The stalked crinoid fauna (Echinodermata)

of the Molucca and Celebes Seas, Indonesia:

taxonomic diversity and observations from
remotely operated vehicle imagery

Verena Tunnicliffe, Michel Roux, Marc

Eléaume & Dustin Schornagel

Marine Biodiversity

ISSN 1867-1616

Mar Biodiv
DOI 10.1007/s12526-015-0369-x

1 23
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 Author's personal copy
Mar Biodiv
DOI 10.1007/s12526-015-0369-x

ORIGINAL PAPER

The stalked crinoid fauna (Echinodermata) of the Molucca

and Celebes Seas, Indonesia: taxonomic diversity
and observations from remotely operated vehicle imagery
Verena Tunnicliffe 1 & Michel Roux 2 & Marc Eléaume 2 & Dustin Schornagel 3,4

Received: 25 November 2014 / Revised: 24 June 2015 / Accepted: 16 July 2015

# Senckenberg Gesellschaft für Naturforschung and Springer-Verlag Berlin Heidelberg 2015

Abstract The seafloor of the Molucca and Celebes Seas,
Indonesia, was investigated using a remotely operated vehicle
with high-definition video during 18 dives to depths of 280 to
3200 m. Irregular slopes and rocky substrata supported abun-
dant megabenthos, including stalked crinoids. Video zoomed
on many individuals provided enough details to assign 770
stalked crinoids to major taxa and many to species level.
These taxa included Guillecrinus neocaledonicus,
Ptilocrinus cf. amezianeae, three species of Hyocrinus, an
unknown phrynocrinid, Naumachocrinus hawaiiensis,
Proisocrinus ruberrimus, Endoxocrinus alternicirrus, seven
phenotypes in the Subfamily Metacrininae and one taxon of
unknown affiliation. A novel observation of a juvenile
Endoxocrinus included the attachment disk to a rocky surface.
Taxa usually considered rare were well represented on hard
substrata, thus expanding knowledge of their depth
distributions. Mostly, stalked crinoids were sparsely distribut-
ed in low densities (to 186 individuals per km), except for
occasional single-species clusters. Metacrinins and P. cf.
amezianeae dominated at depths of 250–600 m, while
E. alternicirrus and G. neocaledonicus were the most abun-
dant species at a depth range of 1000–1400 m. The largest
peak of species richness (with nine of the 17 taxa identified)
occurred at 1000 to 2000 m. Many specimens altered their
feeding posture with current intensity, but a balloon-like
crown posture is common in low or non-existent currents.
The postures of several mobile metacrinin individuals suggest
an active role of arms and cirri in slow crawling movements.
The possible origin of some taxa from either Gondwanaland
or Eurasian margins is discussed using biogeographical and
paleontological data. The location of the Wallace line cannot
be delimited only using extant stalked crinoid distribution.

Keywords Stalked crinoids . In situ imagery . Taxonomic

Communicated by J. Gutt
richness . Behaviour . Abundance . Depth ranges .
Electronic supplementary material The online version of this article Biogeography
(doi:10.1007/s12526-015-0369-x) contains supplementary material,
which is available to authorized users.

* Verena Tunnicliffe
Introduction
verenat@uvic.ca
The Molucca Sea and Celebes Sea of Indonesia lie in a highly
1
complex geological region. The Molucca Plate Collision Zone
Department of Biology and School of Earth & Ocean Sciences,
University of Victoria, Victoria, BC, Canada V8W 2Y2
is formed by tectonic plate subduction on two sides: the
2
Sangihe Volcanic Arc to the west and the Halmahera Arc to
Muséum national d’Histoire Naturelle, Département Milieux et
Peuplements Aquatiques, UMR 7208-BOREA
the east enclose a narrow sea (Widiwijayanti et al. 2004).
MNHN-CNRS-UPMC-IRD, CP26, 57 rue Cuvier, Thus, the seafloor has many abrupt topographic features from
75231 Paris, Cedex 05, France submerged limestone platforms to subsea volcanoes, to deep
3
Department of Biology, University of Victoria, Victoria, BC, Canada abyssal basins (Hall and Smyth 2008). The western edge of
V8W 2Y2 the Sangihe Arc has a wide bathymetric range with geological
4
Department of Biology, Memorial University, St. John’s, NL, Canada features that include hydrothermalism near Kawio Island
A1B 3X9 (McConachy et al. 2004). The shallow parts of this arc and
 Author's personal copy
Mar Biodiv

of the Talaud Ridge (Fig. 1) host deep-water reef structures,
but no prior submersible or remotely operated vehicle (ROV)
work has examined this region.
The INDEX-SATAL 2010 expedition (Indonesia
Exploration of the Sangihe Talaud Region) was a joint initia-
tive of the U.S. Ocean Exploration Program in the National
Oceanic and Atmospheric Administration (NOAA) and of
several ministries of the Indonesian government. The explo-
ration included operations by the NOAA Ship Okeanos
Explorer to produce a multibeam bathymetric map and to
survey selected sites with an ROV in the Molucca and
Celebes Seas. The mission consisted of two legs and executed
a total of 27 dives (for access to cruise track and selected
imagery, see oceanexplorer.noaa.gov/okeanos/explorations/
explorations.html), many of which encountered a diverse
deep-sea fauna that included stalked crinoids. While the
ROV had no collection capability, we used the high resolution
camera to examine the fauna within the study region.
The deep-sea fauna of the intertropical western Pacific was
first investigated between 1875 and 1910 by the Challenger,
Siboga and Albatross cruises. In 1899, the Dutch Siboga
Expedition explored the waters of the Celebes and Banda
Seas with a focus on deep water. The expedition lead, Max
CM Weber, was also interested in testing the nature of the
Wallace Line that extended through these seas; he decided that
the gradual faunal transitions on land and in water did not
support an abrupt Bline^ (Pieters and de Visser 1993). The
many publications from these expeditions describing dredged
fauna provide comprehensive descriptions of benthic animals,
including a diverse stalked crinoid fauna (Carpenter 1884;
Döderlein 1907; Clark 1908; 1910, 1911, 1915; see also
Améziane-Cominardi 1991). Since 1976, new deep-water col-
lections of the French program MUSORSTOM in the region
between the Philippines and New Caledonia have supported
descriptions of new stalked crinoid taxa and studies of
ecophenotypic variations (Roux 1981; Bourseau and Roux

Fig. 1 ROV BLittle Hercules^

dive locations during the INDEX
Ocean Exploration cruise in
summer 2010. Only dives that
located more than ten stalked
crinoids are shown and labelled
with Leg-Dive number.
Multibeam bathymetry was
produced during the cruise by the
National Oceanic and
Atmospheric Administration
(NOAA), courtesy of the Office
of Ocean Exploration and
Research
 Author's personal copy
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1989; Améziane-Cominardi et al. 1990; Améziane-Cominardi
1991; Bourseau et al. 1991; Améziane 1997; Eléaume et al.
2007). In 1989, the CALSUB cruise off New Caledonia using
the submersible Cyana provided the first in situ observations
of this deep-sea crinoid community, with specimens collected
to confirm taxonomic attributions (Lambert and Roux 1991;
Roux 1994).
Macurda and Meyer (1974, 1976) first observed stalked
crinoids in situ using a submersible off Jamaica and compiled
photographs to produce the first key for identifying key taxa.
Subsequently, in situ observations using towed camera sys-
tems or submersibles documented stalked crinoids in variable
concentrations from 150 m to 9000 m (Conan et al. 1981;
Messing 1985, 2004; Messing et al. 1990; David et al. 2006;
Shortis et al. 2007; Oji et al. 2009; Bowden et al. 2010). In situ
observations can reveal the dispersion patterns of crinoids
(Roux 1985; Tyler and Zibrowius 1992) and responses to local
hydrodynamic conditions. Stalked crinoid depth distributions
reflect water mass conditions (e.g., below storm waves or at
ocean fronts), food supply in the water column (low in abyssal
depths), and sea-level changes through geologic time (Roux
1987; Améziane and Roux 1997). Many stalked crinoids are
permanently attached to a rocky substratum by an encrusting
disk. Isocrinids have a distal flat articulation where the stem
can be autotomized; after autotomy, individuals can move,
crawling on sediments and climbing boulders using arm
movements (Messing et al. 1988; Birenheide and Motokawa
1994; Baumiller and Messing 2007).
The western tropical Pacific holds the world’s highest di-
versity of stalked crinoid species (Améziane and Roux 1997),
but, as most species inhabit hard substrata, remote sampling is
difficult (Roux 1994); study of diversity, niches, relative abun-
dances, behaviour and precise depth distributions requires in
situ investigations. Such information is useful in studies of
paleo-environments and paleobathymetry, especially for
post-Paleozoic crinoid limestones that frequently include fos-
sils with ecophenotypic characters similar to those observed in
extant fauna (Bourseau et al. 1988). Paleo-bathymetric esti-
mations from crinoids are useful for the study of sea-level
changes and bathymetric evolution of marginal seas
(Améziane-Cominardi 1991; Roux et al. 1988, 2006). As cri-
noid macro-evolution corresponds to plate tectonic history
and extant stalked taxa are usually holobenthic (Roux 1987),
their current distribution likely reflects origins from either the
Gondwana or Eurasia margins, especially for taxa presumed
to be the most ancient.
Many crinoids were encountered during the INDEX expe-
dition, the first opportunity to observe the deep-water fauna of
this important biogeographic region. This study was undertak-
en to document the taxonomic diversity of the stalked crinoids
over a wide depth range and in several environments with
ROV imagery. As few in situ observations are available on
many poorly known crinoids, we examined animal posture
and site characteristics to interpret likely feeding behaviour
and habitat requirements. We also assessed relative abun-
dances and taxon distribution with depth over the study area.
These data are set in the context of the regional biogeographic
pattern and history of stalked crinoids in the western Pacific.
Methods
Video imagery
The NOAA Ship Okeanos Explorer is equipped with high
speed Internet2 connectivity to stream live video and data to
shore through a VSat link to provide ‘telepresence’ for shore-
side scientists. This approach was tested for the first time on
this expedition using command centres in Jakarta and Seattle.
The ROV Little Hercules was equipped with an ultra-short
baseline navigation system, two 400 watt HMI lights and a
high-definition video camera. The small size of the ROV
allowed very close photography, and a professional videogra-
pher returned high quality imagery. Clips of highlights such as
close-ups were saved shipboard, while full dives were cap-
tured to mp4 video files from the satellite stream. At night,
the ship mapped the study region to return high-resolution
bathymetry (EM302; Fig. 1) that aided both dive site selection
and interpretation of the local topography.
We examined imagery from 17 dives spanning depths from
280 m to 3190 m (Table 1). Dives on bathyal sediments and at
a hydrothermal vent site recorded few to no crinoids. All re-
solvable stalked crinoids on a dive were located by cross-
referencing to navigation data. When possible, the ROV was
halted for close-up imagery video followed by framegrabs of
each animal to facilitate examination. Crinoids were first
grouped into types based on pinnule, arm and stalk character-
istics. Similar types from multiple dives were then compared
to refine taxon identification. Many animals were not possible
to identify, including possible juveniles, as some had too few
distinctive features recognizable in visuals alone, while for
others, imagery was too poor (lighting or distance problems).
Thus, abundances noted here are conservative estimates.
Taxonomic determination
We mainly adhere to the terminology and keys to stalked
crinoid genera in Roux et al. (2002). However, from imagery,
only clear external morphological characters are available.
Video close-ups of isocrinids and hyocrinids can resolve more
detailed characters such as the skeletal architecture, stalk at-
tachment, arm pattern, and filtration network with extended
podia. Small crinoids with ten arms and a xenomorphic stalk
cannot be identified to family level (e.g., either Bathycrinidae
or Septocrinidae). Among metacrinins, ecophenotypic con-
vergences and a wide range of variation in external
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Table 1 INDEX expedition dive

locations where the great majority Dive Region Dive site Launch position Start End
of stalked crinoids was seen. number depth (m) depth (m)
Listing is north to south; dive
number annotation is Leg - Dive # III - 08 Talaud Ridge East Pujada Ridge 5° 24.20′N 126° 35.40′ E 900 807
II - 06 Talaud Flank Paradise Valley 5° 22.64′N 126° 46.02′E 2700 2640
II - 07 Talaud Ridge Black Hole 5° 14.96′N 126° 39.38′E 1170 1200
II - 08 Talaud Ridge K1 5° 04.68′N 126 ° 39.13′E 1630 1513
III - 10 Talaud Ridge Zona Senja 4° 53.47′N 127° 0.90′E 311 280
III - 09 Talaud Ridge BJIV-1 4° 41.77′N 126° 50.81′E 1613 1513
II - 09 Molucca Sea The Eye 4° 56.93′N 125 ° 46.73′E 598 502
II - 04 North Sangihe Arc Bulan 4° 00.76′N 125° 16.80′E 1602 1611
II - 05 Mid Sangihe Arc Nuang Traverse 3° 46.73′N 125° 22.18′E 600 611
II - 11 Mid Sangihe Arc South Mount 2° 51.76′N 125° 02.06′E 490 461
III - 01 Mid Sangihe Arc Site K 2° 50.87′N 125° 03.93′E 632 537
II - 13 Mid Sangihe Arc Site K 2° 50.65′N 125° 03.50′E 466 576
III - 05 Mid Sangihe Arc Site K 2° 49.82′N 124° 58.87′E 3193 3113
III - 13 South Sangihe Arc Gelembung 2° 41.87′N 125° 17.05′E 845 706
III - 14 South Sangihe Arc Gelembung II 2° 41.19′N 125° 15.47′E 1451 1116
II - 12 South Sangihe Arc Site J 2° 37.48′N 125° 05.180′E 1170 1067
III - 04* South Sangihe Arc Site G 2° 16.14′N 124° 49.062′E 2105 1961
III - 03 South Sangihe Arc Site G 2° 15.99′N 124° 50.12′E 1928 1790

*video capture not successful for this dive, but one different crinoid type was documented in a short clip

morphology are very frequent (Améziane-Cominardi 1991).
Therefore, metacrinin identifications correspond to descrip-
tions of ecophenotypes rather than true species. Robust iden-
tification of hyocrinid genera requires detailed characters of
tegmen and genital pinnules. A few close-up views show the
main aspect of tegmen morphology, but genital pinnule archi-
tecture is not accessible from in situ views.
It is difficult to use colour as a taxonomic character. Some
metacrinin species have variable colours (from beige to dark
green) and, within a single specimen, the colour of regenerat-
ed arms differs from older, complete arms. Nevertheless, for
practical reasons, we distinguish different phenotypes using
their colour without prejudging their taxonomic status (varie-
ty, subspecies or species). As specimens were not collected,
we compared our imagery to published studies that have sam-
pled specimens associated with in situ observations, and pro-
pose identifications with relevant references. For Guillecrinus,
Proisocrinus, Porphyrocrinus and Naumachocrinus, compar-
isons to additional unpublished original videotapes and pho-
tographs taken in 1989 during the CALSUB cruise off New
Caledonia were useful. We also examined photographs taken
using a towed video system (Shortis et al. 2007) during cruises
off S E A ustralia and Tasmania, and p ictures of
Porphyrocrinus from the Deep Downunder Expedition off
Queensland.
Studies on crinoid molecular phylogeny (Hemery 2011;
Hemery et al. 2013; Rouse et al. 2013) reveal the need for a
deep reappraisal of extant crinoid classification (Roux et al.
2013); for practical reasons, we use some clades sensu lato
(e.g., Bathycrinidae sensu Gislén 1938, Isocrinidae sensu
Rasmussen 1978) without prejudging their validity. We also
use the term comatulid referring only to the phenotype of
unstalked crinoids attached by cirri.
Abundances
For each site, the relative abundance of each stalked crinoid
taxon was derived from total observations. Seven dives were
selected to examine crinoid abundances, as they were mostly
hard substratum and it was possible to control ROV behaviour
for transecting, although, as observation tasks were performed
for other studies, unbroken transecting was not possible. The
ROV worked in three modes: high fast flight, steady flight
averaging about 2.3 m above bottom (=transect mode), and
slow/stopped with camera zoomed. To estimate crinoid abun-
dances, we used only the transect mode when observation
segments were sustained for more than five minutes. For each
segment, the distance travelled was determined from the x/y
UTM navigation coordinates of the ROV track measured
manually in ArcGIS©. Field of view was estimated from three
video segments with clear laser spots (10 cm separation) on
each dive; area observed in stop frames is about 2.3 m2.
Several factors affected the accuracy of the numbers present-
ed: transponder locations have error, ROV height was variable
and the camera view was not always consistent. We include
only those stalked crinoids that were identifiable. For these
seven dives, we also noted the presence of comatulids; the
diversity of form, colour, arms and size indicated many taxa
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of feather stars were likely present, but it was not possible to
categorize the types with confidence. Thus, all these
comatulids are grouped together.
Results
Crinoid types
We observed over 770 individual stalked crinoids that could
be assigned to one of eight families or subfamilies noted be-
low; about 80 additional animals were not classified due to
poor imagery or lack of distinguishing features.
Family Guillecrinidae
Guillecrinus neocaledonicus Bourseau et al. 1991.
Number observed: 139 individuals on four dives; half the
individuals on dive III-14.
Characters (Fig. 2): Stalk and aboral sides of arms white,
beige or pink-violet white; pinnules and adoral arm face usu-
ally darker, red violet to brown. Aboral cup not clearly differ-
entiated. Five long Bpalm-frond^ arms, sometime nearly as
long as the stalk; arm length 20–25 cm; ratio of pinnule to
arm length < 0.2; high pinnule density with very flexible pin-
nules of nearly the same length. Homeomorphic stalk with
cylindrical columnals and low flexibility (highly flexible
proxistele absent); attachment on hard substratum by
encrusting disk; stalk length 25–40 cm.
Environment: Depth 1244 to 1892 m, probably down to
2000 m, on slopes with blocks (Fig. 2a, c) and hard grounds
with some sediment (Fig. 2b, d); up to seven specimens per
photograph.
Remarks: Other locations include: New Caledonia, 1130 to
1312 m (possibly down to 1520 m) (data in Améziane and
Roux 2005; in situ views in Lambert and Roux 1991; Roux
1994); Vailulu’u Seamount, East Samoa (Staudigel et al. 2006
Fig. 3k). G. reunionensis Roux 1985 from 1775 (possibly
1340 m) to 1880 m (possibly 1980 m) off Réunion Island
and Sunda Strait (1560–1610 m) (Mironov and Sorokina
1998) may correspond to large older specimens of
G. neacaledonicus after Améziane and Roux (2005).
Family Hyocrinidae
Stalk homeomorphic with flexible proximal part; cross sec-
tion circular; attached on hard substratum by encrusting disk.
Aboral cup clearly differentiated consisting of large tall radials
atop low conical basal ring. Usually five arms with long pin-
nules; ratio of pinnule to arm length > 0.4; pinnule density
variable; distal part of arms and pinnules usually curled ab-
orally. Usually bright yellow.
Ptilocrinus cf. amezianeae Eléaume et al. 2012
Number observed: 105 individuals on six dives, with a
majority seen on dive III-1.

Fig. 2 Views of Guillecrinus

neocaledonicus illustrating arm
position in variable currents. a.
Weak current on rocky terrain;
depth 1598 m. b. Weak current
but rippled sediment indicates at
least intermittent strong benthic
current. Tall stalk about 40 cm
high; depth 1787 m. c. Substantial
current upwelling; depth 1240 m.
d. Current from right to left. Stalk
lower left about 35 cm long; depth
1419 m. NOTE: for all figures,
approximate sizes are given when
video included the 10 cm scale
nearby
 Author's personal copy
Fig. 3 Ptilocrinus cf. amezianeae in feeding posture and weak current. a.
General view with current from right to left. Crown 18 cm across; depth
473 m. b, c. Oral views of same specimen with close view of tegmen
showing food grooves converging to mouth, one myzostomid cyst (m)
Characters (Fig. 3): Well-developed filtration fan with
high pinnule density (Fig. 3a, b), 40–60 flexible pinnules per
arm side; arm length 20–25 cm, longest pinnules in mid arm,
ratio of maximum pinnule length to arm length ~0.5,
proximalmost pinnule close to arm base (probably on Br4).
Well-developed aboral cup, radials as wide as high, ratio arm
base to upper radial width < 0.5 (Fig. 3d). Tegmen inflated
with large inter-radial plates, small inconspicuous orals, and
conspicuous anal cone in inner position at top of tegmen, and
substantially higher than mouth (Fig. 3c). Stalk length to
60 cm; proxistele flexible, other parts usually rigid or slightly
curved.
Environment: Depth 422–619 m (may be down to 1871 m),
on slopes with coarse sediment pebbles and talus blocks.
Remarks: The large, conspicuous anal cone at the top of the
tegmen and the small orals place the specimens observed here
closer to Ptilocrinus than to Feracrinus. Although similar to
P. amezianeae, this is likely a new species. P. amezianeae
differs from these specimens in having a more inflated teg-
men, less flexible pinnules, the first pinnule on Br5 and a
larger size. Several specimens of Ptilocrinus cf. amezianeae
have broken or regenerating arms, and are infested by cyst-
forming myzostomids (Fig. 3c, e). P. amezianeae was collect-
ed and observed over a similar depth range, unusually shallow
for hyocrinids, in the Southern Ocean (chiefly 450–600 m, but
as deep as 1680 m) (Eléaume et al. 2012, 2014). Depths of two
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and conspicuous anal cone in SW interradius; depth 460 m. d. Profile
view of another specimen showing the conspicuous aboral cup (arrow
head); depth 465 m. e. Same specimen as d, showing infestation by
myzostomids forming numerous cysts (smooth bumps) on arms
deeper specimens in our study (1570 m and 1871 m) were
similar to the depth range on Tasmanian seamounts of
Feracrinus koslowi Améziane and Roux 2011, a species that
has a similar posture. Unfortunately, assignment of the ob-
served specimens to either Feracrinus or Ptilocrinus was not
possible without close views of the tegmen.
Genus Hyocrinus
Twelve Hyocrinus-like specimens spanning depths from
521 to 1522 m were encountered, but imagery was adequate
to differentiate only five individuals to species level. Usually,
this genus occurs at depths > 2500 m.
Hyocrinus sp. 1
Number Observed: Three individuals on dive III-3.
Characters (Fig. 4): 6–7 pinnules per arm side (Fig. 4a);
pinnules relatively rigid especially in proximal half (genital
inflation); interval between pinnules irregular (probably bra-
chial pairs and triplets); wide interval between tegmen and
first pinnule (probably first pinnule on Br6); ratio of maxi-
mum pinnule length to arm length < 0.5. Tegmen not inflated
(just a few tegminal plates); orals large and triangular; anal
cone small but conspicuous (Fig. 4b). Arm length ~6 cm, and
stalk length ~20 cm.
Environment: Depth (?1242) 1871–1901 m, on blocks
emerging from sediment.
Remarks: The characters are similar to those of
H. bethellianus Thomson, 1876, the type species of
 Author's personal copy
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Fig. 4 Hyocrinus sp. 1. Two
views of same individual showing
filtering posture in weak current;
depth 1901 m. a. Red laser dots
are 10 cm apart. b. Large orals
slightly open; pink isopod below
crown
Hyocrinus. The single species known in the intertropical west
Pacific is H. cyanae Bourseau et al. 1991. H. foelli Roux and
Pawson, 1999 from the north and east-central Pacific (Roux
2004) displays the same feeding posture, but differs in having
smaller orals, a larger anal cone and first pinnule closer to arm
base.
Hyocrinus sp. 2
Number Observed: One on dive II-08.
Characters (Fig. 5): Arms strongly curved aborally in a
singular posture (Fig. 5a); ~7 pinnules per arm side, first pin-
nule on Br5 or Br6, interval between pinnules regular (brachi-
al pairs). Short rounded anal cone adjacent to oral cone; orals
triangular and smaller than in sp. 1; tegminal plates well de-
veloped (Fig. 5b).
Environment: Depth 1538 m on rocky sea floor.
Remarks: Hyocrinus sp. 2 differs from Hyocrinus sp.1 in its
apparently smaller orals and a greater development of
tegminal plates. However, the oral cone is wide open and oral
size is difficult to estimate; open orals suggest active feeding.
The size estimate is 15–20 cm high so it is unlikely to be an
older individual of Hyocrinus sp. 1.
Hyocrinus cyanae Bourseau et al. 1991
Number Observed: One individual on dive III-5.
Characters (Fig. 6): 12–13 flexible pinnules per arm side;
relatively short regular interval between pinnules (probably
brachial pairs); wide interval between tegmen and first pinnule
(probably first pinnule on Br6); ratio of maximum pinnule
Fig. 5 Hyocrinus sp. 2; depth
1538 m. a. Unusual posture with
strongly recurved arms in
negligible current. No scale;
crinoid height estimated at 15–
20 cm. b. Short anal cone lies
adjacent to wide open orals
length to arm length > 0.5. Orals of moderate or small size;
inconspicuous anal cone. Arm length ~8 cm, and stalk length
~20 cm.
Environment: Depth ~3150 m, on block emerging from
sediments.
Remarks: A single specimen (the holotype) of this species
was observed and collected during the CALSUB cruise off
New Caledonia attached on a rocky slab at 2536 m
(Bourseau et al. 1991; Roux 2004). H. cyanae differs from
Hyocrinus sp. 1 and 2 in having a greater number of pinnules
per arm side, successive brachials joined as brachial pairs, and
smaller orals (Fig. 6b).
Family Phrynocrinidae
Unidentified phrynocrinid
Number Observed: One individual on dive III-9.
Characters (Fig. 7a to d): Yellow-green stalk of 40–50 cm
with flexible synarthries in proximal stalk (25 cm below ab-
oral cup); proximal mesistele columnals about as wide as
high, each widest proximally (Fig. 7c); distal columnals wider
than high and more cylindrical (Fig. 7d); attachment on rock
not visible; tegmen, pinnules and food grooves pink-violet to
reddish; stalk, cup and proximal aboral arm face yellow-
green; mid and distal arms probably the same (not visible);
tegmen very inflated, separating adjacent IBr series of proba-
bly > 6 brachials; arms divided rapidly just above tegmen (up
to 20 main branches or more), again at irregular intervals in
mid or distal arm, and frequently forming a small fork at arm
 Author's personal copy
Fig. 6 Hyocrinus cyanae. One
individual in feeding posture;
depth ~3150 m. a. 12–13 pinnules
per arm. Crown about 12 cm
across. b. Orals open wide
end; about one-third of crown regenerating; maximum arm
length about 20 cm; pinnulation dense, short arm tips devoid
of pinnules. Details can be viewed on the video in Online
Resource 1.
Environment: Depth 1565 m, slope with rough rocky sub-
stratum; one isolated specimen displaying a filtration fan.
Remarks: A very similar specimen was photographed off
SE Australia at 1512 m depth (courtesy of A. Williams and F.
Althaus, CSIRO). Xenomophic stalk, arm branching and very
inflated tegmen separating IBr series suggest a new species
belonging to a genus of Phrynocrinidae. We cannot exclude an
attribution to Phrynocrinus, which is considered monospecific
Fig. 7 a-d. Phrynocrinid n. gen., n. sp.; one individual only; depth
1565 m. a. In filtering posture on rocky slope. Crown approximately
40 cm across. b. Crown partly regenerating; arms up to 20 cm long
Mar Biodiv
(P. nudus AH Clark, 1907). P. nudus differs in having more
robust arms, longer pinnules and proximal mesistele with
columnals substantially wider than high. P. nudus is known
from Japan at depths from 700 to 2000 m (Oji and Kitazawa
2006), and between SE Australia and New Zealand at depths
of 980 to 1450 m (McKnight 1979; Donovan and Pawson
1994, Améziane and Roux 2001, Améziane and Roux in
prep.). It is not known between these two areas.
Porphyrocrinus aff. verrucosus Gislén 1925
Number observed: three or four individuals on dive II-09.
Characters: (Fig. 7e) Pale brown xenomorphic stalk with
proximal part more or less white, attached by encrusting disk,
with multiple divisions. c. Detail of the proximal xenomorphic stalk. d.
Detail of the distal stalk. e. Porphyrocrinus aff. verrucosus at 502 m
depth. Red dots to left are 10 cm apart
 Author's personal copy
Mar Biodiv
crown brown orange, five arms with relatively short pinnules,
proximalmost arm without pinnule (probably first pinnule at
Br8). Imagery was poor.
Environment: Depth 500–505 m, rock slabs partly covered
by sediment.
Remarks: Porphyrocrinus verrucosus (included
P. polyarthra AM Clark 1973) was collected at depths of
218 to 400 m in western tropical Pacific (off Palau and the
Indonesian Kai Islands) and eastern Indian Ocean (Gislén
1925, AM Clark 1973; Messing 2007). Porphyrocrinus aff.
verrucosus was photographed at similar depths off
Queensland (courtesy of G. Woerheide), off Tasmania (cour-
tesy of F. McEnnulty and N. Améziane), and down to 616 m
off New Caledonia (Bourseau et al. 1991) .
Family Bathycrinidae (sensu lato)
Naumachocrinus hawaiiensis AH Clark, 1912
Number observed: 37 individuals on several dives.
Characters (Fig. 8): Crown relatively small with respect to
the long stalk; ten arms (one case with 12) proximally divided
at the second primibrachial (IBr2ax), usually with 11–14 pin-
nules per arm side in large specimens (Fig. 8a, d), but three
specimens have 18, 20 and 25 pinnules/arm side; pinnules
with fringe of numerous podia; tips of lateral podia of adjacent
pinnules separated by distinct gap; distal podia extend beyond
Fig. 8 Naumachocrinus hawaiiensis. a. Filtering posture in substantial
current. Usually ten arms with 11–15 pinnules per arm. Crown 8–9 cm
across; depth 1125 m. b. Oral view of same individual with open mouth
and conspicuous anal cone. c. Detail of pinnules with dense podia. d.
pinnule tips (Fig. 8b, c); tegmen extending to IIBr2, conspic-
uous finger-like anal tube substantially higher than oral cone
(Fig. 8b); very high radials (up to five times as long as wide)
and low (sometimes inconspicuous) basals (Fig. 8d); crown
frequently regenerated from basal ring likely after predator
attack; xenomorphic stalk with stout dististele of 7–10
columnals, diameter of following 4–6 columnals rapidly de-
creasing; distalmost columnals with strongly narrow elliptical
articulations; mid stalk columnals barrel-shaped, likely artic-
ulated by synarthries (Fig. 8e); holdfast encrusting hard
substratum.
Environment: Depth 521–1572 m, variable sea floor mor-
phology; in sedimentary environment attached on large peb-
bles, blocks, boulders or hard surfaces.
Remarks: Usually, the radial ring is three times longer than
wide, but in images studied here, the ratio of L/W may equal
five. Three large specimens with stalk length > 300 mm and
features similar to those observed here were collected off
Solomon Islands between 570 and 756 m (French cruise
Salomon 1, 2001) (cat. no. MNHN-IE-2013-10380).
N. hawaiiensis was previously known from Japan, south-
western Pacific and Hawaii Islands at depths of 760 to
1500 m (AH Clark 1912; McKnight 1989; Bourseau et al.
1991; Oji and Kitazawa 2008).
Profile view showing the tall radial ring (arrow). Crown about 12 cm
across; depth 1332 m. e. Distal stalk of specimen in a–c showing well-
differentiated dististele with large columnals wider than tall, and with
strongly oval articulations
 Author's personal copy
Bathycrinid-like specimens
Numerous isolated and grouped individuals with ten
arms, IBr2ax and a xenomorphic stalk attached on hard
substratum were observed at various depths down to
3150 m. Although general morphology and live posture
of Bathycrinidae (sensu stricto) and Septocrinidae are fre-
quently similar (Mironov 2000), they may be distinguished
by the larger size of the primibrachials in Bathycrinidae.
Unfortunately, imagery never yielded views adequate to
distinguish these groups.
Family Proisocrinidae
Proisocrinus ruberrimus AH Clark, 1910
Fig. 9 Proisocrinus ruberrimus. a. Profile view in substantial current;
depth 1608 m. b. Detail on same individual of distinctive aboral cup and
rudimentary cirri on proximal stalk (arrow). c. Distal stalk (of individual
in e, f) with unusual five-fingered basal attachment encrusting rock.
Attachment 2.5 cm across. d. Profile view in weak current. Stalk about
80 cm long; depth 1787 m. e. Oral view of crown with arms open and
Mar Biodiv
Number observed: 48 individuals, nearly all on dive III-3.
Characters (Fig. 9): Colour brilliant light red; 13–20 arms
(usually 16–17) divided at crown base (brachitaxes of two
brachials), one case of ten arms on the youngest specimen
observed; short arm tip without pinnule moderately developed
(Fig. 9a, b), proximal pinnules not covering the tegmen; con-
spicuous aboral cup with cylindrical basal ring and widely
inverted conical radial ring, tegmen swollen interradially;
proximal stalk with rudimentary cirri (Fig. 9b arrow) and pen-
tagonal cross section; mid-distal stalk homeomorphic with
cylindrical cross section; usually attached on hard substratum
by attachment disk (Fig. 9d), sometimes with holdfast of five
pinnules closed (crown about 30 cm across). f. Multiple Proisocrinus in
heterogeneous terrain, likely with local turbulence affecting individual
postures. Sediment-draped slope with large octocorals; arrow head
indicates one Guillecrinus; depth 1755 m. g Profile view in weak
vccurrent on steep slope on Kawio Barat; depth 2324 m
 Author's personal copy
Mar Biodiv
round finger-like extensions (Fig. 9c). Estimated maximum
arm length 15–17 cm and stalk length frequently > 60 cm.
Environment: Depth range 1565–2325 m. P. ruberrimus
inhabits various environments including rocky cliffs, slumped
blocks, and hard outcrops in ripple marked sediments, isolated
or in small clusters (Fig. 9d, f, g). The deepest record is from a
dive that we did not analyze otherwise. Setting and details can
be viewed on the video in Online Resource 2.
Remarks: P. ruberrimus is otherwise known from 960 to
1900 m in the western Pacific from Japan to New Caledonia
(AH Clark 1910; Bourseau et al. 1991; Roux 1994; Oji and
Katazawa 2008), and in the Central Pacific off Hawaii and
Tahiti (Roux 1980).
Family Isocrinidae (sensu lato) Subfamily Diplocrininae
Endoxocrinus (Diplocrinus) alternicirrus (Carpenter, 1882
Number observed: 154 individuals on many dives. Most
characters distinguishing the two subspecies as described by
David et al. (2006) are not accessible on in situ views.
Characters (Fig. 10): Colour white or pale pink (Fig. 10a,
b); stalk attached to rocks or pebbles using the distalmost
cirrus whorl as a terminal claw (Fig. 10c), never with distal
stalk curving onto the substratum; stalk usually short with
noditaxes of 4–10 internodals (usually 5–6); cross section
stellate to pentagonal; arm branching only at base (division
series of 1–2 brachials), number of arms 20 (young specimen)
to about 60 (largest specimens), arm tips short without pin-
nules; proximal pinnules covering the tegmen.
Juvenile characters (Fig. 11): Stalk with nodals bearing
five cirri but attached to rock by encrusting disk; stalk from
attachment disk to distalmost cirrus whorl consisting of ~22
cylindrical columnals with concave lateral surfaces suggesting
articulation by synarthries; other noditaxes of < 15 internodals
cylindrical or pentagonal to stellate proximally; distalmost
cirrus whorl smallest; ten arms with IBr2ax; primibrachials
united by synostosis or syzygy, neither transverse synarthry
at IBr1+2 nor sharp symmorphy at IIBr3+4 excluding an
attribution to Hypalocrinus.
Fig. 10 Endoxocrinus alternicirrus. a. Oral view of individual with
fewer arms (possibly phenotype alternicirrus). Crown about 12 cm
across; depth 1580 m. b. Individual in weak current with large number
Environment: Depth 392 to 1871 m on various substrata
usually blocky rubble and boulders. At depths > 900 m, some-
times associated on rocky substratum with Proisocrinus
ruberrimus. Remarks: E. alternicirrus is a common species
previously known from 364 to 1476 m in the western
Pacific from Tasmania to Japan, and Central Pacific off
Hawaii and Tahiti (review in David et al. 2006).
Family Isocrinidae (sensu lato), Subfamily Metacrininae
Overall, we observed 240 individuals on eight dives, but
most are not possible attributable to genus, as images are too
distant.
General characters: Heteromorphic stalk bearing cirri, al-
ways five cirri per whorl; several noditaxes of distal stalk lying
on substratum and attached by distal cirrus whorls; cirri along
erect portion of stalk; number of arms usually 40–60 (up to
80), arm divisions at various locations; usually 4–7
primibrachials, increasing to 7–11 in secondibrachitaxis, more
than 11 distally; distalmost arms without pinnules; arms usu-
ally flexed aborally into current with conspicuous apinnulate
tips more or less curved orally; proximal pinnules covering the
tegmen.
The two genera (Saracrinus and Metacrinus) are distin-
guished only by the number of primibrachials (mode at four
in Saracrinus and at seven in Metacrinus). Species are diffi-
cult to determine from external morphology alone, because of
high variability and the frequent morphological convergences
corresponding to ecophenotypes (Améziane-Cominardi
1991). In each genus, taxa (species or ecophenotype) differ
mainly by their stalk characters (number of internodals per
noditaxis and ornamentation). Hemery (2011), Hemery et al.
(2013) and Rouse et al. (2013) found Metacrinus and
Saracrinus to be paraphyletic using molecular techniques.
Results from the most extensive data set (Hemery et al.
2011) demonstrated that metacrinin species and genera must
be revised. While colour differences do not necessarily mean
different species, for practical reasons, we distinguish the
main phenotypes using both colour and external characters
of arms (possibly phenotype sibogae). Crown 25 cm across; depth 465 m.
c. Base of same crinoid with basal cirri grasping substratum
 Author's personal copy
Mar Biodiv

Fig. 11 Juvenile attributed to

Endoxocrinus alternicirrus; depth
1175 m. a. Attached juvenile with
crown in balloon-like posture;
estimated 5 cm across. b. Distal
stalk attached on rock by an
encrusting disk

when close images provide detail. Taxonomic attribution is Metacrinin phenotype 1

only suggested. However, short noditaxes (≤ 9 internodals)
exclude these metacrinins: Metacrinus rotundus Carpenter,
1885, M. musorstomae Roux 1981, M. interruptus
Carpenter, 1884, M. cyaneus, Saracrinus nobilis (Carpenter,
1884) (sensu stricto) and S. superbus (Carpenter, 1885). Fujita
et al. (1987) published the first in situ photographs of a
metacrinin (Metacrinus rotundus) taken off Japan at relatively
shallow depths of 100 to 160 m.
Number Observed: Six well-imaged specimens on two di-
ves (III-1 and III-10).
Characters (Fig. 12): Stalk pale green to pink violet, stel-
late in cross section; number of internodals per noditaxis 7–9;
columnals with conspicuous, more or less regular equatorial
ridge; proximal stalk with cirri oriented upward, quickly be-
coming much longer below crown (Fig. 12d); cirri in mid stalk
as long as 3–4 noditaxes and mostly oriented upward,

Fig. 12 Metacrinin phenotype 1

(possibly Saracrinus angulatus).
a. Young individual with arms
closed in very weak to quiet
current; depth 520 m. b. Oral
view of crown with pinnules
closed; depth 542 m. c. Large
individual in very weak to quiet
current (about 50 cm tall); depth
567 m. d. Same individual, detail
of proximal stalk and crown
showing proximal brachitaxes. e.
Same individual, detail of mid
stalk showing cirri pressed to
stem
 Author's personal copy
Mar Biodiv
crowded, and densely flattened against the stalk (Fig. 12a, e);
distal stalk flattened against rocky substratum (Fig. 12b, c),
total stalk length about 50 cm; crown beige of 45–63 arms or
more, robust and moderately serrated (Fig. 12b, d); arm length
about 20 cm, primibrachials 6 (two cases) or 7 (one case),
secundibrachials 5 to 7, tertibrachials 9 to 13 (mode 11),
IVBr 14 or more; arm division pattern isotomous; pinnule
length progressively decreasing distally; approximately distal
fourth of arm slender, with pinnules reduced or absent, but
without angle and not clearly differentiated morphologically.
Young specimen with ~20–25 arms (Fig. 12a).
Environment: Depth 280–605 m, on rocky substrata or
coarse sediment.
Remarks: Characters were examined mainly on one well-
imaged specimen. The very few 6–7 IBr series observed could
suggest a species belonging to the genus Metacrinus, with
other characters close to those of M. serratus Döderlein
1907. However, these specimens could be attributed to a spe-
cies of Saracrinus with variable number of IBr and displaying
other characters that suggest affinities with S. angulatus, a
robust species frequently collected in the NW Pacific from
Japan to Timor and NW Australia at depths from 188 to
545 m (Améziane-Cominardi 1991; Eléaume et al. 2007). It
usually exhibits a moderately long arm tip without pinnules
and a general aspect similar to that of our specimens (see
Améziane 1997, Fig. 12a).
Metacrinin phenotype 2
Number Observed: A single individual see on dive III-1.
Characters (Fig. 13): Stalk pale grey-green and crown
white-beige (Fig. 13a); arm division pattern isotomous
(Fig. 13b); pinnules gradually becoming shorter distally, so
that distal fifth to third of arm has pinnules reduced or absent;
mid and distal stalk cross section subpentagonal; columnals
smooth or with small knobs, 4–6 internodals per noditaxis
(Fig. 13c).
Environment: A single individual on a highly weathered
surface at 473 m depth with dense coverage by diverse com-
munity dominated by suspension-feeders. Many specimens of
Fig. 13 Metacrinin phenotype 2
(possibly Saracrinus moosai);
depth 473 m. a. In filtering
posture in weak current from left
to right. b. Same individual, oral
view of crown. Size is 22 cm
across tips of arms. c. Same
individual, detail of distal stalk
with cirri pressed to stem
this or phenotype 1 seen on dive II-9 between 500 and 600 m
depth and on II-5 (600 to 610 m) most frequently on steeply
sloping surfaces with corals and sponges.
Remarks: Phenotype 2 generally resembles phenotype 1,
but differs in having less robust stalk and crown, shorter
cirri, weak columnal ornamentation and number of
internodals per noditaxis <7. The last character typifies
Saracrinus moosai Améziane 1997, which is closely related
to S. angulatus, but differs mainly in stalk and arm articulation
characters. The two species were collected in the same
environment off Kai Islands at depths between 296 and
349 m. Améziane (1997) transferred three specimens de-
scribed as S. varians by Döderlein (1907) (Kai Islands,
Indonesia, depth 304 m, Siboga expedition) to S. moosai.
Metacrinin phenotype 3
Number Observed: Five individuals seen on dive III-13.
Characters (Fig. 14): Colour blue-green, sometimes darker
in proximal stalk and crown, and substantially lighter in pre-
sumed regenerated part of arms and cirri (Fig. 14b); arm main-
ly divided proximally (likely IBr4ax and IIBr7-9ax); about
35–40 arms, apinnulate arm tips relatively long (up to
10 cm), strongly differentiated and curved adorally at a con-
spicuous angle; maximum arm length 25 cm; stalk length >
60 cm, stalk smooth with pentalobate cross section; number of
internodals per noditaxis 7–9; small and large columnals al-
ternating along the stalk.
Environment: Depth 731–757 m on cliffs, rough and irreg-
ular rocky substratum.
Remarks: Characters suggest affinities with Saracrinus
varians. This latter species was interpreted as an
ecophenotype of S. nobilis more frequent in deepest biotopes
(Améziane 1997). S. nobilis (including phenotypes superbus
and varians) was collected from Japan to New Zealand at
depths from 221 to 1200 m (Améziane-Cominardi 1991;
Eléaume et al. 2007). Attribution of collected specimens to
S. varians was confirmed only from depths > 500–600 m by
Améziane (1997).
Metacrinin phenotype 4
 Author's personal copy
Mar Biodiv

Fig. 14 Metacrinin phenotype 3

(possibly Saracrinus varians). a.
Individual in weak upwelling
current. 50 cm tall; depth about
730 m. b. Second individual with
crown bent in weak current from
right to left. About 20 cm across;
depth 756 m

Number observed: 52 individuals on a single dive, III-14.
Characters (Fig. 15): Stalk green grey; crown beige to pale
green with pale dark yellow bands (Fig. 15c); stalk length
up to 30 cm, cross section pentagonal from proximal to
distal, columnals with mid sharp ridge, number of inter-
nodals per noditaxis 7–10 (mode 9), cirrus as long as 3–4
noditaxes (Fig. 15b); conspicuous globular basals as wide
as high; maximum arm length 15 cm, up to ~40 arms rel-
atively slender and serrated, primibrachials 6 or 7,
secundibrachials 5 to 10, tertibrachials 11 to 15, beyond
IIIBrax free distal arm with relatively short tip without
pinnules but frequently oriented subperpendicular to arm
axis in oral direction (included when arms closed). One
young specimen with ~15 arms.
Environment: Depth 1108–1190 m; on local topographic
mounds and cliffs such as a ledge (appears to be layered
volcaniclastics, Fig. 15a) with up to 15 specimens of various
sizes per view; stalk and crown oriented downward; no cur-
rent. It is the deepest metacrinin seen on this cruise. Setting
can be viewed in video in Online Resource 3.

Fig. 15 Metacrinin phenotype 4

(possibly Metacrinus nodosus). a.
Population attached on a bedded
ledge of volcaniclastics; crown-
downward posture is usual
(associated fauna includes
comatulids possibly belonging to
Thalassometridae, ophiuroids and
hydroids). Centered red dots are
10 cm apart; depth 1175 m. b.
Individual of medium size (stalk
length to 45 cm) at cliff base. c.
Same individual, detail of
proximal stalk and crown
showing proximal brachitaxes
 Author's personal copy
Mar Biodiv
Remarks: May be Metacrinus nodosus Carpenter, 1884,
but this species is poorly known. Two specimens were collect-
ed by the Challenger at a depth of 1152 m and one with crown
missing beyond Br1 at 856 to 884 m off the Philippines
(Bourseau and Roux 1989).
Metacrinin phenotype 5
Number observed: Two individuals on dives II-11 and II-
13.
Characters (Fig. 16): Stalk white-beige or pale green with
relatively long cirri (4–5 noditaxes) mostly oriented upward
(Fig. 16a), noditaxes short (probably about 7–9 internodals);
adoral crown pale orange with arm axis darker orally
(Fig. 16b), ~40 arms or more; disc flattened; long well differ-
entiated white arm tips, with pinnules reduced or absent, al-
ways angled adorally with crown in filtation posture and oral
surface facing down-current; crown orients mouth upward
with arms curved aborally, forming a balloon–like posture
under weak or no current, and with apinnulate arm tips
pointing outward (Fig. 16c).
Environment: Depth 461 to 470 m, rocky substratum.
Remarks: May be a new species of Metacrinus or
Saracrinus, the colour of the crown has not been observed
on collected metacrinin specimens. A similar phenotype with
orange crown and the same posture, but having stouter arms,
was photographed off Tasmania at a depth circa 900–1100 m
(courtesy of K. Gowlett-Holmes, CSIRO).
Metacrinin phenotypes 6 and 7
Number Observed: We assign six individuals from dives II-
10 and II-13 to these morphotypes but other crawling
metacrinins may also belong to these taxa.
Characters (Figs. 17 and 18): Crown white with other
characters like phenotype 5; arm division pattern endotomous;
stalk pale green grey, 7–9 internodals per noditaxis.
Phenotypes 6 and 7 differing only by columnal shape and
ornamentation.
Phenotype 6: distal noditaxes stellate in cross section with
mid columnal ridge thin and moderately developed (Fig. 17b);
Fig. 16 Metacrinin phenotype 5 with orange crown. a. Individual in weak current with crown in balloon-like posture. About 25 cm in height; depth
464 m. b. Detail on recurved arms. c. Second individual assuming open posture in moderate current from right to left; depth 467 m
mid stalk pentalobate in cross section and thicker mid
columnal ridge well developed (Fig. 17c); ratio of noditaxis
to cirri length about 0.25, stalk length ~50 cm.
Phenotype 7: mid and distal noditaxes pentagonal in cross
section; smaller and larger columnals alternating and smooth,
without ridge (Fig. 18b, c); ratio of noditaxis to cirri length
about 0.33.
Environment: Like phenotype 5, but crawling specimens
move on heterogenous sandy sediments with pebbles and
blocks (Fig. 19). Depth of phenotype 6 specimen, 282 m; of
phenotype 7, 465 m; of specimens indistinctly attributed to 6
or 7, both at 258 m.
Remarks: These two morphotypes display about the
same general morphology and colours, but have distinct
noditaxis characters that are visible only in close-up
views. The phenotype 5 differs in crown colour only,
but its noditaxis and other characters were not visible;
thus, affinities to known metacrinin species cannot be
discussed.
Insertae sedis
Purple Umbrella Stalked Crinoid
Number observed: One, and possibly a second young spec-
imen, on dive III-4.
Characters (Fig. 20): White stalk seems to be mostly ho-
meomorphic, but proximal stalk with synarthries; stalk length
about 25–30 cm with distal stalk attached on hard substratum;
short pinnules, purple in adoral view (Fig. 20c); 10 arms prox-
imally divided (may be IBr2ax); robust proximalmost arm
without pinnule (Fig. 20b); first pinnule beyond IIBr2 (prob-
ably beyond IIBr4).
Environment: Depths 2125 m and maybe 1997 m on rocky
substratum dusted by sediment.
Remarks: This taxon seems to be new and its affinities
cannot be clarified from videos. The open proximal crown
of the small specimen (Fig. 20d) somewhat resembles that of
the specimen in Fig. 20a rather than other ten-armed
bathycrinid-like specimens that we observed. It has a stalk
 Author's personal copy
Mar Biodiv

Fig. 17 Metacrinin phenotype 6.

a. Individual in weak current with
crown in ballooning posture.
Laser dots 10 cm apart; depth
296 m. Shimmering water here
suggests fresh water outflow on
this island slope. b. Detail of
distal stalk. c. Detail of proximal
stalk

of ~20 columnals higher than wide suggesting that it is a
juvenile stage of a substantially larger adult.
Substrata
Seafloor character reflected the complex geophysical setting.
The deepest basins were floored in foraminiferal ooze with
occasional erratic blocks providing purchase for sessile ani-
mals. In the southern Sangihe Arc (Fig. 1), volcanoclastics
were common, but deep sandstones, possibly consolidated
ooze, formed extensive outcrops. Volcanic talus and vesicu-
lated extrusive lavas were common in the mid and north arc
from 400 to 700 m depths. Volcanics on the Talaud Ridge
were topped with limestone sediments and outcrops in
shallower portions. There was abundant hard substratum in
steep terrain, much of which appeared relatively recent with
little sediment cover.
We recorded the frequency of substratum type on sev-
eral dives for four of the most abundant crinoids:
G . ne o c a l e d o n i c u s , P t i l o cr i n us c f . a m e z i an e a e ,
N. hawaiiensis, and E. alternicirrus. Stalked crinoids most

Fig. 18 Metacrinin phenotype 7.

a. Individual crawling on rock.
Crown about 13 cm across; depth
465 m. b. Detail of mid-stalk. c.
Detail of distal stalk
 Author's personal copy
Mar Biodiv

Fig. 19 Crawling metacrinin

phenotype 6 or 7. a. Individual
crawling on sediment. Crown
about 10 cm across; depth 258 m.
b. Three individuals on mixed
substratum in high current; depth
259 m

often attached to bedrock (165/189=87.3 %) and less on
talus (19/189=10 %). Observations on mixed (3/189) and
boulder (2/189) substrata were infrequent although uncon-
solidated surfaces were abundant. Local topographic highs
were more likely to be colonized. On steep slopes, the
substratum often appeared unstable, creating highly het-
erogeneous talus scree (see Figs. 9g, 21b). Particulates
may form deposits that drape boulders and the attached
bases of crinoid stalks (Figs. 2b, d, 21b). Hyocrinids usu-
ally colonized rocks or boulders emerging from sediment
(Figs. 4a, 6a). Naumachocrinus occurred on more regular
slope surfaces variably draped in sediment. Their feeding
posture indicates localized upwelling current (Fig. 21a).
The only stalked crinoids found on sediments were
crawling metacrinins (Fig. 19). We encountered substantial
currents around abrupt topography on many dives, espe-
cially above 500 m depth; in deeper water, turbidity
currents may have generated rippled sediments near some
crinoids (Fig. 2b, d).
Distributions, assemblages and density
Taxon distribution and abundance related to depth have bi-
modal distributions (Fig. 22; Online Resource 4 notes all
sightings). The metacrinins and P. cf. amezianeae dominate
the diversity peak at 400 to 600 m, while E. alternicirrus,
G. neocaledonicus and P.ruberrimus are most abundant be-
tween 1000 and 1600 m, along with metacrinin 4 (seen at one
site only). Only G. neocaledonicus is relatively abundant be-
low 1600 m. Both diversity at the family-subfamily level and
total crinoid abundance were highest in the 1000 to 1600 m
interval (Fig. 23). E. alternicirrus and N. hawaiiensis appear
eurybathic down to 1600 m; if the ptilocrinid-like specimens

Fig. 20 Unknown taxon with

homeomorphic stalk (BPurple
umbrella crinoid^). a. Individual
with crown in ballooning posture
before disturbance by ROV.
Estimate of stalk length 30 cm;
depth 1997 m. b. Crown after
disturbance, with resuspended
sediment deposited on arms and
pinnules. c. Detail of pinnules. d.
Young individual on same dive in
weak current presumed to belong
to the same species. Size not
available; depth 2125 m
 Author's personal copy
Fig. 21 Steep unstable slopes. a.
Slumped surface of sediments and
debris; two bathycrinid-like
crinoids (possibly
Naumachocrinus) in feeding
posture indicating substantial
upwelling current, one white
echinothurioid sea urchin in the
lower left corner; depth 1072 m
on dive II-12. b. Single
Guillecrinus on an unstable scree
talus slope; depth 1325 m on dive
III-6
observed at depths of 1500 to 1600 m are P. cf. amezianeae
(rather than Feracrinus cf. koslowi), they represent a third
eurybathic species. From 2100 m to 3185 m, only a few small,
unidentifiable bathycrinid-like individuals occurred on two
dives where sediments predominated, except one Hyocrinus
cyanae, which is the deepest identifiable species at 3150 m.
Depth distributions of the known species of Indonesian cri-
noids fall mostly within prior observations (Fig. 23), with
depth extensions for N. hawaiiensis, P. ruberrimus,
E. alternicirrus and H. cyanae.
Regional distribution (Fig. 24) encompasses several wide-
spread taxa, especially the metacrinins and E. alternicirrus.
While the pattern is confounded by depth, some differences
occur: II-4 encountered many G. neocaledonicus on the north-
ern Sangihe Arc, while this species did not occur on II-8 and
III-9 on the Talaud Ridge at similar depths (~1600 m) where
E. alternicirrus and metacrinines dominated. Some taxa
(P. ruberrimus and metacrinin phenotypes 3 and 4) occurred
at only one site.
For the most part, stalked crinoids were sparsely distributed
in low densities. For several dives, we have intermittent tran-
sect data for total distances of 1 to 2 kilometers (Table 2)
Fig. 22 Numbers of stalked crinoids observed by taxon (over ten
sightings each) with respect to depth intervals. Numbers of dives in
those intervals noted at end of bars. Ptilocrinus and metacinins
dominated in shallow waters, but both diversity at the family or
subfamily level and total crinoid abundance were highest in the 1000–
1600 m interval
Mar Biodiv
where densities reached 186 individuals/km. Comatulid cri-
noids were over three times more abundant overall. A
clumped dispersion was evident for a few species, most nota-
bly metacrinin phenotype 4, which occurred on only the last
segment of one dive, where it was associated with numerous
yellow comatulids presumed to belong to the Family
Thalassometridae (Fig. 15a).
Functional morphology and behaviour
Several taxa were often observed with the crown in a balloon-
like posture and stalk upright (or bent just below the crown) in
weak or absent currents: Hyocrinus sp. 2 (Fig. 5), metacrinin
phenotypes 5 and 6 (Figs. 16 and 17), the purple umbrella
species (Fig. 20), and more rarely, E. alternicirrus (Fig. 11).
Hyocrinus sp. 2 is the first record of such an unusual posture
for hyocrinids; the highly recurved arms are due to relaxed
adoral muscles when aboral ligaments in the brachial articu-
lations contract. Metacrinin phenotype 5 displays posture
changes in different hydrodynamic conditions. In a weak cur-
rent, the oral surface orients more or less upward with cirri
extended outward from the stalk or lying along it, either up-
ward or downward; the flexible proximal stalk bends slightly,
and the arms flex aborally, forming a balloon-like posture with
pinnules moderately open and distal arm tips bent almost per-
pendicularly outward (Fig. 16a, b). When current velocity
increases moderately, the stem bends at least 60° and arms
sprawl out with distal tips curled orally or down-current
(Fig. 16c, with the same tip posture as in phenotype 3,
Fig. 14).
As the ROV settled to film the new (tentative) purple um-
brella species, it was in a stable crown ballooning posture with
pinnules unfolded and stalk erect (Fig. 20a). ROV turbulence
resuspended a cloud of fine sediment, causing tilting of the
crown that maintained the same crown posture as the upper
stalk flexed. Neither arms nor pinnules displayed movement
except retraction of most podia (Fig. 20b, c); then, as sediment
 Author's personal copy
Mar Biodiv
Fig. 23 Bathymetric distribution
of identified taxa. Grey bands:
ranges known from other records
for these taxa. Black bands:
ranges observed in the present
study. Stippled band: hyocrinid-
like specimens observed in the
present study
particles settled, the specimen recovered the same previous
posture. The presumed juvenile of the same species has a
typical filtration fan, with pinnules wide open and arms ab-
orally flexed into the current (Fig. 20d).
Other taxa (Proisocrinus, metacrinin phenotypes 1 and
2, phrynocrinids, Naumachocrinus, bathycrinid-like speci-
mens and hyocrinids) have expanded crowns, maximizing
surface area in weak currents. In substantial current,
P. ruberrimus forms a conical filtration fan and the proxi-
mal stalk is strongly curved (Fig. 9a, b). In moderate cur-
rent, the parabolic filtration fan is open wide with arms
slightly curved up-current (Fig. 9d, g). In low current ve-
locity, the crown was frequently observed opened, but with
pinnules completely folded over ambulacral grooves
(Fig. 9e, f). In moderate current, hyocrinids have the arms
slightly but regularly curved up-current (Figs. 3a, 4 and 6).
In weak to absent water movement, cirri of the metacrinins
(except E. alternicrirrus) were folded against the mid part
of the stalk, which was perpendicular to the substratum
(see Fig. 12b, e); they may adopt a crown ballooning pos-
ture. Nearly all recorded individuals of metacrinin pheno-
type 4 dangled from vertical surfaces, where they were
attached by distal cirri with their crowns more or less
closed in a posture not previously seen.
Guillecrinus never forms a parabolic filtration fan. Instead,
in weak currents, the rigid stalk remains straight or
subvertical, with flexible arms and pinnules tending to curve
downward distally; frequent arm movements suggest a possi-
ble active capture of food (Fig. 2a, b). In stronger currents, the
stalk curves gently (frequently markedly in the proximal
mesistele), with the oral surface oriented down-current and
the straight or outwardly curved arms assuming a conical pos-
ture (Fig. 2c, d). Extended podia were not observed in close
views.
Individuals belonging to metacrinin phenotypes 5 and 6
were observed slowly crawling using slow arm movements.
Cirri are extended perpendicularly to the stalk in a regular
pattern (Figs. 18 and 19a). Tips of the lowest arms and distal
claws of cirri brace against substratum protrusions. Both seem
to contribute to the crinoid displacement. Three crawling
metacrinins were observed in one view (Fig. 19b), suggesting
synchronous response to a stimulus.
Discussion
Crinoids were abundant on most dives in the region, although
about 85 % of the occurrences were in non-stalked crinoids.
Nonetheless, the 770 stalked crinoids examined on 17 dives
represent a broad range of taxa from at least six families with a
diverse group of phenotypes in the subfamily Metacrininae.
While several taxa are present only rarely, observation of their
live characters and habitat extends the limited information on
this poorly accessible group of echinoderms.
Crinoid behaviour
The variety of feeding postures reflects differences in food-
gathering modes. As most individuals occurred on rocky fea-
tures, habitat selection seems to target higher water turbu-
lence; where rippled sediments indicated mean current direc-
tion, we observed filtration fans usually tipped down-current.
Macurda and Meyer (1974) described similar feeding pos-
tures, with arms and pinnules forming a parabolic fan as a
passive filter with the food grooves and mouth orientated
down-current. In contrast, the metacrinin phenotype 4
(?Metacrinus nodosus) nearly always displayed a crown-
downward posture with arms in a cone shape, suggesting that
 Author's personal copy
Mar Biodiv

Fig. 24 Spatial distribution

within the study area of the same
taxa as in Fig. 22. Top numerals in
each pie=dive number, below is
total number of stalked crinoids in
the taxa observed on that dive

local resuspension of particles may be important. In similar Roux 1991; Roux 1994). The postures of metacrinin pheno-
topography of canalized flow off New Caledonia, a high den- types 1 and 2 with conical or subhorizontal crown and pin-
sity of echinoderm suspension feeders occurred (Lambert and nules closed (Figs. 12c and 13b) was first described in

Table 2 Transecting to count animals was possible for sustained distances on select dives

Dive Mean depth (m) Transect area (m2) Transect distance (km) Stalked /km Non-stalked /km Stalked taxa where > 10 encountered

III - 10 296 3721 1.65 20 90 Metacrinins

II - 11 476 3279 1.45 39 321 Metacrinins
III - 01 585 4594 2.03 35 215 P. cf amezianeae
III - 13 776 3567 1.58 10 58 mixed
III - 14 1284 2315 1.03 186 677 G. neocaledonicus, metacrinins
III - 09 1563 4129 1.83 58 44 E. alternicirrus
III - 03 1859 3220 1.43 64 3 P. ruberrimus

The distance noted is the sum of the segments of the dive during which ROV flight maintained a consistent viewing distance over the bottom. Taxa of
non-stalked crinoids were not possible to distinguish. Average depth change for a dive was about 100 m
 Author's personal copy
Mar Biodiv
Metacrinus rotundus by Fujita et al. (1987, Fig. 5), which
adapts the filtration fan posture as hydrodynamic conditions
change (Kitazawa and Oji 2014). The balloon-like crown pos-
ture of metacrinin type 5 (Fig. 16a, b) is analogous to the
unusual crown posture of Hyocrinus sp. 2 (Fig. 5), in which
the gaping orals may form by permanent aboral ligaments
with no counter action (i.e., adoral muscle contraction or sub-
stantial current), or, more likely, active control by aboral mu-
table collagenous tissue when the current regime changes.
In the Order Isocrinida, stalk attachment by an encrusting
disk is a plesiomorphic character that is shared by juveniles of
Balanocrininae and Diplocrininae and adult Proisocrinus,
whereas the development of the cirrus whorl on nodals and
attachment by cirri is a derived character. The two observed
cases of five finger-like holdfasts in Proisocrinus ruberrimus
could reflect the juvenile attachment by the distalmost cirri.
However, as suggested by juvenile attachment in
Endoxocrinus, the lack of cirri and the development of an
attachment disc are more likely related to paedomorphic de-
velopment through ontogeny rather than derived characters.
The juvenile specimen of Endoxocrinus alternicirrus
photographed at 1175 m (Fig. 11a) is the first observation of
a young diplocrinin attached by an encrusting disk. Carpenter
(1884, pl. 30–4) first figured a young specimen of
Hypalocrinus naresianus (Clark 1908) with a distal attach-
ment disk. He also described a juvenile of Neocrinus decorus
(Thomson, 1864) with the distal columnals articulated by
synarthries. These two species belong to the subfamily
Balanocrininae, which is closely related to Diplocrininae
(Roux et al. 2009). Clark (1912, fig. 55) figured a young
specimen of the diplocrinin Teliocrinus springeri liliaceus
(Clark, 1909) with the distalmost noditaxis of columnal fea-
ture suggesting articulation by synarthries. Améziane-
Cominardi (1991) found similar distal stalk synarthries in a
young specimen attributed to Metacrininae.
Slow crawling in the Caribbean species Endoxocrinus
parrae [Gervais (in Guérin, 1835)] and Cenocrinus asterius
(Linné, 1775) was first observed by Messing et al. (1988).
Baumiller and Messing (2007) reported rapid crawling
(10 mm s−1) in Neocrinus decorus, using only its arms.
Birenheide and Motokawa (1994) report very slow crawling
(0.1 mm s−1) in Metacrinus rotundus using its arms during
laboratory observations. Our in situ observations of crawling
metacrinins on sedimentary (Fig. 19) and rocky substrata
(Fig. 18) suggest that cirri may play an active role. In
Metacrinus rotundus, the mutable collagenous tissue that con-
nects cirrus ossicles can develop slow active contractions un-
der nervous control (Birenheide et al. 2000).
The western Atlantic species Cenocrinus asterius also ex-
hibits cirri folded against an erect mid-stalk when arms are
closed, as in metacrinins; Baumiller (2008) suggested that cirri
have an active role to maintain stalk rigidity when the crown is
no longer supported by hydrodynamic lift as currents wane.
This hypothesis assumes that stalk ligaments (including mu-
table collagenous tissue) cannot support an upright posture in
the absence of current. However, our observations of homeo-
morphic stalked crinoids with an upright stalk posture without
cirri in very weak currents (Figs. 2a, 4a, 5a, 6a, 20a) indicate
that paraxial ligaments can provide sufficient stalk stiffness to
support the crown.
Crinoid distribution
We observed abundant metacrinins on the shallowest dives at
250 m. In general, from this depth to the bathyal region, strong
turbulence and high currents induced by storm waves are ab-
sent, while food supply is greater than in the abyssal plains;
however, low oxygen and water mass boundaries may reduce
species richness (Roux 1987; Améziane and Roux 1997).
Most of the isocrinids and five-armed bathycrinids are found
in epibathyal environments, whereas hyocrinids and ten-
armed bathycrinids predominate below 2000 m. The depth
of maximum diversity depends on many oceanographic vari-
ables. It occurs in the lower margin of the continental slope
(2000–3000 m) in the NE Atlantic and on the upper slope
(300–500 m) of the western tropical Atlantic. In the western
tropical Pacific, a major peak occurs at 300 to 500 m and a
secondary peak occurs at 1000 to 2000 m, with a low in
species richness at 800 to 900 m (Roux 1987; Améziane and
Roux 1997). The data from the Celebes and Molucca Seas are
consistent with this pattern, although the largest peak occurs at
1000 to 1600 m depth, where nine of the 17 taxa were record-
ed (Fig. 22). The difference could be due to insufficient inves-
tigations of sedimented and mixed substrata at shallower
depths in the present study, or, for the global model of
Améziane and Roux (1997), to underestimation of species
diversity mainly estimated from specimens collected with
beam trawls that are less successful on irregular rocky floors,
which were common in our study.
We observed small areas with abundant crinoids. However,
densities never constituted the Bcrinoid meadows" that form
when mobile isocrinids converge on ideal environmental con-
ditions at 15–20 individuals per m2 (Conan et al. 1981;
Messing et al. 1990), or with local mass recruitment
(Messing 1985; Oji et al. 2009). Usually, stalked crinoids find
optimal filtration conditions in moderate but substantial cur-
rent velocities and water flow of low turbulence (Roux 1987).
Along with current regime, substratum complexity is cited as
a major factor in determining stalked crinoid abundance
(Conan et al. 1981; Baumiller 2008). Baumiller et al. (1991)
reported that isocrinids climbed objects in flow tank experi-
ments. Hard substrata and boulders may be ideal stable habi-
tats that offer a perch in high currents (Figs. 2a, 3a, 9f, 12c).
While mobile isocrinids may relocate to optimum conditions
(Baumiller 2008), permanently attached stalked crinoids can-
not. The persistence of abundant immobile stalked crinoids
 Author's personal copy
like hyocrinids (Eléaume et al. 2011), Guillecrinus (Fig. 2d) or
Proisocrinus (Fig. 9g) may be indicative of favourable and
relatively stable habitat conditions, sometimes despite sub-
stantial sedimentation over their holdfasts (Fig. 2a, d).
Relevance to biogeography
During the last decade, our knowledge about the biogeograph-
ic distribution of stalked crinoids has made significant prog-
ress. Nevertheless, it remains sketchy because of the hetero-
geneity of deep-sea investigations and the scarcity of detailed
studies using data from submersible vehicles where hard
substratum is abundant. The historical biogeography
proposed by Améziane and Roux (1997) needs testing under
the light of new data. Here, for practical reasons, we subsume
to a single species all specimens identified by Baff^ and also
simplify by considering Guillecrinus neocaledonicus as a ju-
nior synonym of G. reunionensis, as suspected by Améziane
and Roux (2005).
Endoxocrinus alternicirrus and monotypic
Naumachocrinus and Proisocrinus are widespread from
Japan to New Zealand and the central Pacific, whereas they
remain unknown in the Indian Ocean. N. hawaiiensis and
E. alternicirrus are eurybathic species, but P. ruberrimus is
not (Fig. 23). The genus Endoxocrinus has a tethyan-like dis-
tribution (i.e., W to Central Pacific, N-E Atlantic and W-tropical
Atlantic) (David et al. 2006). Moreover, it is closely related to
the diplocrinin genus Teliocrinus from the northern Indian
Ocean (Roux et al. 2009). Diplocrininae shares with
Balanocrininae a tethyan-like distribution, having close affini-
ties (Roux et al. 2009) confirmed by molecular phylogeny
(Hemery 2011; Hemery et al. 2013). The balanocrinin species
Hypalocrinus naresianus has a distribution similar to
E. alternicirrus in the western Pacific, where it occurs in abun-
dance, including in samples taken off the SW Philippines
(Bourseau and Roux 1989; Eléaume et al. 2007). However, it
was not seen in the present study, and is unknown from the
Central Pacific. The widespread tethyan-like distribution of
Endoxocrinus, including the NW, SW and Central Pacific, sug-
gests an ancient origin of the genus from both southern
(Gondwanaland) and northern (Eurasia) Tethys margins with
passive expansion towards the Central Pacific through geolog-
ical time (Roux 1987; Améziane and Roux 1997). An alterna-
tive interpretation is a more rapid larval dispersal than previous-
ly thought and a younger origin of the species. Diplocrininae
are known from the Miocene of Japan (Oji 1990) and the west-
ern tropical Atlantic (Donovan 2001), but Proisocrinus and
Naumachocrinus are unknown from the fossil record.
Guillecrinus reunionensis (including G. neocaledonicus)
and Porphyrocrinus verrucosus are two Indo-Pacific species
that have yet to be found north of SE Philippines and may have
originated on the Gondwanaland margins of the Tethys Sea.
The eastern most site of Guillecrinus distribution is the
Mar Biodiv
Vailulu’u Seamount in the East Samoa Chain (Staudigel et al.
2006, Fig. 3k). Ptilocrinus cf. amezianeae could have the same
history, as suggested by the presence of P. amezianeae on Indo-
W Pacific, Antarctic and subantarctic seamounts (Eléaume
et al. 2011, 2014). Columnals of a xenomorphic stalk attributed
to Porphyrocrinus are known on the Eurasian margin from
Late Miocene of Spain (Roux and Montenat 1977) and
P. thalassae is a large extant NE Atlantic species (Roux
1985). Guillecrinus is unknown in the fossil record.
The oldest metacrinin was found in the Early Eocene of the
Antarctic Peninsula (Baumiller and Gazdzicki 1996).
Therefore, the Gondwanaland margin origin of this subfamily
is likely, and we could expect an Indo-Pacific distribution to-
day. However, extant metacrinins are unknown in the Indian
Ocean (except in its NE boundary between Indonesia and
Australia), but occur in the western Pacific from Tasmania to
Japan. Saracrinus angulatus (the species having close affinities
with our metacrinin phenotype 1) was frequently collected
from Japan to Timor and the NW Australian margin, but never
in the SW Pacific (Améziane 1997; Eléaume et al. 2007).
Metacrinus rotundus is restricted to an area including Japan
and South China Sea, whereas Saracrinus nobilis is known
from the whole western Pacific. It appears that the complex
channels and basins of the Indonesian seas still present a bio-
geographic barrier between the Pacific and Indian Oceans.
Except for Eocene fossils with preserved crowns from
Antarctica, only a few fossil columnals are attributed to
Metacrininae from Early Miocene of New Zealand (Eagle
2004), Middle-Late Miocene of western Europe (Roux and
Montenat 1977; Klikushin 1982) and Pliocene of Seran
(Indonesia) (Sieverts 1933), but such attribution is only tenta-
tive because the structure of stalk articular facets has limited
diagnostic value, and nothing is known about arm division
patterns from these fossils. We conclude that a distribution
restricted to the NW or SW Pacific cannot be used to argue
an origin from Eurasian or Gondwanian tethyan margins with-
out the contribution of robust paleontological data; a wide-
spread distribution in the western Pacific (from the South to
the North) could be due to relatively recent dispersal with re-
spect to geological times, especially for metacrinins. Therefore,
to delimit the location of the Wallace line only using extant
stalked crinoid distribution seems to be highly speculative.
Acknowledgments We are grateful to: Steven Hammond, David
McKinnie, Jeremy Potter and Sugiarta Wirasantosa for facilitating the
evolution of the INDEX plans; Jim Holden and Tim Shank for shoreside
science oversight, and Santiago Herrerra, John Sherrin, Rainer Troa, and
Xerandy as interfaces for telepresence research on board; Cherisse Du
Preez and Rachel Brown for help with dive logging; Meme Lobecker and
Kelley Elliott (NOAA) for the multibeam map; and Jessica Nephin for
some crinoid observations and density estimates. Special thanks to Jona-
than Rose, who mastered video capture, logged dives, captured images
and helped with manuscript production. We thank Nadia Améziane
(MNHN), Franzis Althaus, Karen Gowlett-Holmes, Felicity McEnnulty,
and Alan Williams (CSIRO), who provided information and access to
 Author's personal copy
Mar Biodiv
deep-sea photographs of stalked crinoids off SE Australia and Tasmania.
Thanks are also due to Gert Woerheide (LMU) for pictures and a speci-
men of Porphyrocrinus collected during the Deep Downunder Expedi-
tion (funded by the German Science Foundation Projects Wo896/7 &
LU839/2). The INDEX expedition was funded by National Oceanic
and Atmospheric Administration’s Ocean Exploration Program, and sup-
ported by several agencies of the US and Indonesian governments. VT
was funded by the Canada Research Chairs Foundation. MR and ME
were funded by various MNHN grants: BActions Transversales^:
BBiodiversité actuelle et fossile. Crises, stress, restaurations et
panchronisme : le message systématique^, BEmergences^, BFormes pos-
sibles, Formes réalisées^ and BInteractions Minéral-Vivant^
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