l~ccords of the \:Ycstcm A/lstrallal/ I'vl/lsC/l1l1 Supplement No 69 111~117 (2006)

A redescription of Facelina stearnsi Cockerell, 1901 (Nudibranchia: Aeolidacea: ) with a reassignment of its generic placement

Jamie M. Chan and Terrence M. Gosliner Department of Invertebrate Zoology and Geology, California Academy of Sciences 875 lloward Street, San Francisco, California 94103, USA

Abstract - Facelina steamsi Cockerell, 1901 is known from Santa Cruz (Nelson 1986) to La Jolla, California (Behrens 1991). Several preserved specimens are examined and compared with the original description and other species in the AlIstraeolis Burn, 1962. The coloration, reproductive and radular morphology are described in detail for the first time using SE M, compound microscopy and photographs of live specimens. Observation of several new morphological and anatomical characters suggests a reassignment of Facelina stcamsi to the genus AlIstraeolis Burn, 1962.

INTRODUCTION Features of living were recorded from According to Miller (1974) the Facelinidae is photographs or notes of collectors. divided into subfamilies based on the ceratal arrangement. Genera are further divided on the basis of penial morphology. Cockerell (1901) Austraeolis stearnsi (Cockerell, 1901) described Facelina stearnsi from a specimen collected Facelina stearnsi Cockerell, 1901: 86. Behrens, 1991:97 in San Pedro, California. Cockerell's description Fig. 206. consisted of observations of the external and radular morphology. No details were given for any Phidiana stearnsi McDonald, 1983: 203-204 figure of the internal organ anatomy. There were no plates 111. or drawings in the original description. The type AlIstraeolis stearnsi (Cockerell) comb. novo material has not been located in any American museum and a neotype is here designated. Material examined Additional records of this species have been Neotype: CASIZ 171165, one specimen 2 cm in published by Eliot (1907), O'Donoghue (1926), length. CASIZ 89095 fifteen preserved specimens, Steinberg (1961), Marcus and Marcus (1967), and three dissected, Morro Bay, San Luis Obispo Nelson (1986) but only two publications, County, California, United States of America, 12 m O'Donoghue (1927) and McDonald (1983), included depth, 17 July 1993, Mike Behrens, lengths 2.5 cm, 3 line drawings of radular teeth and one jaw part. cm, 2cm (preserved dissected specimens). The present study examines several specimens of Facelina stearnsi and compares them to previous Distribution descriptions. This species is known from Santa Cruz (Nelson, 1986) to La Jolla, California (Behrens, 1991). MATERIAL AND METHODS Material was obtained from the Department of Etymology Invertebrate Zoology and Geology, California This species was named after Or. Robert Edward Academy of Sciences, San Francisco (CASIZ). The Carter Stearns. specimens were dissected by ventral or dorsal incision. Their internal features were examined and External Morphology drawn under a dissecting microscope with a camera The body is elongate and slender, with a trailing lucida. Systematically important soft parts were posterior end of the foot (Figure 1). The anterior critical point dried for SEM. Special attention was margin of the foot and tentaculiform foot corners paid to the morphology of the reproductive system, are bilabiate and slightly notched. Body color can digestive system and central nervous system. The range from translucent clear to a pink or purple penial hooks and jaw plates were prepared for cast. The moderately long are cylindrical and examination by SEM. Radulae were extracted and taper distally. The tightly grouped cerata are examined using SEM or compound microscopy. arranged in eight arches on each side of the body ------

112 J.M. Chan, T.M. Gosliner

located below the anterior limb of the first ceratal arch.

Anatomy The formula is 24-25xO.1.0 (Figure 3A) for three examined specimens. The rachidian tooth consists of a single large incurved hook-like tooth flanked by 4-6 smaller denticles on each side (Figure 3A, 3B). The jaws have a tan-brown hue to them (Figure 2B, 3C). There is a thickened tooth­ like projection at the anterior corner of each jaw. There are 18-20 denticles on the masticatory edge of each jaw. Some of the larger denticles have distinct papillae on the top (Figure 3D). The ampulla is long and curved (Figure 2C). The ampulla bifurcates to the oviduct and the vas deferens. The oviduct is one quarter the length of ampulla and leads to a large spherical receptaculum seminis. The receptaculum seminis connects to the female gland mass via a thin duct half the size of the oviduct. The female gland mass is composed of a large mucous gland, a small albumen gland and a convoluted membrane gland. The vas deferens is thick, twice diameter and half the length of the ampulla. The vas deferens curves into a fibrous penial sheath. The penis is armed with a ring of 20 spines (Figure 20, 3E, 3F). The spines are papillate in shape.

Discussion Cockerell (1901) described the radula of Facelina stearnsi as being very similar to Hermissenda crassicornis (Eschscholtz, 1831), but Facelina stearnsi can be distinguished from H. crassicornis by the absence of minute denticles lining the ventral side of the primary cusp of the rachidian tooth. The Figure 1 Photograph of living specimen of Austraeolis rachidian tooth of H. crassicornis is more concave stearnsi (CockerelJ, 1901) photo by T. Gosliner. than Facelina stearnsi. Cockerell made no observation of the jaws or reproductive system. (Figure 2A). The largest arch is the most anterior on Eliot (1907) described a Facelina sp. from a each side and consists of 28-30 cerata. The cerata preserved specimen collected in Dunedin Harbour, are translucent from the base terminating with a ew Zealand. Eliot described the Facelina sp. as broad band of orange/vermillion to yellow and having a similar radula and jaw morphology to tipped with white. The are annulate Facelina stearnsi. The ceratal arrangement of this and have 1D-15 annulae each. The upper half of the species was not described. 0 observation was made rhinophores transition from yellow to orange/ of the reproductive system other than the presence vermillion, with a white tip. Both the oral tentacles of penial armature. Data were not collected about the and the tentacular anterior foot corners have an color and appearance of the specimen in its living orange mid-region and an opaque white tip. The state. The reported range of Facelina stearnsi is from long tapering oral tentacles also have a white or Santa Cruz to La Jolla, California (Bemens, 1991). It cream line that runs along the dorsal side and then is unlikely that Eliot's Facelina sp. is Facelina stearnsi across the head to the base of the rhinophores. The based on the collection locality and the line is often orange/yellow toward the base of the biogeographical distinctions of California and New . There are irregular spots of orange/ Zealand marine biotas. vermillion behind the rhinophores and between O'Donoghue (1927) described a preserved each group of cerata. A white to orange line runs specimen of Facelina stearnsi from Laguna Beach, dorsally from the posterior cerata cluster to the tip California. The external morphology and radula of the foot. The anus is located ventral and within described agree with the description by Cockerel!. the second cluster of cerata. The genital is O'Donoghue observed that the jaw had a A B

a 9

c

2 arr"an,geml:nt. abbreviatIOns: bar lmm; 114 J.M. Chan, T.M. Gosliner

Figure 3 Scanning Electron Micrographs of Allstraeolis steamsi (Cockerell, 1901) (CASIZ 89095). A, Radula, scale bar = 100Jlm; B, Radula close up, scale bar = 20Jlm; C, Jaw, scale bar = lOOJlm; D, Jaw close up, scale bar = lOJlm; E, Penis and detail of penial papillae, scale bar =20Jlm, Insert scale bar =3Jlm; F, Detail of penial papillae, scale bar = lOJlm.

Based on Miller's diagnosis and original generic are very incurved and terminate in a fine point. The descriptions the penis of Face/ina steamsi is most penial spines of Face/ina steamsi are shorter and similar to the Favorininae genera Amanda (Macnae more papillate in shape. Other differences include 1954) and Austraeolis Bum 1962 (Table 1). Amanda dramatically different color pattern and more armata is described as having a penis with a sub­ sparsely arranged cerata in Face/ina armata. apical circle of tiny hooks. The rhinophores are The genus Shinanoeolis Baba, 1965 is described as annulate with 3-4 incomplete rings compared to having a more similar penial shape and armature to 10-15 annulae in Facelina steamsi. The radula is Face/ina steamsi. Based on the original type uniserate with denticles and the jaw is denticulate. description Cllthona emllrai (Baba 1937) for The description of the reproductive system of A. Shil1anoeolis the external features and radula do not armata, the type species, indicates penial spines that agree with the current description of Facelina :;.::l t1lp.. t1l tIl n ... >8' -00 ::l o Table 1 A comparison of the anatomical characteristics of Amanda armata Macnae, 1954, Austraeolis ornata Angas, 1864, and Austraeolis stearnsi (Cockerell, 1901)...... Facelinastearnsi ~ Specie/Genus Amanda armata Austraeolis benthicola Austraeolis catina Austraeolis ornata Austraeolis westeralis (Angas, 1864) (Burn, 1966) (Present study) '"~ (Macnae, 1954 ) (Burn, 1966) (Marcus & Marcus, 1964) ::l"0 l:l Annulate and have Rhinophores Annulate Annulate with Annulate and have up Annulate Annulate with 12 '"~ Rhinophores or more annulae each 10-15 annulae each l:l Rhinophores 5 annulae each to 12 annulae each ~ "0'" Subovate jaw with (') Jaw 16 dentides on the Ovate jaw, with 30 conical, knobbed dentides 20 dentides on the o 30 irregularly on the masticatory masticatory edge 20 large conical n masticatory edge ?r of each jaw rounded dentides edge of each jaw of each jaw dentides on the lower edge of ~ each jaw and numerous .... o\D small serrrations above .... 18-20 dentides on the masticatory edge of each jaw

24-25xO.1.0 Radula 16-17xO.1.0 28xO.1.0 23xO.1.0 20-22xO.1.0 23xO.1.0 Radula uniserate with Long slender teeth The cusp of the Prominent central A tapering cusp with Singlelargeincurved hook-like tooth prominent cusp and with a large tapering tooth is prominent, cusp with 4-5 4-5 dentides each side strong lateral dentides cusp and 2-3 flanked by 4-6 lateral dentides flanked by 4-6 dentides each side slightly curved dentides smaller dentides on each side

armed with a ring of 20 Penis sub-apical cirde conical and short, tip cynlindrical and finger-like and Similar to Austraeolis of tiny hooks with a shallow rim curved upward. Ends curved forward, ornata but is spines. The spines are on one side that forms in a disc whose edge transversely demarked shorter and broader papillate in shape a lip from which the vas is beset with 10 by muscular ridges; deferens opens broad warts, each glans beset with a bearing a tiny spine. Two cirdet of six minute further warts on the surface fleshy filaments, of the disc also with spines. without any trace of chitonous or spicular hooks

.... (Jl 116 J.M. Chan, T.M. Gosliner stearnsi. Distinctive differences such as smooth Austraeolis westralis is described from seven rhinophores, color pattern and a rachidian tooth specimens collected in Western Australia. bearing numerous small indistinct denticles suggest Austraeolis westralis has an internal and external its synonymy with Hermissenda crassicornis anatomy similar to A. ornata. Austraeolis westralis (Eschscholtz, 1831). differs from A. ornata in possessing a narrower The penis, radula, and jaw structure of Facelina coiled prostatic vas deferens, larger outer vas stearnsi agree most closely with species in the genus deferens, shorter, broader penis with numerous Austraeolis Bum 1962. Austraeolis is described as denticle-like fleshy serrations at the tip and a more having a conical penis with a fleshy lobe elongate spermatheca. Due to the lack of knowledge (sometimes papillate) or short fleshy processes or on opisthobranch fauna in southern Australia, Bum warts, a rachidian with 4-5 lateral denticles on each considers that Austraeolis westralis might be an side, and a denticulate jaw. There are five described extreme form of A. ornata. species of Austraeolis; A. ornata Bum, 1962, A. fucia Austraeolis catina is described from two Bum, 1962, A. benthicola, Bum 1966, A. westeralis, preserved species. Observations were made of the Bum, 1966 and A. catina Marcus and Marcus, 1967. external and radular morphology. Austraeolis Austraeolis ornata, A. fucia, A. benthicola and A. catina differs from the other species by the absence westeralis are known from Australia. Austraeolis of cuticular hooks on the six filaments of the penis, catina is found in the Caribbean. 9 rhinophoral rings, 20 denticles on the Bum made observations of the external, internal, masticatory border of the jaw and a radular radular and penial morphology of A. ornata. The formula of 20-22xO.1.0. Body color is described as penis is described as '...... finger-like and curved 'transparent greyish .. with brown marks on side forward, transversely demarked by muscular of head and middle of rhinophores' and with ridges; glans beset with a circlet of six minute fleshy white speckling on skin, concentrated in some filaments, without any trace of chitonous or spicular parts such as middle of head and tentacles'. hooks.' This is in contrast to the more than 20 spines Facelina stearnsi can be distinguished from other present in Facelina stearnsi. The radula formula is species of Austraeolis by its external and penial 20-22xO.1.0. The jaw has a denticulate masticatory morphology. border. The body color can range in color from a Based on this study, the following characteristics pale translucent yellow to an intense bright orange. appear to be consistent for all Facelina stearnsi There are usually white patches and spots on the specimens: body often bluish or iridescent. The rhinophores 1. Oral tentacles have a white or cream line that have some prominent folds or ridges and are runs along the dorsal side and then across the usually tipped with white rather than the well head to the base of the rhinophores. The line is developed annulae found in Facelina stearnsi. The often orange/yellow toward the base of the digestive gland within the cerata ranges in color rhinophore. from a pale yellow to a dark brown. (Rudman, 2. Irregular spots of orange/vermillion behind the 2000). rhinophores and between each group of cerata. Austraeolis fucia is described from a single 3. Cerata arranged in eight arches specimen collected in Queenscliff, Australia. 3. Anus is located ventral and anterior to the Observations were made of the external and second cluster of cerata. radular morphology. There are no illustrations of 4. The genital aperture is located below the anterior the internal anatomy. The radula formula is limb of the first ceratal arch. 19xO.1.0. Burn noted that the morphology of Austraeolis fucia is distinct from A. ornata by 'having 5. The penis is armed with a ring of papillate fewer annulae on the rhinophores'. The body color spines. is creamy-white, the cerata are transparent, with the In conclusion, we suggest that Facelina stearnsi is inner digestive gland a creamy-white color. In 1966, more properly placed in the genus Austraeolis due Bum noted that Austraeolis fucia is most likely a to the arrangement of cerata and its penial specimen of Facelina hartleyi Bum, 1962. armature. A thorough study of the Facelinidae is Austraeolis benthicola is described from a single necessary to establish phylogenetic relationships specimen collected in New South Wales. Austraeolis and determine monophyletic clades. benthicola has a conical and short penis, at its tip is a shallow rim on one side that forms a lip from which the vas deferens opens. The radula formula is ACKNOWLEDGEMENTS 28xO.1.0. Austraeolis benthicola is unique from all This research was supported by the California other Austraeolis species in having wide foot Academy of Sciences, the NSF PEET grant #0329054 margins, ovate jaws, slender radular teeth and an and a UNITAS Student Travel Grant that permitted apical rim of the penis. It is the first deep-water the senior author to attend the World Congress of aeolid recorded from Australia. Malacology in Perth, Australia July, 2004. Redescription of Facelilla stearnsi Cockerell, 1901 117

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