Research Article |
Corresponding author: Charles H.J.M. Fransen ( charles.fransen@naturalis.nl ) Academic editor: Sammy De Grave
© 2018 Werner de Gier, Charles H.J.M. Fransen.
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:
de Gier W, Fransen CHJM (2018) Odontonia plurellicola sp. n. and Odontonia bagginsi sp. n., two new ascidian-associated shrimp from Ternate and Tidore, Indonesia, with a phylogenetic reconstruction of the genus (Crustacea, Decapoda, Palaemonidae). ZooKeys 765: 123-160. https://doi.org/10.3897/zookeys.765.25277
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Two new species of palaemonid shrimp associated with ascidian hosts, Odontonia bagginsi sp. n. from Tidore and Odontonia plurellicola sp. n., from Ternate, Indonesia are described and figured. Through phylogenetic analyses based on both morphological and molecular datasets (mitochondrial Cytochrome c oxidase subunit I gene and the 16S mitochondrial ribosomal gene) of the genus Odontonia, the phylogenetic positions of the new species have been reconstructed. Scanning Electron Microscopy has been used to observe additional characters on dactyli of the ambulatory pereiopods. Odontonia plurellicola sp. n. appears to be more closely related to O. simplicipes and O. seychellensis, but it differs most notably in the morphology of the rostrum and mouthparts. Odontonia plurellicola sp. n. appears to be the only Odontonia species living inside a phlebobranch ascidian Plurella sp. Odontonia bagginsi sp. n. is closely related to O. sibogae, but differs markedly in the abundance of setae on the propodi of the ambulatory pereiopods. In the present paper, O. maldivensis Fransen, 2006 is regarded as a junior synonym of O. rufopunctata Fransen, 2002 based on both morphological and molecular aspects.
Ascidians, Caridea , new species, Odontonia , Palaemonidae , symbiosis, Ternate, Tidore
The palaemonid genus Odontonia Fransen, 2002, currently contains 7 species (
In 2009 an expedition was organised by the Naturalis Biodiversity Center (Leiden, the Netherlands) and the Research Center of Oceanography, Indonesian Institute of Sciences (RCO-LIPI) (Jakarta, Indonesia), to the Indonesian islands of Ternate and Tidore (
The following abbreviations are used: COI, mitochondrial gene cytochrome c oxidase subunit I; 16S, 16S mitochondrial ribosomal gene; PoCL, post orbital carapace length; RMNH, Naturalis Biodiversity Center, Leiden (formerly Rijksmuseum van Natuurlijke Historie); MZB, Museum Zoologicum Bogoriense, Research Center for Biology, Indonesian Institute of Sciences, Cibenong, Indonesia.
Specimens were gathered around Ternate, in the northern Moluccas during an expedition organised from 23 October to 18 November 2009, by the Research Center of Oceanography, Indonesian Institute of Sciences (RCO-LIPI) (Jakarta, Indonesia) and Naturalis Biodiversity Center (NBC) (Leiden, the Netherlands). This expedition was part of the Ekspedisi Widya Nusantara project (E-Win expeditions). Specimens were collected using SCUBA equipment. Live specimens were photographed with a Nikon D80 digital camera and preserved in 70% ethanol.
Specimens were studied and drawn using a dissecting stereomicroscope (Zeiss Discovery.V8) and a compound microscope (Olympus BX53) both provided with a drawing tube. Sketches were traced using 2.5 to 3 mm Sakura Pigma Micron-pens and scanned (Canon Canoscan 9000F) with a resolution of 600 dpi. Details of the third pereiopods were photographed with a Scanning Electron Microscope. Pereiopods were dried using Critical Point Drying-methods (CPD) in a Leica EM CPD300 (located in Biopartner, Leiden, the Netherlands) with the following parameters (standard protocol preservation insects): CO2 intake: Auto, Speed slow, Delay 120s; Exchange: Speed 5, Cycles 18; Gas output: Heat medium, Speed slow 100%. The pereiopods were placed on stubs/mounts in pairs of two species, and coated with 20 nm Pl/Pd using a Quorum Q150T S. The dactyli were photographed using a JEOL JSM-S480LV Scanning Electron Microscope (located in the Naturalis Biodiversity Center, Leiden, the Netherlands). Drawings and photographs were edited in Adobe Photoshop (CS6) for better contrast and brightness.
Morphological character state analysis (Appendix I) was based on specimens in the RMNH collection (Table
Specimens used for SEM photography and 16S & COI DNA analyses (roman numerals can be linked back to the phylogenetic trees (Figs
Species | Sample location | Host organism | Registration number | GenBank 16S | GenBank COI | SEM |
---|---|---|---|---|---|---|
Odontonia bagginsi sp. n. | Indonesia, Halmahera, Tidore | Unid. ascidian | MZB Cru 4733 | MH257316 | × | |
Odontonia katoi (Kubo, 1940) | Indonesia, Bali, Tulamben beach | Polycarpa aurata | RMNH.CRUS.D.48689 | MH251614 | ||
Indonesia, SW Sulawesi, Spermonde Archipelago | Polycarpa aurata | RMNH.CRUS.D.46701 | × | |||
Indonesia, NE Sulawesi, Bitung | Polycarpa aurata | RMNH.CRUS.D. 57295 | MH251615 | |||
Odontonia maldivensis Fransen, 2006 | Maldives, S Malé Atoll | Polycarpa cryptocarpa | RMNH.CRUS.D.51001 | × | ||
Maldives, Faafu Atoll, Magoodhoo Island | Polycarpa cryptocarpa | RMNH.CRUS.D. 57296 | MH251616 | |||
Maldives, Faafu Atoll, Magoodhoo Island | Herdmania momus (?) | RMNH.CRUS.D. 57297 | MH251617 | |||
Odontonia plurellicola sp. n. | Indonesia, W Halmahera, Ternate | Plurella sp. | RMNH.MZB Cru 4734 | × | ||
Odontonia rufopunctata Fransen, 2002 | Indonesia, Halmahera mainland, Tanjung Ratemu | Unid. ascidian | RMNH.CRUS.D.53601 | MH257314 | ||
Indonesia, NE Sulawesi, Bitung | Polycarpa sp. | RMNH.CRUS.D. 57298 | MH251618 | MH257313 | ||
Indonesia, NE Sulawesi, Bitung | Polycarpa sp. | RMNH.CRUS.D. 57299 | MH251619 | |||
Indonesia, NE Sulawesi, Bitung | Polycarpa sp. | RMNH.CRUS.D57300 | MH251620 | |||
Indonesia, SW Sulawesi, Spermonde Archipelago | Unid. ascidian | RMNH.CRUS.D.48694 | × | |||
Odontonia seychellensis Fransen, 2002 | Seychelles, E of Mahé | Unid. stolidobranch ascidian * | RMNH.CRUS.D.42762 | × | ||
Odontonia sibogae (Bruce, 1972) | Indonesia, Bali, Sanur | Polycarpa sp. | RMNH.CRUS.D.48691 | MH251621 | ||
Indonesia, Halmahera, Tidore | Unid. ascidian | RMNH.CRUS.D.53558 | MH257315 | |||
Indonesia, Borneo, Sabah | Polycarpa argentata | RMNH.CRUS.D.53964 | JX185703 | |||
Indonesia, Ambon, E coast | Polycarpa sp. | RMNH.CRUS.D.47581 | × | |||
Outgroup species: | ||||||
Pontonia panamica Marin & Anker, 2008 | Panama, Playa Venao | Ascidia sp. | RMNH.CRUS.D.51825 | MH251622 | MH257312 | |
Pontonia pinnophylax (Otto, 1821) | Cape Verde Islands, S coast of São Vicente | Pinna rudis | RMNH.CRUS.D.42607 | KU170692 | ||
Cape verde Island, São Nicocau | Pinna rudis | RMNH.CRUS.D.42608 | MH251623 | |||
Dactylonia ascidicola (Borradaile, 1898) | Indonesia, Bali, Tulamben area | Ascidia sp. | RMNH.CRUS.D.48678 | KU170688 | ||
Indonesia, Borneo, Sabah | Ascidia sp. | RMNH.CRUS.D.53793 | MH257317 |
Data matrix of morphological characters used for phylogenetic analysis. Ordered characters in bold face, other characters unordered.
Species | |||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Outgroup species | Odontonia species | ||||||||||||
Dactylonia ascidicola | Pontonia pinnophylax | Pontonia panamica | Odontonia bagginsi sp. n. | Odontonia compacta | Odontonia katoi | Odontonia maldivensis | Odontonia rufopunctata | Odontonia seychellensis | Odontonia sibogae | Odontonia simplicipes | Odontonia plurellicola sp. n. | ||
Characters | 1 | 0 | 1 | 1 | 2 | 2 | 2 | 0 | 2 | 2 | 2 | 0 | 2 |
2 | 0 | 0 | 0 | 1 | 2 | 2 | 2 | 2 | 1 | 1 | 2 | 1 | |
3 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | |
4 | 1 | 0 | 0 | 1 | 1 | 1 | 1 | 1 | 2 | 1 | 2 | 2 | |
5 | 0 | 0 | 0 | 0 | 1 | 1 | 1 | 0 | 1 | 1 | 1 | 1 | |
6 | 0 | 0 | 0 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 0 | 1 | |
7 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 1 | |
8 | 0 | 0 | 2 | 1 | 0 | 2 | 1 | 0 | 0 | 0 | 2 | 1 | |
9 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 2 | 0 | 0 | 0 | 0 | |
10 | 1 | 0 | 0 | 0 | 1 | 1 | 2 | 1 | 1 | 0 | 1 | 2 | |
11 | 0 | 0 | 0 | 1 | 1 | 1 | 0 | 0 | 1 | 1 | 1 | 1 | |
12 | 0 | 0 | 0 | 0 | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 2 | |
13 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 2 | |
14 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | |
15 | 2 | 0 | 0 | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 1 | |
16 | 1 | 0 | 0 | 2 | 2 | 2 | 2 | 2 | 0 | 2 | 2 | 2 | |
17 | 1 | 0 | 0 | 0 | 1 | 1 | 2 | 2 | 0 | 1 | 0 | 0 | |
18 | 0 | 1 | 0 | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 1 | |
19 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 1 | 1 | |
20 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | |
21 | 0 | 0 | 0 | 1 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | |
22 | 0 | 1 | 1 | 0 | 2 | 1 | 1 | 1 | 1 | ? | 1 | 1 | |
23 | 1 | 0 | 0 | ? | 1 | 1 | 1 | 1 | 1 | 1 | ? | 2 | |
24 | 0 | 0 | 0 | 1 | ? | 1 | 1 | 1 | 1 | 0 | 2 | 2 |
Mitochondrial COI (7 sequences) and 16S (12 sequences) were available for a partial molecular phylogenetic reconstruction of the genus Odontonia (Table
Host records for Odontonia species were assembled from the literature (
(modified from
The type species: Pontonia katoi Kubo, 1940, by original designation, gender feminine.
Known from shallow coastal waters of the Indo-West Pacific.
Associated with Ascidiacea.
Type series. 1 ovigerous female (holotype), PoCL 1.55 mm (MZB Cru 4734), Tarau, W Halmahera, Ternate, Indonesia, 0°50'30"N, 127°22'38.5"E, shallow area with coral followed by sandy slope with coral gardens, 9 m depth, scuba diving, 2-11-2009; in ascidian Plurella sp. (Asc. 68), leg. C.H.J.M. Fransen, photo TER.17.0049 – 76; 1 male, PoCL 1.30 mm, 1 ovigerous female, PoCL. 1.50 mm, 2 non-ovigerous females PoCL 0.90–1.05 mm (paratypes) (RMNH.CRUS.D.53554), same data as holotype.
Rostrum as long as antennular peduncle, with distoventral tooth. Pterygostomial angle produced. Basal segment of antennular peduncle with distolateral tooth minute, medioventral tooth strong, acute. Distolateral tooth of scaphocerite robust, 0.3 length of lamina. Dactylus of ambulatory pereiopods with flexor margin of corpus with few (usually 3) short teeth but without accessory tooth; unguis without terminal scales. Telson with two pairs of medium sized (approx. 0.17 of telson length) submarginal dorsal spines at 0.20 and 0.54 of telson length.
Body (Figs
Abdomen smooth, sixth segment 1.4 times longer than fifth, 1.4 times wider than long, posterolateral angle blunt, slightly produced, posteroventral angle blunt, not produced; pleura of first five segments broadly rounded.
Telson (Fig.
Eyestalk short, broader than long, slightly broader than diameter of hemispherical cornea.
Antennula (Fig.
Antenna (Fig.
Epistome with rather sharp anterior carina; labrum normal.
Paragnath well developed, alae with broad transverse more or less rectangular distal lobes, and small rounded more or less triangular ventromedial lobes; corpus very short, with shallow median excavation, bordered laterally by non-setose, small, oblique, carinae.
Second thoracic sternite (Fig.
Third thoracic sternite with indistinct shallow lateral carinae.
Fourth thoracic sternite with shallowly developed, medially notched plate formed by the lateral carinae.
Fifth thoracic sternite with well-developed lateral plates with medial broadened deep slit, posteromedial to second pereiopod coxae.
Sixth to eight thoracic sternites unarmed, broadening posteriorly.
Mandible (Fig.
Maxillula (Fig.
Maxilla (Fig.
First maxilliped (Fig.
Second maxilliped (Fig.
Third maxilliped (Fig.
First pereiopod (Fig.
Second pereiopods (Fig.
Ambulatory pereiopods short, stout. Dactylus of third pereiopod (Fig.
First pleopod of female (Fig.
Odontonia plurellicola sp. n., ovigerous female PoCL 1.50 mm (A, D), male PoCL 1.30 mm (B, C) (RMNH.CRUS.D.53554). A first pleopod of female with endopod B male first pleopod with endopod C endopod of second pleopod with appendix masculina and appendix interna D detail of left uropod. Scale bars: 1 mm (A, C, D); 0.5 mm (B).
Endopod of second pleopod (Fig.
Number of eggs approximately 11.
This is a small sized species. The maximum PoCL is 1.55 mm in adult females, 1.30 mm in adult males. The minimal PoCL in ovigerous females is 1.50 mm.
(Fig.
Specimens were found inside a colonial ascidian of the genus Plurella Kott, 1973 (Plurellidae, Phlebobranchia).
Only known from the type locality.
The species is named plurellicola after the colonial ascidian genus Plurella Kott, 1973 in which it was found.
The species resembles O. simplicipes, known only by the holotype, in morphological characters. It differs from this species in the length and shape of the rostrum (most notably, O. simplicipes has no distal tooth on its rostrum, while O. plurellicola bears a small distal tooth), in the size of the ventromedial tooth on the basal segment of the antennular peduncle which is larger in O. seychellensis than in the new species, in the distolateral tooth of the basal segment of the antennular peduncle which is well developed in O. seychellensis while minute in the new species, in the amount of plumose setae on the three maxillipedes and the antennular peduncle.
Thus far this is the only species of Odontonia living inside a colonial ascidian. The ascidian genus Plurella has also been recorded as host for Dactylonia holthuisi Fransen, 2002, another symbiotic palaemonid shrimp (
(i) Indo-West Pacific: Indonesia. –1 ovigerous female (holotype) PoCL 3.4 mm (MZB Cru 4733, ex RMNH.CRUS.D.53559), N of Desa Rum, Tidore, Ternate, Indonesia, 0°44'35.8"N 127°23'6.3"E, 27 m depth, scuba diving, reef consisting primarily of boulders and soft corals; 4-11-2009; in solitary ascidian (Asc. 67), leg. A. Gittenberger, photo TER.17.0136–39.
Diagnosis. Rostrum as long as antennular peduncle, with strong distoventral tooth. Pterygostomial angle broadly rounded, produced. Basal segment of antennular peduncle with strong, acute medioventral tooth. Distolateral tooth of scaphocerite robust, 0.4 length of lamina. Dactylus of ambulatory pereiopods with blunt accessory tooth, perpendicular to flexor margin; flexor margin of corpus with strong, acute forward directed proximal tooth and two small denticles between this tooth and accessory tooth; unguis without terminal scales. Telson with two pairs of medium sized (0.13 of telson length) submarginal dorsal spines at 0.15 and 0.48 of telson length.
Description. Body (Figs
Abdomen smooth, sixth segment 1.3 times longer than fifth, 1.4 times wider than long, posterolateral angle blunt, slightly produced, posteroventral angle blunt, not produced; pleura of first five segments broadly rounded.
Telson (Fig.
Eyestalk short, about as long as broad, as broad as diameter of hemispherical cornea.
Antennula (Fig.
Antenna (Fig.
Epistome anteriorly broadly rounded; labrum normal, oval.
Paragnath well developed, alae with broad transverse more or less rectangular distal lobes, and small rounded more or less triangular ventromesial lobes; corpus very short, with shallow median excavation, bordered laterally by non-setose, small, oblique, carinae.
Second thoracic sternite with anterior margin broadly rounded; without median process forming round tubercle.
Third thoracic sternite with indistinct shallow lateral carinae.
Fourth thoracic sternite with developed, bluntly triangular medial plate without median notch.
Fifth thoracic sternite with well-developed lateral plates with medial broadened deep slit, posteromedial to second pereiopod coxae.
Sixth to eight thoracic sternites unarmed, broadening posteriorly.
Mandible (Fig.
Maxillula (Fig.
Maxilla (Fig.
First maxilliped (Fig.
Second maxilliped (Fig.
Third maxilliped (Fig.
First pereiopod (Fig.
Second pereiopods (Fig.
Ambulatory pereiopods short, stout. Dactylus of third pereiopod (Figs
First pleopod with endopod almost half as long as exopod, with plumose setae laterally and distally, with long simple setae distomedially.
Uropods, with short unarmed protopodite; exopod broad, 2.2 times longer than central width, lateral margin feebly convex, without distolateral tooth, with minute spinule distolaterally; endopod exceeding exopod, about as long as telson, 2.8 times longer than wide.
Ovigerous female with about 100 eggs of 0.05 mm in diameter.
(Fig.
Solitary ascidian (A. Gittenberger Asc. 67).
Only known from its type locality at Tidore, Indonesia.
The species is named “bagginsi”, inspired by the famous Hobbit family name “Baggins” featured in the “The Hobbit” and “The Lord of the Rings” books. The fictional characters called “Hobbits” possess hairy feet comparable to this species.
The species bears resemblance to O. sibogae in its morphological characters. It differs in having a strongly developed ventral tooth on the rostrum; a strongly produced pterygostomial margin of the carapace; two pairs of dorsal spines on the telson and a broad upper lacinia of the maxillula. In addition, the pereiopods bear some notable differences: the segments of the first pereiopods are stouter and the cutting edge of the major cheliped bears a broad flattened tooth. The dactyli of the ambulatory pereiopods bear two denticles on the flexor margin. The unguis is devoid of distal scales. Most characteristic is the dense cover of simple setae on the propodi of the ambulatory pereiopods. Life colour patterns of the new species is similar to that of O. sibogae (see Levitt & Shenkar 2018: fig. 2B) with scattered white chromatophores of various sizes scattered over body and appendages. The new species has a reddish overall appearance while O. sibogae is paler.
The morphological phylogenetic analysis (Fig.
The resulting tree from the incomplete COI dataset (Fig.
The resulting tree of the incomplete 16S dataset (Fig.
Most Odontonia species live as endosymbionts in ascidian species in the order Stolidobranchia (Fig.
Both molecular and morphological phylogenetic analyses show good resolution in the distal part of the tree while basal support is low. The sister position of the species pair O. sibogae – O. bagginsi in the morphological tree is confirmed by the COI phylogeny. Low support in the basal nodes of the molecular phylogenies might be an effect of the chosen markers. More conservative markers might give better resolution in the basal part of the tree. However, the low support of the basal branches in the molecular trees and the basal polytomy in the morphological phylogeny could also be an indication of a rapid radiation over the host species as was shown for Caribbean sponge-dwelling snapping shrimps Synalpheus (
The similar 16S sequences of O. rufopunctata and O. maldivensis as well as their highly similar morphology, indicate that these nominal species actually form one species. Differences indicated by
Several Odontonia species (O. simplicipes, O. compacta, and O. bagginsi sp. n.) are only known from a single or few type specimens. Therefore, intraspecific morphological variation is not known and molecular data are not available (except for O. bagginsi sp. n.) which hampers the present phylogenetic analyses. When more material of these rare species comes available a more comprehensive phylogenetic analysis can be performed.
Most Odontonia species live as endosymbionts in ascidian species in the order Stolidobranchia (Fig.
1 | Dactylus of ambulatory pereiopods without proximal teeth on flexor margin or with row of few shallow, forward directed teeth; unguis glabrous | 2 |
– | Dactylus of ambulatory pereiopods with single large forward directed proximal tooth on flexor margin; distodorsal scales on unguis (except in O. bagginsi) | 4 |
2 | Dactylus of ambulatory pereiopods with row of few shallow, forward directed teeth on flexor margin | 3 |
– | Dactylus of ambulatory pereiopods with flexor margin entire | O. seychellensis |
3 | Rostrum with distal ventral tooth; distolateral tooth of antennular basal segment slightly exceeding distal margin of segment | O. plurellicola sp. n. |
– | Rostrum without distal ventral tooth; distolateral tooth of antennular basal segment reaching distal margin of intermediate segment | O. simplicipes |
4 | Unguis of dactylus of ambulatory pereiopods without or with few distal scales (not more than 5); small denticles between accessory and proximalmost tooth | 5 |
– | Unguis of dactylus of ambulatory pereiopods with patch with many distal scales; no denticles between accessory and proximalmost tooth | O. rufopunctata |
5 | Rostrum not overreaching antennular peduncle | 6 |
– | Rostrum overreaching antennular peduncle | 7 |
6 | Telson with 2 pairs of dorsal spines; unguis without scales; propodi of ambulatory pereiopods with many long simple setae | O. bagginsi sp. n. |
– | Telson with 5 pairs of dorsal spines; unguis with few distodorsal scales; propodi of ambulatory pereiopods almost devoid of setae | O. sibogae |
7 | Unguis of dactylus of ambulatory pereiopods with two distal scales, and one denticle between accessory and proximalmost tooth | O. compacta |
– | Unguis of dactylus of ambulatory pereiopods with one distal scale, and (with or without) five small denticles between accessory and proximalmost tooth | O. katoi |
The authors would like to thank Bastian T. Reijnen for his assistance with the SEM, Bertie-Joan van Heuven for her help preparing and coating the pereiopods for the SEM, and Livia Oliveira for her help with the ascidian phylogeny.
1. Ventral teeth: 0, absent; 1, one subdistal; 2, one apical. Character states 0 and 1 are found in the outgroups. An apical tooth (often accompanied by some distal setae) is found in most Odontonia species, while a subdistal tooth is found only in the Pontonia members of the outgroup. No tooth is found in the outgroup Dactylonia ascidicola, in O. simplicipes and O. maldivensis. (Fig.
2. Length of rostrum: 0, falling short of antennular peduncle; 1, around the same length as antennular peduncle; 2, overreaching antennular peduncle. A small rostrum is found in all outgroups, while in most Odontonia species the rostrum is overreaching, or as long as, the antennular peduncle.
3. Relative size rostrum to post-orbital carapace length: 0, around 0.15 to 0.45 times the PoCL; 1, around 0.75 times the PoCL. All outgroup species, as well as most Odontonia species possess a small rostrum-PoCL ratio, while only O. simplicipes has a rostrum-PoCL ratio of 0.75.
4. Relative size rostrum to hemispherical diameter of cornea: 0, around 4.0 times the cornea size; 1, around 2.5 to 3.0 times the cornea size; 2, around 2.0 times the cornea size. A rostrum-cornea ratio of 4.0 is found in the outgroup species Pontonia pinnophylax (Otto, 1821) and P. panamica, while most Odontonia species and the other outgroup species have a smaller ratio.
5. Anterolateral angle: 0, absent; 1, present. No angle is found in the outgroup species, as well as in O. rufopunctata Fransen, 2002 and O. bagginsi sp. n. (Fig.
6. Ventromedial tooth basal segment: 0, small; 1, strongly developed; A strongly developed tooth is found in almost all Odontonia species (hence the name of the genus), while a small tooth is found in the three outgroup species, and in O. simplicipes (Fig.
7. Dimensions of intermediate segment: 0, about as long as wide or slightly broader than long; 1, about twice as long as wide. Most Odontonia species share character state 0 with the outgroups. Odontonia plurellicola sp. n., O. simplicipes, and O. seychellensis Fransen, 2002 have a longer intermediate segment. The character is treated ordered in the analysis.
8. Plumose setae on median margin of intermediate segment: 0, two or more setae; 1, one setae; 2, no setae. The specific setae are absent in Pontonia panamica, as well as in O. katoi (Kubo, 1940) and O. simplicipes. The other two outgroup species have two or more setae, and most Odontonia species share this treat (Fig.
9. Maxilla: 0, upper lacinia and lower lacinia both well developed; 1, upper lacinia well-developed, lower reduced; 2, both lacinia almost completely fused. Dactylonia ascidicola is the only species in the analysis with an uneven development of the two lacinia. The other outgroups share both developed lacinia with almost all Odontonia species. Odontonia rufopunctata has no more distinguishable lacinia, both are almost completely fused.
10. First maxilliped; amount of distal setae on exopod: 0, eight or more setae; 1, five or six setae; 2, four setae. This character is treated as ordered; it is believed that the number of setae decreases in the exopods of Odontonia species due to their symbiotic lifestyle in ascidians.
11. Second maxilliped; angle in median margin of basis: 0, absent; 1, distinct angle. Character state 1 is found in two outgroup species, except for Pontonia panamica. Most Odontonia species do have an angle in the basis, with the exception of O. maldivensis and O. rufopunctata.
12. Second maxilliped; amount of distal setae on exopod: 0, eight or more setae; 1, five or six setae; 2, four setae. This character is treated as ordered (see character 10).
13. Third maxilliped; amount of distal setae on exopod: 0, eight or more setae; 1, five or six setae; 2, four setae. This character is treated as ordered (see character 10).
14. First pereiopod; ratio merus/carpus: 0, merus as long as carpus; 1, carpus shorter than merus. Character state 1 is present in outgroup species Dactylonia ascidicola, O. rufopunctata and O. bagginsi sp. n.
15. Third pereiopod; dactylus; teeth on flexor margin: 0, teeth absent; 1, dactylus with simple teeth; 2, dactylus with minutely spinulate blunt tubercles. Character state 2 is only found in Dactylonia ascidicola, and character state 0 is found in the two Pontonia outgroup species, and in O. seychellensis (Fig.
Variety in dactylar morphology of all currently known Odontonia species (setae omitted). Note the amount of distal scales, presence of an additional and accessory tooth, size of the dactylus, and the presence of denticles. A O. compacta (Bruce, 1996) B O. bagginsi sp. n. C O. katoi (Kubo, 1940) D O. maldivensis Fransen, 2006 E O. rufopunctata Fransen, 2002 F O. seychellensis Fransen, 2002 G O. sibogae (Bruce, 1972) H O. simplicipes (Bruce, 1996) I O. plurellicola sp. n.
16. Third pereiopod; dactylus; teeth on flexor margin: 0, teeth absent; 1, teeth similar, increasing in size; 2, proximalmost tooth strong, large, directed forward. The teeth of Dactylonia ascidicola are very different from the Odontonia and Pontonia species, hence character state 1. All Odontonia with teeth on their flexor margin have one large proximalmost tooth, with or without more teeth following (Fig.
17. Third pereiopod; dactylus, unguis: 0, distal scales absent; 1, unguis with fewer than five distal scales; 2, unguis with more than five distal scales. Scales are absent in the two Pontonia outgroup species, as well as in O. seychellensis, O. simplicipes, O. plurellicola sp. n., and O. bagginsi sp. n. The species O. maldivensis and O. rufopunctata have more than five distal scales (Fig.
18. Third pereiopod; dactylus; dimensions of corpus: 0, more than 2.0 times as long as broad; 1, less than 2.0 times as long as broad. Dactylonia ascidicola, Pontonia panamica and O. seychellensis all share a dactylus that is more than 2.0 times as long as broad (Fig.
19. Third pereiopod; dactylus; accessory tooth: 0, present; 1, absent. Most Odontonia species, as well as the outgroup species, possess an accessory tooth on the distal flexor margin. Odontoniaseychellensis, O. simplicipes, O. maldivensis and O. plurellicola sp. n. all have no accessory tooth (Fig.
20. Third pereiopod; dactylus; additional forward-pointing teeth distal from proximalmost tooth: 0, absent; 1, two additional forward-pointing teeth. Only O. simplicipes and O. plurellicola sp. n. possess the two additional teeth (Fig.
21. Third pereiopod; dactylus; denticles between accessory and proximalmost tooth: 0, absent; 1, one, two or five denticles in flexor margin. Odontonia sibogae and O. compacta are the only ones with one denticle. Odontonia bagginsi sp. n. has two denticles. Odontonia katoi is the only species with accounts of there being a row of five denticles on the flexor margin, but often these denticles are not visible or not present. The outgroup Dactylonia ascidicola has broad denticles considered not homologous with the ones from Odontonia (Fig.
22. Position of distal pair of dorsal spines over telson length: 0, at about 1/2; 1, at about 2/3; 2, in distal 1/4. Character state 0 is present in Dactylonia ascidicola and in O. bagginsi sp. n. The species O. compacta has the distal spines implanted in the most distal 1/4th of the telson length. In O. sibogae there are more than four spines implanted in the dorsal telson side. It is not sure which spines are homologous with the four spines in other species, hence the “?” in the dataset. This character is treated as ordered; it is believed the spines migrate more distally in the evolution of the shrimp.
23. Maximum PoCL of male specimens: 0, PoCL longer than 5.5 mm; 1, PoCL between 5.5 and 2.0 mm; 2, between 2.0 and 0.5 mm. Depending on the availability of the specimens a “?” may be given to missing character states.
24. Maximum length of female specimens: 0, PoCL longer than 5.5 mm; 1, between 5.5 and 2.0 mm; 2, between 2.0 and 0.5 mm. Depending on the availability of the specimens a “?” may be given to missing character states.