Chthamalid Barnacles of Easter Island; Peripheral Pacific Isolation of Notochthamalinae New Subfamily and Hembeli-Group of Euraphiinae (Cirripedia: Chthamaloidea)
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BULLETIN OF MARINE SCIENCE, 41(2): 322-336,1987 CHTHAMALID BARNACLES OF EASTER ISLAND; PERIPHERAL PACIFIC ISOLATION OF NOTOCHTHAMALINAE NEW SUBFAMILY AND HEMBELI-GROUP OF EURAPHIINAE (CIRRIPEDIA: CHTHAMALOIDEA) B. A. Foster and W.' A. Newman ABSTRACT Euraphia devaneyi new species (of the hernbeli-group) is described from Easter Island. Rehderella belyaevi (Zevina and Kurshakova) new genus is given a supplementary description and its range extended from Easter Island to Pitcairn Island. Consideration of the pleiso- morphies and apomorphies of R. belyaevi and related genera necessitates recognition of the Notochthamalinae, a new subfamily of South Pacific species with card setae on some or all of the anterior cirri. Diagnoses are given for Notochtharnalus new genus, for N. scabrosus from South America, Nesochtharnalus new genus, for N. intertextus from widespread west Pacific localities, and Charnaesipho Darwin containing three species from New Zealand and Australia. Both chthamalids of Easter Island are peripheral Polynesian endemics, and the historical biogeography of this in connection with such young islands is discussed. Chthamalid barnacles are characteristic of rocky shores all round the world. Twenty-five species have been assigned to the genus Chthamalus or the closely related genus Euraphia (Newman and Ross, 1976; Dando and Southward, 1980). The distinction between Chthamalus and Euraphia has rested primarily on the nature ofthe mandible; quadridentoid with a combed lower margin in the former and tridentoid with a pectinate lower angle in the latter. Difficulties with this were raised by Pope (1965), particularly with Euraphia intertextus (Darwin) in which characters other than the mandible relate it to Chamaesipho columna (Spengler) which has a quadridentoid mandible. Our attention was refocused on this and related matters when we examined barnacles from Easter Island. The commonest barnacle on the basalt shores there is a chthamalid described and named Chthamalus belyaevi by Zevina and Kur- shakova (1973). It has a Chthamalus-like mandible yet is tridentoid, and shows clear shell relationships with E. intertexta of the tropical western Pacific as well as with Chthamalus scabrosus Darwin and Chamaesipho spp. of southeast and southwest shores of the South Pacific respectively. It is our conclusion that the Chthamalus-Euraphia dichotomy for the bulk of the six-plated chthamalid bar- nacles is not now adequate. The barnacle fauna of Easter Island is meager. Only three shore species are known: a new species of Euraphia described herein, the chthamalid C. belyaevi here given a supplementary description and generically redefined, and a Tesser- opora of small size and similar to specimens that have been collected from Norfolk Island and the Hawaiian Islands, and also reminiscent of T. atlantica from Ber- muda and the Azores in the Atlantic Ocean (Newman and Ross, 1977). Two stalked barnacles are known: Lepas anserifera Linnaeus was collected on floating debris (BAF, pers. obs.) and Temnaspis cf. hawaiensis (Pilsbry) was shown as commensal on the mouthparts of the lobster Panulirus pascuensis Reed by Hol- thuis (1972). This paper pays particular attention to the two Easter Island chthamalids, considers their relationship to other chthamalids of the South Pacific Ocean, and 322 FOSTER AND NEWMAN: NEW PACIFIC BARNACLES 323 discusses the consequences to our understanding of the Chthamalidae. Questions of historical biogeography of how these species came to occur in the relatively young Easter Island have been discussed in general terms by Newman and Foster (1983), and wider implications are considered by Newman and Foster (1987). SYSTEMATIC PART Family CHTHAMALIDAE Darwin, 1854 Subfamily Euraphiinae Newman and Ross, 1976 (Group of Euraphia hembeli Conrad, Nilsson-Cantell, 1921) Diagnosis. -Chthamalid barnacles with six parietal shell plates including rostrum, carina and paired latera and rostrolatera. Scutum higher than wide, interlocked but not concrescent with tergum. Mandible incisiform, tridentoid, and with pec- tinate lower angle. Cirri without card setae. Euraphia devaneyi new species Figures 1A-E, 2 Material Examined. -Easter Island-Two specimens, Hanga Roa, intertidal, 5 February 1965 (I. Elford coli.); several specimens, Hanga Nui, in tide pools, 30 October 1974 (H. Rehder coil., no. E-27, USNM file no. 313647, included 2nd paratype); several specimens, Moto Nui, intertidal rocks, I November 1974, (H. Rehder coil., no. E-33, included holotype and 1st paratype); Hanga Roa, Vinapu, Vaihu, Anakena, 20-23 November 1976 (BAF coil., included 3rd paratype lot); four specimens, from 5 m of depth on dead Pocillopora, Motu Nui Bay, 18 February 1983 (L. H. DiSalvo coli.). Deposition of Types. -(1) Holotype, U.S.N.M. Cat. No. 233266 (dissected); (2) 1st paratype, U.S.N.M. Cat. No. 233267 (in alcohol, undissected); (3) 2nd paratype, Brit. Mus. (Nat. Hist.) Reg. No. 1987-1 (dissected); (4) 3rd paratype lot, Brit. Mus. (Nat. Hist.) Reg. No. 1987-2 (undissected). Distribution. -Easter Island (Fig. 4). Etymology. -Named in honor of the late Dr. Dennis Devaney, Bishop Museum, Honolulu, who provided us with the specimen of E. hembeli he and Dr. J. Randall collected from Pitcairn Island. Description. - In un eroded specimens (Fig. IA), shell conic, apexes of parietes projecting, externally white and strongly ribbed but deeper shell matrix grey-pink. Radii narrow with sinuous margins interlocking with adjacent parietal margin. Alae of carina wide, extending to lateral angle of orifice, midway on latera. Basal edge of parietes radially ridged. Eroded specimens externally showing pink laminae and obliterated radial sutures; internally with secondary calcification lining the entire body chamber of the shell. Base calcareous, white; in larger specimens overlain internally and peripherally with pink ledge of secondary parietal calcification. Scuta and terga (Fig. 1B-E) strongly interlocked but not calcified together, darkly vinaceous internally, brown externally when uneroded. Scutum very narrow, with thick articular margin elevating sinuous inner tergal margin and forming apical edge between inner and outer faces of plate; adductor muscle pit prominent; lateral depressor muscle crests at rostral and tergal angles. Tergum with strong and pectinate depressor muscle crests, spur truncated at basi-scutal angle. Maximum Shell Size. - Forty-five millimeter rostrocarinal diameter, 24-mm high. Labrum (Fig. 2A) shallowly concave, with row of minute teeth and fine setae on edge. Mandible (Fig. 2B) tridentoid, denticulate between, with closely pectinate lower angle. Maxillule (Fig. 2C) with two large spines at slightly protuberant upper 324 BULLETIN OF MARINE SCIENCE, VOL. 41, NO.2, 1987 &!:L1mm L...-lmm ,,,II lmm I BCDE Figure 1, A-E, Euraphia devaneyi new sp, A, shell from left side; B-E, terga (B, D) and scuta (C, E), exterior and interior views; F-I, Rehderella be/yaevi new genus F, juvenile from Pitcairn Island; G, H, shell from Easter Island viewed from above and below; 1, fused opercula viewed obliquely from above. angle, double row of slender spines in central part of blade, and more or less prominent lower angle with short spines. Numbers of segments in rami of cirri of two specimens as follows (anterior ramus first): rostrocarinal diameter I II III IV V VI 15 mm 8, 7 6,10 15, 16 19, 19 21,20 20,21 42mm 8, 7 10,16 17, 19 24, 27 27,28 27,25 FOSTER AND NEWMAN: NEW PACIFIC BARNACLES 325 I kC J-L .25mm IE-I .5mm K Figure 2. Euraphia devaneyi new sp., appendages of a 15-mm basal diameter specimen: A, labrum and mandibular palps; B, mandible; C, maxillule; D-F, cirri I-III with setae omitted but showing posterior distal spines; 0-1, basal parts of cirri IV-VI; J-L, segments 22, 3 and 12 of the anterior ramus of cirrus VI. Cirrus I (Fig. 2D) with short rami of wide segments, very setose medially with many finely serrate setae, with prominent spines on outer face of the anterior ramus, and especially prominent at distal angles at posterior edges of both rami. Cirrus II (Fig. 2E) with the anterior ramus short and like the posterior ramus 326 BULLETIN OF MARINE SCIENCE, VOL. 41, NO.2. 1987 of cirrus I. Posterior ramus longer and narrower with basal five segments with finely serrate setae medially, more distal segments like those of cirri IV-VI. Stout spines present at posterior angles of basal segments, particularly in anterior ramus. Cirrus III (Fig. 2F) with similar length rami, basal three segments of anterior ramus with finely serrate setae medially, and basal five or six segments of both rami with stout distal posterior spines, stouter in anterior ramus. Cirri IV-VI (Fig. 20-L) with similar rami, segments with 3-5 pairs ofprominent setae on anterior edge (more than three in larger specimens) and single short seta between bases of each large pair. Basal five or six segments of anterior rami with stout distal posterior spines. Habitat. -Large specimens were collected from rock surfaces covered with cor- alline and filamentous algae on surf shores, occurring as solitary individuals or in clumps of four or five specimens. Subtidally, E. devaneyi extends to depths of 10m being more cryptic with increasing depth. Subtidal specimens were smaller than intertidal ones. Dr. Luis DiSalvo has observed a crab, Carpi/ius convexus, eating E. devaneyi at 3-m depth, and has recorded the bamacle from the stomach contents of puffer fish, Arothron meleagris. The larger size of shallower specimens may reflect relative immunity from the small-sized rasping and picking fishes of Easter Island. New Subfamily Notochthamalinae Diagnosis. -Chthamalid barnacles with six parietal shell plates, ontogenetically reduced to four in Chamaesipho, including rostrum, carina and paired latera and rostrolatera, in some species concrescent. Scutum elongate; tergum narrow and deeply interlocked, sometimes concrescent with scutum of same side. Mandible with or without a combed lower edge. Card setae develop on some or all of cirri I to III. Etymology. -OK notos = south, with reference to the predominantly South Pacific biogeography of the species included in this subfamily.