Zoosyst. Evol. 99 (2) 2023, 375-389 | DOI 10.3897/zse.99.103682

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Illustrated catalogue of sphaeromatoid isopods (Crustacea,
Malacostraca) in the Canadian Museum of Nature (CMN)

Valiallah Khalaji-Pirbalouty’, Jean-Marc Gagnon?

1 Department of Biology, Faculty of Basic science, Shahrekord University, Shahrekord, Iran

2 Beaty Centre for Species Discovery, Canadian Museum of Nature, Ottawa, Canada

https://zoobank. org/55 1CBC3C-0F 54-4634-9210-27E17AQA4E49

Corresponding author: Valiallah Khalaji-Pirbalouty (khalajiv@yahoo.com)

Academic editor: Luiz F. Andrade # Received 17 March 2023 # Accepted 8 June 2023 @ Published 5 July 2023

Abstract

Zoological collections are major treasures representing the history of animal biodiversity on Earth and are an important resource
for biodiversity and conservation research. The Canadian Museum of Nature (CMN) has one of the oldest crustacean collections in

North America.

Here, we provide an illustrated catalogue of the superfamily Sphaeromatoidea Latreille, 1825, deposited in the Canadian Museum
of Nature’s Crustacea Collection (CMNC). In this paper, we report 18 species, belonging to 3 families and 14 genera. The majority
of species belong to the family Sphaeromatidae with 16 species, followed by the Ancinidae and the Tecticipitidae each with one
species. We present a bibliography of the original description, current taxonomic status, the type locality, geographic distribution,

and an updated illustration for all species.

Key Words

Canada, CMNC, Isopoda, Sphaeromatoidea, taxonomy

Introduction

The superfamily Sphaeromatoidea Latreille, 1825, com-
prising the Ancinidae, Sphaeromatidae and Tecticiptidae,
with almost 111 genera and 649 known species, is one
of the most frequently encountered and diverse isopod
taxa (Boyko et al. 2008 onwards). Within the Sphaero-
matoidea, the family Sphaeromatidae Latreille, 1825, is
the largest family of free-living marine Isopoda, with 622
species belonging to 108 genera occurring in the shal-
low-water marine environments, and many as yet unde-
scribed species and genera. The two small sphaeromatoid
families are the Ancinidae Dana, 1852, with 15 described
species distributed across two genera, and the Tecticipit-
idae Iverson, 1982, with one genus and 12 valid species
(Boyko et al. 2008 onwards).

The history of isopod taxonomy in North America
dates back to Say (1818), Dana (1853), Oscar Harger
(1880) and Verrill et al. (1873), as well as the massive

contributions of, among others, Harriet Richardson,
culminating in her 1905 monograph. While many of
the species described by these early authors are val-
id, the brief species descriptions provided at the time
have resulted in difficulties in recognizing species and
subsequent misidentifications by later authors. Many
species in most families remain to be fully described.
Furthermore, there is a need to review many of the
genera and the placement of species within these. The
Canadian Museum of Nature (CMN) holds large collec-
tions across all animal groups. Based on the Canadian
Museum of Nature database, the museum’s Invertebrate
Collection contains more than seven million specimens
in more than 1.9 million lots. Of these, Amphipoda, Iso-
poda and Mysidacea are the major groups found in the
Crustacean Collection (CMNC). Most of the specimens
were collected by Edward L. Bousfield, who joined the
CMN in 1950. Bousfield was a world authority on the
systematics of Amphipoda; he described more than 300

Copyright Khalaji-Pirbalouty, V. & Gagnon, J-M. This is an open access article distributed under the terms of the CCO Public Domain Dedication.

376

new species (Conlan et al. 2016). Besides his inten-
sive focus on the Amphipoda, his collections cover all
sorts of invertebrates (mostly marine), as well as isopod
crustaceans. With more than 3200 isopod records in the
CMN database, including about 80 genera and 190 spe-
cies (about 85% of which are from marine and brackish
water habitats), in addition to the many samples yet to
be catalogued, this collection is amongst the most 1m-
portant natural history archives for this group in North
America. The present catalogue is a comprehensive,
up-to-date account of the 18 species of sphaeromatoid
isopods represented in the Canadian Museum of Na-
ture, with illustrations of the species. This catalogue is
arranged alphabetically by genus and then by species
names within families, followed by the original combi-
nation, author(s) name and year of publication. For each
species, a current nomenclature (valid names or synon-
ymy) is given.

Material and method

Sphaeromatoid isopods for this study are from the Crus-
tacean Collection of the Canadian Museum of Nature
(CMNC; located at its Natural Heritage Campus, in
Gatineau, Quebec, Canada).

Specimens were examined using Zeiss Stereomicro-
scope (Stemi 508). Color images of the specimens were
taken using a Zeiss AxioCam ERcSs digital camera
mounted on a Zeiss (Stemi 508) stereomicroscope. Pho-
tographs were merged and edited using Adobe Photoshop
CC v.20.0.6.

Results

Systematics
Alphabetical list of taxa

Family Ancinidae Dana, 1852
Genus Bathycopea Tattersall, 1905

Bathycopea daltonae (Menzies & Barnard, 1959)
Fig. 1A, B

Ancinus daltonae Menzies & Barnard, 1959: 31, fig. 25; Schultz 1973:
270-272, fig. 1D, G.

Bathycopea daltonae.— Loyola e Silva, 1971: 217-222, figs 5—7; Kus-
sakin 1979: 369, fig. 229; Shimomura 2008: 26.

Type locality. The shelf off San Miguel Island, California.

Material examined. CANADA. | ovigerous &
(5.2 mm), 2 ¢¢ (up to 5.1 mm); British Columbia, Van-
couver Island, Barclay Land District, Cape Beale; 2 Aug.
1975; E.L. Bousfield leg.; CMNC 1985-0633. 4 6S (up
to 5 mm); British Columbia, Barclay Land District, Trev-
or Ch.; 29 Jul. 1975; E.L. Bousfield leg.; CMNC 1985-

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Khalaji-Pirbalouty, V. & Gagnon, JM.: Sphaeromatoid isopods of CMN

0630. 1 ovigerous 2 (4 mm); British Columbia, Barclay
Land District, Trevor Ch.; 30 May 1977; E.L. Bousfield
leg; CMNC 1985-0634. USA. 2 ovigerous 29 (up
to 4.9 mm), 1 4 (4.25 mm); Washington, Clallam Co.,
Makah Bay; 31 July 1966; E.L. Bousfield leg.; CMNC
1991-2557.

Distribution. San Miguel Islands, southern California
to Vancouver Island.

Family Sphaeromatidae Latreille, 1825
Genus Amphoroidea H. Milne Edwards, 1840

Amphoroidea typa H. Milne Edwards, 1840
Fig. 1C

Amphoroidea typa Milne Edwards, 1840: 22—23; Dana 1853: 783; Han-
sen 1905: 108-126; Menzies 1962a: 140-141, fig. 47D; Hurley and
Jansen 1977: 27 (listed as type species).

Type locality. Chile.

Material examined. CHILE. 30 9 (up to 19.5 mm),
1 3 (16.5 mm); Magallanes-Antarctica Region, Isla Len-
nox; 5 Feb.1970; E.L. Bousfield & J.W. Markham leg.;
CMNC 1992-0567.

Distribution. Known only from Chile.

Genus Cassidinidea Hansen, 1905

Cassidinidea ovalis (Say, 1818)
Fig. 1D

Naesa ovalis Say, 1818: 484-485.

Cassidena lunifrons Richardson, 1900: 222.

Naesa ovalis Richardson, 1900: 224.

Cassidina lunifrons.— Richardson 1901: 533, fig. 14.

Cassidisca lunifrons.— Richardson 1905: 273, figs 283-284.

Cassidisca ovali.— Richardson, 1905: 274, figs 283, 205.

Cassidinidea ovalis— Hansen, 1905: 130; Menzies and Frankenberg
1966: 44 fig. 20; Schultz 1969: 115, fig. 158; Kussakin 1979: 336,
figs 199-200; Heard 1982: 32, fig. 35; Kensley and Schotte 1989:
208, fig. 92; Bruce 1994: 1151, fig. 45; Camp et al. 1998: 136; Kens-
ley and Schotte 1999: 701-702; Khalaji-Pirbalouty and Bruce 2021:
494-502, figs 2—5.

Type locality. St John’s River in Florida.

Material examined. USA. 4 ¢¢ (up to 3 mm), 8 99
(up to 3.2 mm); South Carolina, Georgetown County; 26
Apr. 1965; E.L. Bousfield leg.; CMNC 1992-0582. 1 3
(2.8 mm), 6 99 (up to 3.2 mm); South Carolina, Charles-
ton County, 25 Apr. 1965; E.L. Bousfield leg.; CMNC
1991-2575. 2 99 (up to 3 mm); North Carolina, Dare
County; 11 Apr. 1975; E.L. Bousfield leg.; CMNC 1985-
0643. 1 2 (3.1 mm); North Carolina, Tyrrell County, 11
Apr. 1975; E.L. Bousfield leg.; CMNC 1985-0644.

Distribution. Eastern coast of North America
from New Jersey to Florida (Khalaji-Pirbalouty and
Bruce 2021).

Zoosyst. Evol. 99 (2) 2023, 375-389 377

' 1000 um

Figure 1. A. Bathycopea daltonae (Menzies & Barnard, 1959), male, (CMNC 1985-0633); B. Female, pleotelson; C. Amphoroidea
typa Milne Edwards, 1840, female, (CMNC 1992-0567); D. Cassidinidea ovalis (Say, 1818) male, (CMNC 1992-0582); E. Cas-
sidinopsis emarginata (Guérin-Méneville, 1843), male, (CMNC 1985-0636); F. Pleotelson dorsal view; G. Pleotelson ventral view.

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378
Genus Cassidinopsis Hansen, 1905

Cassidinopsis emarginata (Guérin-Meéneville, 1843)
Fig. 1,G

Cassidina emarginata Guérin-Méneville, 1843: 31; Cunningham 1871:
499, pl. 59, fig. 4; Miers 1879: 204; Studer 1883: 19; Pfeffer 1886:
63-69, pl. 2, figs 9-10, pl. 4, figs 23-30, pl. 6, figs 1-10; Dollfus
1891: 67, pl. 8, fig. 10; Stebbing 1900: 562; Ortmann 1911: 650.

Cassidinopsis emarginata — Hansen, 1905: 82, 87, 128; Hodgson 1910:
4: Tattersall, 1921: 223; Monod, 1931: 25-26; Stephensen 1947:
28; Vanhoffen 1914: 514; Menzies 1962a: 144, fig. 49; Carvacho,
1977: 177; Kussakin and Vasinal980: 356-360; Brandt 1998: 150,
figs 9-11.

Cassidinopsis emarginatus — Giambiagi, 1925: 233; Stebbing 1914: 351.

Type locality. Falkland Island. (Falkland Islands and the
West coast of Patagonia).

Material examined. CHILE. 1 4 (36 mm); Magallanes
y de la Antartica Chilena Region, Isla Navarino, Punta
Wulaia; 3 Feb. 1970; E.L. Bousfield leg.; CMNC 1985-
0636. 18 3d and 9 (up to 28 mm); Magallanes-Ant-
arctica Region, Canal Beagle; 19 Feb. 1970; E.L. Bous-
field leg.,; CMNC 1985-0637. 12 (18 mm), 2 juveniles;
Magallanes-Antarctica Region, Isla Lennox; 6 Feb. 1970;
E.L. Bousfield leg.; CMNC 1985-0635.

Distribution. South Georgia Island, Falkland Islands
to Southern coasts of Chile; Puerto Deseado, Argentina
(Brandt 1998; Gomez Simes 1979).

Genus Dynamene Leach, 1814

Dynamene dilatata Richardson, 1899 (uncertain/
incerta sedis)
Fig. 2A, B

Dynamene dilatata Richardson, 1899: 882-883, fig. 8; Richardson
1905: 304, fig. 327; Schultz 1969: 122, fig. 171; Hatch 1947: 214,
pl. 7, figs 85-86.

Dynamenella dilatata— Kussakin, 1971: 450, fig. 298; Brusca et al.
2007: 521, 537, pl. 242.

Note. This species is retained in the genus Dynamene (in
original combination) and listed as “taxon inquirendum”
in WoRMS (Boyko et al. 2008 onwards). Thus, the cor-
rect generic status of this species 1s still in question. Since
Richardson (1899; 1905) and subsequent authors pro-
vided only a brief description with a figure of the whole
body, a morphological revision is required to correctly
place this species.

Type locality. Monterey Bay, California.

Material examined. CANADA. 3 9 (up to 6.5 mm);
Oregon, Lincoln Co.; 12 Aug. 1966; E.L. Bousfield leg.;
CMNC 1991-2567. 1 ovigerous 2 (8.5 mm); British Co-
lumbia, Long Beach Unit, Long Beach; 5 Aug. 1955; E.L.
Bousfield leg.; CMNC 1985-0648. 1 4 (5.5 mm); Brit-
ish Columbia, Long Beach Unit, Long Beach; 5 August
1955; E.L. Bousfield leg.; CMNC 1985-0660.

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Khalaji-Pirbalouty, V. & Gagnon, JM.: Sphaeromatoid isopods of CMN
Genus Dynamenella Hansen, 1905

Dynamenella sheareri (Hatch, 1947)
Fig. 2C

Dynamene sheareri Hatch, 1947: 164, 262, fig. 173; George and Strom-
berg 1968: 246-248, pl. 2, fig. 9.

Dynamenella sheareri.— Schultz ,1969: 123, fig. 174; Harrison and Hol-
dich 1982: 90.

Note. The true generic status of this species is still
undetermined.

Type locality. Coos Bay, Oregon.

Material examined. CANaDa. 6 9 (up to 3 mm),
1 ¢ (3.8 mm); British Columbia, Renfrew Land Distr.,
Port Renfrew; 1 Aug. 1970; CMNC 1990-0069. 5 °° (up
to 3.5 mm), 2 Jo (4 mm), British Columbia, Vancou-
ver Island, Barclay Land Distr., Bordelais Islets; 9 Aug.
1975; CMNC 1990-0070.

Remarks. Dynamenella sheareri cannot be a Dy-
namenella because of the following characters: having
a single pleonal suture running to posterior margin (in-
stead of two); penial process more fused, not basally
swollen; appendix masculina is evenly slender and long,
not “flask shaped.”

Genus Dynoides Barnard, 1914

Dynoides canadensis Khalaji-Pirbalouty &
Gagnon, 2021
Fig. 2D

Dynoides canadensis Khalaji-Pirbalouty & Gagnon, 2021: 12—20, figs 2—S.

Type locality. Canada, British Columbia.

Material examined. CANADA. Holotype. 3 (4.2 mm);
British Columbia, Barclay Land District, Cape Beale;
19 July 1970; E.L. Bousfield leg.; CMNC 1985-0667.1.
Paratypes. 4 33 (up to 4.2 mm), 1892 (up to 4.3 mm),
1 ovigerous? (4.5 mm), same data as holotype; CMNC
1985-0667.2. 2 do (up to 5 mm), 2 ovigerous 99
(3.5 mm), 2 99 (3.5 mm); British Columbia, Sooke
Land District, Whiffin Spit; 17 August 1955; E.L. Bous-
field leg.; CMNC 1990-0064. 3 2° (up to 5 mm); British
Columbia, Rupert Land Distr., Vancouver Island, Cape
Scott; 18 July 1959; E.L. Bousfield leg.; CMNC 1990-
0066. 20 3&9; British Columbia, Renfrew Land Distr.,
Vancouver Island, Port Renfrew; 1 August 1970; E.L.
Bousfield leg.; CMNC 1990-0068. 12 (3 mm); British
Columbia, Queen Charlotte Islands Land Distr., Gra-
ham Island, 27 July 1957; E.L. Bousfield leg.; CMNC
1990-0067. 5 3S (up to 4.9), 6 YO (up to 4.2); Brit-
ish Columbia, Barclay Land District, Trevor Channel,
Tzartus Island; 21 July 1970; E.L. Bousfield leg.; CMNC
1985-0664. 4 43 (up to 4.5 mm), 5 43 (up to 4.2 mm),
British Columbia, Metchosin Land Distr., Sooke Basin,
Becher Bay; 31 July 1970; E.L. Bousfield leg.; CMNC
1990-0057. 43.4 (up to 4.5 mm), 2 92 (3 mm), British

Zoosyst. Evol. 99 (2) 2023, 375-389

Columbia, Nootka Land Distr., Nootka Island; 20 Au-
gust 1959; E.L. Bousfield leg.; CMNC 1990-0059. 1 3
(4.2 mm), 1 6 (3.5 mm), British Columbia, Rupert Land
Distr., Hope Island; 22 August 1959; E.L. Bousfield leg. ;
CMNC 1990-0060. 1 3 (5.1 mm), 5 4 (up to 4 mm),
1 juvenile, British Columbia, Range 2 Coast Land Disttr.,
Goose Island; 6 August 1964; E.L. Bousfield leg.; CMNC
1990-0061. 6 9° (up to 3 mm), British Columbia, Range
2 Coast Land Distr., Hunter Island; 8 August 1964; E.L.
Bousfield leg.; CMNC 1990-0063. 8 3'¢ (up to 5 mm),
20 9° (up to 4.1 mm), 10 juveniles, British Columbia,
Range 3 Coast Land Distr., Princess Royal Island; 20 July
1964; E.L. Bousfield leg.; CMNC 1990-0065.

Distribution. Western coasts of British Columbia
from Victoria area to Graham Island.

Genus Exosphaeroma Stebbing, 1900

Exosphaeroma gigas (Leach, 1818)
Fig. 2E, F

Sphaeroma gigas Leach, 1818: 346-347; Desmarest 1825: 301;
Milne-Edwards 1840: 205; White 1847: 102; Dana 1853: 775; Miers
1879: 202-203; Haswell 1882: 287; Studer 1884: 17-18; Dollfus
1891: 62, pl. 8a, fig. 6.

Sphaeroma jurinii.— Krauss, 1843: 65.

Sphaeroma propinqua— Nicolet, 1849: 277-278.

Sphaeroma chilensis — Dana, 1853: 195-196.

Sphaeroma obtusa— Hutton, 1879 (in Chilton 1906).

Exosphaeroma gigas.— Stebbing, 1900: 553-558, pl. 39; Chilton 1906:
271-272; Ortmann 1911: 646-647; Vanhoffenl1914: 510-511; Tat-
tersall 1921: 216; Giambiagi 1925: 235; Stephensen 1927: 362;
Nierstrasz 1931: 194; Barnard 1940: 413, fig. 13; Hurley 1961: 269;
Hale 1929: 275, fig. 273; Menzies 1962a: 132-134, fig. 43; Kus-
sakin 1967: 235; Hurley and Jansen 1977: 58, fig. 52; Carvacho,
1977: 177-178; Kussakin and Vasinal980: 355-359; Brandt and
Wagele 1989: 209-214. figs 5-9; Bruce 2003: 368.

Type locality. Unknown; The Natural History Museum,
syntypes: 1941: 6:27: 5 (presented by Leach) and 1979:
420: 1, Sir Joseph Banks collection (Ellis 1981).

Material examined. CHILE. 22 3 (up to 24 mm),
50 2° (up to 18 mm); Magallanes and Chilean Antarc-
tica Region, Picton Island; 7 Feb. 1970; E.L. Bousfield
leg., CMNC 1990-0090. 46 33’ and 29; Magallanes
and Chilean Antarctica Region, Navarino Island; 29 Jan.
1970; E.L. Bousfield leg.; CMNC 1990-0092. 300 ¢¢
and 9 9; Magallanes and Chilean Antarctica Region, Ca-
nal Beagle; 19 Feb. 1970; E.L. Bousfield leg.; CMNC
1990-0094. FALKLAND. 192 (16 mm); Atlantic Ocean,
Falkland Island, East Falkland; 1 Feb. 1969; S.W. Gor-
ham leg., CMNC 1992-0545. 9 ¢¢ and 99 (up to
18 mm); East Falkland; 25 Jan. 1967; S.W. Gorham leg.;
CMNC 1992-0552.

Distribution. Magallanes and Chilean Antarctica Re-
gion, Tierra del Fuego, Falkland Islands, Crozet Islands,
Kerguelen Islands, Peru, South Africa, South Australia,
New Zealand (Chatham Rise, Macquarie Island, Auck-

319

land Island, Campbell Island), Tasmania (Dana 1853;
Vanhoffen 1914; Tattersall 1914; Kussakin 1967; Brandt
and Wagele 1989).

Exosphaeroma rhomburum (Richardson, 1899)
Fig. 2G

Sphaeroma rhomburum Richardson, 1899: 835-836, fig. 12; 1900: 222.

Exosphaeroma rhomburum.— Richardson, 1905: 290, fig. 303; Nier-
strasz 1931: 195; Schultz 1969: 135, fig. 197; Kussakin 1971: 402,
fig. 257; Bruce 2003: 369.

Type locality. Monterey Bay, California.

Material examined. USA. 3 2 (up to 4.1 mm), 2 ovig-
erous 9 9 (up to 4.2 mm); Washington, Clallam Co., Makah
Bay; 31 July 1966; E.L. Bousfield leg.; CMNC 1991-2559.
Canaba. 4 3 (up to 4.5 mm), 4 9 (up to 5 mm), Brit-
ish Columbia, Range 5 Coast; 13 July 1964; E.L. Bousfield
leg.,; CMNC 1984-1535. 3 Jo (up to 4.1 mm), 4 9° (up to
4.5mm); British Columbia, Rupert Land Distr., Hope Island;
22 July 1959; E.L. Bousfield leg.; CMNC 1984-1537. 1 °
(4 mm); Alaska, Prince of Wales-Outer Ketchikan Census
Area; 31 May 1961; E.L. Bousfield leg.; CMNC 1991-2501.

Distribution. Monterey Bay, California, Washington
to British Columbia.

Exosphaeroma russellhansoni Wall, Bruce &
Wetzer, 2015
Figs 2H, 3A, B

Exosphaeroma russellhansoni Wall, Bruce & Wetzer, 2015: 28-33,
figs 9-12.

Type locality. Washington, Puget Sound, Seattle, Smith
Cove.

Material examined. CaNnabDa. 15 @¢ (up to 6.2 mm);
British Columbia, Esquimalt Land Distr., Esquimalt; 30 July
1970; E.L. Bousfield leg.; CMNC 1984-1465. 2 64 (5.1,
5.5 mm); British Columbia, Queen Charlotte Islands Land
Distr., Yakan Pt.; 25 Aug. 1975; E.L. Bousfield leg.; CMNC
1984-1445. 3 do (up to 6 mm); Land Distr., Graham Is-
land; 11 Aug. 1975; E.L. Bousfield leg.; CMNC 1984-1447.
4 33 (up to 5.8 mm), 2 2 9 (up to 4.8 mm); British Colum-
bia, Range 3 Coast Land Distr., Princess Royal Island; 20
July 1964; E.L. Bousfield leg.; CMNC 1984-1448. 5 ¢¢
(up to 6.5 mm), 2 2° (up to 4.8 mm); British Columbia,
Metchosin Land Distr., Witty’s Lagoon; 28 July 1964; EL.
Bousfield leg., CMNC 1984-1449. 16 3 (up to 6 mm),
3 2° (up to 5 mm); British Columbia, Range 3 Coast Land
Distr., Lady Douglas Island; 9 July 1964; E.L. Bousfield
leg., CMNC 1984-1452. 4 3 (up to 6.1 mm), 4 9° (up
to 4.1 mm); British Columbia, Range 3 Coast Land Dis-
tr., Miles Island; 5 Aug. 1964; E.L. Bousfield leg.; CMNC
1984-1452. 2 J (up to 6.5 mm); British Columbia, Range
5 Coast Land Distr., Stephens Island; 12 July 1964; E.L.
Bousfield leg.; CMNC 1984-1455. 4 @ (up to 6.8 mm),

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380 Khalaji-Pirbalouty, V. & Gagnon, J-M.: Sphaeromatoid isopods of CMN

Figure 2. Dynamene dilatata Richardson, 1899, female, (CMNC 1991-2567). A. Lateral view; B. Dorsal view; C. Dynamenella
Sheareri (Hatch, 1947), male, (CMNC 1990-0070); D. Dynoides canadensis Khalaji-Pirbalouty & Gagnon, 2021, (CMNC 1985-0667b).
Exosphaeroma gigas (Leach, 1818), male, (CMNC 1990-0090); E. Dorsal view; F. Lateral view. Exosphaeroma rhomburum (Rich-
ardson, 1899), male, (CMNC 1991-2559); H. Exosphaeroma russellhansoni Wall, Bruce & Wetzer, 2015, male, (CMNC 1984-1465).

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Zoosyst. Evol. 99 (2) 2023, 375-389

1 2 (5.5 mm); British Columbia, Rupert Land Distr.; 7 Aug.
1959: E.L. Bousfield leg.; CMNC 1984-1461. 6 33 (up to
6.1 mm), 3 29 (up to 5 mm); British Columbia, Vancouver
Island, Barclay Land Distr.; 8 September 1975; E.L. Bous-
field leg.; CMNC 1984-1463. USA. 10 3 (up to 6.5 mm);
Washington, Jefferson Co. (WA), Marrowstone Island; 26
July1966; E.L. Bousfield leg.; CMNC 1984-1456.

Exosphaeroma studeri VanhOffen, 1914
Fig. 3C, D

Exosphaeroma studeri Vanhoffen, 1914: 510-511, fig. 44; Menzies
1962a: 132-133, fig. 43; Nierstrasz 1931: 195; Bruce 2003: 369.
Sphaeroma calcarea— Dollfus 1891: 64, pl. 8, fig. 7. [Not Sphaero-
ma calcarea Dana, 1853; misidentification, according to Men-

zies 1962a].

Type locality. Punta Arenas, Chile.

Material examined. Cute. | 3 (17.1 mm), | ovig-
erous 2 (14.5 mm); Banco de las Taeas, Isla, Navarino;
5 Feb. 1970; E.L. Bousfield leg.; CMNC 2023-0242.
1 ¢ (18 mm) 5 8 (up to 12 mm), Canal Beagle; 19
Feb. 1970; E.L. Bousfield leg.; CMNC 2023-0243. 2 ¢¢
(9 mm), 3 28 (up to 6 mm); Isla Navarino; 5 Feb. 1970;
E.L. Bousfield leg.; CMNC 2023-0244. 1 4 (9.8 mm);
Peninsula Scott; W. Bank LW-HW; 20 Feb. 1970; EL.
Bousfield leg.; CMNC 2023-0245. 1 4 (15 mm), 2 sub-
adult ¢¢ (13 mm), 3 2° (up to 13 mm); Islotes, Ma-
mones, off Isla Lennox; 6 February 1970; E.L. Bousfield
leg.; CMNC 2023-0246. 2 29 (up to 9.1 mm); Punta Ro-
balo, Isla, Navarino; 29 Jan. 1970; E.L. Bousfield leg.;
CMNC 2023-0247.

Distribution. Chile,
zies, 1962a).

Straits of Magellan (Men-

Genus Gnorimosphaeroma Menzies, 1954

Gnorimosphaeroma oregonense (Dana, 1853)
Fig. 3E, F

Spheroma oregonensis Dana, 1853: 778, pl. 52x; Richardson 1899: 836;
1900: 223.

Sphaeroma olivacea— Lockington, 1877: 45, pl. 1.

Exosphaeroma oregonensis— Richardson, 1905: 296, figs 315, 316;
Hatch, 1947: 213, Pl. 6, figs. 82, 83.

Neosphaeroma oregonense.— Monod, 1931: 76, fig. 74.

Gnorimosphaeroma oregonensis — Miller, 1968: 12; Hoestlandt 1969:
325; Schultz 1969: 129, fig. 187; Kussakin 1971: 406-408, fig. 260—
262; Hoestlandt 1975: 31; Brusca et al. 2007: 537, pl. 243; Wetzer et
al. 2021: 32, figs 1-9 (Neotype designation).

Gnorimosphaeroma oregonensis lutea— Menzies, 1954: 406, figs 1-4,
6A-P; Riegel 1959: 154-161, fig. 1A.

Gnorimosphaeroma oregonensis oregonensis— Menzies, 1954: 406,
figs 5, 7A—E, 12; Riegel 1959: 154-161, fig. 1B.

Note. Latest synonymies to the species can be found in
Wetzer et al. (2021).

381

Material examined. CANaDa. 50 ¢'d (up to 9.5 mm),
7 22 (up to 6 mm); British Columbia, Sayward Land
Distr., Gowlland Island; 8 Jul. 1983; F. Rafi leg.; CMNC
1985-0715. 16 33 (up to 9.5 mm), 42 29 (up to 7 mm);
British Columbia, Nootka Land Distr.; 13 June 1976;
R.M. O’Clair leg.; CMNC 1987-0201. 75 3 & Q; British
Columbia, Sayward Land Distr.; 30 Aug. 1984; F. Rafi
leg.; CMNC 1986-0206. 6 3'¢ (up to 9 mm), 10 2° (up
to 6 mm); British Columbia, Comox Land Distr.; 28 Jul.
1959; E. L. Bousfield leg.; CMNC 1987-0141. 973&9;
Alaska, Juneau Borough; 13 Jun. 1961; E. L. Bousfield
leg.; CMNC 1987-0146. USA. 214&9; Washington,
Mason Co. (WA); 17 Jul. 1966; E. L. Bousfield leg.;
CMNC 1991-2543.

Distribution. Widely distributed in North America
from Alaska, British Columbia, and Vancouver to Wash-
ington (Kussakin 1979; Wetzer et al. 2021).

Genus [schyromene Racovitza, 1908

Ischyromene menziesi (Sivertsen & Holthuis, 1980)
Fig. 3G, H

Dynamenella menziesi Sivertsen & Holthuis, 1980: 41-48, figs 6-8, pl. 1.

Ischyromene menziesi.— Harrison & Holdich, 1982: 86. [New combina-
tion]; Bruce 2006: 20; Gonzalez et al. 2008: 174.

Dynamenella eatoni.— Menzies, 1962a: 135, fig. 44. [Not Dynamene
eatoni Miers, 1875; misidentification, according to Harrison and
Holdich 1982].

Type locality. Tristan da Cunha Island, south Atlan-
tic Ocean.

Material examined. Cute. | 3 (11 mm); Ma-
gallanes-Antarctica Region, Isla Lennox; 6 Feb.1970;
E. L. Bousfield leg.; CMNC 1985-0652. 1 subadult 3
(9 mm), 1 Ovi. 9 (9 mm), 2 299 (10 mm, 19 mm); Ma-
gallanes-Antarctica Region, Isla Hoste; 4 Feb. 1970; E. L.
Bousfield leg.; CMNC 1985-0653. 2 33 (10 mm), 2 2°
(9 mm); 5 juveniles; Magallanes-Antarctica Region, Isla
Picton; 7 Feb. 1970; E. L. Bousfield leg.; CMNC 1985-
0655. 10 Juveniles; Magallanes-Antarctica Region, Ca-
nal Beagle; 19 Feb. 1970; E. L. Bousfield leg.; CMNC
1985-0656.

Distribution. Chile, South Atlantic Ocean (Tristan da
Cunha, Nightingale, and Stoltenhoff Islands).

Genus Paracerceis Hansen, 1905

Paracerceis sculpta (Holmes, 1904)
Fig. 4A, B

Dynamene sculpta Holmes, 1904: 300-302, pl. 34, figs. 1-7.

Cilicaea sculpta.— Richardson, 1905: 318-319, fig. 349.

Paracerceis sculpta— Richardson, 1905: 9; Menzies 1962b: 340, 341,
fig. 2; Miller 1968: 14, fig. 3.; Brusca et al. 2007: 537, pl. 243.

Note. Latest synonymies to the species can be found in
Martinez-Laiz et al. (2018).

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382 Khalaji-Pirbalouty, V. & Gagnon, J-M.: Sphaeromatoid isopods of CMN

Figure 3. Exosphaeroma russellhansoni Wall, Bruce & Wetzer, 2015, male, (CMNC 1984-1465). A. Lateral view; B. Ventral view.
Exosphaeroma studeri Vanhoften, 1914, male, (CMNC 023-0242); C. Dorsal view; D. Lateral view. Gnorimosphaeroma oregon-
ense (Dana, 1853), male, (CMNC 1985-0715); E. Dorsal view; F. Lateral view. /schyromene menziesi (Sivertsen & Holthuis, 1980),
male, (CMNC 1985-0652); G. Dorsal view; H. Pleotelson, dorsal view.

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Zoosyst. Evol. 99 (2) 2023, 375-389

Type locality. SAN CLEMENTE ISLAND; San Diego,
California.

Material examined. | 3 (5.1 mm); Caribbean Sea;
15 Mar. 1968; N.A. Powell; CMNC 1992-0513. 2 3 (5
mm, 5.2 mm), 8 9 (up to 4.9 mm); Hawaiian Islands,
Coconut Island; 1 Mar. 1962; D.C. Matthews leg.; CMNC
2023-0248. 3 9° (up to 4.5); Radio Island, North Caroli-
na, 31 Mar. 1975; M.C. Govern leg.; CMNC 2023-0249.

Distribution. This species is widely distributed along
the North American Pacific coast from southern Califor-
nia in San Diego to Mexico (Menzies 1962b); Pakistan,
in the Indian Ocean (Yasmeen and Javed 2001); Hawaii,
Hong Kong, Japan, Australia, Argentina, Brazil and the
Azores (mid —Atlantic Ocean); in the Mediterranean
Sea from Tunisia, Italy, Greece, France, Malta, Cyprus
and Turkey. The wide distribution is most likely correct
and results from ship-borne translocations. (Ulman et al.
2017; Martinez-Laiz et al. 2018; Rumbold et al. 2018).

Genus Paradella Harrison & Holdich, 1982

Paradella dianae (Menzies, 1962)
Fig. 4C, D

Dynamenopsis dianae Menzies, 1962b: 341, fig. 3.
Dynamenella dianae.— Menzies & Glynn, 1968: 63, 113, fig. 3.
Paradella dianae.— Harrison & Holdich, 1982: 104, fig. 6.
Paradella quadripunctata— Van Dolah et al. 1984: 52.

Note. A comprehensive synonymy to the species can be
found in Martinez-Laiz et al. (2018: 8-10).

Type locality. The Bay of San Quintin, Baja California.

Material examined. USA. 4 ¢¢ (up to 7.5 mm);
3 98 (up to 5 mm); Carteret County, North Carolina; 8
Apr. 1975; E. L. Bousfield leg.; CMNC 1990-0073.

Distribution. Baja California (Menzies 1962b), Italy,
Egypt, Spain, Cyprus, Turkey, Libya Pakistan (Arabian
Sea); Cadiz Bay, Caleta Vélez, Motril, Barbate (Spain),
Australia (Menzies 1962b; Ulman et al. 2017; Martinez-
Laiz et al. 2018). Its presence in different parts of the world,
especially port cities, indicates a transport via shipping.

Genus Sphaeroma Bosc, 1802

Sphaeroma quadridentatum (Say, 1818)
Fig. 4E

Sphaeroma quadridentata Say, 1818: 400-401; De kay 1844: 44; White
1847: 102; Harger 1873 in Verrill, Smith and Harger 1873: 275, pl.5,
fig. 21; Kensley and Schotte 1989: 234, fig. 10.

Sphaeroma quadridentatum.— Harger 1880: 368-370, pl. 9, figs. 53, 54;
Smith 1964: 103, pl. 15, fig. 27; Miller 1968: 8, fig. 3; Schultz 1969:
128, fig. 183; Kussakin 1971: 394, fig. 248;

Type locality. Saint Catherine’s Island, Georgia.
Material examined. USA. 4 4 (up to 6 mm); 2 9°
(up to 5 mm); Virginia, Gloucester; 17 Apr. 1975; E. L.

383

Bousfield leg.; CMNC 1992-0583. 4 34 (up to 6 mm),
8 2° (up to 5 mm); Pawley’s Greek, Carolina; 17 Mar.
1915; E. L. Bousfield leg.; CMNC 2023-0250. 2 34 (up
to 6 mm); 3 29 (up to 5.5 mm); South Carolina; 16 May
1975; D.R. Calder leg.; CMNC 2023-0251.

Distribution. Georgia, Florida, Long Island Sound,
Connecticut, New Haven County, West Haven, Savin
Rock. It is common on the southern shore of New En-
gland (Harger 1880; Kensley and Schotte 1989).

Sphaeroma terebrans Bate, 1866
Fig. 4F

Sphaeroma terebrans Bate, 1866: 28, pl. 2, fig. 5; Stebbing 1904: 16;
Richardson 1905: 282-286, figs. 294-298; Calman 1921: 217-218;
Baker 1926: 247-278; Nierstrasz 1931: 192; Van Name 1936:
447-449, fig. 279; Barnard 1940: 405; Pillai 1954: 9; Pillai 1955:
129-131, Pl. 6, figs 1-11; Loyola e Silva 1960: 14—28, figs 1, 2;
John 1968: 1-73, pl. 1, figs 1-36; Miller 1968: 11, fig. 3; Harrison
and Holdich 1984: 287-292, fig. 4; Kensley and Schotte 1989: 234,
fig. 10; Wilkinson 2004: 1; Baratti et al. 2005: 225-234; Li et al.
2016: 307, fig. 2.

Sphaeroma destructor Richardson, 1897: 105-107.

Type locality. India.

Material examined. USA. 3 3'¢ (up to 9 mm); South
Carolina, Charleston Co.; 25 Apr. 1965; E. L. Bousfield
leg.; CMNC 1991-2576. NIGERIA. 15 43 (up to 9 mm),
30 9 (up to 7 mm); Mayuku Creek; 5 Oct. 1975; C.
Powell leg.; CMNC 2023-0252.

Distribution. Virginia to Louisiana; Belize; Cuba;
Venezuela to Brazil; Gulf of Mexico; Kenya, Nigeria,
Tanzania, Zanzibar, east coast of southern Africa, Paki-
stan, India, Sri Lanka, Thailand, Indonesia, Philippines,
Australia, China (Kensley and Schotte 1989; Wilkinson
2004; Li et al. 2016).

Family Tecticipitidae Iverson, 1982
Genus Tecticeps Richardson, 1897

Tecticeps convexus Richardson, 1899
Fig. 4G, H

Tecticeps convexus Richardson, 1899: 837-838, fig. 15; Richardson 1905:
278-280, figs 290-291; Richardson 1906: 4, figs 6-9; Schultz 1969:
116, fig. 161; Kussakin 1971: 347, figs 210, 211; Brusca etal. 2007: 538.

Type locality. Monterey Bay, California.

Material examined. ALASKA. 10 ¢¢ (up to 9 mm);
25 9° (up to 10.5 mm); Sitka Borough, Chichagof Is-
land; 30 Jul. 1980; G. Peter & G. Ronald leg.; CMNC
1992-0541. Canapa. 15 @&Q; British Columbia, Range
4 Coast Land Distr., Banks Island;18 Jul. 1964; E. L.
Bousfield leg.; CMNC 1991-2583. 19 ¢&9; British Co-
lumbia, Range 2 Coast Land Distr., Goose Island; 5 Aug.
1964; E. L. Bousfield leg.; CMNC 1991-2584. 2 3&9;

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384 Khalaji-Pirbalouty, V. & Gagnon, J-M.: Sphaeromatoid isopods of CMN

Figure 4. Paracerceis sculpta (Holmes, 1904), male, (CMNC 1992-0513). A. Dorsal view; B. Lateral view. Paradella dianae (Menzies,
1962), male, (CMNC 1990-0073); C. Dorsal view; D. Pleotelson, dorsal view; E. Sphaeroma quadridentatum (Say, 1818); EF. Sphaero-
ma terebrans, Bate 1866. Tecticeps convexus Richardson, 1899, male, (CMNC 1992-0541); G. Dorsal view; H. Head ventral view.

British Columbia, Vancouver Island, Barclay Land Distr., Pt.; 1 Aug. 1966; E. L. Bousfield leg.; CMNC 1991-2562.
Pachena Bay; 22 Jul. 1975; E. L. Bousfield leg.; CMNC 2 29; Oregon, Tillamook Co., Cape Kiwanda; 16 Aug.
1991-2590. USA. 33 6&9; Washington, Clallam Co., 1966; E. L. Bousfield leg.; CMNC 1991-2574.

Crescent Bay; 27 Jul. 1966; E. L. Bousfield leg.; CMNC Distribution. Eastern Pacific, British Columbia, Ore-
1991-2550. 7 O&Q; Washington, Clallam Co.,Shipwreck gon to California (Espinosa-Pérez and Hendrickx 2006).

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Zoosyst. Evol. 99 (2) 2023, 375-389

Discussion

The first contribution to the knowledge of isopod taxon-
omy of North America was Say (1818), who described
Naesa depressus and N. ovalis from the east Coast of
North America; these species were later placed in the
genus Ancinus and Cassidinidea by H. Milne Edwards
(1840) and Hansen (1905), respectively. A third sphaero-
matid isopod, Sphaeroma quadridentatum, was described
by Say (1818) from Georgia and East Florida. Dana
(1853) described Spheroma oregonensis from the Pacific
Coast of North America; this species being transferred to
the combination Gnorimosphaeroma oregonense (Dana,
1853) by Wetzer et al. (2021). Harger (1873) reported
Sphaeroma quadridentatum (Say, 1818) to occur from
Massachusetts to Florida. Historically, the major period
of isopod documentation in North America was from
Richardson’s first major (1899) contribution to Kensley
and Schotte’s (1989) field guide. Prior to 1899, 75 iso-
pod species had been recorded for North America. When
Richardson (1899) worked on the North American Pacific
Coast, she added 22 new species to the isopod fauna, in-
cluding seven new sphaeromatids.

At the turn of the 20" century, in 1905, Harriet Rich-
ardson published her significant monograph on the iso-
pod fauna of North America in the Bulletin of the U.S.
National Museum. This contribution included all terres-
trial, freshwater, and marine isopods and Tanaidacea. Out
of about 240 marine isopod species, 31 species (~13%)
were from the family Sphaeromatidae. The next com-
prehensive account was given by Hatch (1947), who
gave a systematic account of the Pacific Northwest crus-
tacean isopods, reporting 70 species, most of them (51
species) being marine isopods, of which seven species
(~14%) were sphaeromatids. Schultz (1969), in “How to
know the marine isopod crustaceans’, \isted 444 marine
species known from the waters off the coasts of North
America, of which 45 (~10%) are sphaeromatid isopods.
Finally, Kensley and Schotte (1989) reported about 280
shallow-water species from the Caribbean, of which 30
species (~11%) were sphaeromatid isopods.

There are now 110 accepted genera and close to 650
named species of the sphaeromatoid isopods worldwide
(Boyko et al. 2008 onwards). Given that there are approx-
imately 6250 species of marine and estuarine isopods
(Poore and Bruce 2012), sphaeromatoid isopods account
for approximately 10% of the marine species. The ratio
of North American sphaeromatids to the other marine iso-
pods appears to be around 10% as well.

In recent decades, a few taxonomic studies of the North
American marine isopod fauna have described or report-
ed new species. For instance, Paradella tiffany Bruce &
Wetzer, 2004, and P. garsonorum Wetzer & Bruce, 2007, are
both described from Baja California, Mexico. Later, Bruce
and Wetzer (2008) reported a Pseudosphaeroma Chilton,
1909 species from the Pacific coast of North America.

Wall et al. (2015) added three new north-eastern Pacif-
ic species, Exosphaeroma paydenae, E. russellhansoni,

385

and E. pentche. Recently, Wetzer et al. (2021) revised
Gnorimosphaeroma oregonense (Dana, 1852) from the
north-eastern Pacific. Furthermore, Khalaji-Pirbalouty and
Gagnon (2021) described Dynoides canadensis from the
south-western coast of British Columbia, and Cassidini-
dea ovalis, a common species along the Atlantic Coast of
North America, was redescribed by Khalaji-Pirbalouty and
Bruce (2021). At last count, 70 species of sphaeromatid
isopods have been reported from North America (Wall et
al. 2015; Khalaji-Pirbalouty and Gagnon 2021; Wetzer et
al. 2021). This accounts for roughly 10% of all reported
sphaeromatid isopods worldwide. In comparison, Austra-
lia, with the length of its coastline approximately 15% of
that of North America, has a rich sphaeromatid fauna with
203 recorded species, which are nearly 30% of all sphaero-
matid species known worldwide (Poore et al. 2002; Bruce
2003, 2009; Poore 2005). Yet, it 1s estimated that the spe-
cies number for this family in Australia is still about 50%
of the expected total (Poore et al. 2002; Bruce 2003).

The relatively low number of North American sphaero-
matid species may be, in part, related to the lower-diversi-
ty trends observed for most taxa in higher-latitude regions,
particularly in the Northern Hemisphere. The number of
known sphaeromatid species reported from Australia is
about three times that of North America; it appears that
temperate regions are more favorable for sphaeromatid
isopods, as previously mentioned by Poore and Bruce
(2012). There are clearly other major factors at play, such
as the age of the crust around Australia, and also that the
family may have primarily evolved and radiated largely
in the Southern Hemisphere (unpublished results).

Taxonomic diversity at the family level and above, as
reported here, would greatly benefit from the addition of
new, fresh material from the field (particularly from un-
der-collected regions and using finer scale methods that
capture small crustaceans) which, beyond morphological
examination, would provide the additional opportunity
to run genetic analyses. There is, however, still lots of
material to be examined in collections such as that of the
CMN. Work by isopod taxonomists in these collections
will likely result in the description of many new species
(for instance, see Khalaji-Pirbalouty and Gagnon 2021),
especially where there are lots of unexamined samples
with unidentified species, or where previous identifica-
tions have not been reviewed by experts.

Acknowledgements

We express our gratitude to the Canadian Museum of
Nature, Ottawa (Visiting Scientist Awards 2019) and
Shahrekord University, for providing financial support
for this project. Special thanks to Philippe Ste-Marie
(Assistant Collections Manager, Invertebrate Collections,
Zoology) for his assistance during the first author’s visit
to the Canadian Museum of Nature. We deeply appreci-
ate the constructive comments and helpful suggestions
from Wolfgang Wagele (Zoological Research Museum

zse.pensoft.net

386

Alexander Koenig, Bonn), Niel Bruce (Queensland Mu-
seum), and Brenda Doti (Universidad de Buenos Aires,
Argentina) that helped improve the manuscript.

References

Baker WH (1926) Species of the isopod family Sphaeromidae from the
eastern, southern and western coasts of Australia. Transactions and
Proceedings of the Royal Society of South Australia 50: 247-279.

Baratti M, Goti E, Messana G (2005) High level of genetic differentia-
tion in the marine isopod Sphaeroma terebrans (Crustacea Isopoda
Sphaeromatidae) as inferred by mitochondrial DNA analysis. Jour-
nal of Experimental Marine Biology and Ecology 315(2): 225-234.
https://doi.org/10.1016/j.jembe.2004.09.020

Barnard KH (1940) Contributions to the crustacean fauna of South Afri-
ca. XII. Further additions to the Tanaidacea, Isopoda and Amphipo-
da, together with keys for the identification of the hitherto recorded
marine and freshwater species. Annals of the South African Museum
32: 381-543.

Bate CS (1866) Carcinological gleanings. No. I. Annals and Mag-
azine of Natural History, series 3 17(97): 24-31. https://doi.
org/10.1080/00222936608679472

Boyko CB, Bruce NL, Hadfield KA, Merrin KL, Ota Y, Poore GCB,
Taiti S [Eds] (2008 [onwards]) World Marine, Freshwater and Ter-
restrial Isopod Crustaceans database. Sphaeromatoidea Latreille,
1825. World Register of Marine Species. https://www.marinespe-
cies. org/aphia.php? p=taxdetails&id=292950 [on 2023-03-15]

Brandt A (1998) Sphaeromatidae (Crustacea, Isopdoda) of the Beagle
Channel and description of Cymodopsis beageli n. sp. Beaufortia
48(7): 137-161.

Brandt A, Wagele JW (1989) Redescriptions of Cymodocella tubicauda
Pfeffer, 1887 and Exosphaeroma gigas (Leach, 1818) (Crustacea,
Isopoda, Sphaeromatidae). Antarctic Science 1(3): 205-214. https://
doi.org/10.1017/S0954102089000325

Bruce NL (2003) New genera and species of sphaeromatid isopod
crustaceans from Australian marine coastal waters. Memoirs of the
Museum of Victoria 60(2): 309-369. https://doi.org/10.24199/).
mmv.2003.60.28

Bruce NL (2006) A new species of /schyromene Racovitza, 1908
(Sphaeromatidae: Isopoda: Crustacea) from intertidal marine
habitats in New Zealand. Zootaxa 1220(1): 19-34. https://doi.
org/10.11646/zootaxa.1220.1.2

Bruce NL (2009) A new genus and new species of Sphaeromati-
dae (Crustacea: Isopoda) from the Great Barrier Reef, Australia.
Memoirs of the Museum of Victoria 66(1): 35-42. https://doi.
org/10.24199/).mmv.2009.66.5

Bruce NL, Wetzer R (2004) Paradella tiffany sp. nov., a distinc-
tive sphaeromatid isopod (Crustacea: Isopoda: Sphaeromatidea)
from Baja California, Mexico. Zootaxa 623(1): 1-12. https://doi.
org/10.11646/zootaxa.623.1.1

Bruce NL, Wetzer R (2008) New Zealand exports: Pseudosphaeroma
Chilton, 1909 (Isopoda: Sphaeromatidae), a Southern Hemisphere
genus introduced to the Pacific coast of North America. Zootaxa
1908(1): 51-56. https://doi.org/10.11646/zootaxa.1908.1.4

Brusca RC, Coelho VR, Taiti S (2007) Isopoda. In: Carlton JT (Ed.) The
Light and Smith Manual: Intertidal Invertebrates from Central Cali-

zse.pensoft.net

Khalaji-Pirbalouty, V. & Gagnon, JM.: Sphaeromatoid isopods of CMN

fornia to Oregon (4" edn.). University of California Press, Berkeley,
503-542.

Calman WT (1921) Notes on marine boring animals: Crustacea. Pro-
ceedings of the General Meetings for Scientific Business of the
Zoological Society of London 1921(2): 215-220. https://doi.
org/10.1111/j.1096-3642.1921.tb03260.x

Carvacho A (1977) Sur une importante collection d’isopodes des iles
Kerguelen. Comité National Frangais des Recherches Antarctiques
42: 173-191.

Chilton C (1906) List of Crustacea from the Chatham Islands. Transac-
tions of the N.Z. Institute 38: 269-73.

Conlan KE, Bousfield MA, Hendrycks EDA, Mills EL, Cook FR,
Gruchy CG (2016) A tribute to Dr. Edward Lloyd Bousfield,
1926-2016. Canadian Field Naturalist 130(4): 359-372. https://do1.
org/10.22621/cfn.v13014.1932

Cunningham RO (1871) Notes on the reptiles, Amphibia, fishes, Mol-
lusca, and Crustacea obtained during the voyage of H.M.S. Nassau
in the years 1866-1869. Transactions of the Linnean Society of
London 27(4): 465-502. https://doi.org/10.1111/j.1096-3642.1871.
tb00219.x

Dana JD (1853) Crustacea. Part II. In: Wilkes C (Ed.) United States
Exploring Expedition. During the years 1838, 1839, 1840, 1841,
1842. Under the command of Charles Wilkes. U. S. N. C. Sherman.
Philadelphia, 689-1618.

De Kay JE (1844) Crustacea. Zoology of New-York or the New-York
Fauna; comprising detailed descriptions of all the animals hitherto
observed within the State of New-York, with brief notices of those
occasionally found near its borders, and accompanied by appropri-
ate illustrations (Vol. 1, Part 6). Carrol and Cook, Albany, 70 pp.

Desmarest AG (1825) Considérations générales sur la classe des
crustacés, et description des espéces de ces animaux, qui vivent dans
la mer, sur les cOtes, ou dans les eaux douces de la France, Paris,
446 pp. https://doi.org/10.5962/bhI title.6869

Dollfus A (1891) Crustacés isopodes, 55-76. Mission Scientifique de
Cape Horn 1882-1883, Tome VI, Zoologie 55—76. Gauthier- Villars,
Paris.

Ellis JP (1981) Some type specimens of Isopoda (Flabellifera) in the
British Museum (Natural History), and the isopods in the Linnaean
Collection. Bulletin of the British Museum (Natural History). His-
torical Series 40(4): 121-128. https://doi.org/10.5962/p.271708

Espinosa-Pérez MC, Hendrickx ME (2006) A comparative analysis
of biodiversity and distribution of shallow-water marine isopods
(Crustacea: Isopoda) from polar and temperate water in the East Pa-
cific. Belgian Journal of Zoology 136: 219-247.

Giambiagi DC (1925) Resultado de la primera expedicion a Tierra del
Fuego (1921). Enviada por la Facultad de Ciencias Exactas, Fisicas
y Naturales de la Universidad Nacional de Buenos Aires. Crusta-
ceos, Isépodos. Anales de la Sociedad Cientifica Argentina 1925:
229-246.

Gomez Simes E (1979) Algunos isopodos de la Ria Deseado (Santa
Cruz, Argentina). Contribuciones Cientifica Centro de Investi-
gacion de Biologia Marina. Estacion Puerto Deseado, Buenos Aires
166: 5-16.

Gonzalez ER, Haye PA, Balanda MJ, Thiel M (2008) Systematic list
of species of peracarids from Chile (Crustacea, Eumalacostraca).
Gayana (Concepcion) 72(2): 157-177. https://doi.org/10.4067/

S0717-65382008000200006

Zoosyst. Evol. 99 (2) 2023, 375-389

Gueérin-Méneville FE (1843) Crustaces. In: Cuvier G (Ed.) Iconogra-
phie du Régne animal de G. Cuvier, ou représentation d’apreés nature
de l'une des espéces les plus remarquable et souvent non encore
figurées de chaque genre d’ animaux (Vol. 3). Bailliere, Paris.

Hale HM (1929) The Crustaceans of South Australia. Part 2. Handbooks
of the Flora and Fauna of South Australia, issued by the British
Science Guild (South Australia Branch). Adelaide: British Science
Guild (South Australian Branch), 201-380.

Hansen HJ (1905) On the propagation, structure and classification of
the family Sphaeromidae. The Quarterly Journal of Microscopical
Science 49: 69-135. https://doi.org/10.1242/jcs.s2-49.193.69

Harger O (1880) Report on the marine Isopoda of New England and adja-
cent waters. Report of the United States Commission of Fish and Fish-
eries. For 1878(6): 297-462. https://doi.org/10.5962/bh1.title. 1391

Harrison K, Holdich DM (1982) Revision of the genera Dynamenella,
Ischyromene, Dynamenopsis, Cymodocella (Crustacea: Isopo-
da), including a new genus and five new species of eubranchiate
Sphaeromatids from Queensland waters. Journal of Crustacean Bi-
ology 2(1): 84-119. https://doi.org/10.2307/1548115

Harrison K, Holdich DM (1984) Hemibranchiate sphaeromatids (Crusta-
cea: Isopoda) from Queensland, Australia, with a world-wide review
of the genera discussed. Zoological Journal of the Linnean Society
81(4): 275-387. https://doi.org/10.1111/).1096-3642.1984 tb01175.x

Haswell WA (1882) Catalogue of the Australian stalk- and sessile-eyed
Crustacea. Sydney, Australian Museum, 324 pp. https://doi.
org/10.5962/bh1 title. 1948

Hatch MH (1947) The Chelifera and Isopoda of Washington and ad-
jacent regions. University of Washington Publications in Biology
10(5): 155-274.

Hodgson TV (1910) Crustacea IX. Isopoda. In: Harmer SF (Ed.) Nation-
al Antarctic Expedition 1901-1904. Natural History (Vol. 5) (Zoolo-
gy and Botany). British Museum (Natural History), London, 77 pp.

Hoestlandt H (1969) Sur un sphérome nouveau de la céte pacifique
américaine, Gnorimosphaeroma rayi n. sp. (isopode flabellifere).
Comptes rendus hebdomadaires des Séances de |’ Académie de Sci-
ences, Paris, 268D: 325-327.

Hoestlandt H (1975) Occurrences of the Isopoda Flabellifera Gnorim-
osphaeroma rayi Hoestlandt on the coast of Japan, eastern Siberia
and Hawaii, with a brief note on its genetic polychromatism. Pub-
lications of the Seto Marine Biological Laboratory 22(1/4): 31-46.
https://do1.org/10.5134/175891

Holmes S (1904) Remarks on the sexes of Sphaeromids, with a descrip-
tion of a new species of Dynamene. Proceedings of the California
Academy of Sciences (3). Zoology 3: 295-306.

Hurley DE (1961) A checklist and key to the Crustacea Isopoda of New
Zealand and Subantartic Islands. Transactions of the Royal Society
of New Zealand (Zoology) 1: 259-292.

Hurley DE, Jansen KP (1977) The marine fauna of New Zealand: Fam-
ily Sphaeromatidae (Crustacea Isopoda Flabellifera). New Zealand
Oceanographic Institute Memoir 63: 1—80.

John PA (1968) Habits, structure and development of Sphaeroma terebrans
(a wood-boring isopod). University of Kerala Publications 1: 1-73.
Kensley B, Schotte M (1989) Guide to the Marine Isopod Crustaceans of
the Carribean. Smithsonian Institution Press, Washington D.C. and Lon-
don, 114 pp. [204, 232, and 234. ] https://doi.org/10.5962/bh1 title. 10375

Khalaji-Pirbalouty V, Bruce NL (2021) Redescription of the type spe-
cies of the genus Cassidinidea Hansen, 1905 (Crustacea: Isopoda:

387

Sphaeromatidae) with notes on geographic distribution of the New
World species. Marine Biology Research 17(5—6): 494—502. https://
doi.org/10.1080/17451000.2021.1990958

Khalaji-Pirbalouty V, Gagnon JM (2021) A new species of Dynoides
Barnard, 1914 (Crustacea, Isopoda, Sphaeromatidae) from Canada,
with notes on geographic distribution of the north-eastern Pacific
Ocean species. Marine Biology Research 17(1): 12—20. https://doi.
org/10.1080/17451000.2021.1892766

Krauss CFF (1843) Die Stidafrikanischen Crustaceen. Eine Zusammen-
stellung aller bekannten Malacostraca, Bemerkungen tiber deren
Lebensweise und geographische Verbreitung, nebst Beschreibung
und Abbildung mehrerer neuen Arten, Stuttgart, 68 pp. https://do1.
org/10.5962/bhI.title.4825

Kussakin OG (1967) Isopoda and Tanaidacea from the coastal zones of
the Antarctic and subantarctic. In Biological Results of the Soviet
Antarctic Expedition (1955-1958), 3. Isseldovaniia Fauny Morei
4(12): 220-380. https://doi.org/10.2307/1442262

Kussakin OG (1979) Marine and brackish-water Isopoda of cold and
temperate (boreal) waters of the Northern Hemisphere. Part 1. Fla-
bellifera, Valvifera, and Tyloidea). National Academy of Sciences,
USSR, Zoology [Opredeliteli po Faune SSR, Akademiya Nauk,
SSSR] 122: 1-470.

Kussakin OG, Vasina GS (1980) Additions to the marine Isopoda and
Gnathiida of Kerguelen Islands (Southern Indian Ocean). Tethys
9(4): 355-369.

Leach WE (1818) Cymothoadées. In: Cuvier F (Ed.) Dictionnaire des
Sciences Naturelles 12: 338-354.

Li X-F, Chong H, Zhong C-R, Xu J-Q, Huang J-R (2016) Identification
of Sphaeroma terebrans via morphology and the mitochondrial cyto-
chrome c oxidase subunit I (COI) gene. Zoological Research 37(5):
307-312. https://doi.org/10.13918/j.issn.2095-8137.2016.5.307

Lockington WN (1877) Remarks on the Crustacea of the Pacific coast,
with descriptions of some new species. Proceedings of the California
Academy of Sciences 7: 28-36. https://do1.org/10.5962/bhl.part.27534

Loyola e Silva J (1960) Sphaeromatidae do litoral Brasiliero (Isopo-
da-Crustacea). Boletim da Universidade do Parana. Zoologia 4: 1-182.

Loyola e Silva Jde (1971) Sébre os géneros Ancinus Milne Edwards,
1840e Bathycopea Tattersall, 1909, da colecdo U.S. Nat. Mus. (Iso-
poda-Crustacea). Arquivos do Museu Nacional, Rio de Janeiro 54:
209-223.

Martinez-Laiz G, Ros M, Guerra-Garcia JM (2018) Marine exotic iso-
pods from the Iberian Peninsula and nearby waters. PeerJ 6e4408.
https://doi.org/10.7717/peerj.4408

Menzies RJ (1954) A review of the systematics and ecology of the ge-
nus “Exosphaeroma,” with the description of a new genus, a new
species, and new subspecies (Crustacea, Isopoda, Sphaeromatidae).
American Novitates 1683: 1—24.

Menzies RJ (1962a) The zoogeography, ecology and systematics of
the Chilean marine isopods. Lunds Universitets Arsskrift A vd. 2,
57(11): 1-162.

Menzies RJ (1962b) The marina isopod fauna of Bahia de San Quintin,
Baja California, Mexico. Pacific Naturalist 3(11): 331-348.

Menzies RJ, Barnard JL (1959) Marine Isopoda on coastal shelf bot-
toms of southern California: Systematics and ecology. Pacific Natu-
ralist 1(11-12): 1-35.

Menzies RJ, Glynn PW (1968) The common marine isopod crustacea of
Puerto Rico. A handbook for marine biologists. In: Hummelinck W

zse.pensoft.net

388

(Ed.) Studies on the Fauna of Curagaoand other Caribbean Islands
(Vol. XX VII). The Hague Martinus Nijhoff, Leiden, 133 pp.

Miers EJ (1875) Descriptions of new species of Crustacea collected at Ker-
guelen’s Island by the Rev. A. E. Eaton. Annals and Magazine of Natural
History (ser. 4) 16: 73—76. https://doi.org/10.1080/0022293750868 1124

Miers EJ (1879) Crustacea. In: Eaton AE (Ed.) An Account of the Petro-
logical, Botanical, and Zoological Collections Made in Kerguelen’s
Land and Rodriguez During the Transit of Venus Expeditions in the
Years 1874-75. Philosophical Transactions of the Royal Society of
London 168: 200-214.

Miller MA (1968) Isopoda and Tanaidacea from buoys in coastal waters
of the continental United States, Hawaii, and the Bahamas (Crusta-
cea). Proceedings of the United States National Museum 125(3652):
1-53. https://doi.org/10.5479/si.00963801.125-3652.1

Milne Edwards H (1834-1840) Histoire Naturelle des Crustacés, Com-
prenant |’Anatomie, la Physiologie et la Classification de ces Ani-
maux (Vol. III). Librairie Encyclopédique de Roret, Paris, 638 pp.
https://doi.org/10.5962/bh1 title. 16170

Monod T (1931) Tanaidaces et Isopodes Aquatiques de |’ Afrique Occi-
dentales et Septentrional 3e Partie. Sphaeromatidae. Mémoires de la
Société des sciences naturelles de Neuchatel 29: 1-91.

Nierstrasz HF (1931) Die Isopoden der Siboga-Expedition HI. In: Max
Weber (Ed.) Isopoda Genuina, II. Flabellifera. Siboga Expéditie
(Vol. 32c). E. J. Brill, Leiden, 16—227.

Ortmann AE (1911) Crustacea of Southern Patagonia. Reports of
the Princeton University Expeditions to Patagonia, 1896-1899
(Zoology), 635-667. https://doi.org/10.5962/bh1 title. 10517

Pfeffer G (1886) Die Krebse von Siid-Georgien nach der Ausbeute der
Deutschen Station 1882-83. Jahrbuch der Hamburgischen Wis-
senschaftlichen Anstalten 4: 43-150. https://doi.org/10.5962/bhl.
title. 10084

Pillai NK (1954) A preliminary note on the Tanaidacea and Isopoda of
Travancore. Bulletin of the Central Research Institute. University of
Tranvancore, Trivandrum 3(1): 1-21.

Pillai NK (1955) Wood boring Crustacea of Travancore. I. Sphaeroma-
tidae. Bulletin of the Central Research Institute of the University of
Travancore, Series C. Nature and Science 4(1): 127-139.

Poore GCB (2005) Supplement to the 2002 catalogue of Australian
Crustacea: Malacostraca — Syncarida and Peracarida (Vol. 19.2A):
2002-2004. Museum Victoria Science Reports 7: 1-15. https://doi.
org/10.24199/).mvsr.2005.07

Poore GCB, Bruce NL (2012) Global Diversity of Marine Isopods (Ex-
cept Asellota and Crustacean Symbionts). PLoS ONE 7(8): 43529.
https://doi.org/10.1371/journal.pone.0043529

Poore GCB, Lew Ton HM, Bruce NL (2002) Sphaeromatidae Latreille,
1825. 221-252. In: Houston WWK, Beesley P (Eds) Crustacea:
Malacostraca: Syncarida, Peracarida: Isopoda, Tanaidacea, Micta-
cea, Thermosbaenacea, Spelaeogriphacea. Zoological Catalogue of
Australia. CSIRO Publishing, Melbourne, 433 pp.

Richardson H (1897) Description of a new species of Sphaeroma. Pro-
ceedings of the Biological Society of Washington 11: 105-107.

Richardson H (1899) Key to the isopods of the Pacific Coast of North
America, with descriptions of twenty-two new species. Proceedings
of the United States National Museum 21(1175): 815-869. https://
doi.org/10.5479/si1.00963801.21-1175.815

Richardson H (1905) A monograph on the isopods of North America.
Bulletin of the United States National Museum 54, 727 pp. https://
doi.org/10.5479/si.03629236.54.i

zse.pensoft.net

Khalaji-Pirbalouty, V. & Gagnon, JM.: Sphaeromatoid isopods of CMN

Richardson H (1906) Descriptions of new isopod crustaceans of the fam-
ily Sphaeromidae. Proceedings of the United States National Muse-
um 31(1479): 1-22. https://doi.org/10.5479/si.00963801.31-1479.1

Riegel JA (1959) A revision in the sphaeromid genus Gnoriomosphaero-
ma Menzies (Crustacea: Isopoda) on the basis of morphological,
physical, and ecological studies on two of its “subspecies”. The Bi-
ological Bulletin 117(1): 151-162. https://doi.org/10.2307/1539047

Rumbold C, Meloni M, Doti B, Correa N, Albano M, Sylvester F,
Obenat S (2018) Two new nonindigenous isopods in the Southwest-
ern Atlantic: Simultaneous assessment of population status and ship-
ping transport vector. Journal of Sea Research 138: 1-7. https://doi.
org/10.1016/j.seares.2018.04.008

Say T (1818) An account of the Crustacea of the United States, part 7. Jour-
nal of the Academy of Natural Sciences of Philadelphia 1: 374-401.

Schultz GA (1969) How to Know the Marine Isopod Crustaceans. W.M.
C. Brown Company Publishers, Iowa, 359 pp.

Schultz GA (1973) Ancinus H. Milne Edwards in the new world
(Isopoda, Flabellifera). Crustaceana 25(3): 267-275. https://doi.
org/10.1163/156854073X00254

Shimomura M (2008) Bathycopea (Isopoda: Sphaeromatidea: Ancin-
idae) from Japan, with descriptions of two new species and rede-
scription of B. parallela Birstein. Zootaxa 1678(1): 25-49. https://
doi.org/10.11646/zootaxa.1678.1.2

Sivertsen E, Holthuis LB (1980) The marine Isopod Crustacea of the
Tristan da Cunha archipelago. Gunneria 35: 1-128.

Smith RI (1964) Keys to marine invertebrates of the Woods Hole Re-
gion. Contribution No. 11, Systematics-Ecology Program, Marine
Biological Laboratory, Woods Hole, Massachusetts, 208 pp.

Stebbing TRR (1900) On some crustaceans from the Falkland Islands,
collected by Mr. Rupert Vallentin. Proceedings of the General Meet-
ings for Scientific Business of the Zoological Society of London
1900: 517-568. https://do1.org/10.5962/bh1.title. 10309

Stebbing TRR (1904) Gregarious Crustacea from Ceylon. Spolia Zey-
lanica, Bulletin of the National Museum. Ceylon 2: 1-29.

Stebbing TRR (1914) Crustacea from the Falkland Islands collected by
Mr Rupert Vallentin, FLS. Part I. Proceedings of the General Meet-
ings for Scientific Business of the Zoological Society of London
84(2): 341-378. https://doi.org/10.1111/j.1469-7998 1914. tb07042.x

Stephensen K (1927) Papers from Dr. Th. Mortensen’s Pacific expedi-
tion 1914-16. XL. Crustacea from the Auckland and Campbell Is-
lands. Videnskabelige Meddelelser fra Dansk naturhistorisk Foren-
ing 1 Kobenhaven 83: 289-390.

Stephensen K (1947) Tanaidacea, Isopoda, Amphipoda, and Pycnogoni-
da. In: Norske videnskaps-akademi i Oslo (Eds) Scientific Results of
the Norwegian Antarctic Expedition 1927-1928. Oslo, 90 pp.

Studer T (1884) Isopoden, gesammelt wahrend der reise SMS Gazelle
um die erde 1874-76. Kurfiirstlich Akademie der Wissenschaften,
Berlin, 27 pp.

Tattersall WM (1905) The marine fauna of the coast of Ireland. Part
V. Isopoda. Reports of the Department of Agriculture and Techni-
cal Instruction for Ireland. Scientific Investigations of the Fisheries
Branch 1904(2): 1-90.

Tattersall WM (1914) Order Isopoda: Tribe Flabellifera. Transac-
tions of the Royal Society of Edinburgh 49: 880-890. https://do1.
org/10.1017/S0080456800017178

Tattersall WM (1921) Crustacea. Part VI-Tanaidacea and Isopoda. Nat-
ural History Report. British Antarctic Terra Nova Expedition, 1910.
Zoology 3: 191-258. https://doi.org/10.5962/bh1 title. 1679

Zoosyst. Evol. 99 (2) 2023, 375-389

Ulman A, Ferrario J, Occhipinti-Ambrogi A, Arvanitidis C, Bandi A,
Bertolino M, Bogi C, Chatzigeorgiou G, Cigek BA, Deidun A,
Ramos-Espla A, Kogak C, Lorenti M, Martinez-Laiz G, Merlo G,
Princisgh E, Scribano G, Marchini A (2017) A massive update of
non-indigenous species records in Mediterranean marinas. PeerJ 5:
e3954. https://doi.org/10.7717/peerj.3954

Van Dolah RF, Knott DM, Calder DR (1984) Ecological effects of rub-
ble weir jetty construction at Murrells Inlet, South Carolina—Vol. I:
colonization and community development on new jetties. Technical
Report EL-84-4. Prepared by Marine Resources Research Institute,
Charleston, SC, for Coastal Engineering Research Center. Vicks-
burg, U.S. Army Engineer Waterways Experiment Station, 138 pp.
https://doi.org/10.5962/bh1.title.48315

Vanhoffen E (1914) Die Isopoden der deutschen Siidpolar-Expedition
1901-1903. Deutsche Stidpolar-Expedition. Zoologie 15: 447-598.
https://doi.org/10.5962/bh1 title. 10649

Verrill AE, Smith SI, Harger O (1873) Catalogue of the marine inver-
tebrate animals of the southern coast of New England, and adjacent
waters. In: Verrill AE, Smith SI (Eds) Report upon the invertebrate
animals of Vineyard Sound and adjacent waters, with an account of
the physical features of the region. Extracted from: Report of Profes-
sor S.F. Baird, Commissioner of Fish and Fisheries, on the condition
of the sea-fisheries of the south coast of New England in 1871 and
1872. Government Printing Office, Washington, 243 pp. https://dol1.
org/10.5962/bh1 title.3 1963

389

Wall A, Bruce N, Wetzer R (2015) Status of Exosphaeroma amplicauda
(Stimpson, 1857), E. aphrodita (Boone, 1923) and description of
three new species (Crustacea, Isopoda, Sphaeromatidae) from the
north-eastern Pacific. ZooKeys 504: 11-58. https://doi.org/10.3897/
zookeys.504.8049

Wetzer R, Bruce NL (2007) A new species of Paradella Harrison &
Holdich, 1982 (Crustacea: Isopoda: Sphaeromatidae) from Baja
California, Mexico, with a key to East Pacific species. Zootaxa
1512(1): 39-49. https://do1.org/10.11646/zootaxa.1512.1.2

Wetzer R, Wall A, Bruce NL (2021) Redescription of Gnorimosphaeroma
oregonense (Dana, 1853) (Crustacea, Isopoda, Sphaeromatidae),
designation of neotype, and 16S-rDNA molecular phylogeny of the
north-eastern Pacific species. ZooKeys 1037: 23-56. https://doi.
org/10.3897/zookeys. 1037.63017

White A (1847) List of the Specimens of Crustacea in the Collec-
tion of the British Museum. Natural History Museum, London,
143 pp.

Wilkinson LL (2004) The Biology of Spaeroma Terebrans in Lake
Pontchartrain, Louisiana with Emphasis on Burrowing. PhD Dis-
sertation, University of New Orleans, New Orleans, 205 pp. https://
scholarworks.uno.edu/td/205

Yasmeen R, Javed W (2001) A new record of Paracerceis sculpta
(Holmes, 1904) (Sphaeromatidae: Isopoda) from Pakistan, northern
Arabian Sea. Pakistan Journal of Marine Sciences 10: 43-48. https://
www.pakjmsuok.com/index.php/pjms/article/view/45

zse.pensoft.net