A New Species of Megalomma Johansson, 1927 (Polychaeta: Sabellidae: Sabellinae) from Taiwan, with Comments on Sabellid Dorsal Li
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Zoological Studies 42(1): 106-134 (2003) A New Species of Megalomma Johansson, 1927 (Polychaeta: Sabellidae: Sabellinae) from Taiwan, with Comments on Sabellid Dorsal Lip Classification Kirk Fitzhugh Research and Collections Branch, Los Angeles County Museum of Natural History, 900 Exposition Boulevard, Los Angeles, California 90007, USA E-mail: [email protected] (Accepted August 19, 2002) Kirk Fitzhugh (2003) A new species of Megalomma Johansson, 1927 (Polychaeta: Sabellidae: Sabellinae) from Taiwan, with comments on sabellid dorsal lip classification. Zoological Studies 42(1): 106-134. A new species, Megalomma cinctum, is described from the intertidal zone of Hungtou Yu, also known as Orchid Island, located off the southeast coast of Taiwan. The species is most similar to M. coloratum (Chamberlin), M. modestum (Quatrefages), M. quadrioculatum Willey, M. roulei (Gravier), and M. splendidum (Moore), each of which has 1 pair of dorsalmost compound eyes on the radioles, sometimes with additional eyes on the 2nd and 3rd pairs of radioles, and the middorsal margins of the peristomial collar are fused to the lateral margins of the fecal groove, forming dorsolateral pockets. Megalomma cinctum is distinct from these species by the presence of a narrow, white, transverse band on setiger 2, and sometimes also on setiger 3. The number of pairs of radi- olar eyes in M. cinctum is much more variable than has been reported in other species, and is strongly correlat- ed with body size. Additional variability in the species is also reported for the number of pairs of radioles, peri- stomial collar shape, number of thoracic setigers, occurrence of the white transverse bands on setigers 2 and 3, and pigmentation patterns on the crown and thorax. The species is dioecious, but asexual reproduction also occurs by scissiparity. The description of M. cinctum also includes details on the internal construction of the dorsal lip radiolar appendages, necessitating an emendation of the definition of the genus. A brief survey is also presented of dorsal lip internal construction within the Sabellidae, and the implications of those findings for the determination of cladistic relationships among Sabellinae genera are discussed. http://www.sinica.edu.tw/zool/zoolstud/42.1/106.pdf Key words: Annelida, Fan worm, Sabellid, Systematics, Cladistics. The genus Megalomma Johansson, 1927, ourense Knight-Jones, 1997, both from New has been the subject of several recent systematic Zealand. The present paper describes a new studies (Perkins 1984, Fitzhugh 1989, Knight- species from the intertidal zone of Hungtou Yu, Jones 1997). In her review of the genus, Knight- also known as Orchid Island, located off the south- Jones (1997) lists 23 valid species names. east coast of Taiwan. Subsequently, Nishi (1998) described a species During the study of this new species of from the intertidal zone along Phuket Island, Megalomma, it was recognized that a feature of Thailand, and Fitzhugh (2002b) described a the branchial crown dorsal lips has not been fully species from the continental shelf off Thailand, in considered in sabellid systematics. While the the Andaman Sea. Of the species currently occurrence of radiolar appendages in the dorsal described, only 5 have type localities in the west- lips has recently become an integral part of sabel- ern Pacific Ocean: M. acrophthalmos (Grube lid taxonomy (Knight-Jones 1983, Perkins 1984, 1878) from the Philippines, M. pacificum Fitzhugh 1989), the presence or absence of a Johansson, 1927, from the Gilbert Islands, M. tri- branchial skeleton within the appendage has been oculatum Reish, 1968, from the Marshall Islands, infrequently reported (e.g., McIntosh 1918, Nicol and M. suspiciens (Ehlers 1904) and M. kaik- 1931, Orrhage 1980, Fitzhugh 2002b). The pres- 106 Fitzhugh -- New Species of Megalomma 107 ence or absence of the radiolar appendage skele- thoracic notosetae broadly hooded, arranged in 2 ton among species in sabellid genera, and the or more transverse rows. Abdominal neurosetal necessity of including this and other associated fascicles with 2 transverse rows of elongate, nar- characters in descriptions are discussed. As a rowly hooded setae. Thoracic uncini avicular; result, an emended definition is also provided for numerous, very small teeth above main fang; Megalomma. breast well developed; handles of medium length. Companion setae with very thin, tear-drop-shaped distal ends situated perpendicular to shaft. MATERIALS AND METHODS Abdominal uncini similar to thoracic ones, but with shorter handles. Specimens of the new species of Megalomma Remarks: The present definition primarily fol- described in the present paper were collected by lows that given by Perkins (1984) and Fitzhugh hand, fixed in 10% seawater-formalin, then trans- (1989). As was indicated by Fitzhugh (1989), ferred to 70% ethanol. Specimens in the type monophyly of the genus is based on the presence series have been deposited in the following institu- of unpaired, distal, compound radiolar eyes. tions: National Museum of Natural Science, Compound eyes at the distal ends of radioles are Taichung, Taiwan (NMNS), and the Allan Hancock also known in Stylomma palmatum Knight-Jones, Foundation Polychaete Collection of the Los 1997 (see also Knight-Jones and Perkins 1998). Angeles County Museum of Natural History, Los The arrangement of eyes differs, however, in the 2 Angeles, California (LACM-AHF). Additional mate- genera in that they are sessile in Megalomma rial examined in the course of this study came from species and stalked in S. palmatum. The inclusion the LACM-AHF collection and from the Florida of Stylomma Knight-Jones in the cladistic analysis Marine Research Institute, St. Petersburg, Florida of Sabellinae genera by Fitzhugh and Rouse (FSBC). (1999) also showed that the eyes were indepen- For examination of the internal structure of dently derived in the 2 genera. some dorsal lips, simple transverse sections were The emendation provided here is to point out prepared by cutting an isolated lip at mid-length (1) that the radiolar appendages of the dorsal lips with a pair of ophthalmic micro-scissors, then cut- do not have an associated branchial skeleton, and ting off this exposed surface from the remainder of (2) that dorsal pinnular appendages can be pre- the lip as thinly as possible. Sections were then sent or absent. The new species described below placed in glycerol on a microscopic slide, and lacks both the pinnular appendages and a radiolar examined unstained using a compound micro- appendage skeleton, which has prompted this scope. modification of the definition of the genus. Although recent definitions of Megalomma (e.g., Perkins 1984, Fitzhugh 1989, Knight-Jones 1997) SYSTEMATIC ACCOUNT have taken into consideration the presence of radi- olar appendages, this has only been to the extent Genus Megalomma Johansson, 1927, emended of recognizing the general shape of the dorsal lips, indicating the presence of those appendages. The Definition: Medium- to large-sized species presence of radiolar appendages can readily be with numerous radioles; radiolar skeleton with 4 or observed in species of Megalomma by the longitu- more rows of cells. Palmate membrane and radio- dinal ridge or“midrib”(Knight-Jones 1983 1997, lar flanges absent. One or more pairs of radioles Knight-Jones and Walker 1985, Knight-Jones et al. each with a single, sessile, compound eye situated 1991, Knight-Jones and Perkins 1998) extending along inner radiole margin near distal end. Dorsal through the length of the lips. lips triangular, radiolar appendages present, indi- In his investigations on the structural relations cated by longitudinal ridge or“midrib”along entire between sabellid dorsal lip radiolar appendages length; distinct blood vessel and radiolar sheath “( lip-associated radioles”) and radioles, Orrhage tissue “( surrounding sheath tissue”sensu Orrhage (1980) described the internal components of both 1980, see below) extending through length of structures. At least 4 distinct features are common appendages, without a branchial skeleton exten- to the interior of radiolar appendages and radioles: sion; dorsal pinnular appendages present or (1) a coelomic space, (2) a blood vessel branching absent. Ventral lips and parallel lamellae present. from the branchial vessel, (3) surrounding sheath Posterior peristomial ring collar present. Inferior tissue, referred to here as radiolar appendage sur- 108 Zoological Studies 42(1): 106-134(2003) rounding sheath tissue, and (4) a pair of nerves. radiolar eyes), LACM-AHF 2088 (6 specimens, Orrhage also found that in at least 1 species he each with 5-6 radiolar eyes), and LACM-AHF 2089 examined, Sabella penicillus Linnaeus (now S. (28 fragments, consisting of detached branchial pavonina Savigny, see Knight-Jones and Perkins crowns and specimens without crowns). 1998), a branchial skeleton, composed of thick- Etymology: The specific epithet refers to the walled cells, was also present within the radiolar transverse white bands present around setigers 2 appendages. As a result of recent work by and 3. Fitzhugh (2002b) and Fitzhugh et al. (in prep.), it Description: Holotype complete, with 7 tho- has become apparent that taxonomic studies of racic and 33 abdominal setigers. Total branchial sabellids require consideration of the presence or crown length 2.5 mm; total thorax-abdomen length absence of at least the radiolar appendage skele- 7.0 mm; maximum width 0.6 mm throughout most ton, and preferably both the skeleton and the sur-