Tympanicheles ornatus (Holthuis, 1952)

de Gier, Werner & Fransen, Charles H. J. M., 2023, Polka-dotted treasures: Revising a clade of ascidian- and bivalve-associated shrimps (Caridea: Palaemonidae), Contributions to Zoology 92 (3), pp. 179-282 : 254-282

Tympanicheles ornatus (Holthuis, 1952)

(figs. 36A–B, 37, 39A,C)

Paranchistus ornatus Holthuis, 1952: 97–100 , figs. 39, 40 [type locality: Mozambique]; Barnard, 1955 (listed): 5; Barnard, 1958 (listed): 12; Rosewater, 1961: 195; Jacquotte, 1963: 61; Hipeau-Jacquotte, 1967: 153–166; Hipeau-Jacquotte, 1971: 194, figs. 4, 5 9; Bruce, 1972a: 219–221, colour figures (listed); Hipeau-Jacquotte, 1972a: 288, fig. 2a2; Hipeau-Jacquotte, 1972b: 10, 14, 17, figs. 1a2–e2, 3a2, 3b2, 4a2–d2, 6b, 7a2–f2, 8a2–d2; Thomassin, 1972: 311; Hipeau-Jacquotte, 1973a; 96–116, fig. 1a2–e2, 3a2–b2, 4a2–d2, 5, 6b, 7a2–f2, 8a2– d2; Hipaeu-Jacquotte, 1973b: 63–70; Hipeau-Jacquotte, 1974a: 367–382; Hipeau-Jacquotte, 1974b: 383–402; Hipeau-Jacquotte, 1974c: 403–407; Hipeau-Jacquotte, 1974d: 359–386; Thomassin, 1974:306; Bruce, 1975a:162 (listed); Bruce, 1976a: 93; Bruce, 1976d: 471: Monod, 1976a: 839 (listed); Bruce, 1978a: 119; Bruce, 1978b: 279; Bruce, 1980: 179, textfig. 1a, pl. 1 figs. a–c; Müller, 1993: 51 (listed); Fransen, 1994a: 107, 111, 112, fig. 2g, pl. 2 fig. 3; Fransen, 1994b: 117, fig. 2g, pl. 2D; Li, 2000: 111, fig. 121 (listed); De Grave & Fransen, 2011: 357 (listed); Chow et al., 2021: table A.1 (listed).

Material examined. RED SEA: 3 males pocl. 3.90, 4.22, 4.74 mm, 7 ovigerous females pocl. 7.40, 7.92, 8.18, 8.18, 8.31, 8.70, 8.83 mm ( RMNH. CRUS.D.41491), Nocra Island , 18.iii.1962, Israel Red Sea Expedition no. E62/2028. – 1 ovigerous female pocl. 6.10 mm ( RMNH. CRUS.D.41492), Loe Salina , 1965, in Atrina sp. , Israel Red Sea Expedition no. E65/1868. – 3 males pocl. 3.64, 3.90, 4.42 mm, 3 ovigerous females pocl. 7.53, 7.66, 8.70 mm ( RMNH. CRUS.D.41493), off Nocra Island , 17.iii.1962, in Atrina sp. , Israel

Red Sea Expedition no. E62/1350.– 1 male pocl. Island, 4°37’S 55°31’E, 25.xii.1992, depth 10 m; 5.32 mm, 1 ovigerous female pocl. 9.09 mm SCUBA diving, in Atrina vexillum (Born) , coll. ( RMNH. CRUS.D.41494), NE of Nocra Island, C.H.J.M. Fransen. 18.ii.1962, depth 2-3 m, in Atrina sp. (10049), Diagnosis. Rostrum well developed, lat- Israel Red Sea Expedition no. E62/1354. – erally compressed towards distal part, bent 3 males pocl. 4.35, 4.87, 4.94 mm ( RMNH. downwards, reaching end of te second seg- CRUS.D.41495), Nocra Island , 18.iii.1962, Israel ment of antennular peduncle, with 3–5 dor- Red Sea Expedition no. E62/2059. – 1 male sal distal teeth, without ventral distal teeth; pocl. 4.48 mm ( RMNH. CRUS.D.41496), Umm row of setae between dorsal teeth. Carapace Aabak, 4.v.1962, Israel Red Sea Expedition with well-developed antenna tooth, with no. 1452. – 2 males pocl. 3.05, 3.57 mm, 1 small articulating hepatic spine, situated female pocl. 5.52 mm, 1 ovigerous female pocl. somewhat posteroventrally of antennal spine; 6.36 mm ( RMNH. CRUS.D.41497), SW of Umm inferior orbital angle acute, slightly produced; Aabak, near Nocra Island , 22.iii.1962, depth anterolateral angle broadly rounded, slightly 0-3 m, in Atrina sp. , Israel Red Sea Expedition produced. Sixth abdominal segment with posno. E62/1379. MOZAMBIQUE: 1 male holo- terior ventral margin rounded. Telson with type pocl. 5.97 mm ( ZMA. CRUS.D.102827), two pairs of small dorsal spines at 0.58 and published by Holthuis , 1952: 97–100, figs. 39, 0.81 of telson length; lateral pair of terminal 40. SEYCHELLES: 1 ovigerous female pocl. spines placed just behind intermediate pair. 7.7 mm ( RMNH. CRUS.D.42794), NIOP-E stn Basal segment Downloaded of from antennular Brill.com peduncle 06/24/2024 with 12:38:05 AM SEY.748, E of Mahé, E coast of via Sainte Open Anne Access. This one is (seldom an open access two) distal article teeth distributed, anterior under margin the terms of the CC-BY 4.0 license. https://creativecommons.org/licenses/by/4.0 /

almost straight; outer flagellum with basal 4–6 Second pereiopods usually equal in size and segments fused, free shorter ramus of two or shape in females, subequal in size in males; three segments. Scaphocerite well developed dactylus with one large tooth in proximal half;

with slightly convex lateral margin; distolat- fixed finger with 5–6 small teeth in proximal

eral tooth normal, almost reaching the distal half; palm with oblong ventral oval tympamargin of the lamina; slit between distolat- nal organ over entire length. Ambulatory

eral tooth and lamina shallow. First pereio- pereiopods with slender long slightly curved

pod slender, chela sub-spatulate, minutely non-biunguiculate Downloaded dactyli from Brill;.com serrated 06/24/2024 carinae12:38:05 AM pectinate in distal two-third of lateral via margin Open Access. on. This distoventral is an open access margin article of corpus distributed; unguis under not the terms of the CC-BY 4.0 license. https://creativecommons.org/licenses/by/4.0/

flattened, dorsally and ventrally with trans- 2001: 107, 109, 135, fig.; Hayashi, 2002: 224, verse rows of scale-lake spinules. Uropod with figs. 432k, l, 434i, j, 435i, j; Hayashi, 2006: 510, protopodite posterolaterally acute, exopod 512; De Grave & Fransen, 2011: 340 (listed); with small mobile distolateral spine. Radhakrishnan et al., 2012: table 1 (listed); Remarks. Smaller specimens have the dor- Samuel et al., 2016: table 1 (listed).

sal rostral teeth in the distal fourth of the ros- Anchistus misakensis Yokoya, 1936: 136 , fig. 5 trum while larger specimens have them all at [Type locality: Octavia Bay, Nancowry Harbor, the tip. This indicates that the dorsal rostral Nicobars]; Holthuis, 1952: 13; Bruce, 1967: 568 teeth are placed more distally after each moult (key).

as has been observed in Anchistus australis . Material examined. JAPAN: 1 ovigerous Colour (figs. 36A,B). “Uniformly covered female pocl. 3.51 mm, 1 male pocl. 3.90 mm all over body and appendages with small red ( RMNH. CRUS.D.26972), Kanagawa prefecture, chromatophores, with minute yellow and Sagami Bay near Manazuru, S of Odawara, white chromatophores interspersed.” ( Bruce , 25.viii.1968, coll. H. Suzuki. INDONESIA: 1 1978a: 119; see colour figures Bruce, 1972a). non ovigerous female pocl. 1.8 mm ( RMNH.

Hosts. Reported from Pinnidae : Atrina vex- CRUS.D.47465), stn MAL.21, Moluccas, Ambon, illum (Born, 1778) (Hipeau-Jacquotte, 1967, Ambon bay, N coast Cape Hatupero, E of 1973b, 1974b; Bruce, 1976e, 1978a, 1978b, 1980; Lilibooi, 03°44’S 128°02’E, 20.xi.1996, depth Fransen, 1994b); Pinna bicolor Gmelin, 1791 10 m, SCUBA diving, in pectinid shell, coll.

( Rosewater, 1961; Hipeau-Jacquotte, 1967). C.H.J.M. Fransen. VANUATU: 1 non ovigerous Distribution. Distributed along the East female pocl. 1.6 mm ( RMNH. CRUS.D.58094), African coast: Mocambique (Holthuis, 1952); stn LD25, N Aoré, 15°19’29”S 167°06’19”E, Zanzibar and Kenya (cf. Rosewater, 1961; 7.x.2006, scuba diving, depth unknown.

Bruce, 1976d, 1980), Madagascar (cf. Hipeau- Diagnosis. Rostrum well developed, lat- Jacquotte, 1967, 1971, 1972a, 1972b, 1973a, 1973b, erally compressed towards distal part, bent 1974a, 1974b, 1974c, 1974d; Thomassin, 1972, downwards; reaching just beyond basal seg- 1974; Bruce, 1978b), Comoro Islands (cf. Bruce, ment of antennular peduncle, with 3, seldom 1978a); Seychelles (Fransen, 1994b); Nocra 2 distal teeth with setae in between. Antennal Island, Red Sea (Fransen, 1994a). spine present, hepatic spine absent; inferior orbital angle slightly produced; anterolateral Tympanicheles pectinis (Kemp, 1925) angle rounded, not produced. Sixth abdom-

(figs. 36C,D, 38) inal segment with posterior ventral margin Anchistus pectinis Kemp, 1925: 327–330 , figs. rounded. Telson with dorsal pairs of spines 19, 20 [Type locality: Octavia Bay, Nancowry at 0.61 and 0.79 of telson length; lateral pair Harbor, Nicobars]; Holthuis, 1952: 13 (listed); of terminal spines placed just behind inter- Bruce, 1967: 568; Fujino & Miyake, 1967: 293- mediate pair. Basal segment of antennular 296, figs. 1–3; Suzuki, 1971: 101–106, figs. 5–7, pl. peduncle with one small distolateral tooth.

2; Bruce, 1972a: 219 (listed); Bruce, 1975a: 150, Scaphocerite with lateral margin almost 162 (listed); Bruce, 1976d: 465; Bruce, 1977b: straight, slit between distolateral tooth and 175–176, fig. 6e; Bruce, 1983c: 200 (listed); lamina of moderate depth. Chela of first Bruce, 1990: 15, 18 (listed); Bruce, 1991a: 261, pereiopods spatulate, with minutely pect- 275, fig. 24; Bruce, 1991b: 378–381, figs. 56–57, inate cutting edges distolaterally. Second

71 d-f; Müller, 1993: 11 (listed); Fransen, 1994a: pereiopods unequal; dactylus exceeding fixed 107, 111, fig. 2b; Bruce, 1996: 205, figs. 2, 3; Li, finger, distally Downloaded strongly from Brill hooked.com 06, with /24/2024 one12large:38:05 AM 2000: 12, fig. 12 (listed); Yamaguchi via& Open Holthuis Access,. This triangular is an open tooth access in article proximal distributed half, fixed under finger the terms of the CC-BY 4.0 license. https://creativecommons.org/licenses/by/4.0/ with 4-6 small teeth in proximal half of which distalmost largest; palm with oblong oval tympanal organ on entire medio-ventral surface. Ambulatory dactyli simple; corpus with carinated ventral margin with small triangular subdistal denticle, distodorsally transversely wrinckled; unguis strongly hooked, not flattened, with transverse rows of minute scale-like spinules dorsally. Uropod with protopodite posterolaterally acute, exopod with small mobile distolateral spine.

Remarks. The variability in the development of the accessory tooth on the dactylus of the ambulatory pereiopods ranging from its absence in Kemp’s material to well developed in the type of Anchistus misakiensis Yokoya, 1936 and the illustrated specimen from New 1977b), Keppel Bay and Magnetic Island, Caledonia (indicated as Lagon, stn. CC 147) of Australia (Bruce, 1983b); New Caledonia Bruce (1991b) attributes to the opinion that A. (Bruce, 1991a,1991b,1996).Now newly recorded misakiensis is conspecific with Tympanicheles from Ambon, Indonesia, and Vanuatu.

pectinis . In the two specimens studied the

accessory tooth was absent or minute (fig.

38I). Differences in the male first endopod as Key to the genera of the ‘ Anchistus’ clade

noted by Bruce (1991b: 381) could be due to

different developmental stages involved. In 1. Unguis of ambulatory pereiopods with the present male (pocl. 3.90 mm) 13 spinules few scattered minute scales dorsally......2 are present on the proximal median margin. – Unguis of ambulatory pereiopods with

In the male specimen (pocl. 2.7 mm) from dense cover of minute scales dorsally...4 New Caledonia (Bruce, 1991b: fig. 56k) only 2. Ambulatory dactyli without obtuse comsix spinules are present while in a large male pressed ventral process.............................. 3 specimen (pocl. between 3.65 and 5.88 mm) – Ambulatory dactyli with obtuse comfrom Japan ( Suzuki, 1971: fig. 7.4) at least 10 pressed ventral process.................. Dasella spinules are present. 3. Ambulatory pereiopods biunguiculate; Colour (figs. 36C, D). “The body is translu- mobile hepatic spine present.................. cent. A number of reddish circular marking, .................................................... Paranchistus which are encircled with purplish colouration, – Ambulatory pereiopods simple; hepatic are found on the body and appendages; these spine absent....................................... Ensiger marks are variable in size and number. The 4. Rostrum laterally compressed, distally distal portion of the antennular and anten- acute, with teeth in distal part.................5 nal peduncles, the scaphocerite, the second – Rostrum broad, swollen, distally trunchela, the telson and the uropod is purplish in cate, without teeth............... Neoanchistus colour.” ( Suzuki, 1971: 13, textfig. 5, pl. 2). 5. Palm of second chelipeds with tympanal Hosts. Reported from Pectinidae : organ...............................................................6 Bractechlamys vexillum (Reeve, 1853) (Bruce, – Palm of second chelipeds without ven- 1996, as Semipallium vexillum (Reeve, 1853)) ; tral tympanal organ; mobile hepatic Mimachlamys crassicostata (G. B. Sowerby II, spine present........... Polkamenes gen. nov.

1842) (Hayashi, 2002, 2006, as Mimachlamys 6. Tympanal organ on palm of second nobilis (Reeve, 1852) ; Pecten albicans (Schröter, cheliped oblong; dorso-distal part of 1802) (Fujino & Miyake, 1967; Suzuki, 1971; corpus of ambulatory dactyli wrinkled Hayashi, 2002, 2006, as P. (Notovola) albi- ............................... Tympanicheles gen. nov.

cans); Pecten sp. (Kemp, 1925); Yilistrum bal- – Tympanal organ on palm of second loti (Bernardi, 1861) ( Yokoya, 1936; Suzuki, 1971; cheliped restricted to proximal part Bruce, 1977b, 1983b, 1991a, 1991b, 1996; Hayashi, of palm; dorso-distal part of corpus of 2002, 2006, as Amusium ‘ japonicum’ balloti ambulatory dactyli glabrous..... Anchistus (Bernardi, 1861)) ; Pectinid bivalve (Bruce,

1976d). Also one record from an unidentified

bivalve (Bruce, 1977b). Discussion

Distribution. Nicobar Islands (Kemp, 1925);

Zanzibar (Bruce, 1976d); Japan ( Yokoya, 1936; Outgroup placements

Fujino & Miyake, 1967; Suzuki, 1971; Hayashi, All outgroup Downloaded species from, except Brill.com Cuapetes 06/24/2024 tenuipes 12:38:05 AM 2002, 2006); Townsville, Australia via Open(Bruce Access,. This and is Palaemonella an open access rotumana article distributed, are part under of ‘Clade the terms of the CC-BY 4.0 license. https://creativecommons.org/licenses/by/4.0/ 5’ recovered by Horká et al. (2016), which has Allopontonia Bruce, 1972 , which appear to be

a topology that only partly matches the cur- somewhat related (Chow et al., 2021)).

rent results. The coral-associated Actinimenes

inornatus, A. ornatus , and Periclimenes kempi Uncertain placements and polytomies

split off as a separate clade in the phyloge- Although most ingroup clades seem to be

nies resulting from the concatenated and TE recovered in a similar way across the various alignment (figs. 1, 2). This seems to be con- analysis methods, there are a few species with gruent with the results of Horká et al. (2016). aberrant placements. Paranchistus armatus, Here , the present ingroup (consisting of three the type species of Paranchistus , was recovspecies) is a sister of a clade comprised of ered based on the concatenated alignment as a Lipkemenes lanipes , Periclimenes colemani , part of a large polytomy, including various out-

and Zenopontonia rex . In our analyses (TE and groups (see above) and the rest of the ingroup, concatenated datasets), these outgroups seem suggesting a basal placement of the species. In

to be placed elsewhere: L. lanipes is placed the tree based on the TE-approach, the species basal to a clade including both P. colemani , Z. is placed next to a clade including Ensiger cus-

rex , as well as the ingroup (figs. 1, 2); and Z. rex tos and E. custoides , with Dasella being placed

and P.colemani seem to be clustering together basal to this group. The morphological fea-

in all analyses. In the phylogeny reconstruc- tures placed P.armatus with the two species of

tion resulting from the concatenated data- Ensiger . Although this difference in placement

set, Z. soror clusters together with Z. rex and is not interfering with the taxonomic decisions P. colemani , while in the TE-analysis Z. soror made in this study, a more basal placement of

is placed basally to these species and the this species in the TE-approach would probaingroup (figs. 1, 2). Horká et al. (2016) found bly directly influence the low ML-value of the Z. soror to be basal to a clade containing all character state reconstruction of the common three before-mentioned outgroups, as well as ancestor (now p = 0.508; fig. 3) to be higher in

the ingroup. In Chow et al. (2021), the place- favour of a bivalve host.

ment of these species is also different, plac- The genera Anchistus and Paranchistus

ing Z. soror basal to the current ingroup, and were in need of reappraisal after being recov- L. lanipes basal to Z. rex and P. colemani (sim- ered as polyphyletic in the phylogeny recon-

ilar to Horká et al., 2016). One has to keep in struction (figs. 1, 2). Two species, however, mind that the present phylogeny reconstruc- were also represented by more than one lintions are based on two molecular markers, eage. This is only the case in the phylogenetic compared to four and eight markers used tree resulting from the concatenated molec-

by Horká et al. (2016) and Chow et al. (2021), ular dataset (fig. 1). First, Anchistus demani is which partially use the same sequences. represented by three specimens, two of which

In short, the systematic position of the cluster together (KP759379/KP725496 and selected outgroups remains elusive. In order JX185707) next to A. miersi . The other spec-

to explore the systematic position of all mem- imen, making the placement of the species

bers of our outgroup genera, a large-scale paraphyletic, is only represented by a newly systematic review, combining molecular and acquired 16S barcode ( RMNH. CRUS.D.48354), morphological data, is needed, especially and is placed basal to a clade comprising the involving Zenopontonia and related echino- other specimens of A. demani and A. miersi .

derm-associated genera (e.g., Sandimenes Li , This deviating placement is probably caused 2009, various members of Periclimenes , and by the limited Downloaded phylogenetic from Brill.com information 06/24/2024 12of:38:05 AM via Open Access. This is an open access article distributed under the terms of the CC-BY 4.0 license. https://creativecommons.org/licenses/by/4.0/ 16S, when used on its own (supplementary limited morphological data, and can only be fig. S1). In a similar light, the specimens repre- resolved by increasing the number of morphosenting Ensiger custos are grouping together, logical characters (Neumann et al., 2021). The but in two lineages. Two specimens group present polytomous branch splits into three together (KF738360 and KJ584120), while monophyletic clades: Anchistus , Neoanchistus , one other specimen is placed basal to this and Tympanicheles . Considering the molecclade ( RMNH. CRUS.D.57963), in both the tree ular trees (fig. 1, supplementary figs. S1–S3), resulting from the TE-approach (fig. 2), as well the placement of the three Anchistus species

as the tree resulting from the concatenated and T. ornatus seems to be fixed. The placemolecular alignment (fig. 1). Again, this could ment of T. pectinis inside Tympanicheles , be the result from the first two specimens only solely based on morphology, is tentative but being represented by 16S (supplementary fig. convincing. What remains is the placement S1), while the latter is represented by both of Neoanchistus within this group; both spemolecular markers. cies’ placement is based solely on morpholog- Horká et al.(2016) included only three of the ical features, possibly causing the polytomy.

present ingroup species: Dasella herdmaniae Whether Neoanchistus should be placed more was recovered in a basal position to Ensiger basal to Tympanicheles , or closer to Anchistus , custoides and Polkamenes pycnodontae (sensu remains unclear. Acquiring DNA extractions Horká et al., 2016), which complies to the from fresh specimens of Neoanchistus may current results. Similarly, Chow et al. (2021) solve this polytomy in a future study (A.Anker, recovered a clade including E. custoides and pers. comm.).

E. custos as a sister group to a clade including

P. pycnodontae and an unidentified species of Host switching

Paranchistus (probably part of Polkamenes ). An ancestral character state reconstruction This clade was, however, recovered as a sis- of the host affiliations resulted in ML probater to a branch containing D. herdmaniae and bilities for all nodes in the tree based on the Tympanicheles ornatus (Chow et al., 2021) . In TE-approach (fig. 3, supplementary fig. S4, our results, Dasella is always placed basally to supplementary table S3). Most interesting are the other mentioned species and T. ornatus is nodes where host switching might have hapgrouped together with the more derived spe- pened, and the host affiliation of the common cies of the ingroup (figs. 1, 2). The 16S barcode ancestor of the entire ingroup. In this case, a

of T. ornatus featured in Chow et al. (2021) is single host switch seemed to have happened the same as the one used in the current study, in the split from the common ancestor of the only here it is accompanied by a COI-barcode entire ingroup.In contrast to the other bivalveprovided by Aznar-Cormano et al. (2015). and ascidian-associated palaemonid clade (de Based on this, we believe that a more derived Gier et al., 2022), which appeared to have an placement of T. ornatus is correct. ascidian-associated ancestor, the common The phylogenetic trees that are solely based ancestor of the present ingroup was recovered on molecular data (fig. 1, supplementary as a bivalve-associated species, albeit with a figs. S1–S3) do not pose problems regarding low probability (fig. 3, supplementary fig. S4, polytomies. The tree resulting from combined supplementary table S3). Consequently, there molecular and morphological data, however, seems to have been one host switch from a shows one major polytomous split (fig. 2). bivalve to an ascidian in the branch includ- Polytomies are common in trees based on ing Dasella Downloaded. Horká from et al. (Brill.com 2016) were 06/24/2024 not able 12:38:05 AM via Open Access. This is an open access article distributed under the terms of the CC-BY 4.0 license. https://creativecommons.org/licenses/by/4.0/ to recover an ancestral host affiliation from have a similar spotted pattern, although two

the currently studied clade, and Chow et al. groups can be distinguished by their spots (2021) recovered the ancestral node of the and colouration (Bruce, 1972a; supplemenentire clade as a bivalve-associated species. tary appendix S1, supplementary table S2).

This node also included various species of the Minute white, red or yellow spots can be found echinoderm-associated genera Zenopontonia , in Dasella , Tympanicheles ornatus and both Sandimenes , Lipkemenes Bruce & Okuno, 2010 , species of Ensiger . In addition, minute spots

etc., and one free-living species, Periclimenes can also be found in the more closely related josephi Li, 2008 . outgroups, like Zenopontonia , Lipkemenes

and Periclimenes kempi , confirming the basal Phylogenetical significance of placement of some of the before-mentioned

morphological features species. Larger spots, usually a combination

Comparing phylogenetic trees resulting from of both blue and red (or orange) chromato-

the concatenated molecular alignment and phores, can be found in all other ingroup spe-

the morphological character state dataset cies: Anchistus australis , A. demani , A. miersi , could give us an insight in which morpho- Polkamenes pycnodontae , P. spondylis , and logical characters are more phylogenetically additionally to some extent in Tympanicheles significant than others. Previous authors have ornatus . The colour pattern of T. ornatus made some remarks on the subject, which resembles the minute spots of E. custoides

will be discussed below, together with new (figs. 20, 36), although closer inspection (e.g., findings. MNHN-IU-2010-4971) reveals that the yellow Bruce (1975b, 1976a) mentioned that the spots are larger and that additional orange “irrelevant” colour patterns of Anchistus spots are present. Projection of the colour pat-

(s.l.) have little specific relevance due to the terns on the molecular trees show that larger shrimps endosymbiotic lifestyle and their spots are derived characters and that they are monogamous mating system (Neo et al., 2015). only found in symbionts of larger bivalves.

This implies that the conspicuous colourations The presence of minutely pectinate cutting

have no obvious advantages or disadvantages edges of spatulate to subspatulate first cheli- (Bruce, 1976a). Consequently, there would be peds is considered to be a definitive charac-

no strong selection pressure for this character. ter for the studied ingroup genera (e.g., Bruce,

The whole ingroup clade can be seen exhibit- 1995; Fransen & Van der Meij, 2010), but was

ing a spotted (or polka-dotted) colour pattern also found in the related outgroup species

(figs. 4, 17, 20, 33, 36; Bruce, 1972a), with the Lipkemenes , Zenopontonia , Actinimenes and exception of Neoanchistus (fig. 28; A. Anker, both Periclimenes colemani and P. kempi

pers. comm.), and to some extent A. pectinis . (supplementary appendix S1, supplemen-

The colour pattern of Neoanchistus and A. tary table S2). A quick investigation also pectinis somewhat resembles that of other proves that Sandimenes and various memendosymbiotic shrimps, such as those of the bers of Periclimenes and Allopontonia (which genus Platypontonia Bruce, 1968 (see de Gier et were proven to be somewhat related to the

al., 2022). Why these species exhibit a distinct currently studied ingroup by Chow et al., pattern of larger red spots is unknown, but this 2021) also have subspatulate first chelipeds,

may be concordant with the species’ aberrant although the associated minutely pectinate

host choice, consisting of pectinid and cardiid cutting edges cannot be seen in the published bivalves. All other ingroup species appear to illustrations Downloaded (Bruce,from 1971Brill, 1972.com b, 06 1995 /24/2024, 2010 12b:38;:05 AM via Open Access. This is an open access article distributed under the terms of the CC-BY 4.0 license. https://creativecommons.org/licenses/by/4.0/ Li, 2009; Rauch et al., 2019), except for the 39E, F). Although the systematic position of T.

illustrations of P. josephi , which show “a dis- pectinis , A. gravieri , and Neoanchistus cannot tal row of pectinate short setae” (Li, 2008b). be confirmed with the current molecular data, Closer examination of collection material is T. ornatus has a more basal placement than A.

needed in order to see if minutely pectinate australis , A. demani , and A. miersi (fig. 1). Thus, cutting edges are also present in related ecto- it seems that a dinstinctly outlined smaller symbiotic species. The potential function of organ evolved from a less clearly delimited the (sub)spatulate cheliped shape is currently larger organ. More morphological and histounknown, although it may be linked to the logical work is needed in order to see if the dietary habits of the shrimp. Species could surface of the tympanal organ is homologous be using the minor chelipeds to harvest and to the outer surface of the chelae of other spedigest their hosts’ mucus and ‘steal’ food par- cies in the ingroup. The potential function of ticles, such as in E. custos (Johnson & Liang, the organ is unknown, and a potential link 1966), although the cheliped morphology of to the host choice could also not be found E. custos and E. custoides is quite aberrant (with T. ornatus occupying the same hosts (e.g., fig. 27A). Interestingly, De Grave et al. as Ensiger ). Similar drum-looking organs are (2021) concluded that representatives of the used for interspecies communication, host currently studied clade, Actinimenes ornatus , detection, or predator avoidance in various Zenopontonia soror , and Anchistus miersi , groups of insects (described as a ‘tympanum’: were facultative or obligate parasites, feed- Yack, 2004; Strauss & Lakes-Harlan, 2014) and ing from the hosts’ tissue. Comparatively, as a ‘directional air motion sensor’ in the spithe limited information on the bivalve-asso- der genus Hortipes Bosselaers & Ledoux, 1998 ciated species from the other endosymbiotic ( Ramírez, 2014). Perhaps the functionality of palaemonid clade (de Gier et al., 2022) were the drum-like organ for shrimps can be found found to feed from host mucus strands and/ in the field of bioacoustics (e.g., Lovell et al.,

or pseudofeces (Kennedy et al., 2001; Ashelby 2005). In crustaceans, similarly looking struc-

et al., 2015). In addition, Pontonia pinnophylax tures called ‘gas windows’ or also ‘tympana’ (Otto, 1821) was found to be feeding from sim- can be seen in the last walking leg pairs and ilar carbon sources as its host (De Grave et al., sometimes thoracic sternites of semi-ter- 2021), making it a ‘true’ commensal symbiont. restrial sand bubbler crabs (Brachyura: The flat oval-shaped ‘tympanal’ organ Dotillidae ), which they use for osmoregulafound on the underside of the second cheli- tion (Matsumasa et al., 2001). In various other peds was first described and figured by semi-terrestrial crabs, small ‘window-like’ Monod (1976a) for Anchistus miersi . This organs called ‘Barth’s myochordotonal structure can be found in various members organs’ can be found, which play a role in

of the ingroup: Anchistus australis , A. dem- intraspecific communication, being able to ani, A. miersi , and A. gravieri bear major claws respond to vibrations transmitted through with small, distinct tympanal organs (fig. 39B, the exoskeleton from the substrate and the D–F), while both species of Tympanicheles , air (Davie et al., 2015). In male Australian Neoanchistus nasalis , and possibly N. cardi- freshwater crayfish Cherax quadricarinatus odytes , bear major claws with larger, less-pro- (von Martens, 1868), sexually dimorphic ‘red nounced tympanal organs (fig. 39A, C). In A. patches’ on the propodus of the major claws’ australis , the ‘veil’ is much thinner than the can be found. The function of this uncalsurrounding rim and the rest of the claw (fig. cified structure Downloaded from was reviewed Brill.com by 06/24/2024 Karplus 12:38:05 AM via Open Access. This is an open access article distributed under the terms of the CC-BY 4.0 license. https://creativecommons.org/licenses/by/4.0/ et al. (2003) and hypothesized to be a sensory (Fransen, 2002; de Gier & Fransen, 2018). In organ, possibly used in intraspecific social addition, both species of Ensiger have no interactions. Various species of amphipods accessory teeth whatsoever (figs. 22E, 27D).

within the Bogidiellidae , Niphargidae , and Higher up in the tree (see fig. 1), Polkamenes Ingolfiellidae exhibit ‘windows’ in the cuticle pycnodontae (as well as P. spondylis and P.

of the pereiopods, pleon, and pereion, called nobilii ) exhibit a strong accessory tooth (figs.

‘lentiform organs’ ( Pretus & Stock, 1990). 34K, L, 35S), while Tympanicheles ornatus has

As for now, the function of these structures just one minor denticle (fig. 37H). Similar to T.

remains unknown (Vonk & Schram, 2003). ornatus , T. pectinis , A. gravieri , as well as both

In addition, a semi-transparent structure species of Neoanchistus (although the struc-

can be found on the dorsal sides of the heads tures on the illustrations of N. cardiodytes of

of ‘mountain shrimp’ ( Anaspidacea ) in the Bruce (1975a) are difficult to identify) bear genus Allanaspides Swain, Wilson, Hickman none or one small denticle on the flexor mar-

& Ong, 1970, which resembles the structures gin (fig. 29E). The elusive T. liui also has no found in amphipods, mentioned above (Vonk accessory teeth (Li et al., 2004). Interestingly,

& Schram, 2003). the species in the more derived clade of

The walking legs’ (third to fifth pair of Anchistus , comprised of A. australis , A. dempereiopods’) dactyli bear many of the char- ani, and A. miersi , show a presence of a larger acteristic features used for species and gen- accessory tooth, most apparent in A. miersi

era identifications within the present shrimp (figs. 5L, M, 6J, 16G). In contrast, the microclade (Fransen, 2002). In most cases, the scopic spinules which can be found on the dactyli are well-illustrated in taxonomic liter- dorsal surface of the unguis, can exclusively ature, revealing the wide range of morpholog- be found in the more derived lineages, and is

ical ornamentations and setation patterns on not found in Ensiger and Dasella . Paranchistus

the corpus and unguis. Although homology armatus exhibits a few microspinules on the

and sequential evolution in the structures unguis, although preliminary SEM-images

are sometimes hard to dinstinguish (supple- prove that these structures are not homolomentary appendix S1; supplementary table gous (de Gier et al., unpublished observation).

S2), some characters give the impression to

be unique to bivalve- and ascidian-associated Sexual dimorphic and ontogenetic

endosymbiotic shrimp (Fransen, 1994a, 2002; characters

de Gier & Fransen, 2018; de Gier et al., 2022). To keep the dataset uniform, mostly female However, Chow et al. (2021) showed that the ovigerous specimens were used to score overall shape of a ‘biunguiculate’ dactylus has almost all of the character states (with the evolved many times in both endo- and ecto- exception of the exclusively male features), symbionts, and bears no phylogenetic signal both regarding data from the literature as

on a higher taxonomic level. Similarly, in the from collected material (supplementary present clade, traces of convergent evolu- appendix S1, supplementary table S2). Since

tion can be found in the accessory teeth on the species in the ingroup are considered to

the flexor margin of the dactylar corpus: all exhibit reverse sexual dimorphism (Baeza species of Dasella show a blunt, distoventral et al., 2013), the morphological characters process (fig. 19D), while Paranchistus armatus of the bigger female specimens are easier

has a somewhat sharper tooth (fig. 30J; almost to score. The differences between male and resembling species of Pontonia and Odontonia female specimens Downloaded can from Brill mostly.com be 06/24found /2024 12in:38:05 AM via Open Access. This is an open access article distributed under the terms of the CC-BY 4.0 license. https://creativecommons.org/licenses/by/4.0/ the abdominal region (male and female first a juvenile shrimp found in a boring mussel

and second pleopods, size of abdominal seg- ( Spengleria mytiloides (Lamarck, 1818) belongments), which has an inclusion that is limited ing to the Gastrochaenidae ; Fransen & Van

in the morphological analysis. The bigger size der Meij, 2010). The specimen is thought to be

of females may result in a stronger selection related to the species in the currently studied

for ecomorphological characters, due to the ingroup, and was described as an unknown limited size of their hosts’ internal cavities. species of Anchistus (Fransen & Van der Meij,

This link has not been studied in palaemonid 2010), although its juvenile characters could shrimp as for now, although there is strong not be included in the morphological analysis.

evidence for sex-related selection pressure in

other crustaceans with more extreme sexual

dimorphisms (e.g., where males visit seden- Conclusions

tary females in gall and pea crabs (Brachyura:

Cryptochiridae , Pinnotheridae ); Van der Meij, New distributions, hosts, and classification

2014; Vehof et al., 2016; de Gier & Becker, In this study, numerous new records of dis- 2020). Sexual dimorphism in the colouration tribution ranges and host affiliations are

of the chromatophores (dichromatism) was presented, summarized in table 2. In addirecently found in Anchistus miersi , with the tion, this study establishes three new genera,

male showing red spots, and the female blue and resulted in some additional taxonomic spots (Neo et al., 2015). This causes no prob- changes (table 2).

lems in our dataset (supplementary appendix

S1, supplementary table S2). Future perspectives

All chosen characters were picked based on The analyses revealed two genera to be poly-

their visibility in adult specimens, since juve- phyletic and that three genera needed to be

nile anatomy may differ a lot from their adult (re)erected to make a naturally sound classi-

form (Fransen, 1994a). In most cases, these fication. Various species of the present study juvenile characters are easily distinguisha- did not have any molecular data, causing

ble, such as a shorter rostrum and relatively various branches to have low support values larger eyes (Fransen, 2002, 2006). In extreme and their exact evolutionary placement to cases, juvenile characters may be completely remain uncertain. Similar to an earlier study reduced in the adult stages, and can provide (de Gier et al. 2022), additional molecular erroneous character states. Examples can data are needed for this clade to obtain bet-

be seen in the (articulate) hepatic spine of ter supported and resolved phylogeny recon- Paranchistus armatus , which is much larger in structions regarding host associations and juveniles than in adults (Bruce, 1975c). More ancestral state reconstructions, which would extreme examples can be found in the ‘biun- require more field observations and studies guiculate’ juvenile dactyli of the walking legs addressing the feeding habits, host choices,

of Tympanicheles pectinis , which grow away in and distribution ranges.

later stages (Bruce, 1996), and in the juvenile Some authors have suggested that the molrostrum morphology of Anchistus australis , lusc- and ascidian-associated shrimp clades which grows from an acute to a more rounded display various host-specific (eco)morphologshape (Fransen, 1994a). Some juvenile char- ical adaptations (Bruce, 1972a, 1976a; Fransen, acters result in unknown species-identifica- 1994a), although the statistical evidence for tions, as is the case for a single specimen of this is limited Downloaded (Dobson from et Brill al..,com 2014 06, 2016 /24/2024; Chow 12:38:05 AM via Open Access. This is an open access article distributed under the terms of the CC-BY 4.0 license. https://creativecommons.org/licenses/by/4.0/

Species Taxonomic New locality records New host records name changes

Anchistus A. australis – Indonesia: West Papua: Raja Ampat;

NE Kalimantan; Nanaka Is., Sulawesi .

Malaysia: Tioman Is. A. demani – Indonesia: N Sulawesi; NE Kalimantan; Hippopus hippopus ;

Red Sea; Palau; Philippines: Cebu strait. Tridacna deresa ;

Tridacna crocea

(confirmed after

Bruce, 1979a) A. gravieri – Vanuatu Tridacna sp. A . miersi – Micronesia: Caroline Islands. Indonesia: –

NW Java: Thousand Islands ; NE and S

Kalimantan; West Papua: Numfor; SW and

N Sulawesi; Mollucas: NW Seram. Dasella D. ansoni – – – D. brucei – – – D. – Vanuatu: Tutuba Is. Indonesia : N, SW, Polycarpa sp. ; herdmaniae and NE Sulawesi; Mollucas: Ambon; Phallusia sp.

Halmahera; NW Java: Thousand Islands ;

West Papua: Raja Ampat. Ensiger E. custoides Transferred Indonesia: Ambon; Kera Is. near Timor; Atrina pectinata

from Komodo; NW Java: Thousand Islands ;

Anchistus Bali; NE and SW Sulawesi, Halmahera; W

Papua: Raja Ampat; NE Kalimantan: Berau

Islands. Vanuatu. Philippines: Cebu strait. E. custos Transferred Indonesia: NW Java: Thousand Islands; Atrina pectinata

from Sunda Islands ; SW Sulawesi ; West Papua .

Anchistus Japan. Oman: Masicoh (?). Saudi Arabia. Neoanchistus N. – – – cardiodytes N. nasalis – – – Paranchistus P. armatus – Indonesia: Moluccas; Kei Islands, –

Tanimbar. Polkamenes gen. nov.

et al., 2021). Ecomorphological analyses using Eurlings (all Naturalis Biodiversity Center) are

new imaging methods (µCT-scanning, SEM), thanked for their assistance generating the combined with ecological and molecular DNA-barcodes which were used during this

data may help us to understand how exactly study.Collection managers Jeroen Goud, Bram

the currently studied species have adapted to van der Bijl, and Esther Dondorp (all Naturalis

their endosymbiotic way of life. Biodiversity Center) are thanked for their sup-

port during the examinations of the physical specimens. Bertie-Joan van Heuven and Dirk Acknowledgements van der Marel (both Naturalis Biodiversity Center) are thanked for their assistance man-

The authors would like to thank Prof. Dr. Bert ning the SEM and µCT scanner in the Naturalis W. Hoeksema (Naturalis Biodiversity Center, imaging facilities. Research interns Jurriaan Rijksuniversiteit Groningen), Dr. Ronald van den Oever en Pepijn Helleman are Vonk (Naturalis Biodiversity Center), and Dr. thanked for their help in the SEM lab, acquir- Jeroen Hubert (Universiteit Leiden) for their ing images of the dactyli and claws of various helpful comments and feedback during the species studied in this paper. Dr. Arthur Anker writing phase of this project. The lab-techni- (King Abdullah University of Science and cians Frank Stokvis, Roland Butôt, and Marcel Technology Downloaded, Saudi Arabia from) is Brill.com thanked 06/24/2024 for his 12:38:05 AM via Open Access. This is an open access article distributed under the terms of the CC-BY 4.0 license. https://creativecommons.org/licenses/by/4.0/ contributing colour pictures of Neoanchistus (chief scientist Dr. B.W. Hoeksema) and the cardiodytes and N. nasalis (fig. 28), and Jeroen RCO-LIPI (chief scientist Dr.Suharsono),as part Goud (Naturalis Biodiversity Center) for the of the NWO-LIPI East Kalimantan Programme.

identification of the Vasticardium host of The ‘Thousand Islands Expedition, September N. cardiodytes . In addition, two anonymous 2005’, organized by the RCO-LIPI and the NNM.

reviewers are thanked for their helpful com- The ‘ SANTO 2006 Global Biodiversity Survey’ ments and suggestions, which helped to was initiated by the Muséum national d’Hisimprove the manuscript. Mr. H.L Strack organ- toire naturelle, Paris ( MNHN), Institut de ized the ‘Rumphius Biohistorical Expedition’ Recherche pour le Développement ( IRD) and (1990) to Ambon, in cooperation with the ProNatura International. Dr. Philippe Bouchet Nationaal Natuurhistorisch Museum ( NNM; acted as expedition leader for the marine now Naturalis Biodiversity Center), Leiden, theme. The ‘2007 Raja Ampat expedition’ the Netherlands; Oceanological Institute, was organized by the NNM (at that time NCB Indonesian Institute of Sciences ( LIPI, now Naturalis) and RCO-LIPI, under the umbrella RCO-BRIN); and Universitas Pattimura, both of Ekspedisi Widya Nusantara (E-Win) of LIPI.

from Ambon, Indonesia. The R. V. ‘Tyro’ We are grateful to LIPI for granting research ‘Oceanic Reefs Expedition’ to the Seychelles permits and to Dr. B.W. Hoeksema for his (1992–1993) was organized by the NNM, and efforts in organizing the expedition. The the Netherlands Indian Ocean Program. Dr. ‘ Ternate expedition’ in the northern Moluccas

J. van der Land ( NNM) was chief-scientist was organised by the RCO-LIPI and the NNM

at these cruises. The ‘ Sulawesi expedition’ (at that time Naturalis Biodiversity Center).

(1994) was organized was organised by the This expedition was part of the Ekspedisi NNM in cooperation with the Hasanuddin Widya Nusantara project (E-Win expeditions), University at Makassar ( UNHAS) and the field with chief-scientists Dr. B.W. Hoeksema and station of the Oceanological Institute of LIPI Ir. M.I.Y. Tuti ( PPPO-LIPI). The ‘Semporna

at Bitung with chief-scientist Dr. J. van der Marine Ecological Expedition’ (2010) was Land. The ‘Maluku Expedition’ (1996) was jointly organized by WWF-Malaysia, Universiti organised by the NNM in cooperation with the Malaysia Sabah’s Borneo Marine Research Oceanological Institute of LIPI at Ambon with Institute, Naturalis Biodiversity Center and chief-scientist Dr. J. van der Land and counter- Universiti Malaya’s Institute of Biological part Dr. Dwi Listyo Rahayu ( LIPI). The ‘Cebu Sciences. Research permission was granted Strait’ Expedition (2000) by the NNM and the by Economic Planning Unit, Prime Minister’s University of San Carlos ( USC), Cebu, with Department, Economic Planning Unit Sabah, chief-scientists Dr. Bert W. Hoeksema, Dr. Sabah Parks and Department of Fisheries Thomas Heeger ( USC) and Dr. Filipina B. Sotto Sabah. Dr. B.W. Hoeksema and Ms. Zarinah ( USC). The ‘Lombok Strait’ Expedition (2001) Waheed acted as expedition leaders for the by the NNM, the Center of Oceanological biodiversity theme. The ‘Marine Biodiversity Research and Development ( PPPO) of LIPI in Workshop at Lembeh Strait’ (2012) was organ- Jakarta, and the World Wildlife Foundation ized by Universitas Sam Ratulangi ( UNSRAT in ( WWF) Wallacea project at Denpasar, with Manado) and the Bitung field station of the chief-scientists Dr. B.W. Hoeksema and Ir. M.I. Research Centre of Oceanography ( RCO-LIPI) Yosephine Tuti ( PPPO-LIPI). The 2003 ‘Berau in cooperation with Naturalis Biodiversity 2003’ expedition around the Berau Islands, Center. Prof. Dr. Suharsono, former director East Kalimantan was organised by the NNM of the Downloaded Research from Centre Brill.com for Oceanography 06/24/2024 12:38:05 AM via Open Access. This is an open access article distributed under the terms of the CC-BY 4.0 license. https://creativecommons.org/licenses/by/4.0/ ( PPO-LIPI), is acknowledged for his sup- relationships in Caridean decapods. Genetica, port. Prof. Dr. Markus T. Lasut (Universitas 143, 195–205. doi: 10.1007/s10709-014-9807-0. SamRatulangi), Dr. B.W. Hoeksema, and Ir. Baeza, J.A., Ritson-Williams, R. & Fuentes, M.S. Y. Tuti ( PPO-LIPI) acted as workshop coor- (2013) Sexual and mating system in a caridean dinators. Dr. Francesca Benzoni organized shrimp symbiotic with the winged pearl oyster the ‘Marine Biodiversity Workshop’ (2014) at in the Coral Triangle. J. Zool., 289, 172–181. the Marine Research and Higher Education doi:10.1111/j.1469-7998.2012.00974.x. Centre University of Milano-Bicocca, Barnard, K.H. (1955) Additions to the fauna-list Magoodhoo Island, Faafu Atoll, Republic of South African Crustacea and Pycnogonida. of Maldives. Prof. Mike Berumen (Red Sea Ann. S. Afr. Mus ., 43(1), 1–107. Research Center, King Abdullah University Barnard, K.H. (1958) Further additions to the of Science and Technology, KAUST), hosted crustacean fauna-list of Portuguese East Africa. a Naturalis field team under leadership of Mem. Museu Dr.Álvaro de Castro, 4, 3–23. Dr. B.W. Hoeksema in November 2014 off the Berggren, M. (1990) Dasella herdmaniae (Lebour) Red Sea coast of Thuwal, Saudi Arabia. ( Decapoda : Natantia: Pontoniinae ) from

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