Malmgreniella Hartman, 1967

Barnich, R. & Fiege, D., 2001, Mediterranean species of Malmgreniella Hartman, 1967 Polychaeta: Polynoidae: Polynoinae), including the description of a species, Journal of Natural History 35, pp. 1119-1142 : 1121-1123

Genus Malmgreniella Hartman, 1967 , emended

Type species: Malmgreniella dicirra Hartman, 1967 , by monotypy; gender feminine.

Diagnosis. Body dorsoventrally ¯attened, short, with up to 46 segments; body more or less covered by elytra or short tail uncovered (in large specimens). Fifteen pairs of delicate elytra on segments 2, 4, 5, 7, on alternate segments to 23, 26, 29, and 32; ®rst pair rounded, the following oval to kidney-shaped; surface with or without microtubercles and papillae, without macrotubercles; margin smooth or with fringing papillae (e.g. ®gures 1B, C; 2B, C; 6C, D; 7B, C). Dorsal cirri with cylindrical cirrophores and long styles on segments lacking elytra; small nodular dorsal tubercles on cirrigerous segments (e.g. ®gure 3A, D). Ventral cirri short (except

second segment), consisting of cirrophore and style on all segments (e.g. gure 3D). Nephridial papillae usually present from chaetiger 6 to the end of the body. Pygidium with one pair of long anal cirri.

Prostomium bilobed, usually without distinct cephalic peaks, three antennae: ceratophore of median antenna in anterior notch, lateral antennae with ceratophores inserted terminoventrally; one pair of palps; two pairs of eyes, anterior pair usually dorsolaterally in front of widest part of prostomium, posterior pair dorsally near hind margin of prostomium (e.g. ®gure 6A, B). Pharynx with nine pairs of border papillae and two pairs of hooked jaws. First or tentacular segment with a pair of tentaculophore s inserted laterally to prostomium, each usually with few chaetae on inner side (in some species without chaetae), and with a dorsal and a ventral tentacular cirrus (e.g. ®gures 3A; 4A). Second or buccal segment with ®rst pair of elytra, sub-biramous or biramous parapodia, and long ventral cirri.

Parapodia of subsequent segments sub-biramous or biramous, both rami with projecting acicular lobes; neuropodia not deeply notched, usually with supra-acicular process; tips of noto- and neuroacicula penetrating epidermis (e.g. ®gure 2D). Notochaetae about as stout as neurochaetae, usually with rows of spines and blunt

pointed tips (®gures 1E, F; 2E, F); neurochaetae more numerous, with rows of spines only distally and one or two kinds of tips: bidentate with secondary tooth subdistally and / or unidentate with pointed or knob-like tip (e.g. ®gures 1G±J; 2I±L).

Remarks. In the revision of Pettibone (1993) the genus Malmgreniella Hartman, 1967 was included in the subfamily Harmothoinae Willey, 1902 (i.e. Polynoinae Kinberg, 1856 , which has priority over Harmothoinae Willey, 1902 ; see also Muir, 1982 and Barnich and Fiege, in press), while Hartmann-Schr oÈder (1996) considered this genus to belong within the subfamily Arctonoinae Hanley, 1989 . According to Hanley (1989) and Hanley and Burke (1991) members of the Arctonoinae are characterized by lateral antennae inserted terminoventrally, tentaculophore s without chaetae, and parapodia sub-biramous with neuropodia deeply notched dorsally and ventrally, in contrast to those of the Polynoinae ( Harmothoinae sensu Hanley ), which show lateral antennae inserted ventrally, tentaculophore s usually with chaetae, and parapodia biramous with neuropodia not deeply notched. But, in the genus Malmgreniella lateral antennae are inserted terminoventrally, which points to the 1122 Arctonoinae , while tentaculophores usually show few chaetae, and parapodia are sub-biramous or biramous with neuropodia not deeply notched, which rather points

the Polynoinae . Since similar problems considering the subfamily grouping exist also for several other polynoid genera, e.g. Adyte Saint-Joseph, 1899 , Subadyte Pettibone, 1969 (see Hanley, 1989 and Pettibone, 1993), we suggest that a revision

the diOEerent genera grouped within the Arctonoinae and Polynoinae should be undertaken based on type material. Unfortunately such a revision is beyond the scope of this study, thus, we prefer to follow Pettibone (1993) and consider the genus Malmgreniella to belong to the Polynoinae for the time being.

In the previous literature species considered herein have either been included in the genus Malmgrenia McIntosh, 1874 (see also McIntosh, 1874a, 1876 and 1900; Fauvel, 1923; Chambers and Muir, 1997) or in Malmgreniella Hartman, 1967 (e.g. Pettibone, 1993; Hartmann-Schr oÈder, 1996). In accordance with Hanley (1987) and Pettibone (1993), and in contrast to Chambers and Muir (1997), we consider Malmgrenia McIntosh, 1874 indeterminable and use Malmgreniella Hartman, 1967 , although we believe that Chambers and Muir correctly interpreted McIntosh’s intention to establish the genus Malmgrenia based on the new species M. castanea and M. andreapoli s (see McIntosh, 1900:`Malmgrenia McIntosh, 1876’). But unfortunately McIntosh’s paper with the description of the new species M. whiteavesii was published earlier and thus has priority, as acknowledged by Hartman (1959), through the designation of M. whiteavesii as the type species for the genus Malmgrenia . The genus Malmgrenia , however, is indeterminable (see also Pettibone, 1993), because

. whiteavesii McIntosh, 1874 is insu ciently described, and the holotype has been lost since 1973 (personal communication by M. Lowe, BMNH). However, Chambers and Muir (1997) justi®ed the use of the genus Malmgrenia , by the fact that in the original diagnosis Hartman (1967) described several characters of Malmgreniella which are not found in the material from the northeast Atlantic. But, Chambers and Muir did not consider the emended generic diagnosis given in Pettibone (1993), which ®ts the species of the northeast Atlantic, and also those of the Mediterranean Sea.

Malmgreniella Hartman, 1967 is closely related to Harmothoe Kinberg, 1856 , but the following characters clearly separate the two genera: (1) the absence / presence

cephalic peaks; (2) the position of the anterior pair of eyes; and (3) the mode of insertion of the lateral antennae. In Harmothoe distinct cephalic peaks are usually present, the anterior pair of eyes is either situated below the prostomial peaks or at the widest part of the prostomium, and the lateral antennae are clearly positioned ventrally to the median antenna. In Malmgreniella however, distinct cephalic peaks are usually absent, the anterior pair of eyes is usually situated in front of the widest part of the prostomium, and the lateral antennae are inserted`terminoventrally’ or ventrally converging mid-ventrally’ (see Pettibone, 1953, 1993), or`subterminally’ see Day, 1967; Hanley, 1987), or`laterally’ (see Chambers and Muir, 1997). In fact, these diOEerent terms describe the same feature: viewed dorsally, the styles of the lateral antennae are situated more or less laterally with respect to the median antenna (see e.g. ®gure 6A), but viewed ventrally, their ceratophores originate terminally below the median antenna and are fused midventrally at their bases (see gure 6B). Thus, in our opinion the term`terminoventrally’ as used by Pettibone 1993) is the most appropriate to describe the insertion of the lateral antennae in Malmgreniella .

Unfortunately, the term`usually’ has to be used in the description of several characters given in the generic diagnosis of Malmgreniella (see above) and also in that of Harmothoe (see Barnich and Fiege, in press). In our opinion, this reēcts that too many diOEerent species have to be covered by these diagnoses, but only a complete revision of all Malmgreniella , Harmothoe and related species could help in establishing more precise diagnoses.

Branch (1998) described a new species, Malmgreniella W mbria, which she tentatively placed in the genus Malmgreniella because of the similarities in the structure

the parapodia, the antennae, the elytra, and the neurochaetae. But this new species is also characterized by 16 pairs of elytra and the presence of distinct cephalic peaks. Since Malmgreniella has been characterized up to now by 15 pairs of elytra only and cephalic peaks being usually absent, the generic diagnosis ought to be emended regarding the number of elytra to cover this species. This would add one more imprecise character to the diagnosis and therefore, we would rather consider this species to be of uncertain generic status for the moment, until a revision as mentioned above will help to solve this problem.

Key to the Mediterranean species of Malmgreniella

Elytral surface without microtubercles; in anterior elytra some scattered papillae on elytral surface and margin; neurochaetae all unidentate, tapering to long, pointed tips,

few lower with knob-like tips (®gure 1B, C, G±J)....... M. lilianae Elytral surface with microtubercles.............. 2

Elytral surface with patch of microtubercles in anterior part; elytral margin smooth (®gures 2B, C; 3B, C; 4B, C; 5B, C).............. 3 Elytral surface covered more or less completely by microtubercles; elytral margin with fringing papillae (®gures 6C, D; 7B, C)............. 6

Short notochaetae stout, with blunt tips; long notochaetae slender, with pointed tips; upper and middle neurochaetae bidentate, lower neurochaetae unidentate (®gure 2E±L)

..................... M. darbouxi All (short and long) notochaetae stout, tips either all blunt or all pointed.... 4

Neurochaetae usually all bidentate, upper and middle with long, lower with (or rarely without) short secondary tooth; notochaetae with distinct rows of spines and blunt

tips (®gure 3E±J)................ M. lunulata Neurochaetae bi-and unidentate; notochaetae with faint rows of spines.... 5

Notochaetae with blunt tips; upper and lower neurochaetae usually unidentate with knob-like tips, middle neurochaetae bidentate with distinct secondary tooth, tips knob-

like or pointed; antennae and cirri papillate, usually pigmented basally (in preserved material) (®gure 4E±J).............. M. andreapolis Noto- and neurochaetae with pointed tips; upper and middle neurochaetae bidentate with minute secondary tooth, lower neurochaetae unidentate; antennae and cirri smooth, not pigmented basally (in preserved material) (®gure 5E±J).. M. castanea

Outer lateral margin of elytra with many long papillae, posterior margin with shorter papillae; neurochaetae all unidentate, tapering to long, pointed tips (®gure 6F±K)................... M. polypapillata , sp. nov. Elytral margin with few, short, scattered papillae; neurochaetae usually all bidentate (®gure 7E±J.)................. M. ljungmani