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- Hemicaridina Desmaresti. – Barrois 1893: 126–134: Figs 1–3.
- Atyaephyra desmarestii var. orientalis Bouvier, 1913: 65–74, Figs 1, 3C [type locality: Syria].
- Atyaephyra desmaresti. – Annandale and Kemp 1913: 241–244.
- Atyaëphyra Desmaresti. – Bouvier 1925: 84–89 Figs 159–162, partim.
- Atyaephyra desmarestii orientalis. – Holthuis 1961: 5–10, Figs 2C–E, 3C–H; Kinzelbach and Koster 1985: 127–133, Fig. 1, partim.
- Atyaephyra desmarestii mesopotamica Al-Adhub, 1987: 1–4, Fig. 1 [type locality: Shatt Al-Arab River and Hammar Lake, Iraq]. – Salman 1987: 27–42, Figs 1–8.
- Atyaephyra desmarestii. –Gorgin 1996: 662–668, Figs 1–2; Anastasiadou et al. 2004: 5–13, partim; Von Rintelen et al. 2012: 82–96, partim.
Turkey: 3 ♀♀ (CL 4.8–5.0 mm), Antalya, Kirkgoz Spring (Fig. 1, stn 107), 21.6.2006, coll. M. Özbek; 7 ♀♀ (CL 4.5–5.5 mm), SMF 12174, Akbez, Karasu River (Fig. 1, stn 108), 22.9.1982, coll. R.K. Kinzelbach. Syria: 10 ♀♀ (3 ovig.) (CL 5.0–6.0 mm) and 4 ♂♂ (CL 4.0–5.0 mm), SMF 12050, below the dam of Ascharna, Orontes River (Fig. 1, stn 109), 30/31.3.1979, coll. R.K. Kinzelbach; 34 ♀♀ (15 ovig.) (CL 4.1–4.8 mm), SMF 12188, north of M’adan, Euphrates River (Fig. 1, stn 110), 17.8.1978, coll. R.K. Kinzelbach; 3 ♀♀ (2 ovig.) (CL 4.5–5.6 mm), SMF SYR8, Euphrates River (Fig. 1, stn 111), 15/16.6.1998, coll. R. Beck. Israel: 3 ♀♀ (2 ovig.) (CL 4.7–5.3 mm) and 2 ♂♂ (CL 3.9–4.0 mm), SMF IES 1189, Te’o Spring (Fig. 1, stn 112), 16.2.1977; 9 ♀♀ (CL 4.3–6.0 mm) and 4 ♂♂ (CL 3.9–4.0 mm), Hula Lake (Fig. 1, stn 113), 29.1.1981, coll. D. Eurth; 2 ovig. ♀♀ (CL 3.8–3.9 mm), NHM 1918.104.22.168–12, Kinneret Lake (Fig. 1, stn 114), 24.7.1913; 1 ♀ (CL 3.9 mm), Samakh, Kinneret Lake, 6.5.1986, coll. R. Ortal; 1 ♀ (CL 4.4 mm), Zaki River (Fig. 1, stn 115), 6.5.1986, coll. R. Ortal; 1 ♀ (CL 4.0 mm), Jordan River (Fig. 1, stn 116), 6.5.1981, coll. R. Ortal; 1 ♀ (CL 4.2 mm) and 1 ♂ (CL 3.8 mm), NHM 1922.214.171.124.12, Jordan River, 26.1.1938. Jordan: 2 ♀♀ (1 ovig.) (CL 4.0–4.9 mm), SMF 12057, Al-Azraq Oasis (Fig. 1, stn 117), 24.3.1977, coll. H. Damian. Iraq: 12 ♀♀ (CL 5.6–6.8 mm) and 3 ♂♂ (CL 4.5–4.8 mm), Basrah, Garmat Ali marsh (Fig. 1, stn 118), 24.2.1987, coll. A.H.Y. Al-Adhub; 1 ♀ (CL 5.2 mm), NHM 19126.96.36.199–20, Basrah, Shatt Al-Arab River (Robat creek) (Fig. 1, stn 119), 14.11.1919, coll. Capt. Boulenger; 1 ♂ (CL 4.2 mm), NHM 19188.8.131.52–3, Basrah, Shatt Al-Arab River (Robat creek), 28.4.1919, coll. P.J. Barraud; 4 ♀♀ (1 ovig.) (CL 5.2–5.5 mm) and 1 ♂ (CL 4.8 mm), Basrah, Shatt Al-Arab River (Fig. 1, stn 120), 2011, coll. M.D. Naser; 1 ovig. ♀, NHM 19184.108.40.206, Amarah, Tigris River (Fig. 1, stn 121), 12.11.1919, coll. J.O. Cooper Esq.
Amendments to description
Rostrum long, slender, dorsal margin straight, slightly or strongly curved in the middle and pointed upwards or downwards, 6.0–10.0, most frequently (91% of the individuals examined) 6.5–9.25, × as long as high, shorter or equal to, or longer than scaphocerite (longer in 71% of the individuals examined). 14–29 (18–23 in 80% of the individuals) pre orbital teeth on dorsal margin of rostrum arranged to tip. 0–3, most often (85%) 1–3, post-orbital teeth. 3–13 teeth, mostly (96%) 4–10, arranged on ventral margin of rostrum (Fig. 3A). Carapace smooth with pterygostomial angle not protruding and rounded or bluntly produced (Figs 3B–C). Pleuron of fifth abdominal segment pointed ending in an acute or an obtuse posterior angle (Fig. 3D). Telsonwith 3–6, predominantly (93%) 4–5, pairs of dorsal spines arranged in curved fashion (Fig. 3E). Distal border of telson with 7–12, most often (91%) 8–10, spines (4–5 pairs) arranged in a fork-like or a fan-like way. Outermost pair of spines shortest, similar to dorsal spines, adjacent pair stronger terminating beyond, along with or before (beyond and along with in 64% of the individuals) the inner finely setulose pairs (Figs 3E–F). Basal segment of antennular peduncle with long stylocerite, with its tip failing to reach, reaching or overreaching the distal end of basal segment. Anterolateral lobe of basal segment short and pointed (Figs 3H–I). Distal segment of antennular peduncle with 0–3, most often (93%) 1–2, spines (Fig. 3G). Basal lower endite of maxilla densely covered with long simple setae arranged in 11–16 (12–15 in 93% of the individuals) oblique parallel rows. Endite of maxilla 1.75–2.20, mostly (93%) 1.81–2.07, × as long as basal lower endite (Fig. 4G). Basal endite of first maxilliped failing or reaching to distal end of exopod distal margin (Fig. 4F). Distal one-third of terminal segment of third maxilliped bearing 10–36 (14–31 in 84% of the individuals), mesial spines and one subdistal lateral spine near the base of larger terminal spine (Fig. 4H). Armature along flexor margin of dactylus of third and fourth pereiopod consisting of 6–11 (7–10 in 97% of the individuals) and 7–11 (8–10 in 89% of the individuals) spines (including terminal spine) respectively (Figs 4B, 4D). Merus of third and fourth pereiopod with 6–10 (7–9 in 85% of the individuals) and 5–9 (6–7 in 83% of the individuals) spines respectively (Figs 4A, 4C). Dactylus of fifth pereiopod with 33–55 (36–49 in 83% of the individuals) spines arranged in comb-like fashion on flexor margin (Fig. 4E). Endopod of first male pleopod expanded proximally with a distal portion stout and not tapering, often, with a, large protruding lobe in its outer subdistal part. Endopod with 13–38 spines arranged on a strongly curved inner margin and 5–8 setae arranged on outer margin (Fig. 4I, Bouvier et al. 1913: Fig. 1). 32–158 eggs of 0.5–0.75 × 0.35–0.5 mm in size.
Atyaephyra orientalis is a small-medium sized species of Atyaephyra, with maximum carapace length to be 4.8 mm in ♂♂, 6.8 mm in ♀♀ and 5.5 mm in ovig. ♀♀.
Atyaephyra orientalis can be differentiated from all other species of Atyaephyra by molecular characters, as demonstrated by the phylogenetic analysis of mtDNA COI sequences. Additionally, 5 haplotypes, each from a different location, found in Atyaephyraorientalis were not shared by any other species of the genus. It also differs from all the other species in the following nucleotide positions in the COI gene of Atyaephyra desmarestii specimen Dour1, position 273: guanine (G), position 276: guanine (G) and position 369: cytosine (C).
Atyaephyra orientalis is found in freshwater habitats of Middle East, from Turkey to Iraq (see material examined and Fig. 1).
Bouvier (1913) after examining the Atyaephyra material deposited in the MNHN collections he assigned it into two varieties (Atyaephyra desmarestii var. orientalis and Atyaephyra desmarestii occidentalis) based mainly on differences observed in the endopod of first male pleopod. Atyaephyra desmarestii var. orientalis was originally described from Syria (from Orontes River, near the Lake Qattinah (Lake Homs), from a stream in Kousseir (probably Qoussair) near Damascus and from Barada River, Ataibe, East of Damascus) and was elevated to subspecies level by Holthuis (1961). Apart from Atyaephyra desmarestii orientalis, a second subspecies, Atyaephyra desmarestii mesopotamica, was found to exist in the Middle East and was described by Al-Adhub (1987). Al-Adhub (1987) described the new subspecies based on the presence of a distinct subterminal process (vs. absent from Atyaephyra desmarestii orientalis and Atyaephyra desmarestii desmarestii) and the presence of 50 spines on dactylus of fifth pereiopod (vs. 40 in Atyaephyra desmarestii orientalis and Atyaephyra desmarestii desmarestii). Furthermore he noticed that the rostrum of Atyaephyra desmarestii mesopotamica resembles that of Atyaephyra desmarestii desmarestii from Greece but differs in having the distal ventral part always devoid of teeth. Indeed the individuals from Shatt Al-Arab River had the highest number of spines on dactylus of fifth pereiopod ranging from 41–55 but specimens from the River Orontes were also found with up to 47 spines (33–47). Additionally, male individuals having endopod with a distinct subterminal process were found again in River Orontes as well as in other Middle East Rivers. Gorgin (1996), after studying 150 males from two different localities in Iran found individuals with a distinct subterminal process and without inside the same population. Finally, specimens from Greece belonging to Atyaephyra stankoi (as the sample of Holthuis to which Al-Adhub refers to) were found to be also devoid of teeth in the distal part of the rostrum. Even in the illustration included in Holthuis (1961) work, the Greek specimen is devoid of teeth in the distal part of the ventral margin. Although the genetic distances within the Atyaephyra orientalis phylogroup were high (0.9%–10.2%) no firm conclusion could be drawn whether the hypothesis of multiple species is valid or not. Sequences from Orontes River (topotypical location of Atyaephyra desmarestii orientalis) and from Shatt Al-Arab River (topotypical location of Atyaephyra desmarestii mesopotamica) presented a noticeable mean genetic divergence (5.0%) but still not strong enough to support the hypothesis of different species. Detailed future studies on the morphological and genetic variability within the Atyaephyra distributed throughout the Middle East will help clarify the relationships between the populations in this region. However, only one species is currently considered to exist, Atyaephyra orientalis. Therefore, Atyaephyra desmarestii mesopotamica is here proposed as a synonym.
Atyaephyra orientalis appears to be morphologically more similar to Atyaephyra stankoi and Atyaephyra thyamisensis sp. n. by sharing characters such as the presence of numerous mesial spines (10–38) on terminal segment of third maxilliped (Figs 4H, 6H, 8H). It also shares in common with the other two species the presence of fewer rows of setae (12–16) on basal lower endite of maxilla, the endite of maxilla being 1.75–2.24 × as long as basal lower endite (Figs 4G, 6G, 8G) and basal endite of first maxilliped failing or reaching to distal end of exopod distal margin (Figs 4F, 6F, 8F). Atyaephyra orientalis can be separated from Atyaephyra thyamisensis sp. n. and Atyaephyra stankoi by the presence of a pointed antennular lobe (Figs 3H–I) (vs. round in Atyaephyra stankoi and Atyaephyra thyamisensis sp. n. Figs 5H, 7H). Further, Atyaephyra orientalis can be distinguished by the strongly curved and distally stout and not tapering endopod of male first pleopod (Fig. 4I) (vs. slightly curved and distally more or less elongated but always tapering in Atyaephyra stankoi, Fig. 6I; slightly or strongly curved but always its distal part is elongated and tapering (ribbon shaped) in Atyaephyra thyamisensis sp. n., Fig. 8I). Atyaephyra orientalis differs from the other four species of Atyaephyra in having 10–36 spines on terminal segment of third maxilliped (Fig. 4H) (vs. 0–8 in Atyaephyra desmarestii, Atyaephyra strymonensis sp. n., Atyaephyra acheronensis sp. n. and Atyaephyra tuerkayi sp. n. Figs 10H, 12H, 14H).
- Christodoulou, M; Antoniou, A; Antonios Magoulas, ; Athanasios Koukouras, ; 2012: Revision of the freshwater genus Atyaephyra (Crustacea, Decapoda, Atyidae) based on morphological and molecular data ZooKeys, 229: 53-110. doi
- Barrois T (1893) Liste des décapodes fluviatiles recueillis en Syrie suivie de quelques considérations sur le genre Caridine. Revue Biologique du Nord de la France 5 (4): 125–134. http://ia700304.us.archive.org/22/items/revuebiologiqued05lill/revuebiologiqued05lill.pdf
- Annandale N, Kemp S (1913) The Crustacea Decapoda of the Lake of Tiberias. Journal and Proceedings of the Asiatic Society of Bengal, New Series 9 (6): 241–258. http://decapoda.nhm.org/pdfs/24601/24601.pdf
- Bouvier E (1925) Recherches sur la morphologie, les variations et la distribution systématique des crevettes d’eau douce de la famille des Atyidés. Encyclopédie Entomologique 4: 1-365.
- Holthuis L (1961) Report on a collection of Crustacea Decapoda and Stomatopoda from Turkey and the Balkans. Zoologische Verhandelingen 47: 1–67. http://www.repository.naturalis.nl/document/148913
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- Salman S (1987) Larval development of Atyaephyra desmaresti mesopotamica Al-Adhub (Decapoda, Atyidae) reared in the laboratory. Investigacion Pesquera 51 (1): 27-42. doi: 10.1163/156854087X00484
- Gorgin S (1996) The first record of two species of freshwater shrimps (Decapoda, Caridea, Atyidae) from Iran. Crustaceana 69 (5): 662–668. http://www.jstor.org/discover/10.2307/20105244?uid=2129&uid=2134&uid=2&uid=70&uid=4&sid=21101302264147 , doi: 10.1163/156854096X00664
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- Von Rintelen K, Page T, Cai Y, Roe K, Stelbrink B, Kuhajda B, Iliffe T, Hughes J, von Rintelen T (2012) Drawn to the dark side: a molecular phylogeny of freshwater shrimps (Crustacea, Decapoda, Caridea, Atyidae) reveals frequent cave invasions and challenges current taxonomic hypotheses. Molecular Phylogenetics and Evolution 63 (1): 82-96. doi: 10.1016/j.ympev.2011.12.015
- Bouvier E (1913) Les variations d’une crevette de la famille des Atyidées, l’Atyaephyra Desmaresti Millet. Bulletin du Muséum National d’Histoire Naturelle 19 (2): 65–74. http://www.biodiversitylibrary.org/item/27226
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