Revision of Rogneda Uljanin, 1870 (Rhabditophora, Eukalyptorhynchia, Polycystididae) with the Description of Seven New Species

Zoological Journal of the Linnean Society, 2008, 153, 1–28. With 12 ﬁgures

Revision of Rogneda Uljanin, 1870 (Rhabditophora, Eukalyptorhynchia, Polycystididae) with the description of seven new species

TOM J. ARTOIS*

Hasselt University, Centre for Environmental Sciences, Research Group Biodiversity, Phylogeny and Population Studies, Agoralaan Gebouw D, B-3590 Diepenbeek, Belgium

Received 14 May 2007; accepted for publication 10 July 2007

A morphological and taxonomical account of the taxon Rogneda is given. A new interpretation of the stylet morphology is presented and a new terminology pertaining to these hard parts is proposed. All known species are discussed and additional information is provided when necessary. Seven new species are described. Five of these are from the Mediterranean: R. colpaerti sp. nov., R. licyae sp. nov., R. schaereri sp. nov., R. valckei sp. nov. and R. vangronsveldi sp. nov. R. verveckeni sp. nov. is from the Atlantic coast of Spain (Galicia). The seventh, R. martensi sp. nov., is from Indonesia and is the ﬁrst species of Rogneda to be found outside the Northeast Atlantic and the Mediterranean. All new species can be recognized based on the morphology of the stylets in the male system. A discussion on the similarities and differences of the 22 species is provided, resulting in the grouping of the species into seven morphological groups: the capulata-group, the falcata-group, the hibernica-group, the polyrhabdota-group, the steueri-group, the tripalmata-group and the westbladi-group. R. minuta cannot be placed into any of these groups and therefore stands alone within the taxon. An identiﬁcation key is provided to facilitate further study of the taxon Rogneda. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153, 1–28.

ADDITIONAL KEYWORDS: identiﬁcation key – morphology – Platyhelminthes – Rhabdocoela – taxonomy – Turbellaria.

INTRODUCTION of the morphology, biogeography and systematics of the taxon. Thus, he provided a solid basis for later Although the genus Rogneda Uljanin, 1870 is one of descriptions, as the internal morphology shows very the most species-rich taxa within the Polycystididae little variation within the genus, and species descrip- Graff, 1905, only very few studies have been dedi- tions can now easily be made by only describing the cated to it. The ﬁrst species to be described was often very complex hard parts of the male system. Rogneda minuta by Uljanin in 1870. More than Three years later, Ax (1956) described a new sub- 80 years later, Karling (1953) described three new species (R. westbladi gallica). In 1959, Ax described species (R. anglica, R. capulata and R. westbladi) and a new species (R. polyrhabdota), and gave a detailed transferred three species formerly described as Poly- redescription of the enigmatic species R. tripalmata, cystis Kölliker, 1845 species to Rogneda [R. tripal- based on newly collected material. Later, several new mata (Beklemischev, 1927), R. steueri (Steinböck, species were described by Brunet in 1965 (R. falcata), 1933) and R. hibernica (Southern, 1936)]. In that 1969 (R. cincta, R. palula, R. reticulata) and 1979 seminal monographic treatment of Rogneda, Karling (R. acuta, R. exilis). Since then, no more descriptions (1953) gave an extensive and comprehensive overview have followed. All these species were described from the European North Atlantic coasts south to the English Channel and from the Mediterranean. No *E-mail: [email protected] species were ever recorded from other areas.

In this contribution I describe seven new species. not designate a holotype for R. palula, although he Five of them are from the Mediterranean, one is from did do so for all other species he described in that the Atlantic coast of Spain (Galicia) and one is paper. The syntype, however, is still in the collections from Indonesia and is the first representative found of the Swedish Museum of Natural History, and one outside the Northeast Atlantic or the Mediterranean. whole-mounted specimen is chosen as lectotype. The In order to facilitate future identification of species original material of R. minuta, R. steueri and R. hi- within this taxon I give an overview of all species and bernica has never been kept or has disappeared. In add new data for the previously described species if the meantime, however, new material has been col- necessary and where possible. A drawing of both lected. As all conditions for the designation of a stylets is provided for each species. If possible and neotype are fulfilled (ICZN, 1999: Art. 75), I have useful, a photomicrograph of the entire specimen or designated a neotype for each of these species. For (one of) the stylets is also given. I re-measured all R. steueri and R. minuta, these neotypes are chosen stylets based on my new interpretations of the con- from material present in the collection of the Swedish struction of these stylets (see General Morphology). Museum of Natural History. The neotype of R. hibernica is a specimen collected by members of the research group Biodiversity, Phylogeny and Popula- MATERIAL AND METHODS tion Studies of the Hasselt University. Also, the origi- Whole mounts were studied of all species, and, if nal material of R. tripalmata has disappeared, and Ax available, also sectioned specimens were studied. The (1959), when redescribing it, only kept sectioned material is present in the collections of the research material. Sections, however, do not allow species iden- group Biodiversity, Phylogeny and Population Studies tification, as the detailed construction of the stylets of the UHasselt (UH), the Swedish Museum of cannot be studied, and therefore I refrain from des- Natural History (SMNH) and the Museum National ignation of a neotype for this species. R. gallica Ax, d’Histoire Naturelle in Paris (MNHN-P). Of R. tripal- 1956 was originally described as a subspecies of mata, only some sectioned specimens are available R. westbladi, but also without designation of a holo- (present in the collections of the II. Zoologischen type. Only one specimen is left of the syntype. Unfor- Institut-Göttingen), which were studied in great tunately, this specimen is a sectioned specimen, and detail by Ax (1959). Therefore, I did not study this does not provide any useful information as to the material. overall morphology of the hard parts. However, as it Live animals were extracted from sediment using is the only specimen available, it is this specimen that the MgCl2 method (see Schockaert, 1996). They were must be considered lectotype, following article 75.5 of studied alive and whole mounted with lactophenol. If the ICZN (1999). additional specimens were available, they were fixed in hot (50 °C) Bouin’s fixative, embedded in paraffin GENERAL MORPHOLOGY (FIG. 1) and serially sectioned. After sectioning they were stained with Haidenhein’s haematoxylin using eosin Karling (1953) gave an excellent and comprehensive as counterstain. Drawings of the hard parts were overview of the morphology of the taxon Rogneda, and made with a camera lucida on a Reichert Polyvar the same outstanding standard of description is found microscope, using Nomarski interference contrast. in the works by Ax (1956, 1959) and Brunet (1965, The orientation of the drawings of the hard parts is 1969, 1979). For a detailed discussion of the morphol- with the proximal ends towards the top of the page. ogy the reader is referred to these papers. The taxon Measurements were taken along the central axis of is so homogenous as to the internal morphology that the measured object. Drawings without a scale are it is impossible to identify the species based on the freehand. anatomy of the soft parts alone. For the description of Holotypes of the new species will be deposited in the atrial system I use the terminology proposed by the SMNH. All other new material, including Artois & Schockaert (2003) for the male system, and paratypes, will be deposited in the collections of the that proposed by Artois & Schockaert (2005) for the HU. Karling (1953) did not designate any type speci- female system. men, but for R. anglica, R. capulata and R. westbladi Mature specimens of Rogneda are 0.5–1 mm long. the original specimens are available and constitute They have two eyes (Fig. 1B: e). A number of species the type series (syntypes). One specimen of each of have a dark reddish-brown subepidermal pigment in these three species is labelled lectotype, most prob- the form of two or four, sometimes reticulated, dorsal ably by Karling himself on a later occasion, although, stripes (Figs 1A, 7D, 9A, 12A), while others are to my knowledge, this designation has never been completely transparent. Three species (R. westbladi, officially published. Therefore, I designate these R. minuta and R. vangronsveldi sp. nov.) have a specimens as the lectotypes. In fact, Brunet (1969) did parenchymal pigment that gives them a dark brown

Figure 1. A, pigmentation pattern as seen on a whole mount (R. schaereri sp. nov.). B, habitus of live animal (R. colpaerti sp. nov.). C, schematic representation of a stylet. D, schematic reconstruction on sagittal sections of the genital system from the left hand side (R. hibernica). Abbreviations: 1, plate 1; 2, plate 2; cga, common genital atrium; de, ejaculatory duct; e, eye; fd1, female duct type I; gl, glands; gp, gonopore; m, mouth; ma, male atrium; mb, male bursa; mu, muscles; od, oviduct; ov, ovary; p, proboscis; ph, pharynx; pv3, prostate vesicle type 3; stA, stylet A; stB, stylet B; t, testis; ut, uterus; vd, vitelloduct; vde, vas deferens; vi, vitellarium; vs, seminal vesicle. colour, but this is clearly not the granular pigment ph) is found in the ﬁrst body half and is of the normal discussed above. The epidermis is syncytial and com- polycystidid construction, with four hard teeth pletely ciliated, containing numerous rhabdites. around the proximal pharynx opening. As in all Eukalyptorhynchia, there is an anterior The common genital pore (Fig. 1B, D: gp) is situ- proboscis (Fig. 1B: p), ±1/5 of the body length long. It ated ventrally in the last quarter of the body. It opens is of the normal polycystidid construction and has into the common genital atrium (Fig. 1D: cga). The four pairs of retractors, supplemented with one pair of gonads are paired. The testes (Fig. 1B: t) are situated ventral integument retractors. The pharynx (Fig. 1B: at both sides of the body, left and right from the

© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153, 1–28 4 T. J. ARTOIS pharynx, lying rostrally from the ovaries (Fig. 1B, D: to assess on whole mounts, and very good sectioned ov). In R. capulata and R. exilis, the testes lie in the material is needed. Such material is lacking in the caudal body end, even posterior of the ovaries. In species described in this contribution, and therefore R. hibernica and R. falcata the testes are much larger this feature is not further discussed here. than in the other species, extending from the level of Because prostate stylet type III and accessory stylet the proboscis to the caudal body end. The vitellaria type I cannot be distinguished (see Artois & Schock- (Fig. 1B: vi) extend at both sides of the body, lying aert, 2003) I will continue naming them stylets A and dorsally to the testes. The ovaries are oval, situated in B as was proposed by Karling (1953). However, stylets the last body quarter, lying behind the testes (except named the same in the different species are not in R. capulata and R. exilis, see above). necessarily homologous (see Karling, 1953). I have The female system is rather simple. It consists of a kept the naming in each of the species as was pro- muscular female duct type I (the ‘ductus communis’ posed by the authors of the original description, and of Karling, 1953) (Fig. 1D: fd1), which leaves the have named the stylets in the new species in the same common genital atrium caudally and which proxi- way as in those species they most closely resemble. mally splits into the two oviducts (Fig. 1D: od). A For a discussion on the possible homologies see the large bundle of glands enters this bifurcation following section on Comparison of the species. I will (Fig. 1D: gl), which is often swollen and ﬁlled with deviate, however, from the terminology for the differ- sperm, functioning as a seminal receptacle. Each of ent parts of the stylets proposed by Karling (1953) the oviducts receives a vitelloduct (Fig. 1D: vd). The and followed by all later authors. Karling (1953) uterus (Fig. 1D: ut) is of the typical polycystidid con- described the basic shape of a stylet of a species of struction and leaves the common genital atrium from Rogneda as consisting of a proximal stalk (sa for its anterior wall. stylet A; sb for stylet B), and a distal part, normally The male atrium (Fig. 1D: ma) is very broad and itself made up of two parts (da1, da2 and db1, db2, muscular. From its most distal part, just proximal to respectively). At the base of da2/db2 there is often a the common genital atrium, a bursal stalk departs, basal extension (‘Basalscheibe’: pa2/pb2). My observa- which leads to a male bursa (Fig. 1B, D: mb) (not tions, however, suggest that the typical stylet can present in R. minuta). Karling (1953) considered this better be described as consisting of two plates that part of the male atrium the proximal part of the are connected to each other (Fig. 1C). I will refer to common genital atrium and named it ‘atrium supe- these plates as plate A1 and plate A2 (stylet A), and rius’. A prostate vesicle type III (Fig. 1B, D: pv3) plate B1 and plate B2 (stylet B). In a typical stylet, enters the proximal part of the male atrium. The vasa plate A1/B1 is rather slender and long, whereas plate deferentia (Fig. 1D: vde) form two large, weakly mus- A2/B2 is shorter and sturdier and attached more or cular seminal vesicles (Fig. 1B: vs). Distally these less to the midpoint of plate A1/B1. The proximal part vesicles join each other to form the ejaculatory duct of A2/B2 is rather extensive and is actually the (Fig. 1D: de), which enters the male atrium on a small ‘Basalscheibe’ of Karling (1953). The parts of the protrusion (called ‘penial papilla’ by Karling, 1953) plates A1/B1 proximal from the attachment of plate next to and close to the prostate vesicle. In all but one A2/B2 often show a longitudinal striping, caused by species the male atrium contains two hard, plate- superﬁcial foldings, which sometimes even give the shaped stylets (Fig. 1B, D: stA, stB): a prostate stylet impression of being tubiform. This is visible in almost type III and an accessory stylet type I, with the all of the species, except R. minuta (Figs 2, 4A, B), ‘penial papilla’ mostly in between them. In R. minuta R. martensi sp. nov. (Figs 8C, 9D, E) and R. reticulata the accessory stylet type I is lacking. The proximal (Fig. 8E). In the case of R. minuta the striping is parts of these stylets are connected to each other and absent, while in the two other species mentioned it to the wall of the male atrium by a very thick and could be present but not visible because of the poor complex musculature (Fig. 1D: mu). The relative posi- state of the specimens, as is the case in some whole- tion of the prostate vesicle proper is variable. In some mounted specimens of other species. In R. gallica species it is situated in between the proximal parts of (Fig. 8B), the striping is only visible just proximal the two stylets (R. hibernica, R. anglica, R. westbladi; from the attachment of plates A2/B2. see Karling, 1953), while in some other species the The stylets can be very complex, as both parts of prostate vesicle is situated ventrally from the stylets. each stylet can show extremely complex extensions. In these latter cases the prostate vesicle distally In some species, however, the stylets become simpler narrows to a duct (the ‘ductus granulorum’ of Ax, as one of the plates of the stylets can be lost, e.g. in 1959), which enters the male atrium in between both stylet A of R. tripalmata (Fig. 10C) and R. schaereri stylets along with the seminal duct (R. capulata, sp. nov. (Figs 6B, 7E). Not only are the stylets often R. steueri, R. polyrhabdota, R. tripalmata, R. reticu- very complex, but also some details can be difficult lata, R. cincta, R. palula). This feature is impossible to observe. For instance, one important character is

© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153, 1–28 REVISION OF ROGNEDA 5 whether the distal rims of the constituting plates are species of Rogneda, and therefore R. westbladi is serrated or not. These ends are, however, extremely considered unpigmented. The same is true for the thin, and therefore easily overlooked in whole varying pattern of brownish–blackish pigment that is mounts. observed in R. minuta (see Karling, 1953; Brunet, 1969) and for the brownish body colour of R. vangronsveldi. COMPARISON OF THE SPECIES Alternatively, based upon the detailed morphology Apart from the differences found in the construction of the stylets of the male system, the species with two of the stylets and in the pigmentation patterns, the stylets (all except R. minuta) can be split into seven different species are almost identical to each other, groups: the capulata-group, the falcata-group, the except for R. minuta, the type species. This species hibernica-group, the polyrhabdota-group, the steueri- can be differentiated from all other species in two group, the tripalmata-group and the westbladi-group. important ways: it lacks a male bursa and it only has one, albeit complex, stylet (Figs 2, 4A, B). Both features are important in taxonomy, and especially the THE CAPULATA-GROUP presence of an accessory stylet type I, which is unique The capulata-group includes two species from the to the other species of Rogneda. It could therefore be Mediterranean with two dorsal pigment stripes and argued that Rogneda should be split into a monotypic an aberrant construction of stylet A: R. exilis Rogneda (with only R. minuta) and a new genus with (Figs 3A, 4C, D) and R. capulata (Fig. 3B). The rela- all other species. However, such radical taxonomical tionship between these two species has already been decisions should be backed up by a thorough cladis- discussed by Brunet (1979). The distal part of stylet A tical analysis of the Polycystididae as a whole, solving consists of at least two plates: a hook-shaped the question of the possible non-monophyly of the (R. exilis) or rectangular (R. capulata) one, and a taxon Rogneda. Until such analysis is performed, I broader oval one, with a serrated distal rim. In prefer to keep R. minuta and all the other species R. capulata there is a third (and possibly even a together within Rogneda for the sake of nomencla- fourth) plate, which is (are) also oval and with a tural stability. Apart from the morphological differ- serrated distal rim, but somewhat longer. In R. exilis ences mentioned, R. minuta also differs from the the hook-shaped plate has a proximal extension, other species of Rogneda in habitat preference, as it is lying parallel to the proximal plate of the stylet. the only species to be found on algae and sea grass. Therefore, this plate could be homologous with plate All other species are interstitial. A2 of the other species of Rogneda (see the remarks The taxon Rogneda can easily be split into two on R. exilis in the taxonomical account). This exten- morphologically distinct groups: those species with sion may also be present in R. capulata, but this pigment and those without. The simplest pigmenta- cannot be determined with certainty because of the tion is found in R. vangronsveldi, which has a poor orientation of the stylet in the specimens avail- pigmented area around the eyes. In most other pig- able. In this species, the other two (three?) plates are mented species, however, the pigmentation consists of thus probably subdivisions of plate A1. Stylet B of two reddish-brown dorsal stripes and, sometimes con- both species is very simple, with a long and narrow nected to them, two stripes in front of the eyes. This plate B1 and a simple, shorter plate B2. Plate B2 is situation is found in R. anglica, R. capulata, R. exilis, attached to the proximal part of B1 in R. capulata, R. palula, R. steueri, R. tripalmata and R. valckei sp. probably around its midpoint in R. exilis. Another nov. (Fig. 12A). In three species the two dorsal feature that points towards a close relationship pigment stripes split into four anastomosing stripes: between these two species is the caudal position R. reticulata, R. schaereri (Figs 1A, 7D) and of the testes, which is almost unique within the R. polyrhabdota.InR. cincta the situation is some- Eukalyptorhynchia. what intermediate. This last species is also unique in showing a concentration of pigment around the seminal vesicles (indicated with arrows on Fig. 9A) THE FALCATA-GROUP (‘enveloppe pigmentaire’, see Brunet, 1969). The The falcata-group includes two unpigmented species following eight species lack pigmentation: R. acuta, from the western Mediterranean: R. acuta (Figs 3D, R. colpaerti sp. nov., R. falcata, R. martensi, R. gal- 4E, F) and R. falcata (Figs 3C, 4G). These species lica, R. hibernica, R. licyae sp. nov. and R. verveckeni differ from the other species of Rogneda by the fact sp. nov. Karling (1953) mentioned a blackish paren- that stylet A consists of only one plate, and by a chymal pigment for R. westbladi that disappears particular construction of stylet B. According to when the specimen is ﬁxed. This pigment is not Brunet (1979), stylet A of R. acuta can easily be comparable with the dorsal pigment stripes of other derived from that of R. falcata ‘by a simple regression’

© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153, 1–28 6 T. J. ARTOIS of the proximal part of the stylet. Although such a R. hibernica and R. verveckeni is longer than in the process could be imagined, it is highly speculative. other two species and has a relatively longer proximal Actually, stylet A of R. acuta cannot be compared with part. It is distally split into two parts: a broad and a that of any of the other species of Rogneda, while narrow rectangular part, both with a serrated distal stylet A of R. falcata resembles stylet B of R. reticu- rim, in R. hibernica (the most narrow considered a lata (Fig. 8E), but with the distal part of plate B1 spine); and a very narrow rectangular part with a lacking in R. falcata. Stylet B of the species of the serrated distal rim and a hook-shaped part (the spine) falcata-group differs from that of all other species of in R. verveckeni. R. hibernica therefore is the only Rogneda (except the species of the capulata-group) in species within the hibernica-group that has the distal that plate B2 is attached to the proximal end of plate ends of both parts of plate B1 blunt and serrated. B1. In R. falcata, plate B1 is sickle-shaped and plate Therefore, the evidence for sister group relation- B2 is digitiform. In R. acuta, stylet B is more of the ships within this group is somewhat contradictory. typical form, with plate B2 being plate-shaped and With regard to stylet A, R. hibernica from the North distally forked. Sea, R. verveckeni from the Atlantic coast of Spain and R. vangronsveldi from the Mediterranean most resemble each other, whereas the detailed mor- THE HIBERNICA-GROUP phology of the stylet B suggests a close relationship This is a morphologically homogeneous group, con- between the two Mediterranean species R. vangrons- taining four species without dorsal subepidermal veldi and R. licyae. R. verveckeni has a stylet B with pigment stripes: R. hibernica (Figs 5A, 7A), R. licyae a somewhat intermediate morphology between this of (Figs 5C, 7C), R. verveckeni (Fig. 5D) and R. van- R. hibernica on the one hand and R. vangronsveldi/ gronsveldi (Figs 5B, 7B) (the last species is coloured R. lyciae on the other. brownish by a parenchymal pigment and has some dorsal subepidermal pigment at the level of the eyes). Stylet A of these species consists of a relatively simple THE POLYRHABDOTA-GROUP plate A1, and a plate A2 that is very long and narrow The polyrhabdota-group consists of two species with a and distally splits into two sharp ends (the shortest of reticulate dorsal pigmentation: R. polyrhabdota which is considered a spine, indicated with an arrow (Fig. 6A) and R. schaereri (Figs 1A, 6B, 7D, E). They in Fig. 5A–D). In R. hibernica, R. vangronsveldi and have a stylet B resembling that of the steueri-group R. verveckeni plate A1 evenly tapers towards a (see further below). This stylet is almost twice as long pointed distal end, whereas in R. licyae it suddenly in R. schaereri as in R. polyrhabdota. The proximal narrows and is arrowhead-shaped distally (indicated part of plate B2 is sturdier in R. polyrhabdota. More- with a dashed arrow in Fig. 5C). Plate A2 and over, plate B2 is relatively smaller in R. polyrhabdota its spine in R. hibernica, R. vangronsveldi and (± 40% of plate B1; ±50% in R. schaereri). Stylet A in R. verveckeni are very slender. The spine is much both R. polyrhabdota and R. schaereri consists of one shorter in R. hibernica (as compared with the length plate only, which at ﬁrst sight suggests a close of plate A2) than it is in R. vangronsveldi and relationship between the two species. However, in R. verveckeni. Plate A1 in R. verveckeni is relatively R. polyrhabdota it is clearly a plate A1, with the longer than in R. vangronsveldi, with a longer proxi- distal part serrated and a thickened area at about its mal part, and has a serrated distal rim, which was midpoint, at the place where in other species plate A2 not observed in any of the other three species. Plate is attached. This is much less obvious in R. schaereri. A2 and the spine are much more sturdy in R. licyae. Here stylet A is a simple plate, with a stronger The construction of stylet B is also very typical for the proximal end, more resembling plate A2 of the species members of this group, consisting of a plate B1 that of the steueri-group. Therefore, the resemblance is distally split and a plate B2 that is a simple between the two species could as well be superﬁcial rectangular plate with a serrated distal rim. This and not represent a common descent. stylet is very similar in R. vangronsveldi and R. licyae. In these two species, plate B1 has a short proximal part and its distal part is split into a rect- THE STEUERI-GROUP angular part with a serrated rim and a hook-shaped The largest of the groups is the steueri-group, with six part, which I refer to as a spine (this is also the case species: two without pigment (R. gallica, R. martensi), in R. verveckeni; see further below). Plate B2 of both two with two dorsal stripes (R. steueri, R. palula), one species is rather straight and broad, even slightly with a reticulate dorsal pigment pattern (R. reticu- broader distally than proximally. Stylet B differs in lata)andR. cincta (Fig. 9A), which has a pigment the two species in that the proximal part of plate B1 pattern in between that of R. steueri and R. reticu- is much broader in R. vangronsveldi. Plate B1 of lata. The species of this group all have relatively

© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153, 1–28 REVISION OF ROGNEDA 7 simple stylets, with A and B each consisting of two tical, with an S-shaped plate A1, distally ending in a simple plates. The pigmented species of this group all serrated rim and with a rather broad hook at about its have a serrated distal end of plates A1 and A2, which midpoint. Plate A2 ends in a point, just before the is not the case in R. gallica (Fig. 8B) and R. martensi distal end of plate A1. In R. westbladi (Fig. 11A), plate (Figs 8C, 9D, E). For R. steueri (Figs 8F, 9C) this is A1 is not S-shaped and does not have a hook, and plate difficult to assess, but at least plate A2 seems to have A2 is relatively much shorter. Stylet B of the three a serrated distal end. R. steueri is characterized by species is very complex and apparently both consti- very slender plates A1 and A2. R. palula (Figs 8D, 9B) tuting plates have grown together at some places, can easily be recognized by the fact that plate B1 which is most apparent in R. anglica. The construc- distally splits into two parts. The stylets of R. cincta tion of this stylet is, however, comparable in the three (Fig. 8A) and R. reticulata (Fig. 8E) resemble each species. It consists of a curved plate B1, which distally other very closely. There are, however, some clear is rather narrow and rectangular, ending in a serrated differences. Plate A1 of R. cincta shows a bifid distal rim. In R. colpaerti and R. westbladi this plate has two end, with a rectangular and a more pointy part, the extensions (x and y in Figs 11A, B, 12D). Extension y latter one I consider a spine (arrow in Fig. 8A). In is apparently lacking in R. anglica, but extension x R. reticulata the proximal part of plate B2 is elon- could be represented in this species by the broad plate gated and curved towards the distal point of the stylet that seems to be connected to plate B2 (x? in Figs 11C, (arrow in Fig. 8E), which is not the case in R. cincta. 12C). In all three species, plate B2 projects extensively This feature is very conspicuous, even on living speci- at one side of plate B1 (z in Figs 11, 12C, D) and in mens and at lower magnification. Together with the R. colpaerti and R. anglica also a smaller projection at differences in pigmentation, these features make it the other side is observed (arrow in Figs 11B, C, 12C, very easy to distinguish the two species. R. martensi D). Plate B2 in R colpaerti also has a sharp-ending and R. gallica, both unpigmented, can easily be dis- projection near the proximal rim of plate B2 (indicated tinguished from each other by the differences found in by a dashed arrow in Figs 11B, 12D). In all three stylet A, which has a very broad and square proximal species, B2 is relatively broad with a broad, serrated part in R. martensi. distal rim. Thus, also in this group there is conflicting evidence for establishing sister group relationships. The detailed construction of stylet B, especially the THE TRIPALMATA-GROUP presence of the projections at either side of the distal The tripalmata-group includes two Mediterranean part of plate B1, and the lack of pigment suggest a species with two dorsal pigment stripes: R. tripal- close relationship between the two Mediterranean mata (Fig. 10C) and R. valckei (Figs 10A, B, 12A, B). species: R. colpaerti and R. westbladi. The almost Both species are characterized by the unique con- identical construction of stylet A, however, suggests a struction of stylet A, which is no more than a slender relationship between R. anglica (from the Atlantic) plate carrying a spine. This spine is situated at about and R colpaerti. the midpoint of the stylet in R. tripalmata, more As already mentioned in the Introduction and as is distally in R. valckei. Stylet B is of the normal con- obvious from the discussion above, it is extremely struction. Because of the lack of material of R. tripal- difficult to make correct assessments of homology mata, a direct comparison between the species on the between the different (parts of the) stylets within the detailed structure of stylet B is impossible. Judging taxon Rogneda. In some particular cases, however, from the drawing by Ax (1959; here Fig. 10C), R. tri- it seems fairly straightforward to make homology palmata differs from R. valckei in that its plate B1 assessments. For instance, stylet B of the species lacks a serrated distal rim and also lacks the small within the westbladi-group is so typical that it hardly hook just proximal from this rim. Moreover, R. tripal- can be imagined not to be homologous among the mata also lacks a hook on plate B2. These observa- three species of this group (of course the logical tions should, however, be confirmed on new material deduction is that stylets A of these three species are of R. tripalmata. also homologous with each other). The same applies for stylet A in the capulata-group, stylets A and B in the hibernica-group and probably stylet B in the THE WESTBLADI-GROUP falcata-group. Whether stylets A (and B) of the two The westbladi-group consists of three species: two species of the polyrhabdota-group are homologous unpigmented ones (R. colpaerti and R. westbladi), and with each other is doubtful, a problem discussed one with two dorsal pigment stripes (R. anglica). above. Within the steueri-group, homologies are also Stylet A of these species is again very simple, consist- very unclear, and there is little evidence with which ing of two simple plates. In R. anglica (Figs 11C, 12C) to assess homologies. Stylets A of R. cincta, R. palula and R. colpaerti (Figs 11B, 12D, E) it is almost iden- and R. reticulata are possibly homologous; they all

© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153, 1–28 8 T. J. ARTOIS have both plates A1 and A2 with a serrated distal rim. Also plate A2 of R. steueri has the distal rim serrated, but (apparently) plate A1 does not. None of the plates of stylet B has a serrated rim in these species, although Karling (1953) draws a plate B2 with a serrated rim in R. steueri. I, however, cannot confirm this observation, not even after studying the same material he studied. The other two species of this group, R. martensi and R. gallica, have a stylet A with plates that do not have a serrated distal rim. More- over, stylet A of R. gallica is clearly almost identical to stylet B of R. steueri, which suggests a homology of these two structures. It becomes even more difficult to search for homologies between the stylets of species of different groups. Karling (1953) considered stylet A of R. hibernica homologous with stylet A of R. steueri, because of the ‘whip-like’ appearance of what I now call plate A2. However, stylet B of the species of the hibernica- group has plates B1 and B2 with a serrated distal rim, thus completely comparable with stylet A of, for example, R. cincta or R. reticulata. It is almost impossible to assess homologies between groups with one or two aberrant stylets (falcata-group, tripalmata-group) and the other groups. As already mentioned, there is some resem- Figure 2. R. minuta: prostate stylet type III (from the blance between stylet A of R. falcata and stylet B of neotype). R. reticulata. If one imagines plate B1 of R. reticulata without its distal part (i.e. the part distally from the junction with B2), it would be almost identical to Uterus enters the common genital atrium anteriorly, stylet A of R. falcata. Whether this reflects a common separately from the female duct. origin of the two structures remains, however, an open question. Type species: Rogneda minuta Uljanin, 1870 (by original monotypy). TAXONOMIC ACCOUNTS ROGNEDA MINUTA ULJANIN, 1870 In this section, the species are categorized based on (FIGS 2, 4A, B) stylet-morphology groupings as discussed in the ‘Comparison of Species’ section, starting with Rogneda minuta Uljanin, 1870: 22–23, plate 5 fig. 7; R. minuta. The groups are arranged alphabetically. Karling, 1953: 349–350, 352, 359, 361, 364, 366–367; Within each group the species are also arranged Ax, 1959: 119, 164; Brunet, 1969: 217; Evdonin, 1977: alphabetically. 11, 242–243, 387–388; Brunet, 1979: 103–106, figs 2–4. Macrorhynchus minutus von Graff, 1882: 327–328; Pereyaslawsewa, 1892: 282–284, plate 5 fig. 33, plate ROGNEDA ULJANIN, 1870 10 fig. 63g, plate 11 fig. 66a–k. Diagnosis: Polycystididae with a syncytial epidermis. Polycystis minuta von Graff, 1905: 133–136, plate 4 Gonads paired. Gonopore situated ventrally. With a figs 10–18; von Graff, 1913: 335–336, fig. 303; Stein- prostate vesicle type III, with at least its distal end in böck, 1933: 14. between the two stylets. Prostate vesicle and ejaculatory duct enter the male atrium together through a Known distribution: The Black Sea near Sebastopol small penial papilla. Seminal vesicles paired. No (Ukraine), in algae (Uljanin, 1870; von Graff, 1882, other glands present in the male system. Copulatory 1905; Pereyaslawsewa, 1892). Adriatic Sea, Rovinj organ proper consisting of a prostate stylet type III. (Croatia), algae (Steinböck, 1933). Bay of Marseilles Accessory stylet type I present or not. Male bursa (France), Posidonia fields near to the Amphioxus- present or not. Female bursa absent. A large bundle of sand of the ‘Plateau des Chèvres’ (Brunet, 1979) glands opens in the junction of the two oviducts. (type locality).

IDENTIFICATION KEY 1. Male system with only one stylet. Stylet consisting of two plates proximally connected to each other in a complex manner (Figs 2, 4A, B). The longest plate is 133–166 mm long, the shortest one 110–116 mm. Male bursa lacking...... R. minuta – Male system with two separate stylets (stylet A and stylet B) (Fig. 1D). Male bursa present (Fig. 1B, D)...... 2 2. Animals without dorsal pigment lines. Both stylets consisting of two plates (Fig. 1C). Stylet B consisting of a broad plate B1, which distally splits into a rectangular plate and a spine, and a broad plate B2 which is more or less rectangular and has a serrated distal rim. Stylet A with a simple plate A1 and a plate A2 that also splits into two parts, the shortest one considered a spine (Figs 5, 7A–C) (species of the hibernica-group)...... 3 – Animals with or without pigment lines. Stylets not as described...... 6 3. Plate A1 with an arrowhead-shaped distal point (indicated with a dashed arrow in Fig. 5C). Plate A2 with a very broad and triangular spine (indicated with an arrow in Fig. 5C)...... R. licyae sp. nov. – Plate A1 very slender, evenly tapering to a very sharp distal end. Plate A2 slender and sharp-ending, with a rather long and slender spine, which distally ends in a sharp point...... 4 4. Plate A1 125 mm long, with a blunt spine at its midpoint (Fig. 5B). Plate A2 very long, 158 mm, and slender. Also the spine very long (indicated with an arrow in Fig. 5B), 79 mm, and connected to plate A1 somewhat at its midpoint. Plate B1 92 mm long, with a very short and broad proximal end and a 39-mm-long spine. Plate B2 59 mm long...... R. vangronsveldi sp. nov. – Plate A1 127–155 mm, without a spine. Plate A2 94–117 mm. Plate B1 115–145 mm long, with a relatively longer and more slender proximal part. Plate B2 relatively narrow and curved ...... 5 5. Plate A2 with a short spine (19–30 mm) (indicated with an arrow in Fig. 5A), which is connected to plate A2 near its distal tip. Plate B1 distally splits into a narrow (the spine) and a broad rectangular part, each with a serrated distal rim. Plate B2 short, 37–42 mm ...... R. hibernica – Plate A2 with a long spine (68 mm) (indicated with an arrow in Fig. 5D), which is connected to plate A2 at about its midpoint. Plate B2 distally splits into a narrow rectangular part with a serrated distal rim and a hook-shaped part (the spine). Plate B2 long, 70 mm...... R. verveckeni sp. nov. 6. One of the two stylets (stylet B) very complex, with the different plates difficult to distinguish. Plate B1 distally a narrow rectangular plate, with a serrated distal rim. Plate B2 very broad, with a broad serrated distal rim and a large and extended proximal part. Stylet A simple, with plate A1 slender, ending in a serrated distal rim. Plate A2 with a sharp distal end (Figs 11, 12C–E) (species of the westbladi-group) ...... 7 – Stylets not as described ...... 9 7. Animals with two dorsal pigment lines. Plate A1 105–140 mm long, somewhat S-shaped, with a 21–38-mm-long spine ± at its midpoint. Plate A2 46–63 mm long (± 50% of plate A1). Plate B1 mildly curved, 123–157 mm long. Plate B2 very broad, almost square, 44–88 mm long (Fig. 11C) ...... R. anglica – Animals not pigmented. Stylets not as described...... 8 8. Plate A1 98 mm long, without a spine. Plate A2 a simple hook, 34 mm long (± 35% of plate A1). Plate B1 very curved, 116 mm long, with a blunt ending (x in Fig. 11A) and a sharp ending projection (y in Fig. 11A) at about its midpoint and at opposite sides. Proximal part of plate B2 very extensive (z in Fig. 11A), distal part rather thin but very broad. Plate B2 42 mm long ...... R. westbladi – Plate A1 94–98 mm long, somewhat S-shaped, with a ± 20-mm-long spine ± at its midpoint. Plate A2 44–62 mm long (± 50% of plate A1). Stylet B resembling that of R. westbladi. Plate B1 strongly curved, 93–113 mm long, with two projections at about its midpoint and at opposite sides (x and y in Fig. 11B). Plate B2 56–73 mm long, with a sharp-ending projection near its proximal rim (indicated with a dashed arrow in Fig. 11B) ...... R. colpaerti sp. nov. 9. Animals with two or four, sometimes anastomosing reddish-brown dorsal pigment lines (Figs 1A, 7D, 9A, 12A)...... 10 – Animals without dorsal pigment lines...... 19 10. Stylet A very aberrant, consisting of at least one broad, almost oval plate, with a serrated distal rim (plate A1) and a hook-shaped plate (plate A2) (Figs 3A, B, 4D). Stylet B very simple and of the normal construction (Figs 3A, B, 4C). Testes situated caudally (species of the capulata-group)...... 11 – Stylet A not as described...... 12 11. Stylet A with a very broad plate A1, divided into at least two plates; both formed somewhat like a baker’s shovel. Both with a serrated distal rim. The ﬁrst is 72–78 mm long; the second 90 mm. Possibly there is a third plate, also with a serrated rim and also about 90 mm long. Plate A2 triangular, 54–63 mm long (Fig. 3B)...... R. capulata – Stylet A with a less broad plate A1, also with a serrated distal rim. Plate A2 with the distal end hook-shaped. Both plates 60 mm long (Figs 3A, 4D)...... R. exilis

12. Pigmentation consisting of two dorsal pigment lines (Fig. 12A). One of the stylets (stylet A) only consisting of one slender plate, with a spine (Figs 10, 12B) (species of the tripalmata-group)...... 13 – Pigmentation in two lines or four anastomosing lines (Figs 1A, 7D, 9A). Such a stylet lacking ...... 14 13. Stylet A 135–175 mm long, with a distal spine 39–42 mm long. Plate B1 150–192 mm long with a pointed distal end, which is serrated at one side. Just proximal to the serrated edge there is a sharp, 18–19-mm-long hook. Plate B2 100–117 mm long, with 23–30-mm-long hook at about its midpoint (Figs 10A, B, 12B)...... R. valckei sp. nov. – Stylet A ±167 mm long, with the spine attached more precisely at its midpoint. Length of the spine unknown. Detailed morphology of stylet B unknown, plate B1 195 mm long, plate B2 114 mm (?) (Fig. 10C)...R. tripalmata 14. Pigmentation consisting of four lines, which anastomose and form a somewhat reticulating pattern (Figs 1A, 7D, 9A). One of the stylets (stylet A) consisting of one plate only, the other one consisting of two plates (species of the polyrhabdota-group)...... 15 – Pigmentation as above or consisting of two dorsal stripes. Both stylets consisting of two plates each. One stylet (stylet A) mostly with both plates with a serrated distal rim. Plate A1 slender. The other stylet (stylet B) with plates ending in a blunt or sharp point, but never serrated (pigmented species of the steueri-group) ...... 16 15. Stylet A a rather slender plate, 107–129 mm long, with a narrow linguiform distal end. Stylet B sturdy. Plate B1 159–207 mm long. Plate B2 relatively long, 83–98 mm (± 50% of plate B1) (Figs 6B, 7E)...R. schaereri sp. nov. – Stylet A a rather broad plate, 117–118 mm long, with a thickened band at about its midpoint. Its distal rim rather broad. Stylet B sturdy. Plate B1 119–126 mm long (up to 163 mm according to Ax, 1959). Plate B2 relatively short, 47–51 mm (± 40% of plate B1) (Fig. 6A) ...... R. polyrhabdota 16. Pigmentation consisting of two dorsal pigment stripes. One of the stylets (stylet B) with a plate B1 that distally splits into a blunt and a sharp ending part (the spine). Plate B1 80–107 mm long, spine 47–71 mm long. Plate B2 of the normal construction, 52–77 mm long. Plate A1 89–104 mm, plate A2 48–63 mm (Figs 8D, 9B)...... R. palula – Not as above...... 17 17. Pigmentation consisting of two dorsal pigment stripes. Stylet A with a long (104 mm) and slender plate A1, which distally ends in a narrow, serrated rim. Also plate A2 very long and slender, 62 mm. Stylet B very simple, with a slender plate B1, 73–82 mm long, and a sharp ending plate B2, 36–47 mm long (Figs 8F, 9C)...... R. steueri – Pigmentation consisting of four dorsal pigment stripes, sometimes anastomosing and forming a reticulate pattern...... 18 18. With four dorsal pigment stripes. Pigment also concentrated as an envelope around the seminal vesicles (indicated with arrows in Fig. 9A). Plate A1 69–71 mm long, with a distal spine that is 24–26 mm long. Plate A2 35–39 mm long. Stylet B with a slender plate B1, 60–69 mm long and spatulate-ending plate B2, 30–39 mm long, without a hook-shaped proximal part. (Fig. 8A)...... R. cincta – With four dorsal pigment stripes that anastomose and form a reticulate pattern. No pigment envelope around the seminal vesicles. Plate A1 without a spine, 77–92 mm long. Plate A2 34–51 mm long. Plate B1 77–96 mm long. Plate B2 36–45 mm long, with a large and procurving proximal end that forms a hook (indicated with an arrow in Fig. 8E)...... R. reticulata 19. One of the two stylets (stylet A) consisting of only one plate. The other one (stylet B) of an aberrant construction (Figs 3C, D, 4E–G) (species of the falcata-group)...... 20 – Both stylets consisting of two plates (unpigmented species of the steueri-group)...... 21 20. Stylet A a simple plate, 71 mm long, with a slender proximal part and a somewhat broader distal part, which ends in a sharp point. At the transition between the two parts there is a 21-mm-long projection that points distally. Plate B1 sickle-shaped, 88 mm long. Plate B2 somewhat S-shaped, 44 mm long, attached to the proximal tip of B1 (Figs 3C, 4G)...... R. falcata – Stylet A is a simple hook, 72–74 mm long, with a rounded proximal end and somewhat leaf-shaped distally. Plate B1 120–121 mm long, with a circular proximal end. A spine 62–65 mm long is attached to its proximal part. Both plate B1 and spine end in a sharp point. Plate B2 attached to the proximal part of plate B1. It is 44 mm long and distally broadens and splits into two, making the whole plate B2 resemble a kite’s tail (Figs 3D, 4E, F)...... R. acuta 21. Plate A1 with a very broad, square proximal part, distally narrowing abruptly to a stalk with a blunt end. Plate A2 a simple hook, attached to the proximal end of plate A1. Plate A1 70 mm long, plate A2 36 mm. Plate B1 116 mm long, with a rectangular proximal part, evenly narrowing towards a distal sharp point. Plate B2 rectangular, 68 mm long (Figs 8C, 9D, E)...... R. martensi sp. nov. – Both stylets of a relatively simple construction. Plate A1 a simple, slender plate with two blunt ends, 98 mm long. Plate A2 57 mm long, attached to the middle part of plate A1 and ending in a sharp point. Stylet B of the same construction as stylet A, but plate B2 broader, more diamond-shaped. Plate B1 109 mm long, plate B2 56 mm long (Fig. 8B)...... R. gallica

Material examined: Neotype (a whole mount) (SMNH, (considered part of plate A1 and called A1a; see ‘Com- no. 3210). One other whole mount (SMNH, no. 40859) parison of Species’) is ovoid, somewhat in the form of a from Marseilles. One serially sectioned specimen baker’s shovel (‘Brotschaufel’ of Karling, 1953), with (SMNH, no. 40857) from Rovinj. a serrated distal rim. Beneath this plate there is a second, longer plate of the same form (A1b) and also Remarks: Apart from accounts by von Graff (1882, with a serrated rim. Just distal from this plate, a third 1905) and Karling (1953), this species is extensively serrated rim can be seen, probably of a third plate of described by Brunet (1979). In the male genital atrium the same form. Alternatively, this rim could be a fold of there is only one hard part, which consists of two long the rim of the second plate. Both A2b and the addi- plates. One of these plates has a sharp distal end, the tional serrated rim can only be seen in one of the other being much broader distally, and somewhat specimens. The fourth plate (plate A2) is somewhat gutter-shaped. Proximally they are connected to each triangular to rectangular, with one of its sides strongly other in a very complex way. Measured from the pro- curved (the ‘Deckel’ of Karling, 1953). My measure- ximal tip, the sharp-ending plate is 133–166 mm long ments refer to these three plates, and all are taken (N = 2), the broad-ending plate 110–116 mm(N = 2). from the proximal point of the stylet to the distal rim Both Evdonin (1977) and Brunet (1979) doubted of the plate considered: A1a, 72–78 mm(N = 2); A1b, whether Macrorhynchus minutus discussed by Pere- 90 mm; A2, 54–63 mm(N = 2). Karling (1953) yaslawsewa (1892) (Polycystis minuta of von Graff, adequately described stylet B of this species. It consists 1905, 1913) is indeed the same species as Rogneda of two plates. Plate B1 [(sb + db2 of Karling (1953)] is minuta described by Uljanin (1870). The poor drawing 93–107 mm long (N = 2) and curved. Plate B2 is by Pereyaslawsewa (1892) indeed casts some doubts, attached to plate B1 in the proximal half of B1. Its and only shows a stylet that consists of two parts that proximal end strongly protrudes at one side of plate B1 are connected to each other proximally. This problem [(pb2 of Karling (1953)]. It is 52–54 mm(N = 2) long. can, however, only be solved by sampling at the exact same localities as Pereyaslawsewa and von Graff did in ROGNEDA EXILIS BRUNET, 1979 the hope of finding specimens that could contradict or (FIGS 3A, 4C, D) confirm this synonymy. For the time being, I follow the Rogneda exilis Brunet, 1979: 108, figs 8–10. view of Karling (1953), who considered all these records to refer to one and the same species. Known distribution: Bay of Marseilles (France), sandy to muddy gravel (85–95 m) (Brunet, 1979).

CAPULATA-GROUP Material examined: The holotype (a whole mount) ROGNEDA CAPULATA KARLING, 1953 (SMNH, no. 3208). (FIG. 3B) Rogneda capulata Karling, 1953: 350, 352–356, 358– Remarks: Brunet (1979) described stylet A as two 361, 363, 365–366, figs 3, 9–12, 29, 30; Ax, 1959: 121; plates extending from a common triangular proximal Brunet, 1969: 208, 210, 217, 220; Evdonin, 1977: 14, part: one in the form of a hook, the other distally very 122, 243, 245, figs 2, 112.1; Brunet, 1979: 105, 108. broad and round. Proximally, the hook-shaped plate has a second extension, which is also pointed and Known distribution: Adriatic Sea, Island of Ciovo, which was not mentioned by Brunet (1979). Although Split (Croatia), Amphioxus-sand (Karling, 1953) (type it is clear that there are two plates, it is almost locality). Several localities in the Bay of Marseilles impossible to say which plate is A1 and which is A2 (Brunet, 1969). because of the aberrant construction of the stylet. The proximal extension of the hook-shaped plate could be Material examined: The lectotype (a whole mount) homologous to the extension of plate A2 observed in (SMNH, no. 2677). One whole mount and three other species of Rogneda at the place where A2 is serially sectioned specimens from Croatia (SMNH, attached to A1. If this is true, the hook-shaped plate is nos. 43562-65). A2, the rounded one plate A1. Both plates are 60 mm long. Stylet B is very simple with a 104-mm-long, curved and distally pointed plate B1 and an equally Remarks: Stylet A is of an aberrant construction, with simple, 45- m-long, slightly curved plate B2. a narrow proximal point [sa of Karling (1953)] and a m plate-shaped distal stalk. According to Karling (1953), the distal part (his da) only consists of one broad plate FALCATA-GROUP with a basal lid (‘Deckel’). However, close inspection ROGNEDA ACUTA BRUNET, 1979 reveals the presence of three plates that lie on top of (FIGS 3D, 4E, F) each other and are different in length. The first Rogneda acuta Brunet, 1979: 106–107, figs 5–7.

Figure 3. Stylets of the representatives of the capulata- and the falcata-group. A, R. exilis (from the holotype). B, R. capulata (from the lectotype). C, R. falcata (from the holotype). D, R. acuta (from the holotype). Stylet A is indicated by an A, stylet B by a B. Plates B1 and B2 are indicated by a 1 and a 2, respectively. Scale bar = 25 mm.

Known distribution: Bay of Marseilles (France), 62–65 mm long. Plate B2 (pb2 of Brunet, 1979) is medium sand, Amphioxus-sand and sandy to muddy 44 mm long (N = 2) at its longest, and distally splits gravel (8–50 m) (Brunet, 1979). into two pointed ends, resembling a kite’s tail.

Material examined: The holotype (a whole mount) ROGNEDA FALCATA BRUNET, 1965 (SMNH, no. 3206) and one whole mount (SMNH, (FIGS 3C, 4G) no. 3207). Rogneda falcata Brunet, 1965: 153–158, plate 9 fig. 5, plate 10 figs 1–3; Brunet, 1969: 208, 212, 217, 221; Remarks: Stylet A was adequately described by Evdonin, 1977: 122, 243, 245, figs 112.5–6; Brunet, Brunet (1979). It is 72–74 mm long (N = 2). Plate B1 is 1979: 107. 120–121 mm long (N = 2). It carries a shorter spine, which is attached to its proximal half and also ends in Known distribution: Bay of Marseilles (France), a sharp point (db2 of Brunet, 1979). This spine is Amphioxus-sand (8–18 m) (Brunet, 1965).

Figure 4. Stylets of R. minuta and of representatives of the capulata- and falcata-group. A, B, R. minuta (A from the neotype; B specimen from Marseilles). C, D, R. exilis [ C, stylet B; D, stylet A (both from the holotype)]. E, F, R. acuta [E, stylet A; F, stylet B (both from the holotype)]. G, R. falcata (from a specimen from Marseilles). Scale bars = 20 mm.

Material: The holotype (a whole mount) (SMNH, no. Rogneda hibernica Karling, 1953: 350–352, 354–355, 3046). One whole mount (MNHN-P). 357–360, 363–364, 366–367, ﬁgs 4–8, 15, 17, 23–24, plate 1A, D, E, plate 2A, D, E; Brunet, 1969: 213, 217; Remarks: Brunet (1965) described this species Evdonin, 1977: 90, 104, 122, 244–245, ﬁgs 54, 112.10; adequately. Stylet A consists of one plate only. At the Schockaert, Jouk & Martens, 1989: 24. midpoint of this plate there is a small projection, which obviously corresponds to the proximal projection of plate A2 in other species of the taxon. Stylet A Known distribution: Known from several localities in the English Channel, the Irish Sea and the Irish [sa + da1 of Brunet (1965)] is 71 mm long, the projec- Atlantic coast (Southern, 1936; Karling, 1953). Also tion [pa1 of Brunet (1965)] 21 mm. Stylet B consists of two plates with plate B2 attached to the proximal tip recorded from the sublittoral in the Netherlands of plate B1. Plate B1 is 88 mm long, plate B2 44 mm. Delta area (Schockaert et al., 1989).

HIBERNICA-GROUP New localities: Ostend (Belgium), Spuikom, coarse ROGNEDA HIBERNICA (SOUTHERN, 1936) sand from the eulittoral (type locality). KARLING, 1953 (FIGS 1D, 5A, 7A) Material examined: The neotype (a whole mount) Polycystis hibernica Southern, 1936: 45, 59–61, (SMNH, no. 7154). Two whole mounts from Ostend ﬁgs 2–5. (HU). Three whole mounts and ﬁve serially sectioned

Figure 5. Stylets of the representatives of the hibernica-group. A, R. hibernica (from the neotype). B, R. vangronsveldi sp. nov. (from the holotype). C, R. licyae sp. nov. (from the holotype). D, R. verveckeni sp. nov. (from the holotype). Stylet A is indicated by an A, stylet B by a B. For each stylet plate 1 is indicated by a 1, plate 2 by a 2. Scale bar = 50 mm. Arrows: see text. specimens from Port Erin (Isle of Man) (SMNH, nos. further referred to as ‘spine of plate B1’, the broader 43554-561). one I consider the distal end of plate B1 proper. As already mentioned by Karling (1953), plate A2 is also Remarks: Karling (1953) described this species distally split. I will refer to the shortest of both these adequately. On the newly collected material I distal ends as ‘spine of plate A2’ (indicated with an observed a bifurcated distal part of plate B1, some- arrow in Fig. 5A), the longer one considered to be the thing not mentioned by Karling (1953), but easily distal end of plate A2 proper. recognizable on the drawings by Southern (1936). Our measurements on the hard parts are as follows Both distal ends are serrated and one end is much [abbreviations in parentheses are from Karling broader than the other one. The narrowest one is (1953)]. Stylet A: plate A1 (sa + da1): 127–155 mm

(x¯ = 144; N = 5); plate A2 (da2): 94–117 mm(x¯ = 109; long, and distally reaches as far as the distal tip of

N = 5); spine of plate A2 (da2 partly): 19–30 mm plate A1. It evenly tapers towards its distal end,

(x¯ = 24 mm; N = 5). Stylet B: plate B1 (sb + db1): 115– ending rather bluntly. It carries a strong and broad, 145 mm(x¯ = 134; N = 3); spine of plate B1: 34–44 mm 37-mm-long spine somewhat at its midpoint (indi-

(x¯ = 39; N = 2); plate B2 (db2): 37–42 mm (x¯ = 40; cated by an arrow in Fig. 5C). The proximal part of N = 4). These measurements correspond to the mea- plate B1 is rather narrow. Distally plate B1 splits surements given by Karling (1953). into a narrow, rectangular part, which is serrated I deem the designation of a neotype necessary for distally, and a broad, sharp-ending spine. The plate this species, considering the complexity of species is 96 mm long, the spine 29 mm. Plate B2 has a recognition within the taxon Rogneda. The type series rather broad distal end with a serrated distal rim. not longer exists or has never existed, as Southern It is 60 mm long and 25 mm broad at its broadest. At (1936) based his description on live material only, the place where it is attached to plate B1 it shows keeping no slides. Both Karling’s (1953) material a small projection pointing towards the distal tip of (from southern England, the Isle of Man and the the stylet. northern English Channel), and my material (from Belgium) are a considerable distance from the origi- ROGNEDA VANGRONSVELDI SP. NOV. nal locality where Southern (1936) collected his speci- (FIGS 5B, 7B) mens (Valentia Island and Erin, both south-west coast of Ireland). Therefore, I choose a specimen with Holotype: A whole-mounted specimen, Corsica the most clearly observable hard parts as neotype. Its (France), in the harbour of the marine biological hard parts are practically identical to those depicted station STARESO, between 4 and 8 m deep, in coarse by Southern (1936: ﬁg. 5). sand, 19.v.1983 (SMNH, no. 7156).

Other material: Observations on live animals. ROGNEDA LICYAE SP. NOV. (FIGS 5C, 7C) Etymology: Dedicated to Professor J. Vangronsveld, Holotype: A whole-mounted specimen, Corsica head of the Centre for Environmental Sciences of the (France), near Pointe Revellata, sand from a sublit- Hasselt University. toral sample (6 m) off a small sandy beach, 18.ix.1983 (SMNH, no. 7155). Diagnosis: Species of Rogneda with a pigmented area at the level of the eyes. Stylet A with a long Other material: Observations on live animals. and slender plate A1, ending in a sharp point, proximally with a short, blunt spine. A2 long and Other locality: Corsica (France), near Ocellutia, sand slender, ending in a sharp point, with a relatively from 19 m deep, 19.x.1982. long and slender spine that is attached to A2 at about its midpoint and also ends in a sharp point. Etymology: In memory of the late Licy Oeyen, an Stylet B with a plate B1 with a blunt, broad proxi- excellent biologist and even better friend. mal end and a distal end that consists of a rectangular plate with a serrated distal rim and a broad Diagnosis: Unpigmented species of Rogneda. Plate A1 spine. proximally rectangular, distally narrowing abruptly to a slender stalk, which ends in an arrowhead- Description: The specimen studied is 0.9 mm in whole shaped point. Plate A2 a simple hook, with a broad mount. The live animals show an opaque black area and triangular spine. Plate B1 with a slender proxi- around the eyes and a light brown body colour, the mal end and a distal end that consists of a rectangu- latter owing to the presence of parenchymal pigment. lar plate with a serrated distal rim and a broad spine. In the squashed and damaged holotype, pigment Plate B2 broad, distal rim serrated. grains are found outside the body, probably the grains responsible for the black area around the eyes Description: The whole-mounted specimen is 0.7 mm observed in the live animal. long and is unpigmented. Stylet A has a plate A1 that is 125 mm long and very Plate A1 is rather long and broad. Distally it slender. It tapers distally to a very sharp point. More abruptly narrows to a stalk, which ends in an or less at its midpoint it carries a very short (23 mm), arrowhead-shaped point (indicated with a dashed blunt spine. Plate A2 is connected to the short proxi- arrow in Fig. 5C). Proximally it is rather narrow, mal part of A1. It is very long (158 mm) and slender, ending bluntly. The plate is 121 mm long. Plate A2 is tapering to a sharp distal end. At about its midpoint connected to the proximal half of A1. It is 88 mm it carries a very long (79 mm), slender spine (indicated

© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153, 1–28 16 T. J. ARTOIS by an arrow in Fig. 5B), which also evenly tapers Known distribution: Sea of Marmara (Turkey), ﬁne towards its distal and sharp end. Stylet B is almost sand from the littoral zone in Pendik and Florya (Ax, identical to that of R. licyae. Plate B1 has a very 1959). broad and blunt proximal end. Distally it splits into a rectangular part that is serrated distally, and a broad, New locality: Nea Epivates (near Thessalonica, sharp-ending spine. The plate is 92 mm long, the spine Greece), ﬁne sediment (2.5 m deep), left from the 39 mm. Plate B2, as in R. licyae,is59mm long and jetty, 6.viii.2002 (coll. HU). 27 mm broad at its broadest.

Material examined: Two whole mounts from Nea ROGNEDA VERVECKENI SP. NOV. Epivates. (FIG. 5D) Holotype: A whole-mounted specimen, Ferrol (Galicia, Remarks: The species was extensively described by Spain), muddy–sandy sediment from the sublittoral Ax (1959). The description of the hard parts by Ax near the pump of the marine biological station, (1959) is accurate, but none of the plates is distally 26.viii.2006 (SMNH, no. 7157). open and funnel-shaped as he describes them. Stylet A consists only of plate A1, and shows a strong Etymology: Dedicated to Erwin Vervecken, triple transverse band more or less at its midpoint. Distally world champion cyclo-cross (2001, 2006, 2007). it is rather delicate, with a serrated distal rim. It is 117–118 mm long (N = 2) in the specimens from Diagnosis: Species of Rogneda without pigment. Greece, comparable with the measurements given by Stylet A with a long and slender plate A1, distally Ax (1959). Plate B1 is proximally slightly broader ending in very narrow, blunt serrated end. Plate A2 than stylet A. Its distal half is rectangular and also long, ending in a sharp point, with a relatively long, has a serrated distal rim. Plate B2 is attached to plate triangular spine that is attached to A2 at about its B1 almost at its midpoint and has a somewhat spatu- midpoint and also ends in a sharp point. Stylet B with late distal part. It is not serrated. Its proximal part is a plate B1 with a long and relatively slender proximal very thick and appears very sturdy (pb2 of Ax, 1959). end and a distal end that consists of a rectangular In the specimens from Greece plate B1 is 119–126 mm plate with a serrated distal rim and a broad spine. long (N = 2); plate B2 47–51 mm(N = 2). This is Plate B2 relatively narrow and curved. slightly smaller than was mentioned by Ax (1959) (135–163 mm for the whole stylet). Description: The specimen studied is 0.7 mm in whole mount (animal very contracted). Stylet A has a plate A1 that is slender and 153 mm ROGNEDA SCHAERERI SP. NOV. long. It tapers distally to a narrow, blunt end. This (FIGS 1A, 6B, 7D, E) end is serrated, consisting of four finger-like projec- Holotype: A whole-mounted specimen, Lignano (Italy), tions. Plate A2 is connected to plate A1 more or less fine sand taken on a very flat beach at low tide, from at about one-third the length of the latter from the the water edge down to ± 8 cm deep; oxidized top proximal tip. It is rather long (107 mm) and slender, layer (± upper 2 cm). (SMNH, no. 7158). tapering to a sharp distal end. At about its midpoint it carries a long (68 mm), triangular spine (indicated by an arrow in Fig. 5D), which is slightly curved and Paratypes: Two whole-mounted specimens, same data also evenly tapers towards its distal sharp end. Stylet as for the holotype (HU, nos. 344–45). B has a plate B1 with a relatively long proximal end. Distally it splits into a narrow rectangular part that Etymology: Named after Dr Lukas Schärer (Univer- is serrated distally, and a broad, sharp-ending spine. sity of Basel), who kindly gave me the material of this The plate is 141 m long, the spine 56 m. Plate B2 is m m species. 70 mm long, somewhat curved, and distally ends in a serrated rim. Diagnosis: Species of Rogneda with four anastomosing, brown, dorsal pigment stripes. Stylet A very POLYRHABDOTA-GROUP simple, consisting of only one plate. Stylet B with two ROGNEDA POLYRHABDOTA AX, 1959 plates. Plate B1 rather long, with a serrated distal (FIG. 6A) rim. Plate B2 attached to plate B1 at about its mid- Rogneda polyrhabdota Ax, 1959: 48, 119–123, 145, point, distally ending in a blunt point. Plates B1 and figs 107–115; Brunet, 1969: 212, 217, 219, 221. B2 close to each other.

Figure 6. Stylets of the representatives of the polyrhabdota-group. A, R. polyrhabdota (from a specimen from Greece). B, R. schaereri sp. nov. (from the holotype). Stylet A is indicated by an A, stylet B by a B. Plates B1 and B2 are indicated by a 1 and a 2, respectively. Scale bar = 25 mm.

Description: Animals ± 0.9 mm long (measured on between the island of Jarre and the coast (8–10 m), whole mounts). Four irregular, anastomosing longitu- and ‘Pierre de Joseph’, near the island of Plane, fine dinal dark brown pigment stripes mark the dorsal sand (17 m) (Brunet, 1969). side of the body. Stylet A is very simple, 107–129 mm long (x¯ = 115; N = 3). It consists of one plate only, which has a thick Material examined: The holotype (a whole mount) proximal end. It is somewhat bent, with a linguiform, (SMNH, no. 3045). The paratype (a whole mount) slightly serrated distal end. Stylet B is of the typical (MNHN-P). Three whole-mounted specimens construction, with plate B2 always lying next to plate (MNHN-P). B1 and attached to B1 at about its midpoint. Plate B1 is 159–207 mm long (x¯ = 187; N = 3), with a serrated Remarks: Brunet (1969) described this species distal end. Plate B2 tapers towards its distal end, adequately. The brown pigment around the seminal ending in a blunt point. It is 83–98 mm long (x¯ = 92; vesicles is typical of this species (arrows in Fig. 9A). N = 3). The proximal end of each stylet shows many In the holotype the strong distal spine of plate A1 superficial folds. appears not to be spatulate, as was described by Brunet (1969), but pointed (arrow in Fig. 8A). Plate STEUERI-GROUP B2, by contrast, is very broad and distally spatulate, ROGNEDA CINCTA BRUNET, 1969 not as pointed as was drawn by Brunet (1969). My (FIGS 8A, 9A) measurements on the hard parts of this species are Rogneda westbladi ssp.? Brunet, 1965: 135. as follows [the abbreviations in parentheses are Rogneda cincta Brunet, 1969: 208, 214–221, figs these of Brunet (1969)]. Stylet A: plate A1 (sa + da1): 7–9, 13; Evdonin, 1977: 12, 14, 104, 122, 244–245, 69–71 mm(x¯ = 70; N = 3), spine of plate A (part of figs 2, 112.9; Brunet, 1979: 105. da1): 24–26 mm(x¯ = 25; N = 3), plate A2 (pa2 + da2): 35–39 mm(x¯ = 38; N = 3). Stylet B: plate B1

Known distribution: Bay of Marseilles (France), (sb + db1): 60–69 mm(x¯ = 66; N = 4), plate B2

Amphioxus-sand from the ‘Plateau des Chèvres’ (pb2 + dbx): 30–39 mm(x¯ = 34; N = 4).

Figure 7. Stylets of the representatives of the hibernica-group and the habitus and stylet of R. schaereri sp. nov. A, R. hibernica (from the neotype). B, R. vangronsveldi sp. nov. (from the holotype). C, R. licyae sp. nov. (from the holotype). D, E, R. schaereri sp. nov. [D, habitus; E, stylets (both from the holotype)]. Scale bars = A–C, E = 20 mm; D = 100 mm.

ROGNEDA GALLICA AX, 1956 Known distribution: The lagoon of Lapalme (Etang de (FIG. 8B) Lapalme), near La Franqui (French Mediterranean Rogneda westbladi gallica Ax, 1956: 5, 146–148, 165, coast), in pure fine sand, sometimes with some 189, 200, fig. 36; Brunet, 1969: 212; Evdonin, 1977: detritus (Ax, 1956) (type locality). Bay of Marseilles 243, 245, fig. 112.4. (France), near la Mounine (east of the ‘Île de Rogneda westbladi Brunet (1969): 208, 210–212, Maire’), fine sand with some detritus (12 m) (Brunet, figs 2, 14. 1969).

Figure 8. Stylets of the representatives of the steueri-group. A, R. cincta (from the holotype). B, R. gallica (from a specimen from Marseilles). C, R. martensi sp. nov. (from the holotype). D, R. palula (from the lectotype). E, R. reticulata (from the holotype). F. R. steueri (from the neotype). Stylet A is indicated by an A, stylet B by a B. For each stylet plate 1 is indicated by a 1, plate 2 by a 2. Arrows: see text. Scale bar = 25 mm.

Material examined: One sectioned specimen from the not allocate them to one of the two subspecies recog- Etang de Lapalme, designated lectotype (SMNH, no. nized, but suggested that the Marseilles population 5988). One whole mount from Marseilles (SMNH, might represent a taxonomic entity of its own. He no. 43574). mentioned that stylet A of the specimens from Marseilles is identical to that described by Ax (1956) Remarks: The whole mount I studied is one of the for R. w. gallica, but indicated some differences in the specimens already studied and discussed by Brunet detailed morphology of stylet B between his speci- (1969), who referred to them as R. westbladi.Hedid mens and Ax’s (1956) description. These differences

Figure 9. Habitus and stylets of representatives of the steueri-group. A, habitus of R. cincta (from the holotype). B, stylets of R. palula (from the lectotype). C, stylets of R. steueri (from the neotype). D, stylet B of R. martensi sp. nov. (from the holotype). E, stylet A of R. martensi sp. nov. (from the holotype). Arrows: see text. Scale bars = A = 50 mm; B–E = 20 mm.

© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153, 1–28 REVISION OF ROGNEDA 21 are the presence of a short, strong spine at the base Etymology: Dedicated to Dr Paul Martens, who col- of db1 (distal part of plate B1) and, more importantly, lected the material of this and many other kalypto- the fact that in the specimens from Marseilles, db2 rhynchs present in the collections of the HU. (the distal part of plate B2) is somewhat longer than db1 (plate B1) and has a rounded distal end. The first character is, however, difficult to assess, and was not Diagnosis: Unpigmented species of Rogneda. Plate A1 even visible on the specimen I studied. Ax (1956) did proximally very broad, square, distally very narrow, not mention or draw this spine, but it could still be ending bluntly. Plate A2 a simple, curved hook, present. The relative lengths of the two parts of stylet attached more or less at the midpoint of plate A1. B is another doubtful character. Not only is it very Plate B1 curved, ending in a very sharp distal point. difficult to judge from Ax’s (1956) drawings which one Plate B2 attached to the midpoint of plate B1, rect- is the longest, but even in the material from Brunet angular, distally ending in a broad and blunt end. himself the relative position of the two distal tips appears not as constant as he indicated. Finally, the distal parts of these stylets are often very difficult to Description: The only specimen available is unpig- study, and a more or less rounded end is also a mented, about 0.9 mm long (measured on the whole doubtful character. These observations suggest that mount). It is not in an excellent condition, but enough the specimens studied by Brunet (1969) in fact belong detail could be seen to mark it undoubtedly as a new to Ax’s (1956) R. w. gallica. species. Interpretation of the construction of plate A1 According to Ax (1956), stylet A of R. w. gallica is difficult. At first sight it appears to be very broad differs from that of the nominal subspecies in the and square proximally, but narrows abruptly at about relative length of the two subunits: plates A1 and A2. its midpoint to a rectangular plate that ends bluntly. In R. w. gallica plate A2 is rather long and its distal It is 70 mm long. Plate A2 is attached to plate A1 more point projects beyond the distal point of plate A1, or less at the place where the latter narrows. It is a whereas in R. w. westbladi plate A2 is much shorter. simple 36-mm-long, curved hook. However, another Moreover, the distal end of plate A1 is serrated in interpretation could be that plate A2 partly has come R. w. westbladi, but smooth in the specimen of loose from A1 because of the squeezing and is con- R. w. gallica I studied. Stylet B is also very different nected to it by a thin, very narrow clasp. If this is the between the two taxa, a feature barely discussed by Ax case, plate A1 is very narrow, and the square that can (1956). In R. w. gallica, plate B1 is somewhat curved, be seen in the holotype is not a plate (as in the first with a blunt distal point. Plate B2 is rather simple, interpretation), but a hole. As such, stylet A, recon- with a somewhat diamond-shaped distal point, and is structed, would look very much like that of R. gallica. attached to plate B1 more or less at its midpoint. On However, the observations on the live animal (only Ax’s (1956) drawings this plate is depicted as very very slightly squeezed) suggest that the first inter- narrow, almost threadlike. Possibly he only saw the pretation is the correct one. thick rim of the plate, and not the whole plate. Stylet Stylet B is relatively simple and consists of a 116- BofR. w. westbladi is much more complex and indeed mm-long plate B1, which slightly bends at about its not reminiscent of that of R. w. gallica (see further midpoint. Proximally it ends bluntly; distally it ends under R. westbladi). These large and fixed differences in a sharp point. Plate B2 is attached almost at the between the two taxa indicate that the French popu- midpoint of plate B1, and is a rather broad rectan- lation (R. w. gallica) belongs to a separate species from gular plate. It is 68 mm long. the population from the Adriatic Sea (R. w. westbladi). Therefore, I consider R. w. gallica a species of its own: R. gallica Ax, 1956. ROGNEDA PALULA BRUNET, 1969 Measurements on the hard parts of the whole (FIGS 8D, 9B) mount: stylet A: plate A1: 98 mm; plate A2: 57 mm. Rogneda palula Brunet, 1969: 208, 216–221, figs 10– Stylet B: plate B1: 109 mm, plate B2: 56 mm. 11, 15; Evdonin, 1977: 14, 122, 244–245, figs 2, 112.12. Rogneda patula (incorrect subsequent spelling) ROGNEDA MARTENSI SP. NOV. Watson, 2001: 224, figs 20.23, 20.27, table 20.1. (FIGS 8C, 9D, E) Holotype: A whole-mounted specimen, south-west Sulawesi (Indonesia), Kajangan, coral sand from the Known distribution: Bay of Marseilles (France), eulittoral, 22.x.1984 (SMNH, no. 7159). Amphioxus-sand from between the ‘Château d’If’ and the island of Ratonneau (14–16 m) (Brunet, 1969) Other material: Observations on live material. (type locality).

New localities: Roscoff (France), very coarse sand larger plate A1 measuring 90, 90 and 92 mm, respec- from between rocks in the Fucus serratus zone in the tively (77–86 mm in the others), but they do not show ‘Green Island Channel’ (3.vi.2007). Roscoff (France), any difference as to the other measurements. coarse sand from a tide pool in front of the marine station, near to the long jetty (6.vi.2007). ROGNEDA STEUERI (STEINBÖCK, 1933) KARLING, 1953 Material examined: One whole mount, designated (FIGS 8F, 9C) lectotype (SMNH, no. 3047). One other whole mount Polycystis steueri Steinböck, 1933: 14–15, figs 6–7. the type locality (MNHN-P). Three whole mounts Rogneda steueri Karling, 1953: 350, 356, 358–359, from Roscoff. 361, 363, 365–367, figs 2, 13, 19–22, plates 1B–C; Ax, 1959: 121, 123; Brunet, 1969: 217, 221; Evdonin, Remarks: The species is adequately described by 1977: 14, 122, 244–245, figs 2, 112.13; Brunet, 1979: Brunet (1969). Plate B1 is distally split into a blunt- 105. ending plate and a sharp ending one, the latter further referred to as spine of plate B1. The stylets of Known distribution: Adriatic Sea, Split (Croatia), the specimens from Roscoff are identical in shape to Ciovo, Amphioxus-sand (Karling, 1953) (type locality). those of the specimens from the Mediterranean, but Adriatic Sea, Rovinj (Croatia), Cuvi, in coarse sand are slightly larger. Therefore, measurements on speci- and shell-gravel (Steinböck, 1933). mens of both areas are given separately. Measure- Material examined: One whole mount from Split, des- ments on the two specimens from Marseilles: plate A1 ignated neotype (SMNH, no. 5987). Two other whole (sa + da1): 89–90 mm; plate A2 (da2): 48–51 mm, plate mounts from Split, on the same slide as the holotype. B1 (sb + part of db1): 80–82 mm; spine of plate B1 One whole mount, on the same slide as the lectotype (part of db1): 47–56 mm; plate B2: 52–56 mm. Mea- of R. capulata (SMNH no. 2677). One other whole surements on the three specimens from Roscoff: mount from Split (SMNH, no. 43579). plate A1: 100–104 mm(x¯ = 102); plate A2: 57–63 mm (x¯ = 61), plate B1: 102–107 mm(x¯ = 104); spine of plate Remarks: Stylet A of this species is very typical, with B1: 70–71 mm(¯x = 71); plate B2: 71–77 mm(x¯ = 74). a long and slender plate A1, distally ending in a sharp R. palula is the only species of Rogneda that has point, and a very slender plate A2, which distally both a Mediterranean and an Atlantic population. projects beyond plate A1. In some of the specimens, plate A2 has a blunt, serrated distal end, in others it ends in a sharp point. We have measured lengths of ROGNEDA RETICULATA BRUNET, 1969 104 mm for plate A1 (N = 2); 62 mm for plate A2 (FIG. 8E) (N = 1). The length of plate A1 (sa + da1) is compa- Rogneda reticulata Brunet, 1969: 208, 212–214, 216– rable with the length mentioned by Karling (1953), 219, 221, figs 4–6, 14; Evdonin, 1977: 14, 104, 122, whereas the length of plate A2 (pa2 + da2) is ± double 244–245, figs 2, 112.11; Brunet, 1979: 105. the length mentioned by Karling. Judging from his drawings it is clear that the lengths he mentioned for Known distribution: Several localities in the Bay of plate A2 are erroneous. Stylet B is also of a very Marseilles, in Amphioxus-sand and fine and muddy simple construction. Plate B1 is simple, proximally sands (4–17 m) (Brunet, 1969). somewhat broader than distally. Attached to it is plate B2, which also is a simple plate, distally ending New locality: Cerbère (France), Anse de Terrimbou, at the same level as plate B1. In some specimens it 10 m deep; coarse clean sand, 12.iv.2002 (coll. HU). seems to end bluntly, with a serrated edge. In other specimens it seems to end in a sharp point (as in Material: The holotype (a whole mount) (SMNH, no. Fig. 8F). Plate B1 is 73–82 mm long (N = 2), which is 3048). Two whole mounts from Pière de Joseph (bay of slightly smaller than mentioned by Karling (1953). Marseilles) (MNHN-P). Live observations and three Plate B2 is 36–47 mm long (N = 2), which is in com- whole mounts from Cerbère. plete correspondence with the measurements given by Karling (1953). Remarks: Brunet (1969) has described this species adequately. Our measurements (on four specimens) TRIPALMATA-GROUP are as follows: plate A1: 77–92 mm(x¯ = 85); plate A2: ROGNEDA TRIPALMATA (BEKLEMISCHEV, 1927) 34–51 mm(x¯ = 45), plate B1: 77–96 mm(x¯ = 85), plate KARLING, 1953 B2: 36–45 mm(x¯ = 41). These measurements are com- (FIG. 10C) pletely in accordance with those given by Brunet Polycystis tripalmata Beklemischev, 1927: 195–198, (1969). The specimens from Cerbère appear to have a 206–207, plate 1 figs 13–16.

Figure 10. Stylets of the representatives of the tripalmata-group. A, B, R. valckei sp. nov. (A from a paratype, B from the holotype). C, R. tripalmata (after Ax 1959, no scale bar provided). Stylet A is indicated by an A, stylet B by a B. Plates B1 and B2 are indicated by a 1 and a 2, respectively. Scale bar = 25 mm.

Rogneda tripalmata Karling, 1953: 350, 352, 356, (‘pb2’ of Karling, 1953). If the plate is measured 359–361, 367; Ax, 1959: 119, 121–124, 146, 161; axially, as I did for the other species, it probably will Brunet, 1969: 213, 217, 219, 221; Evdonin, 1977: 8, be smaller. 101, 104–105, 121, 244–246, ﬁgs 62, 112.8.

ROGNEDA VALCKEI SP. NOV. Known distribution: The Black Sea, the bight of (FIGS 10A, B, 12A, B) Odessa (Ukraine), in sand (Beklemischev, 1927). Sea Holotype: A whole-mounted specimen, Corsica of Marmara, ﬁne sand from the littoral zone on (France), near Ocellutia, sand from 10 m deep, Heybeli Island (Turkey) (Ax, 1959). 25.x.1982 (SMNH, no. 7160).

Paratypes: Two whole-mounted specimens, same data Material examined: None. as for the holotype (HU, nos. 346–47).

Remarks: See the discussion by Karling (1953) and Ax Other material: Observations on live animals. (1959). The lengths of the stylets given previously (Beklemischev, 1927; Ax, 1959) are as follows (my Other localities: Several localities near Calvi (Corsica, notation used): stylet A: 154–180 mm; plate B1: 190– France): off Pointe Revellata, sand from 6 m deep, 200 mm; plate B2: 114 mm. The measurement of B2, 25.xi.1984; sublittoral sample from the bay of Calvi, however, includes the proximal extension of plate B2 sand from 20 m deep, 30.i.1984.

Etymology: Dedicated to Professor R. Valcke, head of Material examined: Observations on live animals the research group Molecular and Physical Plant from Kristineberg. The lectotype (a whole mount) Physiology of the UHasselt. (SMNH, no. 2676). Four whole mounts (on the same slide) and three serially sectioned specimens Diagnosis: Species of Rogneda with two dorsal, lon- from Plymouth (SMNH, nos. 43575–78). Two gitudinal pigment stripes. Stylet A represented by a whole mounts and three serially sectioned speci- single long, narrow plate with a small distal spine. mens from Kristineberg (coll. HU). Two whole Stylet B consisting of two plates. Plate B1 distally mounts from Galicia (coll. HU). Three of the nine ending in a sharp point, with a small hook not far whole-mounted specimens do not allow accurate from the distal point. Rim between hook and distal measuring. point serrated. Plate B2 connected to plate B1 above the midpoint of B1, distally ending in a sharp point. Remarks: Stylet A is relatively simple, and the basic A small hook present at about its midpoint. construction can still be recognized. Plate A1 is more Description: Animals 0.7–1 mm long, measured on or less S-shaped. It corresponds to Karling’s (1953) whole mounts. Dorsally, at both sides of the body, two sa + the broad ending part of da2. It is 105–140 mm stripes of brown pigment stretch from the level of the long (x¯ = 118; N = 6). At about its midpoint, plate A eyes towards the caudal end of the body. shows a rather broad, sharp-ending hook (da1 of Stylet A is very simply built. It consists of a rela- Karling, 1953), which is 21–38 mm long (¯x = 26; N = 6). tively slender plate that distally carries a small spine. Plate A2 is connected to plate A just proximally from It is 135–175 mm long (¯x = 158; N = 3), with a rela- this hook. It is curved and lies adjacent to the distal tively broad and blunt proximal end and a more part of plate A1. It ends in a sharp point (part of da2 pointed and much narrower distal end. The spine is of Karling, 1953). It is 46–63 mm long (x¯ = 56; N = 6). connected to this main plate between the second and Stylet B is much more complex than is depicted by the distal third. It is 39–42 mm long (x¯ = 41; N = 3), Karling (1953), and apparently the distal parts of and can end in a sharp or blunt point, depending on both constituting plates have grown together at some the specimen. Plate B1 is 150–192 mm long. Its distal places, making it rather difficult to distinguish them. end is serrated at one side, and ends in a sharp point. Plate B1 [sb + part of db1 of Karling (1953)] is rather Just proximal from the serrated rim, the plate shows long, somewhat curved, and distally ends in a rather a 18–19-mm-long hook (N = 2). Plate B2 is attached to narrow, oblong part with a serrated distal rim. Plate plate B1 somewhat above the midpoint of B1 (in one B2 [other part of Karling’s (1953) db1] is connected to of the paratypes it has come loose: see Fig. 10B). It plate B1 more or less at its midpoint, and proximally has a straight distal end, and tapers towards a sharp protrudes extensively at both sides of B1; the projec- point. At about its midpoint it shows a relatively tions are called pb2 (z in Figs 11C, 12C) and db3 broad hook. Plate B2 is 100–117 mm long (x¯ = 108; (indicated with an arrow in Figs 11C, 12C), respec- N = 3); the hook is 23–30 mm long (N = 2). tively, by Karling (1953). Distally it is very broad, and has a relatively long serrated part. At the opposite WESTBLADI-GROUP part of this serrated rim, a broad distal extension of ROGNEDA ANGLICA KARLING, 1953 plate B1 (x? in Figs 11C, 12C) seems to be connected (FIGS 11C, 12C) to plate B2. The lengths of the different parts are as follows: plate B1: 123–157 mm(x¯ = 136; N = 6); plate Rogneda anglica Karling, 1953: 350, 354–356, 358– B2: 44–88 mm(¯x = 69; N = 6). 361, 363–366, ﬁgs 1, 14, 18, 31–32, plates 1F, 2B, C, F; Brunet, 1965: 153; Brunet, 1969: 217, 219–220; Evdonin, 1977: 14, 101, 110, 122, 243, 245, ﬁgs 2, ROGNEDA COLPAERTI SP. NOV. 112.2. (FIGS 1B, 11B, 12D, E) Holotype: A whole-mounted specimen, Corsica Known distribution: The English Channel, several (France), bay of Calvi, off Pointe Revellata, sand from localities near Plymouth (type locality, no more a sublittoral sample (± 40 m), 26.xi.1982 (SMNH, details available) and Millport (UK), mostly in loamy no. 7161). bottoms (10–36 m) (Karling, 1953).

New localities: Kristineberg (Sweden), Gullmarfjord, Paratypes: Two whole-mounted specimens, same data Essvik, muddy sediment, 20 m deep (coll. HU). Ferrol as for the holotype (HU, nos. 348–49). (Galicia, Spain), muddy–sandy sediment from the sublittoral near the pump of the marine biological Other material: Observations on live material by station (26.viii.2006). Dr P. Martens.

Figure 11. Stylets of the representatives of the westbladi-group. A, R. westbladi (from the lectotype). B, R. colpaerti sp. nov. (from the holotype). C, R. anglica (A, B stylets A and B, respectively, from the same specimen from Kristineberg, B′ stylet B from another specimen from Kristineberg). Stylet A is indicated by an A, stylet B by a B. For each stylet plate 1 is indicated by a 1, plate 2 by a 2. Scale bar = 25 mm. Abbreviations: x–z, see text. Arrows: see text.

Etymology: Dedicated to Professor J. Colpaert, distal point. Stylet B complex. Plate B1 with a broad mycologist at the HU. and short proximal part, distally splitting into a short pointed part and a rectangular part with a serrated Diagnosis: Unpigmented species of Rogneda. Stylet A distal rim. Small triangular tooth present at base of simple. Plate A1 narrow, rounded proximally, serrated rectangular part. Plate B2 S-shaped, with a serrated distally, with a strong hook almost at its midpoint. distal rim and one strong, thick wall. Narrow pointed Plate A2 a simple curved plate, ending in a sharp projection present at base of plate B2.

Figure 12. Habitus and stylets of R. valckei sp. nov. and stylets of representatives of the westbladi-group. A, B, R. valckei sp. nov. (A, habitus from a paratype, B stylets from a paratype). C, stylets of R. anglica (from a specimen from Kristineberg). D, E, R. colpaerti sp. nov. (D, stylet B. E, stylet A). Abbreviations: x–z, see text. Arrows: see text.

Description: Animals unpigmented, rather small, very simple, with a broad proximal end, tapering 0.4–0.7 mm long in whole mounts. towards its distal end, which is a very sharp point. It Stylet A is rather simple. Plate A1 is narrow, with is 44–62 mm long (x¯ = 51; N = 3). Stylet B is very a rounded proximal and a serrated distal end. It is complex, and can be best described on the holotype. S-shaped, 94–98 mm long (x¯ = 96; N = 3). At about its Plate B1 is curved and has a short and broad proxi- midpoint, near to the place where plate A2 is mal part, which distally splits into a short and attached, it forms a relatively broad hook, about pointed projection (x in Figs 11B, 12E) and a longer, 20 mm long in the holotype and in one of the rectangular part with a serrated distal rim. At the paratypes (not visible in the other paratype, probably base of the rectangular plate, at the opposite side of because of the poor quality of the slide). Plate A2 is the projection, there is a small triangular tooth (y in

Figs 11B, 12E). Measured from the proximal point of ACKNOWLEDGEMENTS the plate to the serrated rim of the rectangular plate I wish to thank the following persons and institu- the stylet is 113 mm long, with the projection 25 mm tions: Dr L. Schärer (University of Basel) for kindly long and the tooth 12 mm. Plate B2 makes an providing the material of R. schaereri collected by him S-shaped curve. Distally it ends in a serrated rim, in Lignano; Dr P. Martens for providing all the mate- which can only be followed for a short distance. At the rial he collected during several field trips to Corsica opposite side of the pointed projection of plate B1, and during the Snellius expedition to Indonesia; plate B2 shows a narrow and very pointed projection Dr P. Jouk (Antwerp Zoo) for providing sublittoral (indicated with a dashed arrow in Figs 11B, 12E). The sand samples from Cerbère; Professor E. Schockaert proximal part of plate B2 extends beyond the curve of for providing the material of R. verveckeni and R. an- plate B1 (z in Figs 11B, 12E) and has a smaller glica from Galicia and the new material of R. hiber- projection at the opposite side of this (indicated with nica; Professor E. Schockaert, Dr Wim Willems an arrow in Figs 11B, 12E). Plate B2 is 73 mm long. In (SMNH) and Mr Wouter Willems (University of the paratypes, the overall shape of stylet B is easily Ghent) for providing the new material of R. polyrhab- recognizable but some details are somewhat obscured dota; Mr Bart Tessens for providing the specimens of and difficult to find. The measurements in these R. palula from Roscoff; Dr S. Boström for sending all specimens are as follows (some of the measurements the material of Rogneda present in the collections of were impossible to take on one or both of the speci- the Swedish Museum of Natural History; Mrs M-N. mens): plate B1: 93 and 111 mm; projection of B1: Hellouet and Dr G. Boucher for sending the material 25 mm; tooth of B1: 11 mm; length B2: 56 mm. of Rogneda present in the collections of the National History Museum in Paris; Dr N. Watson for the ROGNEDA WESTBLADI KARLING, 1953 critical reading of the manuscript and correction of the English text; Mrs N. Steffanie for the preparation (FIG. 11A) of the collected material. The research was financed Rogneda westbladi Karling, 1953: 350, 354, 358–359, by FWO-VL (project G.0235.02). 361, 363–367, figs 16, 25–28; Ax, 1956: 147, 189; Brunet, 1969: 217, 220. Rogneda westbladi westbladi Ax, 1956: 148, 189; REFERENCES Brunet, 1965: 153; Brunet, 1969: 212, 221. Artois T, Schockaert E. 2003. 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