identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
CA4F87BBB752FF97F3B1F6C6FED95C13.text	CA4F87BBB752FF97F3B1F6C6FED95C13.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Tomlinsonia mclaughlinae Williams & Boyko 2006	<div><p>Tomlinsonia mclaughlinae n. sp.</p><p>(Figs 1-6)</p><p>TYPE MATERIAL. — Philippines. Boracay, Rocky Beach, 11°57’N, 121°56’E, from shell of Drupella cornus (Röding, 1798) inhabited by ovigerous ♀ Calcinus gaimardii (H. Milne Edwards, 1848) (3.4 mm SL), 13.IV.1999, ovigerous ♀ holotype 8.15 mm max. length, 4.3 mm max. width (USNM 1084093), ♂ allotype 0.71 mm (USNM 1084094). — Puerto Galera, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=120.95&amp;materialsCitation.latitude=13.5" title="Search Plazi for locations around (long 120.95/lat 13.5)">Big Lalaguna Beach</a>, 13°30’N, 120°57’E, from shell of Peristernia incarnata (Kiener, 1840) inhabited by ovigerous ♀ C. gaimardii (3.8 mm SL), 21.VII.1997, 1 ♀ paratype with Hemioniscus (USNM 1084095) ; from shell of Drupella cornus inhabited by ovigerous ♀ Calcinus minutus Buitendijk, 1937, 31.VII.1997, 1 ♀ paratype (USNM 1084096) . — Puerto Galera, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=120.933334&amp;materialsCitation.latitude=13.5" title="Search Plazi for locations around (long 120.933334/lat 13.5)">Coco Beach</a>, 13°30’N, 120°56’E, from shell of D. cornus inhabited by ovigerous ♀ C. gaimardii, 12.I.1999, 1 ♀ paratype (USNM 1084165; mantle on one SEM stub, cirri on separate SEM stub) . — Puerto Galera, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=120.88333&amp;materialsCitation.latitude=13.483334" title="Search Plazi for locations around (long 120.88333/lat 13.483334)">Bayanan</a> and <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=120.88333&amp;materialsCitation.latitude=13.483334" title="Search Plazi for locations around (long 120.88333/lat 13.483334)">Haligi Beaches</a>, 13°29’N, 120°53’E, from shell of D. cornus inhabited by ovigerous ♀ C. gaimardii, 13.I.1999, 1 ♀ paratype (ZRC 2006.0001) . — Bataan, Mabayo, 14°44’N, 120°16’E, from shell of Drupa grossularia Röding, 1798 inhabited by ovigerous ♀ C. minutus (2.85 mm SL), 21.II.1999, 1 ovigerous ♀ paratype (ZRC 2006.0002) ; from unidentified gastropod shell inhabited by ♀ C. minutus (2.25 mm SL), 21.II.1999, 1 ♀ paratype (ZRC 2006.0003) ; from shell of Morula granulata (Duclos, 1832) inhabited by ♀ C. minutus (2.2 mm SL), 21.II.1999, 1 ♀ paratype (ZRC 2006.0004); from shell of D. grossularia inhabited by ♀ C. minutus (3.0 mm SL), 21.II.1999, 1 ♀ paratype (USNM 1084097); from shell of Cantharus undosus (Linnaeus, 1758) inhabited by ♀ C. gaimardii (3.02 mm SL), 21.II.1999, 1 ♀ paratype (ZRC 2006.0005); from unidentified gastropod shell inhabited by ovigerous ♀ C. minutus (2.18 mm SL), 21.II.1999, 1 ♀ paratype (USNM 1084098); from unidentified gastropod shell inhabited by ovigerous ♀ C. minutus (2.82 mm SL), 21.II.1999, 1 ♀ paratype (USNM 1084099); from unidentified gastropod shell inhabited by ♀ C. gaimardii (1.69 mm SL), 21.II.1999, 1 ♀ paratype (USNM 1084100); from unidentified gastropod shell inhabited by ♀ C. gaimardii (2.34 mm SL), 21.II.1999, 1 ♀ paratype (USNM 1084101); from unidentified gastropod shell inhabited by ♀ C. minutus (2.98 mm SL), 21.II.1999, 1 ♀ paratype (USNM 1084102); from unidentified gastropod shell inhabited by ovigerous ♀ C. minutus (2.42 mm SL), 21.II.1999, 1 ♀ paratype, 1 ♂ paratype (USNM 1084103); from unidentified gastropod shell inhabited by ovigerous ♀ C. gaimardii (3.31 mm SL), 21.II.1999, 1 ♀ paratype (USNM 1084104; mouthparts on SEM stub); from unidentified gastropod shell inhabited by ovigerous ♀ C. minutus (2.9 mm SL), 21.II.1999, 1 ovigerous ♀ paratype 5.90 mm (USNM 1084105); from unidentified gastropod shell inhabited by ovigerous ♀ C. minutus (2.54 mm SL), 21.II.1999, 1 ♀ paratype 5.9 mm max. length (USNM 1084106; mouthparts on SEM stub); from unidentified gastropod shell inhabited by ♀ C. minutus (2.58 mm SL), 21.II.1999, 1 ♀ paratype 3.69 mm max. length, 2.07 mm max. width, 1 ♂ paratype 0.50 mm (USNM 1084107). — <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=121.933334&amp;materialsCitation.latitude=11.95" title="Search Plazi for locations around (long 121.933334/lat 11.95)">Boracay</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=121.933334&amp;materialsCitation.latitude=11.95" title="Search Plazi for locations around (long 121.933334/lat 11.95)">Rocky Beach</a>, 11°57’N, 121°56’E, from shell of D. cornus inhabited by ♂ C. gaimardii (2.9 mm SL), 12.IV.1999, 1 ♀ paratype (USNM 1084108), 1 ♂ paratype 0.68 mm (USNM 1084109); from shell of D. cornus inhabited by ♀ C. gaimardii (3.85 mm SL), 12.IV.1999, 1 ♀ paratype (USNM 1084110); from shell of Drupella rugosa (Born, 1778) inhabited by ovigerous ♀ C. gaimardii (3.5 mm SL), 15.IV.1999, 1 ♀ paratype (USNM 1084111); from shell of D. rugosa inhabited by ♀ C. minutus (3.85 mm SL), 15.IV.1999, 1 ♀ paratype (USNM 1084112); from shell of D. cornus inhabited by ovigerous ♀ C. gaimardii (3.7 mm SL), 15.IV.1999, 1 ovigerous ♀ paratype (USNM 1084113). — Bataan, Morong, 14°41’N, 120°16’E, from shell of C. undosus inhabited by ♀ C. gaimardii, 25.IV.1999, 1 ♀ paratype with Hemioniscus (USNM 1084114); from unidentified gastropod shell inhabited by ♀ C. minutus (3.0 mm SL), 25.IV.1999, 1 ♀ paratype (USNM 1084115); from unidentified gastropod shell inhabited by ♀ C. minutus (3.25 mm SL), 25.IV.1999, 1 ♀ paratype (USNM 1084116); from shell of D. grossularia inhabited by ovigerous ♀ C. gaimardii (3.6 mm SL), 25.IV.1999, 1 ♀ paratype (USNM 1084117); from shell of C. undosus inhabited by ♀ C. gaimardii (5.2 mm SL), 25.IV.1999, 1 ♀ paratype with Hemioniscus (USNM 1084118).</p><p>ADDITIONAL MATERIAL EXAMINED. — Philippines. Bataan, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=120.26667&amp;materialsCitation.latitude=14.733334" title="Search Plazi for locations around (long 120.26667/lat 14.733334)">Morong</a>, 14°41’N, 120°16’E, from unidentified gastropod shell inhabited by ♀ Calcinus latens (Randall, 1839) (2.2 mm SL), 28.II.1999, ♀ (JDW personal collection). — Bataan, Mabayo, 14°44’N, 120°16’E, from unidentified gastropod shell inhabited by ovigerous ♀ C. minutus (1.75 mm SL), 21.II.1999, 1 ovigerous ♀ (JDW personal collection).</p><p>ETYMOLOGY. — In honor of Patsy A. McLaughlin for her many years in pursuit of new and interesting taxa of hermit crabs, as well as for her gracious help in identifying specimens for those in need, including both of the present authors – for which we are grateful.</p><p>DISTRIBUTION. — Found in shells of Calcinus spp. hermit crabs from Aklan, Bataan, and Oriental Mindoro provinces in the central Philippines; shallow subtidally (&lt;5 m).</p><p>DIAGNOSIS. — Female: mantle laterally compressed, muscular, oriented ventrally to aperture, conforming to spiral of the columella, body bluntly rounded at distal end toward apex or with short lobes; chitinous attachment disc with minute tubercles. Left opercular bar thin with notch on ventral side, dorsal side with shallow, rounded indentation; right opercular bar thick with flattened distal margin bordered by raised ridges along length. Opercular bar outer surfaces with dense horizontal rows of variously shaped denticules. Labrum ends of distal margin strongly produced, one broad and rounded distally and one narrow and tapered distally, medial margin subquadrate. Mandible strongly recurved. Maxilla I with acute upper spine and smaller proximoventral spine. Maxilla II an ovate plate with sparsely distributed stub setae. Cirrus I endopod and exopod of about 1/2 length protopod. Three pairs of terminal cirri uniramous with four segments; basal segment elongate, tubular, reaching to near end of second segment; second segment approximately as broad as basal segment but about 1/2 as long with protuberant “cushion” on distal inner surface, ridges on cushion with numerous regular rows of blade-like denticules; third segment narrower than basal or second, approximately as long as basal segment; fourth segment narrower than third, tapering distally, terminating in pair of bifid hooks and superior short recurved spine. Male: if present, one male per female, attached below mantle aperture. Form of stout boot with dorsal surface nearly straight, anteroventral surface domed and separated from posterodorsal surface by rounded concave area of nearly 90°, all corners rounded, ventral and posterior extensions of body subequal in length, anterolateral corner low.</p><p>DESCRIPTION</p><p>Female (Figs 1; 2A-C, E; 3-6) Maximal length 8.15 mm, maximal width (holotype) 4.3 mm (Fig. 1A). Mantle (sensu Kolbasov &amp; Newman 2005) laterally compressed, perpendicular to the surface of gastropod shell (Figs 1; 2A, C). Flattened part of mantle (“disc”) dorsoventrally compressed, parallel to surface of gastropod shell (Figs 1; 2A, B). Chitinous attachment disc regularly rounded, oval, or irregular, depending on size of barnacle; in some larger specimens disc extends as a spiral covering the barnacle body that is exposed in this region, conforming to the shape of the shell (Fig. 2B, C), chitinous disc covered with minute tubercles. Disc lying in a plane perpendicular to sides of mantle and attaching upper part of animal with cement to wall of burrow along columella in position that orifice leads directly into mantle cavity (Fig. 2A). Aperture on columella of shell approximately straight to slightly curved, tapered slit-like opening 5.5 mm in length and 0.9 mm in width at widest point in holotype, tapering to thin extension (peduncular slit) oriented toward apex of shell (Fig. 2A, E). Mantle muscular, oriented ventrally to aperture, extending toward apex of shell, conforming to spiral of the columella, in larger specimens extending into the shell whorls, body bluntly rounded at distal end toward apex or with short lobes corresponding to shell whorls in larger individuals, burrow position shown by striae in thin layer of shell overlaying barnacle (Figs 1B, C; 2A). Orifice of mantle a narrow fissure-like opening about 1/4 length of barnacle, tapering to thin opening on ventral end with round opening at dorsal end (Fig. 2C). Left opercular bar thin with abrupt rounded end on ventral side giving the appearance of a notch, dorsal side with shallow, rounded indentation preceding connection with right opercular bar (Figs 1C; 3J, K); right opercular bar thick with flattened distal margin bordered by raised ridges along length (Figs 1A; 2C; 3A, B); right opercular bar approximately 1/3 longer than left opercular bar. Left opercular bar outer surface with dense horizontal rows of ovate denticules (some with bifid or trifid tips) (Figs 3J, L; 4E), row of long simple setae on upper margin (Fig. 3J, L), distal portion of denticule row presenting sparser area of short multifid star-shaped denticules and fewer, shorter simple setae (Fig. 3J, M); sloping region of mantle distal to notch in opercular bar with sparse area of multifid denticules similar to those seen in Figure 3M but with slightly longer shafts (Fig. 3J, N). Left opercular bar inner surface with small inner ridge bordered by dense rows of variously simple to multifid finger-like denticules, tapering towards region of mouth cirri (Fig. 3K, P), small area distal to ridge with sparse field of star-shaped multifid short denticules (Fig. 3K, O), small field of long plumose setae ventral to inner ridge (Fig. 3K; similar to those in Figure 3H on inner surface of right opercular bar), area dorsal to mouth cirri with sparse field of elongate variously simple to multifid denticules and few long simple setae (Fig. 3K, Q). Right opercular bar outer surface with sparse irregular field of simple tear-drop-shaped denticules ventral to dorsal margin and long simple setae on edge (Fig. 3A, D), below opercular bar edge dense rows of simple tear-drop-shaped denticules (Fig. 3A, C), denticules appear flat viewed head-on but are curved in lateral view (Fig. 3E). Right opercular bar inner surface with strong ridge bordered by dense rows of large multifid, foot-shaped denticules (with up to seven denticular blades), field of denticules tapering towards mouth cirri (Figs 3B, G; 4D), area dorsal to mouth cirri region with sparse rows of ovate denticules (Fig. 3B, F), distal margin with thin sparse rows of ovate denticules, mostly with bifid tips and interspersed with few short simple setae (Fig. 3B, I), area ventral to inner ridge with field of long plumose setae (Fig. 3B, H).</p><p>Labrum very large, inner edge partially covered by minute denticules and row of long setae proximal to margin along whole length of labrum, ends of distal margin of labrum unequal but both strongly produced, one broad and rounded distally and one narrow and tapered distally (Figs 5C; 6B), both with groups of typically three short simple setae (Fig. 5C inset), medial margin subquadrate (Figs 5C; 6B). Mandible strongly recurved, distally as acute projection, no accessory spines (Figs 5D; 6C). Maxilla I smaller than mandible with acute upper spine and smaller proximoventral spine (Figs 5E; 6C), with rows of five or six tapered denticules along inner margin (Fig. 6E). Maxilla II (maxillule) an ovate plate with rows of tiny setae on mesiodistal edge and a strong stout seta near mediomesial edge (Figs 5E; 6C), with sparsely distributed stub setae, more abundant at the base (Fig. 6D).</p><p>Four pairs of cirri. Cirrus I (mouth cirrus) biramous; protopod elongate and tubular, naked; endopod and exopod of about 1/2 length of protopod; exopod bearing more long plumose setae on surface and tip than endopod (Figs 5B; 6A, B). Three pairs of terminal cirri (IV-VI) widely separated from mouth cirrus, uniramous with four segments (Figs 4F, G; 5A); basal segment elongate, tubular, with row of long setae along inner surface, reaching to near end of second segment; second segment approximately as broad as basal segment but only about 1/2 as long, with protuberant “cushion” on distal inner surface (Fig. 4H), ridges on cushion with numerous regular rows of small tapered, blade-like denticules (Fig. 4H and inset); third segment narrower than basal or second, approximately as long as basal segment, narrow setae along lateral margins; fourth segment narrower than third, tapering distally, terminating in pair of bifid hooks and single superior short recurved spine (Figs 4F inset; 5A inset, bottom) (one specimen with three bifid hooks plus superior spine, see Fig. 5A inset, top), two very long setae positioned subterminally.</p><p>Male (Figs 1A; 2D, F, G)</p><p>Length 0.71 mm (allotype). One male present maximally per female, attached below mantle aperture of female (Fig. 1A). Form of stout boot with dorsal surface nearly straight, anteroventral surface domed and separated from posterodorsal surface by rounded concave area of nearly 90°, all corners rounded, both ventral and posterior extensions of body subequal in length (Fig. 2D, F), anterolateral corner low and rounded. Lateral surface with distinct median bulge (Fig. 2G). Penis not observed. Ganglion on vesicular seminalis connected by nerve to small dark eyespot (Fig. 2D, F, G).</p><p>REMARKS</p><p>Tomlinsonia Turquier, 1985 was proposed for the preoccupied genus name Alcippoides Turquier &amp; Carton, 1976 (non Strand, 1928) (see Turquier 1985; Tomlinson 1987), and the type species is still known from only a single female specimen. The form of the protuberant “cushion” or “button” on the second segment of cirri IV-VI of female trypetesid specimens has been described as “transversely wrinkled by fine, distinctly crenated ridges of unknown function” (Nilsson-Cantell 1978). In fact, these “wrinkles” are composed of regular rows of minute tapered and bladelike denticules and possibly serve a function in feeding (Fig. 3H). As indicated by Turquier &amp; Carton (1976), these “cushions” were suggested to be vestigial endopodites (Darwin 1851), although the complexity of the denticules on the surfaces does not fit well with a vestigial designation.</p><p>Males of Tomlinsonia mclaughlinae n. sp. differ markedly from those reported for Trypetesa species in that their sides form a vaguely equilateral triangle with one concave margin and lack either the elongated penis containing lobe of Trypetesa lampas (Hancock, 1849), Trypetesa habei Utinomi, 1962, and Trypetesa spinulosa Turquier, 1976 or the deeply concave margins of Trypetesa nassarioides Turquier, 1967. The male of Trypetesa lateralis Tomlinson, 1953 has never been figured or adequately described but was said by Tomlinson (1953, 1969) to lack a penis. Although we have found no evidence of a penis in Tomlinsonia mclaughlinae n. sp., we have only seen five specimens.The apparent absence of a penis is not likely due to immaturity of the specimens (they lack the antennules which are characteristic of immature males) (see Turquier 1971). However, the majority of males in many acrothoracican species may appear to lack penes as these organs can develop relatively late in the life of the male (Tomlinson 1969). Only after examination of large series of males, as done by Tomlinson (1969) for Trypetesa lateralis, should any conclusions about presence or absence of penes in acrothoracican species be made.</p><p>The only acrothoracican previously reported from the Philippines was Trypetesa lampas (Rosell 1981) . Study of Rosell’s (1981) description and illustration leaves no doubt that he was dealing with a Trypetesa species, but his description could apply to any species in the genus and the illustrations, particularly of the mandible, strongly suggest that this was not T. lampas, but perhaps an undescribed species. The so-called “caudal appendage” cited by Rosell (1981) is the sixth pair of terminal cirri; there are no caudal appendages (= furca) on apygophoran acrothoracicans.</p><p>The present finding of a Tomlinsonia species in the Philippines extends the distribution of the genus considerably from the type locality of T. asymetrica in Madagascar and represents the first report of this genus in the northern hemisphere. Such an amphitropical distribution is also seen in Trypetesa and may reflect a relictual distribution of the entire family (cf. Newman 1979).</p><p>The proportions of the terminal cirri segments in Tomlinsonia mclaughlinae n. sp. are quite different from those reported in T. asymetrica . In T. mclaughlinae n. sp. the first two segments are subequal in width and the distal two segments are much thinner, while the first and third segments are nearly as long as each other with the second and fourth segments being approximately 1/2 the length of first and third. In T. asymetrica, the first three segments are all about as wide and as long as each other, while only the fourth segment is shorter (although not 1/2 as long) and thinner. The existence of a pair of bifid terminal hooks and superior spine (Fig. 5A) has not been previously reported in trypetesids.</p><p>The shape of the labrum differs between the two species of Tomlinsonia with T. asymetrica having a gently rounded medial margin between the terminal produced lobes while the same area in T. mclaughlinae n. sp. is subquadrate. In T. asymetrica the thinner terminal lobe is longer than the broader lobe, whereas in T. mclaughlinae n. sp. the broader lobe nearly extends as far as the thinner one.</p><p>There are also some differences between the mouthparts of the two Tomlinsonia species. The mandible of T. mclaughlinae n. sp. is more acutely tapered than that of T. asymetrica and lacks the distal small spine; maxilla II of T. mclaughlinae n. sp. is also more acutely tapered than that of T. asymetrica, while the maxilla I is more regularly ovate and possesses a stout inner setae that is lacking in T. asymetrica .</p><p>It is difficult to compare the various denticules on the inner and outer opercular bars of T. mclaughlinae n. sp. with those of T. asymetrica, as Turquier &amp; Carton (1976) described and figured only those from certain portions of the opercular bars. However, the basic form of ovate denticules with simple to multifid tips can be seen in both species. The dense rows of denticules found on the inner opercular bars dorsal to the ridges are similar to those reported for Trypetesa spinulosa Turquier, 1976 (see Turquier 1978: pl. 3, fig. 4) but are more densely packed and with typically four or five denticular blades on a single shaft as compared to typically two or three in T. spinulosa .</p><p>ECOLOGY The prevalence of Tomlinsonia mclaughlinae n. sp. ranged from 0.45 to 7.94% in the samples from the Philippines (overall prevalence = 3.2%, n = 981 hermit crabs collected during 1997 and 1999). The species is known to be associated with the three most common species of hermit crabs ( Calcinus gaimardii [n = 14], C. latens [n = 16], and C. minutus [n = 1]) from the coral reef areas sampled in the Philippines (Williams 2002). Of the 31 Tomlinsonia mclaughlinae n. sp. specimens examined all but one was found associated with female hermit crabs; 15 of these females were ovigerous (50%). Although the sex-ratios of the hermit crabs did not significantly differ from 50:50 (χ2 [n = 325] = 1.63, P = 0.2, df = 1], the distribution of the barnacles among the two most common hermit crabs was significantly different than predicted values (χ2 [n = 14] = 10.29, P = 0.001, df = 1; this and previous χ2 test based on a subset of the 1999 collections for which the sex of all hermit crabs was recorded). Tomlinsonia mclaughlinae n. sp. has been found in six species of gastropod shells (most commonly Drupella cornus). No shells were found to harbor more than one Tomlinsonia individual, although other species of boring barnacles (such as members of the genera Trypestesa and Weltneria Berndt, 1907) were found in the same shells as those occupied by T. mclaughlinae n. sp. The position of the barnacle in the columella and general body shape of T. mclaughlinae n. sp. are similar to that of T. asymetrica and Trypetesa nassarioides Turquier, 1967 in conforming to the helical spiral of the columella. The aperture of T. mclaughlinae n. sp. is oriented so that the ventral, tapered end of the slit is facing the apex of the gastropod shells, often too far inside the shell to be detected until cracked. Tomlinsonia mclaughlinae n. sp. is apparently an obligate commensal of hermit crabs and is the seventh known species of extant trypetesid associated with hermit crab hosts (Williams &amp; McDermott 2004). Additional trypetesids are known from the fossil record based on their borings in shells (Lambers &amp; Boekschoten 1986; Baluk &amp; Radwánski 1991).</p><p>The feeding biology of Tomlinsonia and Trypetesa remains largely unknown. Tomlinson (1987) suggested the reduced cirri of the barnacles could not be extended outside the aperture and were only used for feeding on small particles brought in by pumping action of the thorax. However, Williams (1999, 2002) reported that an unidentified species of the genus Trypetesa was found to ingest the eggs or developing embryos of host hermit crabs from the Philippines. Based on its position within the shell and proximity to eggs or embryos attached to the pleopods of host hermit crabs (similar to that seen in Trypetesa sp. from the Philippines), Tomlinsonia mclaughlinae n. sp. may also be an egg predator. Among the present samples of T. mclaughlinae n. sp. there is no direct evidence (e.g., egg corions from host hermit crabs identified within the stomach of the barnacles) for this behavior. However, the body of some specimens contained material that was the same coloration as host hermit crab eggs and the barnacles were found predominately with female hermit crabs. Since gonadal tissue of the barnacles could be confused with ingested eggs or embryos of the hosts, histological examination of newly collected specimens needs to be completed to determine if the new species is an egg predator. The barnacles might also gain food in the form of dropped food particles by the crab, materials brought in by the respiratory currents, and/or fecal material of the crab (Baluk &amp; Radwánski 1991; Williams &amp; McDermott 2004), or even the consumption of other symbionts or their offspring in gastropod shells inhabited by hermit crabs. Although egg predation would in large part explain the blind gut of trypetesids, how they are able to capture host eggs and what they feed on while inhabiting shells with male hermit crabs remains unknown.</p><p>Tomlinsonia mclaughlinae n. sp. produce fairly large broods of eggs (estimated at c. 100+ eggs) that are 186 ± 5 µm in length and 145 ± 7 µm in width (n = 10). The eggs appear to hatch as nauplii as evidenced by this stage found in the mantle cavity of some specimens of Tomlinsonia mclaughlinae n. sp.</p></div>	https://treatment.plazi.org/id/CA4F87BBB752FF97F3B1F6C6FED95C13	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Williams, Jason D.;Boyko, Christopher B.	Williams, Jason D., Boyko, Christopher B. (2006): A new species of Tomlinsonia Turquier, 1985 (Crustacea, Cirripedia, Trypetesidae) in hermit crab shells from the Philippines, and a new parasite species of Hemioniscus Buchholz, 1866 (Crustacea, Isopoda, Hemioniscidae). Zoosystema 28 (2): 285-305, DOI: 10.5281/zenodo.5402100
CA4F87BBB75CFF89F3B6F229FD865C53.text	CA4F87BBB75CFF89F3B6F229FD865C53.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Hemioniscus pagurophilus Williams & Boyko 2006	<div><p>Hemioniscus pagurophilus n. sp.</p><p>(Figs 7-9)</p><p>TYPE MATERIAL. — Philippines. Bataan, Morong, 14°41’N, 120°16’E, in Tomlinsonia mclaughlinae n. sp. from shell of Cantharus undosus inhabited by ♀ Calcinus gaimardii, 25.IV.1999, ♂ holotype 1.0 mm (USNM 1084119), 1 ♂ paratype (ZRC 2006.0006); in Tomlinsonia mclaughlinae n. sp. from shell of C. undosus inhabited by ♀ C. gaimardii, 25.IV.1999, 5 ♂♂ paratypes (USNM 1084120). — Oriental Mindoro, Puerto Galera, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=120.95&amp;materialsCitation.latitude=13.5" title="Search Plazi for locations around (long 120.95/lat 13.5)">Big Lalaguna Beach</a>, 13°30’N, 120°57’E, in Tomlinsonia mclaughlinae n. sp. from shell of Peristernia incarnata inhabited by ovigerous ♀ C. gaimardii (3.8 mm SL), 21.VII.1997, immature ♀ allotype 0.9 mm (USNM 1084121), 2 immature ♀♀ paratypes (USNM 1084122), 2 ♂♂ paratypes 0.86 mm (USNM 1084123; 1 on SEM stub, 1 in alcohol) .</p><p>ADDITIONAL MATERIAL EXAMINED. — Philippines. Bataan, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=120.26667&amp;materialsCitation.latitude=14.683333" title="Search Plazi for locations around (long 120.26667/lat 14.683333)">Morong</a>, 14°41’N, 120°16’E, in Tomlinsonia mclaughlinae n. sp. from shell of Cantharus undosus inhabited by ♀ Calcinus gaimardii, 25.IV.1999, 1 ♂ (JDW personal collection).</p><p>ETYMOLOGY. — From the Greek, pagouros (= kind of crab) and philos (= who loves), latinized as pagurophilus, and meaning lover of hermit crabs. The name refers to the role played by the isopod in sterilizing the cirripede and therefore helping hermit crabs by avoiding weakening of their shells by the barnacle and for the inordinate fondness for paguroids by Patsy A. McLaughlin, the best friend a hermit ever had.</p><p>DISTRIBUTION. — Found in the mantle cavity of Tomlinsonia mclaughlinae n. sp., in shells of Calcinus gaimardii hermit crabs from Bataan and Oriental Mindoro provinces, central Philippines; shallow subtidally (&lt;5 m).</p><p>DIAGNOSIS. — Male: cuticular surface with distinct striations, prominent on coxal plates. Head widest at posterolateral junction with pereomere 1, eyes present. Antennule of three articles, basal article with seven teeth and cuticular scale-like ridges, article 1 overlapping basal article of antenna. Antennae of nine articles, articles 1-3 cylindrical with cuticular ridges; flagellar articles much narrower than peduncular articles, proximal flagellar article longest, flagellar articles 2-5 of approximately equal length. Oral cone anteriorly directed. Pereomeres 5 and 6 broadest. Body pigmentation lacking. Pereomeres with entire coxal plates. Pereopods 1 and 2 stout, gnathopodal, with dactyli entire; propodus smooth; ventral surfaces of carpus and merus with large flat cuticular ridges. Pereopods 3-7 ambulatory, smooth; dactyli long, straight, slender, smooth margins, with ventral setal comb, tip hooked with long (pereopods 3-5) or short (pereopods 6, 7) seta at base of hooked tip; propodi straight, medially inflated (pereopods 3-5 broader than 6 and 7), with ventral setal comb. Pleon with five pleopods. Ventral abdominal lobe between first pair of pleopods with two lateral projections and median smooth concave margin. <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=120.26667&amp;materialsCitation.latitude=14.683333" title="Search Plazi for locations around (long 120.26667/lat 14.683333)">Pleotelson</a> quadrangular, endopods approximately twice as long as exopods.</p><p>Immature female with posterior four pleon segments fused with lateral undulations.</p><p>DESCRIPTION</p><p>Male (Figs 7A, B, D; 8; 9)</p><p>Length 1.00 mm, maximum width 0.33 mm at pereomere 4, head length 0.12 mm, pleon length 0.44 mm (holotype); body tear-drop-shaped (Figs 7A; 8A). Cuticular surface with distinct striations, prominent on coxal plates (Fig. 8C, D).</p><p>Head anterior margin ovate, posterior margin concave, widest at posterolateral junction with pereomere 1 (Fig. 7A). Eyes diffuse, moderately large (approximately 40 µm in maximal length) located proximolaterally, eyes with irregular pigment most prominent around laterally directed edge (Fig. 7A). Antennule of three articles (Figs 7B; 8B, C), first (basal) article with seven teeth approximately three times longer than broad, cuticular scale-like ridges present, most prominent on mesial margin, proximolateral lobe distally tapered with group of setae along sinuous cuticular ridge; article 1 slightly overlapping basal article of antenna, article 2 with cuticular scale-like ridges along distal margin, 4-6 setae on distal margin, article 3 with two flagella and bundle of aesthetascs (Figs 7B; 8C). Antennae of nine articles (four peduncular and five flagellar) (Fig. 7B); articles 1-3 cylindrical with cuticular scale-like ridges (Fig. 8C); flagellar articles much narrower than peduncular articles, each with a terminal seta, proximal flagellar article longest, flagellar articles 2-5 of approximately equal length, article 5 with four distal setae (Fig. 7B). Oral cone anteriorly directed (Figs 7B; 8B).</p><p>Pereomeres 5 and 6 broadest, tapering anteriorly and posteriorly. Body pigmentation lacking. Pereomeres with entire (not toothed) coxal plates 1-7 (Fig. 8B, D). Pereopods 1 and 2 short and stout, gnathopodal with dactyli entire (non-bifid) and having few large cuticular scale-like ridges and rows of minute setae on dorsal edge (Fig. 8E); propodus smooth with few rows of short setae distodorsally, rows of minute setae along distal edge of propodus around joint with dactylus, with setae along edge apposed to dactylus; ventral surfaces of carpus and merus with large flat cuticular scale-like ridges and few long setae (Fig. 8E). Pereopods 3-7 ambulatory, smooth; dactyli long, straight, slender, smooth margins, with ventral setal comb, tip hooked via ventral indentation of margin with long (pereopods 3-5) or short (pereopods 6, 7) seta at base of hooked tip (Fig. 9A, B); propodi straight, medially inflated (pereopods 3-5 broader than 6 and 7), with ventral setal comb and one large stout ventral seta approximately 1/2 (pereopods 3-5) or 1/3 (pereopods 6 and 7) from distal margin and second single smaller seta near junction with dactylus (Fig. 9B); carpi of pereopods 3-7 with one stout terminal ventral seta; meri with one thin terminal ventral seta.</p><p>Pleon with five pleopods composed of basis (sympod), exopod, and endopod (Fig. 9C). Sympods with two medially directed setae bearing 3-branched tips, endopods and exopods with plumose setae (five setae each on pleopod 1, three setae on endopod of pleopod 5), with laterally directed seta shorter than other setae on exopod. Ventral abdominal lobe between first pair of pleopods with two lateral projections and median smooth concave margin (Fig. 9C). Pleotelson quadrangular, endopods approximately twice as long as exopods, basis with one distolateral seta, endopods and exopods with terminal setae (Fig. 7D).</p><p>Immature female (Fig. 7C)</p><p>Maximal body length 0.90 mm, maximal width 0.31 mm at segment 4, pleon length 0.32 mm. Similar to male in dorsal aspect, except for fused posterior four pleon segments appearing as an irregular sac posteriorly on the body and with lateral undulations indicating position of pleomeres (Fig. 7C).</p><p>REMARKS Identification of this material was hampered by the confused taxonomy of cryptoniscid isopods in general and that of barnacle parasites in particular (e.g., Nielsen &amp; Strömberg 1965, 1973b). The following comparisons with all other species of cryptoniscoids known to parasitize barnacles were necessary to reach our conclusions.</p><p>Cryptothir Dana, 1852 (incertae sedis): the type species, C. minutus Dana, 1852, was incompletely described and figured (Dana 1852, 1855) from an Indo-West Pacific barnacle of the genus Creusia (material lost). It may be synonymous with Hemioniscus or at least belong to the Hemioniscidae (in which case the family would have to be called Cryptothiridae). However, as noted by Grygier (1993), it is probably best to leave Cryptothir as incertae sedis until such time as additional material can be examined.</p><p>Liriopsis Schultz in Müller, 1864 ( Cryptoniscidae): the new material shows no resemblance to either Liriopsis pygmaea (Rathke, 1843) or L. monophthalma Fraisse, 1878. The most obvious difference is that the females of the new species are only posteriorly degenerated, while those of Liriopsis spp. are unsegmented and form a double lobed shape. Males of the new species have only moderately long dactyli on pereopods 6 and 7, whereas those of Liriopsis spp. are exceedingly long and thin.</p><p>Hemioniscus Buchholz, 1866 (Hemioniscidae): the males of the new species are similar to those of the type species Hemioniscus balani balani Buchholz, 1866 (see Goudeau 1967, 1970), but differ in that the coxal plates of H. balani balani have seven or eight teeth on the first plate, and four or five on posterior plates, while the new species has no teeth on any of the coxal plates. Other differences include strong cuticular striations on the anterolateral expansion of the basal antennule segment (weakly striated in H. balani balani), considerably shorter distal teeth on the basal antennular article in H. pagurophilus n. sp., medially inflated propodi on pereopods 3-5 (distally inflated in H. balani balani), pronounced indentation on the ventral margin near the tip of dactyli (no such indentation in H. balani balani), and ventral median lobe between first pair of pleopods with concave margin (trilobed in H. balani balani). Shared characters between H. balani balani and H. pagurophilus include: moderate striation of the dorsal cuticle, seven teeth on the basal article of the antennule (occasionally eight in H. balani balani), cuticular ridges on the second article of the antennule, moderate setation on the terminal article of the antennule, eyes present, cuticular ridges on the dactyli of the first two pereopods, and moderate dorsal expansion of the meri of the pereopods. The development of the females is essentially identical to that seen in H. balani balani . All of the differences cited above between H. pagurophilus n. sp. and H. balani balani are also true when comparing H. pagurophilus n. sp. and H. balani japonica Ogawa &amp; Matsuzaki, 1985 . The description of H. balani japonica is somewhat difficult to interpret, the line drawings are poor and Ogawa &amp; Matsuzaki only compared their material with older descriptions of H. balani balani (e.g., Buchholz 1866; Sars 1899) and not with the more detailed studies (e.g., Goudeau 1967, 1970). Many of the purported distinguishing morphological characters between the subspecies (Ogawa &amp; Matsuzaki 1985: table 1) do not allow for differentiation, as their information for variation in H. balani balani is incomplete. The two taxa do show quite different coxal plate formulae, and this suggests that they represent distinct species rather than subspecies. However, in the absence of specimens of H. balani japonica, we are unwilling to make any definite opinion on the matter and retain the two taxa as subspecies for the present. The only other species ever placed in this genus was H. socialis Pérez, 1900, which is now considered a synonym of H. balani balani (Caullery 1908) . This genus is the type of Hemioniscidae Bonnier, 1900 .</p><p>Leponiscus Giard, 1887 (Hemioniscidae): Giard (1887) did not select a type species from the two he included in this genus. Leponiscus anatifae Giard, 1887 was described by reference to the figures of the unnamed taxon reported by Hesse (1867), and is an available name; L. pollicipedis Giard, 1887 was accompanied by neither description nor illustration and is a nomen nudum. The type species of the genus must therefore be L. anatifae . Unfortunately, the figures of Hesse (1867) show no clear characters to distinguish L. anatifae from Hemioniscus spp. and Leponiscus must be considered inquirenda within the Hemioniscidae . Gruvel (1901) described the only other species of Leponiscus, L. alepadis, but it differs markedly from L. anatifae and appears to belong in the Crinoniscidae (see below). Gruvel (1902) also cited “ Leponiscus hessei Giard ” but as Giard never mentioned such a name, it is presumed that L. hessei is a lapsus for L. pollicipedis, which was not cited by Gruvel (1901, 1902). The conclusions of Bocquet-Védrine &amp; Bocquet (1972) that Leponiscus Giard, 1887 is a nomen nudum, and that the taxon is valid from Gruvel (1901) (with L. alepadis as the type species) are clearly incorrect as one of the originally included species in Leponiscus Giard, 1887 is an available name. However, their statement that L. alepadis is actually allied to Crinoniscus was correct, and their suggestion that Leponiscus Giard, 1887 may be identical to Hemioniscus is a reasonable hypothesis, albeit an untestable one in the absence of lepadid cryptoniscid parasite material.</p><p>Crinoniscus Bonnier, 1900 (Crinoniscidae):the new material does not resemble the type and sole species Crinoniscus equitans (Pérez, 1900), the only species currently placed in the family (Pérez 1900a, b). Males of C. equitans lack teeth on the basal segment of the antennule and females metamorphose into a crossshaped incubatory pouch. Leponiscus alepadis Gruvel, 1901 appears more closely related to Crinoniscus than either Leponiscus or Hemioniscus as the males also lack teeth on the basal antennular segment, and the females show complete metamorphosis. However, the female body shape of L. alepadis resembles an ovoid sac with small lobes at the corners, rather than the elongate cross of Crinoniscus suggesting that L. alepadis is not congeneric with C. equitans and requires its own genus (Boyko &amp; Williams pers. obs.). Such action is outside the scope of this work.</p><p>Gorgoniscus Grygier, 1981 (unplaced): males of the new material differ from males of sole species of Gorgoniscus, G. incisodactylus Grygier, 1981, in the following: basal article of antennule with seven teeth (five in G. incisodactylus) and tapered proximolateral lobe (lobe lacking in G. incisodactylus), moderate cuticular striations on whole body (dense cuticular striations on head and first pereomere in G. incisodactylus), and dactyli of pereopods 6 and 7 with smooth margins (notched in G. incisodactylus). Females of G. incisodactylus are globose and lack most appendages and an obvious head, whereas immature females of the new species retain the anterior portion of the body as recognizably isopod-like. Grygier (1981) left Gorgoniscus unplaced as to family, suggesting that a new family might be required for it. This genus still remains unplaced, and no further specimens have been reported.</p><p>Scalpelloniscus Grygier, 1981 (? Hemioniscidae): males of both S. penicillatus Grygier, 1981, and S. binoculis (Menzies &amp; George, 1972) differ from the new material in having toothed coxal plate formulae similar to that seen in H. balani balani, more setae on the third articles of the antennules, lack of cuticular ridges on the second article of the antennules, strongly dorsally inflated meri on the pereopods, and lack of eyes. Mature females of Scalpelloniscus are unknown. Based on the current classification scheme, Scalpelloniscus appears to belong to the Hemioniscidae rather than the Cryptoniscidae, but all the families of Cryptoniscoidea need clearer diagnoses and mature females are needed for Scalpelloniscus to determine the degree and form of their metamorphosis.</p><p>In conclusion, examination of the new species of cryptoniscid from the Tomlinsonia hosts indicates that they are best placed in the Hemioniscidae, and are rather close to Hemioniscus balani balani . There are at least as many differences between our Philippine material and H. balani balani as between Hemioniscus and Scalpelloniscus . However rather than erect a new genus for this new species, we have conservatively placed it within Hemioniscus as the males share many characters with those of the type species, and differ most obviously only in the dentition of the coxal plates, a variable feature in other cryptoniscoid families as currently defined.The females of H. balani balani and H. pagurophilus n. sp. have very similar development with nearly identical degeneration of the posterior segments into an irregularly shaped incubatory pouch, while retaining the anterior segments in unmodified form. In order to include the new species in Hemioniscus, it becomes necessary to modify Nielsen &amp; Strömberg’s (1973b) definition of Hemioniscidae as follows: dorsal cuticular ridges relatively prominent; eyes present; basal segment of antennule with seven or eight teeth on distal margin; oral cone anteriorly directed; coxal plates with or without toothed ventral margins; posterior margin of pleotelson entire;parasitic on cirripedes. This definition is acknowledged to be a temporary one, as it is certain that future phylogenetic analyses of cryptoniscoid isopods will result in rearrangement of genera based on shared derived characters rather than shared host taxa.</p><p>ECOLOGY Of the 31 potential host barnacles examined during 1997 and 1999, three were found to harbor Hemioniscus pagurophilus n. sp. (total prevalence = 9.7%); numbers of isopods within the mantle cavity of the hosts ranged from two to six. Interestingly, numerous specimens of an unidentified species of Trypetesa from the same localities were not found parasitized by H. pagurophilus n. sp.</p></div>	https://treatment.plazi.org/id/CA4F87BBB75CFF89F3B6F229FD865C53	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Williams, Jason D.;Boyko, Christopher B.	Williams, Jason D., Boyko, Christopher B. (2006): A new species of Tomlinsonia Turquier, 1985 (Crustacea, Cirripedia, Trypetesidae) in hermit crab shells from the Philippines, and a new parasite species of Hemioniscus Buchholz, 1866 (Crustacea, Isopoda, Hemioniscidae). Zoosystema 28 (2): 285-305, DOI: 10.5281/zenodo.5402100
