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Echinoderes (Kinorhyncha: Cyclorhagida) from the Hikurangi
Margin, New Zealand
Katarzyna GRZELAK 1,* & Martin V. SØRENSEN 2
1 Polish Academy of Sciences, Institute of Oceanology, Sopot, Poland.
1,2 Natural History Museum of Denmark, University of Copenhagen, DK-2100 Copenhagen, Denmark.
* Corresponding author: kgrzelak@iopan.gda.pl
2 Email: mvsorensen@snm.ku.dk
1 urn:lsid:zoobank.org:author:ABFE9734-0BD8-4ED9-A780-469D0058FCED
2 urn:lsid:zoobank.org:author:4143D650-12FC-4914-93F5-2C39339A7156
Abstract. Limited data are available for the kinorhynch fauna from the Southern Hemisphere, with little
or no data from New Zealand. Here, we provide a rst comprehensive overview of the diversity of mud
dragons, with an emphasis on species of Echinoderes from the continental slope of New Zealand, from
a variety of habitats such as slopes, canyons and seamounts located in the Hikurangi Margin region. The
study revealed fteen species of Echinoderes. Of these, ten are described as new to science: E. aragorni
sp. nov., E. blazeji sp. nov., E. dalzottoi sp. nov., E. frodoi sp. nov., E. galadrielae sp. nov., E.gandal
sp. nov., E. landersi sp. nov., E. leduci sp. nov., E. legolasi sp. nov. and E. samwisei sp. nov. Moreover,
Echinoderes juliae Sørensen et al., 2018, Echinoderes sp. aff. E. balerioni, Echinoderes sp. aff.
E. galadrielae/beringiensis, Echinoderes sp. aff. E. lupherorum and Echinoderes sp. aff. E. unispinosus
are reported in the investigated region. The most abundant among all was E.gandal sp. nov., but it
was found only in canyons. Interestingly, the second most common species was E. juliae that was found
at several stations in canyons, seamount and on the slope. This species is one of the deep-sea species
originally found on the abyssal plain off Oregon and along the continental rise off California, Northeast
Pacic, recorded in polymetallic nodules in the tropical eastern Pacic, and recently found on the abyssal
plains off Chile, east of the Atacama Trench. These ndings, together with records of Echinoderes sp.
aff. E. lupherorum and Echinoderes sp. aff. E. unispinosus indicate that, despite their low dispersal
abilities, kinorhynchs, similar to other meiofaunal species, may exhibit a wider distribution pattern than
previously assumed. The number of recorded species and numerous new species show that New Zealand
sediments not only are inhabited by a diverse kinorhynch fauna, but Echinoderes, the most speciose
genus, still holds much to discover.
Keywords. Echinoderidae, deep-sea canyons, kinorhynchs, meiofauna, seamounts, taxonomy.
Grzelak K. & Sørensen M.V. 2022. Echinoderes (Kinorhyncha: Cyclorhagida) from the Hikurangi Margin, New
Zealand. European Journal of Taxonomy 844: 1–108. https://doi.org/10.5852/ejt.2022.844.1949
Introduction
Over the past years, studies on the taxonomic composition and distribution of kinorhynchs have intensied
considerably, and the number of reports of new kinorhynch species from the most remote locations
1
European Journal of Taxonomy 844: 1–108 ISSN 2118-9773
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2022 · Grzelak K. & Sørensen M.V.
This work is licensed under a Creative Commons Attribution License (CC BY 4.0).
Monograph
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increases steadily (e.g., Cepeda et al. 2020; Grzelak et al. 2021; Adrianov & Maiorova 2022). However,
most of our knowledge about this group is still concentrated in regions in the Northern Hemisphere
where taxonomists happened to be located, i.e., southern Europe (e.g., Sánchez et al. 2012; Dal Zotto &
Todaro 2016), East Asia (e.g., Sørensen et al. 2012, 2013; Yamasaki & Fujimoto 2014; Yamasaki 2016,
2017), and North and Central America (e.g., Higgins 1983; Pardos et al. 2016a, 2016b; Sørensen et al.
2016b, 2018; Landers et al. 2018, 2022). Therefore, many areas, especially in the Southern Hemisphere,
remain virtually unexplored, although the presence of mud dragons is reported in ecological studies. We
nd one of the largest gaps in known kinorhynch biodiversity in the Oceanian Region, and in particular
around New Zealand – nearly a terra incognita in terms of kinorhynch fauna.
From several studies of the meiofauna of this region, it is known that mud dragons occur in a variety of
shallow waters and deeper continental margin habitats, such as slopes, canyons and seamounts (Coull &
Wells 1981; Kamenev et al. 1993; Pilditch et al. 2015; Rosli et al. 2016), but to date only six species
have been described from New Zealand territory. The rst formally described kinorhynch species
from this region were Echinoderes malakhovi Adrianov, 1999 in Adrianov & Malakhov (1999) and
Pycnophyes newzealandiensis Adrianov, 1999 in Adrianov & Malakhov (1999) from sandy sediments
at a depth of 60 m in the Bay of Plenty (Adrianov & Malakhov 1999). Some years later, Neuhaus &
Blasche (2006) described the new echinoderid genus Fissuroderes Neuhaus & Blasche, 2006 from deep-
sea localities east of New Zealand, including four new species of this genus (Neuhaus & Blasche 2006).
A more recent report of kinorhynchs in the area is restricted to a single species, Campyloderes cf.
vanhoeffeni Zelinka, 1913, found at the Hikurangi Plateau and Chatham Rise at depths of 420–1940 m
by Neuhaus & Sørensen (2013). In addition, Neuhaus (2013) cited Coull & Wells (1981) for reporting
Echinoderes coulli Higgins, 1977 from Pauatahanui Bay, Porirua, north of Wellington. The reported
identity of this species is rather doubtful though, and it is more likely that Coull & Wells (1981) collected
a yet undescribed species and representative of the E. coulli species complex (see Yamasaki & Fujimoto
2014). The most recent overview of New Zealand’s kinorhynch fauna was published by Neuhaus et al.
(2010) in the Biodiversity Inventory series of Gordon (2010). An updated version is expected to be
published in 2023 (Grzelak & Sørensen in prep.)
The present contribution is the result of studies of kinorhynchs found in the samples collected during
the cruise TAN1004, which took place in April 2010, in the Hikurangi Margin region, as part of an
extensive meiofauna sampling campaign conducted by the National Institute of Water and Atmospheric
Research (Bowden et al. 2016; Rosli et al. 2016). The main purpose of this study is to provide the rst
assessment of the species richness and distribution of Kinorhyncha Reinhard, 1881 in the deep waters
on the continental slope of New Zealand. Here, we contribute with the description of ten new species
of Echinoderes. Furthermore, we provide detailed data of ve known species of Echinoderes. It is clear
that the kinorhynch fauna from New Zealand has been largely neglected so far; therefore, these new
descriptions will help to signicantly expand the knowledge of this group in Southwest Pacic.
Material and methods
The study area is located along the Hikurangi Margin, southeast of the Cook Strait that separates North
and South Island, New Zealand (Fig. 1). Sampling was conducted at slope, canyon and seamount sites
from the RV Tangaroa during the National Institute of Water and Atmospheric Research (NIWA) voyage
TAN1004 in April 2010. A multicorer (MUC) with tubes of an internal diameter of 9.52 cm was used to
retrieve bottom sediment from 20 stations, with depths ranging from 670 m to 1561 m (Table 1). From
each deployment, one to three cores were taken from the MUC and dedicated to meiofauna analysis. The
samples were xed with 10% buffered formalin. More information about sampling sites and sampling
methodology can be found in Bowden et al. (2016) and Rosli et al. (2016).
European Journal of Taxonomy 844: 1–108 (2022)
2
In the laboratory, samples were washed in a 45 µm sieve and meiofauna organisms were extracted from
the sieved sediment by LUDOX otation (Somereld & Warwick 1996). First, main meiofauna taxa
were identied under a stereo microscope, then kinorhynchs were picked up and transferred to slides
with glycerine, sealed with parafn and stored for several years at NIWA. After this, the slides with
kinorhynch specimens were shipped to the Natural History Museum of Denmark (NHMD) for further
examination and description. A total of 356 kinorhynchs, including 204 adult individuals, were found.
For preparation of light microscopy (LM) slides, original glass slides were opened, specimens were
transferred directly to Fluoromount-G (mounting medium preferred for Kinorhyncha) and remounted.
The new preparations were either made on regular glass slides, or, preferably, by mounting the individual
specimens between two cover slips and attaching them to an H-S plastic slide. The specimens were
examined with an Olympus BX51 microscope with differential interference contrast, and photographed
with an Olympus DP27 camera. Measurements were made with CellˆD software. Line art gures were
made with Adobe Illustrator CS6. Examined specimens were deposited in the collections of NHMD and
NIWA (see Table 1 for catalogue numbers).
Preparation of specimens for scanning electron microscopy (SEM) was more complex than usual because
the kinorhynchs were originally stored in glycerine. Therefore, at rst, the specimens were rehydrated in
a 50% v/v water-glycerine solution. Then, the specimens were transferred to demineralized water, and
a small amount of liquid soap was added to get rid of the residual glycerine and organic particles. The
detergent was mixed with water, and numerous bubbles were created to help capture particles adhering
to the specimens. The samples were left overnight, after which bubbles were again produced. After three
rounds of washing, specimens were transferred to clean, deionized water to wash off any remaining
Fig. 1. Map showing the sampling stations. Colour codes indicate slopes (red), canyons (green), and
seamounts (yellow). Inset shows New Zealand with a frame marking the sampling area.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
3
Table 1 (continued on next four pages). Coordinates and basic data on sampling stations, and identities, type status and catalogue numbers of species
of Echinoderes Claparède, 1863 from each station.
Location Stn Depth (m) Latitude
(°S)
Longitude
(°E) Species Mounting Type status and catalogue number
Slope 4 1046 41.6837 175.6642 E. aragorni sp. nov. LM 1 ♀, paratype (NIWA-159424)
E. frodoi sp. nov. LM 1 ♂, paratype (NHMD-916331)
1 ♀, 1 ♂, paratypes (NIWA-159415–159416)
E. aff. galadrielae/beringiensis LM 1 ♀, non-type (NIWA-159427)
E. juliae LM 1 ♂, 1 ♀, non-types (NHMD-921715, NHMD-921717)
E. aff. lupherorum LM 1 ♂, non-type (NIWA-159428)
E. samwisei sp. nov. LM 1 ♀, 1 ♂, paratypes (NHMD-917299–917300)
2 ♂♂, paratypes (NIWA-159419–159420)
17 1514 41.6288 175.8682 E. juliae LM 1 ♀,1 ♂, non-types (NHMD-921723–921724)
38 1121 41.5937 175.8532 E. aragorni sp. nov. LM ♀, holotype (NIWA-159423)
E. frodoi sp. nov. LM 1 ♀, paratype (NHMD-916335)
SEM 1 ♀, 1 ♂, non-types
E. aff. galadrielae/beringiensis SEM 2 ♂♂, non-types
E. juliae LM 1 ♂, non-type (NHMD-921729)
E. landersi sp. nov. LM ♀, holotype (NIWA-159402)
E. aff. lupherorum LM 1 ♂, non-type (NHMD-921636)
E. samwisei sp. nov. LM ♂, holotype (NIWA-159418)
1 ♂, paratype (NIWA-159421)
SEM 1 ♀, non-type
E. aff. unispinosus LM 1 ♂, non-type (NHMD-921792)
SEM 1 ♀, 1 ♂, non-types
44 728 41.5258 175.8003 E. aragorni sp. nov. SEM 1 ♀, non-type
E. juliae SEM 1 ♀, non-type
European Journal of Taxonomy 844: 1–108 (2022)
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Location Stn Depth (m) Latitude
(°S)
Longitude
(°E) Species Mounting Type status and catalogue number
Slope E. landersi sp. nov. SEM 1 ♀, non-type
E. legolasi sp. nov. SEM 1 ♂, non-type
E. aff. lupherorum SEM 1 ♀, non-type
E. samwisei sp. nov. SEM 1 ♀, non-type
76 1282 41.6833 175.6500 E. frodoi sp. nov. LM
SEM
1 ♀, paratype (NHMD-916336)
1 ♀, non-type
E. galadrielae sp. nov. SEM 2 ♀♀, non-types
E. aff. galadrielae/beringiensis LM 1 ♀, non-type (NHMD-921503)
SEM 1 ♂, non-type
E. juliae SEM 1 ♀, non-type
E. aff. lupherorum SEM 2 ♂♂, non-types
128 1420 42.0485 174.7000 E. frodoi sp. nov. LM 1 ♀, paratype (NHMD-916337)
E. aff. lupherorum LM 1 ♂, non-type (NHMD-921635)
Pahaua Canyon 12 1350 41.5508 175.7250 E. galadrielae sp. nov. LM 2 ♀♀, paratypes (NHMD-921496–921497)
E. aff. galadrielae/beringiensis LM 1 ♀,1 ♂, non-types (NHMD-921498–921499)
E.gandalsp. nov. LM 1 ♀, paratype (NIWA-159409)
E. juliae LM 2 ♀♀, 2 ♂♂, non-types (NHMD-921718–921721)
E. samwisei sp. nov. LM 1 ♂, paratype (NHMD-917301)
22 1188 41.5100 175.7187 E. frodoi sp. nov. LM ♂, holotype (NIWA-159414)
1 ♀, paratype (NIWA-159417)
E.gandalsp. nov. LM 1 ♀, paratype (NHMD-916362)
1 ♂, paratype (NIWA-159410)
E. juliae LM 1 ♀, non-type (NHMD-921728)
Table 1 (continued). Coordinates and basic data on sampling stations, and identities, type status and catalogue numbers of species of Echinoderes
Claparède, 1863 from each station.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
5
Location Stn Depth (m) Latitude
(°S)
Longitude
(°E) Species Mounting Type status and catalogue number
Pahaua Canyon E. aff. lupherorum LM 1 ♂, non-type (NIWA-1592429)
27 1013 41.4983 175.7043 E. blazeji sp. nov. LM ♂, holotype (NIWA-159400)
1 ♀, paratype (NHMD-917223)
1 ♂, paratype (NIWA-159401)
SEM 1 ♂, non-type
E. dalzottoi sp. nov. LM 1 ♂, paratype (NHMD-917147)
SEM 1 ♂, non-type
E. frodoi sp. nov. SEM 1 ♂, non-type
E. galadrielae sp. nov. SEM 3 ♀♀, 1 ♂, non-types
E. aff. galadrielae/beringiensis SEM 2 ♂♂, non-types
E.gandalsp. nov. LM 1 ♂, paratype (NIWA-159411)
SEM 5 ♀♀, 3 ♂♂, non-types
E. juliae SEM 1 ♂, non-type
E. landersi sp. nov. SEM 2 ♀♀, non-types
E. leduci sp. nov. SEM 1 ♂, non-type
E. aff. lupherorum LM 1 ♂, non-type (NIWA-159430)
SEM 3 ♀♀, 2 ♂, non-types
31 730 41.4962 175.6828 E. dalzottoi sp. nov. LM ♀, holotype (NIWA-159403)
SEM 1 ♂, non-type
E. frodoi sp. nov. LM 1 ♂, paratype (NHMD-916332)
E.gandalsp. nov. LM ♂ holotype (NIWA-159407)
1 ♀, paratype (NHMD-916356)
1 ♀, paratype (NIWA-159408)
E. juliae LM 1 ♂, non-type (NHMD-921725)
Table 1 (continued). Coordinates and basic data on sampling stations, and identities, type status and catalogue numbers of species of Echinoderes
Claparède, 1863 from each station.
European Journal of Taxonomy 844: 1–108 (2022)
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Location Stn Depth (m) Latitude
(°S)
Longitude
(°E) Species Mounting Type status and catalogue number
Pahaua Canyon E. landersi sp. nov. LM 1 ♀, paratype (NHMD-916627)
E. leduci sp. nov. LM ♂, holotype (NIWA-159405)
Honeycomb Canyon 53 948 41.4563 175.8970 E. aff. galadrielae/beringiensis LM 1 ♂, non-type (NHMD-921500)
58 670 41.4080 175.8977 E. aragorni sp. nov. LM 1 ♀, paratype (NHMD-920115)
E. blazeji sp. nov. SEM 3 ♀♀, 1 ♂, non-types
E. dalzottoi sp. nov. LM 1 ♂, paratype (NIWA-159404)
SEM 1 ♀, non-type
E. frodoi sp. nov. SEM 1 ♂, non-type
E. aff. galadrielae/beringiensis LM 1 ♂, non-type (NHMD-921502)
E.gandalsp. nov. LM 2 ♂♂, paratypes (NHMD-916357–916358)
1 ♀, paratype (NIWA-159412)
E. juliae LM 2 ♀♀, non-types (NHMD-921726–921727)
SEM 2 ♀♀, non-types
E. leduci sp. nov. LM 1 ♀, paratype (NIWA-159406)
SEM 1 ♂, non-type
62 1171 41.4760 175.9477 E. aragorni sp. nov. SEM 1 ♂, non-type
E. blazeji sp. nov. SEM 1 ♂, non-type
E. galadrielae sp. nov. SEM 1 ♀, non-type
E.gandalsp. nov. SEM 1 ♀,2 ♂♂, non-types
E. juliae SEM 1 ♀, non-type
E. aff. lupherorum SEM 1 ♀, non-type
E. samwisei sp. nov. SEM 1 ♀, non-type
Table 1 (continued). Coordinates and basic data on sampling stations, and identities, type status and catalogue numbers of species of Echinoderes
Claparède, 1863 from each station.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
7
Location Stn Depth (m) Latitude
(°S)
Longitude
(°E) Species Mounting Type status and catalogue number
Campbell Canyon 92 683 41.8922 174.6347 E. frodoi sp. nov. SEM 1 ♂, non-type
126 1495 42.1422 174.5492 E. frodoi sp. nov. LM 1 ♀, paratype (NHMD-916333)
SEM 1 ♀, non-type
E. galadrielae sp. nov. LM ♀, holotype (NIWA-159425)
E.gandalsp. nov. LM 2 ♂♂, 1 ♀, paratypes (NHMD-916359–916361)
1 ♂, paratype (NIWA-159413)
SEM 1 ♀, non-type
E. aff. lupherorum LM 1 ♀, non-type (NHMD-921634)
Seamount 310 69 670 41.3353 176.1882 E. aff. balerioni LM 1 ♂, non-type (NHMD-921971)
E. aff. galadrielae/beringiensis LM 1 ♀, non-type (NHMD-921501)
E. aff. lupherorum LM 1 ♀, non-type (NHMD-921632)
72 985 41.3657 176.1958 E. legolasi sp. nov. LM ♂, holotype (NIWA-159422)
E. aff. lupherorum LM 1 ♂, non-type (NHMD-921633)
SEM 1 ♀, non-type
E. samwisei sp. nov. LM 1 ♀, paratype (NHMD-917302)
SEM 1 ♂, non-type
Seamount 766 129 1456 42.1345 174.5860 E. frodoi sp. nov. LM 1 ♂, paratype (NHMD-916334)
132 1453 42.1345 174.5850 E. galadrielae sp. nov. LM 1 ♀, paratype (NIWA-159426)
E. juliae LM 2 ♀♀, non-types (NHMD-921730, 9217032)
Table 1 (continued). Coordinates and basic data on sampling stations, and identities, type status and catalogue numbers of species of Echinoderes
Claparède, 1863.
European Journal of Taxonomy 844: 1–108 (2022)
8
detergent. Following this process, specimens were dehydrated through a graded water-alcohol and then
alcohol-acetone series, and nally critical point dried. Dried specimens were mounted on aluminium
stubs, sputter coated with a platinum/palladium mixture and examined with a JEOL JSM-6335F Field
Emission scanning electron microscope.
All specimens of Echinoderes were identied to species level, except for the juveniles. Identication
of kinorhynchs to species level was based on the relevant taxonomic literature (Sørensen et al. 2018;
Yamasaki et al. 2018b; Grzelak & Sørensen 2019) and the interactive identication key to species of
Echinoderidae (Yamasaki et al. 2020a).
Abbreviations
The following abbreviations are used for collections and museums:
INBRIV = National Institute of Biological Resources, Korea
NHMD = Natural History Museum of Denmark, Copenhagen, Denmark
NIWA = National Institute of Water and Atmospheric Research, Wellington
The following abbreviations are used in tables:
ac = acicular spine
fs = fringe-like structure
gco1/2 = glandular cell outlet type 1/2
LA = lateral accessory
LD = laterodorsal
ltas = lateral terminal accessory spine
lts = lateral terminal spine
LV = lateroventral
MD = middorsal
ML = midlateral
MSW-X = maximum sternal width, with X indicating segment with greatest sternal width
n/a = data non-applicable
pa = female papillae
PD = paradorsal
pe = penile spines
pr = protuberance
S = segment length
SD = subdorsal
si = sieve plate
SL = sublateral
so = slit-like opening
ss = sensory spot
SW-10 = standard width, always measured on segment 10
tbl = tubule
TL = trunk length
tu = tube
VL = ventrolateral
VM = ventromedial
* = character not present in all specimens
– = marks missing data
(♀) = female condition of sexually dimorphic character
(♂) = male condition of sexually dimorphic character
(♂/ ♀?) = character with uncertain gender afliation
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
9
The following abbreviations are used in gures:
fpa = female papillae
lagco2 = lateral accessory glandular cell outlet type 2
las = lateral accessory spine, eventually followed by spine number
lat = lateral accessory tube
ldgco2 = laterodorsal glandular cell outlet type 2
ldss = laterodorsal sensory spot
ldt = laterodorsal tube
ltas = lateral terminal accessory spine
lts = lateral terminal spine
lvgco2 = lateroventral glandular cell outlet type 2
lvs = lateroventral spine, eventually followed by spine number
lvt = lateroventral tube
mdgco1 = middorsal glandular cell outlet type 1
mds = middorsal spine, eventually followed by spine number
mdss = middorsal sensory spot
mlgco2 = midlateral glandular cell outlet type 2
mlso = midlateral slit-like opening
mlss = midlateral sensory spot
mlt = midlateral tube
mtj = midtergal junction
pdgco1/2 = paradorsal glandular cell outlet type 1/2
pdso = paradorsal slit-like opening
pdss = paradorsal sensory spot
pe = penile spines
pr = protuberance
pvgco1 = paraventral glandular cell outlet type 1
r-ldt = reduced laterodorsal tube
sdgco1/2 = subdorsal glandular cell outlet type 1/2
sdso = subdorsal slit-like opening
sdss = subdorsal sensory spot
sdt = subdorsal tube
si = sieve plate
slgco2 = sublateral glandular cell outlet type 2
slss = sublateral sensory spot
slt = sublateral tube
spf = secondary pectinate fringe
tbl = tubule
te = tergal extension
vlgco1/2 = ventrolateral glandular cell outlet type 1/2
vlss = ventrolateral sensory spot
vlt = ventrolateral tube
vmgco1 = ventromedial glandular cell outlet type 1
vmso = ventromedial slit-like opening
vmss = ventromedial sensory spot
* = character not present in all specimens
European Journal of Taxonomy 844: 1–108 (2022)
10
Results
Species descriptions
Class Cyclorhagida Zelinka, 1896 sensu Herranz et al. 2022
Order Echinorhagata Sørensen et al., 2015
Family Echinoderidae Carus, 1885
Genus Echinoderes Claparède, 1863
Echinoderes blazeji sp. nov.
urn:lsid:zoobank.org:act:38F17023-13B9-487F-AF98-9F44AE186815
Figs 2–4; Tables 2–3
Diagnosis
Echinoderes with a very minute spine in middorsal position on segment 4 and in lateral accessory
positions on segment 7. Tubes present in lateroventral positions on segment 5, sublateral positions on
segment 8 and laterodorsal positions on segment 10; tubes on segment 10 well-developed in males,
whereas much smaller in females. Glandular cell outlet type 2 present in midlateral positions on
segment 8. Large, elongate sieve plates located midlaterally on segment 9. Lateral terminal spines twice
as long in males as in females.
Etymology
The species is named after Blazej, the son of the rst author – for his love of all dragons.
Material examined
Holotype
NEW ZEALAND • ♂; Pahaua Canyon, stn TAN1004/27; 41.4983° S, 175.7043° E; 1013 m b.s.l.; Apr.
2010; NIWA TAN1004 Voyage; soft sediment; NIWA-159400. Mounted for LM in Fluoromount G on
HS slide.
Paratypes
NEW ZEALAND • 1 ♀; same collection data as for holotype; NHMD-917223 • 1 ♂; same collection
data as for holotype; NIWA-159401. Mounted as holotype.
Additional material
NEW ZEALAND • 1 ♂; same collection data as for holotype; personal reference collection of MVS.
Mounted for SEM • 3 ♀♀, 1 ♂; Honeycomb Canyon, stn TAN1004/58; 41.4080° S, 175.8977° E;
670 m b.s.l.; Apr. 2010; NIWA TAN1004 Voyage; soft sediment; personal reference collection of MVS.
Mounted for SEM • 1 ♂; Honeycomb Canyon, stn TAN1004/62; 41.4760° S, 175.9477° E; 1171 m b.s.l.;
Apr. 2010; NIWA TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted for
SEM.
Description
General. Adults with head, neck and eleven trunk segments (Figs 2–4). Overview of measurements
and dimensions in Table 2. Distribution of cuticular structures, i.e., sensory spots, glandular cell outlets,
spines and tubes, summarized in Table 3. The head morphology could not be examined in detail in any
of the available specimens.
neck. Consists of 16 placids. Midventral placid broadest, 11 µm in width and 12 µm in length, whereas
all others narrower, measuring 7 µm in width at bases (Fig. 2). Trichoscalid plates well developed
(Fig. 3B).
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
11
Fig. 2. Line art illustrations of Echinoderes blazeji sp. nov. A. ♂, dorsal view. B. ♂, ventral view. C. ♀,
segments 10–11, dorsal view. D. ♀, segments 10–11, ventral view. Abbreviations: see Material and
methods.
European Journal of Taxonomy 844: 1–108 (2022)
12
SeGment 1. Consists of complete cuticular ring. Subdorsal and laterodorsal sensory spots present, situated
on anterior half of segment. Sensory spots on this and following segments droplet-shaped, consisting
of central pore surrounded by micropapillae (Fig. 4C, E, G). Glandular cell outlet type 1 not observed.
Cuticular hairs arising from rounded perforation sites, distributed evenly around segment except in
anterior part. Segment terminates in pectinate fringe with relatively long fringe tips (Fig. 4C).
Fig. 3. Light micrographs showing overviews and details of Echinoderes blazeji sp. nov. A, G–H. ♂,
paratype (NIWA-159401). B–C, E–F. ♂, holotype (NIWA-159400). D, I. ♀, paratype (NHMD-
917223). A. Lateral overview of male. B. Segments 1 to 5, dorsal view. C. Segments 1 to 5, ventral
view. D. Lateral overview of female. E. Segments 6 to 9, dorsal view. F. Segments 7 to 11, ventral view.
G. Segments 8 to 9, lateroventral view. H. Segments 9 to 11 of male, lateral view. I. Segments 9 to 11
of female, lateral view. Abbreviations: see Material and methods.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
13
SeGment 2. Consists of complete cuticular ring with sensory spots present in middorsal, laterodorsal,
midlateral and ventromedial positions (Figs 2A–B, 3B, 4B–C). Glandular cell outlet type 1 present in
middorsal position and as a pair in ventromedial positions. Pachycyclus of anterior segment margin
of regular thickness, interrupted in middorsal position on this and following segments. Cuticular hairs
densely covering entire segment. Perforation sites on this and following eight segments appear as band
around segment, easily visible in LM (Fig. 3B–C, E–I). Posterior segment margin straight along dorsal
edge, but markedly extended posteriorly in midventral position (Fig. 4C). Primary pectinate fringe with
tips similar to those of preceding segment in middorsal to ventrolateral position and smaller and thinner
tips in ventromedial and paraventral positions (Figs 2A–B, 4B–C).
SeGment 3. Present segment, and eight remaining ones, consist of one tergal and two sternal plates
(Figs 2A–B, 3A–D). Sensory spots present in subdorsal and sublateral positions (Fig. 3B–C). Glandular
cell outlets type 1 located in middorsal and ventromedial positions. Cuticular hairs on this and following
seven segments densely covering entire segment, except for narrow area in laterodorsal position
(Fig. 2A). Posterior segment margin straight, terminating in pectinate fringe with relatively long and
uniform fringe tips along entire segment margin.
SeGment 4. With minute (~10 µm) middorsal spine (Figs 3B, 4E). Sensory spots located subdorsally
(Figs 3B, 4B). Glandular cell outlet type 1 present in subdorsal and ventromedial positions (Fig. 3B–C).
Segment otherwise as segment 3.
SeGment 5. With tubes in lateroventral positions (Figs 2B, 3C, 4F). Sensory spots present in subdorsal,
midlateral and ventromedial positions (Figs 2A–B, 4E–F). Glandular cell outlets type 1 present in
Character Holotype
NIWA-159400 (♂)
Paratype
NHMD-917223 (♀)
Paratype
NIWA-159401 (♂)
TL 261 318 325
MSW-6 55 n/a n/a
MSW-6/TL 21% n/a n/a
SW-10 46 n/a n/a
SW-10/TL 18% n/a n/a
S1 26 27 27
S2 23 29 23
S3 28 32 30
S4 31 40 35
S5 31 42 34
S6 35 44 39
S7 33 47 43
S8 36 48 48
S9 40 38 46
S10 41 39 41
S11 27 25 28
MD4 (ac) 10 – –
LA7 (ac) 6 – –
LTS 105 54 104
LTS/TL 40% 17% 32%
LTAS n/a 19 n/a
Table 2. Measurements from light microscopy of Echinoderes blazeji sp. nov. (in µm) from the Hikurangi
Margin.
European Journal of Taxonomy 844: 1–108 (2022)
14
Fig. 4. Scanning electron micrographs showing overviews and details of Echinoderes blazeji sp. nov.
A. Dorsolateral overview of female. B. Segments 1 to 4, dorsolateral view. C. Segments 1 to 3, ventral
view. D. Lateral overview of male. E. Segments 4 to 6, dorsal view. F. Segments 5 to 7, ventral view.
G. Segments 8 to 11 of female, laterodorsal view. H. Segments 7 to 8, lateral view. I. Segments 10 to 11
of male, dorsal view. J. Segments 10 to 11 of male, dorsolateral view. K. Segments 10 to 11 of female,
lateral view, with close-up of segment 11 ventral side showing ventrolateral tubules. Abbreviations: see
Material and methods.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
15
subdorsal and ventromedial positions. Cuticular hair covering as on preceding segment except for
hairless paraventral area. Secondary fringe and posterior segment margin as on preceding segment.
SeGment 6. With sensory spots present in subdorsal, sublateral and ventromedial positions (Figs 2A–B,
3E, 4E–F). Glandular cell outlets type 1 as on preceding segment. Cuticular hair covering and secondary
pectinate fringe as on segment 5.
SeGment 7. With minute spines (<10 µm) in lateral accessory positions (Figs 2B, 3F, 4F); spines hardly
visible in both LM and SEM due to dense cuticular hairs covering and their inconspicuous appearance.
Sensory spots located in subdorsal, midlateral, and ventromedial positions. Glandular cell outlets type 1
present in subdorsal and ventromedial positions. Segment otherwise as segment 6.
SeGment 8. With tubes in sublateral positions and glandular cell outlets type 2 located in midlateral
positions (Figs 2A–B, 3E–F, 4G–H). Sensory spots present in subdorsal, laterodorsal and ventromedial
positions; subdorsal pair located closer to paradorsal line than on preceding segments. Glandular cell
outlets type 1 present in subdorsal and ventromedial positions. Pectinate fringe tips slightly shorter than
on preceding segments.
SeGment 9. Without spines or tubes. Sensory spots located in subdorsal, laterodorsal, midlateral and
ventrolateral positions (Figs 2A–B, 4G). Pair of sieve plates, composed of large, elongated sieve area
located anterior to rounded areas with central pore, located in sublateral positions (Figs 2A, 3F–I).
Glandular cell outlets type 1, cuticular hair covering and posterior segment margin as on preceding
segment.
SeGment 10. With laterodorsal tubes, located near posterior segment margin. In males, tubes long
(~15 µm) (Figs 2A, 3H, 4I–J). In females, tubes much shorter (~6 µm) and more exible (Figs 2C, 4K),
without basal part characteristic for tubes in males as well as for tubes described on segments 5 and 8.
Sensory spots present in subdorsal and ventrolateral positions. Glandular cell outlets type 1 present as
middorsal one, and as pair in ventromedial positions. Posterior segment margin of tergal plate straight,
while margins of sternal plates concave, reaching posterior margin of terminal segment. Pectinate fringe
tips signicantly shorter and narrower than on preceding segment.
SeGment 11. With lateral terminal spines; in males, lateral terminal spines twice long as in females
(Figs 2, 3A, D, 4A, D; Table 2). Females with short and relatively thin lateral terminal accessory spines
(Figs 2C–D, 3I, 4K); males with three penile spines, two of them exible and elongated, one short and
Table 3. Summary of nature and location of sensory spots, glandular cell outlets, tubes and spines
arranged by series in Echinoderes blazeji sp. nov.
Position segment MD SD LD ML SL LA LV VL VM
1 ss ss
2 gco1,ss ss ss gco1,ss
3 gco1 ss ss gco1
4 ac gco1,ss gco1
5 gco1,ss ss tu gco1,ss
6 gco1,ss ss gco1,ss
7 gco1,ss ss ac gco1,ss
8 gco1,ss ss gco2 tu gco1,ss
9 gco1,ss ss ss si ss gco1
10 gco1 ss tu ss gco1
11 gco1 ss pe × 3 (♂) ltas(♀) lts tbl(♂/♀?)
European Journal of Taxonomy 844: 1–108 (2022)
16
stout (Figs 2A–B, 3H, 4I–J). Additionally, one female specimen shows pair of fringed tubes-like structures
on ventral side (Fig. 4K); examination of ventral side of segment 11 not possible in other specimens;
therefore, we cannot conclude whether it is a sexually dimorphic character or not. Sensory spots present
in subdorsal positions. Unpaired glandular cell outlet type 1 present middorsally. Segment devoid of
cuticular hairs, but very short cuticular hair-like structures covering tergal extensions and posterior
parts of sternal plates. Very short fringes covering margins of tergal and sternal plates. Tergal extensions
elongated and triangular (Figs 2A, C, 4J). Sternal extensions slightly extended posteriorly, not extending
beyond tergal extensions (Fig. 2B, D).
Distribution
Canyons: Pahaua, Honeycomb, 670–1171 m b.s.l. See Fig. 1 for a geographic overview of stations and
Table 1 for station and specimen information.
Taxonomic remarks on Echinoderes blazeji sp. nov.
The arrangement of spines and tubes, with a minute middorsal spine on segment 4, minute lateral
spines on segment 7 only and lateral tubes on segments 5 and 8, is not present in any other species of
Echinoderes. These characters, combined with the large sieve plates and dense cuticular hairs, make
E. blazeji sp. nov. even more distinctive and narrow the number of potential congeners down to species
belonging to the so-called Echinoderes coulli-group (following the diagnosis of Yamasaki & Fujimoto
2014). Currently, this group accommodates 17 species (Randsø et al. 2019; Yamasaki et al. 2020a;
Cepeda et al. 2022; Kennedy et al. 2022) that share a number of morphological characters and habitat
preferences.
Echinoderes blazeji sp. nov. can easily be distinguished from all other E. coulli-group congeners by its
presence of lateral spines only on segment 7. The group is suggested to share morphological features
such as absence of middorsal spines or, if present, on segment 4 only; lateral spines absent or very
minute and restricted to segments 6 and 7; presence of lateral tubes on segments 5 and 8; and female
lateral terminal accessory spines being either poorly-developed or absent (Yamasaki & Fujimoto 2014).
Therefore, the possession of only one pair of lateral spines makes E. blazeji unique among all other
species of this group. Nevertheless, it should be stressed that these spines are extremely minute and
might easily be overlooked due to the dense cuticular hair covering, especially during LM examination.
But even if the presence of lateral spines on segment 7 had gone unnoticed among the nine species of the
E. coulli group with a middorsal spine on segment 4 (i.e., E. annae Sørensen et al., 2016, E.cyaneactus
Cepeda et al., 2022, E. maxwelli (Omer-Cooper, 1957), E. ohtsukai Yamasaki & Kajihara, 2012, E.
parthenope Cepeda et al., 2022, E. regina Yamasaki, 2016, E. rex Lundbye et al., 2011, E. serratulus
Yamasaki, 2016 and E. teretis Brown, 1999 in Adrianov & Malakhov 1999), only E. annae shows
the absence of lateral spines (Omer-Cooper 1957; Adrianov & Malakhov 1999; Lundbye et al. 2011;
Yamasaki & Kajihara 2012; Sørensen et al. 2016a; Yamasaki 2016; Cepeda et al. 2022). However, other
conditions in E. annae make this species easily distinguishable from E. blazeji. In contrast to the new
species, which has only one pair of relatively big glandular cell outlets type 2 on segment 8, E. annae
possesses numbers of minute glands distributed over several segments. Moreover, the latter species is
characterized by the presence of midlateral tubes on segment 9, which are absent in E. blazeji, and by
very short and stout lateral terminal spines, which cannot be confused with the longer and thinner lateral
spines in E. blazeji.
Furthermore, the lateral terminal spines themselves seem to represent another characteristic feature for
the new species, since they are twice as long in males as in females (♂LTS = 104 µm vs ♀LTS = 50 µm,
respectively). Echinoderid sexual dimorphism is usually displayed in the female presence of lateral
terminal accessory spines, in the appearance of the laterodorsal tubes on segment 10 (i.e., Sørensen
2006; Pardos et al. 2016a; Grzelak & Sørensen 2018; present study) or presence of papillae/pores on the
ventral side in females (Sørensen et al. 2020). Having sexual dimorphism expressed in lateral terminal
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
17
spine lengths is a rather unusual trait in Echinoderes. Differences in length of lateral terminal spines
expressed as sexual dimorphism have been observed for E. aquilonius Higgins & Kristensen, 1988 and
E. lusitanicus Neves et al., 2016 (Higgins & Kristensen 1988; Neves et al. 2016), but more interestingly,
also in E. coulli Higgins, 1977 – a species closely related with E. blazeji sp. nov. Higgins (1977) described
two forms of females in E. coulli: one form with lateral terminal spines similar to those in males, and a
second form with short lateral terminal spines being half the length of those of males. The latter short-
spined form was, however, more abundant in the population and constituted about half of all examined
specimens. In our case, all examined females (4 out of 9 specimens in total) had markedly shorter lateral
terminal spines than males, which suggests that we might have the same kind of female dimorphism
in E. blazeji. Sexual dimorphism expressed in spine lengths has also been reported for E. levanderi
Karling, 1954 and the Arctic population of E. pterus Yamasaki et al., 2018, but in these cases it was
related to the length of lateroventral spines (Karling 1954; Sørensen 2018; Yamasaki et al. 2018a).
In addition, the pattern of glandular cell outlets type 1 on the dorsal side appears to be uncommon for
the new species. The taxonomic signicance of the glandular cell outlet type 1 pattern, in contrast to
glandular cell outlets type 2, is not yet well explored or understood. In fact, it was only quite recently
that Sørensen et al. (2020) drew attention to its potential taxonomic signicance. They showed that a
majority of species of Echinoderes (for which we have sufcient data) show outlets on segments 4 to
9 in paradorsal or paradorsal and middorsal (depends on the segment) positions (Sørensen et al. 2020).
The presence of glandular cell outlets type 1 in subdorsal positions on segments 4 to 9, as observed for
E. blazeji sp. nov., has so far only been reported from ten other, putatively closely related species, all
belonging to the E. dujardinii species group, and in E. worthingi Southern, 1914, a species also closely
related to the E. dujardinii group (Southern 1914; Sørensen et al. 2020). Since E. blazeji cannot be
considered as closely related with the E. dujardinii-group, our observation of glandular cell outlets
type 1 in the new species thus indicates that this morphological trait may still hide some interesting
aspects.
Finally, the new species has been found in a habitat that is quite unusual for species of the E. coulli-
group. The majority of these species have been recorded in intertidal marine or brackish water, with
the exception of four species that inhabit subtidal, but yet shallow, marine waters (Lundbye et al. 2011;
Yamasaki 2016; Kennedy et al. 2022). In this context, E. blazeji sp. nov. is unique within the species
group, since it is so far the only species inhabiting deep-sea waters.
Echinoderes landersi sp. nov.
urn:lsid:zoobank.org:act:3F3C572F-E5CE-4645-AA8F-9472E698B2F4
Figs 5–7; Tables 4–5
Diagnosis
Echinoderes with spines in middorsal position on segments 4 and 6, and in lateroventral positions on
segments 6 to 9. Tubes present in subdorsal, laterodorsal, sublateral and ventrolateral positions on
segment 2, lateroventral positions on segment 5, subdorsal, midlateral and lateral accessory positions on
segment 8, and midlateral positions on segments 9 and 10.
Etymology
The species is named after Dr Stephen C. Landers in recognition of his contributions to kinorhynch
taxonomy and ecology.
European Journal of Taxonomy 844: 1–108 (2022)
18
Material examined
Holotype
NEW ZEALAND • ♀; Hikurangi Slope, stn TAN1004/38; 41.5937° S, 175.8532° E; 1121 m b.s.l.; Apr.
2010; NIWA TAN1004 Voyage; soft sediment; NIWA-159402. Mounted for LM in Fluoromount G on
HS slide.
Paratype
NEW ZEALAND • 1 ♀; Pahaua Canyon, stn TAN1004/31; 41.4962° S, 175.6828° E; 730 m b.s.l.; Apr.
2010; NIWA TAN1004 Voyage; soft sediment; NHMD-916627. Mounted as holotype.
Additional material
NEW ZEALAND • 1 ♀; Hikurangi Slope, stn TAN1004/44; 41.5258° S, 175.8003° E; 728 m b.s.l.; Apr.
2010; NIWA TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted for SEM
• 2 ♀♀; Pahaua Canyon, stn TAN1004/27; 41.4983° S, 175.7043° E; 1013 m b.s.l.; Apr. 2010; NIWA
TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted for SEM.
Description
General. Adults with head, neck and eleven trunk segments (Figs 5–7). Overview of measurements
and dimensions in Table 4. Distribution of cuticular structures, i.e., sensory spots, glandular cell outlets,
spines and tubes, summarized in Table 5. No details regarding scalid arrangement and morphology
could be provided, because introverts of all specimens mounted for SEM fully retracted.
neck. With 16 placids. Midventral placid broadest, 11 µm in width and 16 µm in length, whereas all
others narrower, measuring 7 µm in width at their bases (Fig. 5). Trichoscalid plates well developed
(Fig. 6B).
SeGment 1. Consists of complete cuticular ring. Sensory spots located on anterior half of segment, in
subdorsal, laterodorsal and ventromedial positions; sensory spots on this and following segment with
micropapillae surrounding central pore and long marginal hair (Figs 5A–B, 7C). Glandular cell outlet
type 1 present in middorsal and ventrolateral positions. Cuticular hairs relatively long, distributed evenly
around segment. Posterior segment margin almost straight, forming pectinate fringe with very short,
sawtooth-like fringe tips along dorsal margin and with slightly longer tips along ventral margin.
SeGment 2. Consists of complete cuticular ring, with tubes located in subdorsal, laterodorsal, sublateral
and ventrolateral positions (Figs 5A–B, 6A–B, 7C–D); in one specimen right subdorsal tube missing.
Sensory spots present in middorsal, laterodorsal and ventromedial positions. Glandular cell outlets type 1
not observed. This structure better visible in LM than in SEM but none of LM specimens orientated
in way that allowed detailed examination of segments, especially its ventral side; therefore, for this
and following nine segments presence of glandular cell outlets type 1 in ventral positions can neither
be conrmed nor rejected. Pachycyclus of anterior segment margin of regular thickness. Secondary
pectinate fringe present near anterior segment margin of this and following segments, but usually
covered by preceding segment. Cuticular hairs and pectinate fringe tips as on preceding segment.
SeGment 3. Present segment, and eight remaining ones, consist of one tergal and two sternal plates
(Figs 5A–B, 7D). Segment with subdorsal and midlateral sensory spots. On this and following six
segments, cuticular hairs arranged in three or four rows across tergal plate, except for hairless laterodorsal
areas; paraventral areas devoid of hairs on this and following seven segments (Fig. 5). Posterior
segment margin straight, terminating in pectinate fringe with longer and more slender fringe tips along
dorsal margin than on preceding segments, otherwise as on preceding segment.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
19
Fig. 5. Line art illustrations of Echinoderes landersi sp. nov. A. ♀, dorsal view. B. ♀, ventral view.
Abbreviations: see Material and methods.
European Journal of Taxonomy 844: 1–108 (2022)
20
Fig. 6. Light micrographs showing overview and details of Echinoderes landersi sp. nov. A. ♀,
paratype (NHMD-916627). B–D. ♀, holotype (NIWA-159402). A. Lateral overview. B. Segments 1 to
3, lateral view. C. Segments 7 to 11, lateral view. D. Segments 4 to 7, lateral view. Abbreviations: see
Material and methods.
Table 4. Measurements from light microscopy of Echinoderes landersi sp. nov. (in µm) from the
Hikurangi Margin.
Character Holotype
NIWA-159402 (♀)
Paratype
NHMD-916627 (♀)
TL 333 351
S1 27 24
S2 34 21
S3 40 37
S4 42 38
S5 47 42
S6 52 49
S7 54 50
S8 49 44
S9 44 40
S10 43 40
S11 42 39
MD4 (ac) 31 24
MD6 (ac) 33 30
LV6 (ac) 28 24
LV7 (ac) 29 25
LV8 (ac) 34 29
LV9 (ac) 38 33
LTS 198 173
LTS/TL 59.6% 49.3%
LTAS 70 60
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
21
SeGment 4. With spine in middorsal position. Spine relatively short (27 µm), only slightly exceeding
beyond posterior segment margin (Figs 5A, 6A, D, 7A, E). Glandular cell outlets type 1 present in
paradorsal positions. No other traits observed. Cuticular hairs and secondary pectinate fringe as on
preceding segment.
SeGment 5. With tubes in lateroventral positions (Figs 5B, 7D). Sensory spots present in subdorsal,
midlateral and ventromedial positions (Figs 5A–B, 6D, 7D–E). Glandular cell outlets type 1 present
in paradorsal positions. Tips of pectinate fringe of posterior segment margin slightly longer than on
preceding segment. Cuticular hairs as on preceding segment.
SeGment 6. With spines in middorsal and lateroventral positions (Fig. 5A–B). Middorsal spine, as on
segment 4, relatively short (31 µm), only slightly exceeding beyond posterior segment margin (Figs 6A,
D, 7A, E). Sensory spots present in paradorsal, subdorsal, midlateral and ventromedial positions
(Figs 5A–B, 6D, 7D–E). Glandular cell outlets type 1 present in paradorsal positions (Figs 5A, 6D).
Pectinate fringe of posterior segment margin and cuticular hairs as on preceding segment.
SeGment 7. With spines in lateroventral positions, and sensory spots in subdorsal, midlateral and
ventromedial positions (Figs 5A–B, 6C–D, 7E). Glandular cell outlets type 1 present in paradorsal
positions. Tips of pectinate fringe of posterior segment margin slightly longer than on preceding
segments. Segment otherwise as segment 6.
SeGment 8. With spines in lateroventral positions, and tubes in subdorsal, midlateral and lateral accessory
positions (Figs 5A–B, 6C, 7G–H). Sensory spots present in subdorsal positions only. Glandular cell
outlets type 1 present in paradorsal positions. Pectinate fringe of posterior segment margin and cuticular
hairs as on preceding segment.
SeGment 9. With spines in lateroventral positions and tubes in midlateral positions (Figs 5A–B, 6C,
7G–H). In one specimen additional tube on right side of segment located in subdorsal position (Fig. 7I).
Two pairs of sensory spots located in subdorsal positions and one ventrolateral pair (Figs 5A–B, 7H).
Glandular cell outlets type 1 present in paradorsal positions. Small, rounded sieve plates located in lateral
accessory positions (Fig. 6C). Cuticular hair covering and pectinate fringe of posterior segment margin
as on preceding segment.
Table 5. Summary of nature and location of sensory spots, glandular cell outlets, tubes and spines
arranged by series in Echinoderes landersi sp. nov.
Position segment MD PD SD LD ML SL L A LV VL VM
1 gco1 ss ss gco1 ss
2 ss tu tu,ss tu tu ss
3 ss ss
4ac gco1
5 gco1 ss ss tu ss
6 ac gco1,ss ss ss ac ss
7 gco1 ss ss ac ss
8 gco1 ss,tu tu tu ac
9 gco1 ss,ss tu si ac ss
10 gco1 ss tu ss
11 ss,gco1 ltas(♀) lts
European Journal of Taxonomy 844: 1–108 (2022)
22
Fig. 7. Scanning electron micrographs showing overviews and details of Echinoderes landersi sp. nov.
A. Dorsal overview. B. Lateral overview. C. Segments 1 to 3, lateral view. D. Segments 2 to 6, ventral
view. E. Segments 4 to 7, dorsolateral view. F. Segments 1 to 3 of female with inconsistent tube pattern,
dorsal view. G. Segments 8 to 10, lateral view. H. Segments 8 to 11, dorsal view, with inset showing
close-up of midlateral tube on segment 10. I. Segments 8 to 9 of female with inconsistent tube pattern,
dorsal view. Abbreviations: see Material and methods.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
23
SeGment 10. With minute midlateral tubes located near posterior segment margin (Figs 5A, 6C, 7H).
Sensory spots present in subdorsal and ventrolateral positions (Figs 5A–B, 7H); in one specimen extra
subdorsal sensory spot present on left side of segment (Fig. 7H). Glandular cell outlet type 1 present in
middorsal position. Cuticular hairs less dense on dorsal side than on preceding segment. Central part
of tergal plate devoid of hairs. Hairs on sternal plates shorter than on preceding segments. Posterior
segment margin of tergal plate straight, without fringe tips; margins of sternal plates concave, reaching
posterior margin of terminal segment, with very short fringe tips.
SeGment 11. With lateral terminal spines (Figs 5A–B, 6A). Females with lateral terminal accessory spines
(Figs 5A–B, 6A, 7H); male conditions unknown. Sensory spots and glandular cell outlets type 1 present
in subdorsal positions; in one specimen sensory spot on right side of segment missing. Segment devoid
of cuticular hairs, but with very short cuticular hair-like structures covering paradorsal area and very
short fringes covering margins of tergal and sternal plates. Tergal extensions triangular, with elongate
and pointed tips (Figs 5A, 7H). Sternal extensions do not extend beyond tergal extensions (Fig. 5B).
Distribution
Hikurangi slope and Pahaua Canyon, 728–1121 m b.s.l. See Fig. 1 for a geographic overview of stations
and Table 1 for station and specimen information.
Taxonomic remarks on Echinoderes landersi sp. nov.
Echinoderes landersi sp. nov. is easily distinguished from all other congeners by its combination of
spines and tubes. The spine pattern on its dorsal side, with middorsal spines on segments 4 and 6,
is already an uncommon feature within Echinoderes. This trait itself is shared only with E. astridae
Sørensen, 2014, E. bispinosus Higgins, 1982, E. uozumii Yamasaki et al., 2020 and E. dalzottoi sp. nov.
described below in the present study (Higgins 1982; Sørensen 2014; Yamasaki et al. 2020b). This
character in combination with its tubes on segment 2 makes E. landersi unique among its congeners.
The abovementioned species possess either three pairs of tubes as in E. dalzottoi or only one pair as in
E. astridae, E. bispinosus and E. uozumii. Therefore, having four pairs of tubes on segment 2, located
in subdorsal, laterodorsal, sublateral and ventrolateral positions, is uncommon and found exclusively
in E. landersi. The three latter species furthermore differ from E. landersi by being equipped with
glandular cell outlets type 2; such outlets are absent in E. landersi. The regular hair covering found in
E. landersi also distinguishes it from E. dalzottoi, which is characterized by having a trunk cuticle with
perforation sites only but no cuticular hairs.
In addition, E. landersi sp. nov. can easily be distinguished from these species by other traits, among
which the composition of segment 8 with tubes in subdorsal, midlateral and lateral accessory positions
is the most important, as these are not present in any other species of Echinoderes. Numerous species
have various combinations of tubes on this particular segment, but no other species has three pairs of
tubes. There are several species with two pairs of tubes, in laterodorsal and lateral accessory positions
as observed in, e.g., E. abbreviatus Higgins, 1983, E. belenae Pardos et al., 2016 or E. intermedius
Sørensen, 2006, or in subdorsal and lateral accessory positions as found in E. capitatus (Zelinka,
1928) and E. isabelae GaOrdóñez et al., 2008 (Zelinka 1928; Higgins 1983; GaOrdóñez et al. 2008;
Sørensen 2006; Pardos et al. 2016b); nevertheless, the most common tube pattern is the presence of
only one pair of tubes on segment 8, typically in a sublateral or lateral accessory position. Furthermore,
E. landersi possesses midlateral tubes on segment 9, which is another relatively rare trait, shared with
only four species, i.e., E. andamanensis Higgins & Rao, 1979, E. annae, E. newcaledoniensis Higgins,
1967 and E. serratulus (Higgins 1967; Higgins & Rao 1979; Sørensen et al. 2016a; Yamasaki 2016).
Nevertheless, none of these species can in any way be confused with E. landersi due to their signicantly
different middorsal spine patterns. Hence, besides its uncommon middorsal spine pattern with spines on
segments 4 and 6 only, the new species is very easily recognized by the nature of its cuticular structures
of segments 2, 8 and 9.
European Journal of Taxonomy 844: 1–108 (2022)
24
One noteworthy observation in E. landersi sp. nov. is the inconsistent tube pattern found in one
specimen (Fig. 7F, I). However, in contrast to recent observations of intraspecic variation in species of
Echinoderes where pairs of tubes may be present or absent, it was only one tube missing from the pair in
E. landersi. Variation in tube patterns, regarding their presence or absence, has been documented, e.g.,
in E. daenerysae Grzelak & Sørensen, 2017 in Grzelak & Sørensen (2018) for ventrolateral tubes on
segment 2, in E. eximus Higgins & Kristensen, 1988 for sublateral tubes on segment 9, in E. levanderi
for subdorsal tubes on segment 2, and in E. frodoi sp. nov. (present study) for midlateral tubes on
segment 1 (Grzelak & Sørensen 2018; Sørensen 2018). Among specimens of E. landersi, however,
one specimen lacks the subdorsal tube on the right side on segment 2 (Fig. 7F), but has a subdorsal
tube on the right side of segment 9 (Fig. 7I). Tubes in subdorsal positions on segment 9 were not
observed in other specimens. A similar inconsistency was described by Yamasaki & Dal Zotto (2019)
for specimens of E. capitatus that lack a ventrolateral tube on one side on segment 2 (Yamasaki & Dal
Zotto 2019). Moreover, we observed that one specimen lacks a subdorsal sensory spot on the right side
of segment 10, and that this lacking sensory spot seems to be relocated to segment 9 (Fig. 7H). For
now, we cannot explain the reason for this observed variation, but it seems most likely that it is due to
individual abnormalities, rather than intraspecic variation.
Echinoderes dalzottoi sp. nov.
urn:lsid:zoobank.org:act:CFE372C3-2BAF-4A6B-B3C4-A0A14551747F
Figs 8–10; Tables 6–7
Diagnosis
Echinoderes with spines in middorsal position on segments 4 and 6, and spines in lateroventral positions
on segments 6 to 9. Tubes present in subdorsal, sublateral (might be missing in some specimens) and
ventrolateral positions on segment 2, lateroventral positions on segment 5, sublateral positions on
segment 8, and laterodorsal positions on segment 9. Sexually dimorphic tubes furthermore present in
laterodorsal positions on segment 10 in males; females with fringe-like structure in midlateral positions.
Minute scales present on segments 2 to 10, but regular cuticular hairs absent throughout trunk
Etymology
The species is named after Dr Matteo Dal Zotto in recognition of his contributions to kinorhynch
taxonomy and ecology.
Material examined
Holotype
NEW ZEALAND • ♀; Pahaua Canyon, stn TAN1004/31; 41.4962° S, 175.6828° E; 730 m b.s.l.; Apr.
2010; NIWA TAN1004 Voyage; soft sediment; NIWA-159403. Mounted for LM in Fluoromount G on
HS slide.
Paratypes
NEW ZEALAND • 1 ♂; Pahaua Canyon, stn TAN1004/27; 41.4983° S, 175.7043° E; 1013 m b.s.l.; Apr.
2010; NIWA TAN1004 Voyage; soft sediment; NHMD-917147. Mounted as holotype • 1 ♂; Honeycomb
Canyon, stn TAN1004/58; 41.4080° S, 175.8977° E; 670 m b.s.l.; Apr. 2010; NIWA TAN1004 Voyage;
soft sediment; NIWA-159404. Mounted as holotype.
Additional material
NEW ZEALAND • 1 ♂; same collection data as for holotype; personal reference collection of MVS.
Mounted for SEM • 1 ♂; Pahaua Canyon, stn TAN1004/27; 41.4983° S, 175.7043° E; 1013 m b.s.l.;
Apr. 2010; NIWA TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted for
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
25
Fig. 8. Line art illustrations of Echinoderes dalzottoi sp. nov. A. ♀, dorsal view. B. ♀, ventral view.
C. ♂, segments 10–11, dorsal view. D. ♂, segments 10–11, ventral view. Abbreviations: see Material
and methods.
European Journal of Taxonomy 844: 1–108 (2022)
26
SEM • 1 ♀; Honeycomb Canyon, stn TAN1004/58; 41.4080° S, 175.8977° E; 670 m b.s.l.; Apr. 2010;
NIWA TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted for SEM.
Description
General. Adults with head, neck and eleven trunk segments (Figs 8–10). Overview of measurements
and dimensions in Table 6. Distribution of cuticular structures, i.e., sensory spots, glandular cell outlets,
spines and tubes, summarized in Table 7. Head morphology could not be examined in detail in any of
available specimens.
neck. Consists of 16 placids. Midventral placid broadest, 13 µm in width and 15 µm in length, whereas
all others narrower, measuring 7 µm in width at their bases (Fig. 8). The trichoscalid plates are well
developed (Fig. 9D).
SeGment 1. Consists of complete cuticular ring. Sensory spots present in subdorsal, laterodorsal and
ventromedial positions. Sensory spots relatively large and without marginal hairs, located on anterior
half of segment (Figs 8A–B, 9C–D, 10B–C, E). Glandular cell outlet type 1 present in middorsal position
and in ventrolateral positions. Cuticular hairs or perforation sites not present. Posterior segment margin
almost straight, forming pectinate fringe with short, sawtooth-like fringe tips (Fig. 10B–C).
Table 6. Measurements from light microscopy of Echinoderes dalzottoi sp. nov. (in µm) from the
Hikurangi Margin, including number of measured specimens (n) and standard deviation (SD).
Character n Range Mean SD
TL 3 209–310 267 52.0
MSW-6 3 54–66 60 8.8
MSW-6/TL 3 23–25% 25% 1.5%
SW-10 3 40–46 43 4.5
SW-10/TL 3 16–19% 18% 1.9%
S1 3 24–29 26 2.5
S2 3 22–29 26 3.5
S3 3 23–35 27 7.4
S4 3 26–42 32 8.9
S5 3 28–46 35 10.0
S6 3 27–47 36 10.7
S7 3 30–50 39 11.2
S8 3 34–50 40 9.7
S9 3 34–50 43 8.3
S10 3 36–46 43 6.2
S11 1 20 n/a n/a
MD4 (ac) 3 43–62 51 10.1
MD6 (ac) 2 88–110 99 15.8
LV6 (ac) 3 24–29 27 2.4
LV7 (ac) 3 27–30 29 1.8
LV8 (ac) 3 28–34 31 2.7
LV9 (ac) 3 27–32 30 2.6
LTS 3 174–243 213 35.5
LTS/TL 3 78–83% 80% 2.6%
LTAS 1 40 n/a n/a
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
27
SeGment 2. Consists of complete cuticular ring, with tubes located in subdorsal, sublateral and
ventrolateral positions (Figs 8A–B, 9C–D, 10B–C, E); sublateral tubes missing in one paratype and two
SEM specimens; no sexual or developmental differences explain presence or absence of tubes. Sensory
spots of similar sizes as on preceding segment, present in middorsal, laterodorsal and ventromedial
positions; ventromedial ones with long marginal hair. Unpaired glandular cell outlet type 1 present in
middorsal position and as pair in ventromedial positions. Pachycyclus of anterior segment margin of
regular thickness, interrupted in middorsal position. Secondary pectinate fringe present near anterior
segment margin of this and following segments, but usually covered by preceding segment. This and
following nine segments completely hairless. Cuticular hairs reduced to minute scales distributed
Fig. 9. Light micrographs showing overview and details of Echinoderes dalzottoi sp. nov. A, C–D,
F–G, J–K. ♀, holotype (NIWA-159403). B, E, H–I. ♂, paratype (NHMD-917147). A. Ventral
overview. B. Lateral overview. C. Segments 1 to 6, dorsolateral view. D. Segments 1 to 6, lateroventral
view. E. Segments 8 to 11, lateral view. F. Segments 6 to 8, dorsolateral view. G. Segments 6 to 8,
lateroventral view. H. Segments 10 to 11 of male, dorsal view. I. Segments 10 to 11 of female, dorsal
view. J. Segments 8 to 10, dorsal view. K. Segments 8 to 11, ventral view. Abbreviations: see Material
and methods.
European Journal of Taxonomy 844: 1–108 (2022)
28
Fig. 10. Scanning electron micrographs showing overviews and details of Echinoderes dalzottoi sp. nov.
A. Dorsolateral overview. B. Segments 1 to 2 of specimen with (broken) sublateral tube on segment 2,
dorsolateral view. C. Segments 1 to 3 of specimen without sublateral tube on segment 2, dorsolateral
view. D. Ventral overview. E. Segments 1 to 3, ventral view. F. Segments 4 to 7, dorsolateral view with
inset showing close-up of cuticular hairs that are reduced to minute scales. G. Segments 5 to 8, ventral
view. H. Segments 5 to 9, lateroventral view. I. Segments 6 to 9, dorsolateral view. J. Segments 10 to
11 of female, dorsal view with inset showing close-up of midlateral fringe-like structure on segment 10.
K. Segments 10 to 11 of male, dorsal view. Abbreviations: see Material and methods.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
29
around segment (Fig. 10B–C, E–H), emerging through perforation sites; perforation sites easily visible
in LM (Fig. 9C–K). Posterior segment margin almost straight, but with rounded midventral extension
(Fig. 10D–E); pectinate fringe tips as on preceding segment, except midventral area with slightly
narrower fringe tips.
SeGment 3. Present segment, and eight remaining ones, consist of one tergal and two sternal plates
(Figs 8A–B, 9A, D). Sensory spots present in subdorsal and midlateral positions. Sensory spots on this
and following segments smaller than on preceding segments, all with one long marginal hair. Glandular
cell outlets type 1 as on preceding segment. Perforation sites appear as band around segment, interrupted
in middorsal and laterodorsal areas and in central part of sternal plate on this and following ve
segments (Figs 8A–B, 9C–D, F–G, 10F–G). Posterior segment margin straight, terminating in pectinate
fringe with slightly more slender fringe tips along ventral margin than on preceding segments, otherwise
as on preceding segment.
SeGment 4. With spine in middorsal position; spine relatively long (51 µm), reaching posterior margin
of segment 5 (Figs 8A, 10A, F). Glandular cell outlets type 1 present in paradorsal and ventromedial
positions. No other traits observed. Segment otherwise as segment 3.
SeGment 5. With tubes in lateroventral positions (Figs 8B, 9D, 10G). Sensory spots present in subdorsal,
midlateral and ventromedial positions (Figs 8A–B, 9C, 10F). Glandular cell outlets type 1 present in
middorsal and ventromedial positions. Perforation sites, secondary fringe and posterior segment margin
as on preceding segment.
SeGment 6. With spines in middorsal and lateroventral positions (Fig. 8A–B). Middorsal spine, as
on segment 4, relatively long (99 µm), reaching well beyond posterior segment margin of segment 8
(Fig. 10A, I). Sensory spots present in paradorsal, subdorsal, midlateral and ventromedial positions
(Figs 8A–B, 10F–G). Glandular cell outlets type 1 present in paradorsal and ventromedial positions.
Segment otherwise as segment 5.
SeGment 7. With spines in lateroventral positions, and sensory spots in paradorsal, midlateral and
ventromedial positions (Figs 8A–B, 9G, 10F–G). Glandular cell outlets type 1 present in middorsal
position and as pair in ventromedial positions. Tips of pectinate fringe of posterior segment margin
slightly shorter and more slender than on preceding segments. Segment otherwise as segment 6.
Table 7. Summary of nature and location of sensory spots, glandular cell outlets, tubes and spines
arranged by series in Echinoderes dalzottoi sp. nov.
Position segment MD PD SD LD ML SL L A LV VL VM
1 gco1 ss ss gco1 ss
2 gco1,ss tu ss tu* tu gco1,ss
3 gco1 ss ss gco1
4 ac gco1 gco1
5 gco1 ss ss tu gco1,ss
6 ac gco1,ss ss ss ac gco1,ss
7 gco1 ss ss ac gco1,ss
8 gco1,ss ss tu ac gco1
9 gco1,ss ss tu si ac ss gco1
10 gco1 ss tu(♂) fs(♀) ss gco1
11 gco1 ss pe × 3 (♂) ltas(♀) lts
European Journal of Taxonomy 844: 1–108 (2022)
30
SeGment 8. With spines in lateroventral positions, and tubes in sublateral positions (Figs 8A–B, 9E,
10H). Sensory spots present in paradorsal and subdorsal positions. Glandular cell outlets type 1 present
in paradorsal and ventromedial positions. Band of perforation site patches interrupted in subdorsal area
instead of laterodorsally as on preceding segments. Segment otherwise as segment 7.
SeGment 9. With spines in lateroventral positions. Tubes present in laterodorsal positions, but very close
to midlateral line (Figs 8A, 9E, 10H–I). Sensory spots located in paradorsal, subdorsal and ventrolateral
positions (Figs 8A–B, 9J, 10I). Glandular cell outlets type 1 present paradorsally and ventromedially.
Small, rounded sieve plates located in sublateral positions (Fig. 9K). Perforation sites, secondary fringe
and posterior segment margin as on preceding segment.
SeGment 10. With well-developed laterodorsal tubes, present in males only, located near posterior
segment margin and close to midlateral line (Figs 8C, 9H, 10K). Females without tubes, but with fringe-
like structures present in midlateral positions (Figs 8A, 10J). Sensory spots present in subdorsal and
ventrolateral positions (Figs 8A–B, 10J–K); subdorsal pair located rather close to paradorsal area.
Glandular cell outlets type 1 present as middorsal one, and in ventromedial positions. Perforation sites
restricted to paradorsal area, lateral sides of tergal plate, and lateral halves of sternal plates. Posterior
segment margin of tergal plate straight, without fringe tips (Fig. 10J–K); margins of sternal plates
concave, reaching posterior margin of terminal segment, with short and narrow fringe tips.
SeGment 11. With lateral terminal spines (Figs 8A–B, 9A, 10D). Females with lateral terminal accessory
spines (Figs 8A–B, 9I, 10A, D); males with three penile spines (Fig. 10K). Dorsal and ventral penile
spines slender and tubular, with dorsal ones much longer than ventral; median spines very stout, cone-
shaped (Figs 8C, 10K). Sensory spots present in subdorsal positions. Glandular cell outlet type 1 present
middorsally. Segment devoid of characteristic perforation site patches, but very short cuticular hair-like
structures covering paradorsal area. Very short fringes covering margins of tergal and sternal plates.
Tergal extensions triangular (Figs 8A, C, 10J). Sternal extensions rounded, not extending beyond tergal
extensions (Figs 8B, D, 9K).
Distribution
Canyons: Pahaua, Honeycomb, 670–1013 m b.s.l. See Fig. 1 for a geographic overview of stations and
Table 1 for station and specimen information.
Taxonomic remarks on Echinoderes dalzottoi sp. nov.
See taxonomic remarks for E. dalzottoi sp. nov. below, together with remarks for E. leduci sp. nov.
Echinoderes leduci sp. nov.
urn:lsid:zoobank.org:act:B0BC5E3C-0F52-4854-B0BD-D80712B6EC30
Figs 11–13; Tables 8–9
Diagnosis
Echinoderes with spines in middorsal position on segments 4, 6 and 8 and spines in lateroventral
positions on segments 6 to 9. Tubes present in laterodorsal and ventrolateral positions on segment 2,
lateroventral positions on segment 5, sublateral positions on segment 8, and laterodorsal positions on
segment 9. Sexually dimorphic tubes furthermore present in laterodorsal positions on segment 10 in
males. Minute scales present on segments 2 to 10, but regular cuticular hairs absent throughout trunk.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
31
Etymology
The species is named after Dr Daniel Leduc in recognition of his contribution in collecting Hikurangi
Margin kinorhynchs and making them available for description to the authors of the present study.
Material examined
Holotype
NEW ZEALAND • ♂; Pahaua Canyon, stn TAN1004/31; 41.4962° S, 175.6828° E; 730 m b.s.l.; Apr.
2010; NIWA TAN1004 Voyage; soft sediment; NIWA-159405. Mounted for LM in Fluoromount G on
HS slide.
Paratype
NEW ZEALAND • 1 ♀; Honeycomb Canyon, stn TAN1004/58; 41.4080° S, 175.8977° E; 670 m b.s.l.;
Apr. 2010; NIWA TAN1004 Voyage; soft sediment; NIWA-159406. Mounted as holotype.
Additional material
NEW ZEALAND • 1 ♂; Pahaua Canyon, stn TAN1004/27; 41.4983° S, 175.7043° E; 1013 m b.s.l.;
Apr. 2010; NIWA TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted for
SEM • 1 ♂; same collection data as for paratype; personal reference collection of MVS. Mounted for
SEM.
Description
General. Adults with head, neck and eleven trunk segments (Figs 11–13). Overview of measurements
and dimensions in Table 8. Distribution of cuticular structures, i.e., sensory spots, glandular cell outlets,
spines and tubes, summarized in Table 9. Head morphology could not be examined in detail in any of
available specimens.
neck. Consists of 16 placids. Midventral placid broadest, 12 µm in width and 14 µm in length, whereas
all others narrower, measuring 7 µm in width at their bases (Fig. 11). Trichoscalid plates well developed.
SeGment 1. Consists of complete cuticular ring. Sensory spots present in subdorsal, laterodorsal and
ventromedial positions. Sensory spots relatively large with marginal hairs, located on anterior half of
segment (Figs 11A–B, 12B–C, 13B–C). Glandular cell outlet type 1 present in middorsal position and in
ventrolateral positions. Cuticular hairs or perforation sites not present. Posterior segment margin almost
straight, forming pectinate fringe with short, sawtooth-like fringe tips (Fig. 13B–C).
SeGment 2. Consists of complete cuticular ring, with tubes located in laterodorsal and ventrolateral
positions (Figs 11A–B, 12B–C, 13B–C). Sensory spots of similar sizes as on preceding segment with long
marginal hair, present in middorsal, laterodorsal and ventromedial positions. Glandular cell outlet type 1
present in middorsal position and in ventromedial positions. Pachycyclus of anterior segment margin
of regular thickness, interrupted in middorsal position. Secondary pectinate fringe present near anterior
segment margin of this and following segments, but mostly covered by preceding segment. This and
following nine segments completely hairless. Cuticular hairs reduced to minute scales distributed
around segment (Fig. 13D–G), emerging through perforation sites; perforation sites easily visible in LM
(Fig. 12A–H). Posterior segment margin almost straight (Fig. 13C); pectinate fringe tips as on preceding
segment, except midventral area with slightly narrower fringe tips.
SeGment 3. Present segment, and eight remaining ones, consist of one tergal and two sternal plates
(Figs 11A–B, 12C, E). Sensory spots present in subdorsal and midlateral positions. Sensory spots on this
and following segments slightly smaller than on preceding segments, but still with one long marginal
hair. Glandular cell outlets type 1 present in middorsal and in ventromedial positions. Perforation sites
European Journal of Taxonomy 844: 1–108 (2022)
32
Fig. 11. Line art illustrations of Echinoderes leduci sp. nov. A. ♂, dorsal view. B. ♂, ventral view. C. ♀,
segments 10–11, dorsal view. D. ♀, segments 10–11, ventral view. Abbreviations: see Material and
methods.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
33
appear as band around segment, interrupted in middorsal and laterodorsal areas and in central part of
sternal plate on this and following ve segments (Figs 11A–B, 12B–E). Secondary fringe and posterior
segment margin as on preceding segment.
SeGment 4. With spine in middorsal position (Figs 11A, 12B, 13D). Glandular cell outlets type 1 present
in paradorsal and ventromedial positions. No other traits observed. Posterior segment margin straight,
terminating in pectinate fringe with slightly slenderer fringe tips along ventral margin than on preceding
segments, otherwise as on preceding segment.
SeGment 5. With tubes in lateroventral positions (Figs 11B, 12C, 13F). Sensory spots present in
subdorsal, midlateral and ventromedial positions (Figs 11A–B, 12B–C, 13D, F). Glandular cell outlets
type 1 present in middorsal and ventromedial positions. Perforation sites, secondary fringe and posterior
segment margin as on preceding segment.
SeGment 6. With spines in middorsal and lateroventral positions (Fig. 11A–B). Sensory spots present in
paradorsal, midlateral and ventromedial positions (Figs 11A–B, 13D, F). Glandular cell outlets type 1
present in paradorsal and ventromedial positions. Segment otherwise as segment 5.
Table 8. Measurements from light microscopy of Echinoderes leduci sp. nov. (in µm) from the Hikurangi
Margin.
Character Holotype
NIWA-159405 (♂)
Paratype
NIWA-159406 (♀)
TL 212 234
MSW-6 60 62
MSW-6/TL 28% 27%
SW-10 45 47
SW-10/TL 21% 20%
S1 25 26
S2 23 25
S3 28 28
S4 28 29
S5 28 31
S6 32 33
S7 32 36
S8 35 37
S9 40 39
S10 40 38
S11 23 24
MD4 (ac) 47 52
MD6 (ac) 78 78
MD8 (ac) 82 86
LV6 (ac) 27 28
LV7 (ac) 30 29
LV8 (ac) 37 35
LV9 (ac) 32 35
LTS 172 163
LTS/TL 81% 70%
LTAS n/a 27
European Journal of Taxonomy 844: 1–108 (2022)
34
SeGment 7. With spines in lateroventral positions, and sensory spots in paradorsal, midlateral and
ventromedial positions (Figs 11A–B, 13F). Glandular cell outlets type 1 present in ventromedial
positions only; no glands observed on dorsal side. Tips of pectinate fringe of posterior segment margin
more slender than on preceding segments. Segment otherwise as segment 6.
SeGment 8. With spines in middorsal and lateroventral positions, and relatively long tubes (21 µm) in
sublateral positions (Figs 11A–B, 12D–E, 13E, G). Sensory spots present in paradorsal positions only.
Fig. 12. Light micrographs showing overview and details of Echinoderes leduci sp. nov. A–G. ♂,
holotype (NIWA-159405). H. ♀, paratype (NIWA-159406). A. Dorsal overview. B. Segments 1 to 6,
dorsal view. C. Segments 1 to 6, ventral view. D. Segments 6 to 11, dorsal view. E. Segments 6 to 11,
ventral view. F. Segments 9 to 11 of male, dorsal view. G. Segments 10 to 11 of male, dorsal view.
H. Segments 10 to 11 of female, dorsal view. Abbreviations: see Material and methods.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
35
Fig. 13. Scanning electron micrographs showing overviews and details of Echinoderes leduci sp. nov.
A. Lateroventral overview. B. Segments 1 to 3, lateral view. C. Segments 1 to 3, ventral view.
D. Segments 1 to 6, laterodorsal view. E. Segments 8 to 9, dorsal view. F. Segments 3 to 7, ventral view.
G. Segments 7 to 9, laterodorsal view. H. Segments 10 to 11 of male, dorsal view. Abbreviations: see
Material and methods.
Table 9. Summary of nature and location of sensory spots, glandular cell outlets, tubes and spines
arranged by series in Echinoderes leduci sp. nov.
Position segment MD PD SD LD ML SL LA LV VL VM
1 gco1 ss ss gco1 ss
2 gco1,ss tu,ss tu gco1,ss
3 gco1 ss ss gco1
4 ac gco1 gco1
5 gco1 ss ss tu gco1,ss
6 ac gco1,ss ss ac gco1,ss
7 ss ss ac gco1,ss
8 ac gco1,ss tu ac gco1
9 gco1,ss ss tu si ac ss gco1
10 gco1 ss tu(♂) ss gco1
11 gco1 × 2 ss pe × 3 (♂) ltas(♀) lts
European Journal of Taxonomy 844: 1–108 (2022)
36
Glandular cell outlets type 1 present in paradorsal and ventromedial positions. Segment otherwise as
segment 7.
SeGment 9. With spines in lateroventral positions. Long tubes (20 µm) present in laterodorsal positions
(Figs 11A, 12D, F, 13E, G). Sensory spots located in paradorsal, subdorsal and ventrolateral positions
(Figs 11A–B, 12E–F, 13E, G). Glandular cell outlets type 1 present paradorsally and ventromedially.
Small sieve plates located in midlateral positions (Fig. 13G). Band of perforation site patches interrupted
in subdorsal area instead of laterodorsally as on preceding segments. Secondary fringe and posterior
segment margin as on preceding segment.
SeGment 10. With well-developed laterodorsal tubes, present in males only, located near posterior
segment margin (Figs 11A, 12F, 13H). Females without tubes (Figs 11C, 12H). Sensory spots present in
subdorsal and ventrolateral positions (Figs 11A–B, 12E–F, 13H); subdorsal pair located rather close to
paradorsal area. Glandular cell outlets type 1 present in middorsal and in ventromedial positions. Band
of perforation site patches as on preceding segment but with additional patch present in paradorsal area.
Posterior segment margin of tergal plate straight, without fringe tips (Fig. 13H); margins of sternal plates
concave, reaching the posterior margin of the terminal segment, with short and narrow fringe tips.
SeGment 11. With lateral terminal spines (Figs 11A–B, 12A, 13A). Females with lateral terminal
accessory spines (Figs 11C–D, 12H); males with three penile spines (Figs 12G, 13H). Dorsal and ventral
spines slender and tubular, with ventral ones longer than dorsal ones; median spines very stout, cone-
shaped (Figs 11A, 12G, 13H). Sensory spots present in subdorsal positions. Two unpaired glandular
cell outlets type 1 present middorsally. Segment devoid of characteristic perforation sites patches, but
with very short cuticular hair-like structures covering paradorsal area. Short fringes covering margins of
tergal and sternal plates. Tergal extensions triangular (Figs 11A, C, 13H). Sternal extensions rounded,
not extending beyond tergal extensions (Fig. 11B, D).
Distribution
Canyons: Pahaua, Honeycomb, 670–1013 m b.s.l. See Fig. 1 for a geographic overview of stations and
Table 1 for station and specimen information.
Taxonomic remarks on Echinoderes dalzottoi sp. nov. and E. leduci sp. nov.
Echinoderes dalzottoi sp. nov. and E. leduci sp. nov. can easily be recognized by the nature of their trunk
appearance with cuticular hairs that are reduced to minute scales, and characteristic strong perforation
sites present on segments 2 to 10. The complete lack of cuticular hairs is an uncommon feature among
species of Echinoderes. Within Echinoderidae, the species’ peculiar lack of cuticular hairs is shared only
with Meristoderes glaber Sørensen et al., 2013 (Sørensen et al. 2013). Resemblance with M. glaber is
only supercial though, and the lack of lateral spines on segment 6, together with the presence of glandular
cell outlets type 2 instead of tubes on segment 8 in M. glaber easily distinguish it from E. dalzottoi and
E. leduci. The new species can also be distinguished from each other without any difculty. Despite
the number of similarities suggesting a close relationship, such as presence of lateroventral spines on
segments 6–9, lateroventral tubes on segment 5, sublateral tubes on segment 8 and laterodorsal tubes on
segment 9, it is possible to distinguish the two species based on several characters. The main difference
is the number of middorsal spines, present on segments 4, 6 and 8 in E. leduci in contrast to spines
on segments 4 and 6 only in E. dalzottoi. Furthermore, E. leduci lacks subdorsal tubes on segment 2
(present in E. dalzottoi), but has laterodorsal tubes on this segment instead.
Having tubes on segment 2 in only laterodorsal and ventrolateral positions as in E. leduci sp. nov. is
quite a rare character, and only shared with two other species of Echinoderes, i.e., E. daenerysae and
E. higginsi Huys & Coomans, 1989 (although a laterodorsal pair was not mentioned in the original
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
37
description of the latter species) (Huys & Coomans 1989; Grzelak & Sørensen 2018). Neither of them
can be confused with E. leduci, however. Except for the dense cuticular hair covering observed in both
species, E. daenerysae possesses only two middorsal spines on segments 6 and 8, while E. higginsi,
although having the same number and arrangement of middorsal spines as E. leduci, is characterized by
long, conspicuous spinous tergal extensions of segment 11 and the lack of tubes on segment 9.
Echinoderes dalzottoi sp. nov. is also easily distinguished from all other congeners by its combination
of spines and tubes. Middorsal spines being restricted to segments 4 and 6 only is rare and shared only
with four other species, i.e., E. astridae, E. bispinosus, E. uozumii, and E. landersi sp. nov. (Higgins
1982; Sørensen 2014; Yamasaki et al. 2020). However, E. astridae, E. bispinosus and E. uozumii lack
dorsal and lateral tubes on segment 2, and have instead two pairs of glandular cell outlet type 2 – a gland
type that is absent in E. dalzottoi. Echinoderes landersi, in turn, is characterized by four pairs of tubes
on segment 2 and three pairs of tubes on segment 8 – a so far unique tube combination (see ‘Remarks’
in the description of E. landersi above for more details).
The combination of tubes in subdorsal, sublateral and ventrolateral positions on segment 2 is also quite
rare. Numerous species present combinations of two or four pairs of tubes on this segment, but only four
species have three pairs, among which only E. hispanicus Pardos et al., 1998 and E. peterseni Higgins &
Kristensen, 1988 have tube arrangements as in E. dalzottoi sp. nov. (Higgins & Kristensen 1988; Pardos
et al. 1998; Grzelak & Sørensen 2018). However, both species also have three middorsal spines and
lack laterodorsal tubes on segment 9, which easily distinguish them from E. dalzottoi. Even considering
that sublateral tubes might be absent in some specimens of E. dalzottoi (a variation in the occurrence
of tubes has recently been reported for a number of echinoderid species; see, e.g., Grzelak & Sørensen
(2018) and Yamasaki & Dal Zotto (2019) for further details), none of the species having only subdorsal
and ventrolateral tubes on segment 2 can be confused with E. dalzottoi due to their signicantly different
middorsal spine patterns and trunk cuticle appearance.
Echinoderes gandal sp. nov.
urn:lsid:zoobank.org:act:84D3AEAA-391B-4C5C-81FC-380B198251F2
Figs 14–16; Tables 10–11
Diagnosis
Echinoderes with spines in middorsal position on segments 6 and 8, and spines in lateroventral positions
on segments 6 to 9. Tubes present in ventrolateral positions on segment 2, lateroventral positions on
segment 5, lateral accessory positions on segment 8, and laterodorsal positions on segments 9 and 10. A
protuberance-like structure emerges between segments 10 and 11 in middorsal position. Lateral terminal
spines long, always exceeding trunk length.
Etymology
The species name refers to Gandalf the Grey, one of the main characters in the novel “The Fellowship of
the Ring”, the rst volume of J.R.R. Tolkien’s “The Lord of the Rings”. He helped form the Fellowship
of the Ring to destroy the One Ring.
Material examined
Holotype
NEW ZEALAND • ♂; Pahaua Canyon, stn TAN1004/31; 41.4962° S, 175.6828° E; 730 m b.s.l.; Apr.
2010; NIWA TAN1004 Voyage; soft sediment; NIWA-159407. Mounted for LM in Fluoromount G on
HS slide.
European Journal of Taxonomy 844: 1–108 (2022)
38
Paratypes
NEW ZEALAND • 2 ♀♀; same collection data as for holotype; ♀ NHMD-916356, ♀ NIWA-159408.
Mounted as holotype • 1 ♀; Pahaua Canyon, stn TAN1004/12; 41.5508° S, 175.7250° E; 1350 m b.s.l.;
Apr. 2010; NIWA TAN1004 Voyage; soft sediment; NIWA-159409. Mounted as holotype • 1 ♀,
1 ♂; Pahaua Canyon, stn TAN1004/22; 41.5100° S, 175.7187° E; 1188 m b.s.l.; Apr. 2010; NIWA
TAN1004 Voyage; soft sediment; ♀ NHMD-916362, ♂ NIWA-159410. Mounted as holotype • 1 ♂;
Pahaua Canyon, stn TAN1004/27; 41.4983° S, 175.7043° E; 1013 m b.s.l.; Apr. 2010; NIWA TAN1004
Voyage; soft sediment; ♂ NIWA-159411. Mounted as holotype • 1 ♀, 2 ♂♂; Honeycomb Canyon,
stn TAN1004/58; 41.4080° S, 175.8987° E; 670 m b.s.l.; Apr. 2010; NIWA TAN1004 Voyage; soft
sediment; ♀ NIWA-159412, ♂♂ NHMD-916357–916358. Mounted as holotype • 1 ♀, 3 ♂♂; Campbell
Canyon, stn TAN1004/126; 42.1422° S, 174.5492° E; 1495 m b.s.l.; Apr. 2010; NIWA TAN1004 Voyage;
soft sediment; ♀ NHMD-916360, 1 ♂ NIWA-159413, 2 ♂♂ NHMD-916359 and 916361. Mounted as
holotype.
Additional material
NEW ZEALAND • 5 ♀♀, 3 ♂♂; Pahaua Canyon, stn TAN1004/27; 41.4983° S, 175.7043° E;
1013 m b.s.l.; Apr. 2010; NIWA TAN1004 Voyage; soft sediment; personal reference collection of
MVS. Mounted for SEM • 1 ♀, 2 ♂♂; Honeycomb Canyon, stn TAN1004/62; 41.4760° S, 175.9477° E;
1171 m b.s.l.; Apr. 2010; NIWA TAN1004 Voyage; soft sediment; personal reference collection of MVS.
Mounted for SEM • 1 ♀; Campbell Canyon, stn TAN1004/126; 42.1422° S, 174.5492° E; 1495 m b.s.l.;
Apr. 2010; NIWA TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted for
SEM.
Description
General. Adults with head, neck and eleven trunk segments (Figs 14–16). Overview of measurements
and dimensions in Table 10. Distribution of cuticular structures, i.e., sensory spots, glandular cell outlets,
spines and tubes, summarized in Table 11. No details regarding scalid arrangement and morphology
could be provided, because introverts of all specimens mounted for SEM fully or partially retracted.
neck. With 16 placids. Midventral placid broadest, 8 µm in width and 15 µm in length, whereas all
others narrower, measuring 6 µm in width at their bases and 13 µm in length, similar in size (Fig. 15B–
C). The trichoscalid plates are well-developed (Fig. 15B–C).
SeGment 1. Consists of complete cuticular ring. Sensory spots located on anterior half of segment,
in subdorsal positions. Glandular cell outlet type 1 present in middorsal and ventrolateral positions
(Figs 14A–B, 15B–C). Cuticular hairs relatively long, distributed around segment. Posterior
segment margin almost straight, forming pectinate fringe with very short, sawtooth-like fringe tips.
SeGment 2. Consists of complete cuticular ring, with tubes located in ventrolateral positions (Figs 14A–
B, 15C, 16C). Sensory spots present in middorsal, laterodorsal and ventromedial positions; sensory
spots on this and following eight segments relatively large, with numerous micropapillae surrounding
central pore and long marginal hair, giving them droplet-shaped appearance (Fig. 16B). Glandular cell
outlets type 1 present in middorsal and ventromedial positions (Figs 14A–B, 15B). Pachycyclus of
anterior segment margin of regular thickness, without interruption. Secondary pectinate fringe present
near anterior segment margin of this and following segments, but usually covered by preceding segment.
Fairly long cuticular hairs evenly distributed across tergal plate; paraventral areas on this and following
three segments with thinner and much shorter, non-bracteate hairs, emerging through perforation sites
visible in SEM only. Posterior segment margin almost straight, but with rounded midventral extension;
pectinate fringe tips as on preceding segment.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
39
Fig. 14. Line art illustrations of Echinoderesgandal sp. nov. A. ♂, dorsal view. B. ♂, ventral view.
C. ♂, segments 10 to 11, with lateral terminal spines drawn in full length, dorsal view. D. ♀, segments 10
to 11, dorsal view. E. ♀, segments 10 to 11, ventral view. Abbreviations: see Material and methods.
European Journal of Taxonomy 844: 1–108 (2022)
40
SeGment 3. Present segment, and eight remaining ones, consist of one tergal and two sternal plates
(Figs 14A–B, 15A–C, 16B–C). Pachycyclus of anterior segment margin of medium thickness, interrupted
at tergosternal and midsternal junctions and middorsally, on this and following seven segments.
Segment with middorsal and ventromedial glandular cell outlets type 1; no sensory spots or other traits
observed. Cuticular hairs interrupted by hairless midlateral area, otherwise as on preceding segment.
Posterior segment margin straight, terminating in pectinate fringe with longer and more slender fringe
tips than on preceding segments.
SeGment 4. With glandular cell outlets type 1 present in paradorsal and ventromedial positions.
Segment otherwise as segment 3.
SeGment 5. With tubes in lateroventral positions (Figs 14B, 15C, 16A–B). Sensory spots present in
midlateral and ventromedial positions; the latter, however, missing in some specimens examined under
Fig. 15. Light micrographs showing overview and details of Echinoderesgandal sp. nov. A–D, F. ♂,
holotype (NIWA-159407). E, G. ♀, paratype (NHMD-916362). A. Ventral overview. B. Segments 1 to
6, dorsal view. C. Segments 1 to 9, ventral view. D. Segments 6 to 10, dorsal view. E. Segments 10 to
11 of female, dorsal view. F. Segments 10 to 11 of male, dorsal view. G. Segment 11 of female, ventral
view. Abbreviations: see Material and methods.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
41
Fig. 16. Scanning electron micrographs showing overviews and details of Echinoderesgandal sp. nov.
A. Lateral overview. B. Ventral overview. C. Segments 1 to 4, ventral view. D. Lateral overview.
E. Segments 4 to 10, dorsal view. F. Segments 8 to 9, lateral view. G. Segments 10 to 11 of female,
dorsal view. H. Segments 10 to 11 of male, dorsal view. Abbreviations: see Material and methods.
European Journal of Taxonomy 844: 1–108 (2022)
42
SEM (Fig. 16B) and for some specimens (including holotype) examined under LM presence/absence of
ventromedial sensory spots could not be conrmed. Glandular cell outlets type 1 present in paradorsal
and ventromedial positions. Pectinate fringe of posterior segment margin and cuticular hairs as on
preceding segment.
SeGment 6. With spines in middorsal and lateroventral positions (Figs 14A–B, 15C–D, 16A–B, D–F).
Sensory spots present in paradorsal, midlateral and ventromedial positions (Fig. 14A–B). Glandular cell
outlets type 1 present in paradorsal and ventromedial positions (Figs 14A–B, 15C). Tips of pectinate
fringe of posterior segment margin slightly longer than on preceding segments. Segment otherwise as
segment 5.
SeGment 7. With spines in lateroventral positions, and sensory spots in paradorsal, midlateral and
ventromedial positions (Figs 14A–B, 15C–D, 16B). Glandular cell outlets type 1 present in ventromedial
positions, not observed on the dorsal side. Cuticular hairs covering as on preceding segment except for
hairless paraventral and ventromedial areas, on this and following four segments.
SeGment 8. With spines in middorsal and lateroventral positions, and tubes in lateral accessory positions
(Figs 14A–B, 15C–D, 16D, F). Sensory spots present in paradorsal positions only. Glandular cell outlets
Table 10. Measurements from light microscopy of Echinoderes gandal sp. nov. (in µm) from the
Hikurangi Margin, including number of measured specimens (n) and standard deviation (SD).
Character n Range Mean SD
TL 10 140–222 186 24.5
MSW-6 7 42–55 50 4.8
MSW-6/TL 7 21.6–32.0% 27.6% 4.3%
SW-10 7 37–42 39 1.9
SW-10/TL 7 18.3–26.4% 21.6% 3.2%
S1 9 18–21 19 1.8
S2 9 17–23 19 1.7
S3 9 15–25 21 3.4
S4 9 17–29 23 4.1
S5 9 22–31 26 2.9
S6 9 24–31 27 2.1
S7 9 24–33 30 2.2
S8 9 29–37 32 2.3
S9 9 29–36 33 2.3
S10 9 26–33 30 2.2
S11 9 19–28 23 5.1
MD6 (ac) 10 34–42 39 2.9
MD8 (ac) 8 49–56 52 2.1
LV6 (ac) 10 17–23 20 2.3
LV7 (ac) 10 19–25 22 2.7
LV8 (ac) 10 23–28 25 2.5
LV9 (ac) 10 24–31 26 2.4
LTS 10 191–275 241 26.2
LTS/TL 10 103–169.3% 132.0% 24.1%
LTAS 3 28–34 30 3.2
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
43
type 1 as on preceding segment. Pectinate fringe of posterior segment margin and cuticular hairs as on
preceding segment.
SeGment 9. With spines in lateroventral positions and long (~18 µm) tubes in laterodorsal positions
(Figs 14A–B, 15D, 16D–F). Sensory spots located in paradorsal, subdorsal and ventrolateral positions
(Figs 14A–B, 15B–C, 16F). Glandular cell outlets type 1 present in paradorsal and ventromedial
positions. Small, rounded sieve plates located in lateral accessory positions. Cuticular hairs covering as
on preceding segment, but less dense on dorsal side around sensory spots and tubes. Pectinate fringe of
posterior segment margin with tips slightly shorter than on preceding segments.
SeGment 10. With laterodorsal tubes located near posterior segment margin; tubes well developed in
both sexes (Figs 15E–F, 16G–H). Sensory spots present in ventrolateral positions. Glandular cell outlets
type 1 present as two middorsal ones, and in ventromedial positions. Cuticular hairs signicantly scarcer
than on preceding segment. Central part of tergal palate devoid of hairs; short cuticular hairs lightly
scattered on lateral halves only. Hairs on sternal plates short and present mostly on lateral halves and
near posterior segment margin. Posterior segment margin of tergal plate straight, with shorter fringe tips
than those on preceding segment; margins of sternal plates with longer fringe tips than those on tergal
plate and extend midventrally, almost reaching posterior margin of terminal segment.
SeGment 11. With very long lateral terminal spines, always exceeding trunk length (Fig. 14C; Table 10).
Males with three pairs of penile spines; dorsal and ventral ones are long, relatively thin tubes, whereas
median ones markedly thicker, conical and stout (Figs 14A–B, 16H). Females with lateral terminal
accessory spines (Fig. 14D–E). Sensory spots present in paradorsal positions (Figs 14A, D, 16D); sensory
spots smaller than on preceding segments, without long marginal hairs. Glandular cell outlets type 1
present in subdorsal positions. Middorsal protuberance-like structure extends from intersegmentary
joint (Figs 14A,D, 15E–F, 16G). Segment devoid of cuticular hairs, but with short cuticular hair-like
structures covering paradorsal area and short fringes covering margins of tergal and sternal plates. Tergal
extensions short and pointed, with small tooth at inner margin (Fig. 15G). Sternal extensions broadly
rounded, not extending beyond tergal extensions.
Table 11. Summary of nature and location of sensory spots, glandular cell outlets, tubes and spines
arranged by series in Echinoderesgandalsp. nov.
Position segment MD PD SD LD ML LA LV VL VM
1 gco1 ss gco1
2 gco1,ss ss tu gco1,ss
3 gco1 gco1
4 gco1 gco1
5 gco1 ss tu ss*,gco1
6 ac gco1,ss ss ac ss, gco1
7 ss ss ac ss,gco1
8 ac ss tu ac gco1
9 gco1,ss ss tu si ac ss gco1
10 gco1,gco1 tu ss gco1
11 pr ss gco1 pe 3(♂) ltas(♀) lts
European Journal of Taxonomy 844: 1–108 (2022)
44
Distribution
Canyons: Pahaua, Campbell, Honeycomb, 730–1495 m b.s.l. See Fig. 1 for a geographic overview of
stations and Table 1 for station and specimen information.
Taxonomic remarks on Echinoderes gandal sp. nov.
The spine pattern with middorsal spines on segments 6 and 8 is rare among species of Echinoderes,
and is shared with only three species, i.e., E. daenerysae, E. hviidarum Sørensen et al., 2018 and
E. ultraabyssalis Adrianov & Maiorova, 2019, and an undescribed species, Echinoderes sp. 1, from
the Atacama Trench (Grzelak & Sørensen 2018; Sørensen et al. 2018; Adrianov & Maiorova 2019;
Grzelak et al. 2021). Furthermore, E.gandal sp. nov. possesses laterodorsal tubes on segment 9, which
is another relatively rare trait, shared with all the abovementioned species, which suggests that these
species might represent a group of closely related species. The only other species with tubes in this
position on the dorsal side of segment 9 is E. belenae (see Pardos et al. 2016b) and E. frodoi sp. nov.,
E. dalzottoi sp. nov. and E. leduci sp. nov., all described in the present study. Nevertheless, none of these
species can in any way be confused with E.gandal due to their signicantly different spine patterns.
Echinoderes gandal sp. nov. probably shows most resemblance to E. ultraabyssalis, the deepest
kinorhynch species described so far (> 9000 m water depth) from the Kuril-Kamchatka Trench, and
Echinoderes sp. 1, found in the Atacama Trench at a water depth of 7700 m (Adrianov & Maiorova
2019; Grzelak et al. 2021). Echinoderes gandal shares several features with these hadal/deep-sea
trench species, including the presence of tubes in ventrolateral positions on segment 2, lateroventral
positions on segment 5, and in lateral accessory positions on segment 8, the presence of a middorsal
protuberance between segments 10 and 11, as well as very long lateral terminal spines, the shape of the
tergal extensions and the lack of glandular cell outlets type 2. Nevertheless, despite the overall similarity,
E.gandal can be distinguished from both congeners by its morphometrics. Echinoderesgandal is
markedly smaller in trunk length than both species (TL: 186 µm vs 267 µm and 257 µm, respectively)
but has proportionally longer lateral terminal spines, which always exceed the trunk length, resulting
in a markedly higher LTS/TL ratio in E.gandal in comparison with E. ultraabyssalis and Echinoderes
sp. 1 (LTS/TL: 132% vs 59% and 91%, respectively) (Adrianov & Maiorova 2019; Grzelak et al. 2021).
It appears that also the arrangement and number of sensory spots show some differences. Unique for
E.gandal is the presence of laterodorsal sensory spots on segment 2 but the lack of these structures in
ventrolateral positions on segment 1, and in middorsal position on segment 10 as found in the other two
species. In addition, E. ultraabyssalis lacks well-developed laterodorsal tubes on segment 10, which are
present in both sexes and easy to visualize even with LM in E.gandal.
Echinoderes frodoi sp. nov.
urn:lsid:zoobank.org:act:85C8FA48-24D1-4C55-9B41-179E186C7037
Figs 17–19; Tables 12–13
Diagnosis
Echinoderes with spines in middorsal position on segments 4, 6 and 8, and in lateroventral positions
on segments 6 to 9. Tubes present in midlateral positions on segment 1 (might be missing in some
specimens), subdorsal, laterodorsal, sublateral and ventrolateral positions on segment 2, lateroventral
positions on segment 5, lateral accessory positions on segment 8, and laterodorsal positions on segment 9.
Sexually dimorphic tubes furthermore present in laterodorsal positions on segment 10: male tubes well-
developed; female tubes minute.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
45
Etymology
The species name refers to Frodo Baggins, the main character in the novel “The Fellowship of the Ring”,
the rst volume of J.R.R. Tolkien’s “The Lord of the Rings”.
Material examined
Holotype
NEW ZEALAND • ♂; Pahaua Canyon, stn TAN1004/22; 41.5100° S, 175.7187° E; 1188 m b.s.l.; Apr.
2010; NIWA TAN1004 Voyage; soft sediment; NIWA-159414. Mounted for LM in Fluoromount G on
HS slide.
Paratypes
NEW ZEALAND • 1 ♀, 2 ♂♂; Hikurangi Slope, stn TAN1004/4; 41.6837° S, 175.6642° E; 1046 m b.s.l.;
Apr. 2010; NIWA TAN1004 Voyage; soft sediment; ♀ NIWA-159415, 1 ♂ NHMD-916331, 1 ♂ NIWA-
159416. Mounted for LM in Fluoromount G on glass slides • 1 ♀; Hikurangi Slope, stn TAN1004/38;
41.5937° S, 175.8532° E; 1121 m b.s.l.; Apr. 2010; NIWA TAN1004 Voyage; soft sediment; NHMD-
916335. Mounted as holotype • 1 ♀; Hikurangi Slope, stn TAN1004/76; 41.6833° S, 175.6500° E;
1282 m b.s.l.; Apr. 2010; NIWA TAN1004 Voyage; soft sediment; NHMD-916336. Mounted as
holotype • 1 ♀; Hikurangi Slope, stn TAN1004/128; 42.0485° S, 174.7000° E; 1420 m b.s.l.; Apr. 2010;
NIWA TAN1004 Voyage; soft sediment; NHMD-916337. Mounted for LM in Fluoromount G on glass
slide • 1 ♀; same collection data as for holotype; NIWA-159417. Mounted as holotype • 1 ♂; Hikurangi
Slope, stn TAN1004/31; 41.4962° S, 175.6828° E; 730 m b.s.l.; Apr. 2010; NIWA TAN1004 Voyage; soft
sediment; NHMD-916332. Mounted for LM in Fluoromount G on glass slide • 1 ♀; Campbell Canyon,
stn TAN1004/126; 42.1422° S, 174.5492° E; 1495 m b.s.l.; Apr. 2010; NIWA TAN1004 Voyage; soft
sediment; NHMD-916333. Mounted for LM in Fluoromount G on glass slide • 1 ♂; Seamount 766,
stn TAN1004/129; 42.1345° S, 174.5860° E; 1456 m b.s.l.; Apr. 2010; NIWA TAN1004 Voyage; soft
sediment; NHMD-916334. Mounted as holotype.
Additional material
NEW ZEALAND • 1 ♀, 1 ♂; Hikurangi Slope, stn TAN1004/38; 41.5937° S, 175.8532° E; 1121 m b.s.l.;
Apr. 2010; NIWA TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted for
SEM • 1 ♀; Hikurangi Slope, stn TAN1004/76; 41.6833° S, 175.6500° E; 1282 m b.s.l.; Apr. 2010;
NIWA TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted for SEM • 1 ♂;
Pahaua Canyon, stn TAN1004/27; 41.4983° S, 175.7043° E; 1013 m b.s.l.; Apr. 2010; NIWA TAN1004
Voyage; soft sediment; personal reference collection of MVS. Mounted for SEM • 1 ♂; Honeycomb
Canyon, stn TAN1004/58; 41.4080° S, 175.8977° E; 670 m b.s.l.; Apr. 2010; NIWA TAN1004 Voyage;
soft sediment; personal reference collection of MVS. Mounted for SEM • 1 ♂; Campbell Canyon,
stn TAN1004/92; 41.8922° S, 174.6347° E; 683 m b.s.l.; Apr. 2010; NIWA TAN1004 Voyage;
soft sediment; personal reference collection of MVS. Mounted for SEM • 1 ♀; Campbell Canyon,
stn TAN1004/126; 42.1422° S, 174.5492° E; 1495 m b.s.l.; Apr. 2010; NIWA TAN1004 Voyage; soft
sediment; personal reference collection of MVS. Mounted for SEM.
Description
General. Adults with head, neck and eleven trunk segments (Figs 17–19). Overview of measurements
and dimensions in Table 12. Distribution of cuticular structures, i.e., sensory spots, glandular cell outlets,
spines and tubes, summarized in Table 13. No details regarding scalid arrangement and morphology
could be provided, because introverts of all specimens mounted for SEM fully or partially retracted.
neck. With 16 placids. Midventral placid broadest, 9 µm in width and 16 µm in length. Remaining
placids narrower, 7 µm in width and 15 µm in length, similar in size (Fig. 19B). The trichoscalid plates
well developed.
European Journal of Taxonomy 844: 1–108 (2022)
46
Fig. 17. Line art illustrations of Echinoderes frodoi sp. nov. A. ♂, dorsal view. B. ♂, ventral view. C. ♀,
segments 10–11, dorsal view. D. ♀, segments 10–11, ventral view. Abbreviations: see Material and
methods.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
47
Fig. 18. Light micrographs showing overview and details of Echinoderes frodoi sp. nov. A, C (close-
up). ♂, holotype (NIWA-159414). B–D, F. ♀, paratype (NHMD-916336). E. ♀, paratype (NHMD-
916335). G–H. ♂, paratype (NIWA-159416). A. Ventral overview. B. Segments 1 to 8, dorsal view.
C. Segments 1 to 8, ventral view with close-up showing midlateral tube on segment 1. D. Segments 9 to
10, dorsal view. E. Segments 8 to 10, ventral view. F. Segments 10 to 11, ventral view. G. Segments 9 to
10, lateral view. H. Segments 10 to 11, lateral view. Abbreviations: see Material and methods.
European Journal of Taxonomy 844: 1–108 (2022)
48
Fig. 19. Scanning electron micrographs showing overviews and details of Echinoderes frodoi sp. nov.
A. Lateral overview. B. Ventral overview. C. Segments 4 to 8, laterodorsal view. D. Dorsal overview.
E. Segments 1 to 4, laterodorsal view. F. Segments 2 to 6, lateral view. G. Segments 8 to 10 of male,
lateral view. H. Segments 9 to 11 of female, laterodorsal view. Abbreviations: see Material and methods.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
49
SeGment 1. Consists of complete cuticular ring (Figs 17A–B, 18A–C, 19B). Tubes present in midlateral
positions in holotype and some paratypes; however, this character missing in other specimens and
thus shows variation at population level (Figs 17A, 18C, 19A, E); no morphological or developmental
differences explain presence or absence of tubes. Sensory spots located anteriorly on segment, but
not at anterior margin, in subdorsal and laterodorsal positions. Sensory spots on this and following
segments rounded, with numerous micropapillae surrounding central pore and several longer hairs
along posterior margin. Glandular cell outlet type 1 present in middorsal position, and in lateroventral
positions (Fig. 17A–B). Cuticular hairs lightly scattered on dorsal and lateral sides, and in small cluster
ventromedially. Posterior segment margin almost straight, forming pectinate fringe with short and
pointed fringe tips.
SeGment 2. Consists of complete cuticular ring, with tubes located in subdorsal, laterodorsal, sublateral
and ventrolateral positions (Figs 17A–B, 18A–C, 19A–B, E). Sensory spots present in laterodorsal
and ventromedial positions. Glandular cell outlet type 1 located middorsally. Pachycyclus of anterior
segment margin interrupted in middorsal position. Secondary pectinate fringe present near anterior
segment margin of this and following segments, but usually covered by preceding segment. Cuticular
hairs lightly scattered on ventral side and more densely in dorsal and lateral areas. Pectinate fringe of
posterior margin slightly longer in ventromedial areas, otherwise as on preceding segment.
Table 12. Measurements from light microscopy of Echinoderes frodoi sp. nov. (in µm) from the
Hikurangi Margin, including number of measured specimens (n) and standard deviation (SD).
Character n Range Mean SD
TL 8 161–202 185 16.2
MSW-7 6 42–47 45 1.4
MSW-7/TL 6 22.2–27.3% 24.7% 1.9%
SW-10 6 32–38 36 2.3
SW-10/TL 6 18.8–22.2% 20.1% 1.4%
S1 9 18–23 21 1.7
S2 9 17–21 19 1.2
S3 9 19–24 22 1.4
S4 9 22–28 24 2.0
S5 9 22–31 26 2.8
S6 9 24–32 27 2.4
S7 9 23–34 29 3.3
S8 9 26–35 30 3.0
S9 9 28–35 30 2.2
S10 9 25–30 28 1.9
S11 9 17–24 21 2.1
MD4 (ac) 9 33–40 37 2.7
MD6 (ac) 9 55–60 58 1.9
MD8 (ac) 9 61–66 64 1.6
LV6 (ac) 9 24–27 25 0.9
LV7 (ac) 9 25–30 28 1.8
LV8 (ac) 9 28–33 31 2.0
LV9 (ac) 9 27–32 30 1.5
LTS 8 144–155 148 3.5
LTS/TL 7 70.3–88.2% 79.0% 6.3%
LTAS 5 34–48 43 5.4
European Journal of Taxonomy 844: 1–108 (2022)
50
SeGment 3. Present segment, and eight remaining ones, consist of one tergal and two sternal plates
(Figs 17A–B, 18A, C, 19B). Pachycyclus of anterior segment margin of regular thickness, with
middorsal interruption in addition to interruptions around tergosternal and midsternal junctions, on this
and following segments. Segment with sensory spots in subdorsal and midlateral positons, and glandular
cell outlets type 1 located ventromedially. On this and following four segments, cuticular hairs arranged
in two or three rows across tergal plate, except for hairless midlateral areas and on lateral halves of
sternal plates; paraventral and ventromedial areas devoid of hairs. Pectinate fringe as on preceding
segment.
SeGment 4. With exible spine in middorsal position and glandular cell outlets type 1 in paradorsal and
ventromedial positions (Figs 17A, 18B, 19C). Sensory spots not present. Cuticular hairs and posterior
segment margin as on preceding segment.
SeGment 5. With tubes in lateroventral positions, and sensory spots present in subdorsal, midlateral and
ventromedial positions (Figs 17A–B, 18B–C, 19F). Glandular cell outlet type 1 present in middorsal
position and in ventromedial positions. Pectinate fringe of posterior segment margin slightly longer;
cuticular hairs as on preceding segment.
SeGment 6. With spines in middorsal and lateroventral positions (Figs 17A, 18B, 19C, F). Sensory spots
present in paradorsal and midlateral positions. Glandular cell outlets type 1 present in pairs in paradorsal
and ventromedial positions. Pectinate fringe on this and following two segments as on preceding one.
SeGment 7. With acicular spines in lateroventral positions, and sensory spots in paradorsal, midlateral
and ventromedial positions (Figs 17A–B, 18C, 19C). Glandular cell outlets type 1 present in paradorsal
and ventromedial positions. Cuticular hairs on dorsal side less dense than on preceding segment, with
hairless middorsal and paradorsal patches. Pectinate fringe as on preceding segment.
SeGment 8. With acicular spines in middorsal and lateroventral positions, and tubes in lateral accessory
positions (Figs 17A–B, 18B, E, 19G). Sensory spots present in paradorsal positions only. Glandular cell
outlets type 1 and other structures as on preceding segment.
SeGment 9. With spines in lateroventral positions, and tubes in laterodorsal positions (Figs 17A–B,
18D–E, G, 19G–H). Tubes rather short with truncated tips, not differentiated into thicker proximal and
thinner distal part. Sensory spots present in paradorsal, subdorsal and ventrolateral positions. Glandular
Table 13. Summary of nature and location of sensory spots, glandular cell outlets, tubes and spines
arranged by series in Echinoderes frodoi sp. nov.
Position segment MD PD SD LD ML SL LA LV VL VM
1 gco1 ss ss tu* gco1
2 gco1 tu tu,ss tu tu ss
3 ss ss gco1
4 ac gco1 gco1
5 gco1 ss ss tu ss,gco1
6 ac gco1,ss ss ac gco1
7 gco1,ss ss ac ss,gco1
8 ac gco1,ss tu ac gco1
9 gco1,ss ss tu si ac ss gco1
10 gco1,gco1 ss tu ss gco1
11 gco1 ss pe ×3(♂) ltas(♀) lts
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
51
cell outlets type 1 present in paradorsal and ventromedial positions. Small, rounded sieve plates located
in lateral accessory positions. Cuticular hairs lightly scattered on dorsal side; central part of tergal
plate devoid of hairs. Pectinate fringe of posterior segment margin with tips shorter than on preceding
segments.
SeGment 10. With well-developed laterodorsal tubes in males, and minute, very slender tubes in females,
located near posterior segment margin (Figs 17A, C, 18H, 19G–H). Sensory spots present in subdorsal and
ventrolateral positions. Glandular cell outlets type 1 present as two middorsal ones and in ventromedial
positions. Cuticular hairs scarcer than on preceding segment. Central part of tergal plate devoid of hairs;
hairs on sternal plates present only on lateral halves. Pectinate fringe with very short tips.
SeGment 11. With pair of long lateral terminal spines (Figs 17A–B, 19D). Males with three pairs of penile
spines; dorsal and ventral spines relatively long, slender and tubular, while median ones much stouter
(Figs 17A–B, 18A, H). Females with lateral accessory spines (Figs 17C–D, 18F, 19D). Sensory spots
present in paradorsal positions. One middorsal glandular cell outlet type 1 present. Segment devoid of
cuticular hairs in both sexes, but with short cuticular hair-like structures covering paradorsal area and
short fringes covering margins of tergal and sternal plates. Tergal extensions short and pointed, with two
small denticles at inner margin (Figs 17A–D, 18F). Sternal extensions rounded, not extending beyond
tergal extensions (Figs 17B, D, 18F).
Distribution
Hikurangi Margin, from slope, through canyon, and seamount habitats, 670–1495 m b.s.l. See Fig. 1 for
a geographic overview of stations and Table 1 for station and specimen information.
Taxonomic remarks on Echinoderes frodoi sp. nov.
The spine and tube distribution in the middorsal and lateroventral series of E. frodoi sp. nov., with
middorsal spines on segments 4, 6 and 8, and tubes/spines in lateroventral positions on segments 5 to 9,
is a common pattern observed among the species of Echinoderes and shared by 24 congeners (Yamasaki
et al. 2020a). However, when we combine these characters with the presence of four pairs of tubes on
segment 2, we shorten the list to only one described species, i.e., E. hakaiensis Herranz et al., 2018 from
British Columbia and an undescribed species, Echinoderes sp. 3, from the Atacama Trench (Herranz
et al. 2018; Grzelak et al. 2021). However, E. frodoi is easily distinguished from both species by the
presence of short laterodorsal tubes on segment 9, and the (occasional) presence of midlateral tubes on
segment 1. All three share the presence of tubes on segment 8, the positions of several sensory spots and
glandular cell outlets type 1, as well as the shape of the tergal extensions. However, E. frodoi has tubes
in lateral accessory positions on segment 8, which distinguishes it from E. hakaiensis and Echinoderes
sp. 3, which have their tubes in sublateral positions, making the space between lateroventral spines
and the tubes conspicuously larger. Another signicant difference between E. frodoi, E. hakaiensis and
Echinoderes sp. 3 is evident in morphometric details. Echinoderes frodoi is markedly smaller in trunk
length than both species (TL: 185 µm vs 324 µm and 211 µm, respectively), but has proportionally
longer lateral terminal spines, resulting in a markedly higher LTS/TL ratio in E. frodoi in comparison
with E. hakaiensis and Echinoderes sp. 3 (LTS/TL: 79% vs 40% and 69%, respectively) (Herranz et al.
2018; Grzelak et al. 2021).
The occurrence of cuticular structures on segment 1 is very rare in echinoderid species, and the midlateral
tubes on segment 1 as in E. frodoi sp. nov. is a trait shared exclusively with E. cantabricus Pardos et al.,
1998 (see Pardos et al. 1998). However, E. cantabricus cannot in any way be confused with E. frodoi.
The species has only a single middorsal spine on segment 4 and can also be distinguished by the absence
of lateroventral spines on segment 9 (Pardos et al. 1998). What might make the picture a bit unclear is the
fact that these tubes apparently are not consistently present in all specimens of E. frodoi. We did not nd
European Journal of Taxonomy 844: 1–108 (2022)
52
any clear explanation for this intraspecic dimorphism, since the variation in tube pattern was not related
to the sexual or developmental stage, neither to intraspecic variation between populations: the presence
or absence of tubes were noted for specimens from the same locality. Nevertheless, a variation in spine/
tube pattern in Echinoderidae has recently been documented for several species (e.g., E. arlis Higgins,
1966, E. eximus, E. levanderi and E. rhaegali Grzelak & Sørensen, 2017 in Grzelak & Sørensen 2018)
and it seems that such morphological variation might be more frequent among Echinoderes species than
previously thought (Grzelak & Sørensen 2018, 2019; Sørensen, 2018).
Furthermore, the presence of laterodorsal tubes on segment 9 is also a rather rare feature among
echinoderids. This trait has previously been described for only four species, i.e., E. belenae, E. daenerysae,
E. hviidarum, E. ultraabyssalis and in the yet undescribed Echinoderes sp. 1 from the Atacama Trench
(Pardos et al. 2016b; Grzelak & Sørensen 2018; Sørensen et al. 2018; Adrianov & Maiorova 2019;
Grzelak et al. 2021). The latter four cannot in any way be confused with E. frodoi sp. nov. due to the
presence of only 2 middorsal spines, on segments 6 and 8. The only other species with middorsal spines
on segments 4, 6 and 8 and tubes in laterodorsal position on segment 9 is E. belenae, but this species
is generally very rich in tubes, and carries no less than thirteen pairs on its eleven trunk segments. The
species furthermore has conspicuously short lateral terminal spines (Pardos et al. 2016b). Laterodorsal
tubes on segment 9 were also found during the present study in E.gandal sp. nov., E. dalzottoi sp. nov.
and E. leduci sp. nov. However, the former two species can easily be distinguished from E. frodoi by the
presence of only two middorsal spines. The latter, E. leduci, similarly to E. frodoi, is characterized by
having middorsal spines on segments 4, 6 and 8, but can nevertheless be distinguished from E. frodoi by
the lack of subdorsal and sublateral tubes on segment 2.
Echinoderes samwisei sp. nov.
urn:lsid:zoobank.org:act:9D00BABD-1183-40FA-BF7D-0F54E304DCD7
Figs 20–22; Tables 14–15
Diagnosis
Echinoderes with spines in middorsal position on segments 4, 6 and 8, and spines in lateroventral
positions on segments 6 to 9. Tubes present in lateroventral positions on segment 5, lateral accessory
positions on segment 8, and laterodorsal positions on segment 10. A protuberance-like structure emerges
between segments 10 and 11 in middorsal position.
Etymology
The species name refers to Samwise Gamgee, one of the main characters in the novel “The Fellowship
of the Ring”, the rst volume of J.R.R. Tolkien’s “The Lord of the Rings”. Samwise was a hobbit from
the Shire, Frodo Baggins’ best friend and one of the most loyal members of the Fellowship of the Ring.
Material examined
Holotype
NEW ZEALAND • ♂; Hikurangi Slope, stn TAN1004/38; 41.5937° S, 175.8532° E; 1121 m b.s.l.; Apr.
2010; NIWA TAN1004 Voyage; soft sediment; NIWA-159418. Mounted for LM in Fluoromount G on
HS slide.
Paratypes
NEW ZEALAND • 1 ♀, 3 ♂♂; Hikurangi Slope, stn TAN1004/4; 41.6837° S, 175.6642° E; 1046 m b.s.l.;
Apr. 2010; NIWA TAN1004 Voyage; soft sediment; 1 ♀ NHMD-917299, 1 ♂ NHMD-917300, 2 ♂♂
NIWA-159419 to 159420. 1 ♀ and 2 ♂♂ mounted for LM in Fluoromount G on glass slides, 1 ♂
mounted as holotype • 1 ♂; Pahaua Canyon, stn TAN1004/12; 41.5508° S, 175.7250° E; 1350 m b.s.l.;
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
53
Apr. 2010; NIWA TAN1004 Voyage; soft sediment; NHMD-917301. Mounted for LM in Fluoromount
G on glass slide • 1 ♂; same collection data as for holotype; NIWA-159421. Mounted as holotype • 1 ♀;
Seamount 310, stn TAN1004/72; 41.3657° S, 176.1958° E; 985 m b.s.l.; Apr. 2010; NIWA TAN1004
Voyage; soft sediment; NHMD-917302. Mounted for LM in Fluoromount G on glass slide.
Additional material
NEW ZEALAND • 1 ♀; same collection data as for holotype; personal reference collection of MVS.
Mounted for SEM • 1 ♀; Hikurangi Slope, stn TAN1004/44; 41.5258° S, 175.8003° E; 728 m b.s.l.;
Apr. 2010; NIWA TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted for
SEM • 1 ♀; Honeycomb Canyon, stn TAN1004/62; 41.4760° S, 175.9477° E; 1171 m b.s.l.; Apr. 2010;
NIWA TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted for SEM • 1 ♂;
Seamount 310, stn TAN1004/72; 41.3657° S, 176.1958° E; 985 m b.s.l.; Apr. 2010; NIWA TAN1004
Voyage; soft sediment; personal reference collection of MVS. Mounted for SEM.
Description
General. Adults with head, neck and eleven trunk segments (Figs 20–22). Overview of measurements
and dimensions in Table 14. Distribution of cuticular structures, i.e., sensory spots, glandular cell outlets,
spines and tubes, summarized in Table 15. No details regarding scalid arrangement and morphology
could be provided, because introverts of all specimens mounted for SEM fully or partially retracted.
neck. With 16 placids. Midventral placid broadest, 10 µm in width and 13 µm in length, whereas all
others narrower, measuring 5 µm in width at bases and 12 µm in length, similar in size (Figs 21C, 22D).
Trichoscalid plates well developed.
SeGment 1. Consists of complete cuticular ring. Sensory spots located on anterior half of segment, in
subdorsal positions. Glandular cell outlet type 1 present in middorsal, subdorsal and ventrolateral positions
(Figs 20A–B, 21C). Segment almost completely hairless (Fig. 22B–E). Posterior segment margin almost
straight, forming pectinate fringe. Fringe with well-developed, relatively long tips, homogenous along
segment margin (Fig. 22C, E).
SeGment 2. Consists of complete cuticular ring, without tubes. Sensory spots present in middorsal,
midlateral and ventromedial positions (Figs 20A–B, 22B, E). Glandular cell outlets type 1 present
in middorsal position (Figs 20A–B, 21B). Other structures not observed. Pachycyclus of anterior
segment margin of regular thickness, without interruption. Secondary pectinate fringe present near
anterior segment margin of this and following segments, but covered by preceding segment. Fairly
long, single cuticular hairs sparsely scattered around segment. Posterior segment margin almost straight;
pectinate fringe tips as on preceding segment.
SeGment 3. Present segment, and eight remaining ones, consist of one tergal and two sternal plates
(Figs 20A–B, 21C, E–F, 22D). Pachycyclus of anterior segment margin of regular thickness, interrupted
at tergosternal and midsternal junctions and middorsally, on this and following seven segments.
Segment with middorsal glandular cell outlet type 1 and subdorsal sensory spots only; no other traits
observed. Cuticular hairs more densely distributed across tergal plate than on preceding segment, except
for narrow hairless line in laterodorsal area; sternal plates with very few cuticular hairs; on this and
following segments paraventral areas completely devoid of hairs. Posterior segment margin straight,
terminating in pectinate fringe with fringe tips as on preceding segments.
SeGment 4. With spine in middorsal position; spine relatively short (23 µm), only slightly exceeding
posterior segment margin (Figs 20A, 22A). Pair of glandular cell outlets type 1 present in paradorsal
positions. No other traits observed. Segment otherwise as segment 3.
European Journal of Taxonomy 844: 1–108 (2022)
54
Fig. 20. Line art illustrations of Echinoderes samwisei sp. nov. A. ♂, dorsal view. B. ♂, ventral view.
C. ♀, segments 10–11, dorsal view. D. ♀, segments 10–11, ventral view. Abbreviations: see Material
and methods.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
55
SeGment 5. With tubes in lateroventral positions (Figs 20B, 21C, E–F, 22C, F). Glandular cell outlets
type 1 present in paradorsal positions only. No sensory spots or other structures present. Pectinate fringe
of posterior segment margin and cuticular hairs as on preceding segment.
Fig. 21. Light micrographs showing overview and details of Echinoderes samwisei sp. nov. A, D. ♀,
paratype (NHMD-917299). B, E, H. ♀, paratype (NHMD-917302). C, F, I. ♂, holotype (NIWA-159418).
G. ♂, paratype (NHMD-917300). A. Lateral overview. B. Segments 3 to 11, dorsal view. C. Segments 1
to 5, ventral view. D. Segments 7 to 11, lateral view. E. Segments 2 to 11, ventral view. F. Segments 5 to
8, ventral view. G. Segments 10 to 11 of male, dorsal view. H. Segments 9 to 11 of female, ventral view.
I. Segments 9 to 11 of male, ventral view. Abbreviations: see Material and methods.
European Journal of Taxonomy 844: 1–108 (2022)
56
SeGment 6. With spines in middorsal and lateroventral positions (Figs 20A–B, 21A, 22B). Sensory spots
present in paradorsal, midlateral and ventromedial positions (Figs 20A–B, 21F, 22C, F). Glandular cell
outlets type 1 present in paradorsal and ventromedial positions (Figs 20A–B, 21F). Tips of pectinate
fringe of posterior segment margin as on preceding segments. Segment otherwise as segment 5.
Fig. 22. Scanning electron micrographs showing overviews and details of Echinoderes samwisei sp. nov.
A. Dorsal overview of male. B. Lateral overview of female. C. Segments 1 to 5, lateral view. D. Ventral
overview of male. E. Segments 1 to 5, ventral view. F. Segments 5 to 8, lateral view. G. Segments 8
to 10 of female, lateral view. H. Segments 8 to 11 of male, lateral view. I. Segments 9 to 11 of female,
dorsal view. Abbreviations: see Material and methods.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
57
SeGment 7. With spines in lateroventral positions, and glandular cell outlets type 1 in paradorsal and
ventromedial positions (Figs 20A–B, 21F). Sensory spots not observed. Cuticular hairs covering as on
preceding segment.
SeGment 8. With spines in middorsal and lateroventral positions, and tubes in lateral accessory positions
(Figs 20A–B, 21D–F, 22F–H). Middorsal spine long, reaching posterior part of segment 10 (Figs 21A–
B, 22A). Sensory spots present in paradorsal positions only. Glandular cell outlets type 1 in paradorsal
and ventromedial positions. Pectinate fringe of posterior segment margin as on preceding segment,
except slightly shorter and narrower fringe tips along paradorsal and subdorsal areas of segment margin.
SeGment 9. With spines in lateroventral positions (Figs 20B, 21D). Sensory spots located in subdorsal
and ventrolateral positions; subdorsal pair situated close to paradorsal area (Figs 20A–B, 21E, 22G–H).
Glandular cell outlets type 1 present in paradorsal and ventromedial positions. Small, rounded sieve
plates located in lateral accessory positions. Cuticular hair covering and pectinate fringe as on preceding
segment.
SeGment 10. With laterodorsal tubes located near posterior segment margin; tubes well developed in
both sexes, but slightly longer in males (Figs 21G, 22G–H). Sensory spots present in subdorsal and
Table 14. Measurements from light microscopy of Echinoderes samwisei sp. nov. (in µm) from the
Hikurangi Margin, including number of measured specimens (n) and standard deviation (SD).
Character n Range Mean SD
TL 7 140–191 174 17.2
MSW-6 4 40–41 41 0.5
MSW-6/TL 4 21–24% 32% 1.7%
SW-10 4 36–37 36 0.5
SW-10/TL 4 19–22% 21% 1.0%
S1 7 17–22 19 1.9
S2 7 15–18 16 1.2
S3 7 18–20 19 0.9
S4 7 21–25 22 1.5
S5 7 23–28 26 1.9
S6 7 24–28 26 1.8
S7 7 26–30 28 2.0
S8 7 27–32 29 1.8
S9 7 27–34 31 2.6
S10 7 24–32 28 3.0
S11 7 19–20 20 0.4
MD4 (ac) 7 21–25 23 1.4
MD6 (ac) 7 53–60 57 2.6
MD8 (ac) 7 64–68 66 1.6
LV6 (ac) 6 17–20 19 1.3
LV7 (ac) 6 24–27 26 1.2
LV8 (ac) 6 27–33 29 2.0
LV9 (ac) 7 27–37 33 3.4
LTS 6 78–109 94 15.2
LTS/TL 6 44–75% 55% 11.3%
LTAS 2 35–37 36 1.8
European Journal of Taxonomy 844: 1–108 (2022)
58
ventrolateral positions (Fig. 22I). Glandular cell outlets type 1 present as two middorsal ones. Cuticular
hairs scarcer than on preceding segment. Central part of tergal plate devoid of hairs; short cuticular
hairs lightly scattered on lateral halves only. Posterior segment margin of tergal plate straight, with
much shorter fringe and narrower tips than those on preceding segment; margins of sternal plates extend
midventrally, reaching posterior margin of terminal segment (Fig. 22D).
SeGment 11. With pair of lateral terminal spines (Fig. 20C). Females with relatively strong, stout lateral
terminal accessory spines (Figs 21H, 22I). Males with three pairs of penile spines; dorsal ones of medium
length, ventral ones long and relatively thin, whereas median ones markedly thicker, conical and stout
(Figs 20A–B, 21G, I 22H). Sensory spots present in paradorsal positions (Figs 20A, C, 22I). Glandular
cell outlet type 1 present in middorsal position. Middorsal protuberance-like structure extends from
intersegmentary joint (Figs 20A, C, 22I). Segment devoid of cuticular hairs, but with dense covering
of relatively long hair-like extensions in paradorsal area, as well as small patches of shorter ones in
subdorsal area. Short fringes covering margins of tergal and sternal plates. Tergal extensions short and
triangular (Figs 21H, 22I). Sternal extensions do not extend beyond tergal extensions.
Distribution
Hikurangi Margin, from slope, through canyon, and seamount habitats, 728–1350 m b.s.l. See Fig. 1 for
a geographic overview of stations and Table 1 for station and specimen information.
Taxonomic remarks on Echinoderes samwisei sp. nov.
The spine pattern with middorsal spines on segments 4, 6 and 8 and lateroventral spines on segments 6
to 9 is very common within Echinoderes, and is shared with 33 species (Yamasaki et al. 2020).
Nevertheless, among these only E. anniae Sørensen et al., 2018, E. meteorensis Yamasaki et al., 2018,
and an undescribed species, Echinoderes sp. 3, from Senghor Seamount reported by Yamasaki et al.
(2019) resemble E. samwisei sp. nov. by lacking tubes on segment 2 (Sørensen et al. 2018; Yamasaki
et al. 2018c, 2019). Thus, none of these species can be confused with E. samwisei. Echinoderes anniae
and E. meteorensis differ from E. samwisei in having several glandular cell outlets type 2 – structures
not observed in E. samwisei. Moreover, both species are characterized by very long lateral terminal
spines and a lack of tubes on segments 5 and 8. Echinoderes samwisei differs from Echinoderes sp. 3 in
Yamasaki et al. (2019) as well, because the latter lacks tubes on segment 8 and possesses characteristic
long and spinose tergal extensions, whereas the new species has lateral accessory tubes on segment 8
and relatively short tergal extensions.
Table 15. Summary of nature and location of sensory spots, glandular cell outlets, tubes and spines
arranged by series in Echinoderes samwisei sp. nov.
Position segment MD PD SD LD ML LA LV VL VM
1 gco1 gco1,ss gco1
2 gco1,ss ss ss
3 gco1 ss
4 ac gco1
5 gco1 tu
6 ac gco1,ss ss ac gco1,ss
7 gco1 ac gco1
8 ac gco1,ss tu ac gco1
9 gco1 ss si ac ss gco1
10 gco1 ×2 ss tu ss
11 gco1,pr ss pe ×3(♂) ltas(♀) lts
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
59
Moreover, E. samwisei sp. nov. is generally characterized by a low number of cuticular structures.
The new species not only lacks glandular cell outlets type 2, but also its number of sensory spots and
glandular cell outlets type 1 is relatively low, as compared with other species. It is indeed possible to
miss certain glandular cell outlets type 1, in particular in species with an extraordinary thin cuticle, but
the availability of several specimens for SEM examination makes it less likely that any sensory spots
have been overlooked. The available SEM specimens were generally in a good condition and relatively
clean, which allowed for a thorough examination using SEM. In spite of this, we observed only 13 pairs
of sensory spots on the trunk.
Therefore, the combination of spine and tube patterns, together with the lack of tubes or glandular
cell outlets type 2 on segment 2 and the relatively ‘simple’ overall appearance of the trunk, makes
E. samwisei sp. nov. unique among its congeners.
Echinoderes legolasi sp. nov.
urn:lsid:zoobank.org:act:5FE9F89C-A48F-42E8-9E97-491D7A88CFB9
Figs 23–25; Tables 16–17
Diagnosis
Echinoderes with spines in middorsal position on segments 4, 6 and 8, and spines in lateroventral
positions on segments 6 to 9. Tubes present in lateroventral positions on segment 5 and laterodorsal
positions on segment 10. Glandular cell outlets type 2 in sublateral positions on segment 1, subdorsal,
laterodorsal, sublateral and ventrolateral positions on segment 2 and midlateral positions on segment 8.
A protuberance-like structure emerges between segments 10 and 11 in middorsal position.
Etymology
The species name refers to Legolas – Elf of Mirkwood, one of the characters in J.R.R. Tolkien’s “The
Lord of the Rings”. Legolas was an excellent archer, and another very valuable member of the Fellowship
of the Ring.
Material examined
Holotype
NEW ZEALAND • ♂; Seamount 310, stn TAN1004/72; 41.3657° S, 176.1958° E; 985 m b.s.l.; Apr.
2010; NIWA TAN1004 Voyage; soft sediment; NIWA-159422. Mounted for LM in Fluoromount G on
HS slide.
Additional material
NEW ZEALAND • 1 ♂; Hikurangi Slope, stn TAN1004/44; 41.5258° S, 175.8003° E; 728 m b.s.l.;
Apr. 2010; NIWA TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted for
SEM.
Description
General. Adults with head, neck and eleven trunk segments (Figs 23–25). Overview of measurements
and dimensions in Table 16. Distribution of cuticular structures, i.e., sensory spots, glandular cell outlets,
spines and tubes, summarized in Table 17. No details regarding scalid arrangement and morphology
could be provided, because introvert of specimen mounted for SEM was partially retracted.
neck. With 16 placids. Midventral placid broadest, 9 µm in width and 11 µm in length, whereas all
others narrower, measuring 6 µm in width at bases and 11 µm in length, similar in size (Figs 24C, 25D).
Trichoscalid plates well developed.
European Journal of Taxonomy 844: 1–108 (2022)
60
Fig. 23. Line art illustrations of Echinoderes legolasi sp. nov. A. ♂, dorsal view. B. ♂, ventral view.
Abbreviations: see Material and methods.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
61
Fig. 24. Light micrographs showing overview and details of Echinoderes legolasi sp. nov., ♂,
holotype (NIWA-159422). A. Dorsal overview. B. Segments 1 to 6, dorsal view. C. Segments 1 to 6,
ventral view. D. Segments 5 to 8, dorsal view. E. Segments 5 to 8, ventral view. F. Ventral overview.
G. Segments 8 to 11, dorsal view. H. Segments 10 to 11, ventral view. Abbreviations: see Material and
methods.
European Journal of Taxonomy 844: 1–108 (2022)
62
SeGment 1. Consists of complete cuticular ring. Pair of glandular cell outlets type 2 present in sublateral
positions (Figs 23B, 24C, 25B). Sensory spots located medially on segment, in subdorsal and laterodorsal
positions (Figs 23A, 24B, 25B); sensory spots on this and following segments relatively large, with
tuft of micropapillae surrounding central pore. Glandular cell outlet type 1 present in middorsal, and
ventrolateral positions (Figs 23A–B, 24B–C). Cuticular hairs relatively long, arranged in single transverse
row medially on segment. Posterior segment margin almost straight, forming pectinate fringe. Fringe
with well-developed long and exible tips, homogenous along segment margin (Fig. 25B).
SeGment 2. Consists of complete cuticular ring. Glandular cell outlets type 2 present in subdorsal,
laterodorsal, sublateral and ventrolateral positions (Figs 23A–B, 24B–C, 25B). Sensory spots located
in laterodorsal and ventromedial positions (Figs 23A–B, 25B). Glandular cell outlets type 1 present in
middorsal and ventromedial positions. Pachycyclus of anterior segment margin of regular thickness,
without interruption. Secondary pectinate fringe present near anterior segment margin of this and
following segments (Fig. 25E, G). Cuticular hairs evenly distributed over dorsal part of segment, ventral
areas less hairy; cuticular hairs shorter and thinner on anterior part of segment. Posterior segment margin
almost straight; pectinate fringe tips as on preceding segment.
Fig. 25. Scanning electron micrographs showing overviews and details of Echinoderes legolasi sp. nov.
A. Lateral overview. B. Segments 1 to 4, dorsolateral view. C. Segments 6 to 9, lateral view. D. Ventral
overview. E. Segments 6 to 10, dorsal view. F. Segments 5 to 7, dorsal view. G. Segments 8 to 11, dorsal
view. H. Segments 10 to 11, dorsal view. Abbreviations: see Material and methods.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
63
SeGment 3. Present segment, and eight remaining ones, consist of one tergal and two sternal plates
(Figs 23A–B, 24C, E–F, 25D). Pachycyclus of anterior segment margin slightly thinner than on preceding
segment, interrupted at tergosternal and midsternal junctions and middorsally, on this and following
seven segments. Segment with sensory spots located in subdorsal and midlateral positions (Figs 24B–
C, 25B), and glandular cell outlets type 1 in middorsal and ventromedial positions. Cuticular hairs
distributed across tergal plate as on preceding segment; ventromedial and paraventral areas on this and
following seven segments with thinner and much shorter, non-bracteate hairs. Posterior segment margin
straight, terminating in pectinate fringe with fringe tips slightly shorter, as on preceding segments, but
still long and exible.
SeGment 4. With spine in middorsal position (Figs 23A, 24B). Sensory spots not present. Pair of glandular
cell outlet type 1 present in paradorsal and ventromedial positions. Pachycycli, pectinate fringe and
cuticular hairs as on preceding segment.
SeGment 5. With tubes in lateroventral positions (Figs 23B, 24C, E). Sensory spots located midlaterally.
Glandular cell outlets type 1 present in middorsal and ventromedial positions. Segment otherwise as
segment 4.
Table 16. Measurements from light microscopy of Echinoderes legolasi sp. nov. (in µm) from the
Hikurangi Margin.
Character Holotype
NIWA-159422 (♂)
TL 175
MSW-7 40
MSW-7/TL 23%
SW-10 33
SW-10/TL 19%
S1 19
S2 19
S3 17
S4 17
S5 18
S6 23
S7 27
S8 29
S9 31
S10 29
S11 19
MD4 (ac) 34
MD6 (ac) 51
MD8 (ac) 65
LVS6 (ac) 16
LVS7 (ac) 17
LVS8 (ac) 19
LVS9 (ac) 23
LTS 89
LTS/TL 51%
European Journal of Taxonomy 844: 1–108 (2022)
64
SeGment 6. With spines in middorsal and lateroventral positions (Figs 23A–B, 24D–E, 25D–F). Sensory
spots present in paradorsal, midlateral and ventromedial positions (Figs 23A–B, 25C). Glandular cell
outlets type 1 present in paradorsal and ventromedial positions (Figs 23A–B, 24D–E). Tips of pectinate
fringe of posterior segment margin as on preceding segments. Segment otherwise as segment 5.
SeGment 7. With spines in lateroventral positions (Figs 23B, 24E). Sensory spots present in paradorsal,
midlateral and ventromedial positions (Figs 23A–B, 25C, E–F), and glandular cell outlets type 1 in
middorsal and ventromedial positions (Figs 23A–B, 24E). Cuticular hair covering as on preceding
segment. Pectinate fringe of posterior segment margin with slightly narrower and shorter tips.
SeGment 8. With spines in middorsal and lateroventral positions (Figs 23A–B, 24G, 25E), and glandular
cell outlets type 2 in midlateral positions (Figs 23A, 24E, G, 25C). Sensory spots present in subdorsal
positions, but located close to paradorsal line (Fig. 25E). Glandular cell outlets type 1 in paradorsal and
ventromedial positions. Pectinate fringe of posterior segment margin as on preceding segment.
SeGment 9. With spines in lateroventral positions (Figs 23B, 25D). Sensory spots located in subdorsal,
midlateral and ventrolateral positions; subdorsal pair situated close to paradorsal area (Figs 23A–B,
24G, 25E, G). Glandular cell outlets type 1 present in paradorsal and ventromedial positions. Small,
rounded sieve plates in lateral accessory positions (Fig. 25C). Pectinate fringe as on preceding segment.
Cuticular hairs shorter, but cuticular hair covering similar to as on preceding segment, except middorsal
and paradorsal areas covered with very short and thin non-bracteate hairs (Fig. 25G) similar to ones
present on lateral halves of sternal plates, posteriorly expanding into subdorsal areas.
SeGment 10. With laterodorsal tubes located near posterior segment margin (Figs 23A, 24G, 25G–H).
Sensory spots present in subdorsal and ventrolateral positions (Figs 24G, 25G–H). Glandular cell outlets
type 1 present as one middorsal and pair of ventromedial ones. Cuticular hairs scarcer than on preceding
segment. Central part of tergal plate covered with short and thin non-bracteate hairs as on preceding
segment (Fig. 25G–H). Posterior segment margin of tergal plate straight, with much shorter fringe
and narrower tips than those on preceding segment (Fig. 25G–H); margins of sternal plates extend
midventrally (Fig. 23B).
SeGment 11. With pair of lateral terminal spines (Fig. 23C). Males with three pairs of penile spines;
dorsal and ventral penile spines thin and exible tubes, whereas median ones markedly thicker, conical
and stout (Figs 23A, 24H, 25G–H). Female condition unknown. Sensory spots present in paradorsal
Table 17. Summary of nature and location of sensory spots, glandular cell outlets, tubes and spines
arranged by series in Echinoderes legolasi sp. nov.
Position segment MD PD SD LD ML SL LA LV VL VM
1 gco1 ss ss gco2 gco1
2 gco1 gco2 gco2,ss gco2 gco2 gco1,ss
3 gco1 ss ss gco1
4 ac gco1 gco1
5 gco1 ss tu gco1
6 ac gco1,ss ss ac gco1,ss
7 gco1 ss ss ac gco1,ss
8 ac gco1 ss gco2 ac gco1
9 gco1 ss ss si ac ss gco1
10 gco1 ss tu ss gco1
11 pr ss ss pe ×3(♂) lts
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
65
and subdorsal positions; subdorsal pair located near posterior margin of tergal extensions (Figs 23A,
25H). Middorsal protuberance-like structure extends from intersegmentary joint (Figs 23A, 25G).
Segment devoid of cuticular hairs, but with dense covering of short hair-like extensions in dorsal areas
of tergal plates. Short fringes covering margins of tergal and sternal plates. Tergal extensions short,
sternal extensions do not extend beyond tergal extensions (Figs 23A–B, 24H, 25G).
Distribution
Hikurangi slope and Seamount 310, 728–985 m b.s.l. See Fig. 1 for a geographic overview of stations
and Table 1 for station and specimen information.
Taxonomic remarks on Echinoderes legolasi sp. nov.
Echinoderes legolasi sp. nov. is one of 34 species (including E. frodoi sp. nov., E. samwisei sp. nov.,
E. leduci sp. nov., and E. aragorni sp. nov.) having middorsal spines on segments 4, 6 and 8, but can
easily be distinguished from all other congeners by its unique patterns of spines, tubes and glandular
cell outlets type 2. The most exclusive feature of E. legolasi is the presence of four pairs of glandular
cell outlets type 2 on segment 2. This character is shared only with one other species, i.e., E. anniae
described by Sørensen et al. (2018) from the United States west coast. Except for the identical pattern
of cuticular structures on segment 2, both species also share the presence of glandular cell outlets type 2
in midlateral/sublateral positions on segments 1 and 8. However, E. anniae can be discriminated from
E. legolasi by its tube pattern, lacking the lateroventral tubes on segment 5. Instead, E. anniae has
glandular cell outlets type 2 in this position (Sørensen et al. 2018). Furthermore, E. legolasi possesses
short tergal extensions, which differ considerably from the relatively long and narrow tergal extensions
present in E. anniae, and signicantly shorter lateral terminal spines in relation to the trunk length (LTS/
TL = 51% in E. legolasi vs 99% in E. anniae).
The new species shows most resemblance to Echinoderes hamiltonorum Sørensen et al., 2018.
Echinoderes hamiltonorum is also described from deep waters (> 3000 m deep) off the United States
west coast (Sørensen et al. 2018). The two species have almost identical arrangements of cuticular
structures, a highly similar trunk appearance, and they share the shape of the tergal extensions, the
presence of a protuberance protruding from the intersegmental joint between segments 10 and 11, and
an even length and shape of the pectinate fringes. The main difference between the two species is the
possession of either tubes or glandular cell outlets type 2 in ventrolateral positions on segment 2. Both
species have subdorsal, laterodorsal and sublateral glands on segment 2, but in ventrolateral positions
E. hamiltonorum has tubes instead of glandular cell outlets type 2 as in E. legolasi sp. nov. Both structures
(glands vs tubes) are easily distinguishable in SEM, but the ventrolateral tubes in E. hamiltonorum are
hard to visualize in LM (Sørensen et al. 2018). The attachment point of the tubes might resemble the
margins of the outlets, which might potentially be a source of confusion during the identication process.
Therefore, the easiest way to distinguish the species would be through SEM examination. However, even
without access to SEM information, it should be possible to distinguish E. legolasi and E. hamiltonorum
by the location of type 2 glands on segment 8 and some morphometric details. Echinoderes legolasi has
glandular cell outlets type 2 in midlateral positions on segment 8, whereas these structures are displaced
to sublateral positions in E. hamiltonorum (Sørensen et al. 2018). Furthermore, the new species has a
shorter trunk (175 µm vs 233 µm) and has a proportionally longer middorsal spine on segment 8. In
E. hamiltonorum, the middorsal spines on segments 6 and 8 are comparable in length (74 µm vs 75 µm,
respectively), while the spine on segment 6 is noticeably shorter than the one on segment 8 in E. legolasi
(51 µm vs 65 µm, respectively).
Hence, in summary, E. legolasi sp. nov. and E. hamiltonorum can be distinguished by the presence of
ventrolateral glands type 2 on segment 2 and sublateral glands type 2 on segment 8 in E. legolasi, and in
the general combination of spine and glandular cell outlet type 2 patterns.
European Journal of Taxonomy 844: 1–108 (2022)
66
Echinoderes aragorni sp. nov.
urn:lsid:zoobank.org:act:88723AE9-DE79-4A1B-840E-3E23F83CA477
Figs 26–28; Tables 18–19
Diagnosis
Echinoderes with spines in middorsal position on segments 4, 6 and 8, and spines in lateroventral
positions on segments 8 and 9. Tubes present in lateroventral positions on segment 5. Glandular cell
outlets type 2 present in paradorsal positions on segments 2 and 3, subdorsal and laterodorsal positions
on segments 2 to 9, midlateral positions on segments 3 to 5 and 9, sublateral positions on segments 2 and
6 to 8, and in lateral accessory positions on segments 5, 8 and 9. Furthermore, lateroventral glandular
cell outlets type 2 present on segments 3, 4, 6, 7, and ventrolateral ones on segment 2. Males with
additional pair of glandular cell outlets type 2 in laterodorsal positions on segment 10. A protuberance-
like structure emerges between segments 10 and 11 in middorsal position.
Etymology
The species name refers to Aragorn – King of Arnor and Gondor, one of the characters in J.R.R. Tolkien’s
“The Lord of the Rings”. Aragorn was a condant of Gandalf, a great warrior and the true leader of the
Fellowship of the Ring.
Material examined
Holotype
NEW ZEALAND • ♀; Hikurangi Slope, stn TAN1004/38; 41.5937° S, 175.8532° E; 1121 m b.s.l.; Apr.
2010; NIWA TAN1004 Voyage; soft sediment; NIWA-159423. Mounted for LM in Fluoromount G on
HS slide.
Paratypes
NEW ZEALAND • 1 ♀; Hikurangi Slope, stn TAN1004/4; 41.6837° S, 175.6642° E; 1046 m b.s.l.; Apr.
2010; NIWA TAN1004 Voyage; soft sediment; NIWA-159424. Mounted for LM in Fluoromount G on
glass slide • 1 ♀; Honeycomb Canyon, stn TAN1004/58; 41.4080° S, 175.8977° E; 1171 m b.s.l.; Apr.
2010; NIWA TAN1004 Voyage; soft sediment; NHMD-920115. Mounted as holotype.
Additional material
NEW ZEALAND • 1 ♀; Hikurangi Slope, stn TAN1004/44; 41.5258° S, 175.8003° E; 728 m b.s.l.;
Apr. 2010; NIWA TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted for
SEM • 1 ♂; Honeycomb Canyon, stn TAN1004/62; 41.4760° S, 175.9477° E; 1171 m b.s.l.; Apr. 2010;
NIWA TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted for SEM.
Description
General. Adults with head, neck and eleven trunk segments (Figs 26–28). Overview of measurements
and dimensions in Table 18. Distribution of cuticular structures, i.e., sensory spots, glandular cell outlets,
spines and tubes, summarized in Table 19. No details regarding scalid arrangement and morphology
could be provided.
neck. With 16 placids. Midventral placid broadest, 9 µm in width and 10 µm in length, whereas all
others narrower, measuring 6 µm in width at bases and 10 µm in length, similar in size (Figs 27B–C,
28F). Trichoscalid plates well developed.
SeGment 1. Consists of complete cuticular ring. Sensory spots located medially on segment, in subdorsal
and laterodorsal positions (Figs 26A, 27B, 28B). Glandular cell outlets type 1 present in middorsal and
ventrolateral positions (Figs 26A–B, 27B–C). Furthermore, row of unusual cuticular openings present
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
67
Fig. 26. Line art illustrations of Echinoderes aragorni sp. nov. A. ♀, dorsal view. B. ♀, ventral view.
C. ♂, segments 10–11, dorsal view. D. ♂, segments 10–11, ventral view. Abbreviations: see Material
and methods.
European Journal of Taxonomy 844: 1–108 (2022)
68
near anterior segment margin in paradorsal, subdorsal, midlateral and ventromedial positions. Openings
elongate, or ‘slit-like’, and without any other cuticular characteristic, thus resembling neither sensory
spots, osculi, nor glandular cell outlets of any kind. Due to uncertain nature of these openings, they will
simply be referred to as ‘slit-like openings’ (Figs 26A–B, 27B–C, 28B–C). Segment devoid of cuticular
hairs, except for a few very long hairs arranged around sensory spots. Posterior segment margin almost
straight, forming pectinate fringe. Fringe with well-developed long and exible tips, homogenous along
segment margin (Fig. 28B–C).
SeGment 2. Consists of complete cuticular ring. Glandular cell outlets type 2 present in paradorsal,
subdorsal, laterodorsal, sublateral and ventrolateral positions (Figs 26A–B, 27B–C, 28B, D). Sensory
Fig. 27. Light micrographs showing overview and details of Echinoderes aragorni sp. nov. A–B,
D–F. ♀, holotype (NIWA-159423). C. ♀, paratype (NIWA-159424). A. Dorsal overview. B. Segments 1
to 6, dorsal view. C. Segments 1 to 6, ventral view. D. Ventral overview. E. Segments 5 to 9, dorsal view.
F. Segments 7 to 11, ventral view. Abbreviations: see Material and methods.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
69
spots in middorsal, laterodorsal and ventromedial positions (Figs 26A–B, 27B, 28B, D). Glandular cell
outlets type 1 present in middorsal and ventromedial positions. Pachycyclus of anterior segment margin
of regular thickness, without interruption. Secondary pectinate fringe present near anterior
Fig. 28. Scanning electron micrographs showing overviews and details of Echinoderes aragorni
sp. nov. A. Lateral overview. B. Segments 1 to 2, dorsolateral view. C. Segments 1 to 5, ventral
view. D. Segments 3 to 5, dorsolateral view. E. Segments 3 to 6, lateral view. F. Ventral overview.
G. Segments 5 to 7, dorsolateral view. H. Segments 5 to 7, ventral view. I. Segments 4 to 6 of male,
dorsolateral view. J. Segments 9 to 10, dorsal view. K. Segments 10 to 11 of female, dorsal view.
L. Segments 10 to 11 of male, dorsal view. M. Segments 7 to 8, dorsolateral view. N. Segments 7 to 9,
ventral view. Abbreviations: see Material and methods.
European Journal of Taxonomy 844: 1–108 (2022)
70
segment margin of this and following segments. Cuticular hairs evenly distributed across segment.
Posterior segment margin almost straight; pectinate fringe tips as on preceding segment.
SeGment 3. Present segment, and eight remaining ones, consist of one tergal and two sternal plates
(Figs 26A–B, 27C, E–F, 28D). Pachycyclus of anterior segment margin of regular thickness, interrupted
at tergosternal and midsternal junctions and middorsally, on this and following seven segments.
Segment with glandular cell outlets type 2 in paradorsal, subdorsal, laterodorsal, midlateral and
lateroventral positions (Figs 26A–B, 27B–C, 28C–E). Sensory spots in subdorsal, midlateral and
ventromedial positions (Figs 27B, 28D–E). Glandular cell outlets type 1 not observed. On this and
following three segments cuticular hairs evenly distributed across tergal plate, except for hairless
areas located close to midlateral line; ventromedial and paraventral areas on this and following ve
segments with thinner and much shorter, non-bracteate hairs. Posterior segment margin straight,
terminating in pectinate fringe with fringe tips as on preceding segments.
SeGment 4. With spine in middorsal position (Figs 26A, 27B). Glandular cell outlets type 2 present in
subdorsal, laterodorsal, midlateral and lateroventral positions (Figs 26A–B, 27B–C, 28D–E). Sensory
spots or glandular cell outlets type 1 not present. Pachycycli, pectinate fringe and cuticular hairs as on
preceding segment.
Table 18. Measurements from light microscopy of Echinoderes aragorni sp. nov. (in µm) from the
Hikurangi Margin, including number of measured specimens (n) and standard deviation (SD).
Character n Range Mean SD
TL 3 171–232 196 32.0
MSW-7 2 37–40 38 1.8
MSW-7/TL 2 20–23% 22% 2.2%
SW-10 2 30–33 32 2.5
SW-10/TL 2 16–20% 18% 2.4%
S1 3 22–25 23 1.7
S2 3 19–22 20 1.6
S3 3 22–25 23 1.9
S4 3 22–28 25 3.5
S5 3 25–30 28 2.4
S6 3 27–34 29 3.9
S7 3 28–36 32 3.8
S8 3 32–36 33 1.9
S9 3 33–35 34 0.8
S10 3 26–30 27 2.3
S11 3 18–20 19 1.6
MD4 (ac) 3 34–43 38 4.6
MD6 (ac) 3 60–65 62 2.8
MD8 (ac) 3 67–72 70 2.6
LV8 (ac) 3 25–32 29 3.5
LV9 (ac) 3 29–39 34 5.1
LTS 3 78–98 89 9.8
LTS/TL 3 39–53% 46% 7.1%
LTAS 3 29–36 33 3.8
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
71
SeGment 5. With tubes in lateroventral positions (Figs 26B, 27C, 28C, H). Glandular cell outlets type 2
located in subdorsal, laterodorsal, midlateral and lateral accessory positions (Figs 26A–B, 27C, E,
28E,G–H). Sensory spots present in subdorsal, midlateral and ventromedial positions; in holotype and
other female specimens midlateral sensory spots located closer to sublateral line, on outer/lateral side of
midlateral glandular cell outlets type 2 (Fig. 28E), while in male specimen this pair of sensory spots on
dorsal side of midlateral glands (Fig. 28I). Segment otherwise as segment 4.
SeGment 6. With spine in middorsal position only (Figs 26A–B, 27A, E, 28A, G). Glandular cell outlets
type 2 present in subdorsal, laterodorsal, sublateral and lateroventral positions (Figs 26A–B, 27C, E,
28G–H). Sensory spots located in paradorsal, midlateral and ventromedial positions (Figs 26A–B, 28G–
H). Tips of pectinate fringe of posterior segment margin as on preceding segments. Segment otherwise
as segment 5.
SeGment 7. Without spines or tubes (Figs 26B, 27E). Glandular cell outlets type 2 and sensory spots as
on segment 6 (Figs 26A–B, 27E–F, 28G–H), but ventromedial pair of sensory spots located closer to
ventrolateral line than on preceding segment (Fig. 28H). Cuticular hair covering as on segment 6, except
for paraventral areas with much shorter and thinner hairs than on preceding segments.
SeGment 8. With spines in middorsal and lateroventral positions (Figs 26A–B, 27G, 28E). Glandular cell
outlets type 2 present in subdorsal, laterodorsal, sublateral and lateral accessory positions (Figs 26A–B,
27E–F, 28M–N). Sensory spots present in paradorsal positions only (Fig. 28M). Pectinate fringe of
posterior segment margin with slightly narrower and shorter tips on the dorsal side. Segment otherwise
as segment 7.
SeGment 9. With spines in lateroventral positions (Figs 26B, 27F, 28N). Glandular cell outlets type 2
present in subdorsal, laterodorsal, midlateral and lateral accessory positions (Figs 26A–B, 27E–F, 28J,
N); subdorsal pair situated close to paradorsal area (Fig. 28J). Sensory spots present in paradorsal,
laterodorsal, sublateral and ventrolateral positions; laterodorsal pair located more anteriorly on
segment than other pairs (Figs 26A–B, 28J). Small, rounded sieve plates located in lateral accessory
positions (Fig. 27F). Pectinate fringe with shorter tips than on preceding segment, homogenous along
segment margin. Cuticular hair covering of tergal plate similar to that on preceding segment, but with
broader area of short and thin non-bracteate hairs, covering paradorsal and subdorsal areas; sternal
plates devoid of hairs in paraventral and ventromedial areas.
Table 19. Summary of nature and location of sensory spots, glandular cell outlets, tubes and spines
arranged by series in Echinoderes aragorni sp. nov.
Position segment MD PD SD LD ML SL LA LV VL VM
1 gco1 so ss,so ss so gco1 so
2 gco1,ss gco2 gco2 gco2,ss gco2 gco2 gco1,ss
3 gco2 ss,gco2 gco2 gco2,ss gco2 ss
4 ac gco2 gco2 gco2 gco2
5 gco2,ss gco2 gco2,ss gco2 tu ss
6 ac ss gco2 gco2 ss gco2 gco2 ss
7 ss gco2 gco2 ss gco2 gco2 ss
8 ac ss gco2 gco2 gco2 gco2 ac
9 ss gco2 ss,gco2 gco2 ss gco2,si ac ss
10 gco1 ss gco2(♂) ss
11 gco1 ×2,pr ss ss pe ×3(♂) ltas(♀) lts
European Journal of Taxonomy 844: 1–108 (2022)
72
SeGment 10. With laterodorsal glandular cell outlets type 2 located near posterior segment margin in
males; females without glandular cell outlets type 2 on this segment (Figs 26A,,C, 28K–L). Sensory
spots present in subdorsal and ventrolateral positions (Fig. 28L). Glandular cell outlet type 1 present in
middorsal position. Cuticular hairs sparser than on preceding segment. Central part of tergal plate almost
completely hairless, sternal plate hair covering as on preceding segment. Posterior segment margin of tergal
plate straight, with much shorter fringe and narrower tips than those on preceding segment (Fig. 28K–
L); margins of sternal plates extend midventrally (Fig. 26B).
SeGment 11. With pair of lateral terminal spines (Fig. 27D, F). Males with three pairs of penile spines;
dorsal and ventral penile spines thin and exible tubes, whereas median ones markedly thicker, conical
and stout (Figs 26C, 28L); females with lateral terminal accessory spines (Figs 26A, 27F, 28K). Sensory
spots present in paradorsal and subdorsal positions; subdorsal pair located near posterior margin of
tergal extensions (Figs 26A, 28K). Pair of glandular cell outlets type 1 present in middorsal position,
covered by middorsal protuberance-like structure extending from intersegmentary joint (Figs 26A,C,
28K–L). Segment devoid of cuticular hairs, but with dense covering of short hair-like extensions on
dorsal areas of tergal plates. Short fringes covering margins of tergal and sternal plates; fringe tips
slightly longer near insertion of lateral terminal spines. Tergal extensions triangular, sternal extensions
do not extend beyond tergal extensions.
Distribution
Hikurangi slope and Honeycomb Canyon, 670–1171 m b.s.l. See Fig. 1 for a geographic overview of
stations and Table 1 for station and specimen information.
Taxonomic remarks on Echinoderes aragorni sp. nov.
Echinoderes aragorni sp. nov. can very easily be distinguished from all other congeners by the number
and arrangement of glandular cell outlets type 2. No less than 34 pairs of glandular cell outlets type 2 (35
in males) are present in E. aragorni, which is not present in any other kinorhynch species. The only other
species displaying a somewhat similar trait is E. orestauri Pardos et al., 2016 with 30 fringed tubules
distributed on segments 2 to 9 (Pardos et al. 2016a). It seems likely that glandular cell outlets type 2 and
fringed tubes are homologous, but they still differ morphologically since only the fringed tubes form
actual external appendages. In contrast, the glands in E. aragorni are relatively large openings (although
smaller in male specimens than in females: see Fig. 28I vs Fig. 28J), easily distinguishable in both SEM
and LM and present on segments 2 to 10.
A second rare feature in E. aragorni sp. nov. is the presence of lateroventral tubes/spines on segments 5,
8 and 9 only. This feature is shared with three other species only: E. caribiensis Kirsteuer, 1964,
E. lusitanicus and E. skipperae Sørensen & Landers, 2014 (Kirsteuer 1964; Neves et al. 2016;
Sørensen & Landers 2014). However, none of these species can be confused with E. aragorni, as they
do not possess even a single pair of glandular cell outlets type 2. Moreover, E. caribiensis belongs to
the E. coulli species group and is characterized by the absence of middorsal spines, while in contrast
E. lusitanicus has ve middorsal spines. Only E. skipperae has, similar to the new species, a middorsal
spine arrangement with spines present on segments 4, 6 and 8.
The presence of slit-like openings on segment 1 is also unique to E. aragorni sp. nov. Other species
show an occurrence of cuticular structures on segment 1 like tubes or glandular cell outlets type 2
(e.g., E. frodoi sp. nov., E. legolasi sp. nov., E. anniae, and E. hamiltonorum) (Sørensen et al. 2018;
present study), but the structures observed in E. aragorni have not been observed before. Although it
may represent another type of glandular cell outlets – or a modied kind of glandular cell outlets type 2
– we are currently uncertain about their nature and function. Therefore, it is difcult to discuss this
observation further, but it is obviously a structure that deserves more attention in future works.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
73
Echinoderes galadrielae sp. nov.
urn:lsid:zoobank.org:act:CD592202-3712-43B6-9F4E-DE5670D87EE7
Figs 29–31; Tables 20–21
Diagnosis
Echinoderes with spines in middorsal position on segments 4 to 8, and spines in lateroventral positions on
segments 6 to 9. Tubes present in lateral accessory positions on segment 5. Glandular cell outlets type 2
in subdorsal, laterodorsal, sublateral and ventrolateral positions on segment 2, midlateral positions on
segment 5 and sublateral positions on segment 8. Males with minute laterodorsal tubes on segment 10;
females with nearly reduced tubes. Segment 11 composed of two tergal and two sternal plates.
Etymology
The species name refers to Galadriel – ‘Lady’ of Lothlórien, one of the greatest of the elves in Middle-
Earth, and a character in J.R.R. Tolkien’s “The Lord of the Rings” and “Silmarillion”. Galadriel helped
the Fellowship of the Ring signicantly in achieving their goals, hosting them after their escape from the
mines of Moria and giving each member a valuable gift for their onward journey.
Material examined
Holotype
NEW ZEALAND • ♀; Campbell Canyon, stn TAN1004/126; 42.1422° S, 174.5492° E; 1495 m b.s.l.;
Apr. 2010; NIWA TAN1004 Voyage; soft sediment; NIWA-159425. Mounted for LM in Fluoromount
G on HS slide.
Paratypes
NEW ZEALAND • 2 ♀♀; Pahaua Canyon, stn TAN1004/12; 41.5508° S, 175.7250° E; 1350 m b.s.l.;
Apr. 2010; NIWA TAN1004 Voyage; soft sediment; NHMD-921496 to 921497. Mounted for LM in
Fluoromount G on glass slide • 1 ♀; Seamount 766, stn TAN1004/132; 42.1345° S, 174.5850° E;
1453 m b.s.l.; Apr. 2010; NIWA TAN1004 Voyage; soft sediment; NIWA-159426. Mounted as holotype.
Additional material
NEW ZEALAND • 2 ♀♀; Hikurangi Slope, stn TAN1004/76; 41.6833° S, 175.6500° E; 1282 m b.s.l.;
Apr. 2010; NIWA TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted for
SEM • 3 ♀♀, 1 ♂; Pahaua Canyon, stn TAN1004/27; 41.4983° S, 175.7043° E; 1013 m b.s.l.; Apr. 2010;
NIWA TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted for SEM •
1 ♀; Honeycomb Canyon, stn TAN1004/62; 41.4760° S, 175.9477° E; 1171 m b.s.l.; Apr. 2010; NIWA
TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted for SEM.
Description
General. Adults with head, neck and eleven trunk segments (Figs 29–31). Overview of measurements
and dimensions in Table 20. Distribution of cuticular structures, i.e., sensory spots, glandular cell outlets,
spines and tubes, summarized in Table 21. No details regarding scalid arrangement and morphology
could be provided, because introvert of specimens mounted for SEM was fully or partially retracted.
neck. With 16 placids. Midventral placid broadest, 9 µm in width and 11 µm in length, whereas all
others narrower, measuring 6 µm in width at their bases and 11 µm in length, similar in size (Fig. 30C,
31D). Trichoscalid plates are well developed.
SeGment 1. Consists of complete cuticular ring. Sensory spots located anteriorly on segment, in subdorsal
and laterodorsal positions (Figs 29A, 30B, 31B). Glandular cell outlets type 1 present in middorsal, and
ventrolateral positions. Cuticular hairs lightly scattered on dorsal and lateral sides and absent on ventral
European Journal of Taxonomy 844: 1–108 (2022)
74
Fig. 29. Line art illustrations of Echinoderes galadrielae sp. nov. A. ♀, dorsal view. B. ♀, ventral view.
C. ♂, segments 10–11, dorsal view. D. ♂, segments 10–11, ventral view. Abbreviations: see Material
and methods.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
75
side (Fig. 30B–C). Posterior segment margin almost straight along dorsal edge, but slightly extended
posteriorly in midventral position (Fig. 31C). Pectinate fringe with well-developed fringe tips, slightly
shorter and rounded along dorsal margin, and longer and more pointed along lateral and ventral margins
(Fig. 31B–C).
SeGment 2. Consists of complete cuticular ring. Glandular cell outlets type 2 present in subdorsal,
laterodorsal, sublateral and ventrolateral positions (Figs 29A–B, 30B–C, 31B–C); glands on ventral side
located closer to anterior margin of segment (Fig. 31C). Sensory spots located in middorsal, laterodorsal
(two pairs) and ventromedial positions (Fig. 29A–B); sensory spots on this and following segments very
small. Glandular cell outlets type 1 present in middorsal and ventromedial positions. Pachycyclus of
anterior segment margin of regular thickness, interrupted in middorsal position. Secondary pectinate
fringe present near anterior segment margin of this and following segments. Long cuticular hairs lightly
scattered on dorsal and lateral sides; ventral side with few hairs. Paraventral and midventral areas in
this and following eight segments hairless. Posterior segment margin straight along dorsal side, while
extended posteriorly in midventral position. Fringe tips shorter than on preceding segment.
SeGment 3. Present segment, and eight remaining ones, consist of one tergal and two sternal plates
(Figs 29A–B, 30A, C, 31A). Segment with sensory spots located in subdorsal and midlateral positions
(Figs 30B, 31B), and glandular cell outlets type 1 in middorsal and ventromedial positions. Pachycyclus
of anterior segment margin of regular thickness, interrupted at tergosternal and midsternal junctions
Fig. 30. Light micrographs showing overview and details of Echinoderes galadrielae sp. nov. A. ♀,
paratype (NHMD-921497). B–C. ♀, holotype (NIWA-159425). D. ♀, paratype (NHMD-921496).
E. ♀, paratype (NIWA-159426). A. Ventral overview, with inset showing close-up of segment 11.
B. Segments 1 to 5, dorsal view. C. Segments 1 to 7, ventral view. D. Segments 6 to 8, lateral view.
E. Segments 8 to 11, lateral view. Abbreviations: see Material and methods.
European Journal of Taxonomy 844: 1–108 (2022)
76
and middorsally, on this and following seven segments. Cuticular hairs more densely distributed across
tergal plate, but otherwise as on preceding segment. Posterior segment margin straight, terminating in
pectinate fringe with pointed fringe tips, longer than as on preceding segments.
SeGment 4. With spine in middorsal position (Figs 29A, 31D). Segment without sensory spots, but with
glandular cell outlets type 1 in paradorsal and ventromedial positions. Pachycycli, pectinate fringe and
cuticular hairs as on preceding segment.
Fig. 31. Scanning electron micrographs showing overviews and details of Echinoderes galadrielae
sp. nov. A. Ventral overview. B. Segments 1 to 5, dorsolateral view. C. Segments 1 to 2, lateroventral view.
D. Dorsal overview. E. Segments 5 to 7, ventral view. F. Segments 4 to 7, lateral view. G. Segments 8 to
10, dorsolateral view. H. Segments 10 to 11 of male, dorsolateral view. I. Segments 10 to 11 of female,
dorsal view with insets showing close-up of reduced laterodorsal tubes on segment 10. Abbreviations:
see Material and methods.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
77
SeGment 5. With spine in middorsal position and tubes in lateral accessory positions (Figs 29A–B, 30C,
31E–F). Glandular cell outlets type 2 present in midlateral positions, being similar in size and shape
to those on segment 2. Sensory spots located subdorsally, and glandular cell outlets type 1 present in
paradorsal and ventromedial positions. Segment otherwise as segment 4.
SeGment 6. With spines in middorsal and lateroventral positions (Figs 29A–B, 30A ,C, 31A, D, F).
Sensory spots present in paradorsal, midlateral and ventromedial positions (Figs 29A–B, 30C, 31F).
Glandular cell outlets type 1 present in paradorsal and ventromedial positions (Figs 29A–B, 30C). Tips
of pectinate fringe of posterior segment margin as on preceding segments. Cuticular hairs and posterior
segment margin as on preceding segment.
Table 20. Measurements from light microscopy of Echinoderes galadrielae sp. nov. (in µm) from the
Hikurangi Margin, including number of measured specimens (n) and standard deviation (SD). Left part
of the table shows ranges for E. galadrielae sp. nov., right part shows ranges for Echinoderes sp. aff.
E. galadrielae/E. beringiensis with shorter tergal extensions.
Character E. galadrielae sp. nov. E. sp. aff. E. galadrielae/beringiensis
n Range Mean SD n Range Mean SD
TL 4 210–235 224 8.0 7 196–211 208 8.3
MSW-8 4 47–50 49 1.5 6 41–48 44 2.9
MSW-8/TL 4 21–23% 22% 0.7% 620–23% 21% 1.1%
SW-10 4 41–43 42 0.5 6 32–38 34 1.9
SW-10/TL 4 18–20% 19% 0.4% 616–18% 17% 0.7%
S1 4 26–28 27 0.9 7 22–25 23 0.9
S2 4 24–25 25 0.6 7 19–24 21 1.6
S3 4 24–26 25 0.9 7 20–27 23 2.2
S4 4 25–28 27 1.2 7 25–31 26 2.3
S5 4 30–37 34 2.8 7 24–32 28 2.6
S6 4 33–41 36 3.8 7 28–35 30 2.3
S7 4 37–43 40 2.5 7 29–35 32 2.1
S8 4 38–42 40 1.6 7 32–37 35 2.2
S9 4 36–41 38 1.9 7 30–35 32 1.4
S10 4 30–35 33 2.5 7 24–32 29 3.4
S11 4 37–40 38 1.4 6 24–30 27 3.9
MD4 (ac) 3 25–31 29 2.4 7 29–46 40 5.6
MD5 (ac) 3 33–36 36 2.7 6 45–61 53 6.0
MD6 (ac) 4 46–55 51 3.3 6 58–64 60 2.3
MD7 (ac) 3 54–56 55 0.9 6 61–75 69 5.2
MD8 (ac) 4 62–69 66 2.5 6 77–87 81 3.7
LV6 (ac) 4 29–37 32 3.4 7 32–44 37 4.4
LV7 (ac) 3 45–47 46 0.8 7 30–46 42 6.1
LV8 (ac) 4 45–49 47 1.6 7 38–52 44 7.9
LV9 (ac) 4 41–44 43 1.2 6 36–43 38 3.3
TE 4 28–33 32 2.2 7 14–21 18 2.8
TE/TL 4 13–15% 14% 0.9% 77–10% 9% 1.5%
LTS 4 178–214 199 13.4 7 181–254 220 32.2
LTS/TL 4 76–94% 87% 8.0% 786–125% 106% 15.1%
LTAS 4 50–60 55 3.6 4 35–55 42 8.6
European Journal of Taxonomy 844: 1–108 (2022)
78
SeGment 7. With spines in middorsal and lateroventral positions (Figs 29A–B, 31D–F). Sensory spots
present in paradorsal and midlateral positions (Figs 29A–B, 31F), and glandular cell outlets type 1 in
paradorsal and ventromedial positions (Figs 29A–B, 30E). Cuticular hair covering as on preceding
segment. Segment otherwise as segment 6.
SeGment 8. With spines in middorsal and lateroventral positions (Figs 29A–B, 30E, 31D, G). Glandular
cell outlets type 2 located in sublateral positions; gland very conspicuous, larger than those on segments 2
and 5 (Figs 30D–E, 31G). Sensory spots present in paradorsal positions, and glandular cell outlets type 1
in paradorsal and ventromedial positions (Fig. 30E). Pectinate fringe of posterior segment margin as on
preceding segment.
SeGment 9. With spines in lateroventral positions (Figs 29B, 31A, G). Sensory spots located in paradorsal,
subdorsal, midlateral and ventrolateral positions; subdorsal pair situated more anterior than others
(Figs 29A–B, 31G). Glandular cell outlets type 1 present in paradorsal and ventromedial positions. Small,
rounded sieve plates located in sublateral positions (Figs 30E, 31G). Pectinate fringe as on preceding
segment. Cuticular hair covering and posterior segment margin similar to those on preceding segment.
SeGment 10. With sensory spots in subdorsal and ventrolateral positions (Fig. 31G–H). Glandular cell
outlets type 1 in ventromedial positions and as pair in middorsal position (Fig. 30E). Males with very
short laterodorsal tubes, emerging through slit-like, fringed opening; females with similar slit-like,
fringed opening, but with tubes being even further reduced, and either not visible at all or only visible
as plate-like projection emerging through opening (Fig. 31H–I). Cuticular hairs very scarce on both
tergal and sternal plates (Fig. 31A, G–I). Pectinate fringe of posterior margin on dorsal and lateral sides
with markedly shorter fringe tips as on preceding segments, but with longer tips on paraventral areas.
Margins of sternal plates extend midventrally.
SeGment 11. With pair of lateral terminal spines (Fig. 29A). Females with lateral terminal accessory
spines (Figs 29A, 30A, 31I); males with three pairs of penile spines (Figs 29C, 31H), with dorsal- and
ventralmost penile spines being slender and tubular, with abrupt narrowings about ¼ from their distal
tips; median ones stout and triangular. Sensory spots present in paradorsal positions, near posterior
margins of sternal plate (Fig. 31H). A pair of glandular cell outlets type 1 present in middorsal position.
Segment devoid of cuticular hairs, but has small patches with short and tiny hair-like extensions in
Table 21. Summary of nature and location of sensory spots, glandular cell outlets, tubes and spines
arranged by series in Echinoderes galadrielae sp. nov.
Position segment MD PD SD LD ML SL LA LV VL VM
1 gco1 ss ss gco1
2 gco1,ss gco2 ss,gco2,ss gco2 gco2 gco1,ss
3gco1 ss ss gco1
4ac gco1 gco1
5 ac gco1 ss gco2 tu gco1
6 ac gco1,ss ss ac gco1,ss
7 ac gco1,ss ss ac gco1
8 ac gco1,ss gco2 ac gco1
9 gco1,ss ss ss si ac ss gco1
10 gco1 ×2 ss tu(♂) ss gco1
11 gco1 ×2 ss pe ×3(♂) ltas(♀) lts
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
79
paradorsal positions. Short fringes covering margins of sternal plates. Segment composed of two tergal
and two sternal plates (Fig. 31H–I) Tergal extensions signicantly elongated and posteriorly projecting
(Figs 30A, 31H–I); sternal extensions short and rounded.
Distribution
Hikurangi Margin, from slope, through canyon, and seamount habitats, 1013–1495 m b.s.l. See Fig. 1
for a geographic overview of stations and Table 1 for station and specimen information.
Taxonomic remarks on Echinoderes galadrielae sp. nov.
Echinoderes galadrielae sp. nov. displays a very common spine pattern, with ve middorsal ones on
segments 4 to 8, and lateroventral spines on segments 6 to 9, which is shared with almost 50 species
of Echinoderes. However, what makes E. galadrielae differ from most of its congeners is the presence
of tubes on segment 5 in lateral accessory positions rather than in lateroventral positions. Such tube
displacement is actually a very uncommon trait, observed for only four other species, i.e., E. bathyalis
Yamasaki et al., 2018, E. drogoni Grzelak & Sørensen, 2017 in Grzelak & Sørensen (2018), E. ferrugineus
Zelinka, 1928 and E. beringiensis Adrianov & Maiorova, 2022 (Grzelak & Sørensen 2018; Yamasaki
et al. 2018c; Yamasaki & Dal Zotto 2019; Adrianov & Maiorova 2022). Echinoderes galadrielae
can easily be distinguished from E. bathyalis though, by its number and positions of glandular cell
outlets type 2: E. galadrielae has four pairs of those glands on segment 2, and one pair midlaterally on
segment 5 and sublaterally on segment 8, whereas E. bathyalis shows only one pair on segment 2, an
absence of glands on segment 5 and lateral accessorily on segment 8 (Yamasaki et al. 2018c). Moreover,
E. bathyalis possesses markedly different tergal extensions, and all its acicular spines are longer.
Echinoderes ferrugineus and E. galadrielae sp. nov. share the same number and arrangement of type 2
glands on segments 2, 5 and 8 (on the latter segment in midlateral rather than sublateral position though),
but E. ferrugineus has two additional pairs of glandular cell outlets type 2 on segment 4, in subdorsal
and midlateral positions (Yamasaki & Dal Zotto 2019). Moreover, in addition to its very different tergal
extensions, E. ferrugineus also differs by being larger (304 µm vs 224 µm), and having markedly shorter
lateroventral and lateral terminal spines (for details see Yamasaki & Dal Zotto 2019).
Among the abovementioned species, two Arctic ones, E. drogoni and E. beringiensis, show the closest
resemblance to E. galadrielae sp. nov. The three species share several features, including tube and spine
patterns, number and arrangement of glandular cell outlets type 2 and most sensory spots. They also
share another very characteristic and uncommon feature, which is the middorsal division of the tergal
plate of segment 11 (Grzelak & Sørensen 2018; Adrianov & Maiorova 2022). Nevertheless, despite
overall similarity, E. galadrielae can easily be distinguished from both E. drogoni and E. beringiensis
by its elongated tergal extensions.
Such long and conspicuous tergal extensions, constituting almost 15% of the total trunk length, is in fact
the most prominent trait of E. galadrielae sp. nov. Longer tergal extensions (TE/TL ~ 20%) are known
only for two other species, i.e., E. balerioni Grzelak & Sørensen, 2019 and E. cernunnos Sørensen
et al., 2012 (see Sørensen et al. 2012; Grzelak & Sørensen 2019). Echinoderes balerioni can easily be
discriminated from E. galadrielae by its three middorsal spines on segments 4, 6 and 8 and the lack
of glandular cell outlets type 2. In contrast, E. cernunnos – a species described from Korean waters
– shows a much closer resemblance to the new species. The spine and glandular cell outlets type 2
distributions are almost identical with those in E. galadrielae, and a re-examination of the paratypes
(NHMD-099881 to 099882) and photos of the holotype (INBRIV-0000245082) of E. cernunnos revealed
that both species also have their tubes on segment 5 displaced to lateral accessory positions. The most
conspicuous difference between the species is the presence of midlateral glandular cell outlets type 2 on
segment 7 in E. cernunnos, which are absent in E. galadrielae. Interestingly, E. cernunnos is also one
European Journal of Taxonomy 844: 1–108 (2022)
80
of the few known congeners (together with abovementioned E. drogoni and E. juliae Sørensen et al.,
2018) with a middorsal ssure on segment 11 (Sørensen et al. 2012, 2018; Grzelak & Sørensen 2018)
and is also characterised by tergal extensions not only similar in length, but also in their horn-like shape
(compare Sørensen et al. 2012: g. 6f–h with Fig. 31H–I in the present study). Nonetheless, despite
several similarities and the fact that E. galadrielae and E. cernunnos share several rare character traits,
telling them apart should not be a problem. Except for the different distribution of sensory spots, the
easiest way is to focus on segment 7, which has glandular cell outlets type 2 in E. cernunnos, and on the
lateral terminal spines, which are markedly longer in E. galadrielae (LTS/TL = 87% in E. galadrielae vs
LTS/TL = 23% in E. cernunnos) (Sørensen et al. 2012).
Hence, in summary, E. galadrielae sp. nov. appears to share a number of traits with E. drogoni,
E. cernunnos, and E. beringiensis, including general spine pattern, lateral displacement of the tubes on
segment 5, glandular cell outlet type 2 patterns on segments 2, 5 and 8, and middorsal division of the
tergal plate of segment 11. This combination of rather unusual characters suggests that the four species
are closely related and represent a clade within Echinoderes. Following previous attempts to identify
such species groups within the genus (see, e.g., Yamasaki 2016; Sørensen et al. 2018, 2020), we propose
that these four species should be considered as a monophyletic entity and referred to as the E. cernunnos
species group.
Species with uncertain identities
Echinoderes sp. aff. E. galadrielae/E. beringiensis
Fig. 32; Table 20
Material examined
NEW ZEALAND • 1 ♀; Hikurangi Slope, stn TAN1004/4; 41.6837° S, 175.6642° E; 1046 m b.s.l.;
Apr. 2010; NIWA TAN1004 Voyage; soft sediment; NIWA-159427. Mounted for LM in Fluoromount
G on glass slide • 2 ♂♂; Hikurangi Slope, stn TAN1004/38; 41.5937° S, 175.8532° E; 1121 m b.s.l.;
Apr. 2010; NIWA TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted
for SEM • 1 ♀; Hikurangi Slope, stn TAN1004/76; 41.6833° S, 175.6500° E; 1282 m b.s.l.; Apr. 2010;
NIWA TAN1004 Voyage; soft sediment; NHMD-921503. Mounted for LM in Fluoromount G on HS
slide • 1 ♂; same collection data as for preceding; personal reference collection of MVS. Mounted for
SEM • 1 ♀, 1 ♂; Pahaua Canyon, stn TAN1004/12; 41.5508° S, 175.7250° E; 1350 m b.s.l.; Apr. 2010;
NIWA TAN1004 Voyage; soft sediment; NHMD-921498 to 921499. Mounted for LM in Fluoromount
G on glass slides • 2 ♂♂; Pahaua Canyon, stn TAN1004/27; 41.4983° S, 175.7043° E; 1013 m b.s.l.;
Apr. 2010; NIWA TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted for
SEM • 1 ♂; Honeycomb Canyon, stn TAN1004/53; 41.4563° S, 175.8970° E; 948 m b.s.l.; Apr. 2010;
NIWA TAN1004 Voyage; soft sediment; NHMD-921500. Mounted for LM in Fluoromount G on glass
slide • 1 ♂; Honeycomb Canyon, stn TAN1004/58; 41.4080° S, 175.8977° E; 1171 m b.s.l.; Apr. 2010;
NIWA TAN1004 Voyage; soft sediment; NHMD-921502. Mounted for LM in Fluoromount G on HS
slide • 1 ♀; Seamount 310, stn TAN1004/69; 41.3353° S, 176.1882° E; 670 m b.s.l.; Apr. 2010; NIWA
TAN1004 Voyage; soft sediment; NHMD-921501. Mounted for LM in Fluoromount G on glass slide.
Distribution
Hikurangi Margin, from slope, through canyon, and seamount habitats, 670–1350 m b.s.l. See Fig. 1 for
a geographic overview of stations and Table 1 for station and specimen information.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
81
Brief description and remarks
The examined specimens closely follow the morphology of E. galadrielae sp. nov. in terms of distribution
of spines, tubes, glandular cell outlets, and even the middorsal division of the tergal plate of segment 11.
However, E. galadrielae and E. aff. E. galadrielae/E. beringiensis differ at two important points, i.e.,
spine morphometrics (see Table 20) and the shape of the tergal extensions (compare Figs 30A, 31D,
I with Fig. 32). Middorsal spines were in general longer in this current set of specimens, while the
lateroventral ones were shorter than in E. galadrielae. The unidentied specimens have also markedly
longer lateral terminal spines, which in most cases exceed the trunk length, resulting in markedly higher
LTS/TL ratio in this species compared to E. galadrielae (average LTS/TL: 106% vs 87%, respectively).
The most conspicuous difference between E. galadrielae and these unidentied specimens is expressed
in the tergal extensions, which are exceptionally long in E. galadrielae (Figs 29–31). This stands in
strong contrast to the short, triangular tergal extensions in the unidentied specimens (Fig. 32). These
tergal extensions are much more similar to those in E. beringiensis as well as many other species of
Echinoderes.
We would, on the other hand, hesitate in identifying the specimens as E. beringiensis, since they differ
in the distribution of sensory spots and lack the midlateral glandular cell outlets type 2 that characterizes
females of E. beringiensis (see Adrianov & Maiorova 2022). In addition, E. beringiensis has a longer
trunk (TL: 331 µm vs 208 µm, respectively) and has proportionally shorter lateral terminal spines
(LTS/TL: 50% vs 106%, respectively). In conclusion, these enigmatic specimens appear to represent
an intermediate between E. galadrielae sp. nov. and E. beringiensis, and it would require a molecular
comparison to unveil their true identity.
Fig. 32. Segment 11 and morphological variations of tergal extensions in Echinoderes sp. aff.
E. galadrielae/E. beringiensis. A–C. Light micrographs. D–F. Scanning electron micrographs.
European Journal of Taxonomy 844: 1–108 (2022)
82
Echinoderes sp. aff. E. balerioni
Fig. 33; Tables 22–23
Material examined
NEW ZEALAND • 1 ♂; Seamount 310, stn TAN1004/69; 41.3353° S, 176.1882° E; 670 m b.s.l.; Apr.
2010; NIWA TAN1004 Voyage; soft sediment; NHMD-921971. Mounted for LM in Fluoromount G on
HS slide.
Distribution
Seamount 310, 670 m depth. See Fig. 1 for geographic location of station and Table 1 for station and
specimen information.
Brief description and remarks
Echinoderes with middorsal spines on segments 4, 6, and 8 and spines in lateroventral positions on
segments 6 to 9. Tubes present in subdorsal, sublateral, and ventrolateral positions on segment 2,
lateroventral positions on segment 5, and lateral accessory positions on segment 8. Tergal extensions
conspicuously long and spiniform.
Adult with head, neck and eleven trunk segments. Overview of measurements and dimensions in Table 22.
Only a single specimen was available for LM examination, and thus some cuticular structures such as
sensory spots or glandular cell outlets might not have been identied. Therefore, lack of information
about structures reported in Table 23 should not necessarily be understood as a conrmation of their
absence.
Fig. 33. Light micrographs showing overview and details of Echinoderes sp. aff. E. balerioni, ♂
(NHMD-921971). A. Ventral overview. B. Segments 1 to 5, dorsal view. C. Segments 1 to 5, ventral
view. D. Segments 2 to 5, dorsal view. E. Segments 10 to 11, dorsal view. F. Segments 6 to 10, dorsal
view. G. Segments 6 to 10, ventral view. H. Segments 10 to 11, ventral view. Abbreviations: see Material
and methods.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
83
Segments 1 and 2 composed of complete cuticular rings. Segment 1 with sensory spots in subdorsal
positions, and with glandular cell outlets type 1 in middorsal and ventrolateral positions (Fig. 33B–
C). Posterior segment margin of this and following nine segments with well-developed pectinate
fringe tips. Segment 2 with three pairs of tubes in subdorsal, sublateral and ventrolateral positions
(Fig. 33B–D). Sensory spots observed in ventromedial positions and glandular cell outlets type 1 in
middorsal position only. Cuticular hairs on this and following segments abundant, distributed evenly
around segment. Segments 3 to 11 consists of one tergal and two sternal plates. Segment 3 with sensory
spots in subdorsal positions, and glandular cell outlets type 1 in middorsal and ventromedial positions.
Segment 4 with relatively short (23 µm) middorsal spine (Fig. 33B), and glandular cell outlets type 1
located paradorsally and ventromedially; no sensory spots observed. Pectinate fringe with longer fringe
tips than on preceding segments. Segment 5 with lateroventral tubes (Fig. 33C); sensory spots present in
subdorsal, midlateral and ventromedial positions, and glandular cell outlets type 1 located middorsally
and ventromedially (Fig. 33B–D). Segment 6 with spines in middorsal and lateroventral positions
(Fig. 33F–G), paradorsal, midlateral and ventromedial sensory spots, and glandular cell outlets type 1
located paradorsally and ventromedially (Fig. 33G). Segment 7 with spines in lateroventral positions,
Table 22. Measurements from light microscopy of Echinoderes sp. aff. E. balerioni (in µm) from the
Hikurangi Margin with a comparison of measurements for E. balerioni from its type locality.
Character E. aff. E. balerioni present study E. balerioni Grzelak & Sørensen, 2019
NHMD-921971 (♂) NHMD-202793 (♂)
TL 227 300
MSW-6 47 64
MSW-6/TL 20.7% 21.3%
SW-10 39 52
SW-10/TL 17.2% 17.3%
S1 24 32
S2 21 30
S3 22 31
S4 23 36
S5 24 39
S6 28 45
S7 31 46
S8 34 48
S9 31 47
S10 33 43
S11 52 73
MD4 (ac) 23 23
MD6 (ac) 28 32
MD8 (ac) 50 35
LVS6 (ac) 29 34
LVS7 (ac) 28 40
LVS8 (ac) 33 47
LVS9 (ac) 35 52
TE 40 55
TE/TL 17.6% 18.3%
LTS 148 181
LTS/TL 65.2% 60.3%
European Journal of Taxonomy 844: 1–108 (2022)
84
subdorsal and midlateral sensory spots, and glandular cell outlets type 1 present in middorsal and
ventromedial positions (Fig. 33F–G). Segment 8 with spines in middorsal and lateroventral positions, and
tubes in lateral accessory positions (Fig. 33F–G); sensory spots observed only in paradorsal positions;
glandular cell outlets type 1 present in paradorsal and ventromedial positions. Segment 9 with spines
in lateroventral positions (Fig. 33G); sensory spots located in paradorsal, subdorsal and ventrolateral
positions, and glandular cell outlets type 1 present in paradorsal and ventromedial positions (Fig. 33G);
small sieve plate located in lateral accessory positions (Fig. 33G). Segment 10 with glandular cell outlets
type 1 located in paraventral positions and with two located middorsally (Fig. 33G); no sensory spots
observed. Segment 11 with pair of relatively long lateral terminal spines (Fig. 33A,H). Three pairs of
long penile spines present (Fig. 33E, H). Tergal extensions conspicuously long and spiniform (Fig. 33H).
No cuticular hairs nor any other cuticular structures visible on this segment.
Echinoderes balerioni is a species described from 880 m depth at the Yermak Plateau, north of Svalbard
(Grzelak & Sørensen 2019). There is no other species with acicular spines in middorsal position on
segments 4, 6, and 8, three pairs of tubes on segment 2 and conspicuously long tergal extensions.
The latter constitutes almost 18.3% of the total trunk length and this is the most prominent feature
of E. balerioni. The lengths of the tergal extensions in this species are comparable only with those
of E. cernunnos, E. yamasakii and E. galadrielae sp. nov. – the species with the longest extensions
(Sørensen et al. 2012, 2018). However, all three species are easily distinguished from E. balerioni by
their ve middorsal spines on segments 4 to 8 and the presence of glandular cell outlets type 2.
The individual examined for the present study closely follows the morphology of E. balerioni, in
particular in the appearance of tergal extensions (see Table 22). Most morphometric data, general trunk
appearance and other cuticular structures, such as arrangements of spines and tubes (however, see
below), also followed those reported for the Arctic specimen (compare Table 23 in present contribution
with Grzelak & Sørensen 2019: table 3). Nevertheless, we cannot identify the examined specimen as
E. balerioni with certainty because of certain morphological differences. Considering the geographical
distance between northern Svalbard and New Zealand and the fact that in both cases only a single
specimen was available for LM investigation, we cannot be sure whether the observed discrepancies are
the result of inter-population variations or indicate the presence of two, closely related species.
Due to the fact that no specimen was available for SEM investigation, Grzelak & Sørensen (2019) did
not provide details about sensory spots on segments 2 to 11, and a comparison of their patterns between
Table 23. Summary of nature and location of sensory spots, glandular cell outlets, tubes and spines
arranged by series in Echinoderes sp. aff. E. balerioni.
Position segment MD PD SD ML SL L A LV VL VM PV
1 gco1 ss gco1
2 gco1 tu tu tu ss
3 gco1 ss gco1
4 ac gco1 gco1
5 gco1 ss ss tu gco1,ss
6 ac gco1,ss ss ac gco1,ss
7 gco1 ss ss ac gco1
8 ac gco1,ss tu ac gco1
9 gco1,ss ss si ac ss gco1
10 gco1 × 2 gco1
11 pe × 3 lts
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
85
the specimens from the Arctic and New Zealand is therefore not possible. The major difference between
the specimen from New Zealand and E. balerioni from the Arctic is the presence of tubes on segment 2
in the subdorsal rather than the laterodorsal positions. Both specimens also slightly differ in terms of
morphometric details. The specimen from New Zealand is smaller (TL: 227 µm vs 300 µm) and has a
longer middorsal spine on segment 8 (MDS8: 50 µm vs 35 µm) than E. balerioni from the type locality.
Nevertheless, differences in trunk length do not result in differences in TE/TL or LTS/TL ratios
(Table 22), which conrms the close similarity between specimens. Interestingly, the variation in the
arrangement of the tubes on segment 2 might have resulted from inter-population variations. Recently,
specimens from the Bering Sea, Alaska, clearly similar to E. balerioni, have been investigated by the
rst author of the present study, and all of them possess tubes on segment 2, in subdorsal, laterodorsal,
sublateral and ventrolateral positions (Grzelak, unpubl. obs.). A variation regarding the presence
or absence of tubes has previously been observed for several species, e.g., E. arlis, E. daenerysae,
E. eximus, E. rhaegali and E. dalzottoi sp. nov., and it occurred within the same population, randomly
and independently of developmental stage or sex (Grzelak & Sørensen 2018, 2019; present study).
However, uncovering this variation would require a certain number of specimens, which is why the very
low number of individuals of E. balerioni from the Arctic and the single one examined in the present
study hamper proper comparison, and leave us uncertain about the specimen’s identity. Nevertheless,
our results, as well as the observations made by Grzelak for Alaskan specimens, stress the necessity of
further investigations of this interesting species, both in terms of intra- or interspecic variation and its
geographical distribution.
Echinoderes sp. aff. E. lupherorum
Figs 34–35; Tables 24–25
Material examined
NEW ZEALAND • 1 ♂; Hikurangi Slope, stn TAN1004/4; 41.6837° S, 175.6642° E; 1046 m b.s.l.;
Apr. 2010; NIWA TAN1004 Voyage; soft sediment; NIWA-159428. Mounted for LM in Fluoromount
G on glass slide • 1 ♂; Hikurangi Slope, stn TAN1004/38; 41.5937° S, 175.8532° E; 1121 m b.s.l.;
Apr. 2010; NIWA TAN1004 Voyage; soft sediment; NHMD-921636. Mounted for LM in Fluoromount
G on HS slide • 1 ♀; Hikurangi Slope, stn TAN1004/44; 41.5258° S, 175.8003° E; 728 m b.s.l.; Apr.
2010; NIWA TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted for
SEM • 2 ♂♂; Hikurangi Slope, stn TAN1004/76; 41.6833° S, 175.6500° E; 1282 m b.s.l.; Apr. 2010;
NIWA TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted for SEM •
1 ♂; Hikurangi Slope, stn TAN1004/128; 42.0485° S, 174.7000° E; 1420 m b.s.l.; Apr. 2010; NIWA
TAN1004 Voyage; soft sediment; NHMD-921635. Mounted for LM in Fluoromount G on HS slide •
1 ♂; Pahaua Canyon, stn TAN1004/22; 41.5100° S, 175.7187° E; 1188 m b.s.l.; Apr. 2010; NIWA
TAN1004 Voyage; soft sediment; NIWA-159429. Mounted for LM in Fluoromount G on HS slide •
1 ♂; Pahaua Canyon, stn TAN1004/27; 41.4983° S, 175.7043° E; 1013 m b.s.l.; Apr. 2010; NIWA
TAN1004 Voyage; soft sediment; NIWA-159430. Mounted for LM in Fluoromount G on HS slide •
3 ♀♀, 2 ♂♂; same collection data as for preceding; personal reference collection of MVS. Mounted
for SEM • 1 ♀; Honeycomb Canyon, stn TAN1004/62; 41.4760° S, 175.9477° E; 1171 m b.s.l.; Apr.
2010; NIWA TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted for
SEM • 1 ♀; Campbell Canyon, stn TAN1004/126; 42.1422° S, 174.5492° E; 1495 m b.s.l.; Apr. 2010;
NIWA TAN1004 Voyage; soft sediment; NHMD-921634. Mounted for LM in Fluoromount G on HS
slide • 1 ♀; Seamount 310, stn TAN1004/69; 41.3353° S, 176.1882° E; 670 m b.s.l.; Apr. 2010; NIWA
TAN1004 Voyage; soft sediment; NHMD-921632. Mounted for LM in Fluoromount G on glass slide •
1 ♂; Seamount 310, stn TAN1004/72; 41.3657° S, 176.1958° E; 985 m b.s.l.; Apr. 2010; NIWA TAN1004
Voyage; soft sediment; NHMD-921633. Mounted for LM in Fluoromount G on glass slide • 1 ♀; same
collection data as for preceding; personal reference collection of MVS. Mounted for SEM.
European Journal of Taxonomy 844: 1–108 (2022)
86
Distribution
Hikurangi slope, seamount, Honeycomb Canyon, Pahaua Canyon, Campbell Canyon, 670–1495 m b.s.l.
See Fig. 1 for a geographic overview of stations and Table 1 for station and specimen information.
Brief description and remarks
Echinoderes with middorsal spines on segments 4 to 8, and spines in lateroventral positions on
segments 6 to 9. Tubes present in lateroventral positions on segment 2 and 5, and in midlateral positions
on segment 10. Minute glandular cell outlets type 2 present in subdorsal positions on segment 2, and in
laterodorsal positions on segments 8 and 9. Tergal extensions of segment 11 long, constituting 6% of the
trunk length. Males with three pairs of penile spines, females with lateral terminal accessory spines and
papillae in ventrolateral positions on segment 7 and in ventromedial positions on segment 8.
General. Adults with head, neck and eleven trunk segments (Figs 34–35). Overview of measurements
and dimensions in Tables 24. Distribution of cuticular structures, i.e., sensory spots, glandular cell
outlets, spines and tubes, summarized in Table 25.
Fig. 34. Light micrographs showing overview and details of Echinoderes sp. aff. E. lupherorum. A–C. ♂
(NHMD-921636). D. ♂ (NHMD-921635). E. ♀ (NHMD-921632). A. Lateroventral overview, with inset
showing close-up of segment 11. B. Segments 1 to 6, dorsolateral view. C. Segments 4 to 9, dorsolateral
view. D. Segments 8 to 11, dorsal view. E. Segments 6 to 9, lateroventral view. Abbreviations: see
Material and methods.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
87
Segments 1 and 2 composed of complete cuticular rings. Segment 1 with glandular cell outlets type 1
in middorsal and ventrolateral positions and sensory spots in subdorsal, laterodorsal, sublateral, and
ventromedial positions (Figs 34B, 35B–C). Posterior segment margin of this and following nine
segments with pectinate fringe with well-developed tips. Cuticular hairs relatively long, evenly
distributed around segment. Segment 2 with lateroventral tubes and pair of glandular cell outlets type 2
in subdorsal positions (Figs 34B, 35B–C). Sensory spots present in middorsal, subdorsal, laterodorsal
and ventromedial positions. Glandular cell outlets type 1 observed only in middorsal position. Cuticular
hairs on this and following eight segments densely covering tergal plates and lateral halves of sternal
plates; paraventral and ventromedial areas hairless (Fig. 35A, D). Segments 3 to 11 consists of one tergal
and two sternal plates. Segments 3 with sensory spots in subdorsal positions and glandular cell outlets
type 1 in middorsal and ventromedial positions. Segment 4 with middorsal spine, and sensory spots
and glandular cell outlets type 1 located ventromedially. Segment 5 with middorsal spine, lateroventral
tubes, sensory spots present in subdorsal and laterodorsal positions and glandular cell outlet type 1
located subdorsally and ventromedially. Segment 6 and 7 with spines in middorsal and lateroventral
positions, sensory spots in paradorsal, laterodorsal and ventromedial positions, and glandular cell outlets
Fig. 35. Scanning electron micrographs showing overviews and details of Echinoderes sp. aff.
E. lupherorum. A. Ventral overview. B. Segments 1 to 5, dorsolateral view. C. Segments 1 to 3,
lateroventral view. D. Laterodorsal overview. E. Segments 7 to 8, ventral view. F. Segments 5 to 9,
laterodorsal view. G. Segments 10 to 11 of female, dorsal view. H. Segments 10 to 11 of female, ventral
view. Abbreviations: see Material and methods.
European Journal of Taxonomy 844: 1–108 (2022)
88
type 1 in subdorsal and ventromedial positions (Figs 34C–D, 35F); females with papillae located in
ventrolateral positions on segment 7 (Figs 34E, 35E). Segment 8 with middorsal spine, lateroventral
spines, small glandular cell outlets type 2 in laterodorsal positions, sensory spots in paradorsal and
laterodorsal positions and glandular cell outlets type 1 located subdorsally and ventromedially; females
with papillae in ventromedial positions (Figs 34D–E, 35E–F). Segment 9 with lateroventral spines,
small glandular cell outlets type 2 present in laterodorsal positions, four pairs of sensory spots, located in
paradorsal, subdorsal, laterodorsal and ventrolateral positions, and glandular cell outlets type 1 present
in subdorsal and ventromedial positions (Figs 34D–E, 35F); small sieve plate located in sublateral
positions (Fig. 34E); on this and previous segment glandular cell outlets type 2 and laterodorsal sensory
spots located close to midlateral line. Segment 10 with distinct, long midlateral tubes, sensory spots
in subdorsal and ventrolateral positions and two glandular cell outlets type 1 located middorsally and
Table 24. Measurements from light microscopy of Echinoderes sp. aff. E. lupherorum (in µm) from the
Hikurangi Margin with comparison of measurements for E. lupherorum from its type locality, including
number of measured specimens (n) and standard deviation (SD).
Character E. aff. E. lupherorum present study E. lupherorum Sørensen et al., 2018
n Range Mean SD Range
TL 6 250–339 307 33.1 331–415
MSW-8 3 63–67 65 2.1 75–81
MSW-8/TL 3 21–26% 24% 3.4% 19.3–23.9%
SW-10 3 48–57 54 5.1 65–71
SW-10/TL 3 16–22% 19% 3.3% 15.7–20.2%
S1 5 34–38 35 2.0 35–42
S2 5 29–36 33 2.8 33–37
S3 5 32–39 36 3.2 39–43
S4 5 38–47 42 4.4 42–47
S5 5 40–50 45 6.1 42–52
S6 5 38–57 49 9.6 50–57
S7 5 38–56 50 7.6 54–60
S8 5 43–55 49 5.1 54–62
S9 5 46–53 50 2.8 59–62
S10 5 38–48 43 3.9 44–51
S11 6 43–47 44 2.3 53–58
MD4 (ac) 6 27–33 31 2.0 31–36
MD5 (ac) 6 36–39 38 1.4 40–48
MD6 (ac) 5 38–50 46 4.5 50–67
MD7 (ac) 5 53–60 57 4.6 54–83
MD8 (ac) 5 62–81 71 8.3 90–128
LV6 (ac) 6 28–37 32 3.1 32–43
LV7 (ac) 6 38–44 42 2.1 45–53
LV8 (ac) 6 40–50 46 3.4 53–68
LV9 (ac) 6 45–56 53 4.1 54–74
TE 6 16–20 18 1.3 24–28
TE/TL 6 5–6% 6% 0.6% 6.0–8.2%
LTS 5 253–283 270 12.8 324–355
LTS/TL 5 82–108% 91% 11.6 80.5–100.9%
LTAS 1 54 n/a n/a 62–83
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
89
in ventromedial positions (Figs 34D, 35G–H). Segment 11 with a pair of long lateral terminal spines
nearly reaching trunk length (Table 24) and subdorsal sensory spots. Cuticular hairs not present. Females
with lateral terminal accessory spines, males with three pairs of penile spines. Tergal extensions long,
constituting 6% of trunk length; sternal extensions shorter and rounded (Figs 34A, 35G–H).
Echinoderes lupherorum Sørensen et al., 2018 is one of the deep-sea species originally described from
the abyssal plain off southern Oregon, in the Northeast Pacic, at a depth of 2719–3675 m (Sørensen et al.
2018). With ve additional species, i.e., E. kohni Varney et al., 2019, E. microaperturus Sørensen et al.,
2012, E. spinifurca Sørensen et al., 2005, E. sylviae Landers & Sørensen, 2018 and E. yamasakii Sørensen
et al., 2018, it constitutes the E. spinifurca species group, which is characterized by conspicuous, spinous
tergal extensions (Sørensen et al. 2005, 2012, 2018; Landers & Sørensen 2018; Varney et al. 2019).
Additionally, the group shares other morphological features, e.g., the presence of ve middorsal spines,
the presence of lateroventral/ventrolateral tubes on segment 2 and midlateral tubes on segment 10, with
all tubes being long and very well developed, and the presence of the female papillae on sternal plates of
segments 6, 7 and/or 8. Within the group, the species are most easily distinguished by the length of their
tergal extensions and the presence of glandular cell outlets type 2. Echinoderes spinifurca and E. sylviae
stand out as the species without type 2 glands on segment 2 (although the latter species has glands on
segments 8 and 9), while E. yamasakii is known as the species with the longest tergal extensions (see
Varney et al. 2019 for details). The remaining three species, including E. lupherorum, show the greatest
similarity, including an identical pattern of cuticular structures, a very similar arrangement of sensory
spots and a medium length of tergal extensions (E. microaperturus TE/TL = 5%, E. lupherorum TE/
TL = 7.2%, E. kohni TE/TL = 8.3%) (Varney et al. 2019)
The individuals examined for the present study show the closest resemblance to E. lupherorum. They
closely follow the morphology of E. lupherorum, including all taxonomically signicant characters and
all spine dimensions. Although the spines at rst sight appear longer in the East Pacic specimens of
E. lupherorum, the relative lengths of the spines in the specimens from New Zealand are almost identical
after accounting for the difference in trunk length (New Zealand specimens are slightly smaller with
TL = 307 µm than Pacic ones with TL = 371 µm) (see Sørensen et al. 2018: table 12 and Table 24 in the
present study for details). The most conspicuous difference between E. lupherorum from the type locality
and the specimens in the present study are the lengths of the tergal extensions, which are shorter in New
Zealand individuals (TE/TL = 7.2% vs 6%, respectively). We also notice subtle differences in sensory
spot distribution, which can be seen in the shift of the midlateral sensory spots present in E. lupherorum
Table 25. Summary of nature and location of sensory spots, glandular cell outlets, tubes and spines
arranged by series in Echinoderes sp. aff. E. lupherorum.
Position segment MD PD SD LD ML SL LA LV VL VM
1 gco1 ss ss ss gco1 ss
2 gco1,ss gco2,ss ss tu ss
3 gco1 ss gco1
4 ac gco1,ss
5 ac gco1,ss ss tu gco1
6 ac ss gco1 ss ac gco1,ss
7 ac ss gco1 ss ac pa(♀) gco1,ss
8 ac ss gco1 gco2,ss ac gco1,pa(♀)
9 ss gco1,ss gco2,ss si ac ss gco1
10 gco1 × 2 ss tu ss gco1
11 ss pe × 3 (♂) ltas(♀) lts
European Journal of Taxonomy 844: 1–108 (2022)
90
towards laterodorsal positions in our specimens, and the lack of sublateral and ventromedial sensory
spots on segment 1 in the East Pacic specimens that, in contrast, are present in the latter.
We were not able to compare the glandular cell outlets type 1 distribution. In E. lupherorum from the
East Pacic, the presence of this type of glands, particularly on the dorsal side, could not be conrmed
due to a thin cuticle (Sørensen et al. 2018), and a re-examination of the type material did not provide
any new information. This is unfortunate as specimens from New Zealand show a rather rare pattern of
glandular cell outlets type 1 on the dorsal side, namely the presence of type 1 outlets in the middorsal
positions on segments 1 to 3 and the subdorsal positions on segments 4 to 9. This pattern has so far been
connected with the E. dujardinii species group, and E. worthingi – a species also closely related to the
E. dujardinii group (Sørensen et al. 2020). Nevertheless, a very similar distribution pattern of dorsal
glands has also been observed for E. kohni (with the exception of segment 3 glands that are present as
a pair in subdorsal positions rather than as an unpaired one in middorsal position) (Varney et al. 2019).
This may indicate that species belonging to the E. spinifurca group also have an uncommon distribution
pattern of glandular cell outlets type 1.
To summarize, even though we can observe some minor differences between the East Pacic and New
Zealand populations of E. lupherorum, these are too minor to distinguish and describe a new species
without support from molecular data. Molecular approaches appear to be increasingly necessary in
taxonomic research of Echinoderes nowadays, since the differential characters appear to become
increasingly more subtle the more species we examine.
Echinoderes sp. aff. E. unispinosus
Figs 36–37; Tables 26–27
Material examined
NEW ZEALAND • 1 ♂; Hikurangi Slope, stn TAN1004/38; 41.5937° S, 175.8532° E; 1121 m b.s.l.;
Apr. 2010; NIWA TAN1004 Voyage; soft sediment; NHMD-921792. Mounted for LM in Fluoromount
G on HS slide • 1 ♀, 1 ♂; same collection data as for preceding; personal reference collection of MVS.
Mounted for SEM.
Distribution
Hikurangi slope, 1121 m b.s.l. See Fig. 1 for geographic location of station and Table 1 for station and
specimen information.
Brief description and remarks
Echinoderes with middorsal spine on segment 4 and spines in lateroventral positions on segments 6 and
7. Glandular cell outlets type 2 present in midlateral positions on segment 1, subdorsal, laterodorsal,
sublateral and ventrolateral positions on segment 2, lateral accessory positions on segment 5, and
midlateral positions on segment 8. Males with three pairs of penile spines and glandular cell outlets
type 2 in laterodorsal positions on segment 10; females with lateral terminal accessory spines only.
General. Adults with head, neck and eleven trunk segments. Overview of measurements and dimensions
in Tables 26. Distribution of cuticular structures, i.e., sensory spots, glandular cell outlets, spines and
tubes, summarized in Table 27.
Segments 1 and 2 composed of complete cuticular rings. Segment 1 with glandular cell outlets type 2
in midlateral positions, sensory spots in subdorsal and laterodorsal positions, and with glandular cell
outlets type 1 in middorsal and ventrolateral positions (Figs 36B–C, 37B–C). Posterior segment margin
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
91
Fig. 36. Light micrographs showing overview and details of Echinoderes sp. aff. E. unispinosus, ♂
(NHMD-921792). A. Lateroventral overview. B. Segments 1 to 5, dorsolateral view. C. Segments 1
to 5, lateroventral view. D. Segments 5 to 8, dorsal view. E. Segments 5 to 8, lateroventral view.
F. Segments 10 to 11, dorsal view, with inset showing close-up of tergal extensions of segment 11.
G. Segments 8 to 10, dorsal view. H. Segments 8 to 11, lateroventral view. Abbreviations: see Material
and methods.
European Journal of Taxonomy 844: 1–108 (2022)
92
of this and following nine segments with pectinate fringe with well-developed tips. Cuticular hairs
relatively short, lightly scattered on dorsal side, and almost completely absent on lateral and ventral
sides. Segment 2 with glandular cell outlets type 2 in subdorsal, laterodorsal, sublateral and ventrolateral
positions (Figs 36B–C, 37B–C, E). Sensory spots present in middorsal, laterodorsal and ventromedial
positions. Glandular cell outlets type 1 observed only in middorsal position. Cuticular hair covering
denser on this and following eight segments (Fig. 37A–H). Segments 3 to 11 consist of one tergal and
two sternal plates. Segment 3 with sensory spots in middorsal, subdorsal and midlateral positions, and
glandular cell outlets type 1 in ventromedial positions. Segment 4 with short (> 20 µm) middorsal spine
(Figs 36B, 37D), and glandular cell outlets type 1 located paradorsally and ventromedially; no sensory
spots observed. Segment 5 without spines, but with glandular cell outlets type 2 located in lateral
accessory positions (Figs 36E, 37D, G); sensory spots present in subdorsal, midlateral and ventromedial
positions, and glandular cell outlet type 1 located middorsally and ventromedially. Segment 6 with
Fig. 37. Scanning electron micrographs showing overviews and details of Echinoderes sp. aff.
E. unispinosus. A. Lateral overview. B. Segments 1 to 3, subdorsal view. C. Segments 1 to 3, lateral
view. D. Segments 4 to 6, subdorsal view. E. Segments 1 to 3, lateroventral view. F. Lateral overview.
G. Segments 5 to 7, lateroventral view. H. Segments 6 to 8, laterodorsal view. I. Segments 10 to 11 of
male, dorsal view. J. Segments 9 to 11, ventrolateral view. K. Segments 9 to 11 of female, laterodorsal
view. Abbreviations: see Material and methods.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
93
spines in lateroventral positions, paradorsal, midlateral and ventromedial sensory spots and glandular
cell outlets type 1 located paradorsally and ventromedially (Figs 36D–E, 37D, G–H). Segment 7 with
spines in lateroventral positions, subdorsal and midlateral sensory spots, and glandular cell outlets type 1
present in middorsal and ventromedial positions (Figs 36D–E, 37G–H). Segment 8 with glandular cell
outlets type 2 in midlateral positions, and sensory spots and glandular cell outlet type 1 in paradorsal
and ventromedial positions (Figs 36G–H, 37H). Segment 9 without spines or glandular cell outlets
type 2; sensory spots located in paradorsal, subdorsal, laterodorsal and ventrolateral positions, and
glandular cell outlets type 1 present in paradorsal and ventromedial positions (Figs 36G–H, 37J–K);
small sieve plate located in sublateral positions (Fig. 36H). Segment 10 with sensory spots in subdorsal
and ventrolateral positions, and two glandular cell outlets type 1 located middorsally and in paraventral
positions (Figs 36G–H, 37J–K). Laterodorsal glandular cell outlets type 2 present in males (Figs 36F,
37I). Segment 11 with lateral terminal spines, subdorsal sensory spots and middorsal glandular cell
outlet type 1 (Fig. 36F). Cuticular hairs not present. Females with lateral terminal accessory spines,
males with three pairs of penile spines (Figs 36F, H, 37I, K). Tergal extensions relatively long, sternal
extensions shorter and rounded (Figs 36F, 37J–K).
Echinoderes unispinosus Yamasaki et al., 2018 is a deep-sea species, known for its wide geographic
distribution. It was originally described from the deep-sea plain near Sedlo Seamount in the Northeast
Atlantic at a depth of 2875 m (Yamasaki et al. 2018b), and was subsequently reported from bathyal
Table 26. Measurements from light microscopy of Echinoderes sp. aff. E. unispinosus (in µm) from the
Hikurangi Margin with comparison of measurements for E. unispinosus Yamasaki et al., 2018 from its
type locality.
Character E. aff E. unispinosus present study E. unispinosus Yamasaki et al., 2018
NHMD-921792 (♂) Range
TL 253 265–305
MSW-7 44 44–48
MSW-7/TL 17.4% 15.0–18.2%
SW-10 35 33–38
SW-10/TL 13.8% 11.8–13.9%
S1 27 31–34
S2 24 23–25
S3 26 24–26
S4 29 25–28
S5 32 27–30
S6 33 30–34
S7 35 34–37
S8 36 35–40
S9 34 34–39
S10 36 34–38
S11 19 30–33
MD4 (ac) 19 18–22
LVS6 (ac) 22 15–17
LVS7 (ac) 19 15–19
LTS 177 169–205
LTS/TL 69.9% 60.2–72.9%
European Journal of Taxonomy 844: 1–108 (2022)
94
depths (2298–3708 m depth) in the Gulf of Mexico (Alvarez-Castillo et al. 2020) and cold seep areas
in the Mozambique Channel (> 700 m depth) (Cepeda et al. 2020). Moreover, very similar specimens,
identied as E. cf. unispinosus, were found at several localities in the deep sea (2735–3679 m depth) off
the Pacic coast of the United States (Sørensen et al. 2018).
The combination of spines and glandular cell outlets type 2 makes E. unispinosus easily distinguishable
from other congeners. Echinoderes unispinosus is characterized by having only one, relatively short
middorsal spine on segment 4, and lateroventral spines on segments 6 and 7, as well as glandular cell
outlets type 2 on segments 1, 2, 5 and 8 and by the lack of any tubes (Yamasaki et al. 2018b). The
individuals examined for the present study closely follow the morphology of this species. Morphometric
data and general appearance are almost identical in New Zealand and Atlantic specimens (see Table 26 in
the present contribution and Yamasaki et al. 2018: table 4). Subtle differences were noted in the sensory
spot distribution though, e.g., the presence of a middorsal sensory spot on segment 3 and subdorsal
ones on segment 9, observed in New Zealand specimens only, a lack of ventromedial sensory spots on
segment 7 and the displacement of glandular cell outlets type 2 from sublateral to midlateral positions
in our specimens (Figs 2G, 3H). However, some of these discrepancies (i.e., distribution of sensory
spots on segments 3 and 9) were noted also for the Pacic specimens (Sørensen et al. 2018). The only
signicant difference between E. unispinosus from the type locality and specimens from the present
study is the presence of laterodorsal glandular cell outlets type 2 in males that were not mentioned in
the original description. These structures could potentially have been missed in the original description
since Yamasaki et al. (2018b) did not have specimens available for SEM and glands in this position
on segment 10 may be extremely difcult to visualize in LM. Nevertheless, the specimens found by
Sørensen et al. (2018) off the US west coast also lacked these structures, indicating that the population
from New Zealand might in fact represent a closely related, but yet different species. However, since our
material included only three specimens and we cannot conrm whether the subtle differences observed
between Atlantic and New Zealand specimens are constant within the population, we hesitate to describe
a new species based on this material.
Table 27. Summary of nature and location of sensory spots, glandular cell outlets, tubes and spines
arranged by series in Echinoderes sp. aff. E. unispinosus.
Position segment MD PD SD LD ML SL L A LV VL VM PV
1 gco1 ss ss gco2 gco1
2 gco1,ss gco2 gco2,ss gco2 gco2 ss
3ss ss ss gco1
4ac gco1 gco1
5 gco1 ss ss gco2 gco1,ss
6 gco1,ss ss ac gco1,ss
7 gco1 ss ss ac gco1
8 gco1,ss gco2 gco1,ss
9 gco1,ss ss ss si ss gco1
10 gco1 × 2ss gco2(♂) ss gco1
11 gco1 ss pe × 3(♂) ltas(♀) lts
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
95
Recorded known species
Echinoderes juliae Sørensen et al., 2018
Figs 38–39; Table 28
Material examined
NEW ZEALAND • 1 ♀, 1 ♂; Hikurangi Slope, stn TAN1004/4; 41.6837° S, 175.6642° E; 1046 m b.s.l.;
Apr. 2010; NIWA TAN1004 Voyage; soft sediment; 1 ♀ NHMD-921717, 1 ♂ NHMD-921715. Mounted
for LM in Fluoromount G on glass slides • 1 ♀, 1 ♂; Hikurangi Slope, stn TAN1004/17; 41.6288° S,
175.8682° E; 1514 m b.s.l.; Apr. 2010; NIWA TAN1004 Voyage; soft sediment; 1 ♀ NHMD-921723,
1 ♂ NHMD-921724. Mounted for LM in Fluoromount G on glass slides • 1 ♂; Hikurangi Slope,
stn TAN1004/38; 41.5937° S, 175.8532° E; 1121 m b.s.l.; Apr. 2010; NIWA TAN1004 Voyage; soft
sediment; NHMD-921729. Mounted for LM in Fluoromount G on HS slide • 1 ♀; Hikurangi Slope,
stn TAN1004/44; 41.5258° S, 175.8003° E; 728 m b.s.l.; Apr. 2010; NIWA TAN1004 Voyage; soft sediment;
personal reference collection of MVS. Mounted for SEM • 1 ♀; Hikurangi Slope, stn TAN1004/76;
41.6833° S, 175.6500° E; 1282 m b.s.l.; Apr. 2010; NIWA TAN1004 Voyage; soft sediment; personal
reference collection of MVS. Mounted for SEM • 2 ♀♀, 2 ♂♂; Pahaua Canyon, stn TAN1004/12;
41.5508° S, 175.7250° E; 1350 m b.s.l.; Apr. 2010; NIWA TAN1004 Voyage; soft sediment; 2 ♀♀
Fig. 38. Light micrographs showing overview and details of Echinoderes juliae Sørensen et al., 2018.
A, C. ♀ (NHMD- 921719). B, D. ♂ (NHMD- 921729). E. ♂ (NHMD- 921721). A. Dorsal overview.
B. Segments 1 to 5, dorsal view. C. Segments 1 to 6, ventral view. D. Segments 9 to 11, dorsal view.
E. Segments 8 to 11, lateroventral view. Abbreviations: see Material and methods.
European Journal of Taxonomy 844: 1–108 (2022)
96
NHMD-921718-921729, 2 ♂♂ NHMD-921720-921721. Mounted for LM in Fluoromount G on glass
slide • 1 ♀; Pahaua Canyon, stn TAN1004/22; 41.5100° S, 175.7187° E; 1188 m b.s.l.; Apr. 2010; NIWA
TAN1004 Voyage; soft sediment; NHMD-921728. Mounted for LM in Fluoromount G on HS slide •
1 ♂; Pahaua Canyon, stn TAN1004/27; 41.4983° S, 175.7043° E; 1013 m b.s.l.; Apr. 2010; NIWA
TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted for SEM • 1 ♂; Pahaua
Canyon, stn TAN1004/31; 41.4962° S, 175.6828° E; 730 m b.s.l.; Apr. 2010; NIWA TAN1004 Voyage;
soft sediment; NHMD-921725. Mounted for LM in Fluoromount G on glass slide • 2 ♀♀; Honeycomb
Canyon, stn TAN1004/58; 41.4080° S, 175.8977° E; 670 m b.s.l.; Apr. 2010; NIWA TAN1004 Voyage;
soft sediment; NHMD-921726 to 921727. Mounted for LM in Fluoromount G on glass slides • 2 ♀♀;
same collection data as for preceding; personal reference collection of MVS. Mounted for SEM • 1 ♀;
Honeycomb Canyon, stn TAN1004/62; 41.4760° S, 175.9477° E; 1171 m b.s.l.; Apr. 2010; NIWA
TAN1004 Voyage; soft sediment; personal reference collection of MVS. Mounted for SEM • 2 ♀♀;
Seamount 766, stn TAN1004/132; 42.1345° S, 174.5850° E; 1453 m b.s.l.; Apr. 2010; NIWA TAN1004
Voyage; soft sediment; NHMD-921730, NHMD-921732. Mounted for LM in Fluoromount G on HS
slides.
Distribution
Hikurangi slope, seamount, Honeycomb Canyon, Pahaua Canyon, 670–1514 m b.s.l. See Fig. 1 for a
geographic overview of stations and Table 1 for station and specimen information.
Brief description and remarks
Echinoderes with middorsal spines on segments 4 to 8 and spines in lateroventral positions on segments 6
to 9. Tubes present in lateroventral position on segment 5 only. Glandular cell outlets type 2 present in
Fig. 39. Scanning electron micrographs showing overviews and details of Echinoderes juliae Sørensen
et al., 2018. A. Lateral overview. B. Segments 1 to 4, laterodorsal view. C. Segments 1 to 4, lateral view.
D. Segments 4 to 6, lateral view. E. Segments 7 to 8, lateral view. F. Segments 10 to 11 of female, dorsal
view. G. Segments 10 to 11 of male, ventral view. Abbreviations: see Material and methods.
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
97
subdorsal, laterodorsal, sublateral and ventrolateral positions on segment 2, in sublateral positions on
segments 3 and 8, and in midlateral positions on segments 4 and 5 (Figs 38–39). Tergal extensions long,
with exible tips (Figs 38D, 39G).
Echinoderes juliae is one of the deep-sea species originally found on the abyssal plain off Oregon and
along the continental rise off California, in the Northeast Pacic at depths of 2702 to 3679 m (Sørensen
et al. 2018). Recently, its presence was also recorded on the abyssal plain, east of the Atacama Trench
in the southeast Pacic at a depth of 2560 m (Grzelak et al. 2021).
Despite a signicant geographic distance, the Hikurangi Margin individuals examined for the present
study follow the morphology and morphometrics of E. juliae from the northeast Pacic type locality
closely. Differences were only detected for the glandular cell outlets type 2 on segment 4, which are
Table 28. Measurements from light microscopy of Echinoderes juliae Sørensen et al., 2018 (in µm) from
the Hikurangi Margin with comparison of measurements for E. juliae from its type locality, including
number of measured specimens (n) and standard deviation (SD).
Character
E. juliae present study E. juliae Sørensen et al., 2018
n Range Mean SD Range
TL 8 227–288 258 20.5 285–327
MSW-7 6 46–53 51 2.5 51–54
MSW-7/TL 6 18–21% 20% 1.1% 15.6–18.6%
SW-10 6 36–40 38 1.6 41–47
SW-10/TL 6 13–16% 15% 1.4% 12.8–15.4%
S1 8 23–33 27 3.0 31–34
S2 8 24–32 26 2.6 28–32
S3 8 23–33 27 3.0 31–36
S4 8 28–36 31 3.1 34–39
S5 8 29–41 33 3.9 37–42
S6 8 29–47 36 5.3 39–45
S7 8 32–48 38 5.4 44–48
S8 8 36–48 41 4.2 44–49
S9 8 35–44 39 3.1 45–52
S10 8 31–41 37 3.6 38–46
S11 5 26–30 28 1.6 31–40
MD4 (ac) 7 37–48 42 4.0 31–44
MD5 (ac) 7 47–57 52 3.4 42–56
MD6 (ac) 7 53–63 56 3.2 59–70
MD7 (ac) 7 55–72 64 8.7 67–78
MD8 (ac) 5 65–87 80 10.2 71–98
LV6 (ac) 8 31–42 38 4.5 33–43
LV7 (ac) 8 35–48 40 4.8 41–48
LV8 (ac) 8 33–47 43 6.2 44–61
LV9 (ac) 8 35–47 41 5.2 40–60
LTS 8 175–213 189 12.2 205–248
LTS/TL 8 65–94% 74% 8.7% 68.8–78.8%
LTAS 4 54–70 62 7.0 128–133
European Journal of Taxonomy 844: 1–108 (2022)
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displaced from sublateral to midlateral positions in the New Zealand specimens (Figs 38B, 39C–D), and
the length of the lateral terminal accessory spines, which are twice as long in individuals from the US
west coast (see Table 28). Nevertheless, the distribution and arrangement of the other cuticular structures
were in line with the original description, and thus we feel condent that the recorded specimens are
E. juliae.
Discussion
The present study provides the rst overview of mud dragon species inhabiting deep waters east of New
Zealand. Our material revealed 15 species of Echinoderes, of which ten are described as a new species.
Echinoderes is the most widespread and species-rich kinorhynch genus. By mid-2022, it accommodated
137 valid species representing more than 40% of the total Kinorhyncha diversity. Echinoderes overwhelms
the kinorhynch communities regardless of water depth or latitude (e.g., Dal Zotto & Todaro 2016;
Sørensen et al. 2018; Grzelak & Sørensen 2019; Landers et al. 2020; Grzelak et al. 2021). The ndings
in our study were no different from those observations: out of 204 adult individuals, 194 belonged to
the genus Echinoderes and only ten specimens to other genera. Echinoderes not only dominated in
the Hikurangi Margin material, but also shows a high regional diversity, which is comparable to other
temperate or tropical regions (Pardos et al. 2016a, 2016b; Landers et al. 2018, 2019). Also, it was not
surprising to nd a high number of new species. New Zealand territory has hardly been explored so far
in terms of its kinorhynch fauna, thus conrming the notion that the true diversity of Kinorhyncha is
severely underestimated due to limited sampling effort.
Different habitats usually harbor distinct communities and therefore contribute to overall deep-sea
diversity, as was observed for other meiofaunal taxa (Vanreusel et al. 2010). Our results seem to conrm
this observation only partially, since many recorded species of Echinoderes were not restricted in their
distribution regarding habitat type or water depth (Fig. 40). Six species, i.e., E. sp. aff. E. galadrielae/
beringiensis, E. sp. aff. E. lupherorum, E. frodoi sp. nov., E. galadrielae sp. nov., E. juliae and E. samwisei
sp. nov., were present in every surveyed habitat, and three additional species, i.e., E. aragorni sp. nov.,
E. landersi sp. nov. and E. legolasi sp. nov., occurred in two habitat types. Among all, the canyon
communities differed the most from those on seamounts and slopes due to differences in the number of
total recorded species and the number of unique species. Canyons supported the highest diversity with
twelve out of fteen recorded species being present in this environment, as well as the highest number
of exclusive species (Fig. 40). Among them, E.gandal sp. nov. was the most abundant species of all
recorded during the present study. The dominance of E.gandal in canyons, a habitat associated with
strong hydrodynamic conditions, high sediment transport and accumulation rates (Ingels et al. 2009;
Rosli et al. 2018), may indicate its opportunistic behavior and ability to adapt to and tolerate challenging
environmental conditions better than other species. Another species characteristic of this habitat was
E. blazeji sp. nov. This record seems to be of particular interest, since E. blazeji belongs to the E. coulli
species group, which so far has been considered as typical shallow-water species. There are 17 species
in this group (Randsø et al. 2019; Yamasaki et al. 2020a; ; Cepeda et al. 2022; Kennedy et al. 2022)
and a majority were observed in intertidal marine or brackish water, with the exception of four species
that inhabit subtidal, but yet shallow, marine waters (Lundbye et al. 2011; Yamasaki 2016; Kennedy
et al. 2022). Therefore, the occurrence of a representative of the E. coulli-group in deep waters probably
suggests that physico-chemical properties of the waters are major driving factors affecting the species
distribution of the E. coulli-group rather than bathymetry per se. Canyons are very heterogeneous
habitats, complex systems in terms of topography, hydrography and sedimentology (McClain & Barry
2010; Ingels et al. 2011), where also discharges of fresh-brackish water can occur (Hong et al. 2019;
Paldor et al. 2020). Submarine groundwater discharge is a global phenomenon observed at numerous
shallow and deep-water sites (Burnett et al. 2001; Bratton 2010), thus the presence of E. blazeji in Pahaua
and Honeycomb canyons may indicate that this process occurs in the investigated region. The last two
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
99
species found exclusively in canyons were E. dalzottoi sp. nov. and E. leduci sp. nov. Both species
share a number of morphological features, among which the complete lack of cuticular hairs is the most
conspicuous. Echinoderes dalzottoi and E. leduci co-occur consistently at the stations, indicating that
these species not only share morphological traits but also habitat preferences, which may conrm our
assumption of a close relationship/association between them.
The slope community was characterized by the presence of ten species (Fig. 40), of which only one was
unique for this habitat. Echinoderes sp. aff. E. unispinosus was present at only one station on the slope,
but it is hard to consider this species as an indicator of a particular habitat, not only because it occurred
in very low numbers (3 individuals), but also because E. unispinosus is known for its wide geographic
Fig. 40. Distribution of species of Echinoderes Claparède, 1863 according to (A) habitat type and (B)
depth range. Species that occur only in one habitat type or at one depth range are highlighted in the
corresponding colours.
European Journal of Taxonomy 844: 1–108 (2022)
100
distribution and presence in a range of habitats, including cold seeps and seamounts (Sørensen et al.
2018; Yamasaki et al. 2018b; Cepeda et al. 2020; Alvarez-Castillo et al. 2020). In contrast, our results
are rather in line with observations made for other deep-sea taxa, which suggest that relatively widely
distributed species tend to occur at low densities.
The lowest species pool was noted for the seamount habitat (Fig. 40), which is in contrast to observations
made by Rosli et al. (2018) for nematode diversity. These authors argue that active hydrodynamic
conditions occurring on seamounts may increase spatial variation in sedimentation processes and food
availability in the sediment, thus supporting more diverse communities in seamount habitats of the
Hikurangi region relative to canyon and slope habitats (Rosli et al. 2018). Nevertheless, despite the fact
that the seamount diversity of Echinoderes was the lowest in our study, we noted eight species (Fig. 40),
which signicantly increases the number of species of Echinoderes recorded for this type of habitat.
According to Yamasaki et al. (2019), fteen species of Echinoderes have been reported from seamounts
globally. Of the species recorded in the Hikurangi region, none have previously been observed on other
seamounts (Yamasaki et al. 2019). Additionally, none of the species from the present study were closely
related to other seamount species reported by Yamasaki et al. (2019). However, the species composition
at the investigated seamounts was not unique and seven out of eight species also occurred in other
habitats (Fig. 40). Only E. sp. aff. E. balerioni was exclusively present on a seamount. So far, E. balerioni
has been known only from the Arctic: from the Yermak Plateau north of Svalbard (Grzelak & Sørensen
2019) and the Bering Sea, Alaska (Grzelak, unpubl. obs.).
The results of the present study also provide interesting information about known species of Echinoderes.
In our material we found E. juliae, the second most abundant species in our material, and individuals
that closely followed the morphology of E. lupherorum, E. unispinosus, E. balerioni and E. galadrielae/
beringiensis. The former two species are known from the northeast Pacic, the abyssal plain and the
continental rise off California (Sørensen et al. 2018) and E. juliae was additionally noted in the southeast
Pacic (Grzelak et al. 2021). Echinoderes unispinosus is already considered as one of the species with
the widest geographic distribution, as it was described from the Northeast Atlantic and subsequently
noted in the Gulf of Mexico, off the Pacic coast of the United States and in the Mozambique Channel
in the western Indian Ocean (Sørensen et al. 2018; Yamasaki et al. 2018b; Alvarez-Castillo et al. 2020;
Cepeda et al. 2020). The least is known to date about E. balerioni, considered as an Arctic species
(Grzelak & Sørensen 2019). Therefore, if the species reported from New Zealand is conspecic with
the abovementioned species, these ndings signicantly extend its known distribution, and support
the suggestions about the capability of deep-sea kinorhynch species to disperse over great distances
(Neuhaus & Sørensen 2013; Sørensen et al. 2018; Yamasaki et al. 2018a, 2019; Grzelak et al. 2021),
enabled, e.g., through deep-sea currents or thermohaline circulation (Pawlowski et al. 2007; Ptatscheck &
Traunspurger 2020). Moreover, with the results of the present study it can be concluded that species that
were thought to be limited in their range or typical of a particular environment as, e.g., E. balerioni,
could turn out to have a much wider distribution and known distribution ranges of kinorhynch species
should be treated with caution, since the low number of studies limits available information of diversity
and distribution patterns of species.
Conclusions
Current information on New Zealand kinorhynchs is still very scarce, and with the exception of the
Hikurangi Margin, the kinorhynch fauna in other regions of New Zealand may be considered unexplored.
Future research should focus on the coastal areas, the Bay of Plenty which potentially harbours a
diverse kinorhynch fauna, and other deep-water habitats to the north, west and south. With the limited
knowledge that is currently available, it is difcult to estimate the true kinorhynch biodiversity around
New Zealand, but just by looking at the diversity of Echinoderes and the number of new species that
GRZELAK K. & SØRENSEN M.V., New Echinoderes from the Hikurangi Range
101
have been found only along the Hikurangi Margin, we can expect that New Zealand's territory is a
region with a relatively high biodiversity.
Acknowledgements
We would like to thank Norliana Rosli and Daniel Leduc (NIWA, New Zealand) for providing the
specimens and encouraging us to carry out the present study. We also thank the scientic personnel
of voyage TAN1004, and the ofcers and crew of RV Tangaroa. Sampling was carried out as part
of NIWA’s research project ‘Impact of resource use on vulnerable deep-sea communities’ funded by
the New Zealand Ministry for Business, Innovation and Employment (CO1X0906). The rst author
was supported by the Polish National Agency for Academic Exchange NAWA, the Bekker Programme
Fellowship (PPN/BEK/2019/1/00160/00001) at the Natural History Museum of Denmark and partly by
statutory funds from the Institute of Oceanology, Polish Academy of Sciences (IO PAN).
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Manuscript received: 18 May 2022
Manuscript accepted: 6 September 2022
Published on: 18 October 2022
Topic editor: Tony Robillard
Desk editor: Kristiaan Hoedemakers
Printed versions of all papers are also deposited in the libraries of the institutes that are members of
the EJT consortium: Muséum national d’histoire naturelle, Paris, France; Meise Botanic Garden,
Belgium; Royal Museum for Central Africa, Tervuren, Belgium; Royal Belgian Institute of Natural
Sciences, Brussels, Belgium; Natural History Museum of Denmark, Copenhagen, Denmark; Naturalis
Biodiversity Center, Leiden, the Netherlands; Museo Nacional de Ciencias Naturales-CSIC, Madrid,
Spain; Leibniz Institute for the Analysis of Biodiversity Change, Bonn – Hamburg, Germany; National
Museum, Prague, Czech Republic.
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