Corresponding author: Daniele Salvi (
Academic editor: Pavel Stoev
The rocky northern shores of Kuwait and those of the western, inner shores of Kuwait Bay are dominated by a small, densely encrusting oyster. The identity of this oyster has never been confirmed and was mistaken previously for a small
Al-Kandari M, Oliver PG, Salvi D (2021) Molecular and morphological systematics of a new, reef forming, cupped oyster from the northern Arabian Gulf:
The invertebrate fauna of the northern Arabian Gulf and that of Kuwait has a relatively recent investigation period as evidenced by the dearth of specific literature cited by
Oyster beds and reefs in northern Kuwait
Over the years 2014 to 2018, the Kuwait Institute for Scientific Research carried out a survey of the Kuwait’s intertidal fauna and the results for the
Glayzer, Glayzer and Smythe (1984) mentions an unidentified
Given that these oysters are key components of the intertidal communities and are cited as a feature for a potential world heritage reserve, their precise identity is important. Consequently, the senior author within Kuwait Institute for Scientific Research (
This paper concerns the identity of the oyster listed as
Representative samples of all shell morphs were collected during the
The sampling sites for the oysters considered here are indicated on the map (Fig.
Map of Kuwait indicating known distribution of
Sampling transects and localities of the intertidal oysters studied.
Transect | Location Name | Coordinates | Date | References |
---|---|---|---|---|
1 | Khor Al-Milh |
|
2004–2005 |
|
2 | Boubyan Island (north Khor Al-Subiyah) |
|
17.12.2015 |
|
3 | Khor Al-Subiyah (Al Maghasil) |
|
23.11.2014 | |
4 | Khor Al-Subiyah (Al-Alaimah) |
|
12.2019 | Visited |
5 | Khor Al-Subiyah (Shumaymah) |
|
24.11.2014 | |
6 | Khor Al-Subiyah (Ras Himar) |
|
25.11.2014 | Visited |
7 | Ras Kadmah (Al-Kuwaisat) |
|
17.11.2014 |
|
8 | Kuwait Bay (Ashairij) |
|
03.02.2014 |
|
9 | Between Ashairij and Umm Al-Namil Island |
|
29.03.2021 | Visited |
10 | Umm Al-Namil Island |
|
29.03.2021 | Visited |
Total genomic DNA was extracted from 24 alcohol-preserved tissues following standard high-salt protocols (
Genbank accession number, mitochondrial haplotype and GenSeq nomenclature (after
Specimen Catalogue # | Locality | GenBank accession number | Haplotype | GenSeq Nomenclature | |
---|---|---|---|---|---|
|
|
||||
NMW.Z.2021.009.001 (holotype) | Between Ashairij and Umm Al-Namil Island |
|
|
Hap1 | genseq-1 |
NMW.Z.2021.009.002/1 (paratype) |
|
|
Hap9 | genseq-2 |
|
NMW.Z.2021.009.002/2 (paratype) |
|
|
Hap10 | genseq-2 |
|
NMW.Z.2021.009.002/3 (paratype) |
|
|
Hap1 | genseq-2 |
|
NMW.Z.2021.009.002/4 (paratype) |
|
|
Hap11 | genseq-2 |
|
NMW.Z.2021.009.002/5 (paratype) |
|
|
Hap1 | genseq-2 |
|
NMW.Z.2021.009.002/6 (paratype) |
|
|
Hap12 | genseq-2 |
|
NMW.Z.2021.009.002/7 (paratype) |
|
|
Hap1 | genseq-2 |
|
NMW.Z.2021.009.002/8 (paratype) |
|
|
Hap1 | genseq-2 |
|
NMW.Z.2021.009.002/9 (paratype) | – |
|
na | genseq-2 |
|
NMW.Z.2021.009.002/10 (paratype) | – |
|
na | genseq-2 |
|
NMW.Z.2021.009.002/11 (paratype) |
|
|
Hap13 | genseq-2 |
|
NMW.Z.2021.009.004/1 (paratype) | Khor Al-Subiyah |
|
|
Hap1 | genseq-2 |
NMW.Z.2021.009.004/2 (paratype) |
|
|
Hap2 | genseq-2 |
|
NMW.Z.2021.009.004/3 (paratype) |
|
|
Hap3 | genseq-2 |
|
NMW.Z.2021.009.004/4 (paratype) |
|
|
Hap1 | genseq-2 |
|
NMW.Z.2021.009.004/5 (paratype) |
|
|
Hap4 | genseq-2 |
|
NMW.Z.2021.009.004/6 (paratype) |
|
|
Hap5 | genseq-2 |
|
NMW.Z.2021.009.004/7 (paratype) |
|
|
Hap1 | genseq-2 |
|
NMW.Z.2021.009.004/8 (paratype) |
|
|
Hap6 | genseq-2 |
|
NMW.Z.2021.009.004/9 (paratype) |
|
|
Hap1 | genseq-2 |
|
NMW.Z.2021.009.004/10 (paratype) |
|
|
Hap7 | genseq-2 |
|
NMW.Z.2021.009.004/11 (paratype) |
|
|
Hap8 | genseq-2 |
|
NMW.Z.2021.009.004/12 (paratype) |
|
|
Hap1 | genseq-2 |
Phylogenetic relationships were inferred by the Bayesian Inference method in BEAST 2.6.3 (
Mitochondrial sequences of the oysters from Khor Al-Subiyah and Ashairij represent 13 haplotypes differing from each other by one to three nucleotide substitutions occurring at 17 sites. Given their very limited genetic divergence, all 24 specimens analysed represent a single taxon. Phylogenetic analyses resolve the position of this taxon within the subfamily
Bayesian phylogenetic tree based on
Based on morphological and molecular assessments we assign these oysters to a new
The genus
All type material deposited in the National Museum of Wales (NMW.Z) Kuwait • 20 + specimens in two clumps; Kuwait Bay, between Ashairij and Umm Al-Namil Island;
Kuwait • remainder of shells in clumps; same collection data as for preceding; NMW.Z.2021.009.003. Kuwait • 20 + specimens in three clumps; Khor Al-Subiyah, Al Maghasil;
Kuwait, Kuwait Bay, between Ashairij and Umm Al-Namil Island,
(
Upper (right) valve smaller than and fitting into lower valve (Fig.
(
The lower valve is deeply cupped often with a deeper umbonal cavity related to the extension of the dorsal hinge plate. Attachment area over most of lower valve with interlocking spines and foliations. Outline is mostly oval but can be distorted into many shapes from lingulate to subcircular; the free margin is upturned, weakly convoluted with short blunt spines; except where growing in sheltered or uncrowded condition where the margins can be greatly extended into spathulate spines (Fig.
Upper valve smaller than, and fitting into lower valve. Rather flat but undulating, nacreous free margin very narrow slightly elevated. External surface weakly foliaceous but not raised into commarginal frills. Shells sheltered among others and juveniles frequently display open trumpet-shaped projections as above (Fig.
(
Attachment area small, free area with 5–7 prominent folds extending as furrowed spines (Fig.
Upper valve smaller than, and fitting into lower valve. Rather flat but undulating, nacreous free margin narrow, slightly elevated and extended as spines (Fig.
Anatomy of
The mantle in its preserved and contracted condition shows an array of radial folds (Fig.
Adductor muscle crescent shaped in a posterior ventral position; pericardium immediately dorsal to it (Fig.
Alimentary system (Fig.
The shell morphology of
The morphology and molecular results of
It is perhaps surprising that the Kuwait oyster belongs to the genus
Photographs of
The authors wish to thank all those who made it possible to collect the specimens for this study including Henk Dekker who accompanied us in 2019. We thank Matteo Garzia and Emanuele Berrilli at Salvi’s lab (University of L’Aquila) for their valuable molecular laboratory work. PGO wishes to thank Kenji Torigoe for his literature gift; to Bangor University for continuing electronic access to their library; to Harriet Wood of the National Museum of Wales for handling registration of the type material. Gratitude goes to the Kuwait Petroleum Corporation (KPC) and the Kuwait Institute for Scientific Research (