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

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 is 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.


Introduction
In 1991, scientists from France and Indonesia conducted collaborative research through the Karubar expedition in Indonesia. The acronym for this expedition, which collected 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 . Mortensen suggested 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), Paris and the Institut de Recherche pour le Développement (IRD) (formerly ORSTOM), to explore the deep-sea fauna 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 concern crustaceans, especially crabs, lobsters and shrimps (Richer de Forges et al. 2013). 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.
Diagnosis. Capitulum broadly oval, with five smooth plates; carina not extending to area between terga; peduncle with circles of small protuberances; cirri short.
Distribution. Indo-west Pacific: Indian Ocean; Madagascar through Malaysia, Hong Kong, South China Sea; Taiwan; 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, Dianajonesia amygdalum was found at Tanimbar Island, Indonesia.
Description. Capitulum oval, apex pointed, slightly thick, swollen. Scutum with larger segment strongly bowed, basal margin short, apex pointed; smaller segment bowed, terminating in point at base, tergal margin rounded, fitting exactly 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-segmented); cirri II-VI longer, more slender; cirrus VI with caudal appendages. Penis thick, ringed, especially mid-length, terminating in narrower, curved part. Maxillule notched, two large teeth on upper side; mandible with four teeth, largedistance 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.
Description. Capitulum white, with five calcified plates, surfaces strongly striated. Scutum with basal margin rotated; 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 separated by wide notch; mandible with four teeth, lower angle sharp. Measurements of specimen: basal diameter of capitulum 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, Indonesia, 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, Megalasma striatum was found at Kei Islands and Tanimbar Island, Indonesia.
Diagnosis. Formerly, the subfamily was characterised by a subapical carinal umbo, inflexed carina and subapical 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 between the rostrum and rostrolatus extends over the entire height of both plates.
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 quadrangular, angle at apex between scutal and tergal margins distinctly projecting over scutum. Rostrum small, triangular; 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 angle. Measurements of two specimens: height of capitulum 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.
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 morphology. 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 development (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.  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.

Alcockianum persona
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 remaining 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-segments); cirri II-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 margin, 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 Distribution. Indonesian Seas, eastern Australia, New Zealand; 109-915 m depth (Jones 1992). In this study, Alcockianum persona was found at Tanimbar Island, Indonesia.
Distribution. Eastern Indian Ocean; Northwest and Western Central Pacific; Malay Archipelago; Japan; Taiwan; Indonesia; attached to shell of gastropod, gorgonians, rocks; 805-6,810 m depth (Jones and Hosie 2016). In this study, Annandaleum japonicum was found at Tanimbar Island, Indonesia.

Scalpellum molliculum
Description. Capitulum compressed; 13 plates completely covered by fine, hairless membrane. Peduncle half length of capitulum, stout, cylindrical, armed with small, transversely elongated plates. Scutum subtriangular, lateral margin excavated with tooth above excavation blunt, short, simple; tergum almost triangular, scutal margin excavated, 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 II-VI slender, long, rami almost equal lengths; cirrus VI with long, slender caudal appendages. Maxillule slightly notched, two major setae on upper side; mandible with four teeth. Measurements 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. Distribution. Gulf of Oman, Arabian Sea, Sri Lanka, Japan (Chan et al. 2009b). In this study, Annandaleum laccadivicum was found at Tanimbar Island, Indonesia.
Distribution. Southeast Pacific Ocean (Newman and Ross 1971). In this study, Litoscalpellum walleni was found at Tanimbar Island, Indonesia.
Desciption. Capitulum yellowish, with 13 fully calcified plates. Peduncles short with scales slightly overlapping in the middle part. Scutum with pit for complemental 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 capitulum 12.44-13.88 mm, width 6.97-7.52 mm, thickness 2.77-3.47 mm; length of peduncle 2.99-3.15 mm, width 2.94-3.24 mm.
Description. Capitulum yellowish, elongate-oval shape; surface with distinct lines of growth. Carina large, simply bowed. Scutum with umbo at apex, slightly recurved, projecting slightly over tergum; tergum triangular, stout,  broad, apex recurved, scutal margin almost straight. Upper latus quadrangular, apex slightly projecting over scutum. 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 slender setae on cutting edge; mandible with three teeth excluding 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 (Shalaeva and Boxshall 2014). In this study, Amigdoscalpellum tenue was found at Kei Islands, Indonesia.
Description. Capitulum long, narrow, sparsely covered with hairs, plates separated by narrow, chitinous interspaces, 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, carinal margin concave. Scutum with umbo apical, overlapping occludent margin of tergum. Upper latus triangular; 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. Measurements 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. Java Sea, Indonesia; Philippines (Chan 2009). In this study, Teloscalpellum ecaudatum was found at Kei Islands and Tanimbar Island, Indonesia.
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.

Scalpellum imperfectum
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, elongated; 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. Measurements 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.

Diagnosis.
Shell not depressed; carina and rostrum interlocking with single rib from each plate; movable plates large, scutum with four articular ribs, tergum with six articular 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 axial articular ridge. Carina and rostrum irregular quadrangular, with carina higher, rostrum broader. Fixed tergum with two parts: (1) triangular portion very narrow at apex, slightly broader in its inferior (2) flat and broad part at a rear portion 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 appendages. Maxilla bilobed, fringed with setae, except on the notch; maxillule widely notched, horizontally elongated, 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. (Chan et al. 2010). In this study, Altiverruca navicula was found at Tanimbar Island, Indonesia.

Description.
Movable plates parallel to base, wall of parietal vertically ribbed; fixed scutum without internal pit. Movable scutum with crescentic ridge and longitudinal striations; movable tergum with articular ribs and diagonal rib. Apices of fixed scutum and tergum contiguous. Carina occupying carino-rostral wall, apices marginal. Cirrus I with rami unequal and serrulate setae; cirrus VI with caudal appendages. Maxilla globular, with fringing setae; maxillule notched, two large setae on upper side; mandible with three teeth excluding inferior angle.
Diagnosis. Shell with four or six plates; wall solid or permeated by single row of chitin-filled longitudinal canals; radii absent; one or both rami of cirri I and cirri II sometimes antenniform; labrum without notch in crest.
Description. Shell yellowish, conical, with six plates. Orifice 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 (anterior ramus: 8-segmented; posterior ramus: 12-segmented), dense long setae on surface areas. Cirrus II with equal rami, dense long setae. Cirri IV-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.
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. Radii with pale orange-brown and rust red-brown horizontal striation. Oper- 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 comparatively shorter, due to development of rostrum alone or more upright and comparatively higher, with neither carina nor rostrum 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, delicate 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 distinctly separated from fifth; distance between tips of first and second teeth slightly more than that between those of second and third teeth; third tooth larger; fourth and fifth smaller 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 Australia, Malay Arch.; China; Philippines; S Japan; Fiji Is; attached to coenosarc of gorgonians or antipatharians; littoral-453 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 gorgonian (Darwin 1854).  Diagnosis. Shell with parietes and basis not porose; carino-lateral compartments very narrow, almost same width from top to bottom; radii with smooth sutural edges; scutum externally striated longitudinally.

Conopea navicula (Darwin, 1854)
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; parietal plates studded with calcareous points. Parietal plates pearly white, solid, superficially appearing to possess longitudinal tubes, growth lines horizontal. Alae moderately developed. Basis calcareous. Size small. Rostrum well developed, concave, lying at angle of ~ 45°. Laterals very well developed. Carino lateral parietes thin, radii 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; posterior ramus: 9-segmented). Cirri III-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 numerous large setae. Mandibles with five teeth and inferior angle. Measurements of four specimens: basal length of 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.

MNHN-IU
Description. Shell with plates ribbed longitudinally. Shell colour brownish-pink to dull rose-pink, ribs tending to white, colour often faded with specimens appearing 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, especially distally. Cirri I-II with rami slightly unequal, covered with setae; cirri III-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.
Diagnosis. Shell with smooth, glossy white plates, coloured stripes absent; internal plates with thick, solid, finely ribbed longitudinally; base non-porous, radially ribbed.
Description. Shell plates white, stripes absent. Several specimens with pale pink tinge, one with pale brownish-pink parietes with small, narrow ellipsoidal whitish spots, latter orientated longitudinally producing reticulated effect. Radii whitish, pink tinge along distal borders. Scutal growth lines without longitudinal striations; articular ridge absent; pit for adductor muscle small, round. Tergum with shallow, wide furrow running from apex to base. Cirrus I with unequal (anterior ramus: 7-segmented; posterior ramus: 15-segmented). Cirrus II with rami subequal (anterior ramus: 11-segmented; posterior ramus: 12-segmented). Cirri I and II with very dense, long setae on surface areas. Cirri III-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 setae 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, orifice width 3.24-6.33 mm, basal width 5.66-9.56 mm.

Remarks.
In the type description, Broch (1931Broch ( -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 orientated longitudinally, thus producing a reticulated effect. Radii whitish with pink tinge along distal borders.
Striatobalanus amaryllis. -Jones, 2004: 150. -Chan et al. 2009b   Description. Shell conical; tips of rostrum and carina 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, tergum and soft parts. Measurements of specimen: basal length of shell 16.76 mm, orifice length 9.13 mm, carinal height 9.89 mm, orifice width 7.24 mm, basal width 15.14 mm.

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 tubiferous; scutum with conspicuous adductor ridge; tergum with well-developed depressor muscle crests, growth lines in tergum spur display an obvious change in direction; second maxilla with smooth anterior margin of distal lobe, acuminate setae with enlarged, modified tips.
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 anti-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 reference 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 Danish expedition to the Kei Islands (1922) and deposited in the Zoological Museum of Copenhagen University. In his report, only four species, Euscalpellum rostratum (Darwin, 1851), Lepas (Anatifa) anatifera Linnaeus, 1758, Conchoderma virgatum Spengler, 1789 and Acasta dentifer (Broch, 1922), were explicitly collected in the Kei Islands. 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, Makassar Strait, Tual, Banda Neira, Ambon and Saparua Bay.
The lists of Hoek (1913), Broch (1931Broch ( -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 40 species are now recorded from these Islands.
The present study and previous works on the barnacles of the Kei Islands, Aru Island and Tanimbar Island, especially the works of Hoek (1883Hoek ( , 1907Hoek ( , 1913, Broch (1922Broch ( , 1931Broch ( -1932, Buckeridge (1994Buckeridge ( , 1997, Jones et al. (2001) and Jones and Hosie (2016), enrich our knowledge 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 statement of Hoeksema (2007) that the Indo-Malayan region (which extends from East Indonesia to the Philippines 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 barnacles. 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-Malayan 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 cruises 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 eastern 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.
The results of the studies of barnacles from the Philippines and eastern Indonesian waters reconfirm the Indo-Malayan region as the epicentre of marine biodiversity.