and spicule descriptions are based primarily upon a small (2cm2) fragment of the holotype whic h is incomplete and washed-out. Several non-type specimens were also examined for spicde information. Data in micrometres (pm) f st.dev.~.

Size and shape: Measurements of body features taken from photograph of holotype. Irreguiar cup or funnel shape (Plate 8, Figs. A, G). Total sponge height 18 cm; diarneter of distal opening 14- 1 8 cm; sponge wall thickness 1 52.5 cm; wal 1 of sponge appears pleated or folded. Comprised of a complex network of branching and anastomosing tubes. External surface with large (2-6 mm) and small(0.5- 1.O mm) pores; large pores presumed as accessory oscula arranged in longitudinal. or weakly helical, rows; small pores may be reduced in size or absent in basal portions of sponge; small pores presumed as epirhyses. Longitudinal grooves (2-8 mm wide) and ndges (0.3-0.9 cm wide) alternate regularly on interna1 or "gastrai" surface; small pores (diametre unknown) on ridges. Determination of derma1 and gastral surfaces requires soft-tissue analysis. Skeletal Framework (Plate 8, Figs. H-1): Dictyonai framework is an irregular. three- dimensional, network of hexactin megascleres arranged in distinct rotulate (Plate 8. Fig. H) or "spoke-like" patterns; meshes (80- 140 prn) triangular. Extemai surfaces of framework generally with smaller, more irregular mesh. Tube, or ridge. wall thickness 1 60-340 Fm; layering and dictyonal strands not visible. True and false nodes (lacking central axial cross; more than 6 rays) are unswollen and microspined; synapticula and ankylosis present; spurs on external surface present. Beams (30 pm) thick, lightly microspined. Complex channelization of body wall amararhysis-like (Reid. 1964). however, adequate description is not possible until the complete holotype is examined: circula gaps (Plate 8- Fig. 1; diametre 630 - 950 pm) in framework presumed as aporyhses or epirhyses.

Loose spicules (Plate 8, Figs. B-F): Spined pentactins (Plate 8: Fig. B) with straight. or slightly bowed tangentid rays (1 345224 pm) are abundant on both intemal and extemal surfaces: smooth knob of vestigial sixth ray present: rays terminate distally in a slight swel hg; proximal ray spined and tenninally rounded. Loose spined hesactins (Plate 8: Fig. D) present but may be absent in older (basal) portions of sponge. Rays (87k61 pm) are straight or slightly curved. strongly spined and terminally rounded. irregular tylo- discohexasters (60 Fm diametre) have 15-25 secondary rays radiating fiom a ball-shaped centnim; lightly microspined secondary rays terrninate distally with a faintly perceptible disk. Regular parenchymal discohexasters (Plate 8. Fig. C; diametre 24+3= pm) having 2-3 curved secondary rays per primary ray are common and small oxy- and hemioxyhexasters (Plate 8, Fig. E: 42tGj3 Pm diametre) with 1-3 weak and irregular secondary rays per primary ray are common.

DISTRIBUTION The type location of Iphikon panicea Bowerbank is reported as Porto Rico: exact location and depth unknown. Locations of non-type material used in revision: I. conlpressu (Schmidt) (lectotype) MC2 618 1 n-1 Cuba, Blake 1880. stn. + V. Deptli 5 18m. sand/mud. Exact location of collection unknown. I. compressa MCZ 6824 South of Porto Rico (1 9O48'47" N: 77'23-W). Blake 1880. Stn. # 22 Depth 473 rn 1. con?pressa Yale Peabody 005 19 Cuba, Blake 1880, Stn. # V. Depth 5 18m. sand/mud. Exact location of collection unknown. I. coeloprychioides (Schmidt) (holotype) MC2 6342 Cuba, Blake 1880 (sta. 96?), Depth 28Sm. Exact location of collection unknown. 1. coelopiychioides Schmidt MC2 U-15-21 iii Blake CIE, No. 164, collected off Guadeloupe (1 S055' N; 6 1 "4 1 'W), Depth 270m 1. coeloptychioides Schmidt USNM 0765 1 Porto Rico, Fish Hawk Stn. No. 6056. 1899. Exact location of collection unknown. Plate 8 (Iphireon panicea Bowerbank): Figure A, 'bird's eye' perspective of holotype showing longitudinal ndges on imer surfaces; B, pentactine; C, discohexaster; D. liexactine; E. hernioxyhexaster; F. irregular discohexaster; G, side view of holotype: H. photograph of 'rotulate' fiamework; 1, photograph of secondary fiamework (arrows indicate gaps in secondary framework - primary, 'rotdate', framework visible (in focus) ~vithin gaps); J, geographic distribution (solid dot denotes locality of holotype; open dots represent type locality of generic members; star denotes locality of non-type matenal).

CHONELASMA SCHULZE, 1887~

C/~onelasmaSchulze. 1 887a:76(in part); Schulze, 1887b:m Lendenkld. 1 89O:3 80; Schrarnmen. 19 12: 2 17; Ijima, 1927: 165; de Laubenfels, 1955: E83; Reid, 1964225: Reid. 1964: xcviii; Mehl, l992:62; Reiswig & Mehl, 1994: 153.

Leptop/rragnreiIa Reid, 1963:226; Reid, 1964: xcix; Reiswig & Mehl. 1994: 1 53

TYPE SPECIES Chonelusmu iurnella Schulze. l887a (in part) (lectotype designated by Reid. 1963)

BASIS OF DIAGNOSIS Diagnosis based upon Reiswig & Mehl's (1 994) thorough revision of the genus C'l?onefusrnuSchulze. 1887a.

DIAGNOSIS Body fom unknown; dictyonal wall composed of three layers. a channelized dermal layer perforated by radial epirhyses, a middle unchannelized layer containing regularly spaced. parallel. transverse larnellae. and a gastral layer that rnay or may not be cliannelized by radial aporhyses. Epirhyses and aporhyses. if present. do not overlap. Loose spiculation of both dermal and gastral surfaces includes large pentactins with the external surfaces of tangential rays densely covered with sharp conical spines and medium-size scopules with slight terminal caps. Parenchymal spicules include Iarge uncinates and microscieres dominated by oxyhexactins.

REMARKS The following historical review is taken fiom Reiswig & Mehl's (1994) revision of the

beenus Chonelasma. Scliulze (1 887a) erected the genus to receive four new species obtained during the historic world circuit of H.M.S- 'Challenger'. He designated no type species, hence C. lumcflu is regarded as sucli by being the first described and was so designated by Ijima (1927). Schulze (1 887b) based his original description of C. lumella on fragmentary specimens fiom three widety separate locations: one hand-sized plate (or more?) from near Kermadec 1s. NE of New Zealand (sta. 170a). a macerated plate and fragments from the vicinity of the Crozet 1s.. Indian Ocean (sta. 148a), and possibly a macerated plate from near Bermuda (sta 56). Original spicule figures and most of the description are attributable to the spicule-bearing fragment fkom sta. 170a. The specimens differed rnarkedly in skeletal framework arrangement, the first-listed having vertical channelized layers on both surfaces (two-sided), with a middle unchannelized layer composed of large. loose. rectangular meshes. Both macerated fragments exhibited a channelized layer on one surface (one-sided), the loose rectangular-mesh region forming the other surface. Schulze (1 887b) suggested that the weak, loosely skeletalized middle layer may have ruptured and spread over the surface in the two macerated fragments. He designated no type specimen and introduced further ambiguity by including the macerated fragments frorn stations 148a and 56 under description of Chonelasma sp. as well. Schulze & Kirkpatrick (1 9 19, 191 1) then descnbed a suite of 20+ fragments obtained by the 'Gauss' Expedition as C. lumellu choanoides. The material. which included moderate-sized basal funnels and curved plates. was uniformly chamelized on only one surface (one-sided) and almost completely macerated. A few spicules were obtained fiom washings of the skeletons and from a small patch of one specimen bearing possibly autochthonous dermal pentactins, but the scopules were suspected to be foreign. The main points of distinction bettveen C. lamella lamellu and C. lamelfachoanoides were growth form, lamellar vs bel-shape respectively, and details of spicule form. In addition to designation of the type species. Ijima (1 927) greatly restricted the genus. He moved two of Schulze's original species, C. harnarlrrn and C. calyx. and a later addition. C. tmcrzrm Schulze. 1899. to Iiis new genus. Heterochone. transferred the remaining original Schulze species. C. doederleinii. to Ptychodesia Schrarnmen. and synonymized C. schulzei Topsent, 1 890 wi th Periphragella lusilanica Topsent. 1 890. He rede fined the genus on the basis of its restricted membership. C. lumella lurnellu. C. lumella chounoides, and presumably C. qimae Topsent. 190 1. as lacking channeli~ationof the framework. In this process. he clearly did not reexarnine type material. Reid (1 963. 1964) reviewed the status of the 'Challenger' and 'Gauss' specimens of C. iurnellrr and others collected by the Danish 'Ingolf Expedition in the North Atlantic. Reid raised C. lamellcr choanoides to species status, C. choanoides Schulze & Kirkpatnck, and used it as the type species of his new craticulariid genus LepiophragrneZfu. In this action. he included the macerated, one-sided. 'Challenger' specimens (stas. 148a & 56) as well as a number of 'Ingolf specimens from stas. 19.2 1. 53. 54. and 78, previously misidentified by Burton (1 928) as Hexacfinellu grimaldii Topsent, in L. choanoides. He described the channelized dermal layer which bears epirhyses as a "dictyonal cortex" and noted the conspicuous skeietal transverse lamellae of the main framework. between which aporhyses occur in a quadratic series in some. but not all. of the 'Ingolf specimens. He bnefly (Reid, 1963) summarized spiculation. presumably from the 'Ingolf material, but did not identie individual specimens as the source of his information. By Reid's action. C. hmehwas restricted to the single two-sided 'Challenger' specimen from sta. 170a. bearing soft tissues, which he designated as lectotype. He redescribed channelization of the two outer surface strata, designated "cortical layers". and related distribution of sofi part anatomy to skeletal features. Unfortunately. in this complex and important series of actions. Reid provided no figures, gave no indication of the basis for this redescription of C. lamella. designated no type specimen for his new species and genus Leprophragmella, did not indicate locations and catalogue numbers of the specimens he examined, and failed to identi@ the individual specimens eshibiting the characters used to define his new genus. He also failed to clearly indicate the specific characters serving to distinguish the two forms. which had originally been considered conspecific. but were now to be regarded as members of different families. In his original description. Schulze (1 887a: 322) stressed the numerous small discohexasters in C. /amella and Ijima (1927) used this information in his generic description. In the same publication. however. Schulze (1 887a: 397) implied that osyhexactins were the most characteristic microsclere. Reiswig and Mehl's (1994) analysis of the lectotype indicates that discohexactins and discohexasters are definitely uncommon. Reiswig & Mehl (1994) also point out that both Schulze (1 887a&b) and Reid (1 964) failed to observe the distinct transverse lamellae of the framework, a major diagnostic feature of C. Iumella.

DISTRIBUTION Chonelasma larnefla Schulze. 1887a: collected from the Kermadec Islands. 39'45-S 1 78"1 1 'W. Depth I 153m. 1874 (H.M.S.,'Challenger' stn. l7Oa). Substrate and depth range: volcanic mud, depth 1 153m. Geographic distribution: (Plate 9, Fig. M) known only fiom the type locality. Kermadec Islands, NE of New Zealand.

Chonelasma lamella Schulze (Plate 9, Figs. A-M)

Chonelasma lamella Schulze, 1887a: 76(in part); Schulze. l887b:32 1 ; Ijima. 1 927: 165; Reid, 1964: xcviii; Reiswig & Mehl, 1994:153.

TYPE MATERIAL Chonclasma iarnella Schulze, 1887a; lectotype (designated by Reid, 1963) collected fiom the Kermadec Islands by the H.M.S. ,'Challenger' (stn. 170a),depth 1 153m. Currently stored wet at BMNH (catalogue number 1887.10.20.135).

DESCRiPTION The following description is taken from Reiswig & Mehl's ( 1994) rcvision of C. lcrrnella Schulze.

Size. shape and texture: The alcohol-preserved specimen (Plate 9, Figs. A. B) is a flat, uneven Plate 9.1 x 12.0 cm in lateral dimensions, 7.7 to 12.5 mm in thickness from the lectotype which is of unknown size and body form. Both surfaces are covered by a tissue membrane supported by spiny pentactins; underlying dicytonal channels penetrate vertically toward the middle of the plate. Stony hard texture. Although fùnctionally differentiated as dermal and gastral. the two surfaces cm not be distinguished on the bais of their intrïnsic features.

Skeletal Framework: Descriptions of skeletal and tissue are based upon detailed examination of a small 1 x 1.5 cm fragment of the lectotype. Dictyonal skeleton of three distinct layers (Plate 9, Fig. K). Middle layer 4.5 mm in thickness. unchannelized. consists of transverse, slightly arched larnellae with 1-38 mm spacing. presumed perpendicular to growth axis; interlarnellar spaces (1 .3810.093) are spanned by elongate fused rays of Iarnellar dictyonalia oriented longitudinally (Plate 9, Fig. L); meshes are basically elongate-rectangular but locally obscured by intercalated small hexactins, fusion of uncinates to bearns, and spur development. Middle layer enclosed on both surfaces by vertically channelized layers (=primary cortices of Reid, 1964) 1.4 - 6.5 mm in thickness. Ttieir cylindrical channels are bounded by a tight, triangular and rectangular dictyonal meshwork and extend to the wide-meshed middle layer where they terminate openly. Channels of one outer layer are aligned with interlarnellar spaces and presumed to be aporhyses (gastral)(0.890+0.23 17diametre); those of the opposing layer without apparent relationship to lamellae are presumed epirhyses (dermal)( 0.890+0.23 17 diametre). Definite determination of frarnework orientation requires analysis of sofi parts. Vertical dictyonal strands of the channelized layers Vary in relation to the longitudinal strands of the rniddle layer; some appear to be continuations of middle layer strands bending towards the adjacent surface but others have no relationship with longitudinal strands. Dictyonal beams are uniformly finely spined throughout the framework.

Loose spicules (Plate 9. Figs. C-J): Dermal and gastral spicuies. Pentactins (Plate 9, Fig. C) of both surfaces are similar. varying greatly in size; size-fiequency analysis indicates a unimodal population; large exarnples coarsely spined on outer faces of tangential rays: tangential rays ( 1 76+46i00,9 1-280 length; 1 5.2k6.2 oo width) terminate rather bluntly with proclined spines on distal 1/5 of ray: proximal ray long (3 84+2O 1 100.63-849 length:

1 6.4+6.0100width), ratio of mean pro.u./tang. ray lengths = 2.18. Scopules (2 1 9k27100 length) (Plate 9, Fig. F) located below dermal and gastral surfaces similar; with 3-4-7 straight, slightly divergent tines tipped by indistinct serrated caps and arising from a conspicuous 4-lobed basal capitulurn; tines and shafi finely thorned; short-headed

(22+4 length), ratio head/total length = 0.074-0.101- 0. i23kO.O 16(25).

Parenchymal spicules: Uncinates (Plate 9, Fig, E)(357 1$7j530 length; 1 8.2+3. 1 102 width) with long barbs slightly spreading from shafi; mostly perpendicular to surfaces. occasionally oblique. Regular mesohexactines (Plate 9. Fig. D) (1 23134~~)not abundant: rays finely thomed throughout; thoms proclined. Microscleres with finely roughened surfaces; predominately oxyhexactins (Plate 9, Fig. G)(59%), hemioxyhexasters (Plate 9. Fig. H)( l6%)(92+ 1Oioo oxy- and hemioxyhexaster diametre), less common onycho- (Plate

9. Fig. J) and discoliexactins (Plate 9, Fig. 1) (65+11 100 onycho- and discohexactin diarnetre) and hemihexasters (total 9%). and irregular forms (triactins. tetractins. pentactins, spiral forms. etc (1 6%)). Soft tissues: Tissues of the specimen are not well preserved. The analysis of thick sections by Reiswig and Mehl (1994) is consistent with Schulze's (1887b) description as corrected by Reid (1 964). Porous membranes supported by pentact megascleres bound both surfaces. They are loosely comected by trabecular strands to the main choanosomal stratum consisting of 1-3 layers of ovoid vesicles, presumed to be remnants of choanomere chambers. The continuous choanosomal stratun is highly convoluted. and laterally fused through the wall of the specimen, to delineate interdigitating cylindrical water canals and branches progressing from the two surfaces. The inhalent canals extending from below the dermal membrane occupy the skeletal channels (epirhyses) of that channelized layer, traverse the unchannelized middle layer and extend through most of the opposite channelized layer to end blindly within the wall between aporhyses. Exhalent canals of the gastral layer follow the opposite course, ending blindly in the tight dictyonal meshes of the opposite channelized layer between epirhyses. This organiwtion is compatible with the interpretation that water flow through the wall of the living specimen is unidirectional from one surface to the other. It is noteworthy that skeletal channelization is concordant with the convolutions of the choanosomal stratum in only the respective initial outer layer: there is no skeletal channelization corresponding to water passages circumscribed by sofi tissues through the middle and terminating skeletal Iayers. Thus bearns of the siliceous fiamework and adherent sofi tissues span across water canals through most of their length. Dimensions of the presumed choanomere cliambers are 64-89-1 13 k 14 pm (25) in cross section and 102-145-187+13 pm (25) in lenptli. Dense tissue strands (cord syncytia) do not occur in the sample investigated.

DISTRIBUTION Generic Type Species Chonelasma iarnella Schulze, collected near Kermadec Isl.: 29" 8's; 1 70" TE.

O ther species C. tloederleinii Schulze. collected in Sagami Bay: 35" O' N: 139" 5'E. C. Vimm Topsent, collected near SaoMiguel-Terceir: 38" 4' S: 26" 5' W. C'. choanoidL.s Schulze & Kirkpatrick, collected near the Antarctic Gauss station: 66" 0' S: 89" 6'E. Plate 9 (C'huneZusrna lameHu Schulze): Figures A&%. two surfkces of the lectotype; C. dermal pentactin; D. mesohexactin; E, uncinate; F, scopule and enlarged scopule head; G, osyhexactin; H, oxyhemihexaster; 1, discohexactin; J, onychoexactin; K. clean skeletons of a wall section in perpendicular views, facial to transverse lamellae (lefi) and perpendicular (longitudinal section) to four transverse lamellae (nght); dermal layer (d). middle layer (m), gastral layer (g); L, junction of demal and middle dictyonal layers with parts of four larnellae viewed in longitudinal section; M. geographic distribution (solid dot denotes type locality; dl figures taken fiom Reiswig & Mehl. 1994). kt- ie r. .. LEFROYELLA THOMSON, f 877

Lefroyella Thomson, 1877: 403; Schmidt, 1880: 2 1 ;Schulze, 1887: 15; Lendenfeld, 1 890: 40 1 : Schrarnmen, 19 12: 2 16; Ijirna. 1927: 165; de Laubenfels, 1936: 186; Reid. 1958b: 14; Reid, 1963: 325; Reid, 1964: Ixi: Van Soest & Stentofi, 1988: 1 1 ; Reiswig, 1990: 735; Desqueroux-Faundez & Stone, 1992: 123.

Sjvingidium Schmidt, 1880: 46; Weitner, 1882: 32; SchuIze, 1887: 15; de Laubenfels. 1936: 186; Reid. 1963: 225; Reid, 1964: Ixi; Van Soest & Stentofi, 1988: 1 1; Reiswig, 1990: 744.

TAXONOMIC DECISION FOR SYNONYMY Ijima. 1927 and Wheeler, this publication.

TYPE SPECIES Lqhyella decora Thomson, 1877 (lectotype designated (Wheeler. this publication)).

BASIS OF DIAGNOSIS Diagnosis based upon Thomson's (1 877) original description and reexarnination of the lectotype. Specimens from Schmidt's (1 880) type series and other available material were also used as a bais for diagnosis.

DIAGNOSIS Body form funnel-like, erect; transverse rows of tubular stumps alternate regularly with broad grooves on external surface; interna1 surface with distinct longitudinal hollow ridges alternate regularly with deep longitudinal grooves. Skeletal hework in three layers; middle channelized layer with elongate-rectangular mesh often internipted by large gaps; external channelized dictyonal laycrs with irregular meshes and abundant synapticula; beams smooth throughout; nodes not swollen, spurs common; channelization as amararhysis: loose spicules include mesohexactins; oxyliexactins: strongyloscopules. uncinates, oxyhexasters, onycho hexasters. and discohexasters.

REMARKS The genus Lefloyella is one of the more poorly understood members of the hexactinosan Hexactinellida. Thomson (1 877) erected the genus to receive two new specimens obtained by the Challenger at station 56 near Bermuda and, although he did not declare a type specimen, he did figure the larger (12 cm tall) hel-like sponge, which is here considered as lectotype. Thomson (1 877) correctly recognized the distinct body fom of his new LefioyeZIa decora Thomson and noted: "The outer surface of the sponge is raised into spiral ridges .. . and under the ridges are irregularly spiral lines of large holes. The interior of the cup presents a remarkable character .. . The inner layer is deeply fluted. thrown into a series of altematins vertical grooves and ndges .. .". Thomson ( 1877) also reported both specimens as "water-wom" and no information on spicules was included. Schmidt (1 880) erected another new genus, Syringidium. for several fragments collected in 1878 and several specimens later collected in 1878/9 from five locations in the Gulf of Mexico. Of this type series, Schmidt (1 880) figured two specimens (Schmidt. 1880: PI. VII. Fig. 4A. B), one large (Pl. V11, Fig. B; MC2 6494) and one small (Pl. VII, Fig. A: likely MCZ U- 14- 1 Oc iv). both assigned to Syringidium ritteli Although Schmidt (1 880) remarked on the strong similarity between his Syringidiirm zitteli and Thomson's LefroyeZla decora, he iiladvisedly separated (1 880) the two genera using the defense that Tliomson's material was washed-out and therefore not the best representative for a new genus. Schmidt (1 880) finally remarked that Ductyloca~ycrispus Schmidt. 1870 is a younger Syringidium zitteli and thus synonomyzed the two genera. In his account of material collected by the Challenger expedition. Schuize ( 1887) included a specimen from station 33 as Lefroyella decorrr Thomson and the two specimens described by Thomson (1 877). Schulze (1 887) however. mistakenly cited the Iieiglit of Thomson's larger specimen as 17 cm taIl (Thomson described it as 12 cm) and erroneously concluded Thomson's larger specimen as lost. Schulze (1887) then accurately described Thomson's (1 877) original specimen (lectotype), still. however. without useful spicule information, as Lefioyella decora Thomson. Schulze (1 887: 424) listed S. ziffeliSchmidt as an "Insuficiently defined Form and remarked that the newer Svi-ii7gidittrn zitteli Schmidt is inclined to refer to Lej*oyeZZadecora Thomson. Ijima (1 927) formalized the name Syringidium riffeliSchmidt as a junior synonym of Lef;oyeZZa &com Thomson and described a new specimen, L. ceramensis Ijima, collected by the Siboga expedition (stations 177,226) from the Ceram and Banda Seas. Ijima (1927) also listed Schmidt's synonymy of Dactylocalyx- crispus Schmidt as questionable. Reid (1 958b) commented on the forrn of L. decora and noticed a sirnilarity to Periphragella Marshall. Using "available material" and two 'Challenger' specimens Reid (1 964) assumed, on the basis of subtle body form differences, L. decora and S. rittefi as two separate species and resurrected Schmidt's Syringïdium. However, Reid (1 964: cxiv. footnote 1) remarked: '' It is possible that more adequate materiai would show both names to refer to one species, if the differences taken here as generically significant are actually due to (a) different individual developments of the lateral outgrowths [tubular stumps], and (b) differences due to age in respect of enclosure of the paragastrai furrows [intemal longitudinal grooves]. The loose spicules seen were sufficiently similar for this to be possible, and the Syringidium showed occasional transverse perforations like those of L. decoru." In their table on "Hexactinellida from deep water in the West Indian region with remarks on their status" Van Soest & Stentoff (1988: 11) accepted Schmidt-s clainis for the likely identity of Drrctyfoculyr crispus. L. decora and S. zitteli thus resulting in their recognition of L. crispa as the valid name for al1 three forms. Reiswig (1990). in his correction of Ijima's (1927) final list, cited two valid species of Lefroyella Thomson (L. &cor-a and L. ceramensis) and noted that Ij ima's questionable synonymy of Dacrylocalyx crispus Schmidt is accepted as intent and therefore should not be in Ijima's Final List. In reviewing the works of Oscar Schmidt, Desqueroux-Faundez & Stone (1 992) listed the type series of Syringidiun~zitreli Schmidt and declared a lectotype (MC2 6494) and syntypes. Recent systematic investigation and reexarnination of pertinent material have proved helpful for understanding both Lefroj7ellcr and Syringidium. Schmidt (1 880) grossly confüsed matters by (1) using information fiom several specimens and fragments to describe Syringidium zitteli and. (2)not recognizing the taxonomie priority of Thomson's (1 877) description. These two points are especially relevant for the specimen. which Schmidt illustrated as Figure B (Schmidt. 1880: Pl. VII.. Fig. B), so close in iikeness to Thomson's original specimen (the lectotype). was recently found to be without spicules thereby leaving little known difference between the two specimens. Similarl y. in his important actions (1964) Reid did not use type material of either L. decora and S. zi~eliand failed to provide the locations and catalogue nurnber for the material he examined thereby rendering his descriptions and actions unusable. It is doubtful whether ijima (1927) examined Schmidt's (1 870) type material of Dac~localyxcrispus and it is likely that he based his questionable synonymy upon Schmidt's ( 1880) casual remark. Esamination of several specimens labeled as Schmidt's (1 870) type series (MC2 8 175. Tray 83)of Dûctylocalyx crispus showed little similarity to Syringidium Schmidt; D. crispus and S. zitrcili are not considered as synonyms. The status of D. crispus remains unresolved. Reexamination of Thomson's original specimen (the lectotype) of L. decora has suggested the presence of oxyhexactins, mesohexactuis, uncinates and distinct six- tined stronglyloscopules with noticeably thomed shafts. Similar strongyloscopules were also found upon reexamination of several specimens of Schmidt's S. iitteti paralectotype material fhrther suggesting similarity between Lefioyella and Syringidium. On the bais of similarity of body fom and overall lack of spicular information of the two specimens originally described as Lcfioyeflu decora Thomson (lectotype) and Syringidium ritteli Schmidt (lectotype) the two species are here considered the same, thereby returning S'ringiditrrn zitteli Schmidt to the status of juniour synonym of Lefroyeila decora Thomson. Until al1 of Schmidt's paralectotypes (1 880) are closely exarnined and described they are here considered as unidentified specimens of Le@oyefla Thomson.

DISTRIBUTION Generic Type Lefioyellu decora Thomson, 1877, collected by the Challenger Expedition (station 56) near Bermuda. Other Species: Lefroyellu cerumensis Ijima, 1927, collected by the Siboga Expedition fiom station 177 (one specimen; 2"24'S, 129O38'E. depth 1633 m) and station 226 (two specimens: 5O36.S. 127"36'E, depth 1595 m) fiom the Ceram and Banda Seas.

Lefroyella decora, Thomson (Plate 10. Figs. A-1)

Lefroj~elladecora Thomson, 1577: 403, Fig. 106; Schmidt, 1880: 2 1 ; Schulze, 1 887: 30 1, PI-LXXXII: Schulze, 1889: 108; Schnmmen, 19 12: 2 16; Ijima, 1927: 165; Topsent. 1928: 20: Reid. 1964: cxiii; Reid, 196911: 489; Reiswig, 1990: 735: Desqueroux-Faundez & Stone. 1992: 9 1.

Not Ductylocalyx crispus(u) Schmidt. 1870: 19; Marshall, 1875: 15 1 : Sollas, 1877: 288; Schmidt. 1880: 47: Schulze, 1877: 346; Ijima. 1927: 206; Reid, 1957: 824; Reiswig, 1990: 742: Desqueroux-Faundez & Stone, 1992: 86.

Not LefroyeIIa crispa Van Soest & Stentoff, 1988: 1 1

Lcfroyella ;itteIii Van Soest & Stentoff, 1988: 1 I Syringidium U'fteIi Schmidt, 1880: 46, PI-VII. Fig- 4; Schmidt. 1879: PI-IV. Fig. 9- 10; Schulze, 1887: 3.17; Schulze, 1899: 4: Ijima. 1937: 3 10; Reid. 1964: TextFig. 3 l c: Reid. l969d: 189: Reiswig. 1990: 745; Desqueroux-Faundez & Stone. 1992: 9 1.

TYPE MATEMAL LL./royefladecora Thomson, 1 877 lectotype collected by the Challenger (station 56) near Bermuda (currently stored dry at BMNH 188% 10.127); depth 1967 m.

MATERIAL EXAMINED TYPE MATERIAL: Lefroyella decora Thomson. 1 8 77 (see details above).

OTHER MATERIAL: .Yyringidium zittcfi Schmidt, lectotype stored dry at MC2 6494. S. rittki Schmidt. paralectotype (MC2 6595), stored dry at BMNH 1939.2.10.15. S. rinlei Schmidt, illustrated (Schmidt, 1880: Pl. VII, Fig. 4A) paralectotype (MCZ U-14- I Oc iv) stored dry at MCZ. S zitteli Schmidt, currently stored dry at MC2 (U-14-lob ii), collected by the Blake (station 134) off Frederickstadt, Santa Cruz. S. zitreli collected by Captain A. Morrell fiom St. Vincent fiom a cable, currently stored wet at BMNH 1955.0 1 .O 1-001 (Wet 28-Wb). S. zitrcli collected by JSL I rnanned submersible, dive #2613 (HBO1) from the southwest Florida shelf.

DESCRIPTION The size. shape and form description is based upon Thomson's original description and Schulze's reexamination of the lectotype. Skeletal and spicule descriptions are based upon reexarnination of a small, severely macerated. fragment (2 cm2) of the lectotype.

Size. shape and form: The dry-preserved and incomplete specimen (Plate 10. Fig. A) is an erect vase-like funnel 12 cm tall. Width of the sponge increases from 2 cm- at the basal stalk-like constriction. to 4 cm at the distal margin. A wide (5 cm) basal plate and large, deep central oscule are present. The interna1 surface with numerous (twenty at distal margin), hollow, longitudinal iidges, 2 - 3 mm in width, which alternate regularly with deep longitudinal grooves of similar width. The longitudinal ridges appear to increase in number from the base to distal margin. The external surface is characterized by distinct oblique or transverse ridges which alternate regularly with broad grooves. The transverse ndges consist of rows of neighboring. tubular stumps. which are fused together. Radially directed tubular stumps (4 - 5 mm in diametre) communicate directly with deep longitudinal grooves of the interna1 surface, thus representing amararhyses. Broad external grooves with nurnerous, irregularly arranged, round or oval openings of variable size, communicate lateraily with the intemal ("gastral") hollow, longitudinal ridges. Unable to functionally determine dermal and gzstral surfaces. This system of ndges and grooves may be the result of plication of the body wall (Ijima, 1927).

Skeletal frarnework: Dictyonal skeleton of three distinct layers. Middle layer consists of

regular mesh basically elongate-rectangular (1 69k47 17 pn width; 520+157i9 Fm length) but locally obscured by intercalated small hexactins (Plate 10, Figs. G1H); 1 - 2 weakly discerni ble dictyonal laminae are present (Plate I O? Fig. G - see arrows); dictyonal strands presumed parallel to growth ais; transverse beams perpendicular to growth avis not regularly arranged; regular nodes with microspined and curved spurs. Middle dictyonal meshwork is irregular and triangular in periphery of large (394 pn diametre: not shown in figures) gaps through the framework, presurned as channels. Middle layer is enclosed on both surfaces by vertically channelized (Plate 10. Figs. H - c = channels; m= middle layer; e= external layer; other extemal layer dissected off; gaps measure 0.8-1 -0 mm diametre) layers 2-5 mm in thickness. Their irregular meshwork is triangular to ovoid and significantly smaller than middle layer; synapticula abundant; regular and false (no axial cross) nodes present. Both external layers without laminae. Dictyonal bems

(3 8t1 I 38 pm thickness) are smooth tliroughout the framework.

Loose spicules (Plate 10. Figs. B-F): Measurement ranges generally observed from fewer tlian five spicules per spicule type. Oxyhexactins (Plate 10, Fip. F) have straight. distally microspined rays (40-50 um length). Mesohexactin (Plate 10, Fig. B. E) rays (140 - 160 um length) straight, or curved, smooth and terminally blunt: distal rays often shortened (80 - 1 00 um length) and proximal rays lengthened ( 130 - 160 um length). Large ( 1 100 - 1300 um length) strongyloscopules (Plate 10, Fig. D) present. Six slightly divergent. microspined and distally bald tines (140 um length) arise from a smooth capitulum; proximally directed thorns on distal half of shafi are prominent; shafi distally pointed. straight or slightly curved. Uncinate barbs (Plate 10, Fig. C) prominent and highly divergent fiom the shafi. DISTRIBUTION Lefioyella decora Thomson (= Syringidium ziifeli Schmidt); lectotype of Syringidium currently stored dry at MC2 6494; 32O8'N, 46O59'W; depth 1966 m. Lefioyella sp. = S. zitteli Schmidt (MC2 6595), possible syntype of Syringidium currently stored dry at BMNH 1939.2.10.15, location of collection uncofirmed as Blake Station 343; 39"45'N, 70°55'W. depth 1340 m. Lefroyelia sp. = S. zifteli Schmidt, illustrated (Schmidt. 1880: Pl. VII, Fig. 4A) syntype of Syringidiurn (MC2 U-14-1 Oc iv) currently stored dry at MCZ. collected by the Blake (station 139) off Mount Eagle, Santa Cm;17"46'N, 64O48'W; depth 400 m. LefroyeUa sp. = S. zitteli Schmidt, currently stored dry at MC2 (U- 14- lob ii), collected by the Blake (station 134) off Frederickstadt, Santa Cruz; 17"37'N, 64O48'E; depth 454 m. Lefroyella sp. = S. zifteii collected by Captain A. Morrell from St- Vincent from a cable. currently stored wet at BMNH 1955.01 -01-001 (Wet 28-III-b), exact depth and location unknown. LcfioyelZu sp. = S. zitteli collected by JSL 1 manned submersible. dive #26 13 (HBOI) from the southwest Florida shelf, 110 m West of Naples; 26"04'N, 84"13-W. depth 328 m. Plate 10 (Lefioyelludecora Thomson): Figure A, lectotype (modified from Thomson. 1 8 77: Fig. 106); B. mesohexactin; C. uncinate (enlarged area showing long divergent barbs); D. strongyIoscopule (the enlarged to show microspines; note barbed shah): E, meshohexactin; F. oxyhexactin; G, photograph of dissected regular intemal frarnework (arrows indicate laminae); H. photogiaph of fragment of fiamework of Iectotype (m= middle layer; e= external layer; c = channel); 1. geographic distribution (solid dot denotes lectotype locality; stars denote locality of unidentified specimens of LefioyeZZa).

MYLIUSfA GRAY, 1859

Myliusia Gray. 1859: 439; Gray. 1860: 497; Gray. 1867: 506; Bowerbank. 1869a: 76; Bowerbank. 1869b: 333; Gray, 1872: 452; MarshaIl & Meyer, 1877: 266: Zittel. 1877: 268; Schmidt, 1880: 54; Schulze. 1887: 87; Ijima, 1903: 25: Ijima, 1937: 165; de Laubenfels. 1955: E82. Fig. 652; Reid, 1963: 224: Reid, 1964: PLX, Fig.; Reiswig, 1990: 735.

TYPE SPECIES Myliusiu caliocyathes Gray, 1 859: 439 (ho lotype by monotypy Gray, 1859: 439).

BASIS OF DIAGNOSIS Diagnosis based upon reexamination of the holotype and Schulze's ( 1 887) and Ij ima's ( 1 927) diagnoses of M. cullocyathes.

DlAGNOSIS (GENERIC) Cup or funnel body fom; complex of branching and anastarnosing tubes; 'diarhyses-like' parieta1 channels extend from interna1 to externat surface; rectangular and rotulate frarnework; mesh triangular or rectangular; bearns with conical spines arranged in transverse rows; swollen nodes regular or multiradiate with tuberculate warts; no spurs : loose spicules include distinct "sword-like" hexactins: pentactins: discohexasters or osyhexasters. No sceptrules or uncinates.

REMARKS iMylirisiu is one of the oldest genera within the Hexactinellida and its history is long and complex. Gray (1 859) erected aisgenus for a single specimen he obtained from Dr. McGee whom had collected it from the West Indies. Although Gray (1859) initially had doubts about recognizing the specimen as a sponge, he did briefly describe M. cullocyathes as a îûnnel-shaped form with irregular, confluent simple and subcylindrical tubes and likened it with the genera ~Mcrulndrewia and Dnctyloca~~u.Gray ( 1 859) did include an iIlustration of the specimen and mentioned that Dr. Bowerbank regarded the spicules of M. cullocyathes as very different from those of Dacryiocalyx ptmicem. Gray later (1 867) confrrmed the identity of M. callocynrhes as a sponge of the family Dactylocalycidae and described the genus as: ". ..conid, cup-shaped, pierced with numerous short tnincated tubes, forming raised folded anastomosing lamina on the lowcr surface." Gray (1867) provisionally synonomyzed Lirhospongia rorva Duchassaing De Fonbressin & Michelotti, 1864 with M. callocyathes and included information regarding three specimens at the British Museum which he felt were similar to M. callocyathes: (A) a variety, or possibly another species of Myliusia fiorn the West Indies; (B) a specimen collected from the West Indies by Mr. Scrivener in 1812 ad(C) a smaller specimen collected by Rev. Guilding fiom St. Vincent in 1840. Under the genus Dacrylocalyx Stutchbury, Thomson (1 868, 1869d) listed D. callocyathes Gray but did not offer any explanation for this rnovement of M. callocyathes. The genus Myliusia was then (1 869a. b) thrown into tasonomic confusion by Bowerbank. Initially. he (1 869a) enoneously cited the type locality of M. caZZocyïzthes as St. Vincent Island and then (1869b) synonomyzed the type specimen of M. callocyathes.

and specimen 'BI (Gray, 1867). as Iphiteon callocyathes. Presumably Bowerbank moved M. cullocyarhes to Iphiteon on the basis of skeletal features but no explanation was offered. He did, however, vaguely describe (1 869b) and illustrate I. callocyathes with a spined. rotdate framework with swollen, tubercled nodes, oxyhexactins and discohexasters. but did not indicate upon which specimen he based his observations. Bowerbank (1 869b) then comrnitted a taxonornic error when he reestablished the narne ~bîjdizt~-iufor Gray's (1 867) specirnen 'A' (Gray* 1867) which he (1 869b) named Myliusia riAgain, Bowerbank (1 869b) presurnably based this taxonornic decision upon skeletal characterïstics. Schmidt (1 870) listed M. callocyathes as a 'possible synonym' of Daciyloccdy-Y crispzrs but was uncertain of its true affinities to Ductyloculyr. Carter (1 873). however. distinguished these two genera on the thorny appearance of its tianiework and the 'tleur- de-lis' shape of the discohexasters. In describing M. gruyii, Carter (1 877) noticed distinct 'lantern-nodes' and remarked on their absence in M. cullocycrrhes. Marshall and Meyer ( 1877) then described a new sponge with 'lantem-node' framework from the Philippines as Myliusia zittelli, and Zittel(1887) placed Myliusia into the family Meandrospong idae. Schmidt ( 1880) intended to narne several specimens, collected by Agassiz from the West Indies. as M. hassleri but recognized the similarity in framework to Marshall & Meyer's (1 877) M. rirreli and thus synonornyzed the two species. Based upon material collected from Cuba and the Barbados, Schmidt then (1 88054) synonomyzed M. callocyathes with Ducryloculp callocyafhzis,and reported scopules, oxyhexasters and discohexasters fiom this new material. In his report on material collected by the 'Challenger'. Schulze ( 1887) reviewed the taxonomie history of Myliusia and reported four specimens of M. callocyathes from Little Ki Island, Timor (East Indies) and St. Thomas in the West Indies. Schulze (1887) redescribed M. callocyuthes with demal and gastral 'sword-like' pentactines and hexactines, floricome-like discohexaster, oxyhexactins and oxyhexasters but did not indicate from which specimen his observations were based. Based upon their distinct 'lantem-node' frarnes. M. gruyii and M. zittelii were transferred by Schulze (1 887) to his new genus Aulocystis. Ijima (1 903) transferred Myliusia fiom the hilyMeandrosponçidae to the family DactylocaIycidae and then finally (1927) to the Euretidae. He (1927) described the genus Mylirisia as "Cup-like with osculated outbuigings on sides. Dictyonal skeleton with beams. showing spinules which tend to be arranged in transverse rows. and with swollen nodes bearing spinulated wart-like prominences. Genotype: M. caiiocyuthus Gray. 1 859." Ijima found four specimens and fragments of M. callocyarhrcs at sta. 25 1 of the 'Siboga' and diagnosed the genus as having oxypentactin dermalia and gastralia. sword-shaped hexactins, holo-, hemi- and monoxyhexasters and floricome-like discohexasters but in his description of M. crillocyurhus. Ijima clearly stated that no trace of the loose spicules were preseved in the 'Si boga' specimens. Ij ima ( 1 927) described a new species, M. verrzrcosa, fiom the East Indies and. in his final list. included a total of four species within the genus: (1 ) M. callocyathes Gray; (2)M. verrucosa Ij ima: (3)M. conicu (Schmidt), which Ijima (1 927) renamed from Scleropkgma conicum Schmidt collected from the West Indies and (4) a questionable species, M. subglobosu (Gray). formerly Dactylocal' sztbgloboszrs Gray, possibly fiom Malacca. Ijima (1 927: 21 5) also synonomyzed Thomson's ( 1868) and Schmidt's (1 880) DUC~YIOCUZJTcallocyathzrs back to M. culocyathtis, however. he did not include D. crisprrs Schmidt in ~Myliztsia.Mylisia was used by de Laubenfels (1955) as the type genus for the new family Myliusiidae however. Burton (1959) and Reid (1963) rejected this fmily and moved Mfiiusiu back to the Euretidae. In examining the dictyonal frameworks of several euretid genera. Reid (1 963. 1964) noted that M. cullocyathes had "short, irregularly expanding. lateral outgrowths whose margins unite locally so that lateral views show a network of anastomosing margins", however, he made no significant taxonomie decisions. Of the three other specimens that Gray (1 867) mentioned as similar to M. callocyurhes, only specimen C was not dealt with and its current location and identity are unknown. Similarly, the type specimen of Lithospongia forva Duchassaing De Fonbressin & Michelotti, 1864 is considered lost (Schulze. 1887) and the information contained within the original description and illustrations is minimal. therefore. until the type specimen is rediscovered. L. rotva can not be considered synonymous to M. ccrlZoqvnihcs. as Gray (1 867) had indicated. Schulze ( 1 887) and Ijima ( 1927) did not acknowledge Schmidt's (1 870) provisional synonymy of D. crispzrs with M. ca/focyurhc.s and a preliminary investigation of the type material of D.crispus confirms their decision. In his diagnosis of M. callocyathus, Ijima (1 927: 2 15) was most likely reiterating the spicule compliment of Schulze's specimens and not the 'Siboga' specimens. thereby confirming his remark on the lack of spicules in the 'Siboga' material. Reexarnination of the holotype confirmed the ' fleur-de-lis' discohexasters. as noted by Carter ( 1873), however, similady-sized, straight-rayed discohexasters and 'sword-like' l~exactinswere aiso observed. No sceptrules, uncinates or oxyhexasters were found. The lack of taxonomically important sceptrules within M. callocyaihes suggests that this genus be removed from the Euretidae, however. until a systematic analysis cm be performed upon al1 members of the Euretidae Myliusia should remain within the Euretidae. Furthermore, a preliminary investigation (Reiswig and Wheeler) of numerous (20) specimens previously identified as M. callocyurhes and D~~rylo~dp-cullocyufhes suggests that discohexasters are generally the sole rnicrosclere present and oxyhexasters are rarely found with discohexasters. Both Schulze (1887) and Ijima (1927) did not examine Gray's holotype of M. callocyuthes, and most likely did not recognize this taxonomically relevant feature. Because Schulze (1887) did not indicate from which specimen(s?) he based his observations. we can not be sure which specimens possess oxyhexasters.

DISTRiBUTION Generic Tv~e: MyZiusia callocyuthes Gray, 1859 collected from the West Indies. Exact type locality and depth presently unknown. (see Plate 1 1, Fig. J for geographic distribution)

Other species: A4 subglobosus Gray, cotlected from Malacca (?) 1M. vet-rucosa Ijima collected at Station 97 from the Siboga expedition near Pearl Bank in the Sulu Archipelago. iM. conica Schmidt collected by the Blake Expedition at Morro Light. W. Indies.

Myliusia callocyathes Gray (Plate 1 1, Figs. A-J)

Dactyiucafyx ca/locyuthes(us) Thomson. 1 868: 1 19; Thomson, 1869: 7 13 ; Schmidt. 1880: 54, Pl. VIII.; Schuize. 1887: 353; Ijima, 1927: 2 15; Desquerom-Faundez & Stone, 1992: 85

Not Dactylocalyx crispus(a) Schmidt, 1 870: 19; Marshall, 1875 : 1 5 1 ; Sollas, 1877: 288; Schmidt, 1 880: 47; Schuize, 1877: 346; Ijima, 1927: 206; Reid, 1957: 824; Reiswig, 1990: 742; Desqueroux-Faundez & Stone, 1992: 86.

Iphiteon callocyafhes(us) Bowerbank, 1869a: 75; Bowerbank, 1869b: 333. Pl. XXIII; Gray, 1872: 453; Carter, 1873: 358: Carter, 1876: 465; Schulze. 1887: 345: Ijima. 1927: 215.

Not Lithospongiu forwa Duch. & Mich. 1864: 65, Pl. XII: 3,4; Gray. 1867: 506: Soest et al. 1983: 196.

Myliusia callocyat/ies(us) Gray. 1859: 439. PI. XVI; Gray, 1860: 497; Gray. 1867: 506: Bowerbank. l869a: 76; Bowerbank, 1869b: 333, PI-XXIII. Fig. 4-7; Schmidt. 1870: 19; Carter, 1873 : 3 58, PI. XIII, Fig. 10; Marshall, 1875: 15 1 : Carter, 1876: 465; Marshall, 1876: 123; Carter, 1877: 127; Schulze, 1887: 354, PI-CIII; Ijima. 1927: 2 14; de Laubenfels, 1955: E82, Fig. 65:2; Reid, 1963: 224; Reid. 1964: P1.X.

TYPE MATERIAL hfyliusia cnllocyuthes Gray collected in the West Indies and is currently stored dry at BMNH 1988.6.27.1- Exact location and depth of collection unknown.

MATERIAL EXAMINED TYPE MATERIAL: Myliusiu callocyathes Gray (see above for details).

OTHER MATERIAL Mylizrsia callocyathes Gray St. Vincent, USNM 994 W-Indies M. ccrllocyathes Barbados USNM 5423 A4 cnllocyathes St-Vincent, W-Indies IMCZ 6358 M. cullocyat hes Barbados BMNH 1 886.6.14.3

DESCRIPTION The description of the body fom is based primarily upon observations of the complete holotype (preserved dry). Skeletal ffamework and spicule descriptions are based upon examination of a small (lcm2) fragment (preserved in alcohol) of the holotype. Several fragments of non-type specimens were also examined for supporting evidence. Data in micrornetres (pm)k ~t-dev.~.

Body Fonn (Plate 1 1. Figs. A. D & G): Cup or fume1 shape; diameter of distal margin 9- I O cm; total sponge height 4-5 cm; stalk presumably present but unconfinned. Comprised of a complex network of branching and anastarnosing tubes. Intemal or "paragastral" opening (Plate 1 1, Fig. A) smooth with small(2-5 mm diameter) circular or oblong openings aligned in nearly paralie1 grooves radiating from the center: minute (200-500 pm) pores (aporhyses?) perforate srnooth "paragastral" surfaces; single centrai oscule absent. External or -'dermal" surface (Plate 1 1, Fig. D, G) shows short irreguiarly expanding lateral outgrowths (1 -3 mm thick) whose margins unite locally so that lateral views show a network of anastomosing margins or tubes (accessory oscula). Tubes fom diarhyses-like external openings (Ijima (1 927) does not consider these openings as diarhyses) of parietal channels which extend to intemal swface. From external morphology alone, it is not possible to determine gastral and dermal surfaces.

Skeletal Features (Plate 1 1. Figs. H, 1): Dictyonal framework is a regular and irregular. three-dimensional. network of hexactin megascleres. Externai surfaces of framework show irregular rotulate (spoke-like) patterns (Plate 1 1. Fig. 1); mesh is triangular (Fig. )O: no spurs observed on external framework surfaces. Intemally. the framework is slightly more regular; interna1 framework layered (5-8 layers) and arranged in 'tree-ring' like larninae (Plate I 1. Fig. H - see arrows; 305 pm apart) which are ofien oriented parallel to crowth margin; meshes generally rectangular (width 2O9S7 Fm: length M5+301 pm) C or triangular; smal l pores on "paragastral" surface observed as sl ightl y larger meshes (200- 400 pm diametre) in intemal fiamework (aporhyses?) but it is not possible to determine if they open directly to the external (lateral) surfaces. Tube, or margin wall thickness 1-3 mm; 5-7 fiarnework layers distinguishable; dictyonal strands present (Plate 11. Fig. H). Slightly swollen nodes regular or multiradiate (greater than six rays) and ornamented with broad, tuberculate warts (Plate 1 1, Fig. 1). Beams (50+_22jz w thick) distinctly beset with small (1-3 pm). conical spines which are arranged into transverse rows (Plate 1 1, Fig. 1).

Loose Spicules: Microspined hexactins (Plate 1 1, Fig. C) with equal length straight or slightly curved, terminaily rounded, rays (93GSiO0pn length). are abundant on extemal surfaces. Microspined, "sword-like" hexactins (Plate 11, Fig. B) with reduced distal ray (68fXIoo) and significantly elongated, curved and terminaily pointed, proximal ray (26O+l 8OiO0)are abundant on intemal and extemal surfaces. Tangentid rays are straight or slightly curved, terminatly rounded and are oriented paratangentially on internai surface; distal ray of hexactin is freely projecting. Similar pentactins with reduced. knob- like, distal rays and elongated proximal rays are also cornmon. Microscleres as discohexasters (84+26ioo(40-1 60) prn diametre; size-frequency histogram suggests a bi- modal distribution): 1) 5-8 curved secondary rays arranged in a "fleur-de-lis" form (Plate 1 1. Fig. E) with long (8-13 pm) primary rays; 2) discohexasters with 5-8 straight (Plate 1 1. Fig. F) secondary rays and short (3-8 pm) primary rays are common: 3) irregular discohexasters rare. Discohexasters are commonly attached to the tieely projecting radial ray of the "sword-1 ike" hexactin.

DISTRIBUTION Distribution of lM. callocyafhes Author Date # Expedition Location Dcpth Specimens Gray 1859 1 West Indies HOLOTYPE Gray 1867 Il3 West Indies Schulze* 1887 2? West Indies Challenger Stn. # 24 Schuke* 1887 1 Little Ki Isl. Chal. Stn. # 192

Schulze* 1887 1 Banda Island Chal. Stn.# 194

Ijima* 1927 4 Kei Islands Siboga Stn. # 25 1

* unconfirrned identity p - Gray's (1867) specimen B (see Remarks) Distri bution of other s~ecies M szrbglobosus Gray, collected from Malacca (?) M. serrucosa Ijima collected at Siboga sta. 97 near Pearl Bank in the Sulu Archipelago (SO48'7"N.T 1 19O49'E). M. conica Schmidt coilected Morro Light, W. Indies; (23" 1 1 '4"N., 82O23'W) Plate 1 1 (Myliusia callocyathes Gray): Figure A, 'bird's eye' view of holotype; B, 'sword-like' hexactin; C, regular hexactin; D. lateral view of holotype; E. 'fleur-de-lis' discohexaster: F, discohexaster with straight secondary rays; G, underside of holotype (bright white area is plaster-of-paris previously used for display); H, interna1 portion of dictyonal frarnework, extemal surfaces dissected off (arrows indicate laminae); 1, photograph of rotulate fiamework of external surfaces; J, geographical distribution (solid dot denotes holotype location; star denotes locality of specimen B (see remarks); open dots denote locality of unconfïmed specimens of M. callocyathes; open squares denote locality of other species of Myliusia).

PERIPHRAGELLA MARSHALL, 1875

Peripfiragelh Marshall. 1875: 177: Zittel, 1883: 187; Carter 1885394: Lendenfeld. t 886: 579: Schulze, 1887: 299; Ijima. 1927: 165: Okada. 1932: 50; Reid. 1963: 224: Reid, 1964: ix,.uc: Mehl, 1992: 70; Rigby & Mohanti, 1993: 919.

Proeurete t Schrammen, 190221 (in part) Reid, 1963: 224.

TYPE SPECIES Pet-iphrugeila elisae Marshall. 1875: 177 (holotype by monotypy).

BASIS OF DIAGNOSIS Diagnosis based upon reexamination of the holotype of P. elisae, Marshall's ( 1 875) original description and Ijima's (1927) generic diagnosis of Periphragella.

Dl AGNOSIS Fumel-shaped body of anastamosing tubes; regular, layered, rectangular dictyonal frarnework; dictyonal strands prominent; beams uniformly microspined; megascleres as pentactins and four-tined scopules; microscleres as oxyhexasters and discohexasters. Uncinates may be present.

REMARKS Pcriphrugella Marshall (1 875) is one of the more robust genera within the family Eurtetidae. Marshall (1 875) erected the genus for one wetl-preserved specimen. given to him by Dr. Reinwardt, apparently collected in the Moluccas. Marshall (1 875) accurately described and illustrated this specimen of P. elisae as a club-shaped form of anastomosing tubes with regular, rectangular mesh, gastral and derrnal pentactins. "broom fork" scopules and "rosettes" (oxyhexasters and discohexasters). Zittel (1 883), in his classification of the hexactinellids. included Periphragella in the family Meandrospongidae. Using Marshall's (1 875) original description and slide preparations from the holotype of P. elisae, Carter (1885) confidently idenîified a specimen fiom Tokyo Bay as P. elisae, but was vague in his description of the rnicroscleres and did not include any illustrations. Carter (1 885) also observed "barbula", or uncinates in his specimen of P. elisae, contrary to Marshail's holotype description. Lendenfeld (1 886) moved Periphragella from the family Meandrospongidae to the family Euretidne and Schulze (1887) confirmed this action. Schulze (1887) also noted that arnong the hexactinellids obtained in Japan by Dr. Doderlein there were three specimens of P. elisae. Schluze (1887) descnbed and illustrated these specimens as cup or funnel-shaped and with pentactins, scopules, discohexasters, oxyhexasters and uncinates. In his report on the hexactinelllids collected by the 'Siboga', Ijima (1927) reviewed the genus Periphragella and made several important taxonomie decisions. He (1927): (1) named two srnall specimens, that Schulze (1887) had originally described but left unidentified, as Periphragella challengeri; (2) descnbed two new specimens from the 'Si boga' collection as P. purva Ijima, and P. irreguiaris Ijima; (3) renamed Carter's ( 1 885) and Schulze's ( 1 887) specimens of P. elisae as Periphragella elisae japonica for subtle differences from the Moluccan type in microsclere tips and scopule tine morphology: and (4) included Topsent's P. lusitaniea. Ijima (1 927: 164) diagnosed the cenus as: "Cup-like or vase-like with simple or anastomosing tubo-branches on sides. C Oxyhexaster, commonly with disco hexaster. Genotype: P. elisrre Mars hall, 1 875". Okada (1932) later reported a nearly complete specimen of P. elisue. collected by the 'Albatross'. from Sagami Bay and mentioned that his specimen differed from Marshall's (1 875) and Schulze's (1 887) in microsclere rnorphology. Reid (1 963) noted that the genus Periphragella resem bled Verrtrcocoelia Etallon, a fossil genus. and listed another fossil genus. Proeurete Schrammen, as a junior synonym of Periphragc.llu. Reid (1 964). using material from his own collections, also described and photographed the frarnework of P. elisae and coined the terrn 'euretoid' for this type of Iiexactinosan framework. In her dissertation of Mesozoic hexactinellids. Mehl (1 992) mentioned and illustrated an unconfirmed specimen (BMNH 25.1 1.1.978) of P. disae and included the cenus Periphrugeliu as a member of her "tyloscopule" group. Lastly, Rigby and Mohanti CI (1 993) agreed with Reid (1 963) and suggested the name Periphragellu Marshall be iised for extant forms while Verrircocoeiiu Etallon be used for fossil forms. Reexamination of a fragment of the holotype of P. elisae generally contirmcd Marstiall's (1875) description. however. owing to limits in inicroscopy at the time of Marsliall's description. he rnost likely did not notice the rare 'uncinate-like' scopule shafis. Similarly. Marshall (1 875) did not properly distinguish the commonly observed oxy- and discohexasters. The absence of complete uncinates in the holotype suggests that, in fact, uncinates are not a constituent spicule of P. elisae and it is likely that both Carter ( 1 885) and Schulze ( 1 887) mistakenly identified the 'uncinate-like' scopule shafis as uncinates. The genus PeriphrageffaMarshall is thus considered a valid member within the Euretidae.

DISTRIBUTION Generic Tvpe: Periphragella elisae Marshall. 1875: collected from the Moluccas; exact location and depth unknown. (see Plate 12. Fig. 1)

Generic Species: P. chullengeri Ijima collected off Little Ki Island by Challenger expedition. station 192; depth 256. P. elisae japonica Ijima. collected from Tokyo Bay and Enoshima, Japan. P. irregufuris Ijirna, seven specimens collected by the 'Siboga'. P. hisitanica (Topsent) collected near the Azores. P. puma Ijima. six specimens collected near Great Kei Island.

Periphragella elisae, Marshall (Plate 12. Figs- A-1)

Periphragelfa efisae Marshall, 1875: 177. PI-XII. Fig. 8, Pl-XIII. Fig. 24-25. Pl-XIX, Fig. 26; Carter. 1885: 393: Schulze. 1887: 394; Ijima. 1927: 165: Topsent. 1928: 297: Okada. 1932: 50: Reid . 1963: 224; Reid. 1964: 1xx.u. Fig. 43; Mehl. 1992: 70: Rigby & Mohanti. 1993: 9 19.

TYPE MATERIAL Pet-iphragelh elisae Marshall, 1875: holotype collected from the bloluccas: currently stored dry at the Natural History Museum in Leiden (RMNH Porifera nr 28).

MATERIAL EXAMINED TYPE MATERIAL: Per-iphrugelIa elisae Marshall, 1875 (details above).

OTHER MATERIAL: Po-iphruge /la elisae Marshall, sensu Okada, 1932: USNM 22 13 5 collected by the Albatros Stn. # 5088 in Sagami Bay near Jôgashima; Depth 675 m. Periphragella elisue Marshall, MC2 662 1; no information on collection location Periphragella elisae Marshall, MC2 9027; erroneously labeled; no information on collection location. DESCRIPTION The shape, size and texture description is based upon reexamination of the holotype. Skeletal and spicule descriptions are based upon examination of a small fragment (2 cm') of the type specimen, Several fragments of non-type material, and Marshall's ( 1875) description, were used for supporting evidence. Data in micrometres (pm) k st-dev.~. (range).

Shape. size and texture: The dry-preserved damaged specimen (Plate 1 2. Fis. A. F) is an erect fume1 of anastamosing tubes 14 cm tall, 5 cm diametre with a central atrium 10.5 cm in length and 3 cm in diametre at its oscutar edge. This light gray specimen is secured to the substrate by an irregularly shaped, thick and dense basal plate that rnay or may not be part of the same specimen (although it does appear to be hexactinellid-like). Texture is rough and brittle. The intemal surface is smooth with numerous circular. or oval. apertures, which lead laterally to the external surface, thereby forming channels. Basally. the extemal surface shows small (0.5 - 1.5 cm length) tubes with circular apertures (0.2 - 0.G cm diametre). Tubes lengthen and apertures become irregular in shape on distal portions of the extemal surface. This system of anastarnosing tubes. which passes out Iaterally from the central atrium, increases in thickness distally, from one to four centirnetres- Unable to determine gastral and dermal surfaces from morphological information alone.

Skeletal Framework: Dictyonal framework is a regular three-dimensional network of hesactin rnegascleres; framework layered (3-6 layers); interna1 layers (Plate 12. Figs. G. H) with regular mesh basically elongate rectangular (936k289 um length; 4 l8+lO& um width) but may be locally obscured by intercalated small hexactins: external layers less regular than intemal layers, synapticulation prominent in extemal layers; wall thickness 1.0-2.5 mm. thickness decreases from the base (2.5 mm) to the distal margins (1 .O mm): dictyonal strands prominent. orientation parallel to axis of growth. sometimes with slight lateral spreading; beams connecting dictyonal strmds are arranged to fonn successive transverse lamellae thus misleadingly giving a "farreoid" appearance (Reid. 1964: Ixxx. Fig. 43; Plate 12, Fig. G); dictyonal nodes are smooth; finger-like. spined, spurs commonly present on nodes of external frame surfaces; dictyonal beams (65+1 757 um thick; Plate 12, Fig. H) are unifonnly microspined throughout and are forrned by ankylosis and synapticulation.

Loose spicules: Pentactins (Plate 12. Fig. B) on both intemal and external surfaces are similar. varying in size; size frequency distribution indicates a unimodal population.

Tangentid rays (1 8 1- 1 j7 um length) are uniformly microspined. bowed and terminally swollen; the longer (266k555ium length) straight, or curved. proximal ray is also terrninally swollen. Vestigial sixth ray as knob-like swelling. Tyloscopules (3 75+ 1 05 105 um overal length: unimodal size distribution; Plate 12, Fig. C) regularly on both surfaces. Four slightly divergent, straight, microspined, tines (6711 71~~um length) terminate distally as a club, which is heavily beset with numerous minute thorns. Shaft is commonly smooth, rarely uncinate-like or spined, and terminates proximally with a distinct swelling. Microscleres as oxyhexasters and discohexasters. Oxyhexasters (44k7106um diametre; Plate 12, Fig. D) with 2-4 sharply pointed, straight, secondary rays and long (6-8 urn) pnrnaq rays are comrnon. Discohexasters (22+4i19 um diarnetre; size- frequency distribution suggests a unimodal population; Plate 12. Fig. E) have short (4+ 1 50 um) primary rays which terminate distally in a slight swelling that gives rise to 5-7 short. curved or straight, secondary rays. Secondary rays are tipped with minute serrated disks that appear spherical at low rnagnifications. Discohexasters are abundant.

DISTRIBUTION Distribution of generic tvDe s~ecies Author Date # Expedition Location Dept h Specimens Marshall 1875 1 Dr. Reinwardt Moluccas ? Carter 1885 l(?) Dr. J. Anderson Entrance of Tokyo ? Bay (Yedo) Schulze 1887 3 Mr. Doederlein Enoshima, Japan 146- 366 rn Schulze 1887 2 Challenger Stn. # 192 j049'S, 132O 14'E 256 m Okada 1932 1 Albatross Stn. # 5088 Sagami Bay 675 rn Distribution of other s~ecies P. challengeri Ijima, coilected by the 'Challenger7,station 192; depth 256. P. disuc.japonica Ijima, exact location unknown; depth 146-366m. P. irregrr1ari.s fjirna, seven specimens (0°29'S, 130°S'E); depth 469 m. P. /z&unica (Topsent) collected by the 'Hirondelle' (39'22'N. 33O44.W): depth 1384 m. P. purvu Ijima, six specimens (5O54'S, 1 3S056'E); depth 984 m. Plate 12 (Periphragelia eiisue Marshall): Figure A. lateral view of holotype; B. prntactine; C. tyloscopule; D. oxyhexaster; E. discohexaster; F. lateral view of holotype (central atrium visible); G, dictyonal hnework (external layers dissected off to show regular intemal framework; arrows indicate larnellae); H, higher magnification of regular dictyonal framework (external layers dissected O fQ: 1. geographic distri bution (solid dot denotes holotype locality, open dots denote location of other specimens of P. elisoe. stars denote other species of Periphrugelh).

PTYCHODESIA SCHRAMMEN, 1912

Pfychodesia Schrammen, 1912: 252: Zittel, 1915: 75; Ijima, 1927: 165; Topsent. 1928: 1 : Topsent, 1928: 302; Reid, 1958b: 11 ; Reid, 1962: 74 1 ; Reid. 1963: 225; Reid. 1964.

Tretoclione Reid, 1958b: 1 1; Reid, 1962: 741 ; Reid, 1963: 225; Reid, 1964: ciii, Fig. 52.

TAXONOMIC DECISION FOR SYNONYMY Wheeler, this publication

TYPE SPECIES P~ychodesiapapillutu f Schrammen, 1912 (by monotypy).

TYPE SPECIES FOR MODERN REPRESENTATIVES Ptychodesia duplicata Topsent. 1928 (ho ioty pe by monotyp y).

BASIS OF DIAGNOSIS Diagnosis is based primarily upon Topsent's original description and reexamination of type rnaterial of Ptychodesia duplicata Topsent, 1928. Non-type rnaterial (MC2 U-8-C) considered identical to the holotype of P. duplicata and information fiom Ijima (1927) and Reid (1 962; 1964) was also used for the diagnosis.

DIAGNOSIS Body form funnel-like, erect: finger-like papillae and transverse ridges on internal surface; longitudinal grooves on external surface internal and extemal surfaces anatomically considered inverted; dictyonai fiamework in three layers; middle channelized layer with regular rneshes; extemal layers with irregular rneshes; bearns microspined tiiroughout; nodes with greater than six rays cornrnon. Amararhytic. epirhytic and aporhytic channelization. Loose spicules as pinulated hezcactins. oxyhexactins. pentactins. scopules. uncinates and discohexasters.

REMARKS Schrammen ( 19 12) erected this poorly known genus for the distinct plate-like body form with perforated papiilae and transverse ridges possessed by the fossil sponge, Pfychodesia pupi~laru,from the Cretaceous period. Schrammen also described (19 12) distinct radial canals extending throughout the wall of P. papillata as well as epirhyses and aporhyses. As an aside, Schrammen (1 9 12) mentioned having observed an extant member of the genus Ptychodesia collected from Japan that Ijirna had intended (1904). but never described as Hexactinellidu lorica (a nomen nudum). Ijima (1 927) redefined the genus Prychodesia, using as the recent type specimen the extant Chonelasma doederleinii Schulze (= P. doederleinii), as having pinular hexactin dermalia and gastralia discohexasters and distinct body channelization which Ijima temed amararhysis. Ijirna (1 927) did not redescribe or illustrate P. doederleinii and it must be assumed that he based this important taxonomic decision solely upon Schulze's (1 887: 324) original description of Chonelusmu doederleinii. Using Ijima's concept of the genus Ptychodesiu. Topsent (1 928) described a complete but washed-out and long dead specimen given to him by Mr. Rautenfeld. coilected off Misaki, Japan, as Prychodesia duplicata. Topsent described P. duplicata as a conicaf form with papillae and transverse ndges on the concave surface and longitudinal grooves on the convex surface. Topsent (1 928) also suggested that the intemal and extemal surfaces of P. duplicata were inverted with respect to most hexactinosan body forms. Spicules of P. duplicata were sparse but Topsent rnanaged to describe oxyhexactins, pentactins, pinules, uncinates, scopules and discohexasters while noting that contamination was likely. Reid ( 1958b) based a new genus, Tretochone. on Ijima's concept of Ptychodesia but used Topsent's ( 1 928) P. duplicura (= Tretochone duplicata) as the type specimen. thereby leaving the genus Prychodesia Schrammen monospecific with P. papillacu. Reid later (1962: 739) explained that his euretid genus Tretochone differed from Pydzodesicr (sensu Schrammen) by: (1) the absence of radial outgrowths involving the wliolc laterril wall. (2) the presence of amararhysis, and of purely dermal outgrowths related to amararhysis. and (3) the absence of distinct epirhyses and aporhyses. Reid went on ( 1963: 740) to explain that Chonelasma doederleinii Schulze, as Schulze (1 887) had originally descrïbed it, did not possess the characteristic amararhysis of the genus Ptychodesia and suggested that Ijima (1 927) had described a sponge significantly different to Schulze's C. doederkinii. With great detail, Reid (1964: cvii) descnbed the com plex amararhytic c hannel ization of Tretochone duplicata (= Pfychodesia duplicaru Topsent) but made no important taxonomic decisions concerning P. duplicata in this. his last word on the subject. Presurnably, Reid (1 958. 1962, 1964) did not examine Topsent's original P. dupticuta and based his important taxonomic actions on personal material collected fiom Sagami Sea, Japan (1 962: 740). A review of Schulze's (1 887) description and illustrations of Chonelasma doederleinii confirmed Reid's (1 962) observations and indeed suggests that C doederleinii is not in possession of the characteristic papillae and transverse ndges of Pfychodesia duplicara Topsent; the taxonomic status of C. doederleinii remains unresolved. A recent review of Topsent's (1 928) description and reexamination of a fragment of the holotype of Ptychodesia duplicata confmed: (1) the presence of amararhytic channelization and (2) the presence of epirhyses and aporhyses. As this author could not comprehend Reid's (1962) "radial outgrowths involving the whole lateral wall" their presence in the holotype material could not be confirmed. Since Reid (1958) used the absence of epirhyses and aporhyses as a significant distinguishing feature of iiis TI-srochone.the discovery of both epirhyses and aporhyses in the holotype of P. dtcpticara invalidates Tretochonc. P. dtiplicutu Topsent is thus considered the extant type representative of Ptychodesia Schrammen. The presence of pinular hexactins, pentactins. scopules and oxyhexasters as reported by Topsent (1 928) was not confirmed upon reexamination of holotype material. however. a preliminary examination of non-type material (BMNH 1925.1 1 .O 1 -681 ). nearly identical in body form and skeletal structure

DISTRIBUTION Generic modem type s~ecimen Pfychodesia duplicata Topsent. ho10 type col lected off Misaki. Japan. Exact location unknown. Currently stored dry at MNHN Paris P. 4676, Japan No. 3294-5; L.B.I.M. No. H. T. 109. Ptychodesia duplicata, Topsent (Plate 13, Figs. A-P)

Ptychodesia duplicata Topsent, 1928: 302, PI.11. Fig. 1-6, P1.111, Fig. 3; Topsent. 1 928: 1 ; Reid, 1%Sb: 11 ;Reid, 1962: 739; Reid. 1964: p.lxi.

Tretochone duplicata Reid, 1958b: 1 1; Reid, 1962: 739; Reid, 1964: liii. Fig. 44.

TYPE MATERIAL Ptychodesia duplicata Topsent, holotype col lected by Mr. Rautenfeld off Misaki. Japan (currently stored dry at MNHN Paris P. 4676: Japan No. 3294-5: L.B.I.M. No. H. T. 109). Exact location and depth unknown.

MATERIAL EXAMINED TYPE MATERIAL: Prychodesia duplicala Topsent (see above for details).

OTHER MATEMAL: Pfychodesia duplicala Topsent, part of Dendy Collection. Location and depth of collection unknown (BMNH 1925.1 1.01-68 1). Plychodesia duplicatu Topsent (labeled as Hexactinellida Zorica (nomen nudum)) as part of the Owston Collection (#0C5233) currently strored dry at MC2 # U-8-C.

DESCRIPTION The size. shape and skeletal descriptions are based upon Topsent's (1 928) description combined with information taken from non-type material (MC2 U-8-C) considered identical to P. duplicala Topsent. Spicule descriptions are based upon reexamination of two srnall fragments (1 -2 cm') of the type specimen of Prychodesia drrplicata and Topsent's ( 1 928) original description.

Size and shape: The dry-preserved intact specimen (Plate 13, Figs. A. J) is an erect. thin (4-5 mm thick) plate (40 cm tall. 50 cm wide) with basal lateral margins rolled together to produce a fumel-like growth fom. Basal horizontal plaque as presumabte method of substrate attachment. Internai, or concave, surface (Plate 13, Fig. J) of funnel with finger- like papillae and numerous paralle1 transverse ridges (13 mm height) (Plate 1 3. Fig. K). Ridges (Plate 13, Fig. N) are irregularly shaped, often hollow, and appear to arise by fusion of numerous neighboring papillae (Reid, 1962: 739). Concave surface (Plate 13, Fig. A) with srna11 (600 - 700 urn diametre) round apertures (Plate 13, Fig. M) presumed as ostia. Ostia arranged in nearly longitudinal and parallel rows. Oscule-like apertures. slightl y larger ( 1000 um diameue) than ostia, are commonly located distally. or laterally. on transverse ridges. External, or convex, surface with two different alternating, longitudinal and parallel rows. Rows with numerous, densely arranged. minute pores ( 150 um diametre) altemate regularly with rows of larger (700 - 900 um diameue) vertically aligned pores that often are not fully developed thus Ieaving distinct. variable Iength, longitudinai grooves (0.8 - 1.1 mm width; Plate 13, Fig. B). Vertically aligned pores and longitudinal grooves presumed as amararhrytic external openings (Plate 13. Figs. B. G). Although functionally determinable as demal and gastral. the external and interna1 surfaces are reversed in cornparison to most hexactinosan body forms so that what anatomically appears to be the gastral surface is on the external surface-

Skeleton and channelization: Dictyonal skeleton of three distinct layers (Plate 13. Fig. N). Middle layer 300 - 350 um in thickness. consists of transverse. slightly arched lamdlae with 0.8 - 1 .O mm spacing, presumed perpendicular to growth asis; inter1amelIa.r spaces are spanned by elongate füsed rays of larnellar dictyonalia oriented longitudinally; meshes are basically elongate-rectangular (0.8 - 1.0 mm length: 0.4 - 0.5 mm width: Plate 13. Fig. 0)and are often obscured. completely or incompletely. by circuiar apertures (Plate 13, Fig. O) presumed as aporhytic and epirhyritic channels; regular (six rayed) nodes often with freely projecting microspined rays (spurs); beams formed by ankylosis (axial canals prorninent) are microspined. Middle layer enclosed on both surfaces by vertically channelized layers 80 - 100 um thickness (Plate 13. Fig. N). Their cylindrical channels are bounded by a tight irregular dictyonal meshwork and extend into the middle layer. External layers with irregular dictyonal framework; meshes triangular; longitudinal dicytonal strands perceptible; dictyonal beams (50 um thick). fonned by ankylosis and synapticulation, are uniformly spined (spines 2 - 4 um height): true and false (greater than six rays) nodes are present.

Spicules: Oxyhexactines (Plate 13, Fig. 1) with straight. or curved. pointed rays (55 - 100 pm length) are heavily covered with fine, long, spines (2-7 pm Iength) are common. Pinnulated hexactins (Plate 13, Fig. H) with straight, microspined and terminally pointed tangential rays (75 pilength); slightly longer (80 pm) and more slender radial ray: distal ray (70 pm length; 23 - 30 pm thick) club-shaped with taIl spines (10 Fm length). Pentactins (Plate 13, Fig. C) with short vestigial knob of distal ray are rare; tangentid rays 80 pm in length; proximal mys 100 pm in length; evenly microspined. Disco- scopules (length unknown; Plate 13, Fig. F) with a straight, microspined, shafi have three, straight, nearly parallel, tines that terminate distally with a small button. Oxyhexasters (diarnetre unknown; Plate 13, Fig. D) with long primary rays and three secondary rays are rare. Uncinates (1-2 mm length; Plate 13, Fig. E) may occur fieely or fused to the dictyonal hework; barbs (15 - 20 pm length) are slightly outwardly cuwed and closely held to the shaft. Orientation and location of spicules wiknown.

DISTRIBUTION Distri bution of holotvpe Ptychodesicr duplicata Topsent. collected off Misaki, Japan. Exact location and depth unknown.

Distribution of other specimens Piychodesia duplicata Topsent collected near Japan MC2 # U-8-C. Plate 1 3 (Ppchodesia duplicata Topsent): Figure A. photograph of modern holotype showing extemal sudace (taken fiom Topsent. 1928: PI.11, Fig. 5); B, higher magnification of extemal surface (arrows indicate longitudinal grooves; photograph of MCZ #U-8-C): C, spined pentactin; D, oxyhexaster; E, uncinate; F, discoscopule; G, diagrammatic representation of body fiom and external (convex) surface (s, slit on estemal surface; taken from Reid, 1964d:text fig. 52e'); H, pinnulated hexactine; 1, microspined oxyhexactine: J. photograph of modem holotype showing internai surface (taken from Topsent, 1928: PHI, Fig. 1); K, photograph of papillae and ridges situated on internal surfaces (taken fiom Topsent, 1928: PlII, Fig. 2); L, diagrammatic representation of body from internal (concave) surface (p= papillae situated on internal surface: taken from Reid, 1964d:text fig. 52e1l); M. higher magnification of intemal surface (apertures presumably ostia; photograph of MC2 #U-8-C): N. photograph of cross-section through ridge (e. external dictyonal framework layer: m. middle dictyonal layer: arrows indicate lamellae oriented nearly perpendicular to dictyonal strands: photograph of MC2 #U-8-C): O. dictyonal framework with external surface dissected to show middle layers and c ircular channels (circular channels, presumably epirhyses or aporhyses. visible: arrows indicate laminae: photograph of MCZ #U-8-C); P. geographic distribution (solid dot denotes holotype locality; open dot denotes locality of specimen (MC2 #U-8-C) believed to be identical to the holotype).

"HETERORETE-GROUP

Those members of the subfamily Iphitedae in possession of spined dictyonai beams and lacking pentactin dermalia and'or gastralia.

HETERORETE DENDY, 1916

Heferorete Dendy, 19 16: 2 14; Hentschel, 1923: 350; ljima. 1927: 165: Laubenfels, 1936: 187; Reid, 1957: 908: Reid. 1958: 17; Reid, 1963: 224; Reid. 1964: xciii; Reiswig, 1990: 736.

TYPE SPECIES Neterorete pulchra Dendy, 19 16 (holotype by monotypy, Dendy, 191 6: 2 14)

BASIS OF DIAGNOSIS Diagnosis based upon Dendy's (1 9 1 6) description of ffeterorcrc.pzilchrum and a fragment of the holotype.

DIAGNOSIS Body form irregular, anastamosing and branching cylindrical tubes; tubes presumably open as oscula; walls of tubes thick and rigid; dictyonal frarnework is an irregular three dimensional network: meshes highly irregular: significant intercalation of oxyhexactins : i ntradictyonal epirhyses and aporhyses; beams heavil y spined: hexactin nodes not swollen: spurs present. Loose spicules: spined oxyhexactins. regular and irregular discohexactins. No dennalia gastralia, scopules or uncinates.

REMARKS Dendy (1 9 16) erected this genus for the single, incomplete, specimen collected by the 'Sealark' fiom the Salomon Atoll, near the Maldive Islands. Dendy (1 9 16) described and i llustrated Heterorete pulchrum as a thick-walled, tubular and branching form with oxyhexactin. discohexaster and oxyhexaster spicules and a solid three-dimensional hexactine framework. Dendy ( 19 16) imrnediately noted the conspicuous absence of pentactine dermalia and gastralia, scopules and uncinates, ofien associated with specimens having similar 'euretoid' frameworks. Dendy also took care to emphasise the well-preserved state of the specimen and noted that the absence of these spicules could not have been the result of physical maceration. Taxonomically, Dendy ( 191 6) placed Hererorete into Ijirna's (1903) family Dactylocalycidae (whose generic members inc1 uded Dactylocalyx. Margari~eila.Myliusia. A uZocaljx and Euryplegma). Hen tsc hel later (1 923) confirmed Dendy's placement of Heterorete within the Dactylocalycidae, however, in his report on the 'Siboga7 sxpedition, Ij ima ( 1 927) partial 1y synonomyzed his Dactylocalycidae with the family Euretidae Schulze, and therefore moved Heterorere into that îàmily. AIthough it is unlikely that he exarnined the holotype, Ijima (1 927) likened Hererorete to his Pararete, noted the lack of derrnalia and gastralia, and commented that the rnicroscleres of ffererorete were probably only discohexasters. Reid (1 958b. 1 963. 1964) accepted Ijima's (1 927) movement and viewed Hererorere as a genus, or subgenus of Ezirefe. within the order Scopularia. which had secondarily lost its sceptniles. Reexamination of a fragment of the holotype confirms the lack of pentactins, sceptrules and uncinates, and suggests that discohexasters are the only microscleres present in Heterorete. It is likely that Dendy (1 9 16) misidentified an incomplete discohexaster as an oxyhexaster. The genus Heterorete differs significantly from Eurete. the type genus of the family Euretidae, in several important aspects: 1) absence of derrnalia and gastralia; 2) absence of sceptmles; 3) thick and solid frarnework: and 4) channeIization of the framework. Clearly. these differences suggest that Heterorete be removed from the farnily Euretidae and order Scopularia. Untit a thorough systematic analysis of the Hexactinosa can be performed, the genus Heterorete is provisionally retained within the family Euretidae.

DISTRIBUTION Hererorere pulchrum Dendy, 19 16; holotype collected by the 'Sealark' expedition in 1 905 from the Salomon Atoll. Indian Ocean. (see Plate 14. Fig. J).

Heterorete pulchrum Dendy. 19 16 (Plate 14, Figs. A-J)

HeteroretepuIcI~raDendy, 1916: 214, Pl. 41, Fig. 11-18; Ijima, 1927: 165; Topsent. 1928: 302; Reiswig, 1990: 736.

TYPE MATERIAL hrerarete pulchrum Dendy; single specimen collected by the "Sealark" in 1905 from the Salomon Atoll. CurrentIy stored in alcohol at BMNH l9îO.l2.09.064.

MATERIAL EXAMINED TYPE MATERIAL: Heterorefe pufchmm Dendy; (see details above).

DESCRIPTION Description of body form based upon Dendy'ç (1 9 16) original description and illustration. Skeletai framework and spicule descriptions based upon reexarnination of a small(1cm X 1 cm X 2mm) fragment of the holotype (preserved in aicohol). Data in micrometres (p)

Body Form: The illustrated (Dendy, 19 16; Pl. 41, Fig. 1 1), incomplete. holotype (Plate 14, Fig. A) is a network of irregular, anastamosing and branching tubes (8 mm extemal dimetre); tube walls 2 mm thick. Tubes presumably open (distally, proximally and laterally) as oscules (4 mm wide). Both inner (gastral) and outer (dermal) surfaces are marked with numerous srnaIl pits, which correspond to epirhyses and aporhyses. Numerous. smalt commensal zoanthids (class Anthozoa) on dermal surfaces and in channels.

Skeletal Framework (Plate 14; Figs. E-1): The dictyonal framework is an irregular. chameiized (Plate 14, Fig. E). thick (2 mm). three-dimensional. network of spined hexactin megascleres. Interna1 portions of primary framework (Plate 14. Fig. F) regular and irregular; meshes rectangular and triangular (75 - 225 pm); gaps in prima^ framework as intradictyonal canalization; beams formed by ankylosis and synapticulation (Reid. 1964); dictyonal strands barely perceptible; true and false (no axiai cross) nodes. Gastral surfaces of fiamework (Plate 14, Fig. G.) primarily irregular; meshes rectangular or triangular (55 - 180 pm): short spurs as freely radiating rays of primary dictyonalia; intercalation of oxyhexactins; gaps (284 - 396 pm diametre) in gastral surfaces as aporhyses. Dermal surfaces of frarnework (Plate 14. Fig. H) highly irregular and dense; meshes smaller (30 - 190 pm) and triangular; bem fusion primarily by synapticulation: significant intercalation of oxyhexactins (Plate 14, Fig. 1) and rarely discohexasters: gaps (241 - 376 prn) in demai framework as epirhyses. Beams (15 - 57 pm thick) heavily spined (spines 2 - 1 1 pm length) throughout; dictyonal nodes not swollen.

Loose spicules (Plate 14, Figs. B-D): No pentactine dermalia or gastrdia, scopules or uncinates. Oxyhexactins (Plate 14, Fig. B) with unequal length (54* 13 10 pn ray length) spiny. straight, or curved, and pointed rays; commonly fused to framework, especially on gastral surfaces. Discohexasters (Plate 14, Fig. C) (84+1250 pm diametre) are abundant: primary rays (8kisopm length) give rise to three to five (mode is four) curved secondary rays which terminate distally in a small serrated disk. Oftentimes one secondary ray may initial1y project proximaily thus forming an 'L' shape. Similarily sized uncornmon irregular discohexasters (Plate 14, Fig. D) have numerous, radially projecting (secondary?), rays that terminate distally in serrated discs; pnmary rays not distinguishable; rays appear to emanate fiom an irregular centrum.

DISTRIBUTION Hetcrorere pulchrum Dendy; holotype collected by the "Sealark" in 1905 from the Salomon Atoll (estimated CO-ordinates:4" 17' S. 72 "02' E - accuracy * 25 kms); Depth 220 - 275 m. Plate 14 (ffetet-oretepulchrum Dendy): Figure A, holotype @ = polyps of commensal anthozoans; modified fiom Dendy, 19 16: PL. 4 1); B, spiny oxyhexactin; C. discohexaster; D. irregular discohexaster; E. cleaned fragment of holotype framework (arrow indicates large gap in framework wall); F, interna1 portion of cleaned and dissected framework; G. gastral surface of cleaned framework; H; derma! surface of cleaned framework: 1. intercalated oxyhexactins on dermal surfaces; J. geographic distribution (solid dot denotes type locality).

GYMNORETE IJIMA, 1927

Gymnorete Ij ima 1927: 165; Topsent. 1928: 337; Reid, 1965: 224; Reiswig. 1990: 736.

TAXONOMIC DECISION FOR SYNONYMY Ijima, 1927: 165

TYPE SPECIES Ezrrete alicei Topsent, 1901b (lectotype designated by Wheeler, this publication)

BASIS OF DIAGNOSIS Diagnosis based upon Topsent's (1901 b) original slide preparations. description of Eurete dicei and Ijima's (1927) concept of the genus.

DIAGNOSIS Body form "ear-shaped"; tubular and anastornosing; reduction or absence of tubes in basal regions: tubes short with circular and open apertures; method of attachent to substrate unknown; skeleton as three-dimensional fusion of hexactins: external dictyonal surfaces with irregular meshes; synapticulation of external dictyonalia: interna1 dictyonal surfaces with rectangular meshes: bearns covered in strong spines throughout: nodes of framework not swollen. Dermalia and gastralia lacking. Loose spicules include scopules. uncinates. oxyhexactins; onychohexasters. tylo- and discohexasters.

REMARKS Ijima (1 927) erected the genus Gymnore~eto include those 'Eurete-like' specimens lacking dermalia and gastralia and used Topsent's (1 90 1b) Euï-ere alicei as type specimen. Topsenfs original (1 90 1b) description of Eurefe alicei was based upon one specimen and several fragments (type series) collected by the 'Princess Alice' at several separate stations (sta. 578. 602 and 1349) from the Archipelago of the Azores. Topsent later (1 904, Pl. IV, Fig. 8) illustrated the larger specimen fond at station 578. declared here as the lectotype. Topsent (1 90 la, 1904) described, and illustrated, Eurete alicei as an 'ear-shaped' and tubular fonn with scopules, oxydiacts, onychasters, and discohexaster spicules. Topsent (1 90 1 b) clearly pointed out that both the lectotype (sta. 578) and fragments (sta. 602) were collected with the soft tissues intact. indicating that the lack of dermalia and gastralia could not have been the result of maceration. As previously stated, Ijima (1 927) erected the genus Gymnorere for those specimens lacking demalia and gastralia. Ijima (1 927) also pointed out that the tubes of Gymnorere alicei are often flattened and obliterated and that Topsent's (190 1 b) onyc hasters were 'hexactinoid' . Ij ima ( 1927: 1 20, 1 66) made reference to Gymnorere i~uriolosurnas"a new 'euretid' genus and species, yet undescribed. from Sagami Sea. Under the sme genus is to be placed the species described by Topsent (1 901b) under the narne Eztrete aZiceT9. Like many other genera within the family Euretidae. Reid (1963: 224) relegated Gymno~*ereto subgeneric status of Eurere, based primarily on skeletal architecture. Reiswig ( IWO) interpreted Ijima's citation of Gymnorere vuriolosum (Ijima. 1927: 164) as a nomen nzrdum (as of 1927) and the citation of Gymnodicrytrrn varioloszrrn as a mistaken reference to Gynznorete variolostrm. Esamination of Topsent's original spicule preparations of' E. alkei generally confinned Topsent's original description. however. when viewed under high n~agnification (oil immersion). Topsent 's ( 1 90 1 b) discohexasters appeared as tyloliesasters. Also. Topsent ( 190 1b. 1904) did not pay much attention to the dictyonal framework. noi did he make any mention of the small oxyhexactins commonly found in Iiis preparations. Although a specimen bearing the label Gymnorere variolosum does exist (BMNH), no description or illustration has been reported, thus supporting Reiswig's nomen nudurn status. For the obvious lack of zoological information. Reid's subgeneric designation of Gymnorere, based solely upon skeletat characters. is rejected- Until the lectotype and paratype material. currently stored at the Oceanographic Museum of Monaco, is made available. or new material is collected. the genus G_vmnorere will remain a poorly understood genus within the family Euretidae.

DISTRIBUTION Gymnorere alicei (Topsent); lectotype collected by the 'Prince Albert' Expedition (sta. 578) in 1895 between Sao Miguel and Terceira of the Azores. Gymnoreie alicei (Topsent): paralectotypes collected near Sao Jorge of the Azores (sta. (See Plate 15, Fig. J for geographic distribution.)

Gymnorefe aiicei (Topsent, 1901 b) (Plate 15' Figs. A-J)

Eurete alicei Topsent, 190 1 b: 462; Topsent, i 904: 45, Pl.IVI Fig. 8, PLVII, Fig. 5: Ijima. 1927: 165; Reiswig, 1990: 74 1.

Gymnorete alicei Ijiaa, 1927: 165; Topsent, 1928: 302; Reiswig, 1990: 742.

TYPE MATERIAL Gymnorere alicei (Topsent, 190 1b) = Eurete alicei Topsent, Iectotype collected by the 'Princess Alice' expedition in the Azores; currently stored at the Oceanographic Museum of Monaco (04 0339).

MATERIAL EXAMINED TYPE MATERIAL: Gymnoreie dicei (Topsent); Topsent's (1895) original slide preparations are stored at MNHN LBIM No. H. T. 14.

DESCRIPTION The following description is based upon the original description (Topsent. 190 1 b). illustration (Topsent. 1904) and reexamination of two original (Topsent. 1895) slide preparations of the lectotype. Data in micrometres (pm)-f st.dev.~.

Body Forrn: The illustrated (Topsent, 1904) lectotype (Plate 15, Fig. A) is "ear-shaped", asymmetrical and incomplete. The specimen is a network of fiequently anastomosing tubes and measures 8 cm in height, 4.5 cm in width and 3 cm in thickness at the distal end. Distally, tubes have open and nearly circular apertures 8 mm in diametre; tubes are generally less than 8 mm in length; proximally. tubes are rnuch reduced or altogether absent and this region may be interpreted as a stalk; method of attachment to substrate unknown. External surface with numerous, fine pores that correspond to epirhyses. It is not possible to fimctionally determine gastrai and dennal regions from Topsent's description. Skeletal Framework (Plate 15. Figs. H, 1): Information rqprding the skeletal frarnework is sparse; a three-dimensional network of hexactin megascleres: beams of extemal surfaces (Plate 15. Fig. H) thick (30 pm). synapticulation extensive (Topsent. 1904; PI. VIL Fig. 5c); mesh is polygonal; pores on extemal surfaces of two types: 1) small pores near megascleres (presumably synapticula and 2) larger pores 200-400 pm in diarnetre (possibly epirhyses?); shon spurs present on extemal surfaces. Beams of the interna1 (gastral) surface (Plate 15, Fig. 1) are thin (1 8 pm); meshwork is rectangular; long. thin spurs as freely radiating rays of dictyonalia present. Beams covered with strong thoms throughout; dictyonal nodes not swollen.

Loose spicules (Plate 15, Fig. B-G): No dermalia or gasualia. Scopules appear to be of two sizes, however. a size-frequency distribution does not confirm a bi-modal distribution (overall length 328114530). Both scopules reportedly occur on extemal and intemal surfaces (Topsent, 190 1 b). Smaller subtyloscopules (Plate 15, Fig. F) with four straight and siightly divergent smooth tines. which terminate distally in a small sphere. are common. Tines aise fiom a swollen (axial cross is associated with swelling) capitulurn (8+650 width); capitulum is connected to a straight. or sliphtly curved. proximally pointed. and smooth shafi. Large tyloscopules (Plate 15, Fig. C) are uncornmon. Four straight. microspined and divergent tines are distally adorned with small. proximally oriented, curved spines. The slightly swollen and microspined capitulurn (8I69 width) gives rise to a straight, microspined shafi; proximal terminus of shafi is swollen and club-like. Straight, or highly curved. uncinates (Plate 15. Fig. D) (overal length 509+146jj) witli smooth, or siightly grooved shafts, are abundant and occur peripherally in bundles of tllree to six; bundles are oriented parallel to axis of growth. Microscleres as three types. Small oxyhexactins (Plate 15, Fig E) are rare. The six rays (35 prn length) are smooth, straight and sharply pointed. Onychohexactins (Plate 15, Fig. B) are large (62+653 diarnetre) and abundant. Long (26ISS3length) and srnooth primary rays give rise to 4 short ( 1 5- 1 7 pm long). curved secondary rays, or hooks, which terminate distally in a sharp point. Smaller tylohexasters (Plate 15. Fig. G) are extremely rare. The short primary rays give rise to 4-7 curved secondary rays, which terminate distally in a small sphere. DISTRIBUTION Gyrnnorete alicei (Topsent); lectotype collected by the 'Prince Albert' Expedition in 1895 from the Azores at Station 578: 3g026' N: 26" 30' 45" W. Depth: 1 165m: substrate: sand. Gyrnnot-ete alicei (Topsent); paratype material collected at Station 602: 3 8'38' N: 28" 1 3 ' 45" W. Depth: 1230 m; substrate: rock- Gymnorete alicei (Topsent); paratype material collected at Station 1319: 3S03S' N; 28" 5' 45" W. Depth: 1250 m; substrate: volcanic sand. Plate 15 (Gyrnnoreie alicei (Topsent)): Figure A, lectotype (modified from Topsent. 1904: Pl. IV. Fis. 8); B, onycnohexactin; C, tyloscopule; D. grooved uncinate; E. osyhesactin; F. subtyfoscopule; G. tyloscopule; H, diagrammatic representation of extemal framework surfac (L = large pore; s = small pore); 1, diagrarnmatic representation of interna1 (gastral) fiarnework surface; J, geographic distribution (solid dot denotes lectotype locality; open dot represents paratype localities).

ENDORETE TOPSENT, 1928

Ettdorete Topsent, 1928a: 3; Topsent, l928b: 299; de Laubenfels, 1936: 187; Reid. 1958b: 18; Reid, 1963: 224; Reid, 1964: cxxxvi.

TYPE SPECIES Endorele pertusm Topsent. 1928 (holotype by designation, Topsent. 1928a: 3)

BASIS OF DIAGNOSIS Diagnosis based solely upon Topsent's (1 928a & 1W8b) original descriptions.

DIAGNOSIS Body is a mass of rarnified and anastamosing tubes; tubes hollow. with circular oscular apertures: outer surface of tubes with large depressions or perforations presumed as epirhyses. Dictyonal meshwork regular to irregdar; beams with distinct. large hook-like spines: heavily spined spurs present on external surfaces of dictyonal fiarnework: no dermalia; gastralia possibly as pinnular hexactins: scopules, uncinates. onychohexactins, onycho- and discohexasters comrnon.

REMARKS Topsent (1 928a) erected this monospecific genus for the distinct mas of anastamosing tubes possessed by an apparently well-preserved specimen collected fiom Sagarni Bay. Topsent described this new genus and the holotype of Endorete pertusm Topsent. in two separate pubIications. His earlier publication ("On two Euretids from Japan"; Topseni. 1928) included a reasonable description of the holotype and a bnef diagnosis of the genus

Endorete: "Rarnrjkd, tubular euretid. destitule of dermalia but provicied wifhgustralia ztnder- the form of hexactins rhat, usually, swell ut one of their ruys. The rnicroscleres are discohexasters and onychohexasters. "(Topsent, i 928a: 3). His later ( 1928b) publication expanded the initial description. included two illustrations of the holotype and several uninformative line drawings of the fimework and spicules. Topsent (1 928b) correctly recognized sirnilarities in spiculation between his Endorete. Gyrnnorere Topsent and Heterorete Dendy; most notably in regard to the lack of dermalia. Reid (1 958b) regarded Topsent's Endorete as a subgenus within the genus Errrefe and later (1964: cmvi) hypothesized the lack of derrnalia as a secondary loss and a derived feature. Recent revision of Topsent's descriptions and illustrations (1928a,b) found considerable ambiguity with regard to body and spicule fom. Concerning body form. Topsent's (1 928 a & b) descriptions and illustration gives little information regarding the iinderside and method of attachment of the specimen. Perhaps more importantly, Topsent poorly described, and did not illustrate, the taxonomically relevant "pinnular hexactins'.. S imiIarly, Topsent is unclear about the pinnule-like spurs on the external surface of the frarnework - perhaps these dictyonalia are in fact dermalia that secondarily fbsed to the framework. hence explaining the unusual lack of dermalia. A recent examination of a fragment of the type specimen uncovered spicules and framework very unlike those described by Topsent (1 928a), suggesting that the "mass of anastmosing tubes" of E. pertrrsrn might contain more than one species. Although the noticeable "polygonal" perforations (epirhyses) of E. pet-fzrsnz are more reminiscent of the family Tretodictyidae Schuize (genus Hexactinella Carter) than the family Euretidae, their nature can not be properly assessed until the holotype is reexarnined. or fresh material is collected. Endorete is provisionally retained as a valid genus within the famiiy Euretidae.

DISTWBUTION Endot-efeperrzmn Topent, 1928. Single known specimen (holotype) from Sagami Bay. (see Plate 16, Fig. 1 for geographic distribution)

Endorete pertusrn. Topsent (Plate 16. Figs. A-1)

Etdorete pertusrn Topsent. 1 928x3; Topsent. 1 W8b: 299. PI .I. Fig. 1 -9. Pl. 1 II. Fig. 1 -2. TYPE MATERIAL Endorete perfusrn Topsent, 1928a; holotype thought to be stored dry at the Oceanographic Museum of Monaco; collected fiom Sagami Bay, exact location and depth of collection unknown.

DESCRIPTION The following description is an interpretation based solely upon Topsent's original descriptions (1928 a & 1928b) of the only known specimen of Endorete perfusrn..

Size. shape and colour: The incomplete type specimen (Plate 16. Fig. A; interpreted as a "bird's eye view" looking down upon the specimen) is a mass of ramified and anastamosing tubes 40 cm in length. 18 cm wide (as calculated from illustration) and 30 cm in height. Tubes are hollow, subcylindrical or flattened on one side, and rarely longer than 13 mm; tube apertures generally circular and presumed as oscular: tube walls thin (0.8 - 1.5 mm thick) and with numerous "polygonal alveolar" depressions (1 - 1.8 mm in diarnetre) which ofien form perforations thought to be epirhyses (Topsent, 1928a,b); pedorations separated by straight ndges (Plate 16, Fig. H). Method of attachment to substrate unknown. Colour primarily brown. representing areas not living at the rime of collection. and white desiccated areas presurnably living at time of collection. Interna1 surface of tubes considered gastral surface and extemal surfaces are considercd dermal.

Skeletal Frarnework: Dictyonal framework is an irregular three-dimensional network of hexactin rnegascleres. Meshwork generally narrow and irregular but may become larger. more regular and rectangular internally (length 3 15 pm;width 120 pm); meshwork often obscured by intercalated small hexactins. Beam thickness vanes from internai regions (20 35 pm thick) to external surfaces (30 - 40 pm thick); bearns distinctly ornamented with long (45 pm) spines which ofien are hook-like. Dictyonal hexactins on external surfaces with freely radiating rays (spurs) which are shon (90 - 190 pm), thick (24 - 50 pm), obtuse and ornarnented by short serrated spines. Loose spicules (Plate 16. Figs. A-1): No dermalia(see remarks). Gastralia as pinnular hexactins (Plate 16, Fig. B). Tangential and proximal rays are straight and teminally pointed. Distal ray (perhaps also tangential and proximal rays) commonly pimular but may be scarcely spiny. Pinnular hexactins of varying diarnetre (85 - 270 pm) and ray (tangential and proximal?) thickness (4 - 1 1 pm). Pinnular hexactins rarely fuse together to form dodecactins. Uncinates cornmon in bundles of 3 or 4 on dermal surfaces (Plate 16. Fig. E). Uncinates are straight or slightly curved, pointed at both ends. grooved without barbs, and 400 - 900 pm in length. Tyloscopules of two varieties. Srnaller tyloscopules (Plate 16, Fig. C) (350 - 560 pm overal length) occur on both dermal and gastral surfaces. Tines (tines and capitulum 65 - 95 pm length) Vary between two to three in number. are slightly divergent. slender. microspined and terminate distally in a small .'button" or slight swelling (interpretation Wheeler. 1998). Tines emanate from a slightly swollen and srnooth capitulum from which the smooth shafi proximally tapers to a point. Larger (0.7 - 1 .O mm length) tyloscopules (Plate 16. Fig. D) with 3 - 5 tines are abundant on the gastral surface. Tines (tines plus capitulum 1 00 - 1 20 pm length) are liard1y divergent and terminate distally in a swollen club which is omamented with recurved spines. The shaft is suaight. microspincd or smooth and proximally tapas to a blunt terminus. Microscleres as onychohexactins. onycho- and discohexasters. Onychohesactins (Plate 16. Fig. F) (45 - 70 pm diarnetre) have straight and slender principal rays from which four claws (in the same plane) are distally located. Onychohexasters (40 pm diametre) with short (3 - 4 pm length) principal rays which give rise ro three curved and divergent secondary rays are also common. Secondary rays end distally with what can be interpreted as an intemediate tom of onycho- and disco-tip. Discohexasters (Plate 16. Fig. G) (30 -50 pm diametre) with 4 - 7 secondary rays (6 - 12 pm length). which are arranged in a cup-shape. are abundant.

DiSTRIBUTION Endor-eie perfrtsrn Topsent; holotype collected hmSagarni Bay: (35" N. 139" 30' E); depth unknown. Plate 16 (Endorete perrusm Topsent): Figue A, holotype (modified from Topsent, 1928b: Pl. 1. Fig. 1 ); B, pimular hexactin; C' tyioscopule; D, gastral tyloscopule with diagrammatic representation of tine head: E. grooved uncinate; F. onychohexactin; G. discohesactin; H, fragments of holotype (modified fiom Topsent. 1928b: Pl. 1. Fig. 2); 1. geographic distribution (solid dot denotes type locality).

Discussion

Phylogenetic Analysis Hypothesised phylogenetic relationships (Figure 7) based on binary, unordered (Wagner parsimony) and unweighted characters (Table 1) were established by heuristic search on PAUP 3.1.1 (Swofford, 1993). MacClade 3.0.7 (Maddison & Maddison. 1992) was used for character analysis. Plesiomorphic States were established by outgroup cornparison with Farreu occa Bowerbank. Refer to "Methods" for matrix construction and character selection. The use of the Farreidae as outgroup to the Euretidâe is based upon their proposed (Ijima. 1927) sister group relations, also, to the fact that the Farreidae is the only hexactinosan famil y to have undergone recent revision (in prep. Reiswig). On1y type species were used as terminal taxa. The phylogenetic analysis resulted in three most parsimonious trees (84 steps). and to surnmarize these trees. a strict consensus tree was produced (Figure 7). The three trees di ffered on1y in the Caiyptorefe - Eurete - Pararete clade, resulting in an unresolved polytomy in the consensus tree. The Consistency Index (CI) for al1 three trees was 0.44. indicating significant homopIasy within characters. This CI value is lower than those of most demosponge phylogenetic analyses (Hajdu & Van Soest. 1996). even though sirnilar character categories were employed. As the Consistency Index may be negatively correlated (Forey et al.. 1992) with both the number of taxa and characters, the Retention Index (Ri) was also calculated (RI=0.5). This intermediate value indicates that a proportionally equal number of state changes occurred at internai nodes of the dadogram as towards the terminal taxa. Bathyxiphus 1 Conorete Pleurochorium Verrucocoeloidea

Calyp torete Eurete

Pararete

C honelasma

Ptychodesia

Lefroyella

Iphiteon

Periphragella

Myliusia Endorete

Gymnorete

Heterorete

Farrea

Ficure 7. Hypothesized relationships between genera (generic type species) of Euretidae based upoii computer-generated phylogenetic analysis using parsimony (PAUP 3.1 & MacClade 3.0). Each nurnber on the cladograrn refers to one branch of the cladograin (see Table 2 for character changes). The tree is derived from the strict consensus of 3 niiniiiiuni length trees (84 steps), bascd 011 an unweightcd. uxiordcrcd. biiiaq cliaractcr sct (Consistency Index = 0.44). polarization deter~ninedby outgroup coiiiparisons witli [lie genus firreu Bowqbank. Group A and B are desigi~atedas subfai~~iiics(sec test). Table 1. Characters (a-n) used in phylogenetic analysis. Plesiomorphic States (as judged by outgroup cornparison with Fart-eu occa Bowerbank) are coded as zero; asterix denotes indeterminable polarity. Autapomorphic and constant characters were not used in analysis.

State Nurnber Charpcter State

body aP-*aPe 4 oxyhexastefs O=abs; 1=pres body funnel-shape r onydiohexasters O=abs; 1=pres body rnass of tubes s epirhyses O=abs: 1=presg other body shape t aporhyses O=abs: l=presg pentadins u complex channelization O=abs; l=pres oxyhexacîins v regular intemal framework O=abs: l=pres0 discuhexactins w rotulate frarnework O=abs; 1=pres pinnular hexacüns x longitudinal dits on surfaœ(s) O=abs; l=pres tyloscopules Y ridges on surface(s) O=abs; 1=pres oxysmpules z didyonaf strands O=abs; l=presg strongyloxopules aa dictyonal lamellae O=abs: 1=preso discoscopules bb swollen didyonal nodes O=abs; 1=pres davules CC Iuberded didyonal nodes O=abs; 1=pres bracketed uncinates dd smooth dictyonal beams O=abs: 1=pres0 barbed uncinates ee rnicrospined diayonal beams O=abs: l=pres disahexasters ff spines on didyonal beams O=abs; l=pres

Table 2. Character changes for each branch of cladogram (for branch numbers see Figure 7; for characters see Table 1 ).

Bnnch # 1 1of Changea Changes 1 1 Bmnch # 1 # of Changes 1 Changes I

f!aa la

g 0,l

g 1 k (Pl n (Pl Analysis of C haracters This "post-character analysis" (as deiermined by character topology) focuses on three areas: (1) characters used in phylogenetic analysis (labelled by letters of the alphabet; an asterix (*) denotes ancestral state as judged through outgroup cornparison with Furrea occa; a cross ( X ) denotes those constant, autapomorphic or symplesiomorphic characters); (2) performance of those characters (Consistency Indes. or CI), and (3) general discussion of characters for classification. Body form (Tables 1 & 2, chars. a, b. c, d) variability was panitioned into five characters: (a), cup-shape (CI=0.5); (b), fumel-sliape (CI=0.3): plate-shape ( X ): (c). *mas of tubes (CI=0.3); and, (d), other shapes (CI=0.5) (to include forms such as the blade-1i ke forrn of Bathyxiphlrs and the unique laterd branching of Pleurochorium). Some forms (e.g. Calyprorete ijimai) were considered as both cup-shaped and a mass of tubes. Body form characters performed poorly (low CI values) in the phylogenetic analysis. As there are a number of autapomorphic (and hence easily recognisable) body forms within the family, this result is not surprising. Nor is it surprising that authors studying extant hexactinosans (Schulze. 1 887; Ijima. 1927; Reid. 1964) placed little emphasis on body form in their higher (above genus level) classifications. Obviously. more information about dictyonine sponge development and growth and variability of body forms within genera is needed. Interestingly, Chonelasma and Ptychodesia did croup together (branch nurnber 3 1) in the phylogenetic analysis, thus lending further C support to Schulze's ( 1887b) suggestion that the unknown body form of C'honeZrsma Zurnella is plate-like, sirnilar to that of Prychodesiu duplicara. The plate-like body form of Ptychodesiu was autapomorphic for Euretidae and not considered informative. Other features associated with body form were analysed: longitudinal dits or grooves (character x; CI=1 .O) and ndges (character y; CI=l .O) on either the interna1 or extemal surface of the sponge. These two characters performed well in the analysis and may prove useful for classification. The presence or absence of a basal plate is not known for rnany euretid type specimens and hence was excluded fiorn this analysis. Dermalia and gastralia (Tables 1 & 2, chars. e. h), if present. are generally in the form of: (h), * pentactine megascleres (CI=O.3) or, (e) pinnulated hexactine megasclsres (CI=0.3). Although pentactine dermalia and/or gastralia are fairly widespread. and possibly apomorphic within the Euretidae, they appear to have been lost at least three separate times (PIcurochoriurn. Lefioyella and the clade formed by Endorete. Gyrnnorrte and Heferorere). This lends support to both Ijima's (1927) and Reid's (1 964) speculations that pentactins may be lost secondarily. This phylogenetic analysis also suggests that pinnules developed independently in three separate euretid lineages (Endorete, Ptychodesia and the Conorere - Bathyxiphus clade). As pinnules occur both alone and in combination with pentactins, their development is probably not associated with pentactins. Scopule form (Tables 1 & 2. chars. i, j. k, 1) was divided into five characters: (i) tyloscopules (CI=0.3): 0) oxyscopules (CI=0.5); (k) strongyloscopules (CI=O.3): subtyloscopules ( X ), and. (1) discoscopules (CI= 1 .O). The tyIoscopule is the most widespread of euretid scopules (found in 10 of 16 genera) but evolved independently three times within Euretidae (seven taxon clade formed at branch number 30. Pcri/lhragdiu and Enclvrere - Gymnorete clade). Notable is the presence of tyloscopules in al1 tasa of Group A (Figure 7). Oxyscopules also evolved twice within Group A (Figure 7). Although information on scopule development is lacking. oxyscopules may represent juvenile forrns of other scopuIe types. perhaps tyloscopules. Indeed. osyscopules are found in combination with tyloscopules, and are never found donc, Unlike the oxyscopules. strongyloscopules are found in both Group A and Group B. and tlieir appearance in three separate lineages (Conorele - Bu~hyxiphlrsclade. Catyp~ore~e and Lefi-o~liu)suggests parallel development. Discoscopules, although equally rare as oxyscopules, performed substantially better and are found in only one clade: Choneiasma - Plychodesiu. Subtyloscopules were autapomorphic for the Euretidae. Striking is the complete absence of scopules in three euretid genera (Iphireon. Myiilrsia. and Hem-oi-erc). AI1 three "scopule-less" genera faIl within Group B (Figure 7 j and their positioning suggests tliat scopules were lost independently three times. It should be noted that it is primarily upon the presence or absence of scopules that the six farnilies of the order Hcxactinosa are distinguished. Under the current (Ijima, 1927) classification. the absence of scopules within genera of Euretidae suggests they be moved to another farnily. or given their own family within the order Hexactinosa. In order to properly assess the loss or gain of scopules, a phylogenetic analysis including taxa from dl hexactinosan families must be performed. It is also important to recognise that this phylogenetic anaiysis used Fun-euocca, which does not possess scopules, as an outgroup to the Euretidae, and consequently the plesiomorphic state for scopules was "absent", and not a type of scopule (e-g.tyloscopule). Obviously, an outgroup with scopules must be chosen for comparison with the Euretidae, however? until the taxonomy and description of al1 hexactinosan genera are revised, such an analysis is fùtile. Lrnfortunately, the lack of scopule character C polarity in this analysis restricts comments on the evolution and development of the scopule. and consequently. the evolution of the order Hexactinosa. For example. information on scopule development, or ancestry, can be used to test both Schulze's (1 887) and Lendenfeld's (19 15) theories. which suggest that scopules are derived forms of hexasters. Uncinates (Tables 1 & 2, chars. n, O) were either bracketed (n; CI=0.5) or barbed (*O;CI=0.5). Barbed uncinates are by far the more prevalent among euretids (10 of 16 genera) and this analysis suggests they were Iost (branch number 26; Figure 7) and regained once (Chonelasmu - Piychodesia - Lefioyeiia clade). The bracketed uncinates. if considered nonhomologous with barbed uncinates, appear to have evolved twice (Ccdyptorcle. and the Gymnor-ele - Endorete clade). Regardless of homology, uncinates are homoplastic within the Euretidae and on their own are not usehl indicators for natural groups. Microscleres (Tables 1 & 2. chars. f. g, p. qTr) are of six types: (f). osyhexactins (CI=0.2): (g) discohexactins (CI=O.S): (p) * discohexasters (CI=0.25): (q) oxyiiexasters (CI=O. 17); tylohexasters (X ): and. (r). onych~hex~asters(CI=0.5). Microscleres perfbnned the worst of al1 euretid categories and, are not good indicators of natural çroups. Discohexasters are the most widespread microsclere (1 1 of 16 genera) followed by oxyhexasters (8 of 16 genera) and oxyhexactins (7 of 16 genera). This result suggests that Ijima's (1927) reorganisation of Euretidae, based upon microsclere type. is not appropriate. From what little is known about the development of hexasters (from Okada's (1 928) studies on Furrea sollusi) it may be inferred that discohesasters are formed after oxyhexasters, either (1) directly tiom oxyhexasters. or. (2) originated independently from oxyhexasters. In the first case. it is quite likely then. that the hexasters recognised here as different forms, are. in fact, part of a transformation series and, therefore, homologous. The cornmon occurrence of both disco- and oxyhexasters. in combination or alone, within euretid genera indicates that they are symplesiomorphic, and therefore of limited use in establishing euretid phylogeny. In the second case. if discohexasters are independcntly formed, then they are not homologous to oxyhexasters. and the microsclere characters used in this anaiysis are homoplastic.

Numerous characters of the dictyonal framework were used in this analysis

(Tables 1 & 2, chars. v, W. z, aa, bb. cc, dd, ee, ff). A key assumption involved with the chancter of regular intemal framework (v*, CI= 0.67) was that the regular, one-layered. framework of the outgroup, Farrea occu, is homologous to the regular intemal frarneworks of most euretids. This assumption is supported by Reid (1 964). who suggested that the regular one-layered frarnework of F. occu is the ancestral condition of most hexactinosans. This character (v) perforrned well in the analysis only having been lost twice (Plezrrochoriurn and Heferorete). The character of irregular external frarneworks was excluded from this analysis because it was present for al1 taxa except the outgroup. Rotulate fiames (W. CI=0.5) were observed only in two taxa: iphifcon panicea and 1\4yZizrsia callocyuthus. Although the two taxa did fa11 close to each other (Group B). they did not fonn a clade on their own. This suggests that the rotulate frarnework evolved more than one time. Dictyonal strands (character z: CI=O.S) are observed in the majority of euretids and therefore not a usefd character for higher classification. Only two taxa (Heterorete and Iphifeon) did not possess dictyonal strands. This would suggest a difference between the rotulate frameworks of Iphiteon and Myliusia. as Myliusia did show dictyonal strands. The character of dictyonal lamellae (aa; CI=0.25) is lost and regained in several separate lineages. Swollen (bb; CI=0.5) and tubercled nodes (cc: CI=0.5) were observed in two taxa (Pararete and Myliusia) which are in Group A and B respectively; demonstrating parallel deveIopment. nie use of spurs as a character was rejected in this analysis for ail. except one (A4yliusia), are in possession of spurs. The presence or absence of smooth bearns (dd; CI=0.3) showed an unexpected result. Although this characteristic appears to have been lost or gained several times. six of the seven taxa of Group A (Figure 7), and only one taxon of Group B, have smooth beams. Microspines (ee; CI=0-3) on dictyonal bearns are lost and regained several times. The regular arrangement of microspines on dictyonal beams was autapomorphic (Myliusia) for the Euretidae and consequently ignored. The character of spines (character CI=] .O) on dictyonal beams performed very well, having been gained only once. distinguis hing the Endorete - Gymnorere - fieterorete clade. The channelization (Tables 1 & 2. s, t, u), or lack of, in the intra- or extradictyonal framework is an important euretid Feature. The overall presence or absence of epirhysis (s; CI=0.67) and aporhysis (t; CI=1 .O) in either the intra- or extradictyonal fiarnework are the primary characters separating Group A and Group B (see Figure 7). The character of complex channelization (character u: CI=0.5) is. admittedly. an artificiril and subjective attempt at the categorisation of forms with wall-folding. arnararhysis and/or cavaedial spaces. This character did not perforrn well (low CI) in the analysis but did help group together those forms with complex body forms (Ptychodesia, Lefioyella, iphiteon and Periphrugella). Obviously this character (u) is homoplastic and should be revised for further analyses. It should be noted that the Euretidae has historically (Schulze. 1 887; Ijima, 1927) been diagnosed without fiarnework channelization. However. review of Zittel's (1 877) action in erection of the family Euretidae revealed that he intentionally divided the family into two groups: (1) Group A, for those fonns without channelization. and (2). Group B. those forms with. 'scarcely developed' channels (canals). This phylogenetic analysis supports Zittel's (1 877) original diagnosis that channelization is present within the Euretidae.

The Iow overall CI (0.44) of this analysis suggests that considerable homoplasy exists among the characters used. Little is known about "intra-generic" hexactinellid character variability and it is likely that some character States assigned to euretid generic type species in this study may also be used to differentiate species within the same genus. The lack of knowledge of chancter variability across taxa may account for the high level of homoplasy within these characters. This analysis also lacks confident character polarization. Although the use of Frrrreu Bowerbank as outgroup did serve to polarize many characters. several important characters (Le. scopules) were lefi unpolarized. Future analyses may benefit by incorporating more than one outgroup and ontogenetic information. Post-analysis character examination reveded that single characters did not define natural groups, however, character combinations did serve to separate the Euretidae (Figure 7). The Euretidae is divided into two main clades. Groups A & B, which are di fferentiated primarily by the presence or absence of skeletal channel izat ion. beam omarnentation and tyloscopules. Groups A and B may be regarded as subfamilies of the Euretidae. Also distinguished is the Heterore/e - Gymnorete - Enclorete clade (branch number 32. Table 2). This clade is separated on the presence of spined dicytonal beams and lack of penactine dermalia ancilor gastralia and dictyonal strands.

Classification Euretidae contains two subfamilies. sixteen valid genera. fifiy species and nine subspecies (see Table 3). Four genera are synonomyzed with existing valid genera, two subspecies and one form are elevated to species status, three new species are narned and three new species combinations are presented. This revised classification of Euretidae is unique. for not only is it the first revision of a hexactinellid taxon to incorporate evidence from a cornputer-assisted analysis of phylogeny. it is also based upon a variety. and combination. of morphological characters and does not place undue importance on one category of characters. This contrasts with previous schemes which gave pnority to fiamework morphology and channelization (Zittel, 1877; Reid, 1963), framework composition (Carter, 1887), presence or absence of key spicules (Schulze, 1887) or spicule type (Ijima, 1927). Table 3. List of valid genera, species and subspecies of the Euretidae (type species in bold). Geographic distribution Although several species are found in the Antarctic region, euretid distribution appears largely tropical (Figure 8), confïrrning Tabachnick's (1994) earlier results. Primary concentrations of euretid sponges occur in : (1) the Indo-Pacific; (2) North West (Pacific Rim); and (3) Caribbean region, however. any further attempts here at biogrographical analysis are greatly restricted by the broad geographic range and level of taxonomie rad employed. It is aiso possible that the extent of global ocean-floor sampling is such that the observed distributions are artifactual and not representative. Cornparison of generic type localities (Figure 8) with the cladogm (Figure 7) revealed no "cladistic areas".

Conclusions As pointed out in the 'Literature Review', little information pertaining to basic biology and ontogeny exists for hexactinellid sponges. The lack of ontogenetic information greatly hampers cladistic analyses, for undue weight is placed upon the outgroup method of character polarisation. Consequently, attempts at producing phylogenetically "naturai" classifications are made difficult. Perhaps further studies on "SCUBA accessible" hexactinellids will reveal more detail of their development. Re-examination of the type species of 19 genera included in Euretidae at one time or another found that variability among descriptions of different authors. and overall lack of knowledge of hexactinellid morphology of early authors, were largely responsible for mis-identification of important features and mis-description of several species. Euretidae contains two subfarni lies, sixteen valid genera, fifty species and nine subspecies. Phylogenetic anaiysis of generic type specimens of the Euretidae suggests the majority of available characters (body form, spicules, and fiamework) are prone to homopiasy and are generally not useful for establishing "natural" groups. However. this analysis did help to elucidate "artificial" groups within the Euretidae and served as a template for a revision of the classification and artificial key. The poor performance of many of the available euretid characters in the phylogenetic analysis places renewed importance on biochernical and genetic information to help determine hexactinellid relationships. Figure 8. Geographic distribution of the Eiireridae (6=localit)' of generic type specimens; = type locality of valid euretid species; O = type locality of euretid subspecies; O = iinideiiiificd specimeiis References

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