Research Article |
Corresponding author: Asmaa Haris Elgetany ( asmaa_haris222@yahoo.com ) Academic editor: Greg Rouse
© 2022 Asmaa Haris Elgetany, Torsten H. Struck, Christopher J. Glasby.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Elgetany AH, Struck TH, Glasby CJ (2022) Three new species of the genus Perinereis (Annelida, Nereididae) from Egyptian coasts. ZooKeys 1132: 163-188. https://doi.org/10.3897/zookeys.1132.87629
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Despite being one of the most common groups of polychaetes on intertidal shores, the genus Perinereis (Nereididae) is comparatively poorly known taxonomically, with confusion still existing due to the lack of comprehensive systematic studies. The systematics of Perinereis species from the intertidal Egyptian coasts of the Red Sea, Gulf of Suez and Suez Canal have been investigated using morphology and the mitochondrial barcoding marker cytochrome oxidase subunit I (COI). New sequence data was obtained for 102 Perinereis specimens and analysis included all publicly available COI data from other Perinereis species. The COI data indicate that monophyly of the P. nuntia species group is doubtful, as specimens identified in this species group from south-eastern Asia and Australia form a monophyletic group exclusive of the three new species described in this study from the Red Sea region. A morphometric character set (26 characters) was used to identify and characterize each specimen in the study. Three distinct morphospecies belonging to the P. nuntia species group were found, each differentiated by the number and type of paragnaths on pharyngeal areas V and VI, relative sizes of parapodial lobes, type of notochaetae and neurochaetae, and form of the neurochaetal falciger blades. The three morphospecies were well supported by COI data: two of the three new species, Perinereis suezensis sp. nov. and Perinereis fayedensis sp. nov., are closely similar to P. nuntia sensu stricto, while the other, Perinereis damietta sp. nov., is similar to P. heterodonta. The new species are described and illustrated, and bring the number of species in Perinereis to 97. The new species are compared and contrasted to the closely similar P. heterodonta, P. nuntia and other congeners from the region.
Integrative taxonomy, morphometrics, Perinereis damietta sp. nov., Perinereis fayedensis sp. nov., Perinereis nuntia species complex, Perinereis suezensis sp. nov., systematics
The family Nereididae includes several highly variable species characterized by high intra-specific morphological variation especially associated with the pharynx (e.g., number and arrangement of paragnaths) or associated with the parapodia (relative proportions of lobes/ligules and form of chaetae) and coloration. Often these morphologically variable species also show differences in reproductive biology (
Perinereis Kinberg, 1865 is the second most species-rich genus in the family. It includes approximately 94 worldwide-distributed valid species (
Members of Perinereis have been long recognized based primarily on the number and type of paragnaths on areas V and VI (e.g.,
Until recently, the Perinereis nuntia species group comprised 15 valid species (
The present study investigates the taxonomy of three putative species belonging to the P. nuntia species group sampled from the Gulf of Suez, Suez Canal, and the northern Red Sea using a detailed morphological study and the mitochondrial barcoding marker cytochrome oxidase subunit I (COI). We compare our material with other members of the species group originally described from the region, including Perinereis nuntia (type locality: Gulf of Suez) and P. heterodonta Gravier, 1899 (type locality: Red Sea, Obock, Gulf of Aden, Djibouti,). Our results show that all three species are new to science: two of them, P. suezensis sp. nov. and P. fayedensis sp. nov. from Gulf of Suez (part of Red Sea), are closely similar to P. nuntia, while the other one, P. damietta sp. nov., from Hurghada (northern Red Sea), is more similar to P. heterodonta. The three species are described below.
Sampling was carried out during the period of January 2015 to July 2017 from four localities along the intertidal zone of Egyptian coasts of the Red Sea, Gulf of Suez and Suez Canal (Fig.
Map of Perinereis species localities referred to in this study. Colored squares indicate different species: red – Perinereis heterodonta, green – P. nuntia, yellow – P. damietta sp. nov., P. suezensis sp. nov. and P. fayedensis sp. nov. (see Table
A section of the mid body was removed from the Red Sea specimens and stored in 96% ethanol for molecular analysis. The rest of the animal was fixed in 4% seawater formalin and stored in 70% ethanol for morphological studies.
Specimens were studied and photographed using a stereo microscope, Leica MZ16, with a Planapo 1.0X and Canon EOS 500D, as well as a compound microscope, Leica DFC420 connected to a Leica Computer CTR600 and a DM600B camera. For morphological characterization, we grouped specimens into three putative species, and recorded 26 morphometric characters for a subset of 45 of the 102 specimens in total (Suppl. material
Species used in this study with Sample ID for the new specimens and accession numbers for all specimens.
Genus | Species | Sample ID | Accession # | Genus | Species | Accession # |
---|---|---|---|---|---|---|
Perinereis | damietta sp. nov. | 5-1 | OP610122 | Perinereis | aibuhitensis | KC800611 |
Perinereis | damietta sp. nov. | 5-2 | OP610123 | Perinereis | aibuhitensis | KC800612 |
Perinereis | damietta sp. nov. | 5-8 | OP610124 | Perinereis | aibuhitensis | KC800613 |
Perinereis | damietta sp. nov. | 5-10 | OP610125 | Perinereis | aibuhitensis | KC800614 |
Perinereis | damietta sp. nov. | 12-1B | OP610126 | Perinereis | aibuhitensis | KC840698 |
Perinereis | damietta sp. nov. | 12-2B | OP610127 | Perinereis | aibuhitensis | KF611806 |
Perinereis | damietta sp. nov. | 12-3B | OP610128 | Perinereis | aibuhitensis | KY129885 |
Perinereis | damietta sp. nov. | 12-4B | OP610129 | Perinereis | aibuhitensis | MN256534 |
Perinereis | damietta sp. nov. | 12-5B | OP610130 | Perinereis | aibuhitensis | MN256535 |
Perinereis | damietta sp. nov. | 12-6B | OP610131 | Perinereis | aibuhitensis | MN256536 |
Perinereis | damietta sp. nov. | 12-8B | OP610132 | Perinereis | aibuhitensis | MT511716 |
Perinereis | damietta sp. nov. | 12-10B | OP610133 | Perinereis | aibuhitensis | MT511717 |
Perinereis | damietta sp. nov. | 16-1 | OP610134 | Perinereis | aibuhitensis | MT511718 |
Perinereis | damietta sp. nov. | 16-2 | OP610135 | Perinereis | aibuhitensis | MT712474 |
Perinereis | damietta sp. nov. | 16-3 | OP610136 | Perinereis | aibuhitensis | MW593148 |
Perinereis | damietta sp. nov. | 16-4 | OP610137 | Perinereis | anderssoni | MH143495 |
Perinereis | damietta sp. nov. | R4-1 | OP610138 | Perinereis | anderssoni | MH143497 |
Perinereis | damietta sp. nov. | R4-2 | OP610139 | Perinereis | anderssoni | MH143498 |
Perinereis | damietta sp. nov. | R4-3 | OP610140 | Perinereis | anderssoni | MH143502 |
Perinereis | damietta sp. nov. | R6-1 | OP610141 | Perinereis | anderssoni | MH143503 |
Perinereis | damietta sp. nov. | R6-2 | OP610142 | Perinereis | anderssoni | MH143504 |
Perinereis | damietta sp. nov. | R6-3 | OP610143 | Perinereis | anderssoni | MH143507 |
Perinereis | damietta sp. nov. | R8-1 | OP610144 | Perinereis | anderssoni | MH143508 |
Perinereis | damietta sp. nov. | R8-2 | OP610145 | Perinereis | anderssoni | MH143514 |
Perinereis | damietta sp. nov. | R8-3 | OP610146 | Perinereis | anderssoni | MH143516 |
Perinereis | damietta sp. nov. | R8-5 | OP610147 | Perinereis | anderssoni | MH143520 |
Perinereis | damietta sp. nov. | R8-6 | OP610148 | Perinereis | anderssoni | MH143522 |
Perinereis | damietta sp. nov. | R8-7 | OP610149 | Perinereis | brevicirris | JX503024 |
Perinereis | damietta sp. nov. | R8-8 | OP610150 | Perinereis | brevicirris | JX966314 |
Perinereis | suezensis sp. nov. | 1-9 | OP612948 | Perinereis | brevicirris | KC800628 |
Perinereis | suezensis sp. nov. | 2-3 | OP612949 | Perinereis | brevicirris | KC800630 |
Perinereis | suezensis sp. nov. | 2-4 | OP612950 | Perinereis | brevicirris | KC800632 |
Perinereis | suezensis sp. nov. | 2-5 | OP612951 | Perinereis | camiguinoides | KF850496 |
Perinereis | suezensis sp. nov. | 2-6 | OP612952 | Perinereis | cultrifera | KC800624 |
Perinereis | suezensis sp. nov. | 2-7 | OP612953 | Perinereis | cultrifera | KC800625 |
Perinereis | suezensis sp. nov. | 2-8 | OP612954 | Perinereis | cultrifera | KC800627 |
Perinereis | suezensis sp. nov. | 5-4 | OP612955 | Perinereis | cultrifera | KR916906 |
Perinereis | suezensis sp. nov. | 5-5 | OP612956 | Perinereis | cultrifera | KR916907 |
Perinereis | suezensis sp. nov. | 5-6 | OP612957 | Perinereis | cultrifera | KR916908 |
Perinereis | suezensis sp. nov. | 5-7 | OP612958 | Perinereis | cultrifera | KR916909 |
Perinereis | suezensis sp. nov. | 5-9 | OP612959 | Perinereis | cultrifera | KR916910 |
Perinereis | suezensis sp. nov. | 7-5B | OP612960 | Perinereis | cultrifera | KR916911 |
Perinereis | suezensis sp. nov. | 7-7B | OP612961 | Perinereis | cultrifera | KR916912 |
Perinereis | suezensis sp. nov. | 7-8B | OP612962 | Perinereis | cultrifera | KY129882 |
Perinereis | suezensis sp. nov. | 7-9B | OP612963 | Perinereis | cultrifera | KY129883 |
Perinereis | suezensis sp. nov. | 7-10B | OP612964 | Perinereis | cultrifera | MN256544 |
Perinereis | suezensis sp. nov. | 8-1 | OP612965 | Perinereis | cultrifera | MN256545 |
Perinereis | suezensis sp. nov. | 8-2 | OP612966 | Perinereis | cultrifera | NC_051994 |
Perinereis | suezensis sp. nov. | 8-3 | OP612967 | Perinereis | curvata | MW277905 |
Perinereis | suezensis sp. nov. | 8-4 | OP612968 | Perinereis | euiini | KY249122 |
Perinereis | suezensis sp. nov. | 8-5 | OP612969 | Perinereis | euiini | KY249123 |
Perinereis | suezensis sp. nov. | 8-6 | OP612970 | Perinereis | euiini | KY249124 |
Perinereis | suezensis sp. nov. | 8-7 | OP612971 | Perinereis | falklandica | HQ705184 |
Perinereis | suezensis sp. nov. | 8-8 | OP612972 | Perinereis | falklandica | HQ705185 |
Perinereis | suezensis sp. nov. | 8-9 | OP612973 | Perinereis | gualpensis | HQ705186 |
Perinereis | suezensis sp. nov. | 8-10 | OP612974 | Perinereis | gualpensis | HQ705187 |
Perinereis | suezensis sp. nov. | 10-3B | OP612975 | Perinereis | gualpensis | HQ705188 |
Perinereis | suezensis sp. nov. | 10-4B | OP612976 | Perinereis | helleri | JX420256 |
Perinereis | suezensis sp. nov. | 10-5B | OP612977 | Perinereis | linea | MT511711 |
Perinereis | suezensis sp. nov. | 10-6B | OP612978 | Perinereis | linea | MT511712 |
Perinereis | suezensis sp. nov. | 10-7B | OP612979 | Perinereis | linea | MT511713 |
Perinereis | suezensis sp. nov. | 10-8B | OP612980 | Perinereis | linea | MT511714 |
Perinereis | suezensis sp. nov. | 10-10B | OP612981 | Perinereis | linea | MT511715 |
Perinereis | suezensis sp. nov. | 11-1B | OP612982 | Perinereis | longidonta | HQ705190 |
Perinereis | suezensis sp. nov. | 11-3B | OP612983 | Perinereis | longidonta | HQ705191 |
Perinereis | suezensis sp. nov. | 11-4B | OP612984 | Perinereis | nuntia | JX420257 |
Perinereis | suezensis sp. nov. | 11-5B | OP612985 | Perinereis | nuntia | JX644015 |
Perinereis | suezensis sp. nov. | 12-9B | OP612986 | Perinereis | nuntia | MH337359 |
Perinereis | suezensis sp. nov. | 14-1 | OP612987 | Perinereis | seridentata | JF293314 |
Perinereis | suezensis sp. nov. | 14-2 | OP612988 | Perinereis | singaporiensis | EU835665 |
Perinereis | suezensis sp. nov. | 14-3 | OP612989 | Perinereis | sp. | EU352319 |
Perinereis | suezensis sp. nov. | 14-4 | OP612990 | Perinereis | sp. | KR916903 |
Perinereis | suezensis sp. nov. | 14-5 | OP612991 | Perinereis | sp. | KR916904 |
Perinereis | suezensis sp. nov. | 14-6 | OP612992 | Perinereis | sp. | KR916905 |
Perinereis | suezensis sp. nov. | 14-7 | OP612993 | Perinereis | sp. | KX525487 |
Perinereis | suezensis sp. nov. | 14-8 | OP612994 | Perinereis | sp. | KX525497 |
Perinereis | suezensis sp. nov. | 14-9 | OP612995 | Perinereis | sp. | KX525498 |
Perinereis | suezensis sp. nov. | 14-10 | OP612996 | Perinereis | sp. | KX525499 |
Perinereis | suezensis sp. nov. | 18-1 | OP612997 | Perinereis | sp. | KX840014 |
Perinereis | suezensis sp. nov. | 18-2 | OP612998 | Perinereis | sp. | MH143496 |
Perinereis | suezensis sp. nov. | 18-3 | OP612999 | Perinereis | sp. | MH143499 |
Perinereis | suezensis sp. nov. | 18-4 | OP613000 | Perinereis | sp. | MH143500 |
Perinereis | suezensis sp. nov. | 18-5 | OP613001 | Perinereis | sp. | MH143501 |
Perinereis | suezensis sp. nov. | 18-6 | OP613002 | Perinereis | sp. | MH143505 |
Perinereis | suezensis sp. nov. | 18-7 | OP613003 | Perinereis | sp. | MH143506 |
Perinereis | suezensis sp. nov. | 18-8 | OP613004 | Perinereis | sp. | MH143509 |
Perinereis | suezensis sp. nov. | 18-9 | OP613005 | Perinereis | sp. | MH143510 |
Perinereis | suezensis sp. nov. | 18-10 | OP613006 | Perinereis | sp. | MH143511 |
Perinereis | suezensis sp. nov. | R5-1 | OP613007 | Perinereis | sp. | MH143512 |
Perinereis | suezensis sp. nov. | R5-2 | OP613008 | Perinereis | sp. | MH143513 |
Perinereis | suezensis sp. nov. | R5-3 | OP613009 | Perinereis | sp. | MH143515 |
Perinereis | suezensis sp. nov. | R5-4 | OP613010 | Perinereis | sp. | MH143517 |
Perinereis | suezensis sp. nov. | R6-4 | OP613011 | Perinereis | sp. | MH143518 |
Perinereis | fayedensis sp. nov. | 2-2 | OP605755 | Perinereis | sp. | MH143519 |
Perinereis | fayedensis sp. nov. | 5-3 | OP605756 | Perinereis | sp. | MH143521 |
Perinereis | fayedensis sp. nov. | 7-4B | OP605757 | Perinereis | sp. | MH143523 |
Perinereis | fayedensis sp. nov. | 7-6B | OP605758 | Perinereis | sp. | MH143524 |
Perinereis | fayedensis sp. nov. | 10-1B | OP605759 | Perinereis | sp. | MH143525 |
Perinereis | fayedensis sp. nov. | 10-2B | OP605760 | Perinereis | sp. | MH143526 |
Perinereis | fayedensis sp. nov. | 10-9B | OP605761 | Perinereis | sp. | MN823962 |
Perinereis | fayedensis sp. nov. | 11-2B | OP605762 | Perinereis | sp. | MT528267 |
Perinereis | fayedensis sp. nov. | 12-7B | OP605763 | Perinereis | sp. | OK430976 |
Perinereis | aibuhitensis | GU362686 | Perinereis | suluana | JX392072 | |
Perinereis | aibuhitensis | JX503021 | Perinereis | suluana | JX420245 | |
Perinereis | aibuhitensis | JX503022 | Perinereis | suluana | JX420246 | |
Perinereis | aibuhitensis | JX503023 | Perinereis | suluana | JX420247 | |
Perinereis | aibuhitensis | JX661442 | Perinereis | suluana | JX420248 | |
Perinereis | aibuhitensis | JX661443 | Perinereis | suluana | JX420250 | |
Perinereis | aibuhitensis | JX661444 | Perinereis | suluana | JX420251 | |
Perinereis | aibuhitensis | JX661445 | Perinereis | suluana | JX420252 | |
Perinereis | aibuhitensis | JX661446 | Perinereis | suluana | JX420253 | |
Perinereis | aibuhitensis | JX661447 | Perinereis | suluana | JX420254 | |
Perinereis | aibuhitensis | JX661448 | Perinereis | suluana | JX420255 | |
Perinereis | aibuhitensis | JX661449 | Perinereis | vallata | HQ705192 | |
Perinereis | aibuhitensis | JX661450 | Perinereis | vallata | HQ705196 | |
Perinereis | aibuhitensis | JX661451 | Perinereis | vallata | JX676119 | |
Perinereis | aibuhitensis | JX661452 | Perinereis | vallata | JX676143 | |
Perinereis | aibuhitensis | JX661453 | Perinereis | vallata | MT511721 | |
Perinereis | aibuhitensis | JX661454 | Perinereis | vallata | MT511722 | |
Perinereis | aibuhitensis | JX661455 | Perinereis | vancaurica | MT511719 | |
Perinereis | aibuhitensis | JX661456 | Perinereis | wilsoni | KC800623 | |
Perinereis | aibuhitensis | JX661457 | Perinereis | wilsoni | KC800629 | |
Perinereis | aibuhitensis | JX661458 | Perinereis | wilsoni | KC800631 | |
Perinereis | aibuhitensis | JX661459 | Perinereis | wilsoni | KY129887 | |
Perinereis | aibuhitensis | JX661460 | Perinereis | wilsoni | KY129888 | |
Perinereis | aibuhitensis | JX661461 | Perinereis | wilsoni | KY129889 | |
Perinereis | aibuhitensis | JX661462 | Perinereis | wilsoni | MN256541 | |
Perinereis | aibuhitensis | JX661463 | Perinereis | wilsoni | MN256542 | |
Perinereis | aibuhitensis | JX661464 | Perinereis | wilsoni | MN256543 | |
Perinereis | aibuhitensis | JX661465 | Dendronereis | chipolini | MW5320841 | |
Perinereis | aibuhitensis | JX661466 | Hediste | japonica | MN876864 | |
Perinereis | aibuhitensis | JX661467 | Namalycastis | abiuma | KU351089 | |
Perinereis | aibuhitensis | JX661468 | Platynereis | dumerilii | AF178678 | |
Perinereis | aibuhitensis | JX661469 |
Samples are deposited in the Damietta University - Faculty of Science (DUFS), Damietta, Egypt, and the Senckenberg Forschungsinstitut und Naturmuseum (
Genomic DNA was extracted from three to four segments of the middle section of each worm using the DNeasy Tissue Kit (Qiagen) according to manufacturers’ instructions with at least two elution steps to increase the amount of DNA. For each individual, the nucleotide sequences of the mitochondrial COI were amplified using the primer pair LCO1490JJ (forward, 5’-CHA CWA AYC ATA AAG ATA RYG G-3’) and HCO2198JJ (reverse, 5’-AWA CTT CVG GRT GVC CAA ARA ATC A-3’) (
For the phylogenetic analyses, we included all publicly available COI data from other specimens of Perinereis as well as five nereidid species, who have a complete mitochondrial genome sequenced, as outgroup taxon (Table
The ML tree (logL = -11297.8710) showed that the genus Perinereis is probably not monophyletic as the outgroup Hediste japonica Izuka, 1908 grouped within the genus (Fig.
ML tree (logL = -11297.8710) of the Perinereis species in this analysis shown in three parts (A, B, C). The points, where we cut the branches, are indicated by letters. The new species P. suezensis sp. nov., P. fayedensis sp. nov., and P. damietta sp. nov. are highlighted by boxes and in bold. Perineris nuntia is also highlighted in bold. Bootstrap values equal to and more than 95 indicating strong support are given at the branches. The three relevant bootstrap values are highlighted in bold. The scale bar shows substitutions/position.
Nereididae Blainville, 1818
Nereidinae Blainville, 1818
Perinereis Kinberg, 1856
Perinereis nuntia
species complex Wilson & Glasby, 1993: 259. –
Perinereis species having conical paragnaths on all areas (rarely absent on area V), except area VI with long bars, which can be shield-shaped or pyramidal paragnaths, arranged in a single-arched row; area V with paragnaths displaced posteriorly (on everted proboscis) to those on area VI; area IV rarely with merged paragnaths.
We have restricted the diagnosis of the species complex to include only unique diagnostic features. Some new characters introduced by
Holotype : DUFS 067 Al-Adabiya; west of Port Taofik, Gulf of Suez (Red Sea), intertidal, under coarse sands, at 29°56'06.0"N, 32°28'36.6"E, collection date (15.01.2015) Paratypes: 13 specimens (DUFS 057-066, 068-070) from Al-Qantara, Suez Canal, intertidal, muddy sand bottom, at 30°50'31.5"N, 32°18'54.8"E, Fayed, western shore of Great Bitter Lake, intertidal, silty mud bottom at 30°20'18.0"N, 32°18'14.9"E, and Al-Adabiya (same collection details as holotype). Collection dates (18.02.2015/ 15.01.2015/ 01.07.2017).
2 specimens (
Holotype (DUFS 067) not complete, 53 chaetigers, 50 mm in length, 2 mm wide at chaetiger 10 Paratypes with 33–88 chaetigers, 32–81 mm long, 2.0–4.5 mm wide at chaetiger 10. Epidermis with orange pigmentation on anterior dorsum in some preserved paratypes.
Prostomium with entire anterior margin; as wide as long. Antennae closely set, as long as ~ 1/3 length of prostomium. Eyes black, anterior pair set slightly further apart than posterior pair; lenses not obvious.
Apodous segment ~ 1.2× or 1.6× longer than chaetiger 1. Posterodorsal tentacular cirri extending back to chaetiger 6 (6–7).
Pharynx with jaws translucent, red-brown, with 7 (7–8) teeth. Paragnaths black. Area I with 2 (1–5) conical paragnaths; area II conical paragnaths with 5 (5–10) on left and 8 (7–10) on right, in a triangular patch; area III with 10 (9–17) conical paragnaths in 2–3 rows, with two laterally isolated paragnaths; area IV conical paragnaths with 16 (11–19) on left, 13 (9–16) on right, in two or three rows, in elongated triangle; area V with 4 (2–4) conical paragnaths interspersed with one or two bars, set well proximal (on everted proboscis) to line of area VI paragnaths; area VI with 15 (14–21), shield-shaped bars with pointed tips (very close in appearance to cones), arranged in one arc, with the right and left rows almost touching; area VII-VIII with 44 (37–44) conical paragnaths arranged in a single band of two rows laterally to three or four rows deep medially (Fig.
Perinereis suezensis sp. nov. All pictures are from the holotype if not stated otherwise A anterior end, maxillary apparatus, dorsal view B anterior end, maxillary apparatus, ventral view C posterior end, ventral view D right parapodium, posterior view, chaetiger 7 E right parapodium, anterior view, chaetiger 33 F right parapodium, anterior view, chaetiger 64 G chaetal bundle of a right parapodium homogomph spiniger, heterogomph spiniger & heterogomph falciger, chaetiger 7 H neuropodial sub-acicular heterogomph falciger, chaetiger 7 I heterogomph falciger, chaetiger 7.
Comparison of key characters between forms resembling P. heterodonta (pale grey) and Perinereis nuntia (dark grey) in the Red Sea, the Gulf of Aden, and the Arabian (= Persian) Gulf. Abbreviations: AIII = Area III; AV = Area V; AVI Area VI; AVII-VIII = Areas VII and VIII; p-dTC = posterior extension of posterodorsal tentacular cirri (chaetiger); ratio of lengths of dorsal cirri v dorsal notopodial lobe in posterior chaetigers; HS, presence (p) or absence (a) of heterogomph spinigers in anterior neuropodia; NA = data not available.
Species | Type locality | AIII (lateral group p/a) | AV | AVI | AVII-VIII | p-dTC | p-DC:DNL | HS | Reference |
---|---|---|---|---|---|---|---|---|---|
heterodonta sensu stricto | Djibouti | 6-7 irregular cluster | 0 | 10–16 | 18 | 5 | NA | a |
|
heterodonta sensu stricto | Gulf of Oman | 8-14 (p) cluster | 0–1 | 14–24 | 20–35 | 1–6 | 0.6–1× | a |
|
Perinereis damietta sp. nov. | Gulf of Suez | 3-9 (one transverse row) | 0–1 | 24–40 | 16–32 | 2–4 | 1.1–1.2 | a | Present paper |
djiboutiensis | Djibouti | small rectangular patch (p) | 0 | 6–7 | NA (3 rows) | 10–15 | NA (DC > DNL) | NA |
|
nuntia sensu stricto | Gulf of Suez, Red Sea | 15 | 3(2–4) | 10–12 (8–10) | 41 (36–50) | (4–6) | 4–5X (3–4X) | p | Savigny in Lamarck (1818); |
nuntia sensu |
‘Red Sea’ | 8-14 (p) | 3–4 | 8–13 | 24–31 | 6–14 | ~1.2 | p |
|
nuntia sensu |
Gulf of Oman | 9-15 | 3 | 13–20 | 36–58 | 8–14 | 2X | p |
|
Perinereis suezensis sp. nov. | Gulf of Suez, Red Sea | 9-17 (p) | 2–4 | 14–21 | 37–44 | 6–7 | 1.0–1.3X | p | Present paper |
Perinereis fayedensis sp. nov. | Gulf of Suez, Red Sea | 2-5 (one tranverse row) | 1–4 | 14–17 | 28–40 | 6–8 | 1.1–1.2X | p | Present paper |
Anterior notopodia with conical dorsal and median ligules of equal length in anterior body; dorsal ligule slightly longer in mid- and posterior body. Superior lobes absent. DC length 1.1 (1.0–1.2) × DNL length anteriorly (chaetigers 10–20); posteriorly DC length 1.09 (1.0–1.3) × length of DNL length (chaetigers 75–90). DC and DNL of mid-body parapodia proportionally similar to those of posterior parapodia (Fig.
Dorsal notopodial ligule larger than ventral notopodial ligule anteriorly and posteriorly. Neuropodia with inferior and postchaetal lobes, ventral ligule and ventral cirri. Neuropodial postchaetal lobe lowly rounded, not projecting beyond end of acicular ligule. Ventral neuropodial ligule subconical, ca. as long as median ligule throughout. Ventral cirri extending laterally to reach tip of ventral neuropodial ligule anteriorly, extending to ~ 1/4 length of ventral neuropodial ligule posteriorly (Fig.
Notochaetae with homogomph spinigers throughout, blades long; teeth short. Neurochaetae in upper fascicle with homogomph spinigers with long blades; one heterogomph falciger with short blades throughout, blades serrated. Neurochaetae in lower fascicle with heterogomph falcigers, blades short and thick, teeth long; and two or three heterogomph spinigers, median long blades, teeth short present throughout body. Aciculae black, single in each ramus (Fig.
Pygidium with anal cirri extending to last 6 (6–7) chaetigers, 5 (5–7) mm long, whitish cream without any pigmentation (Fig.
Two specimens: one complete with 105 chaetigers, 57 mm long and 2.8 mm wide, and another with regenerating tail, 107 chaetigers, 71 mm long and 4.3 mm wide. Apodous segment ~ 1.3–1.8× longer than chaetiger 1. Posterodorsal tentacular cirri extending back to chaetigers 5 and 6. Jaws with 4–7 teeth. Paragnaths count: area I with 2; area II with 8–17 on left and 9–17 on right; area III with 11 or 12 in two or three rows; area IV with 17 or 18 on both sides, in two or three rows; area V with three or four; area VI with 8–12 on left and 8–11 on right, shield-shaped bars with pointed tips and cones arranged in one row with the right and left side rows almost touching each other; area VII-VIII with 47 or 48, arranged in a single band of two rows laterally to three or four rows deep medially. Dorsal cirrus length ~ 0.8× length of dorsal notopodial ligule anteriorly and 0.7–0.9× length of dorsal notopodial ligule posteriorly. Ventral cirri extending laterally to reach tip or half-length of ventral neuropodial ligule anteriorly. Neurochaetae in upper fascicle with 1–3 heterogomph falcigers. Neurochaetae in lower fascicle with 1–4 heterogomph spinigers, rarely absent.
Gulf of Suez, Suez Canal including Great Bitter Lake, northern Red Sea; intertidal sand and mud, under stones.
The new species is named after the port city of Suez (Egyptian Arabic pronunciation: (السويس) located on the north coast of the Gulf of Suez.
The molecular data place P. suezensis sp. nov. clearly apart from all other species and the monophyly of the species is very well supported by a bootstrap value of 99 (Fig.
In addition to our sequences, only three additional COI sequences for P. nuntia have been published: JX420257 (Indonesia), JX644015 (South Korea), and MH337359 (Andaman and Nicobar Islands). JX420257 and MH337359 are identical (bootstrap value of 100; Fig.
The new species is most similar to P. nuntia, which was also described from the Gulf of Suez. Although the exact location of Savigny’s specimens has never been established, it is very likely to be from shallow waters of the port city of Suez, as for Savigny’s other polychaetes (see
The larger-sized, non-type specimens generally had more paragnaths in each area compared to the type material, except for area VI. The fewer paragnaths in area VI in the non-type specimens is most likely due to loss, as the ones present were irregularly spaced, with some gaps large enough to accommodate a lost shield-shaped bar or two cones. Another reflection on the condition of the non-type specimens is the unusually short dorsal and ventral cirri; on this point, the cirri appeared withered and many were missing, which we attribute to damage or a fixation artifact.
Holotype : DUFS 0123 Al-Adabiya, west of Port Taofik, Gulf of Suez (Red Sea), intertidal, under coarse sands, at 29°56'06.0"N, 32°28'36.6"E. Paratypes (DUFS 120–122, 124–128): 8 specimens from El-Qantara, Suez Canal, intertidal, muddy sand bottom, at 30°50'31.5"N, 32°18'54.8"E, Fayed, western shore of Great Bitter Lake, intertidal, silty mud bottom, at 30°20'18.0"N, 32°18'14.9"E, Al-Adabiya (same collection details as holotype).
Holotype (DUFS 0123) not complete, 49 chaetigers, 35 mm in length, 3 mm wide at chaetiger 10. Paratypes with 37–88 chaetigers, 30–70 mm long, 1.5–4.5 mm wide at chaetiger 10. Epidermis whitish cream with a longitudinal beige pigmentation stripe on ventral side of posterior chaetigers in some preserved.
Prostomium with entire anterior margin; wide as long. Antennae closely set, as long as ~ 1/3 length of prostomium. Eyes black, anterior pair set slightly further apart than posterior pair; lenses not obvious.
Apodous segment ~ 1.5× longer than chaetiger 1. Posterodorsal tentacular cirri with distinct cirrophores, extend back to chaetiger 7 (6–8).
Pharynx with jaws translucent red-brown, with 8 (7–8) teeth. Maxillary ring of pharynx with paragnaths, arranged in discrete areas, areas II-IV arranged in regular comb-like rows. Area I with 2 (1–2) conical paragnaths in vertical arrangement; area II with 9 (7–10) in left and 9 (7–10) in right conical paragnaths, three or four rows in a triangular patch; area III with 2 (2–5) conical paragnaths in vertical arrangement; area IV with 13 (12–15) in left, 14 (12–15) in right, conical paragnaths without bars; area V with 4 (1–3) conical paragnaths; area VI with 17 (14–17), shield-shaped bars with pointed tip present, cones paragnaths absent; area VII-VIII with 38 (28–40) conical paragnaths with small p-bars interspersed arranged in a single band of 3–5 rows (Fig.
Perinereis fayedensis sp. nov. All pictures are from the holotype if not stated otherwise A anterior end, maxillary apparatus, dorsal view B anterior end, maxillary apparatus, ventral view C posterior end, ventral view D right parapodium, anterior view, chaetiger 16 E right parapodium, posterior view, chaetiger 32 F right parapodium, posterior view, chaetiger 67 G chaetal bundle of a right parapodium, homogomph spiniger and heterogomph falcigers, chaetiger 17 H neuropodial chaetal bundle of a right parapodium, homogomph spiniger & heterogomph falciger, chaetiger 33 I heterogomph falciger, chaetiger 33.
Notopodia with conical dorsal and median ligules of equal length throughout. Superior lobes absent. DC length 1.2 (0.9–1.2) × length of DNL length anteriorly (chaetigers 10–20); posteriorly, DC length 1.2 (1.1–1.2) × length of DNL length (chaetigers 75–90). DC and DNL of mid-body parapodia proportionally similar to those of posterior parapodia (Fig.
Neuropodia with inferior and postchaetal lobes, ventral ligule and ventral cirri. Neuropodial postchaetal lobe lowly rounded, not projecting beyond end of acicular ligule. Ventral neuropodial ligule subconical, ca. as long as median ligule throughout. Ventral cirri extending laterally to halfway to tip of ventral neuropodial ligule in anterior and midbody, extending to ~ 1/3 length of ventral neuropodial ligule posteriorly (Fig.
Aciculae black, single in each ramus (Fig.
Pygidium with anal cirri fine, tapering, extending to last 7 (6–8) chaetigers, 50 (45–55) mm long (Fig.
The molecular data place the new species, P. fayedensis, clearly apart from all other species and as sister to P. suezensis (Fig.
Morphologically, P. fayedensis is intermediate between P. nuntia and P. heterodonta described from Obock, Djibouti, Gulf of Aden. It differs from the former most notably in the number of paragnaths in area III (2–5 vs. ~ 15 in P. nuntia) and area VI (14–17 vs. 8–12 in P. nuntia), and the relative length of the DC (1.1–1.2× DNL in the new species vs. 3–5× DNL in P. nuntia; Table
Gulf of Suez, Suez Canal including Great Bitter Lake; intertidal sand and mud, under stones.
The new species is named after the Egyptian city of Fayed on the western shore of Great Bitter Lake approximately halfway along the Suez Canal.
Holotype : DUFS 055, Hurghada (northern Red Sea), Grand Aquarium beach, subtidal area, clay bottom, at 27°07'59.2"N, 33°49'51.2"E. Paratypes: 22 specimens (DUFS 027–048) and non-type material 6 specimens (DUFS 049-054) from Al-Adabiya, west of Port Taofik, Gulf of Suez (Red Sea), intertidal, under coarse sands, at 29°56'06.0"N, 32°28'36.6"E and from Hurghada, National institute of Oceanography beach, intertidal and upper subtidal area, from muddy and sand bottoms, at 27°17'03.1"N, 33°46'19.8"E (Egypt).
Holotype (DUFS 055) not complete, 94 chaetigers, 62 mm in length, 4.5 mm wide at chaetiger 10. Paratypes with 42–96 chaetigers for 30–115 mm long and 1.5–7 mm wide at chaetiger 10. Epidermis with orange and gold pigmentation on anterior dorsum and ventrum in some preserved samples.
Prostomium with entire anterior margin; relatively large, longer than wide, two pairs of eyes, dark green with black lenses, and two large palps longer than antennae, palpostyles conical. Antennae closely set, as long as ~ 1/3 length of prostomium. Lenses not obvious.
One apodous anterior segment, ~ 1.6× longer than chaetiger 1. Tentacular cirri with distinct cirrophores, longest tentacular cirri extend back to chaetiger 2 (2–4).
Pharynx with jaws black, 4 (4–5) reddish brown teeth. Paragnaths black with light brown base; those of maxillary ring pointed conical paragnaths. Paragnath counts: area I with 0 (0–2); area II with 2 (1–5) on the left side and 3 (2–5) on the right side; area III with 4 (3–9) in one transverse row; area IV with 14 (10–21) on the left side and 16 (10–20) on the right side; arranged in irregular row of unequal paragnaths. Area V with 0 (0–1); area VI with 24 (24–40) conical paragnaths arranged in one arc; area VII–VIII with 24 (16–32), similar in size, arranged in two rows (Fig.
Perinereis damietta sp. nov. All pictures are from the holotype if not stated otherwise A anterior end, maxillary apparatus, dorsal view B anterior end, maxillary apparatus, ventral view C posterior end, ventral view D right parapodium, posterior view, chaetiger 9 E right parapodium, posterior view, chaetiger 41 F right parapodium, anterior view, chaetiger 56 G chaetal bundle of a right parapodium, homogomph spiniger & heterogomph falciger chaetiger 56 H neuropodial chaetal bundle of a right parapodium, heterogomph falciger, chaetiger 56 I heterogomph falciger, chaetiger 67.
Anterior notopodia with conical dorsal and median ligules of equal length in anterior body; dorsal ligule slightly longer in mid- and posterior body. DC length 0.8 (0.7–1.0) × length of DNL length anteriorly (chaetigers 10–20); posteriorly, DC length 1.1 (0.9–1.2) × length of DNL (chaetigers 75–90). DC and DNL of mid-body parapodia proportionally similar to those of posterior parapodia (Fig.
Dorsal notopodial ligule; triangular with conical tip, slightly longer than notopodial ventral ligule throughout. Ventral notopodial ligule rounded triangular. Dorsal and ventral notopodial ligules marked decreasing in size on posterior chaetigers. Neuropodium with dorsal rounded lobe in anterior chaetigers, with one black acicula, less developed posteriorly. Ventral neuropodial ligule digitiform, similar in length to acicular ligule on anterior chaetigers; slightly longer than acicular ligule in posterior chaetigers. Ventral cirri extending to ~ 1/3 length of ventral neuropodial ligule anteriorly and posteriorly (Fig.
Notochaetae with homogomph spinigers, long and thin serrated blade throughout. Neurochaetae dorsal fascicle: homogomph spinigers; median thick serrated blade present and heterogomph falcigers present on anterior and posterior chaetigers, blades serrated. Neurochaetae ventral fascicle: heterogomph falcigers with median long and wide blades with a single terminal tooth, in anterior and posterior chaetigers (Fig.
Pygidium with anal cirri cirriform, cirri extending to last 2 (2–4) chaetigers (Fig.
Gulf of Suez, northern Red Sea; intertidal and subtidal, sand and mud, under stones.
The new species is named after the university of the first author, Damietta University, a noun in apposition. Damietta (Egyptian Arabic: Dumyāț (دمياط) is also a port city located on an eastern distributary of the Nile Delta, ~ 15 km from the Mediterranean Sea.
Perinereis damietta sp. nov. is well supported by the highest bootstrap value of 100 (Fig.
Herein, P. vallata also includes one specimen (JX966314) assigned to P. brevicirris (Fig.
Perinereis damietta is morphologically most similar to P. heterodonta (type locality: Obock, Djibouti, Gulf of Aden). Both species belong to the group of the P. nuntia complex that lack heterogomph spinigers in anterior parapodia, which is unlike P. nuntia. Other key differences between P. damietta /P. heterodonta and P. nuntia are the shorter tentacular cirri and the fewer paragnaths in area V (0–1) (Table
Another species originally described from Djibouti, Perinereis djiboutiensis, is unfortunately poorly known, especially in respect to the presence or absence of heterogomph spinigers in anterior parapodia and numbers of paragnaths in areas III and VII–VIII (Table
A novel character introduced by
The present study supports the finding of
Despite recent advancements in integrative studies in many groups of polychaetes, taxonomic confusion still exists in many groups of Nereididae. Perinereis species are especially problematic due to difficult morphological species differentiation and a lack of detailed systematic studies. This has led to informal denomination of the species complex and recognition of geographic morphs and varieties such as P. cultrifera (
Finally, this study has uncovered further examples of sympatry among polychaetes. All three new species described here were found in the same habitat, viz., intertidal sand and mud, under stones, at the same location. Perinereis damietta appears to have a slightly wider habitat preference as it also occurs sub-tidally, but more intense sampling including exploration of potential microhabitat differences, is required to confirm our observations. Assuming sympatry, identification of the specific isolation mechanism(s) would be interesting. Several studies have suggested the importance of reproductive isolation as an important speciation mechanism in the species group (e.g.,
This work was funded by the Egyptian Government to AHE for a research stay at the NHM of University of Oslo. THS received additional support by the Norwegian Metacenter for Computational Science (NOTUR; project numbers NN9408K & NS9408K). This is NHM Evolutionary Genomics lab contribution No #18. We gratefully acknowledge the use of Google map. CG thanks Dr Dieter Fiege (
Supplementary data
Data type: excel file.
Explanation note: Morphometric characterization of 45 specimens of three putative species of Perinereis, identified by voucher number and sample identification (as for Table