Balanodytes taiwanus Utinomi, 1950

Chan, Benny K. K., Cheang, Chi Chiu, Chen, I-Han & Kolbasov, Gregory A., 2013, Acrothoracican barnacles (Lithoglyptida) in Taiwan, including the taxonomic status of Balanodytes taiwanus Utinomi, 1950 and cryptic diversity of Auritoglyptes bicornis (Aurivillius, 1892), Zootaxa 3694 (3), pp. 221-239 : 225-226

Balanodytes taiwanus Utinomi, 1950 c

Figures 1 A–G, 2B–D, 3A–L, 4A–G, 5A–F

Balanodytes taiwanus Utinomi, 1950c: 454 , 459, fig. 16. — Utinomi 1950a: 1, figs 1–2. Kochlorine habei Tomlinson, 1963: 270 , figs 7–17.

Lithoglyptes habei . — Tomlinson 1969: 61–65, fig. 12.

Armatoglyptes habei . — Kolbasov & Newman 2005: 47–51, figs 5a–f, 6a, d. — Kolbasov 2009: 301, fig. 97. Armatoglyptes taiwanus . — Chan et al. 2012: 5–19.

Material examined. Syntypes, SMBL type 52. 2 specimens, in the base of Megabalanus tintinnabulum, Takao (=Kaoshiung), 1938. CEL- Acro-RYU-01, 1 specimen, in a gastropod shell, Lobster Cave, Siaoliouciou, Pingtung, Taiwan, 16 August 2010. CEL-Acro-RYU-02, same data as CEL-Acro-RYU-01. CEL-Acro-RYU-03, same data as CEL-Acro-RYU-01. CEL-Acro-sp-8, 1 specimen, in Megabalanus shell, He-Ping-Dao, Keelung, Taiwan, 17 January 2010. CEL-Acro-sp-6, 1 specimen, in Megabalanus shell, He-Ping-Dao, Keelung, Taiwan, 14 January 2010. CEL-Acro-sp-16, 1 specimen, in Turbo setosus , Shi-Ti-Ping, Hualien, Taiwan, 21 January 2010. CEL-Acrosp-15, 1 specimen, in Turbo setosus , Shi-Ti-Ping, Hualien, Taiwan, 21 January 2010. CEL-Acro-sp-3, 1 specimen, in dead coral skeleton, He-Ping-Dao, Keelung, Taiwan, date unknown. CEL-Acro-sp-10, 1 specimen, in Thais clavigera , He-Ping-Dao, Keelung, Taiwan, 18 January 2010. CEL-Acro-sp-31, 1 specimen, in the base region of Cyphastrea , He-Ping-Dao, Keelung, Taiwan, 25 February 2010. CEL-Acro-sp-13, 1 specimen, in Turbo setosus , Shi-Ti-Ping, Hualien, Taiwan, 21 January 2010. CEL-Acro-sp-6, 1 specimen, in Megabalanus shell, He-Ping-Dao, Keelung, Taiwan, 14 January 2010. CEL-Acro-sp-4, 1 specimen, in Thais clavigera shell, He-Ping-Dao, Keelung, Taiwan, 13 Jan 2010. CEL-Acro-sp-14, 1 specimen, in Thais clavigera shell, He-Ping-Dao, Keelung, Taiwan, 21 January 2010.

Diagnosis. Opercular bars with bifid teeth, posterior projections of opercular bars pronounced, recurved with 1–3 pairs of sharp spine-like teeth, lateral bars feeble, orificial knob distinct, with curved teeth, terminal cirri consisting of 4 pairs, caudal appendages 2-segmented, pedestals absent.

Descriptions. (based on syntype and our own material). Female ( Figs 1 C–G, 2C, D, 3, 4, 5). Shape of burrow opening varying from narrow oval or drop-shaped to oval ( Fig. 2 B). Opercular bars approximately 400 Μm long, armed with series of bifid teeth and simple setae ( Figs 1 C, 2C, 3B, C, F, G); posterior processes pronounced, recurved (anteriorly bent), with small bifid teeth along their length (teeth number variable, Figs 1 C, E, 3B, C), simple spine-like teeth apically ( Figs 1 E, 3B, C, F, G); comb collar long ( Fig. 3 B), with feather-like cuticular projections fused at bases ( Fig. 3 I, J), pair of bifid teeth located at posterior end of comb collar ( Fig. 3 B). Lateral surface of opercular area deep-orange when alive ( Fig. 3 A), color off-white in ethanol, covered with sparse setae and massive multifid scales ( Fig. 3 K), lateral bars feeble. Orificial knob distinct, globular-shaped, surface with setae and curved, simple or bifid teeth ( Figs 1 D, 3D, E, H, L). Attachment disc (removed in preparation) composed of several layers of striated cuticle.

Terminal cirri consisting of 4 pairs ( Fig. 4 A), orange color when alive, white when preserved in ethanol; anterior surfaces of annuli of terminal cirri armed with 2 pairs of plumose setae, posterior surfaces with sparse serrated setae ( Fig. 4 D, G). Caudal appendages 2-segmented, pedestal absent, basal segment with simple distal seta, terminal segment with 4 distal plumose setae ( Figs 1 F–G, 4B, C, E, F). Mouth cirri ( Fig. 2 D) with 4- segmented posterior ramus shorter than 4-segmented anterior ramus, both rami with long plumose setae.

Labrum saddle-like, with small, fine, blunt teeth on anterior margin ( Fig. 5 E, F). Mandible with 3 teeth in upper two-thirds, lower one-third with row of 4 sharp denticles, inferior angle with sparse setae ( Fig. 5 B). Mandibular palp trapezoid, simple setae distally along exterior margin ( Fig. 5 D). Maxillule with 2 long upper cuspidate setae, notch with short sharp seta, lower two-thirds of cutting edge with numerous short sharp setae ( Fig. 5 C). Maxilla triangular ( Fig. 5 A), long, dense setae on exterior margin, sparse setae at inferior margin.

Geographical distribution. Widespread: Maldives, Seychelles, Red Sea, Aden and Persian Gulf, Japan, Vietnam, Java, New Guinea, northern, eastern and southern Australia, Taiwan, Thailand, Philippines, Hawaii and San-Felix ( Chile), from tidal to 220 m depth (Tomlinson 1969; Kolbasov 2009; Jones et al. 1990; Poore 2012).

Remarks. In the present study, we have re-examined one of the syntypes of Balanodytes taiwanus (note the second syntype had only the opercular bar left), revealing that this species has a pair of caudal appendages (without pedestal). Thus, the redescription by Utinomi (1950a) stating that B. taiwanus lacks caudal appendages is incorrect. This is further supported by additional collections of Balanodytes taiwanus from various locations in Taiwan (including Siaoliouciou Island, which is very close to the type locality at Kaoshiung, Taiwan). All Balanodytes specimens collected in the present study have four pairs of terminal cirri and a pair of caudal appendages. The presence of caudal appendages was also observed in a very small juvenile specimen (CEL-Acro-RYU-03), suggesting that the caudal appendages are present during ontogenetic development. The definition of Balanodytes is, therefore, emended as “lithoglytid species having four pairs of terminal cirri and with a pair of caudal appendages (without pedestals)”.

Kolbasov (2000a) originally described Lithoglyptes balanodytes . Kolbasov & Newman (2005) placed this species in Armatoglyptes , herein treated as a junior synonym of Balanodytes . Newman (2011) treated B. balanodytes as a junior synonym of B. taiwanus , followed by Poore (2012). However, B. balanodytes differs from B. taiwanus in having recurved (hooked posteriorly) posterior projections of the opercular bars in the adult female (decurved – hooked anteriorly in B. taiwanus ) and by the absence of a distinct and armed orificial knob, whereas B. taiwanus has a number of curved teeth and setae on its distinct orificial knob. Therefore, we follow Kolbasov (2000a) and conclude that B. balanodytes is a different species from B. taiwanus . Tomlinson (1969) recorded B. taiwanus from the Marshall Islands, Fiji Sea (a fungi-damaged specimen) and described this species as having four pairs of terminal cirri but no caudal appendages. We have contacted the United States National Museum and confirmed Tomlinson had not deposited his material. We follow the suggestion in Kolbasov (2000a) that B. taiwanus identified by Tomlinson (1969) should be Lithoglyptes balanodytes Kolbasov, 2000 (now Balanodytes balanodytes ) and that the absence of caudal appendages in Tomlinson’s specimen might be a result of the specimen being damaged by fungi. Specimens described by Tomlinson (1963) as Kochlorine habei and later as Lithoglyptes habei (Tomlinson 1969) , and other subsequent descriptions of this species (Kolbasov 2009), correspond to A. taiwanus . Therefore, Armatoglyptes habei is treated as a junior synonym of Balanodytes taiwanus .

Figures

FIGURE 1. (A) Syntype of Balanodytes taiwanus at Seto Marine Laboratory, Kyoto University, Japan. (B) Jar of the syntypes contains the base of a Megabalanus where found Balanodytes were removed, and a vial contained two dissected syntypes of B. taiwanus. (C) Opercular region of syntype of B. taiwanus with opercular bars and orificial knob magnified in (D) and posterior process magnified in (E). (F) A pairs of caudal appendages (ca, magnified in “ G ”) were observed at the last pair of terminal cirri (tc). Abbreviations: ca— caudal appendages, tc — terminal cirri. Scale bars in Μm, except B in mm.

FIGURE 2. (A) Sampling locations in Taiwan. (B) Burrow openings of Balanodytes taiwanus Utinomi, 1950, from various substrates. (C) Balanodytes taiwanus, female, general view. (D) Balanodytes taiwanus, mouth cirrus. Abbreviations: cc — comb collar, tc— terminal cirri, ob— opercular bar, obp — posterior process of opercular bar, ok — orificial knob. Scale bars (except on ‘ B’) in µm.

FIGURE 3. Balanodytes taiwanus Utinomi, 1950, external morphology of female. (A) Lateral view of live female. (B) and (C) Posterior processes of opercular bars, opercular bars and orificial knob, showing variation in teeth number along the margin, lateral view. (D) and (E) Orificial knob, showing variation in teeth on surface, lateral view. (F) Opercular area observed under scanning electron microscope (SEM), upper lateral view. (G) Posterior process of opercular bars under SEM, showing teeth and setae, upper lateral view. (H) Orificial knob under SEM, showing teeth and sparse setae, upper view. (I) Comb collar under SEM, showing plumose projections. (J) Enlarged view of cuticular projections of comb collar. (K) Rows of massive multifid scales on opercular region. (L) Mantle teeth on surface of orificial knob. Abbreviations: cc — comb collar, ms— massive multifid scales, ob — opercular bar, obp — posterior process of opercular bar, ok — orificial knob. Scale bars in Μm.

FIGURE 4. Balanodytes taiwanus Utinomi, 1950, terminal cirri and caudal appendages of female. (A) Posterior end of thorax with bases of terminal cirri and caudal appendages. (B) and (C) Caudal appendages, 2 - segmented, without pedestal. (D) Terminal cirri and caudal appendages under scanning electron microscope (SEM), rear view. (E) and (F) Caudal appendages under SEM. (G) Rear view of terminal cirri under SEM. Abbreviations: ca — caudal appendages, tc 1 - 4 — terminal cirri, th — thorax. Scale bars in Μm.

FIGURE 5. Balanodytes taiwanus Utinomi, 1950, mouth parts of female. (A) Maxillae. (B) Mandible. (C) Maxillule. (D) Mandibular palp. (E) Labrum. (F) Enlarged view distal edge of labrum from rectangle area on “ E ” showing fine teeth. Scale bars in Μm.