Zoosyst. Evol. 96 (2) 2020, 649-698 | DOI 10.3897/zse.96.55733 yee BERLIN New insights gained from museum collections: Deep-sea barnacles (Crustacea, Cirripedia, Thoracica) in the Muséum National d’Histoire Naturelle, Paris, collected during the Karubar expedition in 1991 Pipit Pitriana!:*°, Diana S. Jones’, Laure Corbari*, Kristina von Rintelen! Museum fiir Naturkunde — Leibniz Institute for Evolution and Biodiversity Science, Invalidenstrasse 43, 10115, Berlin, Germany Research Centre for Deep-sea, Indonesian Institute of Science (LIPI), JI. Y. Syaranamual, Poka, Tlk. Ambon, Kota Ambon, Maluku, Indonesia The Western Australian Museum, 49 Kew Street, Welshpool WA 6106, Locked Bag 49, Welshpool DC WA 6986, Australia Muséum national d Histoire naturelle, Institut de Systématique, Evolution, Biodiversité ISYEB — UMR 7205 — CNRS, MNHN, UP MC, EPHE, 57 rue Cuvier, CP 26, 75005, Paris, France 5 Institute of Geological Sciences, Freie Universitat Berlin, Malteserstrasse 74-100 Building C and D, 12249, Berlin, Germany bh WN = http://zoobank.org/DF 25E94F-E DEC-4F D4-BA 1D-E4AC288282AD Corresponding author: Pipit Pitriana (pipit.pitriana@mfn.berlin) Academic editor: Martin Husemann @ Received 23 June 2020 # Accepted 13 August 2020 Published 28 September 2020 Abstract An examination of the deep-sea barnacles (Cirripedia, Thoracica) collected by the Karubar expedition to Indonesia (1991) and deposited in the Muséum National d’ Histoire Naturelle, Paris, identified 40 species contained in three families of stalked and five families of acorn barnacles. Information on these species 1s presented, including descriptions, updated distributions and images to aid species identification. Thirty of the species, treated herein, are new records for the Indonesian Kei Islands and Tanimbar Island, which increases the total number of species recorded from Kei Islands, Aru Island and Tanimbar Island to 40. This study demonstrates the value of museum collections as a resource in biodiversity science. Key Words acorn barnacles, Indonesian biodiversity, new record, stalked barnacles, taxonomy Introduction In 1991, scientists from France and Indonesia conducted collaborative research through the Karubar expedition in Indonesia. The acronym for this expedition, which collect- ed the material reported on herein, is a contraction of the names of the Kei, Aru and Tanimbar Islands. These Islands attracted attention after Professor Th. Mortensen’s Danish expedition to the Kei Islands (1914—16). Mortensen sug- gested that the Islands were an ideal place for a marine laboratory to study deep-sea fauna, as he had found stalked crinoids, elasipods and other abyssal creatures at depths of 200-400 m around the Kei Islands (Crosnier et al. 1997). The Karubar expedition was part of the MUSOR- STOM-Tropical Deep-Sea Benthos programme (1976— present). This programme was a collaboration between the Muséum National d’ Histoire Naturelle (MNHN), Par- is and the Institut de Recherche pour le Développement (IRD) (formerly ORSTOM), to explore the deep-sea fau- na of the tropical Indo-Pacific. As the programme was inspired and guided by carcinologists, it is not surprising that ~ 33% of the papers resulting from these cruises con- cern crustaceans, especially crabs, lobsters and shrimps (Richer de Forges et al. 2013). Publications on the thoracic barnacles collected by the Karubar expedition are those of Buckeridge (1994, 1997) who focused on barnacles from the suborder Verru- comorpha. He reported six species of verrucomorph bar- nacles: Newmaniverruca albatrossiana (Pilsbry, 1912), Altiverruca navicula (Hoek, 1913), Brochiverruca dens Copyright Pipit Pitriana et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 650 (Broch, 1932), Metaverruca recta (Aurivillius, 1898), Rostratoverruca intexta (Pilsbry, 1912) and R. kruegeri (Broch, 1922). Here, we present a study of the remaining barnacle material collected by the Karubar expedition, to cover the entire taxonomic range of deep-sea barnacles collected by this expedition. Material and methods In total, 459 specimens were studied at the Muséum Na- tional d’ Histoire Naturelle (MNHN), of which 89 speci- mens were studied in detail at the Museum fiir Naturkunde (MfN), Berlin, Germany. Species were determined on ex- ternal shell morphology and mouthpart and arthropodal characters, as described in Darwin (1852, 1854), Hoek (1883, 1907, 1913), Pilsbry (1890, 1907, 1916), Annan- dale (1905, 1906, 1908, 1909, 1910, 1913, 1916), Broch (1916, 1922, 1931), Foster (1974, 1978, 1980, 1981), Newman and Ross (1971, 1976), Rosell (1981, 1989, 1991), Calman (1918, 1919), Ren (1989), Chan et al. (2009, 2010) and Shalaeva and Boxshall (2014). Hard body parts, such as the shell (parietes) and the opercular plates, were studied using a Leica M125 stereo- microscope. Soft body parts, such as mouthparts and cir- ri, were studied using an Axioskop 20 light microscope. Mouthparts and cirri were mounted on glass slides and examined under a light microscope. Images were taken using a stereomicroscope with a digital camera (Leica Microsystems M205C and Leica Z16 APo-A). In the “Taxonomic account” section, the diagnosis is followed by description, type species and localities and known distributions for each species are given. Each Species 1s listed under its current valid binomen and all known synonyms are also provided. Photographs of each Species are provided to facilitate their recognition. In the “Material examined” section, there are four station codes dependent on the type of dredge used and are listed as follows: DW for dredge Warén, ED (drague épibenthique) for epibenthic dredge, CP (chalut a perche) for beam trawl and CC (chalut a crevettes) for otter trawl (shrimp). Measurements were made using digital callipers (ac- curate to 0.1 mm) as follows: for acorn barnacles, bas- al length of shell (LB), basal width (WB), orifice length (LO), orifice width (WO) and carinal height (H); for stalked barnacles, total height (TH), capitular height (CH), diameter of base of capitulum (DBC), distance be- tween carina and scutum (CS), scutal length (LS), scutal width (WS), tergal length (LT) and tergal width (WT). Results Morphological analyses of all the samples revealed 40 species from three families of stalked barnacles (Heter- alepadidae: two genera, three species; Poecilasmatidae: four genera, five species; Scalpellidae: twelve genera, 21 species) and five families of acorn barnacles (Verrucidae: zse.pensoft.net Pitriana, P et al.: Deep-sea barnacles from the Karubar expedition (1991) two genera and species; Pachylasmatidae: one genus and species; Archaeobalanidae: three genera, six species; Pyr- gomatidae: one genus and species; Balanidae: one genus and species). A list of the species of Cirripedia collected during the Karubar expedition and deposited in the Muséum National d’ Histoire Naturelle (MNHN) Paris is present- ed, including additional information on substrate, as well as the depth where the sample was found (Suppl. material 1: Table S1). Taxonomic account Class HEXANAUPLIA Oakley, Wolfe, Lind- gren & Zaharoff, 2013 Subclass THECOSTRACA Gruvel, 1905 Infraclass CIRRIPEDIA Burmeister, (= Cirrhipédes Lamarck, 1806) Superorder THORACICA Darwin, 1854 Order LEPADIFORMES Buckeridge & New- man, 2006 Suborder HETERALEPADOMORPHA New- man, 1987 1834 Diagnosis. Capitulum and peduncle without calcareous armament. According to Buhl-Mortensen and Mifsud (2017), Heteralepadomorpha consists of seven families of poorlyknown Pedunculata. Family HETERALEPADIDAE Nilsson-Cantell, 1921 Heteralepadidae Nilsson-Cantell, 1921: 245 Lepadidae Darwin, 1852: 8 (part.) Diagnosis. Capitulum naked, thick-walled, globular, sup- ported by stout peduncle; filamentary appendage at base of cirrus I; caudal appendage multi-segmented; cirri cten- opod or acanthopod, rarely lasiopod. Type genus. Heteralepas Pilsbry, 1907a. Remarks. The family consists of two genera: Heterale- pas Pilsbry, 1907a and Paralepas Pilsbry, 1907a. Genus Heteralepas Pilsbry, 1907 Alepas Darwin, 1852: 156, pl. III, figs 5, 6. Heteralepas (Heteralepas) Pilsbry, 1907a: 100. Heteralepas s. str. — Newman, 1960: 109. Diagnosis. Filamentary appendage present at base of cir- rus I, short; posterior rami of cirri V and VI shorter and more slender than anterior ram1. Zoosyst. Evol. 96 (2) 2020, 649-698 Type species. A/epas rex Pilsbry, 1907c: 186, fig. 3, pl. IV fig. 7; type locality: Kauai Island, Hawaiian waters. Heteralepas japonica (Aurivillius, 1892) Figure | Alepas japonica Aurivillius, 1892: 125. — 1894: 28, pl. II figs 14, 15, pl. VUI, figs 3, 7, pl. IX, fig. 3. Alepas indica Gruvel, 1901: 259. — 1905a: 162, fig. 179. Heteralepas (Heteralepas) japonica. — Pilsbry, 1907a: 101. Heteralepas (Heteralepas) japonica var. alba Kriger, 1911: 34, pl. 1, fig. 2b. Heteralepas (Heteralepas) dubia Broch, 1922: 288, fig. 38. Heteralepas japonica. — Pilsbry 1911: 71, fig. 4. — Zevina et al. 1992: 31, fig. 19. — Chan et al. 2009a: 61. — Chan et al. 2009b: 88-91, figs 2A—D, 3A—D, 4, 5. — Pitriana et al. 2020: 12, fig. 3. Material examined. Tanimbar Island: 1 specimen, MNHN-IU-2019-4877, stn. CP 46, 08°01'S, 132°51'E, 271-273 m depth, 29 October 1991. Diagnosis. Capitulum rounded, plates absent; wall of ca- pitulum thick; orifice crenulated with no more than two crests on carinal region; cirrus VI with caudal appendage; maxillule strongly notched. Description. Capitulum and peduncle yellowish. Orifice slightly protuberant, crenulated, occupying one half to one third capitular length, parallel to or at oblique angle to capitulum; integument thick, chitinous. Cirrus I with anterior rami (20-segmented) shorter than posterior rami (25-segmented); cirrus VI with long caudal appendage, 24-segmented, 2/3 length of anterior ramus. Mandibles with four large teeth excluding inferior angle; maxillule strongly notched, two large setae at upper angle, blade- shaped setae on cutting margin; labrum concave with nu- merous teeth. Measurements of specimen: height of ca- 651 pitulum 18.50 mm, width 12.82 mm, thickness 5.97 mm; length of peduncle 15.26 mm, width 5.55 mm. Distribution. Singapore; Indo-west Pacific: Indian Ocean; Australia; Malay Archipelago; Vietnam; Condor Island; South China Sea; East China Sea; Taiwan, Philip- pines; South Japan; NE New Zealand; fouling hard rock substrata, crabs, gorgonians, antipatharians, deep-sea ca- bles; 48-500 m depth (Jones and Hosie 2016). Recently, the species has also been recorded from the Lifamatola Sea and the Halmahera Sea, Indonesia (Pitriana et al. 2020). In this study, Heteralepas japonica was found at Tanimbar Island, Indonesia. Type locality. Hirado Strait, Japan, 146 m depth (Auriv- illtus 1892). Genus Paralepas Pilsbry, 1907 Alepas Darwin, 1852: 156 (part.). Heteralepas (Paralepas) Pilsbry, 1907a: 100, fig. 34A. Paralepas s. str. — Newman, 1960: 108. Diagnosis. Maxillule slightly notched; filamentary ap- pendages well-developed; posterior and anterior rami of cirri V and VI similar in length; segments of cirri V and VI square-shaped, each armed with semi-circular brush of many setae on anterior face. Type species. Alepas percarinata Pilsbry, 1907c: 185, fig. 2, pl. IV, fig. 8; type locality: Molokai, Hawaiian waters. Paralepas minuta (Philippi, 1836) Figure 2 Alepas minuta Philippi, 1836: 254, pl. 12, fig. 23. Alepas (Paralepas) minuta. — Weltner, 1897: 239. Figure 1. Heteralepas japonica (Aurivillius, 1892) (MNHN-IU-2019-4877). a. left lateral view showing the capitulum and pedun- cle; b. right lateral view showing the capitulum and peduncle; c. anterior view, orifice; d. posterior view, capitulum and peduncle. Scale bar: 6 mm (a-—d). zse.pensoft.net 652 Pitriana, P et al.: Deep-sea barnacles from the Karubar expedition (1991) Figure 2. Paralepas minuta (Philippi, 1836) (MNHN-IU-2019-4882). a. left lateral view, capitulum and peduncle; b. right lateral view, capitulum and peduncle; c. anterior view, orifice. Scale bar: 5 mm (a—c). Heteralepas (Paralepas) minuta. — Broch, 1927b: 18, fig. 4, pl. 1, figs 1, 2. Paralepas minuta nipponica Utinomi, 1970: 342. Paralepas minuta. — Stubbings 1967: 240. — Newman 1960: 109. — Chan et al. 2009b: 66, figs 55-57. Material examined. — Kei Islands: 6 specimens, MNHN- TU-2019-4882, Stn. CP 25, 05°30'S, 132°52'E, 336-346 m depth, 26 October 1991, attached to dead stem of gorgonian. —Tanimbar Island: 4 specimens, MNHN-IU-2019-4883, Stn. CP 85, 09°22'S, 131°14'E, 240-245 m depth, 4 No- vember 1991, attached to spine of sea urchin. Diagnosis. Capitulum yellowish with globular shape; ante- rior and posterior rami of cirrus V and VI similar in length. Description. Capitulum yellowish, globose, small-sized, externally smooth; orifice crenulated; peduncle rather short. Segments of rami of cirrus I wide, anterior ramus (9-segmented) shorter than posterior ramus (13-seg- mented); cirrus VI with short caudal appendage, 8-seg- mented. Penis annulated, setae sparsely distributed over most of surface, a few longer setae towards tip. Maxillule notched, two large and one small setae at upper angle; mandible with four major teeth; labrum concave, with numerous teeth. Measurements of specimen: height of capitulum 8.34 mm, width 6.67 mm, thickness 4.53 mm; length of peduncle 3.84 mm, width 2.60 mm. Distribution. Java Sea (Indonesia); Mediterranean Sea; West Africa; Indo-west Pacific: Indian Ocean; Australia; Malay Archipelago; Taiwan; Japan; Philippines; North- ern New Zealand; 485-736 m depth; attached to spines of cidarids (Jones and Hosie 2016). In this study, Para- lepas minuta was found at Kei Islands and Tanimbar Is- land, Indonesia. zse.pensoft.net Type locality. Mediterranean Sea (Chan et al. 2009b). Remarks. Paralepas minuta (Philippi, 1836) can be dis- tinguished from Paralepas ovalis (Hoek, 1907) by the presence of triangular scutal areas and from Paralepas nodulosa Broch, 1922 by the absence of “well defined, scanty, almost thorn-like warts” (Broch 1922). Paralepas morula (Hoek, 1907) Figure 3 Alepas morula Hoek, 1907: 35, pl. IV, figs 9-12. Heteralepas (Paralepas) morula. — Broch, 1922: 28, fig. 34. Paralepas morula. — Newman, 1960: 109. Material examined. — Tanimbar Island: 45 specimens, MNHN-IU-2019-4878, Stn. CP 54, 08°21'S, 131°43'E, 836-869 m depth, 30 October 1991, attached to spine of sea urchin. Diagnosis. Capitulum globular with numerous tubercles on surface, scuta absent, orifice small, not protuberant, peduncular length more than half capitular length. Description. Capitulum and peduncle yellowish. Capit- ulum globular with thick, chitinous, large warts all over surface. Peduncle rather long, narrower than capitulum; surface bearing tubercles, smaller than those of capitulum. Cirrus I placed rather far from cirrus IT; cirri IVI similar, length and number of segments slightly increasing from second to sixth; cirrus VI with 7-segmented caudal append- age. Penis indistinctly segmented, few long hairs towards tip. Maxillule notched, two large setae on upper notch; mandibles with three major teeth, inferior angle considered Zoosyst. Evol. 96 (2) 2020, 649-698 653 Figure 3. Paralepas morula (Hoek, 1907) (MNHN-IU-2019-4878). a. left lateral view, capitulum and peduncle; b. right lateral view, capitulum and peduncle. Scale bar: 3 mm (a, b). as fourth tooth; labrum with continuous row of numer- ous short, strong, blunt teeth. Measurements of specimen: height of capitulum 5.14 mm, width 3.69 mm, thickness 1.81 mm; length of peduncle 5.08 mm, width 1.31 mm. Distribution. Flores Sea (Indonesia); Philippines; Bass Strait, N of Tasmania; attached to spines of echinoids, for example, Histocidaris elegans (Agassiz, 1879), 182- 538 m depth (Jones and Hosie 2016). In this study, Para- lepas morula was found at Tanimbar Island, Indonesia. Type locality. Bali Sea, North of Lombok; 538 m depth (Hoek 1907). Remarks. Most of the smaller specimens do not show the warts that are characteristic of this species. Suborder LEPADOMORPHA Pilsbry, 1916 Lepadomorpha Pilsbry, 1916: 14 (Lepadidae sensu Darwin, 1852). Lepadoidea Darwin, 1852: 8 (nom. trans. Zevina 1978a, b). Family POECILASMATIDAE Annandale, 1910 Lepadidae Darwin, 1852: 8 (part.). — Pilsbry 1907a: 72 (part.). Lepadinae. — Gruvel, 1905: 104 (part.). Poecilasmatinae Annandale, 1910a: 84. Poecilasmatidae. — Nilsson-Cantell, 1921: 253. Trilasmatidae Nilsson-Cantell, 1934: 40. — Hiro 1937a: 79. — Kriiger 1940: 29. Diagnosis. Capitulum with up to five plates, including tergum and scutum; scutum in some species split into two (resulting in seven plates); some or all plates may be degenerate or absent; umbos of terga apical, those of carina and scuta fundamentally basal; peduncle without calcareous scales; maxillule not stepped; cirrus I widely separated and much shorter than posterior cirri; caudal appendages uniarticulate, spinose. Type genus. Poecilasma Darwin, 1852: 99. Genus Dianajonesia Kocak & Kemal, 2008 Temnaspis Fischer, 1884: 357. — Broch 1931: 30. Poecilasma (part). — Darwin, 1852: 99. — Pilsbry 1907a: 82. — Nils- son-Cantell 1921: 253. Dianajonesia Kogak & Kemal, 2008: 2. Diagnosis. Capitulum with five plates, approximate or variously reduced; scutum divided into two parts; apex of carina extending to, or slightly overlapping basal end of terga, basally terminating 1n disc. Type species. Poecilasma fissa Darwin, 1852: 109, pl. I, fig. 4, pl. X, fig. 29. zse.pensoft.net 654 Dianajonesia amygdalum (Aurivillius, 1894) Figure 4 Poecilasma amygdalum Aurivillius, 1894: 10, pl. 1, figs 4-6, pl. 8, fig. 4. — Nilsson-Cantell 1921: 262, fig. 46d—g, pl. 3, fig. 6. Poecilasma fissum (non Darwin). — Hoek, 1907: 8 (part.). Trilasmis fissum hawaiense. — Pilsbry, 1928: 306, pl. 24, figs 1-8. Trilasmis (Temnaspis) amygdalum. — Hiro, 1937a: 85, fig. 69. Temnaspis amygdalum amygdalum. — Zevina, 1982: 70, fig. 62. Temnaspis amygdalum. — Utinomi, 1966: 5.— Dong et al. 1982: 77. Dianajonesia amygdalum. — Kogak & Kemal, 2008: 2. Material examined. — Tanimbar Island: 2 specimens, MNHN-IU-2019-4861, Stn. CP 79, 09°16'S, 131°22'E, 239-250 m depth, 3 November 1991, attached to crab leg. Diagnosis. Capitulum broadly oval, with five smooth plates; carina not extending to area between terga; pedun- cle with circles of small protuberances; cirri short. Description. Capitulum yellowish, with five tranparent, smooth plates. Scutum bilobed, occludent segment nar- row, bow-shaped; tergum sub-triangular, nearly half as broad as long. Carina very narrow, slightly curved down- wards, end blunt. Peduncle yellow, plainly ringed, spines absent. Cirrus I with anterior ramus wider than posterior ramus. Maxillule notched, with two strong teeth in upper part; mandible with four teeth, fourth pectinated, placed very close to inferior angle; labrum with row of minute teeth. Measurements of specimen: basal diameter of ca- pitulum 0.94 mm; capitular height 7.03 mm; total height 10.40 mm; scutal width 4.11 mm; scutal length 5.85 mm; tergal width 1.00 mm; tergal length 3.13 mm. Distribution. Indo-west Pacific: Indian Ocean; Madagascar through Malaysia, Hong Kong, South China Sea; Taiwan; Pitriana, P. et al.: Deep-sea barnacles from the Karubar expedition (1991) Philippines; South Japan; tropical West and central Pacific Ocean to Fiji and Hawaii; attached to decapod crustaceans; shallow water (Jones and Hosie 2016). In this study, Diana- Jonesia amygdalum was found at Tanimbar Island, Indonesia. Type locality. Nordwachter Island, Thousand Islands, Java Sea (Aurivillius 1894). Dianajonesia excavatum (Hoek, 1907) Figure 5 Poecilasma excavatum Hoek, 1907: 10, pl. I, figs 5-10. Poecilasma tridens. — Weltner 1922: 80, pl. 4, fig. 19 (non Aurivillius). Poecilasma (Temnaspis) excavatum. — Nilsson-Cantell 1925: 16, fig. 5, pl. 1, fig. 1. Trilasmis (Temnaspis) excavatum. — Hiro 1937b: 412. Dichelaspis (Dichelaspis) tridens. — Stubbings 1936: 7, fig. 2. (non Au- rivillius). Trilasmis excavatum. — Nilsson-Cantell 1938: 9. Temnaspis excavatum. — Broch 1931: 31, fig. 10. — Dong et al. 1982: 77. Dianajonesia excavatum. — Kogak & Kemal, 2008: 2. Material examined. — Tanimbar Island: 3 specimens, MNHN-IU-2019-4859, Stn. CP 78, 09°06'S, 131°24'E, 284-295 m depth, 3 November 1991. Diagnosis. Capitulum with scutum divided into two seg- ments parallel to occludent margin; tergum with excava- tion in scutal margin near occludent margin, receiving tip of occludent segment of scutum. Description. Capitulum oval, apex pointed, slightly thick, swollen. Scutum with larger segment strongly bowed, basal margin short, apex pointed; smaller segment bowed, termi- nating in point at base, tergal margin rounded, fitting exactly Figure 4. Dianajonesia amygdalum (Aurivillius, 1894) (MNHN-IU-2019-4861). a. left lateral view, capitulum and peduncle; b. right lateral view, capitulum and peduncle; ¢. anterior view, orifice; d. posterior view, carina. Scale bar: 2 mm (a—d). zse.pensoft.net Zoosyst. Evol. 96 (2) 2020, 649-698 655 Figure 5. Dianajonesia excavatum (Hoek, 1907) (MNHN-IU-2019-4859). a. left lateral view showing the capitulum and peduncle; b. right lateral view showing the capitulum and peduncle; ec. anterior view, orifice; d. posterior view, carina. Scale bar: 2 mm (a—d). into excavation of tergum. Tergum triangular, characteristic excavation at scutal margin near occludent margin. Carina narrow, terminating in spatula-shaped disc. Cirrus I with anterior and posterior rami subequal (each five-segment- ed); cirri JI—VI longer, more slender; cirrus VI with caudal appendages. Penis thick, ringed, especially mid-length, ter- minating in narrower, curved part. Maxillule notched, two large teeth on upper side; mandible with four teeth, largedis- tance between first and second teeth. Labrum convex, with numerous blunt teeth. Measurements of specimen: basal diameter of capitulum 1.06 mm; capitular height 7.12 mm; total height 12.69 mm; scutal width 3.08 mm; scutal length 6.09 mm; tergal width 1.30 mm; tergal length 3.47 mm. Distribution. Indo-west Pacific: East coast of Africa (Zanzibar); Gulf of Aden; Indonesia; Malay Archipela- go; South China Sea; East China Sea; Philippines; South Japan (Goto Island); attached to echinoid spines, crusta- ceans; palinurids and cirripedes; 189-600 m depth (Jones and Hosie 2016). In this study, Dianajonesia excavatum was found at Tanimbar Island, Indonesia. Type locality. Siboga station 253; 5°48 .2'S, 132°15'E; depth: 304 m; bottom: grey clay, hard and crumbly (Hoek 1907). Genus Dichelaspis Darwin, 1852 Dichelasmis Darwin, 1852: 115. Heptalasmis. Agassiz, 1842: 178. Octolasmis Gray, 1825: 100. — Pilsbry 1907a: 93. Diagnosis. Capitulum with five plates; scutum with two distinct segments, united at rostral angle; carina generally extending up between terga, terminating downwards as imbedded disc (fork- or cup-shaped). Type species. Octolasmis warwicki Gray, 1825: 100. — 1830: pl. VI, fig. 16. Dichelaspis orthogonia Darwin, 1852 Figure 6 Dichelaspis orthogonia Darwin, 1852: 130, pl. IL, fig. 10a, b. Dichelaspis versluysi Hoek, 1907: 28, pl. Ul, figs 8-13. Octolasmis (Dichelaspis) orthogonia. — Pilsbry, 1907a: 94. Octolasmis orthogonia. — Kriger, 1911: 462. — Pilsbry 1911: 70, pl. 11, figs 6, 7. — Broch 1922: 279. — 1931: 38, fig. 14. — Nilsson-Cantell 1925: 21, fig. 8. — 1928: 18, fig. 8. — Hiro 1933: 55, fig. 16, pl. 2, figs 5a, 5. — 1937a: 91, fig. 71. — 1937b: 415, fig. 12. — Stubbings 1963: 327, fig. 1. — Dong et al. 1982: 79. — Zevina et al. 1992: 26, fig. 15. — Chan 2009: 68, fig. 2 B and fig. 17A—F. Material examined. — Kei Islands: 4 specimens, MNHN-IU-2019-4864, Stn. CP 27, 05°33'S, 132°51'E, 314-304 m depth, 27 October 1991. — Tanimbar Island: 6 specimens, MNHN- TU-2019-4862, Stn. DW 24, 05°32'S, 132°51'E, 230- 243 m depth, 26 October 1991. zse.pensoft.net 656 Pitriana, PR et al.: Deep-sea barnacles from the Karubar expedition (1991) Figure 6. Dichelaspis orthogonia Darwin, 1852 (MNHN-IU-2019-4864). a. left lateral view capitulum and peduncle; b. right lateral view, capitulum and peduncle. Scale bar: 2 mm (ab). Diagnosis. Capitulum flattened; five plates, separated by thin membrane; scutum consisting of two distinct seg- ments, with point of junction perfectly calcified; base of carina oval. Description. Capitulum and peduncle yellowish. Scutum with basal segment narrow, L-shaped; tergum triangular, three prominent ridges at scutal margin; base of carina crescent-formed cup. Cirrus I with anterior ramus (6-seg- mented) shorter than posterior ramus (8-segmented); cir- rus VI with small caudal appendage. Maxillule notched, three large setae on upper notch; mandible with four teeth, inferior angle produced into single strong spine; labrum concave, with numerous teeth. Measurements of specimen: basal diameter of capitulum 1.89 mm; capitu- lar height 8.69 mm; total height 10.47 mm; scutal width 3.5 mm; scutal length 7.16 mm; tergal width 0.80 mm; tergal length 6.53 mm. Distribution. Indo-west Pacific: East and South African Coast, Indian Ocean, Australia, Indonesia, Malay Ar- chipelago, Vietnam, South China Sea, East China Sea, Philippines, Tatwan, South Japan; attached to hydroid, gorgonians, antipatharians, sea urchin spines, nylon cord, zse.pensoft.net sometimes fouling; 14-818 m depth (Jones and Hosie 2016). In this study, Dichelaspis orthogonia was found at Kei Islands and Tanimbar Island, Indonesia. Type locality. Unknown. Remarks. The type locality of this species is unknown. The species was re-discovered in the Malay Archipelago by the Siboga Expedition in 40-112 m depth (Hoek 1907) and the A/batross Expedition, off Kagoshima Gulf, Japan, in 87 m depth (Pilsbry 1907d). Genus Glyptelasma Pilsbry, 1907 Glyptelasma Pilsbry, 1907a: 87. Diagnosis. Capitulum with five approximate, fully cal- cified plates; scutum quadrangular, umbone subcentral; carina becoming larger towards base; peduncle short. Type species. Megalasma (Glyptelasma) subcarinatum Pilsbry, 1907a: 91, pl. VU, figs 1—5; type locality: Atlan- tic Ocean, East of New Jersey, USA. Zoosyst. Evol. 96 (2) 2020, 649-698 Glyptelasma gracile (Hoek, 1883) Figure 7 Poecilasma gracile Hoek, 1883: 46, pl. I, figs 2-4. Megalasma gracile gracilius. — Pilsbry, 1907a: 88, pl. V, fig. 16, pl. VU, figs 6-9. Megalasma gracile. — Nilsson-Cantell 1938: 10. Megalasma (Glyptelasma) gracile. — Zevina 1982: 84, fig. 74. Glyptelasma gracile. — Jones et al. 2001: 239. Material examined. — Tanimbar Island: 14 specimens, MNHN-IU-2019-4873, Stn. CP 54, 08°21'S, 131°43'E, 836-869 m depth, 30 October 1991, attached to spine of sea urchin; 1 specimen, MNHN-IU-2019-4875, Stn. CC 57, 08°19'S, 31°53'E, 603-620 m depth, 31 Octo- ber 1991; 3 specimens (1 adult, 2 juveniles), MNHN- ITU-2019-4876, Stn. CP 73, 08°29'S, 131°33'E, 840- 855 m depth, 2 November 1991, attached to spines of sea urchins. Diagnosis. Capitulum with five plates; carina enlarged, keel-shaped; tergum with basal point truncated; max- illule notched; caudal appendages with relatively-long spines at tip. Description. Capitulum whitish, peduncle brownish. Capitulum compressed, striated, about twice as long as broad. Scutum with apex pointed; tergum basally trun- cated, flat, oblong. Carina flat, very narrow, enlarged, keel-shaped in basal region. Cirrus I with anterior and posterior rami equal length; cirrus II-VI long, slender; cirrus VI with small caudal appendages. Penis with few 657 long hairs towards tip. Maxillule notched, with strong teeth on upper side; mandible with four teeth; labrum with row of small teeth on crest. Measurements of spec- imen: basal diameter of capitulum 1.45 mm; capitular height 10.44 mm; total height 13.99 mm; scutal width 4.79 mm; scutal length 9.01 mm; tergal width 1.87 mm; tergal length 5.07 mm. Distribution. West-southwest Pacific: from Australia (Sydney) to Indonesia (Sumbawa) and the Philippines (northwest Panay Island); East Indian Ocean; attached to glassy spicule of hexactinellid sponge, fragments of sea urchin tests, spines of cidarids; 395-935 m depth (Jones and Hosie 2016). In this study, G/yptelasma gracile was found at Tanimbar Island, Indonesia. Type locality. Challenger expedition Station 164a (off Sydney, Australia); 34°13'S, 151°38'E; depth 125 m; bot- tom: grey ooze (Hoek 1883). Genus Megalasma Hoek, 1883 Megalasma Hoek, 1883: 50. — Pilsbry 1907a: 87. Diagnosis. Capitulum with five strong, calcified plates; scutal umbones subcentral; apex of carina sometimes slightly overlapping basal end of terga, progressively be- coming broader towards base; peduncle very short. Type species. Megalasma striatum Hoek, 1883: 51, pl. I, figs 5-9, pl. VII, figs 8, 9. Figure 7. Glyptelasma gracile (Hoek, 1883) (MNHN-IU-2019-4875). a. left lateral view, capitulum and peduncle; b. right lateral view, capitulum and peduncle; ¢. anterior view, orifice; d. posterior view, carina. Scale bar: 2 mm (a—d). zse.pensoft.net 658 Pitriana, P et al.: Deep-sea barnacles from the Karubar expedition (1991) Megalasma striatum Hoek, 1883 Figure 8 Megalasma striatum Hoek, 1883: 51, pl. II, figs 5-9, pl. VU, figs 8, 9. — Broch 1931: 270, figs 29, 30. Megalasma (Megalasma) elegans. — Zullo & Newman, 1964: 355, fig. 2a. Megalasma (Megalasma) striatum. — Zevina 1982: 80, fig. 71. — Chan 2009: 66, figs 2A, 16 A-H. Material examined. — Kei Islands: 1 specimen, MNHN- TU-2019-4868, Stn. CP 09, 05°23'S, 132°29'E, 368- 389 m depth, 23 October 1991; 1 specimen, MNHN- IU-2019-4869, Stn. CP 16, 05°17'S, 132°50'E, 315-349 m depth, 24 October 1991, attached to sea urchin spine; 2 specimens, MNHN-IU-2019-4866, Stn. CP 17, 05°15'S, 133°O1'E, 439-459 m depth, 24 October 1991. — Tanimbar Island: 1 specimen, MNHN-IU-2019-4867, Stn. CP 46, 08°01'S, 132°51'E, 271-273 m depth, 29 Oc- tober 1991; 4 specimens (3 attached to sea urchin spine), MNHN-IU-209-4870, Stn. DW 49, 08°00'S, 132°59'E, 206—210 m depth, 29 October 1991; 6 specimens, MNHN- TU-2019-4871, Stn. CP 85, 09°22'S, 131°14'E, 240-245 m depth, 4 November 1991, attached to spines of sea urchin. Diagnosis. Capitulum ovate, occludent margin almost same length as carinal margin; capitular plates heavily calcified, markedly sculptured with radial striations be- tween grooves of growth lines; peduncle short, covered by capitulum. Description. Capitulum white, with five calcified plates, surfaces strongly striated. Scutum with basal margin ro- tated; tergum triangular in lateral view; carina with dorsal roof widening apically on either side of midline groove. Cirrus I with anterior ramus wider than posterior ramus. Maxillule with three strong setae at upper angle separat- ed by wide notch; mandible with four teeth, lower angle sharp. Measurements of specimen: basal diameter of ca- pitulum 2.14 mm; capitular height 9.36 mm; total height 9.36 mm; scutal width 3.81 mm; scutal length 7.28 mm; tergal width 1.58 mm; tergal length 4.29 mm. Distribution. West-southwest Pacific, Indo-west Pacific, East coast of Africa, Indian Ocean, north Australia, Indo- nesia, Malay Archipelago, East China Sea, South China Sea, Taiwan, Philippines, south Japan to New Zealand; attached to echinoid spines, antipatharians, gorgonians, glassy spicule of hexactinellid sponges, corallines; 125— 984 m depth (Jones and Hosie 2016). In this study, Meg- alasma striatum was found at Kei Islands and Tanimbar Island, Indonesia. Type locality. Philippine Archipelago (Hoek 1883). Order SCALPELLIFORMES Buckeridge & Newman, 2006 Scalpelloidea Pilsbry, 1916: 14 (nom. trans. Zevina, 1978). Scalpelliformes Buckeridge & Newman, 2006: 22. Family SCALPELLIDAE Pilsbry, 1907 Pollicipedidae Gray, 1825: 100 (part.). — Annandale 1909: 63 (part.). Lepadidae Darwin, 1852: 8 (part.). Polyaspidae Gruvel, 1905: 8, 16 (part.; rejected by Pilsbry, 1907a, be- cause family group name not derived from generic name). Scalpellinae Pilsbry, 1907a: 3 (part.). Scalpellidae. — Kriiger, 1911: 7. — Pilsbry 1916: 4. — Nilsson-Cantell 1921: 162. — Foster 1978: 38. — Zevina 1978a: 999. — Buckeridge 1983: 27. — Gale 2016: 296. Figure 8. Megalasma striatum Hoek, 1883 (MNHN-IU-2019-4868). a. left lateral view, capitulum; b. right lateral, capitulum; c. anterior view, capitulum and peduncle; d. posterior view, carina. Scale bar: 2 mm (a-—d). zse.pensoft.net Zoosyst. Evol. 96 (2) 2020, 649-698 Diagnosis. Capitulum with fully or partially calcified plates; peducle with calcareous or phosphorus scales. According to Gale (2016), the Scalpellidae have a max- imum of 14 plates 1.e. carina, rostrum, paired scuta, ter- ga, upper latera, carinolatera, rostrolatera and inferior median latera; and rarely 13 plates, due to secondary loss of the rostrum. Subfamily SCALPELLINAE Pilsbry, 1907 Scalpellinae Pilsbry, 1907a: 4. — Zevina 1978a: 1002. — Zevina 1981: 93. — Gale 2016: 296. Pollictpedidae. — Annandale, 1909: 63 (part.). Diagnosis. Formerly, the subfamily was characterised by a subapical carinal umbo, inflexed carina and subapi- cal umbones of the upper and inframedian latus (Zevina 1978a). Gale (2016) characterised the subfamily by the broad, low, straplike and gently incurved rostrolatus. The rostrum is broader than high, rectangular, trapezoidal or triangular and its large, triangular, lateral surfaces contact the interior of the rostrolatus. The articulation surface be- tween the rostrum and rostrolatus extends over the entire height of both plates. Type genus. Scalpellum Leach, 1817: 68. Genus Diotascalpellum Gale, 2016 Diotascalpellum Gale, 2016: 297. Diagnosis. Scalpellines without specialised contact be- tween carina and carinolatus, merely a cresentic concavi- ty on interior of plate that forms a rim slightly overlapping the upper latus; umbo of carinolatus upright, only slightly incurved, dorsal margin gently convex; inframedian sur- face of carinolatus well demarcated, slightly inset. Diotascalpellum rubrum (Hoek, 1883) Figure 9 Scalpellum rubrum Hoek, 1883: 91, pl. IV fig. 18. — Pilsbry 1911: 62, text-fig. 1, pl. VUI, figs 14. — Calman 1918b: 122-123. — Broch 1922: 237. — Nilsson-Cantell 1927: 745-747, text-fig. 2.— 1931: 2. Trianguloscalpellum rubrum. — Zevina 1981: 316-317, fig. 240. — Rosell 1986: 89, text-fig. la. — 1991: 22, fig. 1f, g — Huang 1994: 517. — 2001: 318. —Jones et al. 2001: 254. — Young 2001: 464, fig. 7. — Jones 2007: 292. — Shalaeva and Boxshall 2014: 49, fig. 34. Diotascalpellum rubrum. — Gale 2016: 297, figs 2A, 6U-Y, 11Q—T, 13K—-M, 18A-G. Material examined. — Kei Islands: 2 specimens, MNHN-IU-2019-4925, Stn. CP 09, 05°23'S, 132°29'E, 368-389 m depth, 23 October 1991; 1 specimen, MNHN- TU-2019-4926, Stn. CP 16, 05°17'S, 132°50'E, 330- 350 m depth, 24 October 1991, attached to gorgonian. 659 Diagnosis. Capitulum with 14 plates, smooth, white and reddish coloured; carina simply, strongly bowed, roof slightly convex with umbo at apex; rostrum triangular, distinct; upper latus quadrangular, large; infra-median la- tus triangular. Description. Capitulum flat, rather broad, not covered by distinct membrane. Scutum with occludent margin arched, forming with tergal margin a triangular portion projecting over tergum. Tergum surpassing scutal area with occludent margin almost straight. Upper latus quad- rangular, angle at apex between scutal and tergal margins distinctly projecting over scutum. Rostrum small, trian- gular; rostrolatus very low, quadrangular; infra-median latus small, triangular, umbo at apex; carinal latus larger than other latera with carinal margin arched. Cirrus I with anterior and posterior rami almost same length; cirrus VI with long caudal appendages. Maxillule not notched, with large spine on upper side, cutting edge almost straight; mandible with three large teeth excluding inferior an- gle. Measurements of two specimens: height of capitu- lum 12.51—20.57 mm, width 7.39-10.55 mm, thickness 4.09-6.51 mm; length of peduncle 5.10—7.76 mm, width 4.36-6.62 mm. Distribution. Java Sea, Indonesia. Pacific: western cen- tral and northwest; Philippines; Kagoshima Sea and Sag- ami Bay, Japan; 133-551 m depth (Shalaeva and Box- shall 2014). In this study, Diotascalpellum rubrum was found at Kei Islands, Indonesia. Type locality. Philippines, 12°43'N, 122°10'E; depth, 180 m and 207 m; bottom, mud (near Luzon) (Hoek 1883). Genus Regioscalpellum Gale, 2016 Regioscalpellum Gale, 2016: 298. Diagnosis. Concave scalpellines; internal surface upper latus covered by epidermis, short carinal margin present on carinolatus; carinolatus with incurved umbones, later- al margin strongly convex. Type species. Scalpellum regium Thomson, 1873; type locality: Stn 61: Atlantic, Western Central (North Western Atlantic Basin); 5,210 m depth. Regioscalpellum moluccanum (Hoek, 1883) Figure 10 Scalpellum moluccanum Hoek, 1883: 104, pl. V figs 3, 4; 1907: 56, 85, pl. VI, fig. 13. — Gruvel 1905: 76, fig. 85. — Nilsson-Cantell 1927: 747, fig. 3. Arcoscalpellum moluccanum. — Newman & Ross, 1971: 60. — Jones et al. 1990: 6, 27. — Jones 1992: 172-175, figs 18, 19. Trianguloscalpellum moluccanum. — Jones, 2012: 371, table 2. — Sha- laeva and Boxshall 2014: 45, fig. 31. zse.pensoft.net 660 Pitriana, PR et al.: Deep-sea barnacles from the Karubar expedition (1991) Figure 9. Diotascalpellum rubrum (Hoek, 1883) (MNHN-IU-2019-4925). a. left lateral view showing the capitulum and peduncle; b. right lateral view showing the capitulum and peduncle; c. rostral view showing the capitulum and peduncle; d. carinal view. Scale bar: 5 mm (a-—d). Regioscalpellum moluccanum. — Gale, 2016: 298. Material examined. — Tanimbar Island: 3 specimens, MNHN-IU-2019-4909, Stn. CP 38, 07°40'S, 132°27'E, 620-666 m depth, 28 October 1991; 1 specimen, MNHN- TU-2019-4910, Stn. DW 49, 08°00'S, 132°59'E, 206-210 m depth, 29 October 1991; 1 specimen, MNHN-IU-2019-4911, Stn. CC 57, 08°19'S, 131°53'E, 603-620 m depth, 31 Octo- ber 1991; 1 specimen, MNHN-IU-2019-4912, Stn. CP 52, 08°03'S, 131°48'E, 1244-1266 m depth, 30 October 1991. Diagnosis. Capitulum with 14 plates, surface covered by chitinous, almost smooth membrane; carina simply, not strongly bowed, apex projecting freely, roof not flat; umbo of the carina apical; upper latus almost triangular. Description. Capitulum inflated, subtriangular, apices of plates projecting freely through membrane, growth lines fairly distinct. Scutum trapezioid, umbo apical. Tergum large, oval, truncated, very pointed, extending between upper latus and carina. Carina simply, slightly bowed, roof slightly convex, apex projecting freely. Upper latus quadrangular; rostrum very small, carinate; rostral latus trapeziform; infra-median latus triangular, moderately small; carinal margins of carinal latera convex, touching each other at base and below middle of carina. Peduncle stout, slightly thinner below apex. Cirrus I unequal, an- terior ramus oval, posterior ramus more slender; cirrus VI with long caudal appendages. Maxillule not notched, large spine on upper side, cutting edge almost straight; zse.pensoft.net mandible with three large teeth excluding inferior an- gle. Measurements of five specimens: height of capitu- lum 7.88—14.89 mm, width 4.81—-10.65 mm, thickness 2.15—5.24 mm; length of peduncle 3.38—9.13 mm, width 2.09-6.04 mm. Distribution. Pacific, Central and Southwest; Tasman Sea, north-eastern Australia; 788—2,745 m depth (Jones 2012; Shalaeva and Boxshall 2014). In this study, Re- gioscalpellum moluccanum was found at Tanimbar Is- land, Indonesia. Type locality. Banda Sea (4°21'S, 129°7'E); Challenger stn 195; 2,606 m depth: bottom temp. 3 °C; substrate grey ooze (Hoek 1883). Genus Scalpellum Leach, 1817 Scalpellum Leach, 1817: 68. — Darwin 1852: 21. — Hoek 1883: 59. — Gruvel 1905: 23. — Pilsbry 1907d: 181. — Tarasov and Zevina 1957: 126. — Zevina 1978a: 1002. — 1981: 94. — Gale 2016: 297. Strictoscalpellum Broch, 1924: 14. Diagnosis. Scalpellines with carinal and scutal umbones subapical; carinolatus with horn-like projection, often recurved, extending beyond carinal margin; inframedian latus rectangular to slightly trapezoidal, with low umbo; upper latus rhomboidal with subapical umbo; rostrum rectangular, pyramidal, with large sub-umbonal surface. Zoosyst. Evol. 96 (2) 2020, 649-698 661 Figure 10. Regioscalpellum moluccanum (Hoek, 1883) (MNHN-IU-2019-4909). a. left lateral view showing the capitulum and peduncle; b. right lateral view showing the capitulum and peduncle; ec. carinal view showing the capitulum and peduncle; d. rostral view showing the capitulum and peduncle. Scale bar: 4 mm (a—d). Type species. Lepas scalpellum Linnaeus, 1767: 1109. Type locality. Unknown. Scalpellum sp. Figure 11 Material examined. — Tanimbar Island: 1 specimen, MNHN-UI-2019-4929, Sth. DW 49, 08°00'S, 132°59'E, 206-210 m depth, 29 October 1991. Diagnosis. Capitulum with 14 fully calcified plates; ter- gum triangular; inframedian latus broad, quadrilateral with umbo near to basal; upper latus rhomboidal, umbo sub-apical; carinal lateral horn-shaped, with angle ex- tending beyond carina. Description. Capitulum subtriangular, covered by mem- brane. Tergum triangular, occludent and basal margins slightly convex; scutum quadrangular, basal and upper latus margins concave; upper latus pentagonal; rostrolatus quadrangular; inframedian latus broad, rectangular, umbo close to basal margin; carinolatus horn-shaped with slight- ly deep transverse furrows, umbo at basi-carinal angle; carina convex, umbo apical. Cirrus I with rami unequal, anterior ramus broader than posterior ramus; both rami densely covered with long setae. Maxilla subtriangular, with dense long setae; maxillule notched, with numerous setae; mandible with three teeth, inferior angle with large cuspidate setae. Measurements of specimen: height of ca- pitulum 15.90 mm, width 10.29 mm, thickness 4.58 mm; length of peduncle 6.97 mm, width 5.67 mm. Distribution. In this study, Scalpellum sp. was found in Tanimbar Island, Indonesia. Scalpellum stearnsi Pilsbry, 1890 Figure 12 Scalpellum magnum Darwin, 1852: 18, pl. I, fig. 1. Scalpellum calcariferum Fischer, 1891: 116. Scalpellum stearnsii var. gemina and var. robusta Hoek, 1907: 69, pl. VI, figs 2, 3, 8-12. Scalpellum stearnsii var. inerme Annandale, 1916a: 293. Scalpellum stearnsii forma typica Broch, 1931: 16. Scalpellum stearnsii Pilsbry, 1890a: 96. — 1890b: 441, pl. IV, figs 1-5. — 1907a: 14, pl. 4, figs 1-5. — Nilsson-Cantell 1921: 175. — 1934: 33. — Broch 1922: 235, fig. 6. — Hiro 1933: 22, fig. 4, pl. 1, figs 5, 5a. — 1939a: 237. — Zevina 1981: 98, fig. 68. — Rosell 1991: 15. — Jones 1992: 146, figs 1, 2. — Liu and Ren 2007: 226, fig. 95.— Chan et al. 2009b: 89, figs 73-76. — Chan et al. 2010: 24, figs 2C, 18A-G. Scalpellum stearnsi: Gruvel, 1905: 44, fig. 46. — Gale 2016: fig. 5, table 1. Material examined. — Tanimbar Island: 1 specimen, MNHN-IU-2019-4888, Stn. CP 71, 08°38'S, 131°44'E, 477-480 m depth, 2 November 1991; 1 specimen, MNHN-IU-2019-4889, Stn. CC 58, 08°19'S, 132°02'E, 457-461 m depth, 31 October 1991; 6 specimens, MNHN-IU-2019-4890, Stn. CC 42, 07°53'S, 132°42'E, 350-354 m depth, 28 October 1991; 3 specimens, zse.pensoft.net 662 Pitriana, PR et al.: Deep-sea barnacles from the Karubar expedition (1991) Figure 11. Scalpellum sp. (MNHN-UI-2019-4929). a. left lateral view, capitulum and peduncle; b. right lateral view, capitulum and peduncle; ¢c. rostral view, capitulum and peduncle; d. carinal view. Scale bar: 7 mm (a—d). MNHN-IU-2019-4891, Stn. CP 69, 08°42'S, 131°53'E, 356-368 m depth, 2 November 1991; 5 specimens, MNHN-IU-2019-4892, Stn. CC 41, 07°45'S, 132°42'E, 393-401 m depth, 28 October 1991; 5 specimens, MNHN-IU-2019-4893, Stn. CP 59, 08°20'S, 132°11'E, 399-405 m depth, 31 October 1991, 2 specimens attached to glass rope sponge. — Kei Islands: 8 specimens, MNHN-IU-2019-4894, Stn. CP 06, 05°49'S, 132°21'E, 287-298 m depth, 22 Oc- tober 1991. Diagnosis. Capitulum rhomboidal, compressed; carina strongly angled at umbo; scutum large, umbo at tergo-oc- cludent angle; tergum triangular; inframedian latus pen- tagonal; carinolateral horn-shaped. Description. Capitulum with 14 fully calcified plates. Scutum large, longer than broad, apex slightly projecting over tergum; tergum large, triangular, apex erect, slightly recurved. Carina distinctly bent near middle, umbo dis- tant from apex. Cirrus I with anterior ramus oval (12-seg- ments), posterior ramus slender, long (15-segments); cir- ri II-VI slender, long, rami almost equal length: cirrus VI with short caudal appendages (six segments). Max- illule not notched, with numerous setae; mandible with six teeth excluding inferior angle. Measurements of five specimens: height of capitulum 31.69-52.00 mm, width 28.20—40.11 mm, thickness 11.19-18.56 mm; peduncle length 28.29-86.59 mm, width 11.39-17.94 mm. Distribution. Indo-west Pacific: Indian Ocean, off Nico- bar Island; Indonesia: Java Sea, Bali Straits; north west & zse.pensoft.net north east Australia; Malay Archipelago; Celebes; Sulu Arch.; Sulu Sea; South China Sea; East China Sea; Phil- ippines; Taiwan; east coast of south Japan, Sagami Bay, off Hondo; attached to mollusc shells, anchor filaments of hexactinellid sponges, telegraph cables, carapaces of crabs, stones; 146-2117 m depth (Jones and Hosie 2016). In this study, Scalpellum stearnsi was found at Tanimbar Island and Kei Islands, Indonesia. Type locality. East coast of Japan, between the Bay of Tokyo and the Inland Sea (Jones 1992). Remarks. For the first time, Scalpellum stearnsi was found in Japan and described by Pilsbry (1890). During the Siboga expedition (1899), S. stearnsi was collected from different locations in the Malay Archipelago with the depths varying from 204 m to 450 m. Hoek (1907) found intraspecific variations of the shell plate mor- phology. He then divided S. stearnsi into two groups, i.e. variety robusta and var. gemina, which differed in the shape of the tergum. The species S. stearnsi in this study belongs to the group of var. gemina because of the V-shaped tergum. Scalpellum stearnsi has a low period of larval develop- ment (Ozaki et al. 2008) and a slow growth rate (Yusa et al. 2018). This can result in the broad geographical distribution of this species. Recently, Lin et al. (2020) examined the diversity and genetic differentiation of populations of S. stearnsi from the East China Sea, West Philippine Basin, Sulu Sea and Caroline Trenches, which resulted in four distinct clades of S. stearnsi. Zoosyst. Evol. 96 (2) 2020, 649-698 663 Figure 12. Scalpellum stearnsi Pilsbry, 1890 (MNHN-IU-2019-4888). a. left lateral view, capitulum and peduncle; b. right lateral view, capitulum and peduncle; ec. rostral view, capitulum and peduncle; d. carinal view. Scale bar: 14 mm (a-—d). Subfamily MEROSCALPELLINAE Zevina, 1978 Meroscalpellinae Zevina, 1978b: 1343. Diagnosis. Capitulum with 14 or 13 plates, reduced in differing stages or proportions; carina with two umbo positions; females considered rarer than hermaphrodites; males sac-like, usually without plates, rarely with two or four reduced plates. Type genus. Meroscalpellum Zevina, 1978b: 1346. Genus Alcockianum Zevina, 1978 Alcockianum Zevina, 1978b: 1345. Diagnosis. Plates strongly reduced; partly obscured un- der heavy cuticle; tergum double branched, depressed at base; scutum triangular; upper lateral triangular; mid-lat- eral very small, umbo apical; caudal appendages very long; hermaphrodites. Type species. Scalpellum alcockianum Annandale, 1905: 82; type locality: Gulf of Manaar and Andaman Sea. Alcockianum persona (Annandale, 1916) Figure 13 Scalpellum persona Annandale, 1916a: 295, pl. IV, fig. 3; pl. V, figs 7, 8; pl. VI, figs 3-5. — Calman 1918b: 120. — Nilsson-Cantell 1934: 44, pl. 5, fig. 2. Graviscalpellum persona Foster, 1980: 527, figs le, 2. Alcockianum persona Zevina, 1978b: 1345. — 1981: 150-151, fig. 107. — Jones 1992: 151-154, figs 4, 5. Material examined. — Tanimbar Island: 2 specimens, MNHN-IU-2019-4903, Stn. CP 71, 08°38'S, 131°44'E, 477-480 m depth, 2 November 1991; 5 specimens, MNHN-IU-2019-4904, Stn. CC 21, 05°14'S, 133°00'E, 688-694 m depth, 25 October 1991 attached to glass rope sponge; 1 specimen, MNHN-IU-2019-4905, Stn. CC 40, 07°46'S, 132°31'E, 443-468 m depth, 28 October 1991; 5 specimens, MNHN-IU-2019-4906, Stn. CC 57, 08°19'S, 131°53'E, 603-620 m depth, 31 October 1991. Diagnosis. Capitulum with plates buried or obscure; plates reduced in size; tegum reduced, forming four-point- ed star; scutum small; peduncle armed with distinct cir- cles of large, calcareous scales. Description. Capitulum brownish, large, ovoid, inflated, with 13 capitular plates, including a vestigial rostrum, plates embedded and mostly concealed by thick, opaque membrane. Scutum small, widely separated from all re- maining plates except tergum, margins not excavated or deeply concave; tergum reduced in form as four-pointed star, with two rays greatly and two rays slightly produced. Carina reduced in size, apex approaching terga, widely separated from remaining plates. Peduncle cylindrical, similar length to capitulum, with large calcareous scales arranged in alternating rows. Cirrus I with anterior ramus oval (8-segments), posterior ramus slender, long (12-seg- ments); cirri I-VI slender, long, rami almost equal length; cirrus VI with caudal appendages; caudal appendages 1/3 length of cirrus VI, 15-segmented, tapering distally. Penis rather short, smooth, pointed. Maxilla bilobed, dense setae on margin. Maxillule relatively large, with broad, shallow excavation on lower margin occupying more than half mar- gin, remainder of margin obliquely subtruncate; mandible with three main teeth in addition to inner angle, which is variously divided, broad as a whole. Measurements of five zse.pensoft.net 664 Pitriana, P. et al.: Deep-sea barnacles from the Karubar expedition (1991) Figure 13. A/cockianum persona (Annandale, 1916) (MNHN-IU-2019-4903). a. left lateral view, capitulum and peduncle; b. right lateral view, capitulum and peduncle; ¢. rostral view, capitulum and peduncle; d. carinal view. Scale bar: 9 mm (a-—d). specimens: height of capitulum 17.71—35.06 mm, width 12.91—28.06 mm, thickness 4.96—18.20 mm; length of pe- duncle 13.42—28.67 mm, width 6.66—10.79 mm. Distribution. Indonesian Seas, eastern Australia, New Zea- land; 109-915 m depth (Jones 1992). In this study, Alcock- ianum persona was found at Tanimbar Island, Indonesia. Type stocality. Timor Sea (10°22'30"S, 120°7'30"E); 109-366 m depth: bottom temp. 13.1 °C (Jones 1992). Genus Annandaleum Newman & Ross, 1971 Annandaleum Newman & Ross, 1971: 122. — Zevina 1978b: 1346. Diagnosis.Capitulum of female with 14 partially calcified plates; tergum in shape of inverted V; scutum with moder- ately long apicolateral arm, 1/4 to 1/2 length of tergal mar- gin; basal margin of scutum entire; upper latus pentagonal to triangular or subrectangular, with or without short de- pending arm; carinolatus as high as or slightly higher than wide, umbo at basicarinal angle; inframedian latus higher than wide, vase-shaped, umbo submedial to basal; rostro- latus wider than high, commonly less than twice height of inframedian latus; caudal appendage relatively long. Type species. Scalpellum subflavum Annandale, 1906. Annandaleum japonicum (Hoek, 1883) Figure 14 Scalpellum japonicum Hoek, 1883: 67, pl. Ill, figs 9, 10. — Annandale 1906: 131, fig. 3. — Pilsbry 1911: 66, pl. II, figs 1-3. — Weltner 1922: zse.pensoft.net 69, taf. II, fig. 4. — Hiro 1937 b: 392. — Tarasov and Zevina 1957: 144, figs 46, 47. — Zevina 1969: 68. — 1970: 257-259, figs 5, 6. Scalpellum chitinosum Hoek, 1907: 73. Scalpellum curiosum Hoek, 1907: 49. Scalpellum japonicum biramosum Pilsbry, 1911: 68, fig. 4. — Weltner 1922: 69, taf. II, fig. 5. — Nilsson-Cantell 1938: 7. Scalpellum japonicum metapleurum Pilsbry, 1907b: 360. Annandaleum japonicum biramosum Chan et al., 2009b: 98-99, fig. 80. — Chan et al. 2010: 17-18, figs 2A, 14, 15. Annandaleum japonicum. — Newman & Ross, 1971: 122. — Zevina 1981: 166-167. — Jones et al. 2001: 251. — Young 2001: 465, fig. 8. — 2007: 23, fig. 22. Material examined. — Tanimbar Island: 4 specimens, MNHN-IU-2019-4932, Stn. CP 53, 08°18'S, 131°41'E, 1026-1053 m depth, 30 October 1991. Diagnosis. Capitulum with 14 partly calcified plates; ca- rina with flat roof; apex of tergum curved towards carina; rostral latus trapezoidal in shape; peduncle short. Description. Capitulum yellowish, with 14 plates sepa- rated by broad, chitinous spaces. Peduncle short, curved, scales large, not numerous. Scutum long, narrow; umbo apical, divided into two segments, occludent segment large, increasing in width from upper to lower part, oth- er segment very narrow. Tergum divided into occludent and carinal segments, with triangular, chitinous portion between, umbo apical, distinctly recurved. Cirrus I un- equal, anterior ramus oval, posterior ramus slender long; cirrus VI with caudal appendages. Maxillule slightly notched, two long and one short setae on upper side; mandible with three teeth excluding inferior angle; la- brum cutting edge straight, very fine teeth on cutting edge. Measurements of the four specimens: height of ca- Zoosyst. Evol. 96 (2) 2020, 649-698 665 Figure 14. Annandaleum japonicum (Hoek, 1883) (MNHN-UI-2019-4932). a. left lateral view showing the capitulum and pedun- cle; b. right lateral view showing the capitulum and peduncle; ¢c. rostral view showing the capitulum and peduncle; d. carinal view. Scale bar: 2 mm (a-—d). pitulum 9.56—12.73 mm, width 4.67—7.13 mm, thickness 1.85—3.21 mm; length of peduncle 3.37-4.77 mm, width 1.83—3.27 mm. Distribution. Eastern Indian Ocean; Northwest and Western Central Pacific; Malay Archipelago; Japan; Tai- wan; Indonesia; attached to shell of gastropod, gorgoni- ans, rocks; 805—6,810 m depth (Jones and Hosie 2016). In this study, Annandaleum japonicum was found at Ta- nimbar Island, Indonesia. Type locality. Pacific, Northwest (off Nagoya, south of Japan); 1,017 m depth (Shalaeva and Boxshall 2014). Annandaleum laccadivicum (Annandale, 1906) Figure 15 Scalpellum laccadivicum Annandale, 1906b: 393. — 1908: pl. I, figs 3, 4. — 1913: 235; 1916b: 129, pl. vu, fig. 6. — Calman 1918b: 124. — Hiro 1933: 31, text figs 7, 8, pl. I, figs 11-11b. — 1937b: 392. — Stubbings 1936: 26. — Nilsson-Cantell 1938: 25. Scalpellum subflavum Annandale, 1906b: 397. — Newman and Ross LF Lde 2D: Scalpellum polymorphum Hoek, 1907: 80, pl. 7, figs 9-11. — Weltner 1922: 72. Scalpellum molliculum Pilsbry, 1911: 68, pl. 10, figs 4, 5. Annandaleum laccadivicum Rosell, 1991: 16, fig 1a. — Zevina 1981: 170, fig. 121. —Jones et al. 2001: 251. — Chan et al. 2009b: 102-103. Material examined. — Tanimbar Island: 1 specimen, MNHN-IU-2019-4931, Stn. CP 54, 08°21'S, 131°43'E, 836-869 m depth, 30 October 1991, attached to gas- tropod shell. Diagnosis. Capitulum regularly oval; carina simply bowed; tergum almost triangular; scutum subtriangular; upper latus large, perfectly calcified part irregularly tri- angular, constricted above; rostral latus quadrangular; peduncle at least half as long as capitulum. Description. Capitulum compressed; 13 plates complete- ly covered by fine, hairless membrane. Peduncle half length of capitulum, stout, cylindrical, armed with small, transversely elongated plates. Scutum subtriangular, lat- eral margin excavated with tooth above excavation blunt, short, simple; tergum almost triangular, scutal margin ex- cavated, but not very boldly, occludent margin slightly, regularly convex outwards. Carina simply bowed, umbo subterminal, in contact with terga above or just entering between them. Cirrus I unequal, anterior ramus oval, posterior ramus slender, long; cirri IVI slender, long, rami almost equal lengths; cirrus VI with long, slender caudal appendages. Maxillule slightly notched, two ma- jor setae on upper side; mandible with four teeth. Mea- surements of specimen: height of capitulum 18.88 mm, width 11.41 mm, thickness 6.30 mm; length of peduncle 11.45 mm, width 4.89 mm. zse.pensoft.net 666 Pitriana, PR et al.: Deep-sea barnacles from the Karubar expedition (1991) Figure 15. Annandaleum laccadivicum (Annandale, 1906) (MNHN-IU-2019-4931). a. left lateral view showing the capitulum and peduncle; b. right lateral view showing the capitulum and peduncle; c. rostral view showing the capitulum and peduncle; d. carinal view. Scale bar: 5 mm (a—d). Distribution. Gulf of Oman, Arabian Sea, Sri Lanka, Japan (Chan et al. 2009b). In this study, Annandaleum laccadivicum was found at Tanimbar Island, Indonesia. Type locality. Laccadive Sea, 2,077 m depth; on living shells of Dentalium (Annandale 1906). Annandaleum lambda (Annandale, 1910) Figure 16 Scalpellum lambda Annandale, 1910b: 115. — 1916a: pl. VIL figs 6, 6a, pl. VIUL, figs 12-15. Scalpellum longius Annandale, 1913: 234. Annandaleum lambda. — Newman & Ross, 1971: 122. — Jones, 1992: 154, fig. 6. Material examined. — Tanimbar Island: 10 specimens, MNHN-IU-2019-4915, Stn. CP 52, 08°03'S, 131°48'E, 1244-1266 m depth, 30 October 1991; 1 specimen, MNHN-IU-2019-4913, Stn. CP 53, 08°18'S, 131°41'E, 1026-1053 m depth, 30 October 1991; 1 specimen, MNHN- TU-2019-14, Stn. CP 89, 08°39'S, 131°08'E, 1058-1084 m depth, 5 November 1991, attached to pumice. Diagnosis. Capitulum narrow; carinal margin strongly curved; laterally compressed; bearing fourteen imperfect- zse.pensoft.net ly formed, thin, smooth, translucent plates, eight formed like a Greek lambda. Description. Capitulum with 14 plates. Peduncle shorter than capitulum, cylindrical, rather slender, ex- panded at base. Tergum lambda-shaped, with promi- nent tooth on carinal margin just above point where apex of carina approaches it. Scutum shaped similarly, but occludent branch much stouter, with vertical ridge running nearer to lateral than occludent margin. Cirrus I unequal, anterior ramus oval, posterior ramus slender, long; cirri II-VI slender, long, with rami almost equal length; cirrus VI with long, slender caudal appendag- es. Maxillule not notched, two major setae on upper side; mandible with four teeth including inner angle. Measurements of five specimens: height of capitulum 16.61—24.27 mm, width 8.73—-14.49 mm, thickness 3.17-6.31 mm; length of peduncle 5.54—-15.27 mm, width 3.73-6.26 mm. Distribution. Indo-Pacific from off Zanzibar, Indian Ocean, eastern Australia; Sumbawa, Indonesia; Malay Arch.; SW of Calatagan Pt, Philippines; S Japan (S of Honda I.), SW Pacific; attached to shells of bivalves; 234-2077 m depth (Jones and Hosie 2016). In this study, Annandaleum lambda was found at Tanimbar Is- land, Indonesia. Zoosyst. Evol. 96 (2) 2020, 649-698 667 Figure 16. Annandaleum lambda (Annandale, 1910) (MNHN-IU-2019-4915). a. left lateral view showing the capitulum and pe- duncle; b. right lateral view showing the capitulum and peduncle; ¢. rostral view showing the capitulum and peduncle; d. carinal view. Scale bar: 6 mm (a-—d). Type locality. Eastern Indian Ocean (13°54'15"N, 94°O2'1S5"E), Investigator Stn. 372; 1,176 m depth (Jones 1992). Genus Litoscalpellum Newman & Ross, 1971 Litoscalpellum Newman & Ross, 1971: 108. — Zevina 1978b: 1344. — Liu and Ren 1985: 196. — 2007: 228. Diagnosis. Capitulum with 14 plates, mostly not reduced; tergum with straight basal margin or shallow notch; scu- tum with straight, smooth basal margin; upper latus trian- gular or elongate, commonly with slightly hollowed-out basal margin, rarely with deep notch; inframedian latus narrow, triangular or quadrilateral, umbo apical; caudal appendages present. Type species. Litoscalpellum fissicarinatum Newman & Ross, 1971: 108, pl. XC, text-fig. 55. Litoscalpellum juddi (Calman, 1918) Figure 17 Scalpellum juddi Calman, 1918b: 116, text-figs 5—7. Scalpellum regulus Calman, 1918b: 113, text-fig. 4. Litoscalpellum juddi Zevina, 1981:135—136, fig. 94. Material examined. — Tanimbar Island: 5 specimens, MNHN-IU-2019-4896, stn. CC 56, 08°16'S, 131°59'E, 549-552 m depth, 31 October 1991; 1 specimen, MNHN- IU-2019-4898, stn. CP 71, 08°38'S, 131°44'E, 477-— 480 m depth, 2 November 1991; 6 specimens, MNHN- IU-2019-4901, stn. CP 72, 08°36'S, 131°33'E, 676-699 m depth, 2 November 1991. Diagnosis. Capitulum compressed, oval, notched above; surface covered with very short velvety pubescence, ar- eas of valves defined by grooves, lines of growth strongly marked; carina flattened. Description. Capitulum with 14 plates, cartilaginous thick. Terqum irregularly triangular, apex projecting above cari- na, recurved. Scutum triangular, basal width more than half height, apex overlapping tergum. Carina evenly curved, apex projecting freely for short distance, touching or enter- ing between terga, its base rounded, widely separated from carinal latera; roof strongly convex. Upper latus triangular with base convex. Rostrum triangular; inframedian latus small, triangular, base deeply embedded. Peduncle with large transverse plates, not overlapping (widely spaced). Cirrus I unequal, anterior ramus oval, posterior ramus slen- der, long; cirrus VI with caudal appendages. Maxillule with oral edge sinuous, a large seta at its proximal third, another at its distal end; mandible with four teeth. Measurements of five specimens: height of capitulum 16.94-39.25 mm, width 11.39-28.28 mm, thickness 5.20-19.89 mm; length of peduncle 15.02—52.02 mm, width 5.63—16.87 mm. Distribution. In this study, Litoscalpellum juddi was found at Tanimbar Island, Indonesia. zse.pensoft.net 668 Pitriana, P. et al.: Deep-sea barnacles from the Karubar expedition (1991) Figure 17. Litoscalpellum juddi (Calman, 1918) (MNHN-UI-2019-4898). a. left lateral view showing the capitulum and peduncle; b. right lateral view showing the capitulum and peduncle; ¢. rostral view showing the capitulum and peduncle; d. carinal view. Scale bar: 11 mm (a-d). Type locality. 11°O'S, 121°30'E (Java-Australia), 720 m depth (Calman 1918b). Litoscalpellum recurvirostrum (Hoek, 1883) Figure 18 Scalpellum recurvirostrum Hoek, 1883: 77-79, pl. Ill, figs 11, 12, pl. VII, figs 9, 10. Arcoscalpellum recurvirostrum. — Newman & Ross, 1971: 79-80, fig. 39. Litoscalpellum recurvirostrum. — Zevina, 1974: 214. — 1981: 127, 128, fig. 88. — Shalaeva and Boxshall 2014: 13, fig. 6. Material examined. — Kei Islands: 1 specimen, MNHN- IU-2019-4934, Stn. CP 05, 05°49'S, 132°18'E, 296- 299 m depth, 22 October 1991; 1 specimen, MNHN- IU-2019-4936, Stn. CP 09, 05°23'S, 132°29'E, 368-389 m depth, 23 October 1991. — Tanimbar Island: 4 specimens, MNHN-IU-2019-4935, Stn. CP 83, 09°23'S, 131°00'E, 285-297 m depth, 4 Novem- ber 1991; 16 specimens, MNHN-IU-2019-4937, Stn. CP 69, 08°42'S, 131°53'E, 356-368 m depth, 2 November 1991. Diagnosis. Capitulum plates separated, covered by fine membrane; carina slightly angularly bent; upper latus trapeziform; inframedian latus triangular; peduncle with scattered, calcareous scales. Description. Capitulum with 14 plates, lines of growth not distinct, plates separated by broad, membranous interspac- es. Scutum elongate quadrangular, broader at base than in upper half; umbo apical, slightly projecting outwards. Tergum triangular; carina bowed; upper latus trapeziform; inframedian latus very small, triangular. Peduncle cylindri- cal with numerous calcareous scales scattered over surface. zse.pensoft.net Cirrus I with rami unequal, anterior ramus oval, posterior ramus slender, long, segments very hairy. Maxilla bilobed; maxillule notched, two large setae above notch; mandible with three teeth, inferior angle pectinated. Measurements of five specimens: height of capitulum 16.15—26.05 mm, width 9.24—-16.59 mm, thickness 4.45—8.18 mm; length of peduncle 7.25—20.90 mm, width 4.13—8.95 mm. Distribution. Indian Ocean, Antarctic and Southern (South of the Kerguelen Islands); known depth range 195 to 274 m (Shalaeva and Boxshall 2014). In this study, Litoscalpellum recurvirostrum was found at Kei Islands and Tanimbar Island, Indonesia. Type locality. Between Kerguelen and Heard Islands; 52°4'S, 71°22'E; depth: 270 m; bottom temperature: 1.8 °C; bottom: rocks (Hoek 1883). Litoscalpellum walleni Newman & Ross, 1971 Figure 19 Litoscalpellum walleni Newman & Ross, 1971: 116, pl. X E text-fig. 60. Material examined. — Tanimbar Island: 1 specimen, MNHN-IU-2019-4927, Stn. CP 91, 08°44'S, 131°0S'E, 884-891 m depth, 5 November 1991. Diagnosis. Capitulum with 14 plates; inframedian latus triangular, very small, narrow; carina with flat roof; ros- trum minute; scutum lacking prominent apicolateral arm; maxilla subtriangular, slightly trilobate; maxilulle slight- ly notched with teeth diverging laterally from cutting edge; caudal appendage multi-articulate, slightly shorter than first segment of cirrus VI. Zoosyst. Evol. 96 (2) 2020, 649-698 669 Figure 18. Litoscalpellum recurvirostrum (Hoek, 1883) (MNHN-UI-2019-4934). a. left lateral view showing the capitulum and peduncle; b. right lateral view showing the capitulum and peduncle; c. rostral view showing the capitulum and peduncle; d. carinal view. Scale bar: 6 mm (a—d). Figure 19. Litoscalpellum walleni Newman & Ross, 1971 (MNHN-UI-2019-4927). a. left lateral view showing the capitulum and peduncle; b. right lateral view showing the capitulum and peduncle; c. rostral view showing the capitulum and peduncle; d. carinal view. Scale bar: 4 mm (a—d). Description. Capitulum elongate, higher than broad; plates ornamented with growth lines. Carina bowed, roof slightly convex, bounded by angles. Tergum trian- gular; scutum with lateral margin concave; upper latus roughly quadrangular; carinal latus higher than wide; rostral latus quadrangular, slightly broader than high; rostrum minute, narrow, essentially rectangular. Cirrus I unequal, anterior ramus shorter, broader than posterior ramus; both rami densely covered with long setae. Man- dible with four teeth including inferior angle, inferior zse.pensoft.net 670 angle serrate along basal margin with four spines. Mea- surements of specimen: height of capitulum 14.56 mm, width 8.09 mm, thickness 2.50 mm; length of peduncle 7.52 mm, width 3.89 mm. Distribution. Southeast Pacific Ocean (Newman and Ross 1971). In this study, Litoscalpellum walleni was found at Tanimbar Island, Indonesia. Type locality. northeast of Peter I Island, southeast Pa- cific Ocean; 65°50'S, 88°56'W; depth: 4,502 m (Newman and Ross 1971). Subfamily ARCOSCALPELLINAE Zevina, 1978 Arcoscalpellinae Zevina, 1978b: 1346. —1981: 184. — Liu and Ren 2007: 238. Diagnosis. Capitulum with plates packed very closely; ca- rina slightly convex; umbo of scutum and tergum apical. Type genus. Arcoscalpellum Hoek, 1907: 57. Genus Amigdoscalpellum Zevina, 1978 Arcoscalpellum Hoek, 1907: 85 (in part). - Newman and Ross 1971: 42 (part). Amigdoscalpellum Zevina, 1978b: 1349. — Foster 1980: 527. — Liu and Ren 1985: 206. — 2007: 251. Diagnosis. Capitulum of female or hermaphrodite with 13 or 14 completely calcified plates; scutum and tergum with umbones apical; carina evenly bent, umbo apical or sub- apical; middle latera narrowly triangular, baton-like, small, umbo apical, not reaching upper latera; carinolatera typi- cally with umbo orientated in middle part of carinal area, not extending beyond rim of capitulum; caudal appendag- es usually uni-articulate, but possibly absent or extended to seven segments; basically females with dwarf males. Type species. Scalpellum manum Zevina, 1973: 843, figs 1-7; type locality: Indian Ocean, 12°18'S, 112°43'E. Amigdoscalpellum costellatum (Withers, 1935) Figure 20 Scalpellum elongatum Hoek, 1883: 93, pl. IV, figs 8, 9. — Nilsson-Can- tell 1928: 8, fig. 4. — 1931: 3, text-fig.1. — 1938: 7. non Scalpellum (Arcoscalpellum) elongatum. — Steenstrup, 1837: 409. Scalpellum (Scalpellum) elongatum. — Stubbings, 1936: 25, text-fig. 10. Scalpellum (Arcoscalpellum) costellatum Withers, 1935: 279. Arcoscalpellum buccinum Newman & Ross, 1971: 55, pl. VIII, figs 22, 23. Arcoscalpellum costellatum. — Foster, 1978: 56, pl. 7B, C, fig. 33. Amigdoscalpellum costellatum. — Foster, 1980: 527-529, fig. 3J. — Zevi- na 1981: 270-271, fig. 204. — Jones et al. 1990: 5 — Jones 2012: 371, zse.pensoft.net Pitriana, P. et al.: Deep-sea barnacles from the Karubar expedition (1991) 376 — Poltarukha 2013: 52-53, fig. 1. -Shalaeva and Boxshall 2014: 16, fig. 9. Material examined. — Tanimbar Island: 2 specimens, MNHN-IU-2019-4922, Stn. CC 21, 05°14'S, 133°00'E, 688-694 m depth, 25 October 1991, attached to glass rope sponge. Diagnosis. Capitulum triangular; plates ornamented with radial ridges, covered by sparsely hirsute integument; ca- rina with roof deeply grooved; scutum with apex over- lapping tergum; inframedian latus triangular, very small. Desciption. Capitulum yellowish, with 13 fully calci- fied plates. Peduncles short with scales slightly over- lapping in the middle part. Scutum with pit for comple- mental males, above shallow pit for adductor muscle. Carina wide in lower part, ribbed in upper part. Upper latus with straight sides; rostrum appearing externally as inverted triangle. Cirrus I unequal, anterior ramus oval, posterior ramus slender, long; cirrus VI with very short caudal appendages. Maxillule with notch between two or three stout setae at upper angle, group of more slender setae on cutting edge; mandible with three teeth excluding inferior angle; labrum cutting edge slightly concave, numerous pointed teeth on cutting edge. Measurements of two specimens: height of ca- pitulum 12.44—13.88 mm, width 6.97—7.52 mm, thick- ness 2.77—3.47 mm; length of peduncle 2.99-3.15 mm, width 2.94—3.24 mm. Distribution. Indian Ocean, Eastern and Western; Pacif- ic, Southwest and Western Central; Atlantic, Southeast and Eastern central; 110—2,397 m depth (Shalaeva and Boxshall 2014). In this study, Amigdoscalpellum costel- latum was found at Tanimbar Island, Indonesia. Type locality. 1. Station 135: Island of Tristan da Cunha; depth: 110 m, 137 m, 183 m, 274 m, 1,006 m, 1,829 m, 2,012 m; bottom: rocky, shells. 2. Station 164a: off Syd- ney, 34°13'S, 151°38'E; depth: 750 m; bottom: grey ooze. 3. Station 169: off East Cape, Auckland, 37°34'S, 179°22'E; depth: 1,280 m; bottom temperature: 4.2 °C; bottom: grey ooze (Hoek 1883). Amigdoscalpellum tenue (Hoek, 1883) Figure 21 Scalpellum tenue Hoek, 1883: 119, pl. IV, figs 20, 21, pl. X, fig. 6. — Gruvel 1912: 345. — Nilsson-Cantell 1938: 8. non Scalpellum tenue Annandale, 1906a: 142. Amigdoscalpellum tenue. — Zevina 1981: 292-293, fig. 220 (2). — Sha- laeva and Boxshall 2014: 17, fig. 10. Material examined. — Kei Islands: 1 specimen, MNHN- TU-2019-4921, Stn. CP 20, 05°15'S, 132°59'E, 769- 809 m depth, 25 October 1991. Zoosyst. Evol. 96 (2) 2020, 649-698 671 Figure 20. Amigdoscalpellum costellatum (Withers, 1935) (MNHN-IU-2019-4922). a. left lateral view showing the capitulum and peduncle; b. right lateral view showing the capitulum and peduncle; ¢c. carinal view; d. rostral view showing the capitulum and peduncle. Scale bar: 3 mm (a—d). Figure 21. Amigdoscalpellum tenue (Hoek, 1883) (MNHN-IU-2019-4921). a. left lateral view showing the capitulum and pedun- cle; b. right lateral view showing the capitulum and peduncle; ec. carinal view showing the capitulum and peduncle; d. rostral view showing the capitulum and peduncle. Scale bar: 2 mm (a—d). Diagnosis. Capitulum with 13 fully calcified plates; cari- Description. Capitulum yellowish, elongate-oval shape; na slightly bowed, umbo at apex; upper latus trapeziform; surface with distinct lines of growth. Carina large, sim- inframedian latus small, narrow; carinal latus not project- — ply bowed. Scutum with umbo at apex, slightly recurved, ing beyond carina; peduncle short, cylindrical. projecting slightly over tergum; tergum triangular, stout, zse.pensoft.net 672 Pitriana, PR et al.: Deep-sea barnacles from the Karubar expedition (1991) broad, apex recurved, scutal margin almost straight. Up- per latus quadrangular, apex slightly projecting over scu- tum. Rostral latus quadrangular, scutal and basal margins parallel. Carinal latus quadrangular, carinal margin almost straight. Cirrus I unequal, anterior ramus oval, posterior ramus more slender. Maxillule with notch between two or three stout setae at upper angle, a group of more slen- der setae on cutting edge; mandible with three teeth ex- cluding inferior angle; labrum with numerous blunt teeth on straight, cutting edge. Measurements of specimen: height of capitulum 7.26 mm, width 3.74 mm, thickness 1.40 mm; length of peduncle 3.00 mm, width 1.99 mm. Distribution. Indian Ocean, Antarctic and Southern (North East of Prince Edward Island); known depth 2,516 m (Sha- laeva and Boxshall 2014). In this study, Amigdoscalpellum tenue was found at Kei Islands, Indonesia. Type locality. 46°46'S, 45°31'E; depth: 2,475 m; bottom temperature: 1.5 °C; bottom: globigerina ooze (Hoek 1883). Genus Catherinum Zevina, 1978 Acroscalpellum Hoek, 1907: 85.— Newman and Ross 1971: 42 (part). — Liu and Ren 1985: 202. Catherinum Zevina, 1978b: 1348. Diagnosis. Capitulum with 13 or 14 plates; inframedian latus very narrow, bacilliform, umbo sub-medial; umbo of upper latus apical or sub-apical; carinal latus broad, umbo sub-basicarinal, not extending beyond carinal mar- gin; caudal appendages long. Type species. Scalpellum recurvitergum Gruvel, 1902; type locality: Atlantic Ocean, SW of Azores. Catherinum rossi (Rao & Newman, 1972) Figure 22 Arcoscalpellum rossi Lakshmana Rao & Newman, 1972: 82, fig. 7. Catherinum rossi. — Zevina, 1978b: 1348. — Liu and Ren 1985: 203, fig. 12, pl. 2, figs 11-14. — 2007: 250, fig. 108. — Chan et al. 2009b: 113, figs 92-94. Material examined. — Tanimbar Island: 1 specimen, MNHN-IU-2019-4933, Sth. DW 02, 05°47'S, 132°13'E, 209-240 m depth, 22 October 1991. Diagnosis. Capitulum smooth, elongate, 14 fully calci- fied plates; inframedia latus rectangular, not constricted; rostrum large, ovo-triangular; caudal appendages short. Description. Capitulum long, narrow, sparsely covered with hairs, plates separated by narrow, chitinous inter- spaces, marked with growth lines. Occludent margin strongly convex; carinal margin irregularly straight; apex slightly retroverted towards carinal side. Carina long, simply bowed; roof flat; parietes well developed towards distal half of plate. Tergum triangular, occludent margin short, convex, scutal and basal margins almost straight, Figure 22. Catherinum rossi (Rao & Newman, 1972) (MNHN-IU-2019-4933). a. left lateral view showing the capitulum and pe- duncle; b. right lateral view showing the capitulum and peduncle; c. rostral view showing the capitulum and peduncle; d. carinal view. Scale bar: 3 mm (a—d). zse.pensoft.net Zoosyst. Evol. 96 (2) 2020, 649-698 carinal margin concave. Scutum with umbo apical, over- lapping occludent margin of tergum. Upper latus triangu- lar; carinal latus twice as long as broad; inframedian latus rectangular; rostral latus nearly rectangular in outline; rostrum large, elongate triangular, broad above, pointed below. Cirrus I unequal, anterior ramus oval, posterior ramus more slender; cirrus VI with caudal appendages. Maxillule not notched stout spine along the cutting edge; mandible with three teeth excluding inferior angle. Mea- surements of specimen: height of capitulum 9.67 mm, width 5.52 mm, thickness 2.58 mm; length of peduncle 2.79 mm, width 2.29 mm. Distribution. East China Sea, Pacific Ocean, Tatwan (Chan et al. 2009b). In this study, Catherinum rossi was found at Tanimbar Island, Indonesia. Type locality. Stn. 3, Hess Guyot (17°53.2'N, 174°24.8'W), 1,692—1,735 m depth (Sigsbee beam trawl); Stn. 1, Allison Guyot (18°31.0'N, 179°36.0'W), 1,413— 1,645 m depth (otter trawl) (Rao and Newman 1972). Genus Planoscalpellum Zevina, 1978 Arcoscalpellum Hoek, 1907: 85 (part). Planoscalpellum Zevina, 1978b: 1347. Diagnosis. Upper latus with umbo on scutal margin; in- framedian latus with umbo apical or sub-basal; carinal latus with umbo on carinal part, may be protruding. 673 Type species. Scalpellum planum Hoek, 1883; type lo- cality: 42°42'S, 134°10'E; depth: 4,755 m; bottom tem- perature: 0.2 °C; bottom: red clay. Planoscalpellum distinctum (Hoek, 1883) Figure 23 Scalpellum distinctum Hoek, 1883: 111-112, pl. VI, figs 10, 11. — 1907: 83, pl. VIL, fig. 12 — Nilsson-Cantell 1927: 750, text-fig. 4. Planoscalpellum distinctum. — Zevina, 1981: 186-187. — Foster and Buckeridge 1995 a: 360-361, fig. 7/A—C) — Shalaeva and Boxshall 2014: 36, fig. 22. Material examined. — Tanimbar Island: 2 specimens, MNHN-IU-2019-4928, Stn. DW 49, 08°00'S, 132°59'E, 206-210 m depth, 29 October 1991. Diagnosis. Capitulum with 13 plates separated from each other by distinct chitinous interspaces; scutum almost tri- angular; upper latus hexagonal, large. Description. Capitulum flattened, elongate-oval shape; plates with distinct growth lines, separated from each other by rather broad chitinous interspaces. Scutum tri- angular, occludent margin arched, umbo of scutum api- cal, apex slightly turned upwards. Tergum triangular, occludent margin not very long. Carina simply bowed. Upper latus irregularly hexagonal, rostral latus irreg- ularly quadrangular, infra-median latus in the form of wine-glass, carinal latus irregularly quadrangular. Pe- Figure 23. Planoscalpellum distinctum (Hoek, 1883) (MNHN-IU-2019-4928). a. left lateral view showing the capitulum and pe- duncle; b. right lateral view showing the capitulum and peduncle; c. rostral view showing the capitulum and peduncle; d. carinal view. Scale bar: 2 mm (a—d). zse.pensoft.net 674 Pitriana, P et al.: Deep-sea barnacles from the Karubar expedition (1991) duncle short, with very small, numerous, calcareous scales. Cirrus I unequal, anterior ramus oval, posterior ramus more slender; cirrus VI with short caudal append- ages. Maxillule notched, three stout spines on the up- per side; mandible with three teeth, excluding inferior angle. Measurements of two specimens: height of ca- pitulum 6.80—7.90 mm, width 3.68—4.36 mm, thickness 1.24—1.26 mm; length of peduncle 1.39-—2.49 mm, width 1.80—2.04 mm. Distribution. Pacific, Western Central; known depth range 1,302 to 2,745 m (Shalaeva and Boxshall 2014). In this study, Planoscalpellum distinctum was found at Tanimbar Island, Indonesia. Type locality. 2°33'S, 144°4'E; depth: 1,926 m; bot- tom temperature: 2.1 °C; bottom: Globigerina ooze (Hoek 1883). Genus Teloscalpellum Zevina, 1978 Arcoscalpellum Hoek, 1907: 85 (sectio, part.). — Newman and Ross 1971: 42 (part.); Section V. — Pilsbry 1907: 47 (part.). Teloscalpellum Zevina, 1978b: 1350. — Liu and Ren 1985: 212. — 2007: 257. Diagnosis. Capitulum with 13 or 14 plates; carinal lateral umbo at basi-carinal angle, angle not extending beyond carinal margin; inframedian latus triangular or rod-like- shaped, umbo apical or sub-apical; caudal appendage multi-segmented. Type species. Scalpellum spicatum Zevina, 1975. Teloscalpellum ecaudatum (Calman, 1918) Figure 24 Scalpellum ecaudatum Calman, 1918b: 106, text-fig. 2. Teloscalpellum ecaudatum. — Zevina, 1981: 365, fig. 282. — Chan 2009: Sa iesiE sz Material examined. — Kei Islands: 1 specimen, MNHN- TU-2019-4917, Stn. CP 35, 06°08'S, 132°45'E, 390- 502 m depth, 27 October 1991. —Tanimbar Island: 1 specimen, MNHN-IU-2019-4918, Stn. CP 59, 08°20'S, 132°11'E, 399-405 m depth, 31 Oc- tober 1991. Diagnosis. Capitulum compressed, elongated; 14 plates, sculptured with radiating ribs; inframedian latus triangu- lar; rostro-lateral plates very low; penis and caudal ap- pendages absent. Description. Capitulum with all plates strongly ribbed, lines of growth marked by fine, inconspicuous striation. Tergum with occludent margin straight, carinal margin convex, but straight concave near apex. Scutum with occludent margin convex, other margins straight, apex slightly overlapping tergum. Carina evenly curved with flat roof bordered by strong ridges; upper latus quadran- gular; rostrum small, triangular, overlapped at sides by rostral latera; inframedian latus very narrow. Peduncle shorter than capitulum, covered with strong scales. Cir- Figure 24. Teloscalpellum ecaudatum (Calman, 1918) (MNHN-IU-2019-4917). a. left lateral view showing the capitulum and peduncle; b. right lateral view showing the capitulum and peduncle; ec. carinal view; d. rostral view showing the capitulum and peduncle. Scale bar: 5 mm (a-—d). zse.pensoft.net Zoosyst. Evol. 96 (2) 2020, 649-698 rus I with unequal rami (anterior ramus: 11 segmented, posterior ramus: 13 segmented). Maxilla globular with serrulate setae; maxillule not notched; mandible with three major teeth, lower margin with three to four den- ticles. Measurements of two specimens: height of capit- ulum 12.07—-18.59 mm, width 6.17—11.47 mm, thickness 2.68—-5.41 mm; length of peduncle 6.30—8.33 mm, width 3.21—5.41 mm. Distribution. Java Sea, Indonesia; Philippines (Chan 2009). In this study, Teloscalpellum ecaudatum was found at Kei Islands and Tanimbar Island, Indonesia. Type locality. Java Sea; 7°35'S, 114°30'30"E; 132-315 m depth (Calman 191 8b). Genus Trianguloscalpellum Zevina, 1978b Arcoscalpellum Hoek, 1907: 85 (part). — Pilsbry 1907a: 47. - Newman and Ross 1971: 42 (part). Trianguloscalpellum Zevina, 1978b: 1349. — Liu and Ren 1985: 205. Diagnosis. Capitulum of female or hermaphrodite with 13 or 14 completely calcified plates; tergum and scutum with umbones apical; middle latera triangular, sometimes quadrangular, with apical umbo reaching upper latera; ca- rinolatera triangular or subtriangular with apical umbo; caudal appendages distinctly long. Type species. Scalpellum balanoides Hoek, 1883: 129, pl. V, fig. 15, pl. X, fig. 11, pl. XI, figs 1-3; type locali- 675 ty: 5°42'S, 132°25'E; depth: 236 m; bottom: mud; it was found attached to an arm of a Comatula or Pentacrinus. Trianguloscalpellum balanoides (Hoek, 1883) Figure 25 Scalpellum balanoides Hoek, 1883: 129, pl. V, fig. 15, pl. X, fig. 11, pl. XI, figs 1-3. — Broch 1922: 242, fig. 10 — Weltner 1922: 63, taf. II, fig. 7, taf. III, fig. 8. — Nilsson-Cantell 1931: 2. — Hiro 1937b: 42, fig. 33. Scalpellum gonionotum Pilsbry, 1907b: 360. — Pilsbry 1911: 65, pl. IX, figs 2-4. — Hiro 1937a: 43. Trianguloscalpellum balanoides. — Zevina, 1978b: 1349. — 1981: 294, fig. 221. —Liu and Ren 1985: 210, fig. 16, pl. XI, figs 15—17. — 2007: 256-257, fig. 111. — Rosell 1991: 20-22, fig. 2c, d. — Huang 1994: 517. — 2001: 318. — Jones et al. 2001: 253. -Young 2001: 464, fig. 7. — Jones 2007: 292. — Chan et al. 2009b: 116, figs 95, 96. — Shalaeva and Boxshall 2014: 40, fig. 26. Material examined. — Kei Islands: 1 specimen, MNHN- TU-2019-4923, Stn. CP 16, 05°17'S, 132°50'E, 330- 350 m depth, 24 October 1991, attached to crinoid. — Tanimbar Island: 2 specimens, MNHN- TU-2019-4924, Stn. CP 86, 09°26'S, 131°13'E, 223-225 m depth, 4 November 1991. Diagnosis. Capitulum with 13 fully calcified plates, cov- ered by thin membrane; carina and extremely short, sim- ply bowed without distinct roof; umbo of carina at apex; upper latus triangular; carinal latus almost as high as ca- rina, umbo apical, carinal latus very large, almost similar size to tergum. Figure 25. Trianguloscalpellum balanoides (Hoek, 1883) (MNHN-IU-2019-4923). a. left lateral view showing the capitulum and peduncle; b. right lateral view showing the capitulum and peduncle; c. rostral view showing the capitulum and peduncle; d. carinal view. Scale bar: 2 mm (a—d). zse.pensoft.net 676 Description. Capitulum flat, narrow at base, same breadth as peduncle. Scutum about twice as long as broad; occlu- dent margin almost straight, umbo at apex. Tergum trian- gular, the same size as scutum. Carina very short, simply, not very strongly, bowed, umbo at apex. Upper latus tri- angular, carinal margin absent. Rostral latus quadrangu- lar, umbo at apex of angle formed by rostral and scutal margins. Infra-median latus elongate, triangular, umbo apical. Carinal latus very large, convex, umbo at top of carinal margin. Cirrus I unequal, posterior ramus more slender than anterior ramus; cirrus VI with caudal ap- pendages. Maxillule distinctly notched, two stout spines on upper side; mandible with three teeth excluding inferi- or angle. Measurements of specimen: height of capitulum 8.20 mm, width 4.40 mm, thickness 1.75 mm; length of peduncle 5.06 mm, width 2.10 mm. Distribution. Pacific, Western Central; South and East China Sea; South of Sumatra, Banda Sea, Indonesia; Vietnam; Philippines; Taiwan; South of Japan; attached to crinoids, hydroids; 220—1,097 m depth (Jones et al. 2001; Chan et al. 2009b; Shalaeva and Boxshall 2014). In this study, Trianguloscalpellum balanoides was found at Kei Islands and Tanimbar Island, Indonesia. Type locality. 5°42'S, 132°25'E; depth: 232 m; bottom: mud; attached to arm of a Comatula or Pentacrinus (Hoek 1883). Trianguloscalpellum hirsutum (Hoek, 1883) Figure 26 Scalpellum hirsutum Hoek, 1883: 88, pl. IV, fig. 19. — Gruvel 1905: 66, fig. 74. — Pilsbry 1907a: 25. Arcoscalpellum hirsutum. — Newman & Ross, 1971: 62-64, fig. 28. Trianguloscalpellum hirsutum. — Zevina, 1981: 309, fig. 233. — Chan et al. 2009b: 119-121, figs 100-102. — Chan et al. 2010: 13, Pitriana, P et al.: Deep-sea barnacles from the Karubar expedition (1991) figs 1F, 9A—-H. — Jones 2012: 371, 376. — Shalaeva and Boxshall 2014: 44, fig. 29. Material examined. — Tanimbar Island: 1 specimen, MNHN-IU-2019-4908, Stn. CP 91, 08°44'S, 131°05'E, 884-891 m depth, 5 November 1991. Diagnosis. Capitulum with 14 plates covered by mem- brane, covered by very long hairs; carina simply bowed, umbo at apex, roof flat; upper latus triangular; rostrum very narrow stripe distinctly visible at surface. Description. Capitulum small, long hairs covering plates. Scutum elongate, convex, more than twice as long as broad. Tergum large, elongate rhomboid, umbo at apex. Carina simply bowed, roof flat. Upper latus al- most triangular, carinal-basal margin arched. Rostrum small, linear-shaped. Rostral latus quadrangular, basal and scutal margins parallel. Infra-median latus triangu- lar, umbo apical. Carinal latus with irregular shape, umbo near apex. Peduncle short. Cirrus I with rami unequal, an- terior ramus oval, posterior ramus more slender; cirrus VI with caudal appendages. Maxillule not notched, two stout spines on upper side; mandible with three teeth excluding inferior angle. Measurements of the specimen: height of capitulum 12.18 mm, width 6.50 mm, thickness 3.74 mm; length of peduncle 4.10 mm, width 3.43 mm. Distribution. Borneo, Indonesia; Pacific, Western Cen- tral and Southeast; Australia; Taiwan; Antarctica; at- tached to gastropod shells, rocks; 1,502—1,965 m depth (Chan et al. 2010; Jones 2012; Shalaeva and Boxshall 2014). In this study, 7rianguloscalpellum hirsutum was found at Tanimbar Island, Indonesia. Type locality. Pacific, Western Central (Moluccas Sea); 0°48'S, 120°58'E; depth: 252 m; bottom temperature: 2.4 °C; bottom: rock (Hoek 1883). peduncle; b. right lateral view showing the capitulum and peduncle; ec. carinal view; d. rostral view showing the capitulum and peduncle. Scale bar: 5 mm (a-—d). zse.pensoft.net Zoosyst. Evol. 96 (2) 2020, 649-698 Genus Verum Zevina, 1978 Arcoscalpellum Pilsbry, 1907: 47 (part). - Newman and Ross 1971: 42 (part). Verum Zevina, 1978b: 1348. Diagnosis. Capitulum of female or hermaphrodite, with 13-14 completely calcified plates; tergum with carinal margin straight or slightly to moderately recurving; scutum with or without short apico-lateral arm, umbo apical; carina with apical or subapical umbo, evenly curved or angular- ly flexed; upper latera sub-triangular or pentagonal, umbo apical or subapical; middle latera narrow, triangular or quadrangular, umbo basal or sub-basal; carinolatera short, umbo basal to sub-basocarinal, usually not or slightly pro- jecting beyond rim of capitulum; caudal appendages short. Type species. Scalpellum zenkevichi Zevina, 1972: 44, fig. 3; type locality: Pacific Ocean, 24°27'S, 70°42'E. Verum australicum (Hoek, 1883) Figure 27 Scalpellum australicum Hoek, 1883: 118, pl. V, fig. 11. — Weltner 1922: 63. — Nilsson-Cantell 1938: 7. Verum australicum. — Zevina, 1981: 223-224, fig. 160. — Jones et al. 1990: 5. — Jones 1992: 156-160, figs 7-8. — Jones 2012: 371, 376. — Shalaeva and Boxshall 2014: 52, fig. 36. 677 Material examined. — Tanimbar Island: 1 specimen, MNHN-IU-2019-4930, Stn. CP 86, 09°26'S, 131°13'E, 223-225 m depth, 4 November 1991. Diagnosis. Capitulum with 13 plates closely locked to- gether, surface smooth with slightly prominent ridges; carina simply bowed with flat roof; upper latus trapezi- form; inframedian latus elongate, narrow; carinal latus with umbo projecting beyond carina. Description. Capitulum elongate-oval, flat with distinct ridges and furrow. Scutum slightly convex, umbo apical, oc- cludent margin arched. Tergum triangular, umbo apical, dis- tinctly recurved. Rostral latus irregularly quadrangular; infra median latus narrow, umbo near base; carinal latus quadran- gular. Peduncle slightly conical, short, scales near capitulum larger than those near base. Cirrus I unequal, anterior ramus oval, posterior ramus more slender; cirri II to VI very long, rami subequal, segments elongate; cirrus VI with caudal ap- pendages. Maxillule not notched, large spine on upper side, cutting edge almost straight; mandible with three large teeth excluding inferior angle. Measurements of specimen: height of capitulum 10.33 mm, width 6.09 mm, thickness 2.08 mm; length of peduncle 4.62 mm, width 2.24 mm. Distribution. Indian Ocean, Western; Pacific Western Central and Southeast; Zanzibar; known depth range 463- 2,561 m (Shalaeva and Boxshall 2014). In this study, Ver- um australicum was found at Tanimbar Island, Indonesia. Figure 27. Verum australicum (Hoek, 1883) (MNHN-IU-2019-4930). a. left lateral view showing the capitulum and peduncle; b. right lateral view showing the capitulum and peduncle; c. rostral view showing the capitulum and peduncle; d. carinal view. Scale bar: 3 mm (a-d). zse.pensoft.net 678 Pitriana, P et al.: Deep-sea barnacles from the Karubar expedition (1991) Type locality. between New Guinea and Australia (12°08'S, 145°10'E): Challenger station 184: depth: 2,561 m: bottom temperature: 1.8 °C; substrate grey ooze (Hoek 1883). Verum carinatum (Hoek, 1883) Figure 28 Scalpellum carinatum Hoek, 1883: 76, pl. II, figs 7, 8. — Pilsbry 1907a: 53, fig. 18. — Gruvel 1920: 20. — Barnard 1925: 3. — Broch 1953: 7. — Weisbord 1977: 243-244, pl. 27, figs 3, 4. Scalpellum imperfectum Pilsbry, 1907a: 75, fig. 30, pl. IV, figs 15-18. — Annandale 1913: 233. — Barnard 1924: 47. — MacDonald 1929: 537. — Broch 1953: 9. — Stubbings 1961: 11, text-fig. 2. — 1967: 234. Meroscalpellum imperfectum. — Newman & Ross, 1971: 119, text-fig. 62. Verum carinatum. — Zevina, 1981: 225-226, fig. 163. — Shalaeva and Boxshall 2014: 53, fig. 37. Material examined. — Kei Islands: 1 specimen, MNHN- TU-2019-4920, Stn. DW 18, 05°18'S, 133°01'E, 205- 212 m depth, 24 October 1991. Diagnosis. Capitulum with 14 plates separated by chitin- Ous interspaces, surface smooth; carina bowed, angularly bent, with flat roof; upper latus irregular pentagonal; in- framedian latus large, wine-glass-shaped. Description. Capitulum elongate, plates covered by thin, chitinous membrane. Scutum elongated, apex pointed, occludent margin very convex. Tergum flat, triangular, apex very recurved, occludent margin very arched. Carina with umbo at top of flat roof. Upper latus flat, irregular pentagonal: rostrum narrow, elon- gated; rostral lateral convex with rostral margin short; inframedian latus wine-glass-shaped; carinal latus flat, large. Peduncle short, calcareous scales distinct. Cirrus I unequal, anterior ramus broader than posterior ramus; cirri II to VI long, rami equal; cirrus VI with caudal appendages. Maxillule not notched, two large spines on upper side, cutting edge almost straight; mandible with three large teeth excluding inferior angle. Mea- surements of specimen: height of capitulum 9.14 mm, width 4.53 mm, thickness 1.74 mm; length of peduncle 2.54 mm and width 2.20 mm. Distribution. Atlantic, excluding polar areas; Pacific, Southeast. Known depth range 600 to 2,400 m (Shalaeva and Boxshall 2014). In this study, Verum carinatum was found at Kei Islands, Indonesia. Type locality. Station 135, near the Island of Tristan da Cunha; depth: 1,800 m; bottom: rock, shells (Hoek 1883). Verum novaezelandiae (Hoek, 1883) Figure 29 Scalpellum novae-zelandiae Hoek, 1883: 124, pl. V, figs 7, 8. — 1907: 100, pl. VHI, figs 4, 4a; 1913: 14. — Weltner 1897: 249. — 1922: 71. — Annandale 1905: 83. — 1906b: 389. — 1908: pl. 5, fig. 7. — 1913: 231. — Broch 1927a: 540, fig. 512. Scalpellum Novae-Zealandiae. — Gruvel, 1905: 88, fig. 99. Scalpellum (Scalpellum) novae-zelandiae. — Calman, 1918b: 123. Arcoscalpellum novaezelandiae. — Foster, 1978: 65, pl. 8D, fig. 39. > pets WT ° ~ Npr es La: ey ve = a os Figure 28. Verum carinatum (Hoek, 1883) (MNHN-IU-2019-4920). a. left lateral view showing the capitulum and peduncle; b. right lateral view showing the capitulum and peduncle; ¢. rostral view showing the capitulum and peduncle; d. carinal view. Scale bar: 3 mm (a-d). zse.pensoft.net Zoosyst. Evol. 96 (2) 2020, 649-698 79 Figure 29. Verum novaezelandiae (Hoek, 1883) (MNHN-IU-2019-4919). a. left lateral view showing the capitulum and peduncle; b. right lateral view showing the capitulum and peduncle; c. carinal view; d. rostral view showing the capitulum and peduncle. Scale bar: 3 mm (a-—d). non Scalpellum Novae-Zelandiae. — Gruvel, 1902: 54, pl. 2, figs 12, 13, 15. —Gruvel 1912: 346 (= K. parazelandiae Young, 1998b). Verum novaezelandiae. — Zevina, 1978b: 1348. — Zevina 1981: 228, fig. 165. — Chan et al. 2009b: 17, figs 11, 12A—H, 13A—D. — Chan et al. 2010: 17, figs 11-13. — Shalaeva and Boxshall 2014: 53, fig. 39. Material examined. — Tanimbar Island: 1 specimen, MNHN-IU-2019-4919, Stn. CP 87, 08°47'S, 130°49'E, 1017-1024 m depth, 5 November 1991. Diagnosis. Capitulum with 13 plates covered by very thin, chitinous membrane; carina simply bowed, flat roof with umbo apical; upper latus quadrangular or rather pentagonal, lower edge being truncated; inframedian la- tus elongate, quadrangular; carinal latus large, elongate, umbo near base; peduncle short. Description. Capitulum flatted, elongate, plates separat- ed by narrow, chitinous interspaces. Scutum quadrangu- lar, umbo apical, occludent margin arched. Tergum tri- angular with umbo slightly produced apex. Carina with bowed, flat roof, increasing little in width from upper to lower. Upper latus irregularly trapeziform; rostral latus with umbo at top of rostral margin; infra-median latus irregular, quadrangular, elongate, narrow; carina latus quadrangular. Cirrus I unequal, anterior ramus broad- er than posterior ramus; cirrus VI with short caudal ap- pendages. Maxillule slightly notched, two large spines on upper side; mandible with three large teeth excluding inferior angle; labrum slightly convex, with numerous teeth. Measurements of specimen: height of capitulum 13.46 mm, width 6.98 mm, thickness 2.99 mm; length of peduncle 6.02 mm, width 3.23 mm. Distribution. Indo-west Pacific: Eastern Africa, Gulf of Aden, eastwards to New Zealand; Philippines; Taiwan; at- tached to sunken wood, gorgonians, glassy spicule of hex- actinellid sponges; 822-4850 m depth (Shalaeva and Box- shall 2014; Jones and Hosie 2016). In this study, Verum novaezelandiae was found at Tanimbar Island, Indonesia. Type locality. 37°34'S, 179°22'E; depth: 1,260 m; bottom temperature: 4.2 °C; bottom: grey ooze (Hoek 1883). Order SESSILIA Lamarck, 1818 Suborder VERRUCOMORPHA, Pilsbry 1916 Verrucomorpha Pilsbry, 1916: 14. - Newman et al. 1969: 281. — New- man and Ross 1971: 135.— Newman 1987: 8.— 1996: 501. — Bucker- idge 1994: 89. — Young 1998: 74. Diagnosis. Shell asymmetrical with four plates (carina, ros- trum, tergum and scutum), base membranous or calcareous. Family VERRUCIDAE Darwin, 1854 Verrucidae Darwin, 1854: 495. — Gruvel 1905: 169. — Withers 1935: 323. — Newman et al. 1969: 281. - Newman and Ross 1971: 135. — Buckeridge 1994: 89. — Young 1998: 74. zse.pensoft.net 680 Pitriana, P. et al.: Deep-sea barnacles from the Karubar expedition (1991) Diagnosis. Scutum and tergum without depressor mus- cles; movable only on one side; other side immovable, united with rostrum and carina. Genus Alteriverruca Pilsbry, 1916 Verruca Section D Altiverruca Pilsbry, 1916: 40. Altiverruca Pilsbry, 1916: 40 — Newman et al. 1969: 282. —- Newman and Ross 1971: 135. — Ren 1984b: 172. — Buckeridge 1994: 92. — Young 1998: 77. Diagnosis. Verrucids with erect form; bases of plates not inflected. Operculum close to vertical, myophore absent. Type species. Verruca hoeki Pilsbry, 1907. Altiverruca navicula (Hoek, 1913) Figure 30 Verruca navicula Hoek, 1913: 134, figs 4-6. — Nilsson-Cantell 1927: 778, figs a-f. Altiverruca navicula. — Buckeridge 1994: 100, fig. 5. — Chan et al. 2009b: 280, figs 245-247. — Chan et al. 2010: 29, figs 2E, 21, 22. Material examined. — Tanimbar Island: 3 specimens, MNHN-IU-2019-4941, Stn. CP 91, 08°44'S, 131°0OS'E, 884-891 m depth, 5 November 1991, attached to pedun- cle of Alcockianum persona (Annandale, 1916). Diagnosis. Shell not depressed; carina and rostrum inter- locking with single rib from each plate; movable plates large, scutum with four articular ribs, tergum with six articu- lar ribs, growth lines very distinct; caudal appendages long. Description. Shell yellowish. Movable scutum elongately triangular, apex distinctly beaked, projecting freely; sur- face with numerous articular ridges. Movable tergum large, quadrangular; surface with strongly developed, curved ax- ial articular ridge. Carina and rostrum irregular quadrangu- lar, with carina higher, rostrum broader. Fixed tergum with two parts: (1) triangular portion very narrow at apex, slight- ly broader in its inferior (2) flat and broad part at a rear por- tion of shell. Fixed scutum pointed with distinctly beaked apex; composed of broader, nearly flat, triangular portion and narrower inflected portion, only widening towards its inferior. Base of shell elongatedly oval-shaped. Cirrus I with rami very unequal (anterior ramus: 12-segmented, posterior ramus: 28-segmented); cirrus VI with caudal ap- pendages. Maxilla bilobed, fringed with setae, except on the notch; maxillule widely notched, horizontally elongat- ed, two large spines above notch, numerous dense setae at notch; mandible with three teeth excluding inferior angle; labrum slightly concave, conical teeth on cutting margin. Distribution. Pacific Ocean (Chan et al. 2010). In this study, Altiverruca navicula was found at Tanimbar Is- land, Indonesia. Type locality. (1) 9°3.4'S, 119°56.7'E; depth 959 m; bot- tom: globigerina ooze. (2) 3°37.7'S, 131°26.4'E; depth 924 m; bottom: fine grey mud (Hoek 1913). Genus Newmaniverruca Young, 1998 Verruca Section B: Verruca, Group of Verruca alba Pilsbry, 1916: 25 (part.). Verruca Section B: Verruca, Group of Verruca calotheca Pilsbry, 1916: 30 (part.). Verruca (Verruca): Foster, 1979: 68 (part.). — Zevina 1987: 1812 (part. — Buckeridge 1994: 90 (part.). Newmaniverruca Young, 1998: 77. Diagnosis. Shell with four plates (carina, rostrum, scu- tum and tergum); operculum parallel to base. Figure 30. A/tiverruca navicula (Hoek, 1913) (MNHN-IU-2019-4941). a. left lateral view; b. right lateral view; c. upper lateral view; d. lower lateral view. Scale bar: 7 mm (a-d). zse.pensoft.net Zoosyst. Evol. 96 (2) 2020, 649-698 Type species. Verruca albatrossiana Pilsbry, 1912: 292. — 1916: 47; type locality: East of Luzon, Philippines. Newmaniverruca albatrossiana (Pilsbry, 1912) Figure 31 Verruca albatrossiana Pilsbry, 1912: 292. — 1916: 47. — Buckeridge 1994: 91, fig. la-f. — Buckeridge 1997: 129. Verruca grex Hoek, 1913: 142, pl. XI, figs 7-13, pl. XII, figs 11-13. Verruca (Euverruca) albatrossiana. — Broch, 1931: 45. Verruca (Verruca) albatrossiana. — Ren, 1984: 168, fig. 2, pl. 1 (7-11). Newmaniverruca albatrossiana. — Young, 1998: 77. — Chan 2009: 74, figs 2G, 22A-I. Material examined. — Kei Islands: 1 specimen, MNHN- TU-2019-4942, Stn. CC 21, 05°14'S, 133°00'E, 688-694 m depth, 25 October 1991, attached to glass rope sponge. Diagnosis. Rostrum and fixed scutum with unusual length; rostrum low, fixed scutum larger than fixed tergum. Description. Movable plates parallel to base, wall of parietal vertically ribbed; fixed scutum without internal pit. Movable scutum with crescentic ridge and longi- tudinal striations; movable tergum with articular ribs and diagonal rib. Apices of fixed scutum and tergum contiguous. Carina occupying carino-rostral wall, api- ces marginal. Cirrus I with rami unequal and serrulate setae; cirrus VI with caudal appendages. Maxilla glob- ular, with fringing setae; maxillule notched, two large setae on upper side; mandible with three teeth exclud- ing inferior angle. Distribution. Indo-west Pacific: NW Australia; South China Sea; China; Philippines; attached to cidaroid spines (e.g. Stereocidaris indica philippinensis Mortensen, 1928), 345-620 m depth (Jones and Hosie 2016). In this study, Newmaniverruca albatrossiana was found at Kei Islands, Indonesia. 681 Type locality. East of Luzon, Philippines (Pilsbry 1912). Suborder BALANOMORPHA Pilsbry, 1916 Operculata tribe Symetrica Gruvel, 1905: 189. Balanomorpha Pilsbry, 1916: 47 (Balanidae sensu Darwin, 1854). Diagnosis. Peduncle absent; shells bilaterally symmetri- cal on either side of rostrocarinal axis; shell with parietal plates of carina, rostrum and one to three pairs of lateral plates, all separate from one another, variously fused or totally concrescent; base membranous or calcareous; her- maphroditic or with dwarf males. Family BATH YLASMATIDAE Newman & Ross, 1971 Bathylasmatidae Newman & Ross, 1971: 138 (part.). — Newman and Ross 1976: 37, 45 (part.). Bathylasmatinae Newman & Ross, 1976: 37, 45. — Buckeridge 1983: 68. — Newman 1996: 502. — Buckeridge 1999: 522. — Jones 2000: 23¥,,tables-1, 2. Diagnosis. Shell with four or stx plates; wall solid or per- meated by single row of chitin-filled longitudinal canals; radii absent; one or both rami of cirri I and cirri II some- times antenniform; labrum without notch in crest. Subfamily HEXELASMINAE Newman & Ross, 1976 Hexelasminae Newman & Ross, 1976: 37, 46. Diagnosis. Shell with six plates, infiltrated by chitin-filled tubes; basis calcareous; scuta parallel to basis; tergum with distinct spur; cirrus II resembling cirrus I more than cirrus III. Type genus. Hexalasma Hoek, 1913. Figure 31. Newmaniverruca albatrossiana (Pilsbry, 1912) (MNHN-IU-2019-4942). a. left lateral view; b. right lateral view; ec. up- per lateral view; d. lower lateral view. Scale bar: 3 mm (a—d). zse.pensoft.net 682 Pitriana, P. et al.: Deep-sea barnacles from the Karubar expedition (1991) Genus Hexelasma Hoek, 1913 Hexelasma Hoek, 1913: 244 (part.). — Utinomi 1965: 13. — Jones 2000: 240, figs 51, 64, tables 28-30. Aaptolasma Newman & Ross, 1971: 158. Diagnosis. Parietes solid; radii absent; plates with a ca- rina, paired carinolatera and latera and a rostrum without any sign of fusion; caudal appendages absent. Type species. Hexelasma velutinum Hoek, 1913: 246 (part.); type locality: Indonesia, 6°08'S, 121°19'E (Sibo- ga Station 105). Hexelasma arafurae Hoek, 1913 Figure 32 Hexelasma arafurae Hoek, 1913: 251, pl. 25, figs 12-16. — Utinomi 1965: 11.— Newman and Ross 1971: 155. — 1976: 46. — Foster 1981: fig. 6F—-H. — Jones 2000: 246, fig. 51, tables 28-31. Aaptolasma arafura. — Foster, 1978: 79. Material examined. — Kei Islands: 2 specimens, MNHN- TU-2019-4851, Stn. DW 18, 05°18'S, 133°01'E, 205— 212 m depth, 24 October 1991. Diagnosis. Shell with very wide orifice; alae broadly-tri- angular with summits oblique; scutum folded longitudi- nally, articular ridge slightly prominent, deep pit for ad- ductor muscle; tergum beaked, with prominent articular ridge, spur very broadly rounded; inner side of scutum and tergum orange-coloured. Description. Shell yellowish, conical, with six plates. Or- ifice diamond-shaped; scutum triangular elongated with protruding growth-ridges; tergum smaller than scutum, apex beaked, carinal margin rounded, growth-ridges less distinct than on scutum. Cirrus I with unequal rami (an- terior ramus: 8-segmented; posterior ramus: 12-segment- ed), dense long setae on surface areas. Cirrus II with equal rami, dense long setae. Cirri [VW—VI with equal rami with numerous segments; segments almost without exception furnished with two pairs very long, stiff, needle-like spines along inner faces. Measurements of specimen: basal length of shell 14.32 mm, orifice length 8.00 mm, carinal height 12.39 mm, orifice width 6.49 mm, basal width 13.76 mm. Distribution. Arafura Sea, Indonesia; 205-560 m depth (Jones and Hosie 2016). In this study, Hexelasma arafu- rae was found at Kei Islands, Indonesia. Type locality. Indonesia; Siboga stat. 262; 5°54'S, 132°49'E; 560 m depth; bottom: solid bluish-grey mud (Hoek 1913). Superfamily BALANOIDEA Leach, 1817 Balanidae Leach, 1817: 68 — Pilsbry 1916: 48 — Newman et al. 1969: 284 — Newman and Ross 1976: 38, 49 — Newman 1993: 408. Balaninae — Darwin, 1854: 175. Balanoidea — Newman, 1996: 502. Diagnosis. Shell wall composed of four or six plates (rostrum, carina and one to three pairs of laterals); pa- rietes solid or tubiferous, when tubiferous rarely sec- ondarily filled; radii solid or tubiferous. Basis com- monly calcareous, solid or permeated by tubes, rarely membranous; when basis calcareous, internal surfaces of compartments commonly with uniform ribs; when calcareous commonly forming complex interdigitations with wall. Opercular plates occlude aperture; articu- lations between the pairs generally shallow or fused. Cirrus I with rami subequal or grossly unequal; cirri I Figure 32. Hexelasma arafurae Hoek, 1913 (MNHN-IU-2019-4851). a. view from above; b. right lateral view. Scale bar: 4 mm (a, b). zse.pensoft.net Zoosyst. Evol. 96 (2) 2020, 649-698 and III with rami never antenniform; cirrus HI resem- bling Hl more than IV; caudal appendages absent; penis with basidorsal point. Labrum thin, never bullate; crest with pronounced medial incision; mandible quadri- or quinquedentoid; second and following teeth with one or more subsidiary cusps; fifth tooth often vestigial; inferi- or angle commonly molariform. Family ARCHAEOBALANIDAE Newman & Ross, 1976 Archaeobalanidae Newman & Ross, 1976: 38, 49. Diagnosis. Shell with four or sixplates; parietes solid, rarely tubiferous; tubes uniformly or irregularly arranged; radii solid; basis commonly calcareous, rarely tubiferous. Subfamily ARCHAEOBALANINAE Newman & Ross, 1976 Archaeobalaninae Newman & Ross, 1976: 38, 49. Diagnosis. Shell with four or six plates; parietes solid or tubiferous; when tubiferous, tubes uniformly arranged in single row; interlaminate figures simple; basis calcareous or membranous, when membranous wall solid. Type genus. Archaeobalanus Menesini, 1971: 19. Genus Conopea Say, 1822 Conopea Say, 1822: 323. Conoplea Gray, 1825: 98, 103. Balaninus Costa, 1839: 181. Balanus Section B Darwin, 1854: 216. Patella-Balanus Hoek, 1913: 160, 162, 221. 683 Diagnosis. Shell firm, strong, with six thick compart- ments; parietes with or without pores; radi solid with summits parallel to basal margin of parietes and dentic- ulated sutural margins; basis calcareous, elongated along carino-rostral axis, boat-shaped; orifice smooth, not dentated; scutum with simple growth ridges; penis with basidorsal point; attached to gorgonians or antipatharians. Type species. Conopea elongata Say, 1822; type locality: eastern Florida, USA. Conopea cymbiformis (Darwin, 1854) Figure 33 Balanus cymbiformis Darwin, 1854: 221, pl. 3, fig. 5a, b. — Broch 1931: 85, fig. 29a, b. — Nilsson-Cantell 1938: 55, pl. 2, fig. 3. — Utinomi 1962: 219, fig. 2.— Dong et al. 1982: 103. Balanus proripiens Hoek, 1913: 228, pl. 24, figs 1-3, pl. 24, figs 1-3. — Nilsson-Cantell 1921: 331, fig. 70c, d. Pyrgoma jedani Hoek, 1913: 262, pl. 27, figs 3-8. Conopea cymbiformis. — Newman & Ross, 1976: 55. Material examined. — Kei Islands: 9 specimens, MNHN-IU-2019-4823, Stn. DW 30, 05°39'S, 132°56'E, 111-118 m depth, 26 October 1991; 1 specimen, MNHN- TU-2019-4824, Sth. DW 22, 05°22'S, 133°01'E, 82 m depth, 25 October 1991. Diagnosis. Shell irregularly conical; orifice oval with swollen border; basis irregularly cup-formed; scutum and tergum triangular, not calcified together; spur of tergum feebly developed. Description. Shell yellowish with orange rust-brown in proximal areas. Carina, carinolatera and latera with pale orange-brown and rust red-brown longitudinal stripes, latter may have oblique white spots. Radi with pale or- ange-brown and rust red-brown horizontal striation. Oper- Figure 33. Conopea cymbiformis (Darwin, 1854) (MNHN-IU-2019-4823). a. left lateral view; b. right lateral view; c. view from above; d. basal view. Scale bar: 3 mm (a—d). zse.pensoft.net 684 cular plates with scutum pink-brown, transparent; tergum transparent white. Shell may appear longer and lower, due to elongation of carina and rostrum or low and comparative- ly shorter, due to development of rostrum alone or more up- right and comparatively higher, with neither carina nor ros- trum elongated. Cirrus I with unequal rami (anterior ramus: 7-segmented; posterior ramus: 12-segmented). Cirri II-VI with equal rami, numerous segments. Penis very long, del- icate hairs scattered over surface, a few more disposed near tip. Labrum deeply notched, two small teeth on each side of notch. Mandibles with five teeth, inferior angle not distinct- ly separated from fifth; distance between tips of first and second teeth slightly more than that between those of sec- ond and third teeth; third tooth larger; fourth and fifth small- er than others. Maxillule with straight edge and numerous large setae. Measurements of specimen: basal length of shell 7.73 mm, orifice length 4.60 mm, carinal height 8.70 mm, orifice width 3.88 mm, basal width 4.94 mm. Distribution. Indo-west Pacific: Indian Ocean; Gulf of Aden, India, east to Fiji and NW to Indonesia, N Aus- tralia, Malay Arch.; China; Philippines; S Japan; Fiji Is; attached to coenosarc of gorgonians or antipatharians; Iit- toral453 m depth (Jones and Hosie 2016). In this study, Conopea cymbiformis was found at Kei Islands, Indonesia. Type locality. Near Madras, India; attached to a gorgo- nian (Darwin 1854). Conopea navicula (Darwin, 1854) Figure 34 Balanus navicula Darwin, 1854: 221, pl. 3, fig. 6a-d. — Hoek 1913: 223, pl. 22, fig. 26, pl. 23, figs 1-3. — Stubbings 1936: 48. — Utinomi 1962: 74, fig 1.— Dong et al. 1982: 103. Acasta spinitergum Foster, 1982: 209, fig. 4d. Pitriana, PR et al.: Deep-sea barnacles from the Karubar expedition (1991) Conopea navicula. — Newman & Ross, 1976: 55. Material examined. — Tanimbar Island: 12 specimens, MNHN-IU-2019-4821, Stn. CP 82, 09°32'S, 131°02'E, 215-219 m depth, 4 November 1991 attached to gorgo- nian; 4 specimens, MNHN-IU-2019-4822, Stn. DW 49, 08°00'S, 132°59'E, 206-210 m depth, 29 October 1991, attached to gorgonian. Diagnosis. Shell with parietes and basis not porose; cari- no-lateral compartments very narrow, almost same width from top to bottom; radii with smooth sutural edges; scu- tum externally striated longitudinally. Description. Specimens covered with coenosarc of coral, except orifice. Easily recognisable species due to narrow carino-lateral plate, which is nearly same width at top as bottom; scutum externally longitudinally striated; pari- etal plates studded with calcareous points. Parietal plates pearly white, solid, superficially appearing to possess longitudinal tubes, growth lines horizontal. Alae moder- ately developed. Basis calcareous. Size small. Rostrum well developed, concave, lying at angle of ~ 45°. Later- als very well developed. Carino lateral parietes thin, ra- dii and alae well developed. Carina tall, about half width of rostrum. External surfaces of all parietes with very small, calcareous studs, regularly spaced, arranged along horizontal growth lines. Opercular plates sunk down into orifice. Cirrus I with unequal rami (anterior ramus: 5-segmented; posterior ramus: 7-segmented). Cirrus II with unequal rami (anterior ramus: 6-segmented; pos- terior ramus: 9-segmented). Cirri II-VI with subequal rami more slender, longer, with segments more elongate. Penis very long, tapering towards tip, bearing few, very minute hairs. Maxilulle with straight edge with numer- ous large setae. Mandibles with five teeth and inferior angle. Measurements of four specimens: basal length of Figure 34. Conopea navicula (Darwin, 1854) (MNHN-IU-2019-4822). a. left lateral view; b. orifice view. Scale bar: 1 mm (a, b). zse.pensoft.net Zoosyst. Evol. 96 (2) 2020, 649-698 Shell 2.23—-4.22 mm, orifice length 1.09—2.02 mm, carinal height 2.04—3.09 mm, orifice width 0.94—1.59 mm, basal width 1.79—2.80 mm. Distribution. Indo-west Pacific, from Gulfs of Aden and Persia, India, Malaysia, Indonesia, Gulf of Siam, to southern Japan; 45—220 m depth (Jones and Hosie 2016). In this study, Conopea navicula was found at Tanimbar Island, Indonesia. Type locality. Madras, India; attached to gorgonian (Dar- win 1854). Genus Solidobalanus Hoek, 1913 Solido-Balanus Hoek, 1913: 159, 192. Balanus (Solidobalanus). — Pilsbry, 1916: 220. Solidobalanus. — Newman & Ross, 1976: 23, 50. Diagnosis. Shell parietes solid, six plates; radii solid, well developed, with denticulate sutural edges; basis cal- careous, solid; complemental male, when present, in pit of rostral plate of hermaphrodite. Type species. Balanus auricoma Hoek, 1913: 198, pl. XVIII, figs 20-22, pl. XIX, figs 1-7; type locality: Ter- nate, Indonesia (Siboga station 136). Subgenus Solidobalanus Hoek, 1913 Solido-Balanus Hoek, 1913: 159, 192. Balanus (Solidobalanus). — Pilsbry, 1916: 220. Solidobalanus. — Newman & Ross, 1976: 23, 50. Diagnosis. Parietes and radii rather thick, smooth, with- out pores; basis flat, calcareous; rostrum not very elongat- ed; radii with strongly septate sutural edges; tergal spur narrow; crests for lateral depressor muscles of scutum absent; adductor ridge weak or absent; tergum flat or with 685 shallow furrow without infolded sides; conspicuous teeth on anterior margins of fourth cirri absent. Type species. Balanus auricoma Hoek, 1913: 198, pl. XVIII, figs 20-22, pl. XIX, figs 1-7; type locality: Ter- nate, Indonesia (Siboga station 136). Solidobalanus auricoma (Hoek, 1913) Figure 35 Balanus (Solidobalanus) auricoma Hoek, 1913: 198, pl. XVIII, figs 20-— 22, pl. XIX, figs 1-7. — Broch 1922: 323, fig. 62. — 1931: 71. — Foster 1978: 100, fig. 60. — Rosell 1981: 303. Solidobalanus (Solidobalanus) auricoma. — Newman & Ross, 1976: 50. Solidobalanus auricoma. — Foster ,1981: 364, fig. 2G. — Rossel 1991: 38. Material examined. — Tanimbar Island: 2 specimens, MNHN-IU-2019-4842, Stn. CP 45, 07°54'S, 132°47'E, 302-305 m depth, 29 October 1991, shell only; 10 spec- imens, MNHN-IU-2019-4841, Sth. DW 49, 08°00'S, 132°59'E, 206-210 m depth, 29 October 1991, attached to sea urchin spines; 1 specimen, MNHN-IU-2019-4838, Stn. CP 82, 09°32'S, 131°02'E, 215-219 m depth, 4 No- vember 1991, attached to antipatharian; 7 specimens, MNHN-IU-2019-4847, Stn. CP 85, 09°22'S, 131°14'E, 240-245 m depth, 4 November 1991, attached to spines of sea urchin; 63 specimens, MNHN-IU-2019-4850, Stn. CP 86, 09°26'S, 131°13'E, 223-225 m depth, 4 Novem- ber 1991, attached to spines of sea-urchins. — Kei Islands: 1 specimen, MNHN-IU-2019-4849, Stn. CP 05, 05°49'S, 132°18'E, 296-299 m depth, 22 October 1991, attached to lateral plate of Chirona te- nuis, 1 specimen, MNHN-IU-2019-4844, Stn. CP 12, 05°23'S, 132°37'E, 413-436 m depth, 23 October 1991, attached to antipatharian; 6 specimens, MNHN- ITU-2019-4848, Stn. CP 16, 05°17'S, 132°50'E, 330- 350 m depth, 24 October 1991, attached to dead gorgo- nian, spines of sea-urchins, arm of crinoid; 3 specimens, MNHN-IU-2019-4840, Stn. CP 35, 06°08'S, 132°45'E, 390-502 m depth, 27 October 1991; 1 specimen, Figure 35. Solidobalanus auricoma (Hoek, 1913) (MNHN-IU-2019-4850). a. upper lateral view from above; b. left lateral view; c. right lateral view; d. basal view. Scale bar: 2 mm (a—d). zse.pensoft.net 686 MNHN-IU-2019-4843, Stn. CP 36, 06°0S'S, 132°44'E, 210-268 m depth, 27 October 1991. Diagnosis. Shell flatly-conical, colour reddish basally, whitish apically; radii narrow, summits slightly oblique; opercular plates with golden hairs along occludent mar- gins; scutum with articular ridge slightly prominent, ad- ductor ridge absent; tergum narrow, scutal margin dis- tinctly dentated. Description. Shell with plates ribbed longitudinally. Shell colour brownish-pink to dull rose-pink, ribs tend- ing to white, colour often faded with specimens appear- ing uniform white. Parietes of carinolatera very narrow, with single, conspicuous, longitudinal ridge. Scutum with occludent margin straight, surface indistinctly ridged, pit for adductor muscle scarcely visible. Tergum short, narrow, scutal margin straight, unusually distinctly dentated, carinal margin short, convex, depressor muscle crests moderately well developed. Opercular plates with long, golden setae fringing occludent margins, especial- ly distally. Cirri I-II with rami slightly unequal, covered with setae; cirri II-VI longer, more slender, dense setae on inner face. Mandible with four teeth, second to fourth with accessory cusps, lower angle molariform with three blunt cusps in series, lower edge with row of stiff setae. Measurements of five specimens: basal length of shell 6.11-8.18 mm, orifice length 4.18-5.83 mm, carinal height 4.11-6.30 mm, orifice width 2.58—3.19 mm, basal width 4.51—5.96 mm. Distribution. Banda Sea (Moluccas, Indonesia); SW Australia; New Zealand; New Caledonia; Philippines to southern Japan; Malaysian water; Gulf of Oman, Persia. 27-502 m depth (Jones and Hosie 2016). In this study, Solidobalanus auricoma was found at Kei Islands and Tanimbar Island, Indonesia. Type locality. Ternate anchorage; 27 m depth; bottom: mud and stone; numerous specimens on the surface of pieces of rock (Hoek 1913). Pitriana, P. et al.: Deep-sea barnacles from the Karubar expedition (1991) Solidobalanus pseudauricoma (Broch, 1931) Figure 36 Balanus (Solidobalanus) pseudauricoma Broch, 1931. —Utinomi 1949: 97, fig. 4. Solidobalanus (Solidobalanus) pseudauricoma. — Newman & Ross, 1976: 51. — Jones 2007: 294. Material examined. — Kei Islands: 46 specimens, MNHN-IU-2019-4818, Stn. CP 25, 05°30'S, 132°52'E, 336-346 m depth, 26 October 1991, 30 specimens at- tached to gorgonian, 16 specimens attached to cor- al; 6 specimens, MNHN-IU-2019-4825, Stn. CP 27, 05°33'S, 132°51'E, 304-314 m depth, 27 October 1991; 1 specimen, MNHN-IU-2019-4826, Stn. CP 05, 05°49'S, 132°18'E, 296-299 m depth, 22 October 1991, attached to crinoid; 13 specimens, MNHN-IU-2019-4827, Stn. CP 16, 05°17'S, 132°50'E, 330-350 m depth, 24 October 1991; 2 specimens, MNHN-IU-2019-4828, Stn. DW 22, 05°22'S, 133°01'E, 82 m depth, 25 October 1991, 1 spec- imen attached to gorgonian stem; 8 specimens, MNHN- TU-2019-4830, Stn. CP 05, 05°49'S, 132°18'E, 296-299 m depth, 22 October 1991, 5 specimens attached to gor- gonian, 1 specimen attached to crinoid. — Tanimbar Island: 11 specimens, MNHN- TU-2019-4839, Stn. CP 86, 09°26'S, 131°13'E, 223-225 m depth, 4 November 1991, several specimens attached to spines of sea urchins (associated with Solidobalanus au- ricoma (Hoek, 1913)). Diagnosis. Shell with smooth, glossy white plates, co- loured stripes absent; internal plates with thick, solid, fine- ly ribbed longitudinally; base non-porous, radially ribbed. Description. Shell plates white, stripes absent. Sever- al specimens with pale pink tinge, one with pale brown- ish-pink parietes with small, narrow ellipsoidal whitish spots, latter orientated longitudinally producing reticulated effect. Radi whitish, pink tinge along distal borders. Scutal growth lines without longitudinal striations; articular ridge absent; pit for adductor muscle small, round. Tergum with Figure 36. Solidobalanus pseudauricoma (Broch, 1931) (MNHN-IU-2019-4818). a. left lateral view; b. orifice view; c. basal view; d. right lateral view. Scale bar: 4 mm (a-—d). zse.pensoft.net Zoosyst. Evol. 96 (2) 2020, 649-698 shallow, wide furrow running from apex to base. Cirrus I with unequal (anterior ramus: 7-segmented; posterior ramus: 15-segmented). Cirrus IT with rami subequal (an- terior ramus: 11-segmented: posterior ramus: 12-segment- ed). Cirri I and I with very dense, long setae on surface areas. Cirri II-VI with rami slightly subequal, rounded. Penis sturdy, not long. Labrum with very shallow notch, three or four irregularly arranged, blunt teeth on each side. Maxillule with distinct, narrow notch with two large se- tae on upper side. Mandibles with five teeth, second and third bifid and fifth is rudimentary. Measurements of five specimens: basal length of shell 6.23—12.08 mm, orifice length 4.63—9.62 mm, carinal height 3.48—13.22 mm, ori- fice width 3.24—6.33 mm, basal width 5.66—9.56 mm. Distribution. Manado Bay (Indonesia); Japan (Broch 1931-1932). In this study, Solidobalanus pseudauricoma was found at Kei Islands and Tanimbar Island, Indonesia. Type locality. Manado Bay, Indonesia (1°31'N, 124°47'E), 500 m depth; Japan (32°25'N, 128°33'E), 400 m depth (Broch 1931-1932). Remarks. In the type description, Broch (1931-1932) commented that the specimens were white, without stripes. However, several of the specimens collected by KARUBAR had a pale pink tinge and one specimen (from station DW22) had pale brownish-pink parietes with small, narrow ellipsoidal whitish spots, the latter ori- entated longitudinally, thus producing a reticulated effect. Radii whitish with pink tinge along distal borders. Genus Striatobalanus Hoek, 1913 Balanus Section F (part.) Darwin, 1854: 277. Chirona Gray, 1825: 37. — Pilsbry, 1916: 204. Balanus (Striato-Balanus) Hoek, 1913: 159, 179. 687 Chirona (Striatobalanus). —Newman & Ross, 1976: 23, 50 (as subgenus). Striatobalanus. — Jones, 2004: 150. Diagnosis. Parietes and radii without pores; radii usual- ly narrow; scutum striated longitudinally with adductor ridge; tergum furrow deep; cirrus I with rami very un- equal in length. Type species. Balanus amaryllis var. (a), var. (b) Darwin, 1854: 279, pl. 7, fig. 6a—c. Type locality. Unknown. Striatobalanus amaryllis (Darwin, 1854) Figure 37 Balanus amaryllis var. (a), var. (b) Darwin, 1854: 279, pl. 7, fig. 6a—c. — Hoek 1913: 179, pl. 15, figs 17-21, pl. 16, figs 1-4. — Pilsbry 1916: 217. — Nilsson-Cantell 1925: 329. — Hiro 1936: 624. — 1939c: 243. — Stubbings 1936: 174. — Utinomi 1962: 216. — 1969: 88. Balanus amaryllis dissimilis Lanchester, 1902: 369, pl. XXXIV, figs 3—3C, with var. clarovittata Lanchester, 1902: 370. Balanus amaryllis var. a (= Balanus roseus Lamarck, 1818). — Gruvel 1905: 250, with var. b (niveus). — Gruvel, 1905: 250 (= Balanus amaryllis var. b Darwin, 1854: 279)). Balanus (Chirona) amaryllis. — Nilsson-Cantell 1921: 329, pl. 3, fig. 9. — Rosell 1981: 302. Balanus amaryllis forma euamaryllis. — Broch, 1922: 321. Balanus amaryllis forma laevis Broch, 1931: 67, fig. 24. Balanus sp. Dong & Mao, 1956: 290, fig. 8. Chirona (Striatobalanus) amaryllis. — Newman & Ross, 1976: 50. Chirona amaryllis. — Foster, 1981: 350. Balanus amaryllis. — Dong et al., 1982: 95. Balanus amaryllis euamaryllis Dong et al., 1982: 96, fig. A-C. Chirona amaryllis: Zevina et al., 1992: 72, fig. 49. Striatobalanus amaryllis. — Jones, 2004: 150. — Chan et al. 2009b: 222, 190-192. Figure 37. Striatobalanus amaryllis (Darwin, 1854) (MNHN-IU-2019-4814). a. orifice view; b. right lateral view. Scale bar: 3 mm (a, b). zse.pensoft.net 688 Pitriana, P. et al.: Deep-sea barnacles from the Karubar expedition (1991) Material examined. — Tanimbar Island: 1 specimen (only shell), MNHN-IU-2019-4814, Stn. CP 65, 09°14'S, 132°27'E, 174-176 m depth, 1 November 1991, attached to broken sea urchin shell. Diagnosis. Shell conical; orifice large, pentagonal, toothed; shell colour pinkish, lighter transparent stripes run longitudinally over surface; radii very narrow, reddish, upper margins very oblique; alae broader, lighter colour. Description. Shell conical; tips of rostrum and cari- na slightly curved inwards. Orifice large, pentagonal, toothed. Colour yellowish-white, with slightly darker longitudinal lines on main parts of plates. Radii with very oblique summits, broadest a little distance from the orifice, narrower towards basis. Alae broader than radii, summits rounded. Specimen without scutum, ter- gum and soft parts. Measurements of specimen: basal length of shell 16.76 mm, orifice length 9.13 mm, cari- nal height 9.89 mm, orifice width 7.24 mm, basal width 15.14 mm. Distribution. Arafura Sea, Indonesia. Indo-west Pa- cific: South and East Africa, Indian Ocean; Australia (N); Singapore; Malay Arch.; Cambodia; Gulf of Thai- land; Mouth of Bassac; Vietnam (Bay of Nhatrang; Lien Chien, Tourane; Codor Is); Hong Kong; S China Sea; China; Philippines; Taiwan; Japan (S); attached to rocks, gastropod shells, rusting iron, fouling species; sublittoral, 5-500 m depth (Jones and Hosie 2016). In this study, Striatobalanus amaryllis was found at Ta- nimbar Island, Indonesia. Type locality. Unknown. Striatobalanus tenuis (Hoek, 1883) Figure 38 Balanus tenuis Hoek, 1883: 154, pl. 13, figs 29-33. — 1913: 190, pl. 17, figs 14-19, pl. 18, fig. 1. - Gruvel 1905: 247, fig. 275. — Pilsbry 1916: 216. — Barnard 1924: 74. — Nilsson-Cantell 1925: 34, fig. 13, pl. 1, figs 5, 6. — 1927: 785. — 1938: 46. — Broch 1931: 70. — Hiro 1937b: 439, fig. 24. — Utinomi 1962: 216. — 1968: 174. — 1969: 88, fig. 6. — Utinomi and Kikuchi 1966: 6. Balanus albus Hoek, 1913: 185, pl. 16, figs 12, 13, pl. 17, figs 1-6. - Stubbings 1936: 41, fig. 18. Chirona (Striatobalanus) tenuis. — Newman & Ross, 1976: 50. — Rosell 1989: 33, pl. 10g. — 1991: 38. Balanus (Chirona) tenuis. — Ren & Liu, 1978: 161, fig. 22, pl. 7, figs 6-10. — Nilsson-Cantell 1921: 302. Balanus tenuis. — Dong et al.. 1982: 97, fig. A-C. Chirona tenuis. — Zevina et al.. 1992: 74, fig. 50. Striatobalanus tenuis. — Jones. 2004: 152. — Liu and Ren 2007: 363, fig. 161. — Chan 2009: 74, fig. 2H and fig. 23A—G. Material examined. — Kei Islands: 5 specimens, MNHN- TU-2019-4820, Stn. CC 10, 05°21'S, 132°30'E, 329- 389 m depth, 23 October 1991; 1 specimen (shell only), MNHN-IU-2019-4835, Stn. CP 16, 05°17'S, 132°50'E, 330-350 m depth, 24 October 1991. — Tanimbar Island: 1 specimen, MNHN-IU-2019-4832, Stn. DW 49, 08°00'S, 132°59'E, 206-2010 m depth, 29 Oc- tober 1991; 1 specimen (shell only), MNHN-IU-2019-4833, Stn. CP 65, 09°14'S, 132°27'E, 174-176 m depth, 1 Novem- ber 1991; 4 specimens (shell only), MNHN-IU-2019-4834, Stn. CP 79, 09°16'S, 131°22'E, 239-250 m, 3 November 1991; 1 specimen (only shell), MNHN-IU-2019-4813, Stn. CP 85, 09°22'S, 131°14'E, 240-245 m, 4 November 1991, attached to shell of gastropod; 2 specimens, MNHN- TU-2019-4819, Stn. CP 86, 09°26'S, 131°13'E, 223-225 m depth, 4 November 1991, attached to gastropod shells. Diagnosis. Shell colour whitish to yellowish; sur- face smooth, glossy; orifice pentagonal shaped, deep- ly toothed; radii narrow, summits very oblique, slightly concave; basis solid; scutum with longitudinal striations; tergum with short, rather broad spur. Description. Shell with six plates, conical; orifice large, distinctly toothed; radii well-developed; basis thin, ribbed, solid. Scutum triangular, not elongated, with longitudinal Figure 38. Striatobalanus tenuis (Hoek, 1883) (MNHN-IU-2019-4820). a. orifice view; b. left lateral view; ¢. right lateral view; d. basal view. Scale bar: 10 mm (a—d). zse.pensoft.net Zoosyst. Evol. 96 (2) 2020, 649-698 striations; tergum slightly beaked, exhibiting traces of lon- gitudinal striations, spur short, slightly narrow, depressor crests distinctly developed. Cirri I to VI with rami slight- ly equal in length, transparent and orange-coloured. Penis rather long, sparse, minute hairs on the surface, tapering to- wards tip, distinctly hairy at tip. Maxillule with edge slight- ly straight, all the setae similar sized. Mandibles with five teeth, fourth and fifth small. Measurements of five speci- mens basal length of shell 18.92—28.30 mm, orifice length 13.78-16.29 mm, carinal height 18.02—20.24 mm, orifice width 10.15—13.70 mm, basal width 20.51—28.03 mm. Distribution. Arafura Sea, Indonesia; Indo-west Pacif- ic: S Africa; Persian Gulf; Indian Ocean; Australia (N); Singapore; Malay Arch.; Vietnam; Hong Kong; S Chi- na Sea; E China Sea; Philippines; Taiwan; Japan (S); W Pacific; attached to crabs, gastropod, bivalve, shells sol- itary coral, bark of coconut, gorgonians, antipatharians, stones; 7-551 m depth (Jones and Hosie 2016). In this study, Striatobalanus tenuis was found at Kei Islands and Tanimbar Island, Indonesia. Type locality. West of Mindoro, Philippines; 6°8'N, 121°19'E; depth: 275 m, at coral-bottom (Hoek 1913). Family PYRGOMATIDAE Gray, 1825 Balanidae Leach, 1817: 68 (part.). — Darwin 1854: 33 (part.). Balaninae. — Nilsson-Cantell, 1921: 306 (part.). Pyrgomatidae Gray, 1825: 102. —- Newman 1996: 503. Pyrgomatinae. — Ross & Newman, 1973: 149. Diagnosis. Wall of four plates wholly concrescent; pa- rietes solid or tubiferous; when tubiferous, tubes occur between outer lamina and sheath or between external ribs of wall; interlaminate figures complex, essentially arbo- rescent; radii solid; basis calcareous, rarely tubiferous, membranous in Pyrgopsella. 689 Subfamily MEGATREMAOTINAE Holthuis, 1982 Bosciinae Newman & Ross, 1976: 59. Megatrematinae Holthuis, 1982: 319. Diagnosis. Wall of four plates wholly concrescent; oper- cular valves normal; tergum with weakly developed lat- eral depressor muscle crests or crests lacking; when shell concrescent, sheath with paired sulci. Type genus. Megatrema Sowerby, 1823, by original des- ignation (Holthuis 1982). Tribe PYRGOMINI Ross & Pitombo, 2002 Pyrgominini Ross & Pitombo, 2002: 58. Type genus. Pyrgomina Baluk & Radwanski, 1967. Genus Pyrgomina Baluk & Radwanski, 1967 Pyrgomina Baluk & Radwanski, 1967b: 691, pl. 1-2. Type species. Pyrgomina seguenzai Baluk & Radwanski, 1967a: 485. Type locality. Pliocene (Piacenzian Stage); Gournes, Iraklion nomarchia, Island of Crete, Greece, 35°06'N, 25°47'E; host coral unknown (Ross and Pitombo 2002). Pyrgomina sp. Figure 39 Material examined. — Tanimbar Island: 1 specimen, MNHN-IU-2019-4817, Stn. CP 86, 09°26'S, 131°13'E, 223-225 m depth, 4 November 1991, attached to coral. ee Figure 39. Pyrgomina sp. (MNHN-UI-2019-4817). a. orifice view; b. left lateral view; ec. right lateral view. Scale bar: 2 mm (a—c). zse.pensoft.net 690 Diagnosis. Shell purple-pinkish, tall, conical; oriface oval; base permeated by pores, expanded out of the coral; tergum and scutum subtriangular. Description. Shell stands exserted on the coral, externally furnished with ribs, lower part rounded with radiating ribs; oriface oval, small and narrow. Scutum triangular, basal margin curved; tergum triangular with narrow spur. Cirrus I with rami unequal (anterior ramus: 10-segmented; pos- terior ramus: 15-segmented); posterior ramus of cirrus [V without spines. Maxilla globular; maxillule not notched, eight large setae on straight cutting edge; mandible with five major teeth. Measurements of specimen: basal length of shell 7.46 mm, orifice length 2.42 mm, carinal height 10.45 mm, orifice width 1.41 mm, basal width 6.15 mm. Distribution. Pyrgomina sp. was found at Tanimbar Is- land, Indonesia. Family BALANIDAE Leach, 1817 Balanidae Leach, 1817: 68. — Gray 1825: 104. — Darwin 1854: 33 (part.). — Pilsbry 1916: 48. — Nilsson-Cantel 1921: 306. - Newman and Ross 1976: 59. — Foster 1978: 95. — Buckeridge 1983: 103. — Newman 1996: 503. — Pitombo 2004: 262. — Chan et al. 2009b: 229. Diagnosis. Shell with four or six plates; parietes tubif- erous; tubes arranged in single uniform row formed be- tween inner and outer lamina, supplementary tubes may form basally; interlaminate figures complex, arborescent; radii solid or tubiferous, more-or-less developed; basis calcareous, commonly tubiferous; caudal appendages ab- sent; penis commonly with basidorsal point. Subfamily AMPHIBALANINAE Pitombo, 2004 Amphibalaninae Pitombo, 2004: 263. Diagnosis. Shell with four or six plates; parietal tubes with one or more rows, commonly transverse septa; radii with transverse teeth on sutural edge with denticles on lower side only; alae not cleft; basis with single tubif- erous; scutum with conspicuous adductor ridge; tergum with well-developed depressor muscle crests, growth lines in tergum spur display an obvious change in direc- tion; second maxilla with smooth anterior margin of dis- tal lobe, acuminate setae with enlarged, modified tips. Genus Amphibalanus Pitombo, 2004 Amphibalanus Pitombo, 2004: 263. Diagnosis. Shell with four or six plates; parietal tubes with one or more rows, commonly transverse septa; radii with transverse teeth on sutural edge, with denticles on zse.pensoft.net Pitriana, P et al.: Deep-sea barnacles from the Karubar expedition (1991) lower side only; alae not cleft; basis with single tubif- erous; scutum with conspicuous adductor ridge; tergum with well-developed depressor muscle crests, growth lines in tergum spur display an obvious change in di- rection; second maxilla with smooth anterior margin of distal lobe, acuminate setae with enlarged, modified tips. Cirrus III with inner face of endopod with pinnate setae, rarely bifurcate (complex) setae. Type species. Balanus amphitrite Darwin, 1854: 240 (part.), pl. 5, fig. 2a-d, i-k, m—o; type locality: Natal, South Africa. Amphibalanus amphitrite (Darwin, 1854) Figure 40 Balanus amphitrite Darwin, 1854: 240 (part.), pl. 5, fig. 2a-d, i-k, m-o. — Weltner 1897: 264. — Hoek 1913: 167. — Pilsbry 1916: 89. — Zevina et al. 1992: 89, fig. 61. — Puspasari et al. 2001: 7. Balanus amphitrite var. (1) communis Darwin, 1854: 240, pl. 5, fig. 2e, h, 1. Balanus amphitrite communis. — Nilsson-Cantell, 1921: 311, fig. 64. Balanus amphitrite forma hawaiiensis Broch, 1922: 314, fig. 56 (part.). Balanus amphitrite forma denticulata Broch, 1927b: 133, fig. 14 (part.). Balanus amphitrite hawaiiensis. — Hiro, 1937c: 432, figs 20, 21. Balanus amphitrite cochinensis Nilsson-Cantell, 1938b: 43, fig. 11a-e. Balanus amphitrite var. fluminensis Oliveira, 1941: 21, pl. 4, fig. 4, pl. 5, figs 1, 2, pl. 8, figs 1-5. Balanus amphitrite var. aeratus Oliveira, 1941: 22, pl. 4, fig. 5, pl. 9, figs 1-4. Balanus amphitrite herzi Rogers, 1949: 28, pl. 1, figs 6, 12-15. Balanus amphitrite franciscanus Rogers, 1949: 29, pl. 1, figs 5, 7, 16-19. Balanus amphitrite var. columnarius Tarasov & Zevina, 1957: 179, 184, fig. 68 a-e. Balanus amphitrite denticulata Henry, 1959: 192, pl. 1, fig. 5, pl. 3, fig. 7, upper row right. Balanus amphitrite amphitrite. — Harding, 1962: 274, pl. la—g, pl. 2a—k. — Dong et al. 1982: 90, fig. A-E. — Rosell 1981: 302. Balanus amphitrite var. Hawaiiensis. — Stubbings, 1963b: 15. Amphibalanus amphitrite. — Pitombo, 2004: 263, 274, figs 2A, B, 7A, B, 8C. — Chan et al. 2009b: 241. — Chen et al. 2014: 1071. — Pochai et al. 2017: 27, fig. 9. — Pitriana et al. 2020: 42, fig. 21. Material examined. — Tanimbar Island: 4 specimens, MNHN-IU-2019-4815, Stn. CP 52, 08°03'S, 131°48'E, 1244-1266 m depth, 30 October 1991. Diagnosis. Primary parietal tubes with transverse septa; exterior of shell with longitudinal purple striations, hori- zontal striations absent; tergum short with wide spur; cirri II-VI with erect teeth below posterior angles of distal; cirrus IIT without complex setae. Description. Shell six-plated, conical, round; external- ly smooth, white with groups of well-spaced, dark pur- ple vertical stripes; horizontal striations on shell surface absent. Interior of paries with single row of tubes; radii solid, wide; alae with summits moderately oblique; basis Zoosyst. Evol. 96 (2) 2020, 649-698 691 Figure 40. Amphibalanus amphitrite (Darwin, 1854) (MNHN-IU-2019-4815). a. orifice view; b. right lateral view; ¢. basal view. Scale bar: 4 mm (a-c). porose, calcareous. Scutum externally striped, occludent margin toothed, lateral depressor muscle pit small. Ter- gum with spur wider than long, less than its own width from basi-scutal angle. Cirrus I with unequal rami (ante- rior ramus: 10-segmented; posterior ramus: 16-segment- ed). Cirrus II with slightly equal rami (anterior ramus: 10-segemented; posterior ramus: 12-segmented). Cirrus II with rami subequal (anterior ramus: 12-segmented; posterior ramus: 13-segmented. Cirri IV—VI with rami subequal, longer, more slender. Penis moderately long, tapering distally, sparse, minute hairs on surface, distinct- ly hairy distally. Labrum deeply notched, numerous teeth on each side. Maxilla without notch below upper pair of spines, nine spines between upper and lower pairs, few short spines on inferior angle; lower pair of spines usually on slight prominence, which may be sometimes moder- ately strong or strong. Mandible with five teeth, inferi- or angle with second tooth bifid. Measurements of four specimens: basal length of shell 5.27—12.10 mm, orifice length 3.81—7.89 mm, carinal height 3.64—5.65 mm, or- ifice width 2.72—5.22 mm, basal width 4.62—12.03 mm. Distribution. Cosmopolitan in tropical and subtropical waters. Bermuda & SE USA to Brazil; England and West Europe to South coast of Africa Red, Black and Medi- terranean Seas; Suez Canal; SE Africa; Indian Ocean; Australia; Singapore; Malaysia; Réam (Cambodia); Gulf of Siam; Vietnam; Condor Island; Tang Trien (South An- nam); Cauda Nhatrang; Hongay, Tonkin; S China Sea; Hong Kong; China; Bohai Sea; Taiwan; Philippines; Japan; South Honsyu, Kyusyu & Ryukyu Island; Vlad- ivostok; Hawaii; central California to SW Mexico; foul- ing species; lower littoral to sublittoral (Jones and Hosie 2016). Recently, also recorded from Ambon Island and Saparua Island, Indonesia (Pitriana et al. 2020). In this study, Amphibalanus amphitrite was found at Tanimbar Island, Indonesia. Type locality. Natal, on a piece of bamboo (Darwin 1854). Remarks. Known as an important fouling species of ships and marine installations. The suggestion of an- ti-fouling paint on the bases of the specimens examined suggests that these specimens were probably knocked off the ship during trawling operations, explaining the great depth at which these specimens were collected, as the normal depth range is 0-9 m. Discussion Prior to the Karubar expedition, 24 species of barnacles had been collected from the Kei Islands and Aru Island by the Siboga expedition (Hoek 1913). Other pertinent refer- ence works to the barnacles from these islands are Jones et al. (2001) and Jones and Hosie (2016), who recorded 15 species from the Kei Islands and Aru Island. In addition to the works of Hoek (1913), Jones et al. (2001) and Jones and Hosie (2016), Broch (1931-1932) reported on 67 species of barnacles collected by the Dan- ish expedition to the Kei Islands (1922) and deposited in the Zoological Museum of Copenhagen University. In his report, only four species, Euscalpellum rostratum (Dar- win, 1851), Lepas (Anatifa) anatifera Linnaeus, 1758, Conchoderma virgatum Spengler, 1789 and Acasta den- tifer (Broch, 1922), were explicitly collected in the Kei Is- lands. The other barnacle species recorded were collected at other places along the route of this expedition, such as Lampung Bay, Krakatau, Java Sea, Sunda Strait, Makas- sar Strait, Tual, Banda Neira, Ambon and Saparua Bay. The lists of Hoek (1913), Broch (1931-1932), Jones et al. (2001) and Jones and Hosie (2016) record a total of 25 species from the Kei Islands, Aru Island and Tanimbar Island. The results currently recorded herein reveal that AO species are now recorded from these Islands. Of the 40 species herein, ten are recorded in previ- ous studies: Megalasma striatum Hoek, 1883 (listed in Hoek (1913)); Scalpellum stearnsi Pilsbry, 1890 (listed in Hoek (1913)); Annandaleum japonicum (Hoek, 1883) zse.pensoft.net 692 (listed in Hoek (1913)); Annandaleum laccadivicum (An- nandale, 1906) (listed in Hoek (1913)); Regioscalpellum moluccanum (Hoek, 1883) (listed in Hoek (1913)); A/- tiverruca navicula (Hoek, 1913) (listed in Hoek (1913) and Buckeridge (1994)); Newmaniverruca albatrossiana (Pilsbry, 1912) (listed in Hoek (1913) and Buckeridge (1994)); Hexelasma arafurae Hoek, 1913 (listed in Hoek (1913) and Jones and Hosie (2016)); Solidobalanus au- ricoma (Hoek, 1913) (listed in Hoek (1913) and Jones et al. (2001)); Striatobalanus tenuis (Hoek, 1883) (listed in Hoek (1913) and Jones and Hosie (2016)).The remaining 30 species can, therefore, be considered as new records for the Kei Islands and Tanimbar Island. The present study and previous works on the barna- cles of the Kei Islands, Aru Island and Tanimbar Island, especially the works of Hoek (1883, 1907, 1913), Broch (1922, 1931-1932), Buckeridge (1994, 1997), Jones et al. (2001) and Jones and Hosie (2016), enrich our knowl- edge of the barnacle fauna of these islands. This study demonstrates once more the value of museum collections as a resource in biodiversity science. The result of this study also strengthens the state- ment of Hoeksema (2007) that the Indo-Malayan re- gion (which extends from East Indonesia to the Philip- pines and the Solomon Islands) is a centre of maximum marine biodiversity. Darwin (1854) demonstrated that this area had greater species richness than elsewhere in the world at the time. He named it the East Indian Archipelago (including the Philippines, Borneo, New Guinea, Sumatra, Java, Malacca and the eastern coast of India) and categorised it as his third province of bar- nacles. In this province, he found 37 barnacle species, the largest number known at that time, compared with the other provinces. Regarding the biodiversity of barnacles, the Indo-Ma- layan region as the centre of benthic biodiversity has not been replaced by other areas. In recent times, many studies and expeditions have been conducted in this area, revealing many more species of barnacles. For example, three expeditions have been undertaken within Philippine waters from 1976 until 1985 through MUSORSTOM Cruises and the collections the U.P. Marine Biological Laboratory at Puerto Galera, Oriental Mindoro (Rosell 1991; Chan 2009). Overall, the three of scientific cruis- es of MUSORSTOM collected 78 species of barnacles, 43 of which are new records and 12 species are new to science (Rosell 1991). Through the Philippine Panglao expedition (2005), Chan (2009) has also increased the number of barnacles from the Philippines, reporting 20 barnacle species with two new to science. Similar to the Philippine waters, eastern Indonesian waters also have a high diversity of barnacles. Recently, it has been revealed that the Moluccan Islands in east- ern Indonesia have 97 species of barnacles, 23 of which are new records and two species are still awaiting their species descriptions (Pitriana et al. 2020). Furthermore, this number will increase with the results of the study of the barnacles from Karubar expedition (1991) that have revealed 40 species of barnacles. zse.pensoft.net Pitriana, P. et al.: Deep-sea barnacles from the Karubar expedition (1991) The results of the studies of barnacles from the Phil- ippines and eastern Indonesian waters reconfirm the In- do-Malayan region as the epicentre of marine biodiversity. Acknowledgements The first author would like to thank Prof. Frank Riedel from the Freie Universitat Berlin for his general support and Bani Wahyudi and Abdul Basit for their support and comments on picture editing. Thanks to Bernhard Schuri- an for supporting the morphological work at the Museum ftir Naturkunde Berlin. We would like to thank Dr. Martin Husemann, Prof. Benny Kwok Kan Chan and anonymous reviewer for valuable comments on the manuscript. The first author would also like to thank Prof. Keiji Baba, Dr. Enrique Macpherson, Ferran Palero, PhD, Pau- la Martin-Lefévre and Sébastien Soubzmaigne for the great time working in the Crustacean Laboratory of the Muséum National d’Histoire Naturelle (MNHN), Paris and to Thomas von Rintelen (MfN Berlin) for providing funds for the research visit to Paris. The second author would like to acknowledge Dr Alain Crosnier (retired, MNHN) for his generous support to enable her study of the KARUBAR cirripede collec- tion in Paris. This study was funded by the Ministry of Research, Technology and Higher Education, the Republic of Indo- nesia within the Program for Research and Innovation in Science and Technology (RISET-Pro), World Bank Loan No. 8245-ID. 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Pacific Science 18: 355-372. zse.pensoft.net Pitriana, P et al.: Deep-sea barnacles from the Karubar expedition (1991) Supplementary material | Tables S1—S39 Authors: Pipit Pitriana, Diana S. Jones, Laure Corbari, Kristina von Rintelen Data type: Checklist table of species and morphometric for each species Explanation note: Table S1. Checklist of barnacle spe- cies collected during the Karubar expedition (1991) and deposited in MNHN, Paris. Tables S2—39. Mor- phometric/measurements for each species. Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons. org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow us- ers to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/zse.96.55733.suppl1