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European Journal of Taxonomy 506: 1–25 ISSN 2118-9773
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2019 · Kelly M. et al.
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Research article
urn:lsid:zoobank.org:pub:0D5F8DFB-C1AC-47F5-9129-C9241DF3DB04
1
Molecular study supports the position of the New Zealand endemic
genus Lamellomorpha in the family Vulcanellidae
(Porifera, Demospongiae, Tetractinellida),
with the description of three new species
Michelle KELLY 1,*, Paco CÁRDENAS 2,*, Nicola RUSH 3, Carina SIM-SMITH 4,
Diana MACPHERSON 5, Mike PAGE 6 & Lori J. BELL 7
1,3,4 Coasts and Oceans National Centre, National Institute of Water and Atmospheric Research,
P.O. Box 109–695, Newmarket, Auckland, New Zealand.
2 Pharmacognosy, Department of Medicinal Chemistry, BioMedical Centre, Husargatan 3,
Uppsala University, 751 23 Uppsala, Sweden.
5 Coasts and Oceans National Centre, National Institute of Water and Atmospheric Research,
Private Bag 14901, Kilbirnie, Wellington, New Zealand.
6 Coasts and Oceans National Centre, National Institute of Water and Atmospheric Research,
P.O. Box 893, Nelson, New Zealand.
7 Coral Reef Research Foundation, Box 1765, Koror, 96940 Palau.
* Corresponding authors: michelle.kelly@niwa.co.nz 1, paco.cardenas@ilk.uu.se 2
3 Email: Nicola.Rush@niwa.co.nz
4 Email: carina@clearsight.co.nz
5 Email: Diana.Macpherson@niwa.co.nz
6 Email: Mike.Page@niwa.co.nz
7 Email: crrfpalau@gmail.com
1 urn:lsid:zoobank.org:author:F9B821F7-90D0-40C5-8FB3-E96FB0502A4D
2 urn:lsid:zoobank.org:author:9063C523-49FC-427E-9E84-DBC31C5DB6D3
3 urn:lsid:zoobank.org:author:D3B1B062-6550-46C0-B562-DDDC42EEE215
4 urn:lsid:zoobank.org:author:F0205A9D-64B1-4561-8D6B-13429DC01FF3
5 urn:lsid:zoobank.org:author:106CF6B0-9E37-40BB-A85B-0C08010FFEFB
6 urn:lsid:zoobank.org:author:75F24D6D-DB93-4CFC-8978-55BE8404BEB3
7 urn:lsid:zoobank.org:author:4D42296F-6565-4E8F-AEBA-202E240B320C
Abstract. Due to the possession of huge contort strongyles, and a lack of triaenes in an otherwise
‘astrophorine’ spicule complement, the phylogenetic position of the endemic, monospecic New Zealand
sponge genus, Lamellomorpha Bergquist, 1968, has remained enigmatic. The genus was established
within Jaspidae de Laubenfels, 1968 (in the abandoned order Epipolasida Sollas, 1888), but it was
not until 2002 that the genus was transferred formally to Astrophorina Sollas, 1887, albeit incertae
sedis, by Hooper & Maldonado (2002). In this study, we recognise specimens of Lamellomorpha from
the Subantarctic New Zealand region and Chatham Rise, considered by Bergquist to be conspecic
with the type species, L. strongylata Bergquist, 1968, rst described from the Three Kings-Spirits Bay
European Journal of Taxonomy 506: 1–25 (2019)
2
region of Northland, as the new species, L. australis Kelly & Cárdenas sp. nov. These two species of
Lamellomorpha have differences in external morphology and colour, skeletal architecture and spicules,
natural products, geographical distribution, and depth ranges. Sequencing of the COI Folmer barcode/
mini-barcode and of 28S (C1–C2 domains) of these two species suggests phylogenetic afnities of
Lamellomorpha with the tetractinellid suborder Astrophorina and the family Vulcanellidae Cárdenas
et al., 2011. Two Subantarctic New Zealand species of the vulcanellid genus Poecillastra Sollas, 1888,
P. ducitriaena Kelly & Cárdenas sp. nov. and P. macquariensis Kelly & Cárdenas sp. nov., provide
further support for the close relationship of Lamellomorpha and Poecillastra.
Keywords. New species, Poecillastra, Porifera, secondary loss, Subantarctic New Zealand.
Kelly M., Cárdenas P., Rush N., Sim-Smith C., MacPherson D., Page M. & Bell L.J. 2019. Molecular study
supports the position of the New Zealand endemic genus Lamellomorpha in the family Vulcanellidae (Porifera,
Demospongiae, Tetractinellida), with the description of three new species. European Journal of Taxonomy 506:
1–25. https://doi.org/10.5852/ejt.2019.506
Introduction
Lamellomorpha strongylata Bergquist, 1968 (class Demospongiae Sollas, 1885, order Tetractinellida
Marshall, 1876, suborder Astrophorina Sollas, 1887 incertae sedis) was rst described from the Three
Kings Islands to the north of New Zealand and recently recollected by the National Institute of Water &
Atmospheric Research (NIWA) and the Coral Reef Research Foundation, Republic of Palau (CRRF)
from Middlesex Bank to the north of Three Kings Islands and Spirits Bay on the northern tip of the
North Island. In the original description, Bergquist (1968) included two specimens from the Campbell
Plateau in the Subantarctic New Zealand region (New Zealand Oceanographic Institute (NZOI) Stations
B176 and B184, 84 m and 188 m depth, respectively) and considered these to be the same species.
While the specimens have not been relocated within the NIWA Invertebrate Collection (NIC), a recent
opportunity to examine older material in NIC revealed a surprising number of the ‘southern form’ of
L. strongylata Bergquist, 1968 from Bounty Platform (NZOI Station A751, 155 m depth), Solander
Trough, Campbell Platform, and Macquarie Ridge, all in the Subantarctic New Zealand region, and
Mernoo Bank on Chatham Rise, to the east of the South Island. One specimen from the Snares Island
Platform was of great interest as it had rare calthrop-like triaenes amongst what appeared to be a spicule
complement almost identical to that of L. strongylata, suggesting a relationship with calthrop-containing
tetractinellid species, such as in families Calthropellidae Lendenfeld, 1907 or Pachastrellidae Carter,
1875.
The systematic position and phylogenetic afnity of Lamellomorpha Bergquist, 1968 has been the
subject of some debate since it was rst described. Bergquist (1968) established the genus within
Jaspidae de Laubenfels, 1968 in the abandoned order Epipolasida Sollas, 1888, based on the possession
of exclusively monaxon megascleres with asterose microscleres, i.e., lacking triaenes. Bergquist stated
that “L. strongylata would be a typical Jaspis were it not for the completely different microsclere content
of microstrongyles and streptasters”, which she likened to those in Triptolemma simplex (Sarà, 1959) in
the Pachastrellidae.
In 2002, the genus was transferred from Jaspidae to Astrophorina Sollas, 1887 incertae sedis by Hooper &
Maldonado (2002), who explored a plethora of hypotheses on the relationship of Lamellomorpha with
Ancorinidae Schmidt, 1870, Pachastrellidae, ‘Lithistid demospongiae’, Hadromerida Topsent, 1894, and
Halichondrida Gray, 1867. Bergquist’s initial thoughts were that Lamellomorpha was closely comparable
to Coppatias (Ecionemia) baculifera (Kirkpatrick, 1903), due to the joint possession of microstrongyles,
and to Jaspis serpentina Wilson, 1925 on the joint possession of microstrongyles and contort oxeas, and
KELLY M. et al., Revision of Lamellomorpha
3
thus had afnities with species of Jaspis Gray, 1867 and Coppatias Sollas, 1888 (Jaspis) in the current
family Ancorinidae. Hooper & Maldonado (2002) rejected Bergquist’s initial hypotheses on the basis
that both species possess euaster microscleres (Lamellomorpha has streptasters), a relatively strong
synapomorphy for Ancorinidae sensu stricto. Hooper & Maldonado (2002) also rejected Bergquist’s
suggestion of possible afnity with Triptolemma simplex, and thus the family Pachastrellidae, because
Lamellomorpha lacks the triaene megascleres that are used to judge the afnity with other genera.
They also considered any similarity with the form, dimensions and disposition of the megascleres and
microscleres to be articial.
Kelly, in Cryer et al. (2000), tentatively assigned specimens of L. strongylata and ‘Lamellomorpha
n. sp. 1’ (Kelly et al. 2009) to the desma-bearing, lithistid family Theonellidae Lendenfeld, 1903,
after discussions with Professor Murray Munro, University of Canterbury, on the potential origins of
the secondary metabolite chemistry of the specimens (Dumdei et al. 1997; Li et al. 1998; Hickford
2007): Munro’s group identied calyculins, calyculinamides and swinholide H (Dumdei et al. 1997),
and cyclic peptolide theonellapeptolide IIIe (Li et al. 1998), compounds which are related to those
found in Discodermia calyx Döderlein, 1884 and Theonella swinhoei Gray, 1868 suggesting a broad
relationship with the lithistid family Theonellidae. Hooper & Maldonado (2002) also put forward the
suggestion that L. strongylata might be a “lithistid demosponge” that has “lost” its desma megascleres.
Indeed, several theonellid lithistids are known with only rudimentary desmas (Kelly-Borges et al. 1994:
Discodermia dissoluta Schmidt, 1880, in the tropical Western Atlantic) or no desmas at all (Australian
species Theonella deliqua Hall et al., 2014 and Theonella maricae Hall et al., 2014). However, the
possession of extremely long, contort strongyles, not seen in other lithistids, and a lack of triaenes in
L. strongylata precludes this assignment.
Cárdenas et al. (2011) suggested that L. strongylata might be phylogenetically close to Characella
Sollas, 1886, or to Pachastrella Schmidt, 1868, based on the possession of small ectosomal monoaxial
spicules. Cárdenas et al. (2011) and Cárdenas & Rapp (2012) concluded that Pachastrellidae sensu
Maldonado (2002) was polyphyletic and reorganised the genera possessing streptasters into three families
(Pachastrellidae; Theneidae Carter, 1883; Vulcanellidae Cárdenas et al., 2011) and three incertae sedis
groups (Lamellomorpha incertae sedis, Characella incertae sedis, Neamphius de Laubenfels, 1953
incertae sedis) within the Astrophorina.
Neither Cárdenas et al. (2011) nor Cárdenas & Rapp (2012) sampled L. strongylata for molecular
sequences. Here, we sequence, for the rst time, the COI Folmer barcode/mini-barcode and 28S (C1–
C2 domains) of L. strongylata and the new species L. australis Kelly & Cárdenas sp. nov. We also
describe a second, less common Subantarctic New Zealand species, initially identied as a third species
of Lamellomorpha with rare, calthrop-like triaenes, but now considered to be a species of Poecillastra
Sollas, 1888, in the family Vulcanellidae: Poecillastra ducitriaena Kelly & Cárdenas sp. nov. The
systematic and phylogenetic implications of these species are considered with respect to the broader
phylogenetic position of Lamellomorpha, Poecillastra, and the wider Tetractinellida.
Material and methods
Collections and morphological systematics
Specimens were collected by rock and cone dredges as well as by beam and Agassiz medium trawls,
from several research vessels between 1962 and 2010. The majority of specimens were collected
onboard the National Institute of Water & Atmospheric Research (NIWA) research vessels RV Tangaroa
and RV Kaharoa; numerical voyage identier and associated stations cited as NIWA Stn TAN(voyage
number)/(station number) and NIWA Station KAH(voyage number)/(station number), respectively.
European Journal of Taxonomy 506: 1–25 (2019)
4
Several specimens from the Three Kings Islands were collected by dredge from RV Kaharoa in 1999,
on a voyage chartered by the CRRF.
Specimens were frozen immediately upon collection and then preserved in 70% ethanol or preserved
immediately into 70% ethanol (CRRF). Histological sections of the sponges were prepared by embedding
a small piece of the sponge in parafn wax and then sectioning with a microtome at 70 μm. Spicule
slides and SEM spicule preparations were made following the methods of Kelly & Sim-Smith (2012).
Clean spicules for SEM examination were spread on a plastic disc, air-dried, and coated with platinum
for 600 s. Spicules were viewed on a Philips XL30S FEG SEM. Spicule dimensions were measured
using a Meji MT5300L compound microscope tted with a Leica DFC420 microscope camera that was
connected to Leica Application Suite imaging software (Leica Microsystems (Switzerland) Ltd.). Spicule
measurements in the species descriptions are given as the mean length (range) × mean width (range)
of twenty spicule measurements per specimen unless stated otherwise and are based on measurements
from the holotype or paratypes and conrmed through examination of all other specimens. Collection
information on specimens examined in this study and previous records were gathered in a table made
available in the PANGAEA data repository (https://doi.pangaea.de/10.1594/PANGAEA.895370).
Primary and secondary type material of the new species, and additional material, are accessed within
the NIWA Invertebrate Collection (NIC) at NIWA, Greta Point, Wellington, using the prex ‘NIWA-’.
Pieces of the holotypes of L. strongylata, L. australis sp. nov., and Poecillastra ducitriaena sp. nov. are
also stored at the Zoological Museum in Uppsala, Sweden (prex ‘UPSZTY-’).
Additional abbreviations used in the text include CRRF (Coral Reef Research Foundation, Palau). The
taxonomic authority for the new taxa described in this paper is restricted to the authors Michelle Kelly
and Paco Cárdenas.
Systematics
General classication and the names of class, order, suborder, and family follow the classication
proposal by Morrow & Cárdenas (2015). The systematics of the family Vulcanellidae follows Cárdenas
et al. (2011). Terminology for the streptaster microscleres follows Sollas (1888), as used in Cárdenas &
Rapp (2012): streptasters are categorised as spirasters (small, many actines, twisted, long shaft),
metasters (intermediate morphology), plesiasters (large, few actines, short or disappearing shaft), and
amphiasters (where actines radiate from both ends of a straight shaft).
Molecular systematics
DNA was extracted using a DNeasy Blood and Tissue kit (Qiagen). PCRs were carried out in 25 μl
solutions using PuReTaq Ready-To-Go PCR beads (GE Healthcare). Due to poor preservation of the
specimens for molecular work (storage in 70% ethanol, instead of the recommended 96% ethanol), the
DNA quantities were very low, and it was very degraded (observation on a gel of 1 µl of DNA extract).
The complete Folmer fragment could not be sequenced using the standard animal barcoding primer pair
LCO1490/HCO2198 (Folmer et al. 1994) so it was sequenced in two parts: the universal minibarcode
(130 bp, without primers) was obtained using the primer pair LCO1490/Tetract-minibarR1 (Cárdenas &
Moore 2019). Then the second part of the Folmer fragment (539 bp, without primers) was amplied
using the primer pair VulcanF2/HCO2198. VulcanCOI-F2 (5’-GGGGATGACCAACTTTATAATG-3’)
is a new specic primer made to amplify COI in Vulcanellidae species. PCR conditions were (5 min/94
°C; 37 cycles (15 s/94 °C, 15 s/46 °C, 15 s/72 °C); 7 min/72 °C). The 28S fragment (C1–C2) of 308–369
bp, was obtained using the primer pair C1’/Ep3 (Chombard et al. 1998) and the same PCR program
as for COI except that we used 50°C for the annealing temperature. We pruned the comprehensive
Tetractinellida COI alignment from Kelly & Cárdenas (2016), to keep only species of Astrophorina.
We added additional sequences of Astrophorina from the Galapagos (Schuster et al. 2018) along with
KELLY M. et al., Revision of Lamellomorpha
5
the new sequences. The COI data matrix included 115 sequences (with eight Spirophorina Bergquist
& Hogg, 1969 outgroups). For 28S, we built an alignment based upon the Astrophorina 28S (C1–D2)
alignment from Cárdenas et al. (2011) and added Astrophorina 28S (C1–D2) sequences (Thacker et al.
2013; Schuster et al. 2015, 2018). The 28S data matrix included 126 sequences (with seven Spirophorina
outgroups) and was automatically aligned using MAFFT v.7 (Katoh & Standley 2013), L-INS-i option,
implemented in AliView 1.18 (Larsson 2014). Phylogenetic analyses were conducted on the CIPRES
science gateway v. 3.3 (http://www.phylo.org) (Miller et al. 2010): RAxML 8.2.10 (Stamatakis 2014)
for maximum likelihood (ML) and MrBayes v. 3.2.6 (Ronquist et al. 2012) for Bayesian analyses.
Bayesian analyses were run with BEAGLE, and consisted of two runs of four chains, each for 5 000 000
generations and sampled every 1000 tree after a 25% burn-in.
Results
Class Demospongiae Sollas, 1885
Order Tetractinellida Marshall, 1876
Suborder Astrophorina Sollas, 1887
Family Vulcanellidae Cárdenas, Xavier, Reveillaud, Schander & Rapp, 2011
Diagnosis (modied from Cárdenas et al. 2011)
Astrophorina with calthrops, short-shafted triaenes or long-shafted triaenes, in addition to large oxeas and
contort or sinuous strongyloxeas. Aster microscleres include several categories of streptasters (spirasters,
metasters, amphiasters, and plesiasters). Monaxonic spicules consist of one to three categories of spiny
microxeas or microstrongyles.
Lamellomorpha Bergquist, 1968
Lamellomorpha Bergquist, 1968: 30.
Diagnosis (modied from Hooper & Maldonado 2002)
Massive, lamellate stalked-palmate, or paddle-shaped sponges, with a relatively smooth, granular, or
eshy, slightly conulose surface. Ectosomal skeleton a skin-like membrane packed with microstrongyles.
Choanosomal skeletal architecture a core of megascleres, which are straight, curved, sinuous, or contort
oxeas, frequently modied with one or both ends rounded as in strongyloxeas. These radiate through
the stalk and fan. Straight oxeas arise as short subectosomal tracts that emerge oblique to the surface.
Roughened microstrongyles or microxeas and streptasters (amphiasters, metasters, and spirasters)
scattered throughout the body.
Type species
Lamellomorpha strongylata Bergquist, 1968 (by monotypy).
Lamellomorpha strongylata Bergquist, 1968
Figs 1–2, 6; Tables 1–2
Lamellomorpha strongylata Bergquist, 1968 (in part): 31–32, pls 4a, 11e–f, g. 10.
Lamellomorpha strongylata — Cryer et al. 2000: 21, appendix 8a–b. — Hooper & Maldonado 2002:
165–167, g. 1. — Hickford 2007: 40. — Kelly et al. 2009: 42. — NABIS 2017: 1–4.
‘Lamellomorpha n. sp. K & W’ in Cryer et al. 2000: 42 (NIWA 51169 leg.).
European Journal of Taxonomy 506: 1–25 (2019)
6
Type material
Holotype
NEW ZEALAND • Northeast of Three Kings Islands, NZOI Station B93; 33.983° S, 172.350° E; depth
54–109 m; 22 Oct. 1958; NIWA 356 (NZOI H–33) leg.; beam trawl; UPSZTY 178600 (a piece of the
holotype preserved in 70% ethanol, as well as a spicule preparation), NIWA.
Other material examined
NEW ZEALAND – Northeast of Three Kings Islands, NIWA Station Z9678 (KAH9901/27); 34.360° S,
172.712° E; depth 48 m; 26 Jan. 1999; NIWA 51169 and 51172 leg.; UPSZMC 178601 (fragment of
NIWA 51172 leg. preserved in 70% ethanol), NIWA • Northeast of Three Kings Islands, NIWA Station
Z9686 (KAH9901/43); 34.361° S, 172.686° E; depth 48 m; 27 Jan. 1999; NIWA 51267 leg.; UPSZMC
178603 (fragment preserved in 70% ethanol), NIWA • Northeast of Three Kings Islands, NIWA Station
Z9699 (KAH9901/67); 34.360° S, 172.673° E; depth 41 m; 28 Jan. 1999; NIWA 51438 leg.; NIWA •
Northeast of Three Kings Islands, NIWA Station Z9710 (KAH9901/85); 34.353° S, 172.765° E; depth
54 m; 28 Jan. 1999; NIWA 51582 leg.; dredge; NIWA • Three Kings Islands, 2.5 nm east of Great
Island, NIWA Station Z15944; 34.170° S, 172.210° E; depth 200 m; 16 Apr. 1999; CRRF, NIWA 93474
leg.; dredge; NIWA • Spirits Bay, Northland, NIWA Station KAH0606/D3; 34.36° S, 172.847° E; 15
Fig. 1. Study area showing the collection localities for: Lamellomorpha strongylata Bergquist, 1968,
around Middlesex Bank, Three Kings Islands, and Spirits Bay (triangles); L. australis Kelly & Cárdenas
sp. nov., Bounty Platform, Campbell Plateau, Solander Trough, and Macquarie Ridge (Australian EEZ)
(circles); Poecillastra ducitriaena Kelly & Cárdenas sp. nov., on the eastern edge of the Snares Platform
(square); and Poecillastra macquariensis Kelly & Cárdenas sp. nov., on Seamount 5, Macquarie Ridge
(star).
KELLY M. et al., Revision of Lamellomorpha
7
Fig. 2. Morphology, megascleres and microscleres of Lamellomorpha strongylata Bergquist, 1968.
A. Deck image showing midnight blue colouration and palmate morphology (NIWA 93474 leg., image
Lori J. Bell, CRRF). B. Loss of colouration and radiating skeleton (NIWA 73253 leg.). C. Range of
roughened microstrongyles showing centrotylote forms and more typical curved forms (penultimate:
NIWA 356 leg., rest: NIWA 93474 leg.). D. Sinuous oxeas (NIWA 93474 leg.). E. Metaster- to
amphiaster-like streptasters (left: NIWA 93474 leg., center and right: NIWA 356 leg.); F. Metaster- to
amphiaster-like streptasters (left and center: NIWA 356 leg., right: NIWA 93474 leg.).
European Journal of Taxonomy 506: 1–25 (2019)
8
May 2005; NIWA 52375 leg.; dredge; NIWA. • Middlesex Bank, Three Kings Rise, NIWA Station
TAN1105/43; 33.988° S, 171.751° E; depth 170–174 m; 28 Mar. 2011; NIWA 73243, 73253 leg.; beam
trawl; NIWA • Western Continental Slope, Northland, NZOI Station J954 (I808); 34.633° S, 172.225° E;
depth 204–192 m; 18 Jun. 1981; collected by rock dredge; specimen now lost, donated by Dame P. R.
Bergquist to Dr P. Karuso, Macquarie University, Sydney.
Description
The holotype was described by Bergquist (1968) as a “massive, thick, sometimes folded and incurved
lamellate sponge”, 130 mm high, 102 mm wide, and 18–22 mm thick, supported by a stout stalk 30 mm
in diameter. The surface was described as smooth where the dermal membrane was intact, otherwise
ragged due to projecting clumps of oxeas and strongyles. Oscules, 1–2.6 mm in diameter, were found
on the convex surface of the lamella and lie ush with the surface (Bergquist 1968). Examination of the
numerous preserved specimens in NIC reveal occasional membranous oscules, but it is difcult to tell
whether they are restricted to one side of the sponge. However, in the holotype, pores were observed on
the opposite side to the oscules, in cribriporal areas, separated by small ridges, or with no boundaries,
making a continuous pore surface; each pore is 40–80 µm in diameter. The texture was described as,
“rm but compressible, crisp, easily broken”. The colour in life was described as “bright green” and the
colour in spirit, “blue green or yellowish green” (Bergquist 1968). The most recent collection was by
the Coral Reef Research Foundation in 1999 (NIWA 93474 leg.; Fig. 2A), who described a “dark, royal
blue (not navy blue), (palmate) fan sponge with pointed tips, 20 cm high and about 1 cm thick, that tears
easily, and which has a eshy surface”.
Skeleton
The description by Bergquist (1968) of the choanosome as “lax and confused with slight traces of radiate
construction discernible”, is accurate, but in NIWA 93474 leg. the contort strongyles strongly radiate
through the plane of the fan. Bergquist described a “subectosomal region”, in which there were tracts of
megascleres, variable in thickness, that curved outward and intersect with the surface at an acute angle;
in NIWA 93474 leg. these are predominantly oxeas (Fig. 6A). The ectosome is densely packed with
microstrongyles and streptasters, which also occur throughout the sponge, but in much less abundance.
Table 1. Megascleres and dimensions (µm) of Lamellomorpha strongylata Bergquist, 1968 and
L. australis Kelly & Cárdenas sp. nov., given as length [mean (min.–max.)] × width [mean (min.–
max.)], n = 10–17 unless stated otherwise.
Specimen
Straight megascleres
Contort megascleres
L. strongylata
NZOI Holotype 33 (Bergquist 1968)
1980(1000–2808) × 26(14–33)
NIWA 00356 (NZOI Holotype 33)
1993(1520−2416) × 27(21−31)
1556(1306–1787) × 17(11–24)
NIWA 93474
1968(1373−2552) × 22(9–30)
1564(1288−1816) × 15(8−21)
L. australis sp. nov.
NZOI Station B176 (Bergquist 1968)
1482(1161–1937) × 23(17–28)
NIWA 89736 leg. (holotype)
1655(1130–1981) × 26(17–41)
not present
NIWA 93483 leg. (paratype) 1754(1454−2223) × 26(15−36) 2159(1499−3079) × 25(14−31)
NIWA 93484 leg. (paratype) 1562(1403−1988) × 21(13−27) 2657(2243−3058) × 23(17−33)
NIWA 93485 leg. (paratype) 1640(1044−2026) × 19(6−25) 3164(2672−3555) × 23(18−27)
NIWA 93486 leg. (paratype) 1725(1231−2320) × 19(13−28) 2257(1702−3575) × 20(13−32)
NIWA 93487 leg. (paratype)
1752(1226−2613) × 24(16−31)
2195(1658−3560) × 23(17−34)
NIWA 93499 leg.
1882(1572−2400) × 27(21−39)
2664(2098−3100) × 25(16−35)
KELLY M. et al., Revision of Lamellomorpha
9
Spicules
Megascleres (Table 1; Fig. 2D)
Bergquist (1968) considered the megascleres of L. strongylata (Bergquist 1968: 31, 32 (table of spicule
dimensions)) to be “strongyles, oxeas and strongyloxeas”, all of similar range in length and width,
varying only in relative frequency in the two specimens (presumably the Three Kings holotype and
the NZOI Station B176 specimen from Campbell Plateau), with oxeas being dominant in the latter.
Re-examination of the holotype megascleres, and those of more recent collections, indicate that there
are probably two forms of megascleres: 1) straight to slightly curved oxeas that are common in the
subectosomal tracts, ranging from about 1500–1750 µm long and up to 25 µm thick; and 2) massive
sinuous or contort oxeas that are usually very thick and frequently modied with one or both ends
rounded as in strongyloxeas, rarely as in true strongyles, ranging from about 1600–2375 µm long and
up to 40 µm thick. However, it is difcult to distinguish the various megascleres in some specimens, and
in some the spicules are much less contort.
Microscleres (Table 2; Fig. 2C–F)
Microstrongyles are “squat, evenly rounded spicules, slightly roughened and occasionally centrotylote”
(Bergquist 1968: 31, 32 (table of spicule dimensions)) and range from about 21–34 µm long (Table 2).
Bergquist described the streptaster microscleres of L. strongylata (Bergquist 1968: 31, 32 (table of
spicule dimensions)) as “plesiasters, small spicules with 3–12 smooth, sharply pointed rays”. A re-
examination of the holotype (Fig. 2E–F) using scanning electron microscopy has revealed that the
streptaster microscleres are metasters and occasionally amphiasters with relatively long microspined
rays, all in one size category, following the denition of Sollas (1888), and as used in Cárdenas &
Rapp (2012). We describe these spicules as metaster- to amphiaster-like streptasters with heavily spined,
relatively long rays in one size category, ranging in length from about 7–15 µm long (Table 2).
Distribution
Northeast of New Zealand.
Substrate, depth range and ecology
Attached to rocky reefs and sediment and rubble-covered rocky platforms, depth 41–200 m.
Table 2. Microscleres and dimensions (µm) of Lamellomorpha strongylata Bergquist, 1968 and
L. australis Kelly & Cárdenas sp. nov., given as length [mean (min.–max.)] × width [mean (min.–
max.)], n = 10–17 unless stated otherwise.
Specimen
Microstrongyles
Streptasters
Spirasters
L. strongylata Bergquist, 1968
NZOI Holotype 33 (Bergquist 1968)
25(23–28) × 3(2–4)
10(8–11)
not present
NIWA 00356 (NZOI Holotype 33)
23(21−28) × 3(2−4)
10(8−15)
not present
NIWA 93474
27(24−34) × 2(2−3)
9(7−12)
not present
L. australis sp. nov.
NZOI Station B176 (Bergquist 1968)
27(24–30) × 3(2–4)
10(8–11)
not given
NIWA 89736 leg. (holotype)
37(31–42) × 3(2–4)
12(10–13)
10(8–11)
NIWA 93483 leg. (paratype)
32(25−41) × 2(2−4)
10(8−12)
10(9−15)
NIWA 93484 leg. (paratype)
32(29−37) × 2(1–3)
8(8−9)
10(8–12)
NIWA 93485 leg. (paratype)
28(21−38) × 3(2−5)
10(9−12)
11(9−14)
NIWA 93486 leg. (paratype) 36(31−40) × 2(2−3) 9(7−11) 11(9−13)
NIWA 93487 leg. (paratype) 27(19−34) × 3(1−5) 9(8−12) 10(8−14)
NIWA 93499 leg. 32(26−41) × 2(1−2) 9(8−15) 10(8−12)
European Journal of Taxonomy 506: 1–25 (2019)
10
DNA barcodes
COI. NIWA 51172 leg. (minibarcode, MK033624) and NIWA 51267 leg. (MK033623): no bp differences.
28S (C1-C2). NIWA 51172 leg. (MK033143) and NIWA 51267 leg. (MK033142): no bp differences. We
failed to get sequences from the holotype.
Remarks
Lamellomorpha strongylata was originally described in considerable detail by Bergquist (1968), and
the holotype was redescribed without re-examination more recently by Hooper & Maldonado (2002).
No further material was examined. Here, for the rst time, we illustrate the sponge as it appears
upon collection, showing the beautiful royal blue colouration (Fig. 2A), and illustrate the detail and
ornamentation of the microscleres (Fig. 2C–F) using scanning electron microscopy. There is little to add
to the original description, consequently the description and skeletal details are provided in comparative
prose. Lamellomorpha strongylata is restricted to the northernmost tip of New Zealand and beyond to
the Three Kings Rise, and is easily recognised in the eld by the palmate, tree-like shape and the deep
blue to green colouration.
Lamellomorpha australis Kelly & Cárdenas sp. nov.
urn:lsid:zoobank.org:act:2A7459B3-0FBA-441B-806C-7C988075843A
Figs 1, 3, 6; Tables 1–2
Lamellomorpha strongylata — Bergquist 1968: 31–32, pl. 11f (in part). — Dumdei et al. 1997: 2636–
2639. — Li et al. 1998: 724–728. — Hickford 2007: 29–41. — Blunt & Munro 2008: 285, 1798,
1854. — Buckingham et al. 2010: 349. — Bycroft & Payne 2013: 429, 1642.
Etymology
Named for the Chatham Rise and Subantarctic New Zealand distribution of this species (‘australis’,
south, Latin).
Type material
Holotype
NEW ZEALAND • Subantarctic region of New Zealand, Bounty Platform, NZOI Station A751;
47.743° S, 179.124° E; depth 155 m; 16 Nov. 1962; NIWA 89736 leg.; Agassiz medium trawl; UPSZTY
178605 (a small piece of the holotype preserved in 70% ethanol, as well as a spicule preparation), NIWA.
Paratypes
NEW ZEALAND – same collection data as for the holotype; NIWA 92896 to 92900, 93483, 93484, and
93487 leg.; NIWA • same collection data as for preceding; NIWA 93485 leg.; UPSZTY 178606 (a small
piece of the paratype preserved in 70% ethanol), NIWA • same collection data as for preceding; NIWA
93486 leg.; UPSZTY 178607 (a small piece of the paratype preserved in 70% ethanol), NIWA.
Type locality
Subantarctic region of New Zealand, Bounty Platform; depth 155 m.
Other material examined
NEW ZEALAND • Bounty Platform, NZOI Station I711; 47.833° S, 179.250° E; depth 139 m; 22
Mar. 1979; NIWA 89717 leg.; rock dredge; NIWA • Bounty Platform, NZOI Station A714; 47.725°
S, 179.067° E; depth 165 m; 5 Nov. 1962; NIWA 86733 leg.; cone and mesh dredge; NIWA • Bounty
Platform, NZOI Station A715; 47.683° S, 179.051° E; depth 121 m; 5 Nov. 1962; NIWA 89720 leg.;
KELLY M. et al., Revision of Lamellomorpha
11
Fig. 3. Morphology, megascleres, and microscleres of Lamellomorpha australis Kelly & Cárdenas
sp. nov. A. Bilamellate form (NIWA 89736 leg., holotype). B. Club-shaped morphology showing
both sides (NIWA 93483 leg., paratype). C. Sinuous oxeas (NIWA 44388 leg.). D. Curved, roughened
microstrongyles (rounded ends) and microxeas (blunt-pointed ends) (NIWA 44388 leg.). E. Metaster-
like streptasters with heavily spined, relatively long rays (NIWA 89736 leg. and NIWA 89717 leg.).
F. Spirasters with dense, short, microspined rays (NIWA 44388 leg. and NIWA 89736 leg.).
European Journal of Taxonomy 506: 1–25 (2019)
12
cone and mesh dredge; NIWA • Bounty Platform, NZOI Station A757; 47.693° S, 179.058° E; 17 Nov.
1962; NIWA 113894 leg.; NIWA • Solander Trough, NZOI Station D39; 50.967° S, 165.750° E; depth
549 m; 7 May 1963; NIWA 44388 leg.; gear dredge, cone mesh with bag; NIWA • Campbell Platform,
NZOI Station B184; 52.615° S, 169.117° E, depth 344 m; 11 Oct. 1959; NIWA 93499 leg.; dredge;
NIWA • Macquarie Ridge, NIWA Station TAN0803/69; 52.398° S, 160.657° E; depth 451–438 m; 9 Apr.
2008; NIWA 40328 leg.; epibenthic sled; NIWA • Chatham Rise, North Mernoo Bank, NIWA Station
W427; 43.077° S, 175.272° E; depth 180–237 m; 20 Feb 1995; NIWA 44240 leg.; Agassiz Trawl; NIWA
• Chatham Rise, North Mernoo Bank, NIWA Station W446; 43.245° S, 175.444° E; depth 71–76 m; 22
Feb 1995; NIWA 44261 leg.; rock dredge; NIWA • Chatham Rise, North Mernoo Bank, NIWA Station
W447; 43.245° S, 175.458° E; depth 80–85 m; 22 Feb 1995; NIWA 44263 leg.; rock dredge; NIWA
• Chatham Rise, North Mernoo Bank, NIWA Station W448; 43.240° S, 175.458° E; depth 74 m; 22
Feb 1995; NIWA 44267 leg.; rock dredge; NIWA • Chatham Rise, North Mernoo Bank, NIWA Station
W446; 43.247° S, 175.444° E; depth 71–76 m; 22 Feb 1995; NIWA 44272 leg.; rock dredge; NIWA •
Chatham Rise, North Mernoo Bank, NIWA Station W435; 43.172° S, 175.340° E; depth 108–113 m; 20
Feb 1995; NIWA 137202 leg.; rock dredge; NIWA • Chatham Rise, North Mernoo Bank, NIWA Station
W446; 43.247° S, 175.444° E; depth 71–76 m; 22 Feb 1995; NIWA 137201 leg.; rock dredge; NIWA •
Chatham Rise, South Mernoo Bank, NIWA Station W452; 43.450° S, 175.135° E; depth 120–180 m; 22
Feb 1995; NIWA 44241 leg.; rock dredge; NIWA • Chatham Rise, South Mernoo Bank, NIWA Station
W454; 43.451° S, 175.109° E; depth 126–130 m; 22 Feb 1995; NIWA 44262 leg.; rock dredge; NIWA.
Description
Uni- to bilamellate fan sponge, table tennis bat-shaped (Fig. 3A) or club-shaped (Fig. 3B), 130–200 mm
high with a short, broad stalk, 2–3 cm thick, and a relatively thick lamella (up to 2 cm thick in places),
attenuating towards the margins, which are frequently incised. The specimen from NZOI Station B176
was described by Bergquist (1968) as being 160 mm high, 89 mm wide, and 32–58 mm thick, supported
by a stalk that was broken and thus was not measured. Oscules were not visible in the holotype or any
other specimen. Pores are inconspicuous and compressed (probably due to the xation) and were 40–
80 µm in diameter (measured on the holotype). Surface relatively smooth with low ridges radiating from
the stalk to the fan margins. Texture, relatively soft, compressible. Colour in life and preservative, tan.
Skeleton
Choanosome disorganised, with megascleres orientated more or less parallel with the axis of the fan
and stalk (Fig. 6B–D), with single or a couple of megascleres extending beyond the surface from the
subectosome. The ectosome is extremely thick and packed with microstrongyles and streptasters, which
also occur in great density throughout the sponge.
Spicules
Megascleres (Table 1; Fig. 3C)
Bergquist (1968) considered the megascleres of the “subantarctic specimen” (presumably the NZOI
Station B176 specimen from Campbell Plateau) to have, “predominantly oxeas, some of which are
curved, but most are contort”. Our examination of new material reveals that oxeas dominate the
megasclere complement; these are rarely to never modied; all have sharp attenuated tips. The majority
are straight to slightly curved and contort, but not to the degree seen in L. strongylata. The megascleres
reach their greatest length in L. australis sp. nov., up to 3575 µm long in the specimen NIWA 93486 leg.
(paratype).
Microscleres (Table 2; Fig. 3D–F)
Bergquist did not differentiate between the microscleres of the holotype of L. strongylata (from the
Three Kings) and the subantarctic Campbell Plateau specimens (L. australis sp. nov.), calling them all
“plesiasters” in the table of spicule dimensions. However, in pl. 11, gs E2 and F2–3, a clear difference is
KELLY M. et al., Revision of Lamellomorpha
13
obvious between the illustrations of the streptasters: they are metasters in pl. 11, g. E2 (L. strongylata)
and larger metasters (pl. 11, g. F3) and “abnormal microrhabds” in pl. 11, g. F2 (L. australis sp. nov.).
The “abnormal microrhabds” of Bergquist (1968: pl. 11, g. F2) are most likely spirasters (as in our
Fig. 3F), the ornamentation of which would not have been visible under light microscopy available at
the time.
Thus, L. australis sp. nov. has three forms of microsclere: a microxea (Fig. 3D) with attenuating, hastate
ends that is usually straight, but may be slightly curved, ranging in length from about 19–40 µm;
metaster-like streptasters with heavily spined, relatively long rays (Fig. 3E), ranging in length from
about 7–15 µm; spirasters with abundant, short, microspined rays that emanate from a long, spiral axis
(Fig. 3F), ranging in length from about 8–14 µm.
Distribution
Subantarctic region of New Zealand: Mernoo Bank (depth 71–237 m), north-western Chatham Rise
(Dumdei et al. 1997; Li et al. 1998); Bounty Platform (depth 121–165 m), Solander Trough (depth
549 m), Campbell Platform (depth 344 m), and Macquarie Ridge (depth 451–438 m).
Substrate, depth range and ecology
Attached by a thick stalk to sediment covered rocky substrate, depth 71–549 m.
DNA barcodes
COI. Holotype (minibarcode, MK033625), no bp differences with the COI minibarcode of L. strongylata.
Remarks
Bergquist (1968) listed two specimens from Campbell Plateau in the Subantarctic region of New Zealand,
from NZOI Station B176 (46 fathoms = 84.12 m) and NZOI Station B184 (103 fathoms = 188.4 m).
Unfortunately, neither specimen was found in the NIWA collections and both are presumed lost.
However, we did nd a specimen from NZOI Station B184 (NIWA 93499 leg.) from a depth of 344 m.
Bergquist (1968) considered the two specimens she examined to be conspecic with L. strongylata,
despite the obvious disjunct distribution, but noted that the subantarctic specimens had predominantly
oxeas, an observation we agree with.
Examination of numerous specimens uncovered in NIC allows us to convincingly separate L. australis sp.
nov. from the type species on geographic distribution, morphology, and skeletal details, despite the COI
minibarcodes not differentiating them (Fig. 7). The most obvious difference that separates L. australis
sp. nov. from L. strongylata is the markedly disjunct distribution and depth ranges: L. strongylata has
only been recorded to the north of New Zealand, 41–200 m depth, while L. australis sp. nov. is only
found on and south of Mernoo Bank on the Chatham Rise, ranging in depth from 71 m on Mernoo
Bank, to 549 m in the Solander Trough. In terms of morphology and colouration in life, L. strongylata
forms a relatively soft, dark royal blue, palmate sponge, supported by a relatively narrow stalk, while
L. australis sp. nov. forms a distinctive, tan, paddle-shaped sponge, with thin, incised margins, on a
thick, short stalk. In terms of skeletal architecture, the choanosome of L. australis sp. nov. is much
more densely packed with microscleres than L. strongylata, and the former species lacks the relatively
distinct subectosomal tracts of the latter. As noted by Bergquist (1968), the megascleres of L. australis
sp. nov. differ from those of L. strongylata in being predominantly oxeas (straight and contort) with no
modications of the tips to strongyloxeas as in L. strongylata. In addition, we note that the contort forms
are much longer on average, and have a greater size range, than in L. strongylata. Finally, microscleres
also discriminate L. australis sp. nov. from L. strongylata. Lamellomorpha strongylata has stubby, often
centrotylote roughened microstrongyle, while L. australis sp. nov. has a relatively ne, curved, slightly
European Journal of Taxonomy 506: 1–25 (2019)
14
longer roughened microxea. Lamellomorpha strongylata has metaster- to amphiaster-like streptasters
with heavily spined, relatively long rays in one size category, while L. australis sp. nov. has metaster-like
streptasters with heavily spined, relatively long rays and spirasters with abundant, short, microspined
rays that emanate from a long, spiral axis. Spirasters are absent in L. strongylata.
As part of their ongoing investigations into New Zealand marine natural products in sponges, professors
Murray Munro and John Blunt and their group in the Department of Chemistry, University of Canterbury,
Christchurch, collected what was identied by the late professor Patricia Bergquist as L. strongylata,
from Mernoo Bank on the Chatham Rise. Vouchers of these sponge specimens were donated to NIC for
their preservation and future study and have been re-identied here as L. australis sp. nov., extending the
known distribution of L. australis sp. nov. north to the Chatham Rise. Thus, it is L. australis sp. nov., and
not L. strongylata, from which biologically active secondary metabolites were isolated by the University
of Canterbury group, including calyculins (A, B, E, and F), calyculinamides (A and B), swinholide H
(Dumdei et al. 1997), and theonellapeptolides (Li et al. 1998; Hickford 2007); identical and related
compounds are found in sponges in the genus Theonella Gray, 1868 and Discodermia du Bocage, 1869
(family Theonellidae).
It has been shown that calyculins and its derivatives (e.g., calyculinamides) could be produced by the
lamentous bacteria ‘Entotheonella’ spp. in Discodermia (Wakimoto et al. 2014). ‘Entotheonella’
spp. are especially abundant in Theonella swinhoei, as well as in many other demosponges (Wilson
et al. 2014). In her PhD thesis, Hickford (2007) noticed that lamentous heterotrophic (Gram positive)
bacteria were very abundant in L. australis sp. nov. and were associated with several theonellapeptolides.
The producer of theonellapeptolides is currently unknown but the results of Hickford (2007) suggest
that L. australis sp. nov. may be a host for theonellapeptolides-producing ‘Entotheonella’-like bacteria.
Hickford (2007) also isolated unicellular bacteria from the same specimens and showed these were
associated with swinholide H. This result concurs with previous results from Bewley et al. (1996), who
identied swinholide A in unicellular bacteria isolates from Theonella swinhoei from Palau. However,
it is an apparent contradiction with Ueoka et al. (2015) who convincingly show that misakinolide A
(swinholide-like compound) from another Theonella swinhoei chemotype (chemotype WA from
Japan) is produced by ‘Entotheonella serta’. Therefore it seems that swinholide-type compounds may
be produced by bacteria other than ‘Entotheonella’ in L. australis sp. nov. and Theonella swinhoei
(chemotype Palau). Hickford (2007) further states that specimens from the “northern population of
L. strongylata” (L. strongylata) not only had very low quantities of lamentous bacteria (apparently
limited to the surface of the sponge), but also missed the biological activity, and therefore may not
produce the above mentioned compounds. Thus, to conclude, L. strongylata and L. australis sp. nov.
also clearly differ in terms of natural products and microbial communities.
Poecillastra Sollas, 1888
Poecillastra Sollas, 1888: 105.
Diagnosis (Cárdenas et al. 2011)
Vulcanellidae with spiny microxeas in a single category, triaenes are pseudocalthrops or short-shafted
triaenes.
Type species
Poecillastra compressa (Bowerbank, 1866: 55) (by original designation).
KELLY M. et al., Revision of Lamellomorpha
15
Poecillastra ducitriaena Kelly & Cárdenas sp. nov.
urn:lsid:zoobank.org:act:E3C570FA-832A-4FDB-97D3-5CDA250A3797
Figs 1, 4, 6
Etymology
Named for the possession of triaenes in addition to the apparent spiculation of Lamellomorpha, and their
guide to the phylogenetic origins of this species (‘duci-’ in the sense of a guide).
Type material
Holotype
NEW ZEALAND • Subantarctic region of New Zealand, east of Snares Island Platform, NIWA Station
TRIP3072/8; 48.5° S, 168.0° E; depth 125–213 m; 21 Oct. 2010; NIWA 61944 leg.; sh bottom
trawl; UPSZTY 178604 (a small piece of the holotype preserved in 70% ethanol, as well as a spicule
preparation), NIWA.
Type locality
Subantarctic region of New Zealand, Snares Island Platform, depth 125–213 m.
Description
Multilamellate, foliose, fan sponge (Fig. 4A), 160 mm high, 104 mm wide, with a short thick stalk
about 2 cm thick. Lamella up to 2 cm thick in places, attenuating to curled margins. Oscules were
not visible on the holotype. Cribriporal pore areas are widespread between parallel tangential tracts of
oxeas; individual pores 80–160 µm in diameter, on both sides of the lamella. Texture rm, compressible,
exible, granular and smooth to the touch. Colour in preservative tan.
Skeleton
Choanosome composed of huge swathes of long straight oxeas, radiating through the lamella, terminating
below the surface (Fig. 6E). Contort oxeas are found in the stalk. Ectosome, relatively thick, packed with
microxeas and perforated by ostia.
Spicules
Megascleres (Fig. 4B–C)
Oxeas with long ne attenuated tips, 1725(1150−2271) × 16(8−22) µm; contort oxeas in the stalk,
1801(1095−2671) × 13(5−17) µm; short-shafted triaenes, relatively uncommon, rhabd straight,
attenuating, 290(260−300) µm, clads curved or acutely bent, occasionally with bifurcating tips,
232(200−250) µm, cladome width 400−500 µm long.
Microscleres (Fig. 4D–F)
Microxeas, heavily microspined, sometimes faintly centrotylote and acutely centrally bent, sharp ended,
abundant, 38(28−46) × 3(2−4) µm; metaster- to amphiaster-like streptasters with long, microspined
rays, rare, 8(5−12) µm long; spirasters with dense, short rays, rare, 5−7 µm long.
Distribution
East of Snares Island Platform.
Substrate, depth range and ecology
Attached by a thick stalk to sediment covered rocky substrate, depth 125−213 m.
European Journal of Taxonomy 506: 1–25 (2019)
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Fig. 4. Morphology, megascleres and microscleres of Poecillastra ducitriaena Kelly & Cárdenas sp. nov.
(NIWA 61944 leg., holotype). A. Preserved holotype showing stalked morphology (both sides). B. Short-
shafted triaenes with curved clads and short rhabd. C. Sinuous and straight oxeas. D. Microspined
microxeas, sometimes faintly centrotylote, sharp ended. E. Metaster- to amphiaster-like streptasters
with long, microspined rays. F. Spirasters with dense, short rays.
KELLY M. et al., Revision of Lamellomorpha
17
DNA barcodes
COI. Holotype (MK033626); 28 bp difference with the COI of L. strongylata; 24 bp difference with the
COI of Poecillastra compressa (Bowerbank, 1866) (HM592675). 28S (C1-C2). Holotype (MK033144);
5 bp difference with the 28S (C1-C2) of L. strongylata; 3 bp difference with the 28S (C1-C2) of
Poecillastra compressa (HM592757).
Remarks
This remarkable sponge was rst identied as a third species of Lamellomorpha, as it appeared to have
an almost identical form (stalked, multilamellar fan), a megasclere complement of straight and contort
oxeas (more or less restricted to the stalk), small centrotylote microxeas, and metasters (albeit rare).
Because the short-shafted triaenes were relatively uncommon, it was initially hypothesised that this was
a species of Lamellomorpha with rudimentary triaenes that ‘showed the way’ to the true afnity of the
genus with other triaene-bearing Tetractinellida. However, molecular sequencing consistently linked
Poecillastra ducitriaena sp. nov. with other Poecillastra species (Fig. 7). Despite its consistency with
two independent markers, we note that this grouping is not supported (bootstrap of 60 for COI, of 10
with 28S). This may be due to the absence of other subantarctic Poecillastra species in our sampling
which Poecillastra ducitriaena sp. nov. may be closer to (P. Cárdenas, unpublished results).
Although not fully documented (Kelly et al. 2009), our knowledge of Poecillastra in the New Zealand
region is reasonable and includes what we consider to be Poecillastra laminaris (Sollas, 1886) (Zeng
et al. 2016) and Poecillastra schulzei (Sollas, 1886). While several undescribed species are known
from the New Zealand EEZ, no specimens are known that contain the characteristic contort oxeas of
Poecillastra ducitriaena sp. nov.
Poecillastra macquariensis Kelly & Cárdenas sp. nov.
urn:lsid:zoobank.org:act:78B12A37-93E0-4CA9-9317-81F835B112FE
Fig. 5
Etymology
Named for the type location of this species, the Macquarie Ridge.
Type material
Holotype
NEW ZEALAND • Subantarctic region of New Zealand, Seamount 5, Macquarie Ridge, NIWA Station
TAN0803/48; 51.096° S, 161.976° E; depth 462–524 m; 4 Apr. 2008; NIWA 52640 leg.; epibenthic sled;
NIWA.
Description
Solid stalk of sponge of unknown morphology, 15 mm in diameter, 20 mm high, expanding on the
broken, upper surface, sides of stalk sculpted, attachment base contains patches of substrate (Fig. 5A).
Surface hispid and scratchy to the touch; texture rm, incompressible. Colour in preservative tan.
Skeleton
Stalk composed of huge swathes of contort oxeas and triaenes between which are abundant microscleres.
Spicules
Megascleres (Fig. 5D–E)
Abundant contort to sinuous oxeas (Fig. 5D) with slightly rounded tips, 3725(2125−5750) × 53(30−70)
µm; medium-shafted triaenes (Fig. 5E), rhabd slightly curved, tapering to a sharp tip, 852(550−1225) µm,
European Journal of Taxonomy 506: 1–25 (2019)
18
Fig. 5. Morphology, megascleres and microscleres of Poecillastra macquariensis Kelly & Cárdenas
sp. nov. (NIWA 52640 leg., holotype). A. Stalk, showing smooth attachment base with specks of remaining
substrate, and broken edge of the upper part of the sponge. B. Microxeas. C. Detail of microxea showing
roughened surface. D. Contort oxea of choanosome. E. Range of triaenes with medium to short shafts
(calthrops). F. Plesiasters. G. Spiraster to metaster-like streptasters.
KELLY M. et al., Revision of Lamellomorpha
19
Fig. 6. Skeletal arrangement. A. Lamellomorpha australis Kelly & Cárdenas sp. nov. showing dense,
crustose ectosome packed with microrhabds and other microscleres, a rough megasclere bundle in
the subectosome, emerging perpendicular to the surface, and the choanosome densely packed with
microscleres (NIWA 89736 leg., holotype). B. L. australis sp. nov. showing the subectosome densely
packed with microscleres, and megascleres radiating in the plane of the sponge body (NIWA 89717 leg.).
C. L. strongylata Bergquist, 1968 showing a thin, eshy ectosome packed with microrhabds and other
microscleres, through which projects an ‘extra-axial’ tract of megascleres, traversing the subectosome
and projecting through the surface (NIWA 93474 leg.). D. L. strongylata showing the ‘axial’ arrangement
of contort strongyloxeas in the deep choanosome (NIWA 93474 leg.). E. Poecillastra ducitriaenea
Kelly & Cárdenas sp. nov. showing a densely packed, crustose ectosome through which subectosomal
megascleres project, a relatively cavernous subectosomal region packed with microxeas, and in the deep
choanosome, huge swathes of oxeas (NIWA 61944 leg., holotype). Scale bars = 500 µm.
European Journal of Taxonomy 506: 1–25 (2019)
20
clads of slightly uneven length, slightly curved downwards, 578(450−680) µm, overall cladome width,
about 900−1360 µm long, ranging to pseudocalthrops. Broken true oxeas are evident but unmeasurable.
Microscleres (Fig. 5B–C, F–G)
Microxeas (Fig. 5B–C), straight to slightly curved, roughened, abundant, 332(260−420) × 7(5−8) µm,
n = 20; plesiasters (Fig. 5F), with 3−5 microspined blunt-tipped rays, overall 67(50−100), ray length
37(25−60) µm, n = 10; metaster- to amphiaster- to spiraster-like streptasters (Fig. 5G), with long,
microspined rays, abundant, 19(15−20) µm long.
Distribution
Macquarie Ridge.
Substrate, depth range and ecology
Attached to rock substrate; depth 462–524 m.
Remarks
The specimen is the attachment base of a sponge of unknown morphology, but it clearly differs from the
holotype of Poecillastra ducitriaena sp. nov. in having a very hispid, crisp, scratchy surface, indicating
a reduction of the ectosomal crust of microscleres, and the abundance of large megascleres. It is similar
to Poecillastra ducitriaena sp. nov. in the possession of abundant contort oxeas in the stalk, but differs in
the lack of straight oxeas in the stalk and the much larger dimensions of all the spicules: the contort oxeas
are up to 2000 µm longer, on average, in Poecillastra macquariensis sp. nov., and the triaenes are about
double the size of those in Poecillastra ducitriaena sp. nov., and much more abundant, the microxeas are
about ten times larger, and the sponge contains plesiasters, absent in Poecillastra ducitriaena sp. nov.
Because our knowledge of Poecillastra in the New Zealand region is reasonable (see above), we have
made the decision to record and name this second Poecillastra species, despite our lack of information
on the body shape, and because surface texture, spicule types and dimensions are so different from those
of Poecillastra ducitriaena sp. nov.
Discussion
We were not successful in obtaining sequences from the holotype of L. strongylata (NIWA 356 leg.)
but we obtained one Folmer COI sequence from non-type specimen NIWA 51172 leg. and one COI
mini-barcode (130 bp) from non-type specimen NIWA 51267 leg., both from the same type locality, the
Three Kings Islands. The 28S fragment (C1–C2 domains) was also obtained for the latter two specimens
(369 bp each). We obtained one COI mini-barcode from the holotype of L. australis sp. nov. (NIWA
89736) but failed to amplify 28S for L. australis sp. nov. We obtained the Folmer (661 bp) and the 28S
fragments (308 bp) from the holotype of Poecillastra ducitriaena sp. nov. (NIWA 61944 leg.).
In the COI tree (Fig. 7), the two species of Lamellomorpha, which have identical sequences (at least
identical minibarcodes) were a sister group to Vulcanella Sollas, 1886; this grouping with Vulcanella
was poorly supported (bootstrap of 61). Lamellomorpha + Vulcanella were sister to a poorly supported
Poecillastra clade (bootstrap of 60), which included Poecillastra ducitriaena sp. nov. Despite these poorly
supported nodes, the Vulcanellidae was a very well supported clade suggesting that Lamellomorpha is
clearly part of this family. The 28S tree conrms the position of Lamellomorpha in the Vulcanellidae
but with no true support. Vulcanella appeared paraphyletic (poorly supported), while the grouping of
Lamellomorpha was uncertain with respect to Vulcanella or Poecillastra. This poor resolution may be
due to the short sequences we obtained (308–369 bp), which are in fairly conserved domains of 28S and
so with few bp differences to differentiate species.
KELLY M. et al., Revision of Lamellomorpha
21
Fig. 7. Astrophorina COI and 28S maximum likelihood (ML) trees reconstructed with RAxML under
the generalized time-reversible Gamma (GTRGAMMA) model. ML bootstrap supports (100 bootstrap
replicates). GenBank accession numbers are given after each taxon name. In green are the new sequences
produced during this study.
European Journal of Taxonomy 506: 1–25 (2019)
22
The grouping of Lamellomorpha with the Vulcanellidae suggested by the molecular markers is in
accordance and supported by the morphological revision of this study. Indeed, the lamellate external
morphology, the oscule/pore morphologies and distributions, as well as the skeleton organization in
Poecillastra and Lamellomorpha are similar. Some spicules are also clearly homologous, such as
the metasters to spirasters. The key discovery in this work is the conrmation that Lamellomorpha
has integrity as a genus within the family Vulcanellidae, separate from Vulcanella and Poecillastra.
This result generates new hypotheses about the evolution of spicules within the Vulcanellidae. The
microrhabds in Lamellomorpha which earlier confused taxonomists, can now be considered as reduced
Poecillastra/Vulcanella microxeas. Secondary losses of spicules have already been shown to be quite
common in the Astrophorina (Cárdenas et al. 2011). Here, the short-shafted triaenes, which can be scarce
(e.g., Poecillastra compressa) to quite rare (e.g., Poecillastra ducitriaena sp. nov.) in Poecillastra,
would have been completely lost in Lamellomorpha. As for plesiasters (the largest streptaster category),
they are absent in Lamellomorpha, as in some Poecillastra (e.g., Poecillastra ducitriaena sp. nov.) or
Vulcanella (e.g., Vulcanella horrida (Schmidt, 1870)).
Acknowledgements
Specimens were provided by the NIWA Invertebrate Collection (NIC), and for material collected from
the following research projects: Voyage KAH0606 — specimens were collected as part of a Ministry for
Primary Industries funded voyage (NIWA project ENV2005–23), titled ‘The Effects of Fishing on the
Benthic Community Structure between North Cape and Cape Reinga’; Voyage TAN0803 — specimens
were collected during the interdisciplinary New Zealand-Australian MacRidge 2 research voyage
(TAN0803), the biological component of which was part of NIWA’s research project, ‘Seamounts:
their importance to sheries and marine ecosystems,’ funded by the New Zealand Foundation for
Research, Science and Technology, and CSIRO’s Division of Marine and Atmospheric Research project,
‘Biodiversity Voyages of Discovery,’ funded by the CSIRO Wealth from Oceans Flagship; Voyage
TRIP3072 — specimens collected under the Scientic Observer Program funded by the New Zealand
Ministry for Primary Industries; Voyage TAN1105 — specimens were collected as part of the Biogenic
Habitats on the Continental Shelf project (voyages TAN1105 & TAN1108), funded by New Zealand
Ministry of Fisheries (Biogenic Habitats: ZBD200801), New Zealand Foundation for Research, Science
and Technology (CCM: CO1X0907), NIWA Capability Fund (CF111358) and Oceans Survey 20/20 RV
Tangaroa days funded by Land Information New Zealand; NIWA Stn Z15944, collector CRRF: Coral
Reef Research Foundation under contract to the US National Cancer Institute (N02-CM-77249).
We thank emeritus professors Murray Munro and John Blunt, Department of Chemistry, University of
Canterbury, Christchurch, for the donation of their sponge vouchers to NIC, for their continued study.
We also thank Dr Bruce Marshall, Museum of New Zealand Te Papa Tongarewa, Wellington, for the
loan of specimens from their collections. We are grateful to John Rosser MA LTCL for his assistance
with suggestions for new species names and advising on the correct form of the new taxon names.
Satya Amirapu, Auckland University, carried out our histological work. PC received support from the
European Union's Horizon 2020 research and innovation program through the SponGES project (grant
agreement No. 679849). This document reects only the authors’ view and the Executive Agency for
Small and Medium-sized Enterprises (EASME) is not responsible for any use that may be made of
the information it contains. This research was funded by NIWA under Coasts and Oceans Research
Programme Marine Biological Resources: Discovery and denition of the marine biota of New Zealand
(2016/2017 to 2018/2019 SCI).
KELLY M. et al., Revision of Lamellomorpha
23
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Manuscript received: 23 October 2018
Manuscript accepted: 18 February 2019
Published on: 14 March 2019
Topic editor: Rudy CAM Jocque
Desk editor: Alejandro Quintanar
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; Real Jardín Botánico de Madrid CSIC, Madrid, Spain; Zoological Research Museum Alexander
Koenig, Bonn, Germany.
