Diversity and distribution of the Isopoda (Crustacea, Peracarida) of Kuwait, with an updated checklist.

Thirty-eight species of Isopoda, belonging to 13 families and 29 genera, are listed from Kuwait based on previous literature records (of 17 species) and collections carried out along Kuwait's coastal and subtidal zones during the present study. The majority of species belongs to the suborder Cymothoida (23), followed by Sphaeromatidea (9), Oniscidea (3), Valvifera (2), and Asellota (1). In total, 25 species were collected and identified from 12 families and 22 genera from Kuwaiti coastal and subtidal areas. These include eight families, 15 genera, and 21 species recorded for the first time from Kuwait. Isopod diversity was highest in the sandy rock areas, including southern Kuwait, particularly in Al-Khiran and Al-Nuwaiseeb, and in mixed habitat (muddy, rocky, and sandy) intertidal transects such as in Failaka Island. The species number increased from the subtidal and lowest zones into the high tidal zone. Isopods were found in sandy substrata, among shells, cobbles, rocks, dead corals, and algae. Manal Abdulrahman Al-Kandari, Valiallah Khalaji-Pirbalouty, Hadeel Abdulkhaliq, Weizhong Chen.

Introduction

The isopod fauna in Kuwait’s intertidal and subtidal habitats have received little attention. The few significant accounts of Kuwait’s marine isopods are those of Bowman and Tareen (1983), describing six new species of . In addition, Abu-Hakima (1984) recorded a bopyrid, Chopra, 1923, and Jones (1986) in ‘Field Guide to the Seashores of Kuwait’ recorded six marine isopods from Kuwait. However, Stebbing, 1900, Roux, 1828; and Nobili, 1906, were misidentified in his guide. They are reidentified as sp., Khalaji-Pirbalouty & Wägele, 2010, and Khalaji-Pirbalouty, Bruce & Wägele, 2013, respectively, in this work.

Kensley, Schotte & Poore, 2007 and Kensley, Schotte & Poore, 2007 have been reported by Kensley et al. (2007) from the coasts of Kuwait and Saudi Arabia and, most recently, Jones and Nithyanandan (2012) reported four species of the genus Leach, 1815 from Kuwait and Saudi Arabia. In contrast, the isopod fauna along the Iranian coast of the Gulf has received more attention than adjacent regions (e.g., Khalaji-Pirbalouty and Wägele 2009, 2010a, b, c, 2011, 2012; Khalaji-Pirbalouty et al. 2013; Khalaji-Pirbalouty and Bruce 2014; Khalaji-Pirbalouty and Raupach 2014, 2016).

In 2013, a large-scale survey covering Kuwait’s entire coastline and offshore islands was initiated to document biodiversity, species distribution, and species abundance of the intertidal fauna. This survey was completed in 2017 (Al-Kandari et al. 2017). A further complementary sampling of four sites was conducted from 2016 to 2018. Survey results for molluscs, decapods, and polychaetes have been published (Al-Kandari et al. 2019a, b, 2020a, b), and summaries on other taxa are in progress. Here we report the results for the crustacean order .

Materials and methods

Intertidal and subtidal sampling

Thirty-eight intertidal transects and two subtidal sites were sampled quantitatively and qualitatively for macrofauna (Fig. 1, Table 1). Transects were located between Khor Al-Subiya in the north and the border with Saudi Arabia in the south. The surveys were conducted in daylight during the late autumn and winter seasons from December 2013 to December 2016. The sampling dates (see Table 1) and time for each site coincided with the lowest tides (as near to 0 chart datum as possible) using the Kuwait Port Authority’s Tide Tables for 2013, 2014, 2015, and 2016. Kuwait’s intertidal areas consist of coral, rocky, sandy, and/or muddy habitats or combinations thereof. At some transects, sandy mud or muddy sand covered a hard stratum throughout the intertidal range. Other transects consisted of combinations of sand and rocks, with some rocks lose and resting on the surface and others being part of the bedrock. All sandy beaches, rocky beaches, underneath stones, rubble, algal turf, and/or seagrass beds were sampled at each transect. Samples were left in seawater for a day before the fauna was collected from the bottom of the containers. Additionally, fauna living within porous rocks was collected by breaking the rocks with a hammer, placing the resulting debris in isotonic magnesium chloride solution, and collecting the fauna after their relaxation. For soft substrates, a 25 × 25-cm square metal box corer, 15 cm deep, was placed randomly, and sediment was collected by spade from inside the corer. These samples were sieved with seawater using 0.3-mm mesh sieves 45- and 75-cm in diameter, and all sediment and organisms remaining were preserved with 5% buffered formalin for subsequent picking and identification. Isopod specimens were also collected qualitatively from under rocks and among intertidal vegetation. Sand was sieved further samples were collected from rocks broken by a hammer, washing algae, sponges and seagrass, turning over stones, as well as collecting directly in the habitat. Material was rinsed under seawater, and all the washings passed through a 0.3-mm mesh sieve to collect any isopod specimens. The collected isopods were fixed in a 75–95% ethanol solution for subsequent morphological and molecular analyses. All specimens were deposited in the Kuwait Institute for Scientific Research () reference collection.

Figure 1.

Map of the 40 sampling sites established in Kuwait’s intertidal and subtidal zones; site numbers corresponding to Table 1.

Table 1.

Sampling sites studied in the intertidal and subtidal zones of Kuwait with habitat details (*KPC = Kuwait Petroleum Corporation).

Site No.	Site Name (north to south)	Position
		Sampling Dates	Coordinates	Area	Substrate
Khor Al-Subiya: north and west of Boubyan Island (BI)
1	Umm Al-Shajar (north Khor Al-Subiya), (BI1)	29.12.2015	29°54.263'N, 48°01.475'E	BI	Mud
2	Khor Al-Subiya (Al-Magasel)	23.11.2014	29°44.476'N, 48°05.740'E	North	Mud-rock
3	Khor Al-Subiya (Al-Shumaima)	24.11.2014	29°39.403'N, 48°07.850'E	North	Mud-rock
4	Khor Subiyah (south)	25.11.2014	29°34.849'N, 48°10.248'E	North	Mud
Kuwait Bay
5	Mudairah	30.12.2014	29°32.672'N, 47°55.394'E	Bay-mud	Mud
6	Al-Kuwaisat	17.11.2014	29°22.677'N, 47°42.480'E	Bay-mud	Mud
7	Al-Judailiat	02.02.2014	29°22.497'N, 47°45.183'E	Bay	Sand-rock
8	Aushairij	03.02.2014	29°23.047'N, 47°50.192'E	Bay	Sand-rock
9	Sulaibikhat Bay	06.11.2014	29°19.702'N, 47°49.670'E	Bay	Mud
10	Shuwaikh (KPC*), subtidal	22.02.2016	29°21.401'N, 47°56.390'E	Bay	Sand-rock
11	Kuwait Bay (Al-Salam Beach)	09.12.2013	29°21.631'N, 47°57.204'E	Bay	Sand-rock
12	Kuwait Bay (Ras Ajuza)	08.12.2013	29°23.481'N, 47°59.800'E	Bay	Sand-rock
East Kuwait Bay
13	Al-Sha'Eab	19.01.2014	29°21.979'N, 48°01.344'E	Middle 1	Sand-rock
14	Al-Salmiya	19.12.2013	29°20.313'N, 48°05.775'E	Middle 1	Sand-rock
South Kuwait Bay
15	Al-Messilah	18.12.2013	29°16.496'N, 48°05.407'E	Middle 1	Sand-rock
16	Al-Funaitees	19.12.2013	29°11.519'N, 48°06.938'E	Middle 1	Sand-rock
17	Abu Halifa	04.01.2014	29°08.154'N, 48°07.985'E	Middle 2	Sand-rock
18	Al-Mangaf	01.02.2014	29°06.041'N, 48°08.323'E	Middle 2	Sand-rock
19	Masfat Al-Ahmadi	10.12.2014	29°04.431'N, 48°08.676'E	Middle 2	Sand-rock
20	North Oil loading terminal, subtidal	28.09.2014	29°8.043'N, 48°09.139'E	Middle 2	Sand-rock
21	Mina Abdullah	16.02.2014	29°00.071'N, 48°09.853'E	Middle 2	Sand-rock
22	Al-Julaia'Ea	17.02.2014	28°49.480'N, 48°16.812'E	South	Sand-rock
23	Dohat Al-Zour	02.03.2014	28°46.100'N, 48°18.210'E	South	Sand-rock
24	Ras Al-Zour	08.01.2015	28°44.502'N, 48°22.950'E	South	Sand-rock
25	Al-Khiran	03.03.2014	28°38.813'N, 48°23.429'E	South	Sand-rock
26	Al-Nuwaiseeb	04.03.2014	28°34.794'N, 48°24.078'E	South	Sand-rock
Islands
27	Umm Al-Maradim Island, east (UI1)	11.11.2014	28°40.778'N, 48°39.207'E	UI1	Sand-rock
28	Umm Al-Maradim Island, northeast (UI2)	11.11.2014	28°40.939'N, 48°39.196'E	UI2	Sand-rock
29	Umm Al-Maradim Island, northwest	11.11.2014	28°40.960'N, 48°39.173'E	UI3	Sand-rock
30	Qaruh Island (north), (QI1)	10.11.2014	28°49.105'N, 48°46.553'E	QI	Sand-rock
31	Qaruh Island (south), (Q2)	10.11.2014	28°49.022'N, 48°46.607'E	QI	Sand-rock
32	Kubbar Island (east), (Q3)	09.11.2014	29°04.278'N, 48°29.655'E	KI	Sand-rock
33	Kubbar Island (west)	09.11.2014	29°04.377'N, 48°29.472'E	KI	Sand-rock
34	Auha Island (northwest), (AI)	10.02.2016	29°22.726'N, 48°26.269'E	AI	Sand-rock
35	Failaka Island (east 2), (FI1)	25.12.2014	29°23.710'N, 48°24.136'E	FI	Sand-rock
36	Failaka Island (east 1), (F2)	24.12.2014	29°23.629'N, 48°23.958'E	FI	Sand-rock
37	Failaka Island (south), (FI3)	23.12.2014	29°25.625'N, 48°20.307'E	FI	Mud-rock
38	Failaka Island (northwest), (FI4)	22.12.2014	29°28.049'N, 48°17.838'E	FI	Mud-rock
39	Boubyan Island (south), (BI2)	24.01.2015	29°38.993'N, 48°18.830'E	BI	Mud
40	Boubyan Island (Ras Al-Gayed), (BI3)	25.01.2015	29°48.093'N, 48°21.975'E	BI	Mud

Species identification

For identification, morphological studies were conducted using a Leica DFC450 camera mounted on a Leica M125 Stereomicroscope equipped with an imaging system that was employed to obtain colour images of the specimens. For greater depth of field, we merged 10–20 source images of a single specimen taken at different focus distances into one final image with the software LAS V4.5. The final image was edited using Adobe Photoshop. Isopods were identified to the lowest possible taxonomic level.

Results

In total, 25 species representing 12 families and 22 genera were identified from specimens collected in the present study. These species were collected from 31 intertidal transects, including 17 mainland and 14 island transects, and two subtidal sites (Table 2).

Table 2.

List of isopod species recorded in Kuwait in the present survey (* indicates a new record to Kuwait) and from literature records.

Suborder	Family	Species	Reference
CYMOTHOIDA	Anthuridae	Amakusanthura sp. *	This study
CYMOTHOIDA	Expanathuridae	Eisothistos sp. *	This study
CYMOTHOIDA	Cirolanidae	Atarbolanaexoconta*	This study
CYMOTHOIDA	Cirolanidae	Baharilanakiabii*	This study
CYMOTHOIDA	Cirolanidae	Cirolanatarahomii*	This study
CYMOTHOIDA	Cirolanidae	Eurydicearabica	Jones and Nithyanandan (2012)
CYMOTHOIDA	Cirolanidae	E.marzouqui	Jones & Nithyanandan, 2012
CYMOTHOIDA	Cirolanidae	E.peraticis	Jones and Nithyanandan (2012); This study
CYMOTHOIDA	Cirolanidae	Metacirolana sp. *	This study
CYMOTHOIDA	Corallanidae	Lanociragardineri*	This study
CYMOTHOIDA	Cymothoidea	Anilocramonoma	Bowman & Tareen, 1983
CYMOTHOIDA	Cymothoidea	Catoessagruneri	Bowman & Tareen, 1983
CYMOTHOIDA	Cymothoidea	Cymothoaeremita	Bowman & Tareen, 1983
CYMOTHOIDA	Cymothoidea	Jorymasawayah	Bowman & Tareen, 1983
CYMOTHOIDA	Cymothoidea	Mothocya sp.	Bowman & Tareen, 1983
CYMOTHOIDA	Cymothoidea	Nerocilaarres	Bowman & Tareen, 1983
CYMOTHOIDA	Cymothoidea	N.kisra	Bowman & Tareen, 1983
CYMOTHOIDA	Cymothoidea	N.sigani	Bowman & Tareen, 1983
CYMOTHOIDA	Cymothoidea	N.phaiopleura	Bowman & Tareen, 1983
CYMOTHOIDA	Gnathiidae	Gnathia sp.*	This study
CYMOTHOIDA	Gnathiidae	Elaphognathia sp.*	This study
CYMOTHOIDA	Bopyridae	Epipenaeonelegans	Abu-Hakima, 1984
CYMOTHOIDA	Bopyridae	Parabopyrella sp.*	This study
ONISCIDE	Ligiidae	Ligiapersica*	This study
ONISCIDE	Olibrinidae	Olibrinusantennatus*	This study
ONISCIDE	Tylidae	Tylosmaindroni	Taiti and Ferrara(1991); this study
SPHAEROMATIDEA	Sphaeromatidae	Cymodocedelvarii*	This study
SPHAEROMATIDEA	Sphaeromatidae	C.fuscina*	This study
SPHAEROMATIDEA	Sphaeromatidae	C.waegelei*	This study
SPHAEROMATIDEA	Sphaeromatidae	Dynamenellagranulata*	This study
SPHAEROMATIDEA	Sphaeromatidae	Heterodinamccaini*	This study
SPHAEROMATIDEA	Sphaeromatidae	Sphaeromakhalijfarsi*	This study
SPHAEROMATIDEA	Sphaeromatidae	S.walkeri*	This study
SPHAEROMATIDEA	Sphaeromatidae	S.annandalaei	This study
SPHAEROMATIDEA	Sphaeromatidae	Sphaeromopsissarii*	This study
VALVIFERA	Arcturidae	Arcturinoidesangulata	Kensley et al. (2007); this study
VALVIFERA	Arcturidae	Astacillamccaini	Kensley et al. (2007); this study
ASELLOTA	Paramunnidae	Heterosignum sp.*	This study

Latreille, 1825 was the best-represented family with five genera and eight species, followed by the family comprising five genera and five species. Two species were recorded in each of the families and . The remaining seven families were represented by single species (Table 2). In descending order, the most widely distributed isopod species were sp. from 20 transects, sp., and Khalaji-Pirbalouty & Wägele, 2009, from 18 transects, Kensley, Schotte & Poore, 2007, from 15 transects; Schotte & Kensley, 2005, from 12 transects; Khalaji-Pirbalouty, Bruce & Wägele, 2013, occurred at 12 transects, and Stebbing, 1904, was collected from ten transects. Interestingly, some species occurred in their 100s from single qualitative samples. Such high numbers for and were obtained from randomly collected at Al-Nuwaiseeb and Failaka Island. Similarly, high numbers of occurred on algal turfs from Kubbar Island. Other species found in high numbers were found from rocks, dead coral, or dead shells and included sp., , and Stebbing, 1905.

Thirty-eight isopod species under five sub-orders, 13 families, and 29 genera are listed in taxonomic order, including Kuwait’s previous records (17 species), type localities, and geographical distributions.

Taxonomy

Order

Suborder Wägele, 1989

Superfamily

Family Leach, 1814

Genus Nunomura, 1977

sp.

BD136C53-97FA-5215-89AD-A64A56933741

Figure 2A

Figure 2.

A sp. from Kubbar Island B sp. from Failaka Island C Bruce & Javed, 1987 from Masfat Al-Ahmadi D Khalaji- Pirbalouty & Wägele, 2011from Al-Nuwaiseeb E Khalaji-Pirbalouty & Wägele, 2011 from Quaruh Island F Jones, 1974 from Alkhiran G sp. from Um-Almaradim H Stebbing, 1904 from Al-Shamaimah.

­— Jones, 1986: 148, pl. 40 [not

Material examined.

Kuwait. 3 specimens; St. 2; ; 23 Nov. 2014; ♀♀, 2 juveniles; St. 3; ; 24 Nov. 2014; (5 ♀♀); St. 4; ; 25 Nov. 2014, 2 ♂♂; St. 8; ; 3 Feb. 2014; 1 ♀; St. 11; ; 9 Dec. 2013; 2 ♀♀; St. 12; ; 8 Dec. 21013; (1 ♀), 1 juvenile; St. 18; ; 1 Feb. 2014; 3 ♀♀; St. 19; ; 10 Dec. 2014; 2 ♀♀; St. 22; ; 17 Feb 2014, 1 juvenile, 6 ♀♀; St. 24; ; 08 Jan. 2015; 1 ♀; St. 25; ; 3 Mar. 2014; 5 ♀♀, 2 juveniles; St. 26; ; 4 Mar. 2014; 5 ♀♀, 3 juveniles; St. 27; ;; 11 Nov. 2014; 3 ♀♀; St. 28; ; 11 Nov. 2014; 1 ♀; St. 29; ; 11 Nov. 2014; 2 ♀♀; St. 30; ; 10 Nov. 2014; 7 ♀♀; St. 32; ; 9 Nov. 2014; 12 ♀♀; St. 34; ; 10 Feb. 2014; 3 ♀♀; St. 36; ; 24 Dec. 2014; 2 ♀♀; St. 38; ; 22 Dec. 2014.

Remarks.

(Negoescu, 1980) is the only species of this genus recorded from the nearest locality (Gulf of Aden). The specimens examined here differ from in the shape of the pleon with different lengths of pleonites 1–5 (vs. pleonites 1–5 similar to each other in ), the setation of pereopods, uropods and pleotelson; antenna and antennular articles are narrower than in .

Distribution.

New record for Kuwait.

Family Poore, 2001

Genus Haswell, 1884

sp.

7CD2EB0A-C6D9-5CE4-9198-A38DC4429B7E

Figure 2B

1 ♂; St. 38; ; 22 Dec. 2014.

New record for Kuwait

Family Dana, 1852

Genus Bruce & Javed, 1987

Bruce & Javed, 1987

E94991AF-292B-5446-9AF9-F11CBBB65E01

Figure 2C

Bruce & Javed, 1987: 145, figs 1, 2, Manora Island, Pakistan (type locality); Khalaji-Pirbalouty & Raupach, 2016: 155–162, figs 2–6.

4 ♂♂, 5 ♀♀; St. 19; ; 10 Dec. 2014; 1 ♂, 8 ♀♀; St. 21; ; 16 Feb. 2014; 2 ♀♀: St. 27; ; 11 Nov. 2014.

Pakistan, Oman Sea (Bruce and Javed 1987; Khalaji-Pirbalouty and Raupach 2016), new record for Kuwait.

Genus Bruce & Svavarsson, 2003

Khalaji-Pirbalouty & Wägele, 2011

1C5A4C11-46D3-5467-8122-CDFF9EAAD063

Figure 2D

Khalaji-Pirbalouty & Wägele, 2011: 34–39, figs 1–4; Qeshm Island, Iran (type locality).

1 ♀, 1 juvenile; St. 19; ; 10 Dec. 2014; 2 ♀♀; St. 25; ; 3 Mar. 2014; 3 ♂♂, 5 ♀♀; St. 26; ; 4 Mar. 2014; 1 ♂, 1 juvenile; St. 27; ; 11 Nov. 2014; 1 ♀, 1 juvenile; St. 32; ; 9 Nov. 2014; 1 ♀; St. 35; ; 25 Dec. 2014.

Qeshm Island, Hengam Island, Iran (Khalaji-Pirbalouty and Wägele 2011), new record for Kuwait.

Genus Leach, 1818

Khalaji-Pirbalouty & Wägele, 2011

DD5AF440-9865-5202-8592-7B94A07E966C

Figure 2E

Khalaji-Pirbalouty & Wägele, 2011: 39–45, figs 5–8; Qeshm Island, Iran (type locality).

7 ♀♀, 3 juveniles; St. 30; ; 10 Nov. 2014; 1 ♀, St. 32; ; 9 Nov. 2014.

Qeshm Island, Iran (Khalaji-Pirbalouty and Wägele 2011), new record for Kuwait.

Genus Leach, 1815

Jones, 1974

F4282042-FE79-5136-8194-D78A524C055F

Jones, 1974: 202, fig. 2, Red Sea (type locality); Bruce, 1986: 221.

Kuwait, Al-Ahmad Sea City waterways, Bahrain, Mashtan Island (Jones and Nithyanandan 2012).

Jones & Nithyanandan, 2012

8B98574E-A443-5EC8-9C30-9F05FEBC460C

Jones & Nithyanandan, 2012: 47–48, figs 1–4; Tarut Bay, Saudi Arabia (type locality).

Sabah Al-Ahmad Sea City Waterways, Kuwait; Manifa, Saudi Arabia (Jones and Nithyanandan 2012).

Jones, 1974

6D4CFC03-17B5-54A5-9E94-152340F678DE

Figure 2F

Jones, 1974: 204, fig. 3, Dammam, Saudi Arabia (type locality); Eleftheriou & Jones, 1976: 387; Bruce, 1986: 221;

1 ♂; St. 8; ; 3. Feb. 2014; 2 ♀♀; St. 19; ; 10 Dec. 2014; 2 ♀♀; St. 21; ; 16 Feb. 2014; 2 ♂♂, 3 ♀♀; St. 25; ; 3 Mar. 2014; 1 ♀; St. 34; ; 10 Feb. 2016; 3 ♂♂, 7 ♀♀; St. 39; ; 24 Jan. 2015; 6 ♂♂, 9 ♀♀, 2 juveniles; St. 40; 25 Jan. 2015.

Saudi Arabia, Bahrain, India, Pakistan, Kuwait (Eleftheriou and Jones 1976; Kazmi et al. 2002).

Genus Nierstrasz, 1931

sp.

A7FAE570-155F-5549-B803-D7A0A4AD3BE2

Figure 2G

1 ♂; St.2; ; 23 Nov. 2014; 4 ♀♀; St. 3; ; 24 Nov. 2014; 1 ♂; St.27; ; 11Nov. 2014; 1 ♂, 2 ♀♀; St.30; ; 10 Nov. 2014; 1 ♀; St. 34; ; 10 Feb. 2016; 2 ♂♂, 2 ♀♀; St. 36; ; 24 Dec. 2014.

New record for Kuwait.

Family Hansen, 1890

Genus Hansen, 1890

Stebbing, 1904

55F0F564-7242-5C0F-B779-859B28D02BF8

Figure 2H

Stebbing, 1904: 706, pl. LI, A, Mahlosmadulu Atoll, Maldive Islands (type locality).

A comprehensive synonymy to the species can be found in

1 ♂, 4 ♀♀, 3 juveniles; St. 3; ; 24 Nov. 2014; 2 ♀♀; St. 12; ; 8 Dec. 2104; 1 ♀; St. 22; ; 17 Feb. 2014; 1 Juvenile; St. 30; ; 10 Nov. 2014; 2 ♂♂, 5 ♀♀; St. 3; ; 24 Nov. 2014; 3 ♂♂, 6 ♀♀; St. 35; ; 25 Dec. 2014; 2 ♂♂, 5♀♀, 2 ovigerous ♀♀, 5 juveniles; St. 36; ; 24 Dec. 2014; 1 ♀; St. 37; ; 23 Dec. 2014; 1 ♂; St. 38; ; 22 Dec. 2014; 2 ♂♂; St. 40; ; 20 Jan. 2015.

Maldives, Kenya, Madagascar (Delaney 1989); Western Australia (Bruce and Sidabalok 2011); Iran (Khalaji-Pirbalouty, unpublished), new family for Kuwait.

Of the cymothoid isopods (parasites of fishes), the following species have been reported by Bowman and Tareen (1983).

Bowman & Tareen, 1983

8367960F-7CF2-54CE-A2DB-79DAE6C2E23F

Bowman & Tareen, 1983: 1, figs 3, 4, Kuwait (type locality).

Distribution.

Kuwait (Bowman and Tareen 1983).

Bowman & Tareen, 1983

694C969C-D652-555F-B7DB-C7F13E789E78

Bowman & Tareen, 1983: 18, figs 14, 15, Kuwait (type locality).

Kuwait (Bowman and Tareen 1983).

Bowman & Tareen, 1983

FBFB526F-3534-5348-907A-F9CE9AB9B78E

Bowman & Tareen, 1983: 21, figs 16–18, Doha, Kuwait (type locality).

sp., Mathews & Samuel, 1987: 144.

Kuwait (Bowman and Tareen 1983).

Bowman & Tareen, 1983

2D168B58-ADBD-52F9-B36E-B291ED09017A

Bowman & Tareen, 1983: 12, figs 10–12; Kuwait (type locality).

Bowman & Tareen, 1983: 8, figs 6–8.

Kuwait (Bowman and Tareen 1983).

Bowman & Tareen, 1983

73AEEDC7-A449-58F0-BBDE-15CC5216E0E1

Bowman & Tareen, 1983: 12, fig. 9; Kuwait (type locality).

Kuwait (Bowman and Tareen 1983).

Bleeker, 1857

620F65E4-F224-50E2-9312-C63DDE1885FC

Bleeker, 1857: 25–26, fig. 3, Java (type locality); Bowman & Tareen, 1983: 5, fig. 5.

A widespread species, recorded in the Indian Ocean from Hong Kong to South Africa (Bowman and Tareen 1983).

sp.

F08CC190-A0CE-5E01-90B4-69E19C3F68CC

sp., Bowman & Tareen, 1983: 25, fig. 19.

? (Brunnich, 1783), Bowman & Tareen, 1983: 25, fig. 20, India (type locality).

India (Bowman and Tareen 1983)

Genus Leach, 1814

sp.

83AAF206-E7E5-55F9-AA45-62A06C2BB501

Figure 3A

Figure 3.

A sp. from Al-Nuwaiseeb B sp., from Al-Shamaimah C from Al-Nuwaiseeb D Giordani Soika, 1954 from Kubbar Island E Khalaji-Pirbalouty, Bruce & Wägele, 2013 from Al-Nuwaiseeb F Schotte & Kensley, 2005 from USNM G Khalaji-Pirbalouty & Raupach,, 2014 from Al-Nuwaseeb H Javed & Ahmed, 1988 from Um-Almaradim.

1 ♂, 2 ♀♀; St. 7; ; 02 Feb. 2014; 1 ♀, 6 praniza larvae; St. 8; ; 3 Feb. 2014; 1 ♀, 6 praniza larvae; St. 10; ; 22 Feb. 2014; 1 ♂; St. 11; ; 9 Dec. 2013; 1 ♀; St. 12; ; 08 Dec. 2013; 3 ♀♀, 8 praniza larvae; St. 19; ; 10 Dec. 2014; 2 ♀♀, 1 praniza larva; St. 21; ; 16 Feb 2014; 1 ♀, 3 praniza larvae; St. 25; ; 3 Mar. 2014; 3 ♀♀, 1 ♂, 50 praniza larvae; St. 26; ; 4 Mar. 2014; 5 ♂♂, 9 ♀♀, 6 praniza larvae; St. 27; ; 11 Nov. 2014; 1 ♂, 3 praniza larvae; St. 28; ; 11Nov.2014; 8 ♂♂, 16 juveniles, 50 ♀♀, 3 praniza larvae; St. 30; ; 10 Nov. 2014; 3 ♀♀, 6 juveniles; St. 31; ; 10 Nov. 2014; 50 ♀♀ and praniza larvae; St. 32; ; 9 Nov. 2014; 3 ♀♀; St. 35; ; 25 Dec. 2014; 4 ♂♂, 2 sub adults ♂♂, 3 praniza larvae; St. 36; ; 24 Dec. 2014; 4 praniza larvae; St. 38; ; 22 Dec. 2014.

The specimen is closely related to Kensley, Schotte & Poore, 2009 from Khawr Musharraba, Saudi Arabia, Persian Gulf. However, it differs from by having a larger and conical mediofrontal process and bifid superior frontolateral process instead of a conical process. Also, the supraocular lobe is blunt and oblique rather than simply rounded.

New record for Kuwait.

Genus Monod, 1926

sp.

251C742C-9BF2-5015-A9AE-21787F90D6B3

Figure 3B

1 ♂; St. 3; ; 24 Nov. 2014.

The specimen is similar to Kensley, Schotte & Poore, 2009 in having the long, thin saber-like mandible from Somalia. However, it differs from in having a mandible with only one conical lobe at its base rather than two and having an acute mediofrontal process (vs. absent in ).

New record for Kuwait.

Family Rafinesque, 1815

Genus Nobili, 1906

Chopra, 1923

888CFCE7-4395-5342-BDA3-5F26B0A3FA2E

Chopra, 1923: 454–456, figs 6–11, Ganges Delta, India (type locality); Dawson, 1958: 240; Tareen, 1982: 159–160;

India; Kuwait; Abu Ali and Tarut (Saudi Arabia); Boushehr port (Iran).

Genus Markham, 1985

sp.

94B883AF-CCC5-5770-8B46-314535D72329

1 ♂, 1 ♀; St.8; ; 3 Feb. 2014.

Parasite, found on the gill of the common alpheid shrimp in Kuwait the De Haan, 1849.

New record for Kuwait.

Suborder Latreille, 1802

Family

Genus Fabricius, 1798

Khalaji-Pirbalouty & Wägele, 2010

36ED24AF-1E34-5741-BE18-9E5F4A6F18E9

Figure 3C

Khalaji-Pirbalouty & Wägele, 2010b: 136–149, figs 2–7; Kish Island, Iran (type locality).

Roux, 1828. – Jones, 1986: 148, pl. 40.

Material examined.

1 ♀; St. 7; ; 2 Feb. 2014; 4 ♂♂, 2 ♀♀; St. 8; ; 3 Feb. 2014; 3 ♂♂, 17 ♀♀; St. 12; ; 8 Dec. 2013 20 ♂♂ and ♀♀; St. 13; ; 19 Jan. 2014; 4 ♂♂, 6 ♀♀; St. 26; ; 4 Marc. 2014; 4 ♂♂, 4 ♀♀; St. 28; ; 11 Nov. 2014.

Iran, Oman, and United Arab Emirates (Taiti and Checcucci 2011; Khalaji-Pirbalouty and Wägele 2010), new record for Kuwait.

Genus Budde-Lund, 1913

Budde-Lund, 1902

CEC8FE68-199B-5DFA-8EC8-5054BDA01561

Budde-Lund, 1902: 379, Malaysia (type locality); Schmalfuss, 2003: 182; Taiti & Ferrara, 2004: 223, pl. 4.

1 ♀; St. 3; ; 24 Nov. 2014.

Indian Ocean (Taiti and Ferrara 2004), coastal waters of Iran (Khalaji-Pirbalouty, unpublished data), new record for Kuwait.

Genus Audouin, 1826

Giordani Soika, 1954

2B8EF8DB-1845-56F5-95A5-744FDBE7D46B

Figure 3D

Giordani Soika, 1954: 76, figs 8, 9, pl. 10, Oman Sea, Muscat (type locality); Ferrara & Taiti, 1986: 94; Taiti & Ferrara, 1991: 213, fig. 3;

sp. Jones, 1986: 149, pl. 40.

2 juveniles; St. 4; (1 ♀); St. 28; 11 Nov. 2014; 2 ♂♂, 2 ♀♀; St. 33; ; (1 ♀, 3 juveniles); St. 35; ; 25 Dec. 2014; 1 ♀, 4 juveniles; St. 38; ; 22 Dec. 2014.

Oman, Kuwait (Taiti and Ferrara 1991); Bandar-e-Charak, Bandar-e Bostanoo, Iran (Khalaji-Pirbalouty, unpublished data).

Family

Genus Leach, 1814

Khalaji-Pirbalouty, Bruce & Wägele, 2013

DBA64C0D-E100-5445-8D34-02E985AEE844

Figure 3E

Jones, 1986: 149, pl. 40 [not

1 ♂, 1 ♀, 1 subadult ♂, 1 juvenile; St. 3; ; 24 Nov. 2014; 6 ♂♂ 25 ♀♀, 6 sub adult ♂♂; St. 12; ; 8 Dec. 2013; 1 juvenile St.15; ; 18 Dec. 2013; 1 ♀, 1 sub-adult ♂, 1 juvenile; St.18; ; 1 Feb. 2014; 1 ♀; St.19; ; 10 Dec. 2014; 2 juveniles; St. 25; ; 3 Mar. 2014; 4 ♀♀, many juveniles; St. 26; ; 4 Mar. 2014; 1 ♀; St. 28; ; 11 Nov. 2014; 1 ♀; St. 32; ; 9 Nov. 2014; 1 ♂, 15 ♀♀; St. 34; ; 10 Feb. 2016; 1 ♀; St. 35; ; 25 Dec. 2014; 2 ♂♂, 26 ♀♀, many juveniles; St. 36; ; 24 Dec. 2014.

Bousher Province, Iran (Khalaji-Pirbalouty, Bruce and Wägele 2013), new record for Kuwait.

Schotte & Kensley, 2005

55D33660-9034-5770-8A22-2C93AF2D382E

Figure 3F

Schotte & Kensley, 2005: 1245–1248, figs 19–20, Safaniya and Manifa, Saudi Arabia (type locality);

sp. Jones, 1986: 149, pl. 40.

2 ♂♂ and 3 ♀♀; Kuwait Fishery Station (from Smithsonian Natural History Museum collection, USNM 1145230).

Saudi Arabia, United Arab Emirates, the Mediterranean basin, Greece (Schotte and Kensley 2005; Ulman et al. 2017), new record for Kuwait.

Khalaji-Pirbalouty & Raupach, 2014

8DD9F7F1-B0BE-5722-BBAC-E38D943EA7B5

Figure 3G

Khalaji-Pirbalouty & Raupach, 2014: 242–249, figs 7–12, Boushehr Province, Iran (type locality);

2 ♂♂, 3 ♀♀; St. 25; ; 3 Mar. 2014; 4 ♂♂, 9 ♀♀; St. 26; ; 4 Mar. 2014; 1 ♂, 1 ♀ St. 27; ; 11 Nov. 2014.

Bousher Province and Hengam Island, Iran (Khalaji-Pirbalouty and Raupach 2014; Khalaji-Pirbalouty et al. 2015), new record for Kuwait.

Genus Hansen, 1905

Javed & Ahmed, 1988

48B0A4BE-9C21-5DFB-B0B9-8B2B1092CA88

Figure 3H

Javed & Ahmed, 1988: 234–236, figs 1–3, Karachi coast, Pakistan (type locality).

Materials examined.

1 juvenile; St. 25; ; 3 Mar. 2014; 1 sub-adult ♂, 2 ♀♀, 1 juvenile; St. 28; ; 11 Nov. 2014; 4 sub-adults ♂♂, 5 ♀♀, 5 juveniles; St. 33; ; 9 Nov. 2014.

Pakistan and Iran coasts (Javed & Ahmed, 1988; Khalaji-Pirbalouty unpublished data), new record for Kuwait.

Genus Schotte & Kensley, 2005

Schotte & Kensley, 2005

1B083E25-E086-5B70-8594-2ACCD1A7A653

Figure 4A

Figure 4.

A Schotte & Kensley, 2005 from Al-Nuwaiseeb B Stebbing, 1905 from Al-Zhor C Khalaji-Pirbalouty & Wägele, 2010 from Boubyan Island D Khalaji-Pirbalouty & Wägele, 2009 from Kubbar Island E Kensley, Schotte & Poore, 2007 ♂, from Al-Doha F ♀ from Al-Doha G Kensley, Schotte & Poore, 2007 ♂, from Failaka Island H ♀, from Failaka Island.

Schotte & Kensley, 2005: 1259–1261, figs 27, 28, Manifa, Saudi Arabia (type locality).

> 100 ♂♂ and ♀♀; St. 7; ; 2 Feb. 2014; 1 ♀; St. 8; ; 7 ♀♀; St. 12; ; 8 Dec. 2013; 8 ♂♂, > 100 ♀♀ and Juveniles; St. 19; ; 10 Dec. 2014; 1 ♀; St. 21; ; 16 Feb. 2014; 5 ♂♂, 23 ♀♀; St. 25; ; 3 Mar. 2014; > 100 ♂♂ and ♀♀; St. 26; ; 4 Mar. 2014; 2 ♀♀, 1 juvenile; St. 32; ; 9 Nov. 2014; 2 ♀♀; St. 34; ; 10 Feb. 2016; 1 ♂, 1 ♀; St. 37; ; 23 Dec. 2014; 3 ♀♀; St. 38; ; 22 Dec. 2014.

Manifa and Ras Tanajib, Saudi Arabia (Schotte and Kensley 2005), new record for Kuwait.

Genus Bosc, 1802

Stebbing, 1905

1ADA4099-E967-5A0A-BFDD-345FEBBEE3A2

Figure 4B

Stebbing, 1905: 31–33, pl. VII, Jokkenpiddi Paar, Sri Lanka (type locality). Latest synonymies to the species can be found in

8 ♂♂, 5 ♀♀, 10 juveniles; St. 24; ; 8 Jan. 2015; 9 ♀♀; St. 25; ; 3 Mar. 2014.

is one the most widespread species of the marine isopods, reported along the Indian, Atlantic, and Pacific oceans coastal zones (Khalaji-Pirbalouty and Wägele 2010c; Martínez-Laiz et al. 2018).

Khalaji-Pirbalouty & Wägele, 2010

27441AEF-1DE5-5C61-B4B2-F39E5B5CDE28

Figure 4C

Khalaji-Pirbalouty & Wägele, 2010c: 3–9, figs 1–5, Qeshm Island, Iran (type locality).

3 ♀♀, 25 juveniles; St. 4; ; 25 Nov. 2014; 1 juvenile; St. 17; ; 4 Jan. 2014; 2 ♀♀; St. 26; ; 4 Mar. 2014; 2 ♂♂, 6 ♀♀, 6 juveniles; St. 39; ; 24 Jan. 2015; 4 ♂♂, 25 ♀♀, 21 juveniles; St. 40; ; 25 Jan. 2015.

Qeshm Island, Bandare Abbas, Bandare Kolahi, Iran (Khalaji-Pirbalouty and Wägele 2010c), new record for Kuwait.

Stebbing, 1911

AE51027D-796A-53C8-9555-BD3459ED4948

Stebbing, 1911: 181, pl. X, West Bengal, India (type locality); Barnard, 1936: 174; Barnard, 1940: 405; Pillai, 1955: 134, figs 23–35, pl. VII; Joshi & Bal, 1959: 62; Kensley, 1978: 113; Jones, 1986: 149, pl. 40; Khalaji-Pirbalouty & Wägele, 2010: 31–37, figs 1–5.

Ahmed, 1971: 77–79, fig. 1.

India, Habbanyyah Lake, and Shat Al- Arab River (Iraq); Arvand Kenar (Iran); Kuwait.

Genus Holdich & Jones, 1973

Khalaji-Pirbalouty & Wägele, 2009

C99F521E-F560-5F6C-944A-2B3CB12CF4DC

Figure 4D

Khalaji-Pirbalouty & Wägele, 2009: 34–42, figs 1–5, Kish Island, Iran (type locality).

1 ♀; St. 4; l; 25 Nov. 2014; 3 ♀♀l; St. 12; ; 8 Dec. 2013; 100 ♂♂ and ♀♀; St. 15; ; 2 ♂♂, 15 ♀♀; St. 18; ; 1 Feb. 2014; 1 ♂, 10 ♀♀; St. 21; ; 16 Feb. 2014; 2 ♂♂, 12 ♀♀, 2 Juveniles; St. 24; ; 8 Jan. 2015; 3 ♂♂, 8 ♀♀; St. 25; ; 3 Mar. 2014; 1 ♀; St. 26; ; 4 Mar. 2014; 22 ♂♂, 9 ♀♀, 2 juveniles; St. 27; ; 11 Nov. 2014; 28 ♂♂, 31 ♀; St. 28; ; 11 Nov, 2014; 35 ♂♂ and ♀♀; St. 29; ; 11 Nov. 2014; > 100 ♂♂ and ♀♀; St. 30; ; 10 Nov. 2014; 3 ♂♂, 43 ♀♀; St. 31; ; 10 Nov. 2014; > 100 ♂♂ and ♀♀; St. 32; ; 9 Nov. 2014; 9 ♂♂, 12 ♀♀, 3 juveniles; St. 33; ; 9 Nov. 2014 13 ♂♂, 14 ♀♀; St. 34; ; 10 Feb. 2016; > 100 ♂♂ and ♀♀, St. 36; ; 10 Feb. 2016; > 100 ♂♂ and ♀♀; St. 37; ; 23 Dec. 2014.

Kish, Qeshm, Hengam Islands, Iran (Khalaji-Pirbalouty and Wägele 2009; Khalaji- Pirbalouty et al. 2015), new record for Kuwait.

Suborder Sars, 1882

Family Sars, 1897

Genus Kensley, 1977

Kensley, Schotte & Poore, 2007 Figures 4E, 7F

11F249DD-7105-5B83-B945-BA0F3C648AD5

1 ♀; St. 7; ; 2 Feb. 2014; 1 ♀; St. 8; ; 3 Feb. 2014; 2 ♂♂; St. 34; ; 10 Feb. 2016; 1 ♂; St. 35; ; 25 Dec. 2014.

United Arab Emirates, Kuwait Bay, Kuwait (Kensley et al. 2007).

Genus Cordiner, 1793

Kensley, Schotte & Poore, 2007

71B9B655-C38B-5842-93D7-D2DC3F04315E

Figure 4G, H

10 ♂♂; St. 20; ; 28 Sep. 2014; 1 ♀; St. 21; ; 16 Feb. 2014; 1 ♂; St. 34; ; 10 Feb. 2016; 1 ♂; St. 35; ; 25 Dec. 2014; 6 ♂♂, 2 ovigerous ♀♀, 2 juveniles; St. 36; ; 25 Dec. 2014.

Manifa Bay, Saudi Arabia; Kuwait Bay, Kuwait (Kensley et al. 2007).

Family

Gamô, 1976

Type species.

Gamô, 1976

sp.

352BF6D4-077A-589A-AA7C-4781C592FD11

3 ♀; St. 28; ; 11 Nov, 2014; 1 ♀; St. 25; ; 3 Mar. 2014.

New record for Kuwait.

Discussion

Bowmen and Tareen (1983) were the first to study Kuwait’s isopod fauna, recording nine species of , all ectoparasitic on marine fishes (Table 6). Jones (1986) included six isopod species in his ‘Field Guide to the Seashores of Kuwait’: , , and are reidentified as sp., , and , respectively. Moreover, sp. of Jones (1986) is reidentified as and sp. is identified as . The widespread supratidal isopod was previously reported from Kuwait by Taiti and Ferrara (1991). However, Stebbing, 1911 was not found in the current study: the known distribution of is from the West Bengal estuaries in India to the Arvandroud (Shatt-Al-Arab) riverbanks between Iran and Iraq (Khalaji-Pirbalouty and Wägele 2010).

Two additional species, and , were collected from Kuwait Bay by Kensley et al. (2007). In monitoring the fauna of recently dredged canals in the Al-Khiran area of Kuwait, Jones and Nithyanandan (2012) discovered and described two new isopod species from Kuwait and mentioned the occurrence of a third, increasing the valid species of recorded from Kuwait to 17. With the present survey, we now count 38 species of , more than doubling Kuwait’s known isopod fauna. Twenty-one of the 25 species collected for this study represent first records for Kuwait (Table 2). Only four of these 25 species were reported previously: , , , and .

The geographical distribution of isopod species in Kuwait waters show very different patterns. The burrowing isopod was found living in soft rocks in the high intertidal area of the Al-Zour coast. The type locality of this species is Sri Lanka, and it has been considered restricted to the northern Indian Ocean. This thermophilic species is also tolerant to a range of salinities, and its distribution is worldwide in the tropics (Ríos-Touma et al. 2017). The ranges of other species are also limited to the Indian Ocean. For example, , is widely distributed from western Australia (Bruce and Sidabalok 2011) and the Maldives, Kenya, and Madagascar (Delaney 1989). Delaney (1989) recorded it from the Khor Abdullah estuary, Iraq, in the northwestern Gulf. Tolerance of salinity fluctuations is believed to be a primary reason for the wide distribution of this species throughout the Indian Ocean. Other species, such as , and , are widely distributed along the northeastern coast of the Gulf and along the Pakistani coast. Their distribution pattern is similar to some brachyuran decapods as suggested by Apel and Türkay (1999) and Naderloo et al. (2011). According to this distribution pattern, the fiddler crab fauna of the southern and western Gulf is similar in East Africa, the Gulf of Aden, and the Red Sea. At the same time, the fauna of the northeastern parts of the Persian Gulf is also somewhat similar to that of the northeastern Arabian Sea coasts of Pakistan and India. Finally, some of the known species of (e.g., ; , and ) are indigenous to the Gulf.

The new results reveal a low species richness of in Kuwait waters compared to the adjacent regions of the Indian Ocean. Based on Kensley’s (2001) isopod checklist, the Indian Ocean exhibits a high species diversity of more than 1000 species. Of these, 268 species inhabit the Indian coastal region, and fewer than half that number, 121 species, has been recorded from Pakistan’s coast by Kazmi et al. (2002). The apparently low species richness of in the Kuwait region compared to that of other areas of the Indian Ocean is due to Kuwait’s limited coastline, less than 200 km, but also to the Gulf’s young age, less than 6,000 years BP (Sheppard et al. 2010), and the harsh environmental conditions. The age of the environment is an essential factor for the evolution of diversity (Gaston and Chown 1999). The seabed regions of the Gulf presently at depths of 4–6 m have only been submerged for 3,000–4,000 years (Sheppard et al. 2010). Therefore, the current coastal habitat development is comparatively young.

The harsh environmental conditions in Kuwait coastal zone arise from high temperatures and high salinity. Salinities exceed 40 PSU, and summer temperatures often exceed 35 °C. For instance, from 2000 to 2013, the mean seawater temperature in Kuwait Bay was 23.6 °C with a range of 9.7–36.0 °C, and salinity ranged from 30–46 PSU (Al-Yamani et al. 2004). Furthermore, extreme air temperatures with highs up to 55°C in the summer months and winter lows around freezing are known from Kuwait (Jones 1986).

However, a comparison between this study and restricted localities of similar size suggested no lower diversity in Kuwait. Brusca (1987) reported 36 species of marine isopods from the Galapagos. Seventeen species of these were shallow-water species from the littoral to a depth of 100 m. Furthermore, Kensley (1984) identified only 24 species of isopods from the Belizean reef crest. The low species composition of these studies may arise from limited sampling. This study focuses on the Kuwaiti shoreline; therefore, many species living in sub-tidal depths were not collected.

Some isopod species appear to be introduced into Kuwait Bay from outside of the Gulf. For example, and were found in the subtidal zone of the Iranian and Arabian coasts of the Gulf, but were also recently reported from the Mediterranean basin, Greece (Ulman et al. 2017) and Egypt (pers. obs.). This distribution supports the hypothesis of a human- assisted introduction, such as through ballast water discharge. According to the Public Relations Department of Hormozgan Ports, Iran, ca. 53,000 tanker and cargo ships enter the Gulf annually and ca. 40% of the world’s total oil transportation passes through the Strait of Hormuz (Al-Yamani et al. 2015). In this context, ships transport a billion tonnes of ballast water annually, so although this intertidal study was comprehensive, it was only limited to sampling in the intertidal zone. Repeated sampling during different seasons as well as subtidal investigations would certainly increase Kuwait’s known isopod fauna.

The present study provides a baseline account of Kuwait’s coastal zone isopod fauna. The next step will be evaluating their ecology and conservation status. As Kuwait is one of the major oil exporters, invasive species are a significant issue, mainly due to the discharge of ballast water from oil tankers and cargo ships. Therefore, prevention is crucial for decision-making and implementation of invasion control and detection of new exotics. The results of this study highlight the need for further morphological as well as molecular studies to clarify the taxonomic status of some specimens, and a larger sampling effort in deeper waters of this area.