FAU Institutional Repository http://purl.fcla.edu/fau/fauir This paper was submitted by the faculty of FAU’s Harbor Branch Oceanographic Institute. Notice: ©1987 The Biological Society of Washington http://www.brynmawr.edu/biology/BSW/. This manuscript is an author version with the final publication available and may be cited as: Rice, S. A. (1987). Reproductive biology, systematics, and evolution in the polychaete family alciopidae. Bulletin of the Biological Society of Washington, 7, 114‐127. BIOL. SOC. WASH. BULL. NO.7, 1987, pp. 114-127 REPRODUCTIVE BIOLOGY, SYSTEMATICS, AND EVOLUTION IN THE POLYCHAETE FAMILY ALCIOPIDAE Stanley A. Rice Abstract.-Fourteen species of aiciopid polychaetes were collected from the western Atlantic Ocean near Grand Bahama Island. Specimens were collected using a submersible equipped with a special zooplankton-capturing device. Eight of the nine known aiciopid genera were represented in the collections along with a new species in the genus Krohnia. This new species is described and the rela­ tionships among aiciopid genera are reexamined in light ofnew information from field observations and reproductive biology. Six families within the Class Polychaeta are and has provided an avenue for in situ obser­ considered to be holopelagic spending their en­ vation of behavior. tire life cycle in the water column. These six fam­ The present paper contains an account of al­ ilies include: Alciopidae, Iospilidae, Lopador­ ciopid species collected from the western Atlan­ rhynchidae, Pontodoridae, Tomopteridae, and tic Ocean including a description ofa new species. Typhloscolecidae. A number ofother polychaete New information is presented on the structure families contain one or more species that may of aiciopid sperm and their relation to sperm be truly pelagic, such as some species within the storage mechanisms in the females.The evolu­ Polynoidae (Dales and Peter 1972) while many tionary relationships among genera within the species of other families spend part of their life Aiciopidae are reexamined in light of this new cycle in the water column either as larvae or information. adults but at some point return to a benthic hab­ Materials and Methods itat and thus are not truly pelagic. Each of the holopelagic polychaete families The specimens described and illustrated in this shows special morphological and reproductive paper were collected from the northwest Provi­ adaptations to life in the water column. A trans­ dence Channel, Bahama Islands between Sep­ parent body that renders individuals nearly in­ tember 1979 and April 1980. Collections were visible in the natural habitat has evolved in the made using the Johnson Sea-Link submersible Aiciopidae, Tomopteridae, and Typhloscoleci­ operated out ofHarbor Branch Foundation, Inc., dae. Swimming aids have evolved such as flat­ Fort Pierce, Horida. Most specimens were col­ tened, composite setae in the Lopadorrhynchi­ lected at depths between 90 and 250 m. All col­ dae, and parapodial paddles without setae in the lections were made during darkness so that spec­ Tomopteridae. The Aiciopidae have developed imens could be attracted to the submersible by enlarged, complex eyes, grasping eversible pro­ use of floodlights. boscides and sperm storage organs in females as Living specimens were returned to the surface pelagic adaptations. and immediately fixed for electron microscopy Most pelagic polychaetes are fragile and do not (EM) or for taxonomic studies. Specimens in­ stand up well to physical abuse. As a result, spec­ tended for EM studies were fixed for 2 hours at imens collected in plankton nets are often dam­ room temperature in a 2.5% glutaraldehyde buff­ aged and fragmented, making taxonomic work ered with 0.2 M sodium phosphate, followed by difficult and live observation nearly impossible. postfixation in I% osmium buffered with 1.25% The development and application ofadvanced sodium bicarbonate. Thick and thin sections were sampling gear operated from a non-tethered sub­ cut on a DuPont MT2-B ultramicrotome. Scan­ mersible has resulted in the collection ofundam­ ning electron microscope (SEM) specimens were aged living specimens of the family Alciopidae fixed and postfixed as above followed by dehy- NUMBER 7 115 Table I.-Currently recognized genera in the Family Alciopidae, following revisions ofSt0p-Bowitz (1948). Fauvel (1923) Step-Bowitz (1948) Alciopa Audouin and Milne-Edwards (1833) *Naiades Delle Chiaje (1830) Vanadis Claparede (1870) Vanadis Oaparede (1870) Greeffia McIntosh (1885) *Alciopa Audouin and Milne-Edwards (1833) Asterope Oaparede (1870) Torrea Quatrefages (1850) Callizonella Apstein (1891) Krohnia Quatrefages (1865) Corynocephalus Levinsen (1885) Alciopina Oaparede and Panceri (1867) Rhynchonerella Costa (1862) Plotohelmis Chamberlin (1919) Callizona Greeff (1876) Rhynchonerella Costa (1862) *Watelio St0p-Bowitz (1848) * Monotypic genera (Fauchald 1977). dration to acetone. Critical point drying was ac­ Presently, there are nine recognized genera in complished using liquid CO2 as the transition the Alciopidae (Table I) and 31 species, includ­ solvent. Dried specimens were mounted and ing the new species described herein. The prin­ coated with gold-palladium and viewed on a Zeiss cipal diagnostic characteristics used to separate Novascan 30 scanning electron microscope. genera and species within the Alciopidiae in­ clude: I) the prostomial shape and appendages; 2) the proboscis morphology; 3) number and Systematics of the Family Alciopidae shape of tentacular cirri on the first three seg­ The family Alciopidae Ehlers, 1864, contains ments; 4) parapodial morphology and 5) type and a relatively homogeneous assemblage ofspecies distribution of setae. that resemble, in many characteristics, the Phyl­ Proslomium.-A pair of enlarged eyes are lodocidae (Dales 1955). The single unifying char­ prominent on all species and often obscure other acteristic ofall alciopids is the possession oftwo features ofthe head (Fig. lA-F). The angle ofthe greatly enlarged, complex eyes. The structure and eyes may be downward, upward, forward, or function ofthese organs have been described by slanted to the side. Dales (1957) used the angle Hermans and Eakin (1974) and Wald and Ray­ ofthe eyes as a diagnostic character but this has port (1977). In general, alciopids have long, thin been criticized by Tebble (1962) as being unre­ bodies that are semi-transparent to transparent liable. Located anteriorly between the eyes is a in life. Five small antennae are present on the small truncated ·extension of the prostomium prostomium; four antennae on the anterior mar­ bearing four short antennae. These antennae vary gins and one median antenna between the eyes. from inconspicuousto prominentand often have The first three segments bear 3-5 pairs of ten­ sensory tufts that are visible with the SEM (Fig. tacular cirri. The proboscis is eversible with dis­ IB). A single median antenna is present between tal papillae or fleshy lateral horns for grasping. the eyes. Parapodia are uniramous with conical setigerous Proboscis. -An eversible proboscis is present lobes and leafy dorsal cirri. Setae may be simple in all species but shows considerable variation or compound. Segmental glands are present lat­ in size and shape between species (Fig. 2A-F). erally or dorsally on most parapodia. Chitinizedjaws are never present; however, some Early systematic accounts of the Alciopidae species have hardened papillae lining the inner are often confusingand may contain descriptions surface of the everted proboscis (Fig. 2E-F). based upon incomplete and damaged specimens. Fleshy marginal papillae are often present on the Important references in the early literature in­ distal border of the proboscis and these may be clude Greeff (1876), Apstein (1900) and Fauvel accompanied by long fleshy horns in some species (1923). St0p-Bowitz (1948) revised the genera (Fig. 2A-B). Proboscis horns and papillae are within the Alciopidae (Table I). Recent literature generally equipped with mucus glands that pre­ on the family includes: Dales (1955); Tebble sumably aid in prey capture. The relative length (1960, 1962); Day (1967); Dales and Peter (1972); ofthe proboscis along with the numberand shape Uschakov (1972) and Fauchald (1977). of papillae and horns are diagnostic. 116 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. I. Anterior view ofselected alciopid polychaetes showing orientation ofeyes, prostomial antennae and tentacular cirri. A, A/ciopina parasitica; B, Krohnia /epidoa; C, Vanadis crystal/ina; D, v. formosa; E, A/ciopa reynaudii; F, Rhynchonerella moebii. Tentacular cirri. - The first three segments in ofthe tentacular cirri on the first three segments. all alciopids are asetigerous and apodous. These This formula is represented by segments bear tentacular cirri that may be elon­ I I gate and cylindrical, flattened, or conical. Day I+--+---~ (1967) has developed a tentacular formula for o or I 0 or I or N describing the number, arrangement and shape where I = tentacular cirrus present; 0 = tenta- NUMBER 7 117 Fig. 2. Anterior views ofselected alciopid polychaetes showing eversible proboscides. A, A/dopa reynaudii; B, Vanadis formosa; C, Rhyru:honerella ange/ini; D, R. perersi; E, Torrea candida; F, T. candida close-up of chitinized papillae on inner surface of proboscis. cular cirrus absent and N = normal lamellar cir­ lamellar cirri on segment three. The tentacular rus present. For example, Krohnia /epidota has formula is then a single pair of tentacular cirri on segment I;