(Crustacea: Decapoda: Homoloidea) from the Early Eocene of the Iberian Peninsula

Home , Dromiacea, Homolidae, Homoloidea, Paromola cuvieri

This is anopenaccess article undertheCCBY-NC-SAThis Universidad NacionalAutónoma de México. Peer Reviewing under the responsability of Manuscript accepted: Corrected manuscript received: Manuscript received: BSGM2020v73n3a311220 73 (3), A311220.http://dx.doi.org/10.18268/ Boletín de la Sociedad Geológica Mexicana, the Iberian Peninsula: the early Eocene of (Crustacea: Decapoda: Homoloidea) from a homolid crab recordA., 2021, First of Ferratges, F.A., Domínguez,J.L., Ossó, How tocitethisarticle: [email protected] * Corresponding author:(A.Ossó) gona, Catalonia. 3 B, 50011,Zaragoza, Spain. 2 goza, Spain. CA, Universidadde Zaragoza, E-50009, Zara 1 Fernando A. temprano delaPenínsulaIbérica Primer registrodeuncangrejohomólido(Crustacea:Decapoda:Homoloidea)delEoceno Eocene oftheIberianPeninsula First recordofahomolidcrab(Crustacea:Decapoda:Homoloidea)fromtheearly license(https://creativecommons.org/licenses/by-nc-sa/4.0/) José María Muñoz Damián,6,bloque 1, bajo Llorenç de Villalonga, 17B, 1-1, 43007 Tarra Departamento de Ciencias de la Tierra-IU Departamento Ferratges 1 , José Luis Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín - - - Domínguez Homolidae, Ypresian, Pyrenees. Keywords:Systematics,Brachyura, Pyrenees. southern Eocene basinsof the the richdecapod fossil record of the IberianPeninsula, andexpands homolid reported in the Cenozoic of ment. thissystematic assign 1815) confirms the extant of Directwith specimens comparison this genus. with the extant species of similarities morphological based on in Wood-Mason1891, andAlcock, the genus left cheliped, allowit inclusion within ments, and inparticular the complete specimen, somepreserved frontal ele the sole condition of fragmentary the (Aragon, Spain). In spite of HuescaprovinceRoda Formation of crab fromthe Ypresian (earlyEocene) homolid We describe a newspeciesof ABSTRACT Paromola bretoni 2 , Álex Paromola Paromola cuvieri Ossó Wood-Mason n. sp.n. the first is 3,* (Risso, - - /73(3)A3112202021 el único ejemplar se halla parcialmente España). A pesar de que Huesca (Aragón, (Eoceno temprano) de la provincia de homólido, dela Formación Roda, Ypresiano especie de cangrejo Se describe una nueva RESUMEN Pirineos quiura, Homolidae, Ypresiano, Palabras clave: Sistemática, Bra del EocenoenlascuencasSurpirenaicas. la riqueza sobre de registros decápodos fósiles Cenozoico de la Península Ibérica y amplia n. sp. enel es el primer homólido reportado esta ubicación sistemática. la validez(Risso, de 1815)confirmarían con ejemplares actuales de Mason and Alcock, 1891.Comparaciones el género ubicarloquelípedo izquierdo, en permite frontal y en especial el completo de la región especies actuales de los elementos conservados fragmentado, la similitud morfológica con Paromola Wood-Mason Wood- Paromola bretoni Paromola cuvieri 1

Homolid crab from the early Eocene of the Iberian Peninsula ABSTRACT INTRODUTION/ Homolid crab from the early Eocene of the Iberian Peninsula GEOLOGICAL SETTING 1. Introduction 2 2 et al De Haan, 1839have been described (De Angeli Homolidae where several genera and speciesof Italy,are for not uncommon, instanceinnorthern Europe fossil homolidsinthe Eocene of Records of 1839 are summarized by Guinot (2019,p. 753). Homoloidea De Haan, and current systematics of fossil Explanationspecimen dwelt. onthe former 76), similartothepalaeoenvironment where the inhabit shallower ( depths among homolids,common albeit they canalso fig. 1995, Forges,70), usuallyinhabitingthe upper bathyal zone de Richer and (Guinot wide ford, 2008(seeNyborg and Garassino, 2017). Nyborg and Garassino, 2017, and twoonly other fossilspecies known: the extant genus fossilspecies of a clearaffinitywithHomolidae. the as well the presenceas of the left cheliped, of and the peculiar morphology cuticle, preserved partially with flank, extralineal the left the intralineal carapace and of remains of left cheliped and partial ambulatorylegs. The reveal important elements, suchthe complete as preparation, allowed us to inadequate attempt of the holotype, partially destroyed bya previous of Aragon, Spain). Aproper mechanical preparation Roda deIsábena(Huescaprovince, the village of pods, recovered from early Eocene strata near and preserving somecuticle remains andpereo carapace fragmentarily preserved, decorticated, Peninsula. material The is represented by acrab occurrence the Iberian fossilhomolid crabsinthe Eocene of first the report we Here Peninsula. the Iberian report for the family inthe Eocene of the Iberian Peninsula. of Therefore, this is the first have never been reported to date from the Eocene et al (Collins / / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín xatseiso Extant species of Here we describe ,20,p 92) eod fthe Homolidae ., 2005, p. 19–20). Records of ., 2019,p. 20),aswell as inthe British Isles et al ., 2020,p. 32) and Denmark (Collins Paromola bretoni Paromola ibid linea homolica are spread world p 4941 fig. 489–491, p. , http://dx.doi.org/10.18268/BSGM2021v73n3a311220 P. Paromola P. n.sp., anew

vetula rosenburgensis , indicate Craw , with /73(3)A3112202021 - - - /73(3)A3112202021 2. Geologicalsetting unconformities related to tectonic activity (Lerenunconformities San Esteban (Figure 1); and it is bounded by Serraduy, Puebla Limestone, Roda, Morilloand in this area in the consists following formations: Upper Palaeocene1999). The to Lower Eocene thePyrenees,tropical in latitudes (Hay of the axial zone limit of located on the southern Biscay, connected to the west by the Bay of gulf Eocene, these corresponded basins to an elongated Paleogeographically, duringthe Paleocene– lower Cuisian (lower Ypresian, lower Eocene). records marine sedimentsfrom the Thanetian– Tremp-GrausThe Pyrenees) Basin (southern Barcelona (Spain). InstitutICMD: deCiènciesdelMar (ICM-CSIC), Naturales de laUniversidad deZaragoza (Spain). Abbreviations and Molenaar, 1991). mixed siliciclastic-carbonate (Martinius deposits shallow-marine, of upto 200 mthickformed of Tosquella datedlower as Ypresianage (Samsó in Martinius, 1996;Leren Yang, 1991; Lopez-Blanco, 1996; Molenaar and al S and SW (Puigdefàbregas and show to ageneralthe progradational pattern lowing upwards(Molenaar and Martinius, 1990), 2012). 2013a, b;Schweitzer 1973; Artal and Castillo, 2004;Artal pod assemblage (e.g. VíaBoada, 1959; Vía,1969, providedand has sandstones aninteresting deca marls and exposesof a thick succession formation Huesca,Spain). This de Isábena(northeast of Roda Formation (Figure 1), located near to Roda prodelta marls corresponding to the outcrop of et al ., 1988;Lopez-Blanco This formation has been biostratigraphically hasbeen biostratigraphically formation This RodaThe Formation, andshal iscoarsening studiedmaterialThe fromcomes the exposed ., 2010). et al ., 1990) and consists of a succession asuccession of ., 1990)andconsists . MPZ:Museode Ciencias et al et al et al ., 2007;Van Bakel et al ., 2010). ., 1991,2003;Nioand ., 1985; Molenaar et al et al ., 2005, ., 1990; et al et al et ., ., - - 3. Systematicpalaeontology Figure 1 ino, 2017(fossil); 1908 (extant); macrochira 1815) (extant); Richer de Forges, 1995(extant); de Forges, 1995(extant); Species inal designation. Type species Section Homoloida Karasawa, Schweitzer and Genus Maplocationand explanation offormations. Red starindicatesoutcropwhere Superfamily HomoloideaDeHaan,1839 Infraorder Brachyura Linnaeus, 1758 Family HomolidaeDeHaan,1839 . Paromola Sakai, 1961(extant); Paromola bathyalis P. roseburgensis . P. japonica Dorippe cuvieri and Alcock,1891 Feldmann, 2011 Wood-Mason inWood-Mason P. vetula 2011 Crawford, 2008(fossil). Boletín de la Sociedad Geológica Mexicana Mexicana Mexicana Geológica Geológica Sociedad Sociedad la la de de Boletín Boletín Parisi, 1915(extant); P. crosnieri Guinot and Richer Nyborg andGarass Risso, 1815,by orig http://dx.doi.org/10.18268/BSGM2021v73n3a311220 P. rathbunae P. cuvieri Guinot and Porter, (Risso, .P. - - Material. French palaeocarcinologist. Etymology ambulatory legs long, spiny, subcircular in section. circular very in section,spiny; elongated; carpus lateral reentrants. Cheliped extremely long, sub orbits. Posterior margin concave axially with two upwards, originating just above upper margin rostrum; pseudorostralupturned directed spines in anterior third. Long, pointed, and moderately the mesobranchial region, narrowing est at level of carapace; wid directed spines at anterior third of low grooves, incised someculminated byupward rectangular; regions fairly defined, marked by shal Diagnosis. urn:lsid:zoobank.org:act:D020CE6C-4633-4712- Paromola breton HolotypeMPZ 2020/615,decorti /73(3)A3112202021 . Honoring the late. Honoring Gérard Breton, / 73(3)A3112202021 Dorsal carapace longitudinallysub Dorsal 9C38-6237E281CA04 Paromola bretoni (Figures 2and3) i n.sp.wasrecovered. n. sp 3 3 - - - - -

Homolid crab from the early Eocene of the Iberian Peninsula SYSTEMATIC PALAEONTOLOGY Homolid crab from the early Eocene of the Iberian Peninsula SYSTEMATIC PALAEONTOLOGY equal to5mm.(Photographs byIsabel Pérez Urresti). legs; r=rostrum; rl=rostrallobe;s1 =pleonalsomite1.Photographs byIsabel PérezUrresti. ScalebarA,B,C,D,equal to10mm;E, of dactyli. Abbreviations: ba = possibly antennular basal article; lh = article; basal antennular possibly = ba Abbreviations: dactyli. of Figure 2 4 4 / /

Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Paromola bretoni n. sp.,holotype MPZ 2020/615. A:dorsal view; B:frontal view;C:left lateralview;D;ventralE:close-up http://dx.doi.org/10.18268/BSGM2021v73n3a311220 /73(3)A3112202021 /73(3)A3112202021 linea homolica ; psr = pseudorostral spines; P2-P4 = ambulatory = P2-P4 pseudorostralspines; = psr ; extant Figure 3 ae rbcrpc rsrigfwrmiso cated crab carapace preserving fewremains of Ypresian, early Eocene. position and age Stratigraphic Locality Type ICMD122/1998, male, juvenile, Catalonia. lonia; ICMD121/1998,male, juvenile, Catalonia; Namibia; ICMD120/1998, male, juvenile, Cata Namibia; ICMD002503-2, female, juvenile, nile, Namibia; ICMD002503-1, male, juvenile, juvenile, Namibia;ICMD002500, male, juve male, juvenile,male, Namibia;ICMD002501, ICMD002499, female, Namibia; ICMD002498, female,alonia; ICMD119/1998, Namibia; nile, Barcelona; ICMD002023, female, Cat male, Namibia;ICMD002502,male, juve female, juvenile, Namibia;ICMD002505, the catalog/accession number: ICMD002504, celona, Spain) (see Guerrero Institut de Ciències del Mar (ICM-CSIC, Bar Biological Reference Collections (CBR) at the Fourteen samples of Extant comparativematerial. examined left ambulatory legs. cuticle, andcomplete left cheliped and remains of Paromola Idealized reconstructionof spp.(withoutscale). . Roda de Isábena, Huesca, Spain. Paromola cuvieri Boletín de la Sociedad Geológica Mexicana Mexicana Mexicana Geológica Geológica Sociedad Sociedad la la de de Boletín Boletín . Roda Formation, late Paromola bretoni et al http://dx.doi.org/10.18268/BSGM2021v73n3a311220 ., 2020), under from the n. sp., the shaded parts takenfrom n.sp.,theshadedparts takenfrom thespecimen,and lightparts - - - - -

ly-projecting spines. Posterior margin rimmed, lateral margin convex, withat least two lateral upper margin rimmed. margin, directed upward. Orbital cavityrounded, sulcate; pseudorostral spines present on anterior rostraland moderately lobe axially upturned, rounded insection,presumably well developed in intralineal areas. Rostrum simple, pointed, longitudinally and weakly vaulted transversely carapace, dorsal weakly vaulted eral margins of upward directed Convex(scars). acute spines lat granulate smooth, bearing presumably large remains gently surface thin, appears dorsal cuticle of of regions; carapace defined fairly third; at metabranchial region, narrowinganterior in rectangular, longitudinally ovate in outline; widest Description 59.52 x10.16mm(propodus +dactyli). chelax 6.7mm; = 27.72mm = carpus 6.17mm; x 2).Left chelipedmm x (P1): merus =52.39mm carapace 17.72 mm (without lateral half spines; apace length = 36.05mm:carapace width: 35.44 Measurements Anterolateral margin not preserved. Postero /73(3)A3112202021 / 73(3)A3112202021 . Carapace longer than wide, sub . Holotype MPZ 2020/615.Car 5 5 - - - - -

Homolid crab from the early Eocene of the Iberian Peninsula SYSTEMATIC PALAEONTOLOGY Homolid crab from the early Eocene of the Iberian Peninsula SYSTEMATIC PALAEONTOLOGY/ DISCUSSION 4. Discussion 6 6 sub-triangular, withthree swellings, two anterior region Cardiac depressed. weakly defined, poorly versely(broken).aligned spines Urogastric region rior swellings. Protogastric region with four trans ante two with region Epigastric defined, well not length: one axial, and two lateral. Gastric regions three concave arcs, almostequal in of consisting differ substantially from substantially differ specimen. studied the For a simple possess rostrum and long chelipeds, they cheliped. Although someother also homolids anextremely long of spines, andthe long carpus the acute simple rostrum, developed pseudorostral Richer de Forges (1995, p. 358–359),in particular diagnosis for the genus provided by Guinot and lar to that of within theHomolidaewithconfidence. spines, andthe the pointed simple rostrum, the pseudorostral well-preserved andextremely longleft cheliped, served, somediagnosticcharacters, such asthe partially is Although the prestudied specimen thelower margin. of along thewholespaced linear arrangement length the upper margin,in evenlyin andsmallspines to P4,veryelongated, with threelarge to six spines P2 in occlusalmargin; polex edentulous. Meri of several setal pits; dactyluswith one proximal tooth small spines. Dactylisubequal,darkcolored, with andpropodus,gated, both, carpus covered with very elon section; merus strongly spinose;carpus ous. Left cheliped extremely long, subcircular in shallow, groove medially. interrupted Branchiocardiac the carapace. of flanks spines two small (broken). at least with inflated, weakly (broken); spine one small metabranchial regions least at with inflated, weakly region, urogastric of Mesobranchial regions positionedoneither side depressed. terior.defined, poorlyregion Intestinal separated bya shallow and onepos depression, / / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín h obnto fthese characters issimi combinationThe of Cervical groove pronounced, directed to the Paromola linea homolica , and fits perfectly with the with perfectly fits and , , allow to assign it Linea homolica http://dx.doi.org/10.18268/BSGM2021v73n3a311220 sinu /73(3)A3112202021 ------/73(3)A3112202021 n. sp. (seeJenkins, 1977). longer 134), has pseudorostralthan spines genus byto the former Schweitzer as South Australia, described formerly Oligocene of pritchardiDagnaunus 28–33). figs. 376–391, p.Forges, 1995, de Richer preservedthe studiedspecimen(seeGuinotand in spines, whichlong andsharp are absentornot pseudorostral spines, carapace with anddorsal well, species of Richer de Forges, 1995, p. 335-338, fig. 11a-b). As are shorter than inthe new species(see Guinot and acute possess rostrum, and long chelipeds, but they instance, some speciesof directed spines, and linea homolica well defined, well homolica linea and spines, directed regions fairly defined, some culminated by upward abovetral spines upper orbital margin, dorsal species, bya simplerostrum,sharing pseudoros inimola williams those of rostral and pseudorostral features, both similar to Essex (UK),drawsattention byits size, and of Collins andSaward, 2006,from the lower Eocene vetula carapace preserved aregranulated less than in dorsal tina) by itslarger size, anditsremains of vetula 4). Moreover, 3, figs. 140–142, p. 2017, Garassino, and Nyborg cheliped, whichshorter in is of shape different and spines, pseudorostral shorter Oregonhaving (USA)in larger size, apparently of from differs serve the entire carapace. However, the studied specimen, which doesnothelp to pre the fossilhad thesamefragile species cuticle, as (P. Abelló, pers. comm.),soit can be inferred that Extant species of it further with the fossilspecies above indicated. Paromola bretoni thesystematic position. new material confirms specimen within that and therelatively large size, theplacement of Paromola pritchardi h tt f rsraino tecrpc f thecarapace of preservation of stateThe of Based uponthe above-mentioned characters, (see Crawford, 2008). RíoNegro (Argen , from the Oligocene of Paromola bretoni P. roseburgensis Paromola bretoni i is the closestfossili is to our species Moloha n. sp., precludes us to compare Paromola Paromola (Jenkins, 1977), from the early , and subsequently , and subsequently transferred Barnard, 1947, possess long 1947,possess Barnard, n.sp. Apparently, , seems appropriate, seems until possess a thin carapace possess from the early Eocene Homola n. sp. differs from differs sp. n. Londinimola williamsi P. roseburgensis et al P. bretoni Leach, 1816, . (2004,p. P. bretoni n. sp. Lond (see .P. P. - - - - Acknowledgements differentiation between them. Other comparative Other them. between differentiation extant cheliped with respect to the carapace size than which meansthat the new specieshaslonger left 0.600, whereas in the carapace length/merus P1 length ratio is than ratios different the extant of specimens In thissense, examination and measurements of cheliped span withrespect to the carapace length. thoseextant species seemtohave suchleft none of bretoni However, despite the crushed carapace of the genus extant species of 2006, p. 69–70,pl.1,figs. 1a-c,2a,b). detailed comparison (see Collins andSaward, a more the new species, doesnotpermit vation of than in legs of ambulatory the preserved cheliped and remains of as well, upturned; rostrumdownturned instead of but it fromdiffers FAFsupported byis projects CGL2017-85038-P and provided useful comments. research The of Javierlanguage Luquehelped with theEnglish us reviewsthat improved theoriginalmanuscript. and Francisco Vega (UNAM, Mexico) by their Garassino (LomaLinda University, CA,USA) the sample.graphing We alsothankAlessandro to Isabel Pérez Urresti, for her help in photo providing usefulcommentsandrelevant data, Santos-Bethencourt (ICM-CSIC, Barcelona) for We are to Pere grateful Abelló and Ricardo for theAragonesespecimen. a new species extant species, justifythe erection of temporal span separating P. cuvieri ratio, do not show substantial differences: 0.505 in n. sp.,length/merus andlength/ such ascarpus of measurements regarding the carpus Similarly, comparing n p ol igietetu egho it, n.sp.the true coulddisguise length of P. bretoni P. cuvieri vs. 0.529 inP L. williamsi wih ol fvu te specific the favour would which , n. sp. are clearly longer and robust P. bretoni Paromola bretoni . Inanycase, the poor preser . bretoni P. bretoni P. cuvieri Boletín de la Sociedad Geológica Mexicana Mexicana Mexicana Geológica Geológica Sociedad Sociedad la la de de Boletín Boletín P. bretoni n. sp. that ratio is 0.688, P. bretoni n. sp. In any case, the n.sp., for instance Paromola http://dx.doi.org/10.18268/BSGM2021v73n3a311220 above listed, show n.sp. inhavinga n. sp. with the n. sp. from the s difficult. is P. bretoni Paromola - - References Artal, P., Ossó, A.,Domínguez,J.L., 2013a, Artal, P., Castillo, J., 2004, Aragón)-FEDER. of ernment the gov RecursosPaleoambientesGeológicosy of and Innovation), andProject E18 Aragosaurus: Science of Fund, FPU Grant (Spanish Ministry Innovation, theEuropean Regional Development Science and subsidized by of the Spanish Ministry Collins, J.S.H., Mellish,C.J.T., Ross, A.J., Crabb, newspecies K.H.,1947,Descriptionof Barnard, Artal, P., van Bakel, B.W.M., Fraaije, R.H.B., Artal, P., van Bakel, B.W.M., Castillo, J., 2005, Batalleria, 12,39–44. 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