The Massive Death of Lobsters Smothered Within Their Thalassinoides Burrows: the Example of the Lower Barremian from Lusitanian Basin (Portugal)

Home , Glypheoidea, Linuparus, Mecochirus, Thalassina

Versão online: http://www.lneg.pt/iedt/unidades/16/paginas/26/30/209 Comunicações Geológicas (2016) 103, Especial I, 143-152 ISSN: 0873-948X; e-ISSN: 1647-581X

The massive death of lobsters smothered within their Thalassinoides burrows: the example of the lower Barremian from Lusitanian Basin (Portugal)

A morte em massa de lagostas sepultadas no interior das suas galerias do tipo Thalassinoides: o exemplo do Barremiano inferior da Bacia Lusitânica (Portugal)

C. Neto de Carvalho 1,*

Abstract: Some years ago an exceptional site with the preservation of the evolutiva demonstrada pela ocorrência de Thalassinoides associados a lagostas mecochirid lobster Meyeria rapax (Harbort) within their burrows was glyphoídeos desde o Jurássico Superior ao Aptiano, e à sua progressiva described in the Early Cretaceous of Cabo Espichel sector, Lusitanian Basin, substituição pelos thalassinídeos, com algumas excepções registadas para as Portugal. The large number of articulated specimens preserved at the bottom lagostas queladas, a partir do Cretácico Superior. Esta tendência para a of Thalassinoides burrow systems in, at least, four different beds of the Boca convergência evolutiva no hábito fossorial é comparável com os padrões do Chapim Formation (lower Barremian) suggests that a sudden cyclic event evolutivos de diversificação das lagostas e dos thalassinídeos reconhecidos was responsible for the collective killing below the bottom of a confined recentemente. lagoon, and led to the very rare preservation of both burrows and producers. Palavras-chave: Meyeria rapax ; Thalassinoides suevicus ; morte colectiva; The explanation may be found in the highly heterogranular, coarse-grained convergência comportamental; Barremiano inferior; Bacia Lusitânica clastic filling of the burrow systems which represent influxes of freshwater and large volumes of siliciclastics that may had suddenly drop the salinity 1 Geopark Naturtejo da Meseta Meridional – UNESCO Global Geopark. Geology of the carbonate lagoon and smothered the endobenthic communities. These Office, Municipality of Idanha-a-Nova – Raiano Cultural Centre. Av. Joaquim climate-related events seem to have been spread in a wider region of the Morão, 6060-101 Idanha-a-Nova (Portugal) Lusitanian Basin during, at least, the lower Barremian, since previous *Corresponding author/Autor correspondente: [email protected] . findings took us to discover a new site in the correspondent Ribeira de Ilhas Formation, at Ericeira sector. In this study we also document the evolutionary trend in the occurrence of Thalassinoides associated to glyphoid lobsters since the Early Jurassic to the Aptian, and their replacement in soft substrates 1. Introduction by thalassinideans, with few exceptions for chelate lobsters, after the Late Trace fossils may be used for many different purposes, from Cretaceous. This trend to behavioral convergence of the fossorial habit paleoenvironmental indicators in basin analysis to tangible matches lobster and thalassinidean diversity and evolutionary patterns evidences of behavioral interactions with substrate, and thus, through time recognized recently. valuable tools for paleobiological studies. But for a large majority Keywords: Meyeria rapax ; Thalassinoides suevicus ; mass mortality; of ichnotaxa the same question always arise: “Who did it?” Most behavioral convergence; lower Barremian; Lusitanian Basin of the times the diagenetical conditions, such as bacterial decaying or ion-rich fluid percolation, that made trace fossils more clear Resumo: Há alguns anos atrás descrevemos um sítio paleontológico excepcional onde a lagosta mecochirídeo Meyeria rapax (Harbort) é encontrada within the sedimentary background, are the same unfriendly for the no interior dos seus sistemas de galerias, no Cretácico Inferior do sector do preservation as body fossils of the soft-bodied, or low-mineralized Cabo Espichel, Bacia Lusitânica, Portugal. O elevado número de espécimes skeletal organisms responsible for them. Using morphological articulados, preservados no fundo de galerias do tipo Thalassinoides em, pelo features of burrows, borings, trackways and coprolites one may menos, quatro camadas da Formação de Boca do Chapim (Barremiano compare with similar structures from the present and try to get a inferior), indicia que um evento cíclico instantâneo terá sido responsável pela match that may relate, at least, with few or one clade, many times a morte colectiva abaixo do fundo da laguna confinada, e que levou à muito rara Phyla or a Class only. Although not vital for the development of preservação dos sistemas de galerias conjuntamente com os seus produtores. Ichnology, is frequently really frustrating for the scientist or the A explicação pode ser encontrada no preenchimento clástico heterogranular, science communicator to discuss about the behavior and fundamentalmente grosseiro dos sistemas de galerias, que representa descargas evolutionary significance of a trace fossil having no clues about the de água doce e de grandes volumes de detritos siliciclásticos que terão levado organism to which have physiological relations, also knowing that à descida brusca dos níveis de salinidade na laguna carbonatada e ao sepultamento das comunidades endobentónicas. Estes eventos climáticos the same life form can be responsible for different behavioral parecem ter tido um impacte mais alargado na Bacia Lusitânica, pelo menos interactions with the substrate, and the same trace fossil could be durante o Barremiano inferior, uma vez que achados antigos isolados levaram- the result of convergent behaviors for phylogenetical distant clades, nos à descoberta de um novo sítio na correspondente Formação de Ribeira de reinvented over and over again along the geological record. And Ilhas, no sector da Ericeira. Neste estudo também documentamos a tendência there are the common cases that the behavior become extinct with 144 C. Neto de Carvalho / Comunicações Geológicas (2016) 103, Especial I, 143-152 the producer and only somewhat vague comparisons with other trace et al. , 1989) dated as Late Triassic-Early Jurassic, Late Jurassic and fossils are allowed to be made. So, as the konservate lagerstätten are Lower Cretaceous. This crustal extension was related to the opening so rare but sometimes fundamental to understand the functional of North Atlantic. During the Oxfordian-Tithonian stretching episode morphology of an organism, or the whole paleocommunity the Lusitanian Basin was segmented into a number of rapidly dynamics, well preserved evidences of behavior frozen with the subsiding subbasins, major half-grabens tilted blocks, with species responsible for it open new windows to understand subsequent facies differentiation. The western Iberian margin biological evolution, or at least corroborate with clear evidences constitutes a passive margin during the 4 th rifting episode (late hypotheses or ideas in the making. Berriasian-late Aptian) with the filling of the subbasins and the Since Thalassinoides ichnogenus was erected by Ehrenberg opening of the shelf to full marine conditions southwards (Kullberg (1944, see also 1938) these very common burrows in Mesozoic and et al. , 2013). The general filling of the basin is marked by siliciclastics Cenozoic shallow marine deposits have been mostly attributed to transported by rivers from the Iberian Massif positioned to the east. the work of decapod crustaceans, and among these particularly Unconformities and major cycles during the Lower Cretaceous were thalassinoidean (most of Axiidea and Gebiidea) shrimps ( e.g. , controlled mostly by the opening of the Atlantic (Rey et al. , 2006). A Ehrenberg, 1938; Mertin, 1941; Fiege, 1944; Glaessner, 1947; second order T-R cycle was identified for the interval Valanginian- Bromley, 1967; Miller and Curran, 2001; Mourik et al. , 2005; lower Barremian. A transgressive open carbonate platform was Seilacher, 2007; table 1 with occurrences of fossil callianassid developed, with a rapid rise in relative sea-level (Dinis et al. , 2008). remains preserved within burrow structures or associated with Several somewhat hypersaline lagoons were developed and strongly burrows of Hyžný, 2011; Hyžný and Klompmaker, 2015), but also confined by rudist barriers located to west. These lagoons were often and less commonly to stomatopods, crabs and lobsters ( e.g. , invaded by coarse-grained fluvial siliciclastics (Rey, 1992). Bromley and Frey, 1974; Pemberton et al. , 1984; Neto de Carvalho The northern coast of Cabo Espichel exposes a continuous et al. , 2007). Common integradation between the ichnogenera Lower Cretaceous (lower Berriasian-to-Albian) succession (Rey; Ophiomorpha , Thalassinoides and Spongeliomorpha (O-T-S) has 1972) of mixed carbonate-siliciclastic shallow marine and continental been repeatedly described ( e.g. , Bromley and Frey, 1974; Fürsich, deposits 350 m thick (Fig. 1). The lower Barremian depositional 1973, 1974; Schlirf, 2000), and they mostly result from burrowing sequences of third order are composed by Areia do Mastro (partim), for dwelling and feeding in different substrate consistencies, from Papo Seco, Boca do Chapim and Regatão (partim) formations, soup- to soft/stiff- and firmgrounds, by the same kind of producer, showing eustatic oscillations of one million years range (Rey, 1992, and not represent different burrowing behaviors for distinct 2006). Boca do Chapim Formation, 16 m thick, is composed by burrowers. It should be rare to find well-preserved crustacean bodies quartz-rich clayish limestones, passing to highly bioturbated marls within burrows, unless a sudden event (storm, flood, anoxia, salinity with Choffatella decipiens (Schlumberger, 1904) and a bioclastic- drop) caused the animal to die in its burrow. Also, the thin- rich limestone bed. The unit shows other body fossils such as mineralized cuticles of these endobenthic decapods only rarely are Ampullela leviathan , nerineid gastropods, rudists, regular echinoids, preserved within burrows and they are mostly exuvia or chela only. oyster and serpulid patches, wood remains, bivalves, pycnodontid Some authors describing these rare direct relations between O- teeth, and turtle bones. Some ammonites were recorded 3,5 m and T-S and body fossils have tried to list the species and their burrows 4,3 m below the top of Boca do Chapim Formation. (e.g. , Neto de Carvalho et al. , 2007). Lately, Hyžný and Klompmaker The succession is composed by 16 beds, with all the soles (2015) presented a very complete list of occurrences of fossil ghost bioturbated by Thalassinoides . The main units with M. rapax are shrimp remains preserved in burrows. Unfortunately, those burrows located 4,6 m and 5,25 m from the base of the Boca do Chapim are not treated ichnotaxonomically or described which could Formation, a succession of bioclastic marls and limestones provide more useful information on the environmental distribution thickening upwards, and quartz calcarenite, respectively. and evolutionary paleobiology of fossorial behavior. Bioclastic marls and limestones, including abundant poorly The exceptional discover of numerous articulated Meyeria sorted quartz and lidite grains measuring less than 4 cm, show very rapax within burrow systems attributed to Thalassinoides suevicus abundant Choffatella decipiens. Rare Rhizocorallium commune (Rieth, 1932) in the lower Barremian of the Lusitanian Basin, Schmid, 1876 are associated in the same beds. The sequence Portugal (Neto de Carvalho et al. 2003, 2007, 2010) in the context represents a transgressive period and high system tract and it is of few other important sites discovered in the same period (Tsujita, limited by an erosional boundary with the coarse-grained fluvial 2003; Vega et al. , 2008) may have served as trigger for studies on beds of the overlying Regatão Formation. the evolution of fossorial behavior in important groups of decapods, The lower Barremian of Ericeira region (Fig. 1) is composed, such as lobsters and thalassinideans. In Portugal, the analysis of old from bottom to the top, by Ribamar (partim), Ribeira de Ilhas and collections hosted in the National Museum of Natural History and Regatão (partim) formations (Rey, 1992). Ribeira de Ilhas Formation Science, permitted that a new site with in situ preservation of is located between Porto do Cavalinho beach, at north, where Ferreira numerous Meyeria rapax within Thalassinoides suevicus would be (1955) found a single M. rapax , and Ribeira de Ilhas beach, at south, found in Ribeira de Ilhas cliffs, Ericeira. This new site and Cabo where we found a new level with M. rapax within Thalassinoides Espichel are described and compared in this paper to find the causes suevicus . The succession, 50 m thick, is composed by highly of collective death and highly unusual preservation of these lobsters bioturbated ( Thalassinoides suevicus ) clayish limestones and marls in the same time frame, in a basinal and paleoenvironmental context with carbonate nodules. In the lower part of the sequence sediments where Thalassinoides ichnofabrics are widespread. Here we also often show high abundance of poorly sorted quartz grains. In the review the other exceptional occurrence of Thalassinoides suevicus upper part, succeeds fine beds of sandstones or quartz-rich limestones with their producers and relate them with the latest phylogenetic with planar oblique stratification, dolomites and lignite-rich dark reviews of lobsters and thalassinidean shrimps so to contribute for marls. Body fossils are abundant, including Chofatella decipiens and the understanding of O-T-S producers across the geological times. other foraminifers, charophytes, dasycladacean algae and echinoids (Rey, 1992). Ribeira de Ilhas Formation is dated from the lower 2. Studied sections in the Lusitanian basin Barremian and shows lateral sequential correspondence with the Boca do Chapim Formation (Rey, 1972; 1992; Fig. 1), with same basal The Lusitanian Basin, located along the western Iberian Margin, was transgressive sequence with upward fine-graining and high system defined by four episodes of Mesozoic lithospheric stretching (Wilson tract with intercalations of prograding sandstones and dolomites. The Lobsters within Thalassinoides in the Lusitanian Basin 145

Fig. 1. Location of the studied sections at Cabo Espichel (A) and Ericeira (B) and stratigraphy of Boca do Chapim and Ribeira de Ilhas formations (adapted from the Geological Map of Portugal 1:500000, and Rey (1972), respectively). Key: 1 – Cross-bedded sandstones; 2 – Planar-bedded sandstones; 3 – Marls and sandstones; 4 – Marlstones; 5 – Limestones and sandstones; 6 – Reefal limestones; 7 – Limestones; 8 – Limestones and marls; 9 – Limestones and dolostones; 10 – Dolostones. Fig. 1. Localização dos afloramentos estudados no Cabo Espichel e Ericeira e estratigrafia das formações de Boca do Chapim e Ribeira de Ilhas (adaptados da Carta Geológica de Portugal à escala de 1:500000, e de Rey (1972), respectivamente). Litologias: 1 – Arenitos com estratificação entrecruzada; 2 – arenitos com estratificação planar; 3 – Margas e arenitos; 4 – Margas; 5 – Calcários e arenitos; 6 – Calcários recifais; 7 – Calcários; 8 – Calcários e margas; 9 – Calcários e dolomitos; 10 – Dolomitos. important (mid?) Barremian regression with coarse siliciclastics was a) Meyeria rapax in the lower Barremian of the Lusitanian Basin due to regional uplift created by the onset of seafloor spreading in the The characteristic feature of the Mecochiridae family is the Iberia sector (Dinis et al. , 2008). extraordinary extension of the first pereiopods (Förster, 1971). The synonymy of Mecochirus Germar, 1827 and Meyeria M’Coy was 3. Mass mortality in lobsters: the exceptional case of the suggested by Förster (1971), and the first identifications in the lower barremian from lusitanian basin Lusitanian Basin were attributed to the first genus (Ferreira, 1955; Neto de Carvalho et al. , 2003, 2007), but differences between these In both Cabo Espichel and Ericeira sections Meyeria rapax show two mecochirid genera have been noted (Aguirre-Urreta, 1989; 3D preservation, sometimes compressed, of the articulated Astrop, 2011). Neverthless, the species M. rapax was synonymized cylindrical bodies within their burrows. Systematic search for fossil with the genus Meyeria by Feldmann et al. (1995). Meyeria have lobsters in the Boca do Chapim Formation allowed to determinate the first pereiopods extremely long, in some species like in Meyeria 27 fallen blocks in the tallus of the cliff composing sole beds with rapax , longer than or as long as the abdomen and cephalothorax a total area of 169 m 2, where 95 specimens attributed to Meyeria combined (Astrop, 2011; Gonzalèz-León et al. , 2014). The absence rapax were recorded, with an average rate of 1 lobster:1,77 m 2 (Fig. of claws suggests that mecochirid lobsters were probably more 2). Non-systematic work rendered several more specimens, with adapted for scavenging (Feldmann et al. , 2002). At Cabo Espichel, hundreds of individuals observed in this section. Most of the the first pereiopods may reach up to 195 mm long, are laterally specimens seem to be within burrows corresponding to large-sized compressed and saw-toothed in the edges. The body length varies Thalassinoides suevicus , but there are some found apparently between 90-160 mm. We found different body sizes but no clear without any relation with burrows that were buried and preserved ontogenetic stages were identified. by the same event. These specimens show often a “hands up” Bordering the cephalothorax there is a marginal carina that displacement of the first pereiopods resulted from the dragging of extends along the saw-toothed medial rostrum evidencing lateral the corpses on the substrate during the deposition of the sediments. crests. This structure is separated from the branchial area by a deep, In order to be found in the field, M. rapax must have been preserved narrow groove, and is ornamented with asymmetric coarse granules in the bottom of the burrow at the sole of the beds, where the fallen (pointing backwards; Fig. 3C). The posterior edge of cephalothorax rocks usually splits to expose the Thalassinoides meshworks (Fig. shows a concavity enclosing a well developed abdomen composed 2B). The presence of a lobster within the burrow may also be of six smooth somites culminating in a fan-shaped telson, which is detected by the concretionary swelling (Fig. 2C), but these only rarely preserved. Given their long and slender pereiopods, it is occurrences were not counted that certainly would increase the common to see portions of the merus and carpus (more frequent on density of M. rapax populations closer to the original number of, at the second and third pairs), which are dorso-ventrally flattened. The least, one individual per burrow system. pleopods are not preserved. The existence of lateral carina in the 146 C. Neto de Carvalho / Comunicações Geológicas (2016) 103, Especial I, 143-152

Fig. 2. Meyeria rapax in Thalassinoides suevicus from the Boca do Chapim Formation (Cabo Espichel section). A – Boca do Chapim Formation in its type section; B – One of the typical boulders fallen from the cliff that provide bedding plane conditions to analyze the Thalassinoides meshworks with often curvilinear burrows; C – Swellings at the Thalassinoides burrows may indicate the location of their producer when it is not preserved at the bottom of the burrow; D – Cross-cutting relationship between two generations of Thalassinoides , with the later one being much smaller and having a very regular network geometry related with the eodiagenetical increase of the substrate cohesiveness; scale bar is 10 cm; E – Meyeria rapax within Thalassinoides suevicus in rigore mortis position showing the first two pairs of pereiopods; scale bar is 1 cm; F – An example of Meyeria rapax within it burrow filled by coarse grained siliciclastics and cemented by micrite; G – Typical nearly complete preservation of Meyeria rapax ; scale bar is 1 cm; H – Filling of Thalassinoides by Choffatella decipiens -rich carbonate. Fig. 2. Meyeria rapax em Thalassinoides suevicus from the Boca do Chapim Formation (Cabo Espichel section). A – Formação de Boca do Chapim na sua secção-tipo; B – Um dos típicos blocos resultantes da erosão da arriba que possibilitam a análise dos planos de estratificação com os sistemas do tipo Thalassinoides , que frequentemente possuem galerias curvilíneas; C – Alargamentos das galerias do tipo Thalassinoides indicam normalmente a localização do seu construtor quando este não se encontra preservado no fundo da galeria; D – Relação de intersecção entre duas gerações de Thalassinoides , com a subsequente sendo de menor diâmetro e possuindo uma geometria regular de galerias como resultado do aumento eodiagenético da coesão do substrato; escala gráfica de 10 cm; E – Meyeria rapax no interior de Thalassinoides suevicus em posição rigore mortis mostrando os primeiros dois pares de pereiópodes; escala gráfica de 1 cm; F – Exemplo de Meyeria rapax no interior da sua galeria preenchida por siliciclásticos de grão grosseiro e cimentada por micrite; G – A típica preservação quase completa de Meyeria rapax ; escala gráfica de 1 cm; H – Preenchimento de Thalassinoides por carbonatos ricos em Choffatella decipiens . Lobsters within Thalassinoides in the Lusitanian Basin 147

Fig. 3. Meyeria rapax in Thalassinoides suevicus from the Ribeira de Ilhas Formation (Ericeira section). A – Ribeira de Ilhas Formation at Praia do Covalinho; B – Meyeria rapax within Thalassinoides suevicus found at the base of the section of Praia do Covalinho; C – Detail of the preservation of Meyeria rapax at Ribeira de Ilhas Formation; scale bar is 1 cm. Fig. 3. Meyeria rapax in Thalassinoides suevicus from the Ribeira de Ilhas Formation (Ericeira section). A – Formação de Ribeira de Ilhas na Praia do Covalinho; B – Meyeria rapax no interior de Thalassinoides suevicus encontrado na base da secção da Praia do Covalinho; C – Detalhe da preservação de Meyeria rapax na Formação de Ribeira de Ilhas; escala gráfica de 1 cm. rostrum, a deep, narrow cervical groove, a posterior border on the with the major axis parallel to the bedding planes due to sedimentary cephalothorax with carina, and strong granular ornamentation on the compaction; this confirms the common compressed preservation of branchial region identify the Portuguese mecochirid forms as Meyeria M. rapax carapaces. Burrow walls are smooth and unlined and the rapax (sensu Feldmann et al. , 1995) . The elongated form of the burrow fill is passive (Fig. 2H, Fig. 4), differing from the limestone carapace, flattened appendages, absence of claws, sawtooth crests, or fine sandstone sedimentary host rock by the more intense and asymmetrically disposed granular ornament pointing backwards carbonate cementation. These features suggest that burrowing was are characteristics for fossorial scavenging; the long first pereiopods, developed in stable soft-to-stiffgrounds. The burrow fill is also and with the dactyl extended outward, might have served as tactile organ frequently composed of coarse-grained quartz or bioclasts (Fig. 2F, for sensing and locating food (Feldmann et al. , 2002). 3B). The burrow diameter is presently at least three times wider than the width of M. rapax that are usually found inside Thalassinoides . b) Thalassinoides suevicus in association with M. rapax However it is common to overestimate Thalassinoides diameter due The burrows attributed to Thalassinoides suevicus are predominantly to the concretionary enlargement of the burrow during early horizontal systems, and visible at sedimentary interfaces preserved diagenesis (Fürsich , 1974; Fig. 2C). as convex hyporelief or providing a nodular fabric to bedding sections. The burrows are large and characterized by an irregular c) Causes for mass mortality in the Lusitanian Basin width that can exceed 110 mm, and their length can reach more than Vega et al. (2008) described numerous articulated (i.e., no separation 1 m bending occasionally. Burrows branch at acute angles, are Y- between the cephalothorax and the abdomen) specimens of Meyeria shaped and frequently have swellings in divergence areas (i.e., magna in concretions from a single horizon from the upper Aptian turning chambers). Transverse sections of the burrow are elliptical, of Colombia. Uniform size found in this example suggests a mass 148 C. Neto de Carvalho / Comunicações Geológicas (2016) 103, Especial I, 143-152

Fig. 4. Main microfacies found in the Thalassinoides suevicus fillings from Boca do Chapim Formation (Neto de Carvalho and Viegas, 2007). A – Thalassinoides fill of microfacies #2 type (intramicrosparite), with large mud clasts and Miliolidae. Compare with the pelmicritic host sediment. B – Biomicrite with Choffatella (microfacies #3) and the surrounding host sediment (mudstone). Note the lack of burrow wall. C – Microfacies #4: Intrabiomicrosparite packstone with sub-angular quartz grains showing a metamorphic (Variscan) source for the siliciclastics. Bar scale = 1 mm. Fig. 4. Principais microfácies definidas nos preenchimentos de Thalassinoides suevicus da Formação de Boca do Chapim (Neto de Carvalho and Viegas, 2007). A – Preenchimento de Thalassinoides correspondente à microfácies #2 (intramicrosparite), com grandes intraclastos de composição argilosa e Miliolidae. Compare-se com o sedimento pelmicrítico que envolve a galeria. B – Biomicrite com Choffatella decipiens (microfácies #3) e o sedimento que envolve a galeria (sem fósseis). De salientar, a falta de parede na galeria. C – Microfácies #4: Intrabiomicrosparite packstone com grãos de quartzo e quartzito sub-angulosos evidenciando uma fonte metamórfica (Varisca) próxima para os sedimentos detríticos. Escala gráfica = 1 mm. mortality event. They were described as preserved in concretions of the Lusitanian basin) to the lagoon sector. Freshwater runoff and also in Northern Spain and Isle of Wight, UK (see below). M. rapax increased clastic influx toward lagoons protected by rudist reefs led from the Lusitanian Basin represent dead bodies based on the to rapid reduction of salinity and the sudden filling of the lobster articulated side position of repose. They are corpses since they burrow systems. This environmental condition was optimal for the preserve the natural articulation of the cephalothorax and pleon, development of the euryhaline Chofatella decipiens but may have with the later commonly enrolled against the former. The preservation caused the instantaneous death to entire populations of Meyeria rapax of the crustacean producer within Thalassinoides has been smothered in their burrows. This explanation follows in general the understood since Bromley (1967) and Sellwood (1971) as involving subsurface obrution model of Tsujita (2003) for the burial and rapid burial of the whole burrowed level becoming almost preservation of nearly complete lobster populations. instantaneously filled with sediment and causing the death of the Most of the lobsters preserved within Thalassinoides found in the occupants (Tsujita, 2003). literature and also in the Lusitanian Basin are related with What special environmental causes were then responsible for eodiagenetical development of nodules, sometimes the only remaining the collective and instantaneously entombment of Meyeria populations evidence from the burrow systems (Fürsich, 1973; Tsujita, 2003) within their burrow systems, several times repeated during as much masked by the homogenization of the sediment due intensive as 1 Ma (Rey 1992), to apparently never happen again in all the bioturbation. Unusual circumstances, such as the one already described Lusitanian basin record? for obrutionary deposits, may have to exist to protect the corpses from Thin sections analysis performed by Neto de Carvalho and Viegas dissociation by scavengers, bacterial decay and bioturbators. Early (2007) to Thalassinoides fillings allowed to discriminate 4 diagenetic concretions may be the only media capable of preserving microfacies, all defined by the clear absence of abundant organic evidence of within-burrow preservation of decapods. remains (wood fragments) or pyrite, indicators of anoxia. Curiously, Favreina was not reported which could be the result of a predatory 4. Implications for behavioral convergence in thalassinoides or scavenging feeding habit outside the burrows for the producer, and during the mesozoic indicative of frequent interaction with the seafloor. Microfacies #1 is composed by biomicrite with abundant shell fragments and Thalassinoides is probably the dominant arthropod burrow in the dasycladaceans. On the other hand, microfacies #2 is formed by geologic record (Seilacher, 1986), ranging from Early Paleozoic to intramicrite (Fig. 4A). Both microfacies have sparse fine-grained the present. Low rates of deposition and low energy are optimal siliciclastics and almost no Choffatella. These are evidences for a fast conditions for burrowing crustaceans. Particularly, the Mesozoic infill of Thalassinoides with dense mud and pelleted mud flows from was a pivotal time in decapod evolution with adaptations to new the surroundings during storm events or strong tidal currents. environments including infaunalization (Klompmaker et al. , 2013), Biomicrites with Choffatella decipiens from the intermediary so O-T-S burrows became the commonest ones in shallow marine microfacies #3 (Fig. 4B), show more influence from coastal marine non-lithified substrates. In the Lusitanian Basin, since the Kimmeridgian sediments. On the other hand, microfacies #4, composed by Thalassinoides suevicus becomes the predominant ichnofabric until biomicrite wackestones and intrabiomicrosparite packstones (Fig. Late Cretaceous (Neto de Carvalho et al. , 2007, 2010). 4C), reveals an increasing influence of heterogranular siliciclastics Rift basins from the Mesozoic of Portugal record very interesting (abundant quartz and quartzite, angular to sub-rounded grains not and important contributions for understanding the phylogeny and larger than 4 cm). This microfacies indicates episodic flows of coarse biogeography of decapod crustaceans and relations with the evolution siliciclastics from adjacent fluvial braided systems (evidencing also of fossorial behavior and infaunalisation for several clades (Fig. 5). contributions from nearby alluvial fans located in the eastern border The decapod crustacean body fossils are restricted to the Oxfordian- Lobsters within Thalassinoides in the Lusitanian Basin 149

Fig. 5. Time-abundance chart of decapod body fossils and decapod-related trace fossils in the Mesozoic of Portugal (adapted from Mateus et al. , 2013). Fig. 5. Tabela comparativa Idade-Abundância para os somatofósseis e icnofósseis descritos no Mesozóico de Portugal (adaptado de Mateus et al ., 2013).

Kimmeridgian and Lower Cretaceous deposits from the Lusitanian radiation of thalassinidean decapods taking place during the Jurassic Basin. At least the following genera are known: Longodromites , and the Callianasoidea occurring at least at 173 Ma. So, there is Pithonoton , Eodromites , Magila, Meyeria, Mecochirus, and Glyphea some probability that the earliest O-T-S burrows in Late Paleozoic (Mateus et al. , 2013). The first three of those are brachyurans with a were already produced by decapods, besides shrimplike possible non-burrowing way of life. The axiid shrimp Magila sp. was malacostracans (Chamberlain and Baer, 1973; Carmona et al. , 2004; reported by Werner (1986) from the Thracia-Corbulomima Netto et al. , 2007; Baucon et al. , 2014). After the P-T extinction, assemblage (Early Kimmeridgian) at Consolação (Peniche Thalassinoides became very common in low latitude, shallow marine municipality). This is a small anomuran that can hardly be compared carbonate substrates, and this is correlated by Carmona et al. (2004) with the large size of most of Thalassinoides suevicus found in the with the appearance of callianassid body fossils. According with the Lusitanian Basin, inclusive in the same succession where it was phylogenetic analysis of Hyžný and Klompmaker (2015), ghost found. Three lobster genera are known: Mecochirus Germar, 1827 shrimps of Callianassidae and Ctenochelidae were major bioturbators (Mecochiridae) was reported from the Kimmeridgian Alcobaça only after Early Cretaceous. Also, very important during the Early Formation at Barrio (F.T. Fürsich, 1999 in http://paleodb.org); Mesozoic was the evolution of Glypheoidea (Karasawa et al. , 2013). Glyphea von Meyer, 1835 (Glypheidae) was documented from The fossorial superfamily Glypheoidea have a stratigraphic distribution Kimmeridgian sediments from Praia dos Salgados (Mateus et al. , between Permian and the present, the family Mecochiridae spanning 2013). The Valanginian-Hauterivian and lower Barremian from Hettangian to Maastrichtian, with climax of diversification “Mecochirus aff. longimanatus ” described in three different localities and geographical distribution in the Early Cretaceous (Karasawa et from Lisbon region (Ferreira, 1955: Mata, Algueirão and Praia do al. , 2013). Cavalinho, respectively) were revised as M. rapax by Förster (1971); Fiege (1944) suggested that either glypheoids lobsters or Meyeria rapax is also known from the Barremian Boca do Chapim thalassinideans shrimps produced Thalassinoides in the Muschelkalk . Formation ( Mecochirus rapax in Neto de Carvalho et al. , 2007) and The Pliensbachian of the Calcari Grigi Formation, in Trento, Ribeira de Ilhas (this work). These lobster genera are found within Northern Italy, is highly bioturbated with Thalassinoides suevicus large Thalassinoides suevicus , as it was described before, or in and Ophiomorpha irregulaire ; here Monaco and Garassino (2001) successions where this ichnospecies is common. found a single exuvia of the Erymid Phlyctisoma Bell, 1863. Thalassinoides is a good example of behavioral convergence. Bromley and Asgaard (1972) found abundant fragments of Glyphea It is known since the Cambrian (Myrow, 1995), and well established exuviae in Thalassinoides from the Lower Jurassic (Toarcian) of after Middle Ordovician (Droser and Bottjer, 1988), but most east Greenland. Sellwood (1971) described a single Glyphea udressieri probably was produced by other groups than decapod crustaceans (Meyer) lying on its left side at the bottom of a Thalassinoides from until Late Paleozoic (for a review, see Carmona et al. , 2004). the Bathonian of Oxfordshire, England. An exuvia of Glyphea According to decapods divergence time chronogram estimated rostrata have been found in Thalassinoides in the upper Oxfordian using topology of ML tree of Porter et al. (2005), members of the of Dorset (Fürsich, 1974). Mecochirus and their burrows were infraorder Thalassinidea first appeared 325 Ma ago, with the figured by Kühn (1973) from the Late Jurassic of Solnhofen. 150 C. Neto de Carvalho / Comunicações Geológicas (2016) 103, Especial I, 143-152

Lobsters were more diverse in the Mesozoic, especially during the Triassic and Jurassic and may have been the most common producer of Thalassinoides -type ichnofabrics in Late Jurassic (peaks of diversity in the Oxfordian and Tithonian, according to Schweitzer and Feldmann, 2014) and, as we register in the Lusitanian Basin and in previous works by other authors, extending in the Early Cretaceous up to the Aptian with the Mecochiridae. Mud shrimps only become major bioturbators of marginal-to-fully shallow marine substrates after the Hauterivian (Hyžný and Klompmaker , 2015). The genus Meyeria had a fairly wide distribution during Early Cretaceous times. M. rapax is known in Portugal, England, Germany and Argentina. Meyeria rapax is Valanginian-Hauterivian (Förster, 1971; Aguirre- Urreta, 1989, 2003; Astrop, 2011) to Barremian (Neto de Carvalho et al. , 2007) in age. Vega et al. (2008) described a remarkable example where abundant articulated Meyeria magna where found in nodules from a single bed from the upper Aptian of Colombia. Unfortunately, the authors did not provide indications of clear connection of the nodules with the burrowing activity of this species. Three Meyeria Fig. 6. Reconstitution of Meyeria rapax within Thalassinoides suevicus from the lower magna and Haploparia longimana carapaces were reported in the Barremian of the Lusitanian Basin (taken from Neto de Carvalho et al. , 2007). same site with Thalassinoides and other burrows in the Aptian of the Fig. 6. Reconstituição de Meyeria rapax no interior do seu sistema de galerias do tipo Oxfordshire (Goldring, 1996). Pemberton et al. (1984) reported large Thalassinoides suevicus para o Barremiano inferior da Bacia Lusitânica (retirado de Thalassinoides suevicus (15 cm in diameter) at the top of the Cardium Neto de Carvalho et al ., 2007). Formation, with a single lobster Linuparus canadensis Whiteaves (1885), the possible producer, found some meters above the burrowed We compared the occurrences in the Lusitanian basin with others horizon. Upper Cretaceous and Cenozoic Thalassinoides were mostly previously described, and with recent phylogenetic studies, for produced by callianassidae ( e.g. , Ehrenberg, 1938; Mertin, 1941; better understanding evolutionary trends of the fossorial behavior Glaessner, 1947; Fürsich, 1974; Hayward, 1976; Murray and Hanley, for several different groups of crustacean decapods, i.e., macrurans, 1986; Aguirre-Urreta, 1989; Grimm and Föllmi, 1994; Swen et al. , chelate and achelate lobsters and mud shrimps. A trend for 2001; Hyžný , 2011). However, Tsujita (2003) recorded numerous behavioral convergence seemed to have been developed for several palinuran lobsters of the species Palaeonephrops browni (Whitfield) different clades of crustacean decapods. Among them, several preserved in concretions from Late Campanian-Early Maastrichtian members of the Glyphoidea may have been the major architects for of Bearpaw Formation, Alberta. During this period, lobsters were Thalassinoides (Ophiomorpha and Spongeliomorpha included) outcompeted by the radiation of the thalassinideans and crabs, since, at least, the Triassic and up to Lower Cretaceous, although especially in carbonate environments (Karasawa et al. , 2013; being rapidly supplanted by the Mesozoic Marine Revolution Schweitzer and Feldmann, 2014; Hyžný and Klompmaker , 2015), agents, particularly the Eubrachyura and the thalassinideans. Our and most of the fossorial clades became extinct before the K-T study corroborates and remark previous works that feature extinction. Thalassinoides suevicus as a fine example of behavioral convergence, with different types of producers along its geological spanning time. 5. Conclusions It is expected to find new sites in the Lusitanian Basin, and elsewhere, with the joint preservation of O-T-S and their producers The joint preservation of any trace fossil and its producer is an whenever analogous environmental conditions to the ones found for exceptional phenomenon that opens a window to evolution. The the exceptional preservation of Meyeria rapax were reunited, in occurrence of Ophiomorpha-Thalassinoides-Spongeliomorpha mixed siliciclastic-carbonate shallow marine deposits. burrow systems, some of the commonest trace fossils found along the Phanerozoic times, together with their producers, are still very rare and worth to be described in detail. They provide not only Acknowledgments explanations for the exceptional macro- and microenvironmental We would like to thank to the National Museum of Natural History causes for the preservation of body and trace fossil records together, and Science (NMNHS) and the Geological Museum of Lisbon, in but also the evolutionary causes for the development of this specific particular to Vanda Santos, and Miguel Ramalho and Jorge type of fossorial behavior. Following these main reasons, we review Sequeira, respectively, to allow permission and provide support to the remarkable discover of hundreds of Meyeria rapax in study the collections with Mesozoic lobsters. The photographer Luis Thalassinoides suevicus from the lower Barremian of the Lusitanian Quinta is greatly acknowledged for providing the photo of the Basin (Fig. 6), and describe a new site from the same age and similar Meyeria kept in the collections of NMNHS. The colleague Mariana depositional condition, but separated by more than 70 km. In both Vilas Boas (UNESCO Naturtejo Geopark) is greatly appreciated for sites, the climate and tectonic causes for the collective burial and providing the Fig. 1. preservation of the lobsters in their burrows may be related with somewhat cyclic runoffs of freshwater together with the influx of terrigenous sediments that were responsible for sudden drops in salinity and cover of the confined carbonate lagoons with blankets of coarse grained sands mixed with mud clasts and biomicrite cement. This natural concrete filled the burrow systems almost instantaneously, and soon started diagenetical carbonate processes similar to those responsible for the development of concretions, that protected the articulated bodies from scavenging and deeper tiering bioturbation, wide bacterial decay, and skeletal dissolution. Lobsters within Thalassinoides in the Lusitanian Basin 151

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