Integrative taxonomy of introduced Haplosclerida and four new species from Hawaiʻi
Author
Vicente, Jan
Hawai‘i Institute of Marine Biology, School of Ocean and Earth Science and Technology, University of Hawai‘i at Ma ̄ noa, Ka ̄ ne‘ohe, HI, 96744, USA.
Author
Rutkowski, Emily
Hawai‘i Institute of Marine Biology, School of Ocean and Earth Science and Technology, University of Hawai‘i at Ma ̄ noa, Ka ̄ ne‘ohe, HI, 96744, USA.
Author
Lavrov, Dennis V.
Department of Ecology, Evolution, and Organismal Biology, Iowa State University, 343 A Bessey, IA, 50011 - 1020, USA.
Author
Martineau, Gabrielle
Hawai‘i Institute of Marine Biology, School of Ocean and Earth Science and Technology, University of Hawai‘i at Ma ̄ noa, Ka ̄ ne‘ohe, HI, 96744, USA.
Author
Timmers, Molly
Hawai‘i Institute of Marine Biology, School of Ocean and Earth Science and Technology, University of Hawai‘i at Ma ̄ noa, Ka ̄ ne‘ohe, HI, 96744, USA. & Pristine Seas, National Geographic Society, Washington, DC 20036, USA.
Author
Toonen, Robert J.
Hawai‘i Institute of Marine Biology, School of Ocean and Earth Science and Technology, University of Hawai‘i at Ma ̄ noa, Ka ̄ ne‘ohe, HI, 96744, USA.
text
Zootaxa
2025
2025-01-08
5566
2
243
272
https://doi.org/10.11646/zootaxa.5566.2.2
journal article
10.11646/zootaxa.5566.2.2
1175-5326
14702682
85B55E49-BBC7-4321-8CC4-CBD49D29ED43
Haliclona (
Reniera
)
kahoe
sp. nov.
LSIDurn:lsid:zoobank.org:act:
10EF032A-6090-464A-BC51-12059BCD97A5
(
Fig. 4–5
,
Table 2
)
Haliclona
sp.
JV 1;
Vicente
et al.
, 2022b
, Vicente
et a
l., 2022a: Sup.
Fig. S7
Holotype
and type locality.
BPBM
C1539
-ARMS on
reef at Moku o Loʻe
(
Coconut Island
),
Pūpūkea
,
Kāne‘ohe Bay
,
Oʻahu
(
21.4335 °N
, -
157.7863 °W
);
0.3 m
, coll.
Jan Vicente
,
2018-03-16
.
Paratypes
.
BPBM
C1570
,
BPBM
C1540
,
BPBM
C1538
-
ARMS
on
reef at Moku o Loʻe
(
Coconut Island
),
Kāne‘ohe Bay
,
Oʻahu
(
21.4335 °N
, -
157.7863 °W
);
3 m
, coll.
Jan Vicente
,
2017-11-21
,
2018-03-16
,
2018-06-11
, respectively
.
BPBM
C1553
,
BPBM
C1554
,
BPBM
C1552
,
BPBM
C1551
, and
BPBM
C1537
-ARMS in
mesocosms at the Hawai‘i Institute of Marine Biology
(
HIMB
),
Moku o Loʻe
(
Coconut Island
),
Kāne‘ohe Bay
,
Oʻahu
(
21.4334 °N
, -
157.7868 °W
);
0.3 m
, coll.
Jan Vicente
,
2016-12-19
,
2017-02-13
,
2017-08-01
,
2018-01-19
,
2018-03-16
, and
2018-06-11
respectively.
Additional vouchers with metadata can be found in
Table S1
.
Diagnosis.
A soft, thin to thickly encrusting
Haliclona
(
Reniera
)
displaying a variety of irregular, and regular cushion shaped growth morphologies with apical oscula that are mainly light brown in color. The skeleton is exclusively composed of oxeas (154–197 x 1– 9 µm) arranged mainly in unispicular, isotropic, isodictyal, reticulation throughout the choanosome and ectosome.
Description (
Fig. 4
):
Thin to thickly encrusting with erect regular to irregular oscular lobes. Individuals spread laterally measuring
1–4 cm
in length, width of ≤
1 cm
and a thickness up to
0.5 cm
. Oscula measure
1–3 mm
in diameter and may rise
0.5 cm
in height. Some colonies have multiple oscular lobes stemming from one base. The base is thinly encrusting, spreading laterally. Base’s surface is smooth, even, and occasionally irregular with few microscopic pores. In some specimens multiple, long, thin anastomosing branches project outward from the base. Oscula may spread laterally across branches. Superficial canals are slightly pronounced along the base of some individuals but are rarely present. Consistency is soft, delicate, compressible, and easily torn. Color in live specimens ranges from light brown, light purple to greyish yellow. A gradient from dull yellow to purple can be observed concurrently on the same individual.
Skeleton (
Fig. 5a–h
):
Ectosome is ill defined in some specimens but when present, is composed of an isotropic unispicular, isodictyal reticulation of oxeas. Rectangular (80–120 µm in diameter) or quadrangular meshes (up to 150 µm in diameter) are composed of 5–10 oxeas which meet at the nodes with very little spongin. The lack of spongin gives the ectosome a translucent appearance. Choanosome in some specimens is less organized and consists of an isotropic to subisotropic reticulation forming meshes similar in size and shape to those found in the ectosome. Few discernable unispicular ascending primary tracts (spaced 100–150 µm apart) are visible in some individuals but are not connected by a regular frequency of secondary tracts (
Fig. 5b
). Spongin and small auxiliary oxeas is scattered sporadically throughout the choanosome.
Spicules (
Fig. 5
i-j);
Table 2
):
Oxeas are straight or slightly curved with acerate tips measuring 154–162–197 x 1–4.5– 9 μm (
Fig. 5
i-j) Oxeas with both round and acerate ends are rare.
Taxonomic remarks.
The delicate unispicular, isotropic to subisotropic reticulation of oxeas in both the choanosome and ectosome of
Haliclona
(
Reniera
)
kahoe
agrees with both
H.
(
Reniera
)
(
Schmidt, 1862
) and
H.
(
Halichoclona
)
de Weerdt, 2002
. Yet, the soft consistency, compressibility, and the lack of subectosomal or choanosomal spaces is more similar to those of
H
. (
Reniera
)
than of
H
. (
Halichoclona
). The skeleton lacks a ladder-like morphology with primary lines connected irregularly (characteristic of
H
. (
Rhizoniera
)
Griessinger 1971
or regularly (characteristic of
H
. (
Haliclona
)
de Weerdt, 2002
) by unispicular secondary lines. Skeleton reticulation is also not subhalichondroid with multispicular lines as defined for
H.
(
Gellius
)
Gray 1867
nor does it tend to form rounded meshes in the ectosome by paucispicular lines as defined for
H
. (
Soestella
)
de Weerdt, 2002
. Although,
H.
(
Reniera
)
is the most appropriate classification for this species, the ectosomal skeleton when present is not “very regular” nor does the presence of spongin conform to its definition. Spongin is not only present at the nodes of spicules that build the skeletal framework but is also abundant throughout the choanosome.
FIGURE 4.
In situ growth variations of
Haliclona
(
Reniera
)
kahoe
sp. nov.
a, holotype BPBM C1539; b, paratype BPBM C1538; c, paratype BPBM C1551; d, paratype BPBM C1537; e, paratype BPBM C1540; f, paratype BPBM C1570; g, paratype BPBM C1554; h, BPBM C1553; i, paratype BPBM C1552. Scale bars: a-f, 1 cm; g, 0.5 cm; h-i, 1 cm.
FIGURE 5.
Skeletal architecture of
Haliclona
(
Reniera
)
kahoe
sp. nov.
Perpendicular section through the ectosome and choanosome in a-b, holotype BPBM C1539; c-d, paratype BPBM C1537; e-f, paratype BPBM C1551. Tangential section of the ectosome in g-h, paratype BPBM C1537. Light microscopy images of i, first oxea from holotype, second oxea from BPBM C1538 and third from BPBM C1540. SEM images of oxeas from holotype j. Scale bars: a, c, e, f, 500 µm; b, d, g, 300 µm; h, 100 µm; i–j 50 μm.
TABLE 2.
Spicule measurements of oxeas for
Haliclona
(
Reniera
)
kahoe
holotype (h) and paratypes (p). Measurements are expressed as minimum–mean (±1 standard deviation)–maximum. N=50.
|
Voucher
|
Length (µm)
|
Width (µm)
|
| BPBM C1539 (h) |
153.9–(174.9±9.5)–197.4 |
2.5–(5.7±1.4)–8.6 |
| BPBM C1538 (p) |
125.9–(165.8±16.5)–197.3 |
1.1–(5.0±1.7)–8.2 |
| BPBM C1537 (p) |
137.0–(155.6±10.5)–185.3 |
2.5–(4.5±0.8)–5.9 |
| BPBM C1551 (p) |
111.8–(150.8±16.2)–186.0 |
1.1–(3.9±1.4)–7.1 |
There are 23
Haliclona
(unknown subgenera) spp., two
Haliclona
(
Haliclona
)
spp., three
Haliclona
(
Reniera
)
spp. and one
Haliclona
(
Rhizoniera
)
spp. sharing a thin to thick, irregularly encrusting morphology, similar color patterns and spicule composition to
H.
(
Reniera
)
kahoe
(
Table S2
). However, these species can be discarded as possible matches by 1. the presence of smaller oxeas (as in
H. carteri
Burton 1954
(40 x 8 μm),
H. hydroida
Tanita & Hoshino 1989
(120 –145 x 7–14 µm),
H. innominata
(
Kirkpatrick 1900
)
(108 x 2.5 µm),
H. isodictyalis
Bergquist 1961
(130 x 7 μm),
H. macropora
(
Thiele 1905
)
(118–125 × 4–5.2–8 μm),
H. minima
(
Lendenfeld 1887
)
(67 x 3 µm),
H. nitens
Desqueyroux-Faúndez 1990
(100–118 × 1.6–4 μm),
H. offerospicula
Hoshino 1981
(75–82–90 x 2–2.8–3 μm),
H. rectangularis
(
Ridley & Dendy 1886
)
(88 x 9 μm),
H. reversa
(
Kirk 1911
)
(100 x 5 μm),
H. tenuis
Hoshino, 1981
(83–100 x 5–8 μm),
H. translucida
Desqueyroux-Faúndez, 1990
(94–116 x 6–7 μm),
H. venustina
(
Bergquist, 1961
)
(100 x 4 μm),
H
. (
Haliclona
)
tonggumiensis
Kang
et al.
, 2013
(60–110 x 1– 5 μm),
H
. (
Reniera
)
cinerea
(
Grant 1826
)
(76–113 x 5–10 μm),
H
. (
Reniera
)
clathrata
(
Dendy 1895
)
(107 x 6 μm),
H.
(
Rhizoniera
) e
namela
de Laubenfels 1930
) 2. larger oxeas (as in
H. densaspicula
Hoshino, 1981
(187–250 x 3–15 μm),
H. maxima
Bergquist & Warne, 1980
(274–417 μm),
H.
(
Haliclona
)
ieoensis
Kim
et al.
2017
(160–230 × 2.5–12.5 μm), 3. Firm or hard consistency (as in
H. glabra
Bergquist, 1961
,
H. rapanui
(
Desqueyroux-Faúndez, 1990
)
,
H. sataensis
Hoshino, 1981
) and 4. Presence of multispicular tracts throughout the skeleton (as in
H. madagascarensis
Vacelet
et al.
1976
, and
H. tenacior
Bergquist, 1961
) Other clearly distinguishable characters setting
H.
(
Reniera
)
kahoe
apart from the remaining species are the absence of a conulose surface (characteristic of
Haliclona lentus
Hoshino, 1981
), rough oxeas with an uneven surface (diagnostic of
Haliclona scabritia
Tanita & Hoshino, 1989
and the absence of a “very regular” continuous ectosome (as observed in
Haliclona
(
Reniera
)
venusta
(
Bowerbank 1875
)
. Of the
H
. (
Halichoclona
) spp.,
H.
(
Halichoclona
)
mokuoloea
(
de Laubenfels, 1950
)
from Kāneʻohe Bay shares similar consistency and an isodictyal reticulation of oxeas within the skeletal framework. Nevertheless,
H.
(
Halichoclona
)
mokuoloea
is set apart by the yellow, reddish color, smaller oxeas (120–135 x 6 μm) and massive growth morphology.
Habitat and ecology.
All specimens were common in confined cryptic environments of Autonomous Reef Monitoring Structures (ARMS). Specimens were collected from ARMS deployed on the patch reef slope adjacent to Moku o Loʻe (Coconut Island), and ARMS inside mesocosms supplied with unfiltered flow through seawater at the
Hawai‘i
Institute of Marine Biology (HIMB) in Moku o Loʻe (Coconut Island). Time series observations show quick growth and abundance of
Haliclona
(
Reniera
)
kahoe
throughout pioneering and climax stages of succession (Sup.
Fig. S3
, Sup.
Fig. S
7
in
Vicente
et al.
, 2022a
).
Distribution.
Moku o Loʻe (Coconut Island), Kāneʻohe Bay on the island of Oʻahu,
Hawaiʻi
.
Etymology.
The given name is based on Lo‘e’s faithful brother Kahoe, who was a farmer that regularly provided crops he had grown to his brother Pahu. We use the feminine
kahoe
following the feminine gender of
Haliclona
and Article 31.2 of the International Code for Zoological Nomenclature (http://www.iczn.org/, accessed on
October 16, 2023
).