OBSERVATIONS on SETAL MORPHOLOGY in the PHYLLODOCIDAE (Polychaeta: ANNELIDA), with SOME TAXONOMIC CONSIDERATIONS

BULLETIN OF MARINE SCIENCE, 48(2): 530-543,1991

OBSERVATIONS ON SETAL MORPHOLOGY IN THE PHYLLODOCIDAE (pOLYCHAETA: ANNELIDA), WITH SOME TAXONOMIC CONSIDERATIONS

Danny Eibye-Jacobsen

ABSTRACT With few exceptions (principally within the genera Eteone. Hesionura and Paranaitis), the setae of the Phyllodocidae have been regarded as being so homogenous as to lack taxonomic value. Descriptions and figures in the literature have been both imprecise and confusing. An attempt has here been made to carry out systematic observations of the setae in several different genera, using both differential interference contrast light microscopy and scanning electron microscopy. The results indicate that subtle but important differences do exist, and may be very helpful in a more precise delimitation of genera within the Phyllodocidae. These conclusions are compared to remarks in the literature in an attempt to bring some order to setal observations. Potentially valuable characters seem to be present; along these lines, the need for further research is pointed out, above all a thorough revision of the genus Eulalia.

The setae are usually of great taxonomic importance in the identification of polychaetes. However, setae have not been used to any great extent as taxonomic characters in the Phyllodocidae. This is because the compound spinigers, which are the dominant setal type in the family, have been perceived as having a very homogenous morphology (apart from a few exceptions noted below). In this paper the Phyllodocidae are defined as in Fauchald (1977), and exclude the Lopadorhyn- chidae, Pontodoridae and lospilidae. This definition corresponds to the Phyllo- docinae ofUschakov (1972). The few notably characteristic setae fall into one offour groups: 1) Compound spinigers in the genera Eteone s. lat., Hesionura and Paranaitis, 2) Simple noto- podial setae in connection with biacicularity in Notophyllum and Austrophyllum, 3) Hooks on the anterior segments in Chaetoparia, and 4) Long, simple, neuro- podial setae on epitokous individuals of certain species of Mystides, Pseudomys- tides, Protomystides and Eulalia. As this study is concerned solely with the compound spinigers, examples of only the first of these four items will be treated here. In this paper I demonstrate that subtle generic differences exist, making it possible to specify a basic setal morphology for most genera. The differences are primarily to be found in details of the distal portion of the shaft. The shaft is usually widened apically and bears a rostrum on either side (anterior and posterior), each of which may bear teeth of varying number, size and shape (U schakov, 1972, fig. 22D). Bergstrom (1914) included the morphology of the distal portion of the shaft (his "Verschlusseinrichtung") as one of 14 important systematic characters in the Phyllodocidae. However, apart from his remarks on Eteone s. lat. and Paranaitis (as Anaitis), there is no indication of a variation of systematic value within Phyl- lodocidae s. str. Uschakov (1972) did not use setal characters to any great extent in delineating genera or species, apart from the four exceptions mentioned above. A couple of quotations (Uschakov, 1972: 33) describe the problems discussed here: " ... con- ventional microscopes do not permit detailed study of the fine structure of the setae, particularly of details of the structure and serration of the rostrum, which may prove of great importance for the identification of species and lower taxonomic units." "The use of the electron scanning microscope will probably give 530 EIBYE-JACOBSEN: SETAL MORPHOLOGY IN PHYLLODOCIDAE 531

Figure I. Photographs of setae of Eumida sp. 1. Scale equals IO ~m_ A: Normal light microscopy. B: Differential interference contrast microscopy_ much new information. The drawings of the setae of Phyllodocidae are usually very diagrammatic and not exact .... " An example of how difficult it is to make out the details of the setal articulation in light microscopes may be seen in Banse's changing opinion as regards Pterocirrus macroceros (Grube, 1860). Describing animals from the Adriatic Sea, he characterized the rostrum as being "wenig ausgezogener ... kaum geziihnt" (Banse, 1959a: 424). Banse and Nichols (1968: 225) gave the following description of the same structures on specimens from Puget Sound: "rounded with numerous small spines which are slightly bent inward." Banse may, of course, have been making observations on two different species, but it is quite probable that differences in technique or thoroughness of observation account for the different descriptions. Standard light microscopy is not generally adequate to observe the fine details of the compound setae (compare Fig. lA, B). However, with access to a microscope equipped with differential interference contrast (DIC) optics, I was able to record systematically the morphology of the setal rostrum in over 30 phyllodocid species. This raised the possibility that these structures could yield information, both for practical taxonomic and theoretical phylogenetic work, for phyllodocid genera other than Eteone s. lat., Hesionura and Paranaitis. Using DIe, several of them 532 BULLETIN OF MARINE SCIENCE, VOL. 48, NO.2, 1991 were noted to have a characteristic uniform rostral shape slightly different from that found in other, closely related genera. I later had the opportunity to verify and supplement these observations on four genera (Phyllodoce, Eumida, Sige and Pterocirrus) using scanning electron microscopy (SEM). Subsequent routine comparisons of a large number of different phyllodocid species indicate that supplementary characters, useful at the generic level, are present.

MATERIALS AND METHODS

The material studied came from a variety of sources: M. Lindholm Andersen (MLA); P. Garwood (PG); B. O'Connor (BOC); M. E. Petersen (MEP); E. Stikvoort (ES); Allan Hancock Foundation, Los Angeles (AHF); The Australian Museum, Sydney (AM); Zoologisches Museum, Hamburg (HZM); Musee National d'Histoire Naturelle, Paris (MNHN); National Museum of Natural History, Wash- ington D.C. (USNM); Zoological Museum, University of Copenhagen (ZMUC). The following genera and species were studied using DIC-microscopy (unless otherwise indicated, the specimens were collected by the author): Eteone Savigny, 1820: E. barbata (Malmgren, 1865) (ES: E. Oosterschelde, SW of the Netherlands), E. foliosa de Quatrefages, 1865 (BOC: Galway Bay, W Ireland), E. piela de Quatrefages, 1865 (Roscoff, English Channel, France) and E. syphodonta (delle Chiaje, 1822) (ZMUC: Naples, Italy). Hesionura Hartmann-Schroder, 1958: H. elongata (Southern, 1914) (PG: NW North Sea). Pseudomystides Bergstrom, 1914: P. limbata (de Saint-Joseph, 1888) (Tjamo, W Sweden) and Pseudomystides sp. (undescribed) (Tjamo and Frederikshavn, N Denmark). Phyllodoce Lamarck, 1818: P. citrina Malmgren, 1865 (0resund, E Denmark), P. groenlandica 0rsted, 1843 (0resund), P. longipes Kinberg, 1866 (Tjiirno) and Phyllodocesp. (indeterminable) (MLA: Oman). Paranaitis Southern, 1914: P. kosteriensis (Malmgren, 1867) (BOC: Galway Bay) and P. speciosa (Webster, 1879) (MEP: Bogue Sound, North Carolina). Eulalia Savigny, 1818: E. aurea Gravier, 1896 (Roscoff), E. bilineata (Johnston, 1840) (0resund and Tjiirno), E. mustela Pleijel, 1987 (Tjiirno), E. ornata de Saint-Joseph, 1888 (Roscoff) and E. viridis (Linne, 1767) (0resund and Frederikshavn). Eumida Malmgren, 1865: E. bahusiensis Bergstrom, 1914 (0resund and Tjamo), E. ockelmanni Eibye- Jacobsen, 1987 (0resund and Tjiirno), E. sanguinea (0rsted, 1843) (0resund, Frederikshavn and Tjamo), E. venustissima (Banse, 1959) (MEP: Naples), Eumida sp. 1(under description) (AM: New South Wales) and Eumida sp. 2 (under description) (AM: New South Wales). Sige Malmgren, 1865: S. fusigera Malmgren, 1865 (0resund and Tjamo) and Sige sp. 1 (undescribed) (AM: New South Wales). Pterocirrus Claparede, 1868: P. macroceros (Grube, 1860) (MEP: Naples) and P. marginata (Claparede, 1868) (MEP: Naples). Nereiphylla de Blainville, 1828: N. fragilis (Webster, 1'879) (MEP: Bogue Sound), N. lutea (Malmgren, 1865) (Tjiirno) and Nereiphylla cf. nana (de Saint-Joseph, 1906) (MEP: Naples). Notophyllum 0rsted, 1843: N. foliosum (Sars, 1835) (Tjiirno). The following species were studied using SEM (all species collected by the author during a 2-week stay at Carrie Bow Cay, Belize): Hesionura elongata (Southern, 1914), Phyllodoce cf. erythrophylla (Schmarda, 1861), Eumida sp. 3 (under description), Sige sp. 2 (under description) and Pterocirrus foliosus (Treadwell, 1924). An Olympus Vanox AHB-S research photomicroscope was used for the DIC studies. Photographs were not always of publishable quality, but were suitable for production of line drawings. Almost all observations were carried out with the 40x objective. The reasons for this are twofold. First, it was hoped that significant results for practiCal routine work could be achieved without the extra time involved in employing oil immersion. Secondly, I quickly found that the end of the setal shaft is so thick that meaningful observation at higher magnification was difficult. Specimens for SEM were critical point dried and platinum-coated. Observations were made with a Cambridge Stereoscan 250 MK2.

RESULTS AND DISCUSSION In this paper I deal almost exclusively with the morphology of the distal portion of the shaft to which the appendage of the compound spiniger is attached. How- ever, in a few cases characters of the distal appendage provide interesting supplementary information. For purposes of comparative equivalence, observations were primarily confined to the middle setae of median parapodia, in posterior view. Mounting a detached para podium in posterior view is easy since the setae are postacicular in position. The preacicular lobe is always more strongly developed than the postacicular lobe EIBYE-JACOBSEN: SETAL MORPHOLOGY IN PHYLLODOCIDAE 533 and usually divided into supra- and subacicular lips. Furthermore, dorsoventral orientation of the parapodium may be determined by characters of the setae themselves. According to Bobin (1944: 45, 51) and Uschakov (1972: 32) the "inner," toothed margin of the distal appendage of the seta is turned downward. In reality this is rarely the case as almost all setae have the cutting edge dorsally oriented. The only exceptions I have observed are cases in which the very dorsalmost seta of a given fascicle may have the opposite orientation of the others (seen on a few species of Eumida and Sige). There does not appear to be any significant variation in setal morphology from different parapodia, although the setae of the most posterior parapodia often appear to be rather less differentiated than those on segments further forward (Wilson, 1988: own observations). On the other hand, Hutchings and Murray (1984) were able to observe some differences in the rostral morphology between dorsal and ventral setae within a single fascicle and between setae from anterior, median and posterior parapodia in their description of Compsanaitis injlata. One final problem in correctly evaluating the setal details treated here should be mentioned. Bobin (1944) remarks on the confusion that may arise in deter- mining the form of the rostrum if the orientation of the seta is not correct (i.e., perpendicular to the distal appendage, either anterior or posterior), as for example in a more or less dorsal view. An example of this is given in Figure 2C and D of Sige sp. 2 from Belize, showing the difference in appearance that may result from viewing setae of the same form at different angles. In this regard SEM is superior because the angle of orientation may be controlled and there is no confusion arising from the inner structures, thus allowing concentration on the morphology of the rostrum itself. In the following, the genera of Phyllodocidae will be treated individually, my own results (where applicable) being compared to relevant observations in the literature. Concerning the genera Lugia de Quatrefages, 1865, Pseudeulalia Elia- son, 1962, Chaetoparia Malmgren, 1867, Prochaetoparia Bergstrom, 1914 and Austrophyllum Bergstrom, 1914, which contain about 15 species it may be said that what little information is available is both confusing and contradictory. New, original observations are necessary.

Eteone Savigny, 1820 (44 species) In his revision of the genera Eteone, Hypereteone Bergstrom, 1914, and Mysta Malmgren, 1865, Wilson (1988) pointed out the taxonomic importance of rostral morphology in distinguishing species. The rostrum has the shape of a large, beaklike tooth, which may bear small, basal accessory teeth. The setae are called heterogomph or homogomph, depending on whether the rostrum is so developed on only one or both sides of the distal article. My own observations agree com- pletely with those of Wilson and earlier authors.

Hesionura Hartmann-Schroder, 1958 (8 species) The neuropodium bears four or five setae; each of them has both rostra developed as a large, straight, elongate teeth (bifid setae) and may additionally have a third, mediodorsal tooth of almost the same size (trifid setae). The distribution of these two types within the fascicle is of primary importance in species deter- mination. Concerning details of the setal morphology in Hesionura, Laubier (1967: 586) stated " ... c'est meme Ie seul eU:ment morphologique permettant une identification specifique certaine." Although this is somewhat overstated (see for example the description of H. japonica in Yamanishi, 1980), my own experience 534 BULLETIN OF MARINE SCIENCE, VOL. 48, NO.2, 1991

Figure 2. SEM micrographs. Scale equals 2 ILm. A: Phyllodoce cf. erythrophylla. B: Eumida sp. 3. C: Sige sp. 2. View slightly dorsal. D: Sige sp. 2. View posterior. E: Pterocirrus foliosus. EIBYE·JACOBSEN: SETAL MORPHOLOGY IN PHYLLODOCIDAE 535 indicates that certain identification cannot be made without a detailed study of the setae.

Mystides Theel, 1879 (about 7 species) Blake (1988) illustrated setae of M. borealis Theel, 1879 (fig. 3b) and M. caeca Langerhans, 1880 (fig. 3d), two species that have often been synonymized. He demonstrated that the rostrum of the former has three or more distal teeth, whereas the latter has a single tooth. The difference appears significant, suggesting that use of these features as species characters could be useful in this genus. Hartmann- SchrOder (1983: fig. 3) illustrated the setae of M. bathysiphonicola.

Pseudomystides Bergstrom, 1914 (about 5 species) The distal articulation of the setal shaft of Pseudomystides limbata and P. sp. is very similar (Fig. 3A). These setae are distinctive in having a fan oflarge, blunt- tipped teeth radiating from both the ventral and the dorsal edge of the rostrum and an interruption (here termed a heel) in the edge that joins the shaft to the ventral margin of the rostrum. The setae are also distinguished by having the distal portion of the shaft only slightly swollen, in contrast to the majority of phyllodocid genera. Southern (1914: fig. 18B) showed the setae of P. limbata (as Mystides (Meso- mystides», which closely resemble those of P. sp. (Fig. 3A). He specifically mentioned the "fan" of blunt, apical teeth on the rostrum, but appears to have over- looked the presence of a heel.

Protomystides Czerniavsky, 1882 (about 8 species) Hartmann-Schroder (1979: fig. 25) illustrated the setae of P. uschakovi. The distal portion of the shaft appears to be only slightly inflated, with seven very large, distal teeth of equal size and a row of smaller teeth proximally. As far as can be judged, the seta depicted resembles that of Eumida.

Phyllodoce Lamarck, 1818 (about 95 species) The four species of Phyllodoce studied using DIC showed the same basic morphology of the distal part of the setal shaft. The rostrum resembles a pine cone with the apex turned dorsally (Fig. 3B). The teeth on the rostrum are thinner than in Pseudomystides and distinctly acuminate. The heel is less distinct than in the latter, but visible. Study of the anterior setal rostrum in Phyllodoce shows that, although it is of the same maximum length as the posterior one, its ventral apical teeth are relatively longer. A portion of the anterior rostrum may thus be visible in the light microscope, even if the parapodium is mounted to show a posterior view (Fig. 3C). Furthermore, only a slight rotation of the angle of view is necessary to make the anterior teeth very obvious, giving the appearance of two groups of teeth bending slightly towards one another. This phenomenon may be verified by varying the focus and is sufficiently common and characteristic to deserve note. A similar condition is also found in the genus Eumida, where it is even more prominent (see below). SEM micrographs of the setae of Phyllodoce cf. erythrophylla confirm the observations detailed above (Fig. 2A). No significant variation in morphology within a setal fascicle was found. Pleijel (1988), in a revision of the species of Phyllodoce from northern Europe, pointed out that the use of ordinary compound microscopes is inappropriate for interpreting the small differences in setal morphology within the genus. My own 536 BULLETIN OF MARINE SCIENCE, VOL. 48, NO.2, 1991

A 8 c E

G H

Figure 3. Schematic drawings of distal portion of setal shaft and basal portion of distal appendage in genera ofPhyllodocidae. The setae are drawn at approximately the same size for ease of comparison, as setal width is basically a function ofthe size of the animal, not a generic characteristic. Dorsal edge of setae to the left. All figures drawn on the basis of DIC micrographs. Striations on the shaft and distal appendage have not been drawn to avoid confusion. All views posterior. A: Pseudomystides sp. B: Phyllodoce groenlandica. Only elements strictly belonging to the posterior side of the seta are shown. C: Phyllodoce groenlandica. The actual aspect of the seta as seen in the light microscope, showing elements of the anterior rostrum. D: Eulalia aurea. E: Eulalia bilineata. F: Eumida bahusiensis. G: Sige fusigera. H: Pterocirrus macroceros. I: Nereiphylla lutea. J: Notophyllumfoliosum. EIBYE-JACOBSEN: SETAL MORPHOLOGY IN PHYLLODOCIDAE 537 observations agree with this; however, the setae are sufficiently characteristic to be useful in delineating the genus. Phyllouschakovius armigerum Blake, 1988 has, according to Blake's figure 6q (verified on type material; USNM 104925, 104926 and 104929), setae characteristic for Phyllodoce. P. rosea (McIntosh, 1874) has, as illustrated by Hartmann-Schroder and Stripp (1968: fig. 4c, called A. subulifera Eliason, 1962), setae very similar to those described here for other species of Phyllodoce, although the teeth on the rostral "cone" are unusually large.

Paranaitis Southern, 1914 (14 species) Moore (1909: 341), in his description of the setae in Paranaitis polynoides (as Anaitis), observed that "the form of the articulation resembles the Eteone type." Ditlevsen (1917) used the same phrase while comparing his Anaitis sp. with Moore's species. Thus, one or both rostra are drawn out as a large beak-like tooth with a number of smaller teeth at the base. Eliason (1962) used this character to help separate P. wahlbergi (Malmgren, 1865) from P. kosteriensis (Malmgren, 1867). My own observations ofthe latter agree in that the setae are heterogomph, with the large tooth much shorter than on the homogomph setae depicted by Southern (1914: fig. 16) for P. wahlbergi.

Eulalia Savigny, 1818 (about 55 species) The setal morphology offive species of Eulalia may be grouped into two types. Eulalia viridis, E. aurea and E. ornata are very similar (Fig. 3D). The distal portion of the shaft is strongly inflated, especially apparent because of the short, slender distal appendage. The rostrum is distally rounded truncate, making it appear similar to the form caused by the anterior rostrum in Phyllodoce (see Fig. 3C). The most prominent character of the posterior rostrum is the presence of two large teeth that bend towards one another and have a considerable gap between them. A heel is lacking. The setal morphology of Eulalia bilineata and E. mustela is significantly different (Fig. 3E). The shaft is only weakly inflated distally, a heel of the same development as in Phyllodoce is present, and the distal appendage has the relative proportions present in most members of the family. However, the two species differ from one another in the relative size of the rostral teeth; in E. bilineata the apical teeth are only slightly larger than the proximal ones; in E. mustela one or two distal teeth are significantly larger than the others (Pleijel, 1987b: fig. 2C). Thus, there appear to be at least two different basic setal morphologies within Eulalia as it is presently defined. Several species in this genus conform closely to the generic diagnosis and bear a great resemblance to the type species, E. viridis. Among other things, they share the setal morphology described above (Fig. 3D). Hartman (1936: 122) gave the following description of the setae in E. aviculiseta: "shaft with slightly thickened articulating end, with numerous fine hairs and two beaklike teeth." This is one of the characters by which she separated her species from E. viridis. However, as may be seen from Figure 3D, this difference is not valid, although an examination of the holotype of E. aviculiseta (USNM 20341) demonstrated that it is distinct from E. viridis for other reasons. A study of the types of E. microphylloides (HZM P-15494 and P-15495) show that the setae are of the E. viridis type as also illustrated in the original description (Hartmann- Schroder, 1979: fig. 20). The E. viridis-group includes E. ornata, E. aurea, E. quadrioculata Moore, 1906, and E. hanssoni Pleijel, 1987. The setae within this group appear to be quite invariant (i.e., my Fig. 3D; see also Babin (1944: fig. 538 BULLETIN OF MARINE SCIENCE, VOL. 48, NO.2, 1991

14.1», although E. hanssoni may be an exception (fig. 5 in Pleijel, 1987a, drawn from an SEM micrograph). Another group of species within Eulalia has several characters in common with E. bilineata (see below). The morphology of the distal portion of the shaft may be useful in delineating these species more precisely. For example, Pleijel (1987b) showed significant differences between E. expusilla and E. mustela, based on SEM studies. Likewise Fauchald (1972), in his description of E. mexicana and E. anoculata, I demonstrated remarkable differences in the setal morphology ofthese two species. An early example of the use of these features is given by McIntosh (1908); he used the length and shape of the rostral teeth as some of the characters separating E. bilineata from E. trilineata de Saint-Joseph, 1888. Study by the author of the following species showed them to belong to the E. bilineata-group and to possess setae of the same general type described above: E. levicornuta Moore, 1909 (holotype USNM 17288), E. californiensis (Hartman, 1936) (holotype USNM 20342) and E. pachycirra Hartman, 1978 (holotype USNM 46735 and paratypes). Pleijel (1987b) identified a group of species within Eulalia, including E. expusilla n. sp., E. bilineata (Johnston, 1840) and E. tripunctata McIntosh, 1874, characterized by a rounded prostomium, short median antenna, rounded dorsal, ventral and anal cirri, a median pygidial papilla and species-specific pigmentation patterns. In addition, species in this group are characterized by a greater inter- specific variation in the morphology of the rostrum of the setal shaft than species in the E. viridis-group. Other species belonging to the E. bilineata-group include E. anoculata Fauchald, 1972, E. mexicana, E. hutchinsonensis (Perkins, 1984) (most of the type material, including the holotype USNM 80515, has been studied, showing the first tentacular segment to be dorsally well developed), and probably E. mustela Pleijel, 1987. It is probable that Eulalia will eventually be divided into two genera, partly based on setal characters. This is not advisable at the present time, as the fun- damental problem of defining significant autapomorphies for the E. viridis-group has not yet been solved. The name Hypoeulalia Bergstrom, 1914 will have priority for the E. bilineata-group.

Eumida Malmgren, 1865 (21 species) The setae of the six species of Eumida as observed with DIe microscopy are virtually indistinguishable from one another. Their shape most closely resembles that found in Phyllodoce, although the rostrum is apically less acuminate (Figs. lA, B, 3F) and the heel is more strongly developed. The appearance of two groups of apical teeth that bend towards one another, as previously described for Phyl- lodoce, is also often seen in Eumida (not figured). However, this phenomenon does not appear to be the result of differences in the anterior and posterior rostra. Rather, in this cas€ only a slight rotation of the longitudinal axis ofthe seta seems to be necessary to bring it about. The SEM microscope also shows a striking agreement in the rostral morphology in Eumida and Phyllodoce (compare Fig. 2A, B), although this should be confirmed through the study of additional species. The setae of Eumida differ from Phyllodoce in that the teeth are more rounded and apically concentrated and the heel is more strongly developed (partially cov- ered with debris on Fig. 2B).

I Eulalia anoculata Fauchald, 1972 is a junior homonym of E. anoculala Hartman and Fauchald, 1971; a new name for Fauchald's species will not be proposed here, pending a revision of cenain genera presently being carried out by Fredrik Pleijel. EIBYE-JACOBSEN: SETAL MORPHOLOGY IN PHYLLODOCIOAE 539

Several useful figures and remarks on the setal articulation within Eumida may be found in the literature. Banse (1959b: 297) noted about E. venustissima as Eulalia (Sige): "Der Verschlui3 der Borsten ist homogomph und mit einigen Reihen gekriimmter zahne bedeckt, die "auf Liicke" stehen." These observations have been verified (holotype, HZM V-12890). Ditlevsen (1917: pI. IV, fig. 12), in his description of E. minuta (as Eulalia), noted the presence of a heel (called a step by him) on the distal portion of the setal shaft. In her description of E. longicirrata Hartmann-Schroder (1975: fig. 18) showed a seta that appears typical of Eumida, although no heel is indicated. Hartmann-Schroder and Stripp (1968) illustrated setae of E. sanguinea (fig. 4f) (as Pirakia punctifera (Grube, 1860» and E. bahusiensis (fig. 4i). Both have setae characteristic of this genus, including two groups of teeth pointing somewhat towards one another; however, a heel is not indicated on these figures either. A setal morphology typical of Eumida has also been found in E. longicornuta (Moore, 1906) (holotype USNM 5515) and in E. tubiformis Moore, 1909 (holotype USNM 16879). Pirakia Bergstrom, 1914 is a junior synonym of Eumida Malmgren, 1865 (Eibye-J acobsen, 1987). My own observations on more than 20 specimens of the type species Pirakiapunctifera (Grube, 1860), present in the collection ofMNHN Gars A62, A63, A64, A408, A490, A491, A857), show the setae to be of the Eumida type.

Sige Malmgren, 1865 (10 species) The rostral morphology in Sige, as revealed in the DIC microscope, is illustrated in Figure 3G. The distal portion of the shaft is weakly inflated, giving it a very delicate appearance. Fewer teeth are present and the heel is very strongly developed, causing the rostrum to appear unusually long and thin. These setae most closely resemble those of Eulalia bilineata and E. mustela. Further study of Sige with SEM confirmed these observations, although the setae of the species used (an undescribed species from Belize) seem to be extreme in having very few, apically concentrated rostral teeth, leaving most of the rostrum itself exposed (Fig. 2D). Most authors have confused Sige and Pterocirrus (for a history of this problem, see Eibye-Jacobsen, 1987); only one valid observation has been published. Perkins (1984, fig. Id-e) illustrated setae of S. parvicirrus (as Eumida); these show six, short slender apical teeth present on the rostrum. S. brunnea (Fauchald, 1972) (holotype AHF 13775) has similar setae with a very narrow rostrum.

Pterocirrus Claparede, 1868 (8 species) The distal part ofthe shaft in Pterocirrus (Fig. 3H) is recognized by its apparent lack of teeth. Close examination with DIC, however, reveals that small apical teeth are present. A heel is developed, but is much less obvious than in Sige (compare Fig. 3G, H). SEM micrographs corroborate these impressions (Fig. 2E). The general shape of the distal portion of the shaft appears to be similar to that in Eumida. The rostral teeth, however, are both shorter and more apically concentrated in Pterocirrus. In the species treated here, an unusually large infrafas- cicular variation is present in the relative dimensions of the rostrum. Although most of the setae have the shape shown on Figure 2E, the rostrum is noticeably more drawn out in the dorsalmost setae, while it is much broader and more truncate in the ventral ones. The remarks of Banse (19 5.9a) and Banse and Nichols (1968) on the setal morphology of P. macroceros were mentioned above. McIntosh (1908: 62) char- 540 BULLETIN OF MARINE SCIENCE, VOL. 48, NO.2, 1991 acterized the rostrum of P. macroceros (as Eulalia) as: "a little dilated and bevelled, ... [with] traces of spikes at the end, the translucent region being otherwise structureless, except for a single oblique striation" (my brackets). The latter is the line of articulation with the distal appendage. That the teeth present on the setal rostrum are difficult to perceive is also evident in Hartman's (1936: 126) description of P. montereyensis (as Sige): "articulation slightly thickened, ornamented with a few inconspicuous spines but without teeth." This corresponds with my own observations on the holotype (USNM 20340).

Bergstroemia Banse, 1973 (2 species) Banse (1973) described the teeth on the setal rostrum of B. nigrimaculata (Moore, 1909: fig. d) as long, fine, and of uniform width. According to his figure, the seta has a morphology very similar to that of Notophyllumfoliosum (Fig. 3J), although the rostrum would appear to be relatively shorter in Bergstroemia.

Clavadoce Hartman, 1936 (3 species) Banse (1973) described the setal rostrum of C. splendida Hartman, 1936 as bearing long, fine teeth of uniform width (about 13 according to his fig. a). This morphology is quite unlike that shown by Blake (1988: fig. Ic) ofa seta from C. cristata (Hartman and Fauchald, 1971),1 which displays a slender rostrum with 6-8 distal teeth.

Nereiphylla de Blainville, 1828 (about 15 species) The setae of Nereiphylla (Fig. 31) most closely resemble those found in Eulalia viridis and related species (compare Fig. 3D). The distal part of the shaft is strongly and abruptly widened, while the appendage is unusually slender and short. A prominent, apical crown of long, acuminate teeth is present; as a group they give the rostrum a very rounded appearance. Although not figured, the striae of the rostrum are exceptionally prominent, running dorsally towards the distal crown of teeth. McIntosh (1908) noted the strong dilation of the distal portion of the shaft in both N. rubiginosa (de Saint-Joseph, 1888) (as Phyllodoce) and in N. lutea (Malm- gren, 1865) (as Genetyllis). For the latter species he also commented on the fine but very noticeable oblique striation on the rostrum. Other authors have commented on the appearance of the distal part of the shaft as abruptly inflated in N. castanea (von Marenzeller, 1879) (as Phyllodoce (Genetyllis» (Berkeley and Berkeley, 1948; Hutchings and Murray, 1984). The setae of Phyllodoce pancerina (=Nereiphylla paretti (delle Chiaje, 1828) fide Hartman, 1959) are illustrated by Claparede (1870: pl. 9, fig. IC). The distal portion of the shaft is unusually swollen and very broad in relation to the basal portion of the distal appendage. The setae of the rather enigmatic Phyllodoce mernoensis Knox (1960: fig. 134) most closely resemble those found in Nereiphylla. This is the genus to which Knox' species on the basis of other characters (primarily the lack of a nuchal papilla and the development of the tentacular segments shown in his figure) most likely should be moved. However, the ventral cirri do not appear to be characteristic of Nereiphylla in the sense of Banse (1973).

Notophyllum 0rsted, 1843 (about 13 species) Only one species of Notophyllum was studied in the DIC microscope (Fig. 3J), and although observations were rather poor, some features were apparent. The EIBYE-JACOBSEN: SETAL MORPHOLOGY IN PHYLLODOCIDAE 541 shape of the distal portion ofthe shaft generally resembles that found in Eumida (compare Fig. 3F), the principal differences being the apparent lack of a heel and the strongly developed apical group (7-10) of long, acuminate teeth. The setae of N. splendens (Schmarda, 1861) appear to be somewhat intermediate between that depicted here for Eumida and Notophyllum (Hartmann-Schrooer 1979: fig. 31). Both Hartmann-Schroder (1971) and Uschakov (1972) remark on the large, apical teeth on the rostrum of N. foliosum, agreeing with my own observations. The results presented here should clearly be corroborated on a larger number of species, especially in large genera such as Phyllodoce and Eulalia. However, this preliminary study allows the following conclusions to be drawn: a) Distinct differences in setal morphology exist in genera other than those in which this has traditionally been recognized (i.e., Eteone s. lat., Hesionura and Paranaitis); b) Many of these genera appear to have a more or less distinct setal morphology. The variation between species is in this regard presumably small, possibly fur- nishing supplementary generic diagnostic characters (i.e., Pseudomystides, Phyl- lodoce, the Eulalia viridis-group, Eumida, Pterocirrus and Nereiphylla); c) In a few groups (i.e., Mystides, the Eulalia bilineata-group, Sige and Clavadoce), the infrageneric variation of setal details may be greater than has generally been acknowledged and thus taxonomically useful, and d) A revision of the genus Eulalia, in which more attention is given to details of the setal morphology, is necessary.

ACKNOWLEDGMENTS

The following persons are thanked for providing me with specimens used in this study: L. Harris (AHF), P. A. Hutchings, S. Perry and R. Paterson (AM), G. Hartmann-SchrOder (HZM), J.-L. Justine (MNHN), K. Fauchald and L. Ward (USNM), J. Kirkegaard and M. E. Petersen (ZMUC), B. O'Connor (University College Galway, Ireland), Ed Stikvoort (Rijkswaterstaat Dienst Getijdewateren. Middel- burg, the Netherlands), P. Garwood (Dove Marine Laboratory, Cullercoats, U.K.), and M. Lindholm Andersen (Marin ID, Denmark). I would also like to express my appreciation to G. Hartmann-SchrOder, E. Diedrich and G. Schulze for giving me working space and logistic assistance during a period of study at HZM and to J.-L. Justine for giving me access to the collections of MNHN. My special thanks go to K. Fauchald and L. Ward for their great support during a 6-month stay at USNM, for giving me access to the rich material there and for arranging the funding of my participation in an expedition to Carrie Bow Cay, Belize. The staffs of the marine biological stations Helsing0r, Frederikshavn, Tjiirno, Roscoff, and Carrie Bow Cay should all be acknowledged for help with practical problems in the field. Special credit is due to C. Buch (BICO A/S) for the kind loan of the Olympus Vanox AHB-S microscope, which started this whole project. B. Kahn and W. Brown (USNM) are also thanked for qualified technical assistance during SEM-sessions. I also wish to recognize the help of the following persons, who discussed the problems treated here with me: K. Fauchald, L. Ward, Y. Shirayama and R. T. O'Grady (all USNM); J. Kirkegaard, C. Nielsen and M. E. Petersen (all ZMUC); P. A. Hutchings (AM). My warm thanks go to R. Wilson (Museum of Victoria, Melbourne) and F. Pleijel (Naturhistoriska Riksmuseet, Stockholm, Sweden) for giving me access to manuscript material before publication. The assistance received from J. Rask in preparing the figures for publication is gratefully acknowledged. This work would not have been possible without a doctoral study grant given by The Danish Natural Science Research Council (11-6465) and a travel grant from The Danish Research Academy (F- 880073), which allowed me to study at the National Museum of Natural History, Washington D.C. for 6 months.

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DATE ACCEPTED: June I, 1990.

ADDRESS: Zoological Museum. Universitetsparken 15, DK-2100 Copenhagen @, Denmark.