Earth-Science Reviews 194 (2019) 306–326

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Earth-Science Reviews

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Muscles and muscle scars in fossil malacostracan crustaceans T ⁎ Adiël A. Klompmakera,b,c, , Matúš Hyžnýd,e, Roger W. Portellb, Clément Jauvionf,g, Sylvain Charbonnierf, Shane S. Fussellc, Aaron T. Klierh, Raymond Tejerac, Sten L. Jakobseni a Department of Integrative Biology & Museum of Paleontology, University of California, Berkeley, 1005 Valley Life Sciences Building #3140, Berkeley, CA 94720, USA b Florida Museum of Natural History, University of Florida, 1659 Museum Road, Gainesville, FL 32611, USA c Department of Geological Sciences, University of Florida, 241 Williamson Hall, Gainesville, FL 32611, USA d Department of Geology and Paleontology, Faculty of Natural Sciences, Comenius University, Mlynská dolina G1, Ilkovičova 6, SVK-842 15 Bratislava, Slovakia e Geological-Paleontological Department, Natural History Museum Vienna, Burgring 7, 1010 Vienna, Austria f Muséum national d'Histoire naturelle, Centre de Recherche en Paléontologie - Paris (CR2P, UMR 7207), CNRS, Sorbonne Université, 57 rue Cuvier, F-75005 Paris, France g Institut de Minéralogie, de Physique des Matériaux et de Cosmochimie, CNRS UMR 7590, Sorbonne Université, Muséum national d'Histoire naturelle, IRD UMR 206, 61 rue Buffon, F-75005 Paris, France h Department of Biology, University of Florida, 220 Bartram Hall, Gainesville, FL 32611, USA i Geomuseum Faxe, Østervej 2, DK-4640 Faxe, Denmark

ARTICLE INFO ABSTRACT

Keywords: Exceptionally preserved specimens yield critical information about the soft-part anatomy and the evolution of Crustacea organisms through time. We compiled the first global dataset of exceptionally preserved muscles in malacos- Decapoda tracans consisting of 47 occurrences, including 18 new records, predominantly preserved in Mesozoic Konservat- Exceptional preservation Lagerstätten (> 70% of occurrences). Early diagenetic mineralization through phosphatization is the dominant Lagerstätten process for exceptional preservation of muscles in malacostracans. Over 70% of taxon occurrences with muscles Malacostraca preserved are compressed. Rarer, three-dimensionally preserved specimens allow more detailed study of mus- Muscle cles. One example are specimens of the mid-Holocene ghost shrimp Sergio sp. from Panama, showing exquisitely preserved strings of muscle fibers attached to the shell interior, resembling the muscle arrangement ofmodern ghost shrimps. Other fossil malacostracans, including the oldest known fossil shrimp, also show musculature similar to modern taxa. We hypothesize that this muscle conservatism may be related to the confined space within the malacostracan shell in conjunction with the relatively stable body plan of several clades. We also assembled the first dataset on muscle attachment scars in malacostracans. Unlike muscles, muscle scars aremore common. Approximately 24% of the 357 articles analyzed yielded evidence of muscle scars, but such scars were only recognized for 19% of the 162 taxon occurrences that showed muscle scars. Muscle scars are common from the Late Jurassic onwards, are found primarily in Brachyura and Axiidea, and do not suffer from a Lagerstätten- effect. Rocks with well-preserved specimens should yield an additional wealth of information on thesoftpart anatomy of malacostracans. Similarly, muscle scars represent an almost untapped, complementary source of information on muscle evolution.

1. Introduction 1930; Feldmann and Schweitzer, 2010; Wilby and Briggs, 1997). The most diverse clade among malacostracans are the Decapoda, with a The preservation of soft tissues in fossils provides exceptional in- fossil record of ~3700 species (Schweitzer and Feldmann, 2016) and sight into the biology of ancient metazoans. Muscles have been used to major species- and genus-level diversifications from the Mesozoic on- infer the functioning of organisms, their taxonomic identity, and evo- wards (Klompmaker et al., 2013; Schweitzer and Feldmann, 2015; lutionary relationships among clades (e.g., Bergström et al., 1989; Kaji Sepkoski, 2000). Other, less diverse malacostracan clades with a mac- et al., 2018; Parry et al., 2015; Trinajstic et al., 2007; Young and rofossil record are Isopoda, Phyllocarida, and Hoplocarida. Muscles in Vinther, 2017). Muscles are preserved in a variety of arthropods (e.g., fossil malacostracans are almost exclusively mentioned as part of sys- Briggs et al., 2005; Hu et al., 2017; Siveter et al., 2010; Vannier et al., tematic descriptions thus far, but have the potential to provide insight 2014), including in multiple clades of Malacostraca (e.g., Beurlen, into the evolution of myoanatomy through time.

⁎ Corresponding author. E-mail address: [email protected] (A.A. Klompmaker). https://doi.org/10.1016/j.earscirev.2019.04.012 Received 21 December 2018; Received in revised form 8 April 2019; Accepted 8 April 2019 Available online 10 April 2019 0012-8252/ © 2019 Elsevier B.V. All rights reserved. A.A. Klompmaker, et al. Earth-Science Reviews 194 (2019) 306–326

Fig. 1. Thoracic/carapace muscles (A) and muscle scars (B) in the brachyuran decapod crustacean Dromia personata (Linnaeus, 1758). Modified from McLaughlin (1980: Fig. 53A) and redrawn from Glaessner (1969: Fig. 224).

Muscle attachment traces or scars on the hard parts of fossil me- 2.1. Thoracic muscles tazoans can provide complementary information on many aspects of metazoan biology in deep time. Muscle scars have helped gain insight Thoracic muscles (Fig. 1) provide support for the animal body and into the evolution of digestive tracts, the function of skeletal elements, interconnect body parts such as the thorax and pleon. The thorax in the size of soft tissues, life habit, and the relatedness among organisms decapods is rather inflexible and contains mainly thoracopleonal mus- (e.g., Doguzhaeva and Mutvei, 1991; Dzik, 2010; Knight and Morris, cles, in addition to those connecting the internal with the external 2019; Kröger and Mutvei, 2005; Lerosey-Aubril and Peel, 2018; Richter skeleton (Glaessner, 1969). Nevertheless, the thoracic musculature is and Fischer, 2002; Topper and Skovsted, 2017). Such scars have been extraordinarily complex and the number of individual muscles is large found in fossil malacostracans (e.g., Glaessner, 1960; Hof and Schram, in lobsters and most anomurans, whereas the whole arrangement in 1998; Schweigert and Koppka, 2011), but, again, they are primarily brachyurous crabs is less complicated (Pilgrim, 1973). In hoplocar- mentioned as part of taxon descriptions. idans, the thorax is more flexible and contains also bundles of flexors Here we review muscles and muscle scars in fossil malacostracans to and extensors (Pilgrim, 1964). In decapods, various paired bundles of evaluate (1) muscle preservation through geologic time based on dorsal thoracic muscles are situated either between the inner side of the known and new occurrences, (2) how fossil muscles compare to those in carapace and the endophragmal skeleton or between elements of the extant representatives, (3) the taxa with muscle attachment scars and endophragmal skeleton (Keiler et al., 2016). The attachment areas of their location, and (4) the frequency and the degree of recognition of muscles are recognizable on the inner surface of the carapace muscle scars. (Glaessner, 1960, 1969). Institutional abbreviations. GBA: Geologische Bundesanstalt, In decapods, the most important groups of muscles attached to the Vienna, Austria; MAB: Oertijdmuseum, Boxtel, The Netherlands; carapace are the epimeral attractor muscle (muscle inserting along the MGUH: Natural History Museum of Denmark, Geological Museum, branchiocardiac groove that connects the epimeral wall of the ventral University of Copenhagen, Denmark; MNHN: Muséum national thorax to the dorsal carapace), mandibular muscles (operating mand- d'Histoire naturelle, Paris, France; NHMW: Department of Geology and ibles), gastric muscles (responsible for moving the gastric mill), and a Palaeontology, Natural History Museum Vienna, Austria; SMF: small portion of the thoraco-pleonal muscles (interconnecting thorax Forschungsinstitut Senckenberg, Frankfurt am Main, Germany; UF: with pleon). Ventral thoracic muscles are extrinsic muscles (flexors + Florida Museum of Natural History at the University of Florida, extensors) of the bases of appendages and are located directly above the Gainesville, USA; UMJGPA: Geological-palaeontological Department, sternum (Keiler et al., 2016). There is considerable variation in the Universalmuseum Joanneum, Graz, Austria. development of mandibular muscles. In lobsters, the posterior (ad- ductor) muscle is very strong and its area of attachment covers a large 2. Major muscle groups and muscle scars of extant malacostracans part of the lateral internal surface of the carapace. In squat lobsters and with a fossil record crab-like decapods, this muscle is very weak and the lateral adductor muscle occupies more space on its surface (Glaessner, 1960). Skeletal muscles coupled with the nervous system provide the ne- Of the external gastric muscles, only the group of the posterior cessary equipment for organisms to actively move. The neuromuscular muscles is attached to the carapace. It is connected with two small morphology in malacostracans is responsible for distinct ways of mo- calcareous projections of its apodemes close to the median line of the tion, including swimming, tail-flipping (e.g., escape response, see carapace and recognizable from the outside as a pair of minute pits, also Heitler et al., 2000), digging, and burrowing. Skeletal muscles in the called gastric pits. The anterior gastric muscles are attached to the body regions and the appendages are generally organized into antag- lower surface of the rostrum; in brachyuran crabs, they are attached to onistic pairs, such as extensor/flexor, levator/depressor, and promotor/ small flat calcareous projections underneath the epigastric regions of remotor muscle pairs. Skeletal muscles work against a mechanical the carapace. system of levers formed by the exoskeleton, apodemes, and joints of the segments; the movement of appendages results from combinations of 2.2. Pleonal muscles movements of several joints (Evoy and Ayers, 1982). Muscles in ma- lacostracans typically are striated muscles, with actin and myosin fila- Within malacostracan pleons (Figs. 2–3), there are dorsal extensor ments. The fiber type, whether fast (phasic), slow (tonic) or inter- muscles that straighten the pleon and larger ventral flexor muscles mediate, in homogeneous muscles is largely genetically determined (Herrick, 1909) that are responsible for swimming in shrimps (Onnen (Wahle et al., 2012). and Zebe, 1983) and the effective stroke of the “tail flip” escape

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Fig. 2. Thoracic and pleonal muscles in decapod crustaceans. A. Thoracic and pleonal musculature in lobsters as exemplified in an astacid crayfish (redrawn from Schmidt, 1915: Fig. 1). B. Muscles of the telson and the sixth pleonal segment in lobsters and shrimps (redrawn from Paul et al., 1985: Fig. 1). Top = cambarid crayfish; middle = upogebiid mud shrimp; bottom = pandalid caridean shrimp. Left column = axial muscles; right column = appendage muscles. C.Transverse section through the fourth pleonal segment in the squat lobsters Munida Leach, 1820, (top) and Galathea Fabricius, 1793, (middle) and a porcelain crab Petrolisthes Stimpson, 1858, (bottom) (redrawn from Keiler et al., 2017: Fig. 13). response (caridoid escape reaction, typical for shrimps and lobsters). 1980; Pilgrim, 1973). Dorsal, ventral, and lateral pleonal muscles are recognized depending The pleonal muscles differ greatly between the higher malacos- on their position. The dorsal pleonal muscles belong to the extensor and tracan taxa. They are most pronounced in stomatopods, shrimps, and the ventral muscles to the flexor system of the pleon (Herrick, 1909; lobsters, occupying most of the pleon. In astacidean lobsters, the pleon Huxley, 1880; Keiler et al., 2017; Young, 1959). The lateral muscles are contains strong muscular bundles. Well-developed musculature of the relatively thin, fan-shaped, and apparently function to retain the posi- sixth pleomere and telson in astacideans (Herrick, 1909) is used in tion of the cuticle during movements of the animal (Young, 1959). The rapid escape flexions of the pleon and tail fan. The pleonal musculature pleonal muscles are bilaterally symmetrical in most malacostracans, is similar in axiideans and gebiideans, except for the absence of the except in hermit crabs and their relatives (Glaessner, 1969; McLaughlin, ventral telsonal flexor muscles (Paul et al., 1985). The arrangement of

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Fig. 3. Pleonal muscles in the stomatopod Squilla empusa Say, 1818. Modified from Hessler (1964: Fig. 44) and redrawn Moore and McCormick, 1969: Fig. 29).

Fig. 4. Pereopod muscles in decapod crustaceans. A. Apodemes of a brachyuran claw. B. Dorsal view of the distal musculature of a walking leg (top) and a cheliped (bottom) of a majoid crab (redrawn from Vidal-Gadea and Belanger, 2009: Fig. 2). C. Inner and outer sides of the cheliped of the extant ghost shrimp Gourretia phuketensis Sakai, 2002, (SMF 29520) showing muscle bundles arranged in a reticulate pattern through the translucent cuticle (from Hyžný and Klompmaker, 2015: Fig. 8A–B). the sixth pleonal and telson neuromusculature in axiideans and gebii- but highly conserved (Abrahamczik-Scanzoni, 1942; Vidal-Gadea and deans is considered a secondary adaptation to their burrowing behavior Belanger, 2009). Proximal muscles originate from the endophragmal (Dworschak et al., 2012). A reduction of pleonal and tailfan muscles is skeleton and its mass relative to the distal musculature is limited (Vidal- observed in most anomurans (Keiler et al., 2015, 2016, 2017). The Gadea and Belanger, 2009). In malacostracans, pereopods typically proportional volume of pleonal and tailfan musculature is even lower in consist of six segments. For muscles in distal segments, all but two crab-like (carcinized) anomurans and in brachyurans. muscles have common features between the different pereopods. In most cases, the pereopod segments articulate via bicondylar joints, re- 2.3. Walking leg muscles stricting the range of motion of each segment to a single plane. The relatively simple muscle arrangement consists of openers, closers, The muscles of the walking legs (thoracic appendages) are attached stretchers, benders, extensors, flexors, and reductors (Vidal-Gadea and to the internal (endophragmal) skeleton and do not reach the carapace Belanger, 2009). (Glaessner, 1960)(Fig. 4B). The musculature of walking legs is complex

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Fig. 5. The raw number of muscle taxon occurrences 20 Malacostracan clade per period per malacostracan clade. Data are not normalized for the unequal duration of periods. For Achelata Gebiidea all data, see Table 1. Not included are the phyllocarid Aeschronectida Glypheidea in Briggs (1978) that has been reassigned to a non- 15 Astacidea Penaeoidea malacostracan clade (e.g., Aria and Caron, 2017); the Axiidea Peracarida muscles in Zatoń and Broda (2015) may have been of phyllocarid origin, but this could not be confirmed; Brachyura Phyllocarida and an unidentified decapod from La Voulte-sur- 10 Decapoda indet. Polychelida Rhône (Wilby et al., 1996) because this specimen Erymida Stomatopoda could be conspecific to a taxon in Table 1.

5 Occurrences with muscles preserved Occurrences with muscles

0

Silurian Permian Triassic Jurassic Cambrian Devonian Neogene Ordovician Cretaceous Paleogene Quaternary Carboniferous

2.4. Cheliped muscles Allmon, 2015). A larger apodeme area allows for more muscle attach- ments and, hence, to a large extent, the respective muscle scars on the The claw represents a relatively simple motor system with only two propodal surface. We hypothesize that the surface area of the closer antagonistic muscles, an opener (abductor) and a closer (adductor), muscle scars on the lateral surfaces of the cheliped propodus may be which control the movements of the dactylus (Huxley, 1880; Kaji et al., positively correlated with the force of the claw. Muscles in arthropods 2018; Pillai, 1990)(Fig. 4A, C). Decapod claws in heterochelous taxa in general are mechanically linked to the epidermis and cuticle by a often differ in the type of muscle fibers. Average sarcomere length complex network of cytoskeletal and junctional elements (Žnidaršič determines whether muscle fibers are slow- or fast-moving; muscles et al., 2012), and these can be expressed as the muscle scars on cuti- with a shorter sarcomere length tend to be faster in their contraction. cular surfaces. The fibers of the muscle attachments go through the The major (crusher) claw is stout and its closer muscle is composed of cuticle; the fibers traverse the entire new cuticle during the molting slow fibers, so the claw closes strongly but slowly, whereas the minor process, and extend from the tendon cell apex up to the exocuticular (cutter) claw is slender and closes very rapidly (Govind et al., 1987); its layer in the old cuticle (Žnidaršič et al., 2012). Unfortunately, data on closer muscle is made up of predominantly fast fibers and a small how the muscle attachment sites change the properties of the crusta- ventral band of slow fibers. However, sarcomere length undergoes cean cuticle are very limited (see Buchholz and Buchholz, 1989; changes during development (Govind et al., 1987). In fact, differ- Yamada and Keyser, 2009; Žnidaršič et al., 2012). entiation of fiber type in lobster claws occurs during the post-larval period, resulting in an asymmetrical ganglion development that gives 3. Muscles rise to a major claw with exclusively slow fibers and a cutter claw with predominantly fast fibers (Wahle et al., 2012, and references therein). 3.1. Processes of muscle preservation The closer muscle in the paired claws of lobsters, crayfishes, and snapping shrimps undergoes transformation of its fiber composition Muscles in metazoans are among the most commonly preserved soft during development, regeneration, and reversal (Govind et al., 1987). tissues, even though they are among the most decay-prone parts (Parry To enable withdrawal of the chelipeds from the molt, claw muscle et al., 2018). When preserved, they are often phosphatized (e.g., Briggs atrophy occurs (Mykles, 1999). The mass of the claw muscle may de- et al., 1993; Trinajstic et al., 2007; Wilby et al., 1996; Wilson et al., crease about 40% in about one month before ecdysis; mass is restored in 2016), but sometimes silicified (Robin et al., 2018) or preserved in car- a month after successful exuviation (Skinner, 1966). This temporary bonates (Briggs et al., 1997). Phosphatization is the most common type decrease may affect the preservation potential of muscles if the animal of muscle preservation, which has been thoroughly explored and re- dies during that time. Atrophy is associated with a decrease in myo- produced in taphonomic experiments (Briggs et al., 1993; Briggs and fibrilar cross-sectional area (Mykles and Skinner, 1981), but the number Kear, 1994, 1993; Kear et al., 1995; Sagemann et al., 1999). These ex- of myofibrils and muscle fibers remain relatively constant throughout periments show that extensive and high-quality phosphate muscle pre- the molting process. Atrophy occurs in the claw muscle, but not in servation (mineral morphology replication) is mainly driven by a few muscles of the walking legs (El Haj, 1999, and references therein). Slow interconnected factors: favoring conditions include a high microbial ac- fibers, restricted to the major claw, show more evidence of atrophy than tivity, a low pH, and high ion availability. One of the most prominent slow fibers in the minor claw (Mykles, 1997). factors controlling phosphatization is pH: in a closed, lower pH (micro) environment, phosphates tend to form, whereas in an open, more alka- 2.5. Muscle scars line environment carbonate precipitation dominates (Briggs and Wilby, 1996; Sagemann et al., 1999). Three-dimensional preservation of soft Muscle scars are attachment spots for the muscles to the cuticle of tissues, including muscles, is most frequently found in combination with the appendages (including apodemes), carapaces, and pleons of mala- phosphatization (Allison and Briggs, 1991). Several taphonomic experi- costracans. In decapods, the cheliped dactylus functions as a lever. The ments have been conducted on malacostracans (Briggs and Kear, 1994; lateral closer apodeme surface area, together with the claw geometry, Hof and Briggs, 1997; Klompmaker et al., 2017: Table 1; Sagemann et al., the angle of pinnation of closer muscles, and the resting sarcomere 1999). Consequently, the conditions under which muscles can preserve length, is positively correlated with the force produced (Kosloski and in malacostracans are relatively well-known.

310 ..Kopae,e al. et Klompmaker, A.A. Table 1 All malacostracan taxa with muscles preserved based on the literature and new records arranged by age. * indicates that specimen(s) of this taxon were previously studied in the accompanying paper, but muscles are reported here for the first time. § Taxon may be conspecific to the taxon in the following row(see Schweigert, 2001).

Clade Taxon Locality Stratigraphic unit Preservation Preservation Konservat- Age (young to old) Part with muscle Source specimen: compressed process Lagerstätte? preserved (C) or three- dimensional (3D)

Axiidea Sergio sp. Amador and Farfan beaches, – 3D Phosphatization No mid-Holocene Major cheliped herein near Panama City, Panama propodi

Brachyura Pinnixa sp. Cape Blanco, Oregon, USA – 3D No Late Pleistocene Venter Zullo and Chivers (1969)

⁎ Stomatopoda Squilla taulinanus Ahyong Taulignan (La Croix de – Compressed No Miocene (Burdigalian) Pleon Ahyong et al. (2013); pers. et al., 2013 Bouchet), France comm. Carolin Haug 25 April 2017

Gebiidea Upogebia aronae Haug Bressi Ranch project site, Santiago Fm Compressed ?Phosphatization Not middle Eocene Pleomere 6 Haug et al. (2013) et al., 2013 California, USA mentioned

Stomatopoda Trelde Næs near Fredericia, Lillebælt Clay Fm 3D No Eocene (late Ypresian- Pleon herein Denmark middle Lutetian)

Astacidea Aenigmastacus crandalli McAbee locality, British Kamloops Gr, Compressed Y Eocene (Ypresian) Shadows of muscle Feldmann et al. (2011) Feldmann et al., 2011 Columbia, Canada McAbee beds in propodus and pleon

Stomatopoda Lysiosquilla antiqua Monte Bolca, Italy Monte Bolca Compressed Y Eocene (Ypresian) Pleon Hof and Briggs (1997); 311 (Münster, 1842) Lagerstätte Secrétan (1975)

⁎ Penaeoidea Aeger sp. Sahel Alma, Lebanon Sahel Alma Compressed Y Late Cretaceous (late Pleon Charbonnier et al. (2017) Lagerstätte Santonian)

⁎ Penaeoidea Palaeobenthonectes Sahel Alma, Lebanon Sahel Alma Compressed Y Late Cretaceous (late Pleon Charbonnier et al. (2017) arambourgi (Roger, 1946) Lagerstätte Santonian)

⁎ Penaeoidea Carpopenaeus garassinoi Sahel Alma, Lebanon Sahel Alma Compressed Y Late Cretaceous (late Pleon Petit and Charbonnier, 2012 Charbonnier in Petit and Lagerstätte Santonian) Charbonnier, 2012

Penaeoidea Palaeobenthesicymus Sahel Alma, Lebanon Sahel Alma Compressed ?Silicification and Y Late Cretaceous (late Mainly in pleonal Audo and Charbonnier (2013) libanensis (Brocchi, 1875) Lagerstätte ?phosphatization Santonian) region and pereiopods

Achelata Scyllaridae Rancho El Pilote, Coahuila, Eagle Ford Gr Compressed Phosphatization Not Late Cretaceous (early- Pereopods Vega et al. (2007) Mexico mentioned middle Turonian) Earth-Science Reviews194(2019)306–326

Brachyura Cenomanocarcinus Rancho El Pilote, Coahuila, Eagle Ford Gr Compressed Phosphatization Not Late Cretaceous (early- Carpus to dactylus Vega et al. (2007) vanstraeleni Stenzel, 1945 Mexico mentioned middle Turonian)

Brachyura Opthalmoplax spinosus Villa de Leiva - Samacá, San Rafael Fm Compressed Y Late Cretaceous (early Propodi Feldmann et al. (1999: Feldmann et al., 1999 Department of Boyacai, Turonian) Fig. 3.2), according to Vega Colombia et al. (2007)

Penaeoidea Cretapenaeus berberus Djebel Oum Tkout, Morocco Kem Kem beds Compressed Phosphatization Y Late Cretaceous Pleonal somite 2 Garassino et al. (2006); Garassino et al., 2006 (Cenomanian) Gueriau et al. (2014: soft tissue)

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Clade Taxon Locality Stratigraphic unit Preservation Preservation Konservat- Age (young to old) Part with muscle Source specimen: compressed process Lagerstätte? preserved (C) or three- dimensional (3D)

⁎ Penaeoidea Epipenaeus abisaadorum En Nammoura, Lebanon En Nammoura Compressed Y Late Cretaceous Pleon Charbonnier et al. (2017) Charbonnier et al., 2017 Lagerstätte (Cenomanian)

⁎ Achelata Palinurus teruzzii Hakel, Lebanon Hakel Lagerstätte Compressed Y Late Cretaceous Pleon Charbonnier et al. (2017) Charbonnier et al., 2017 (Cenomanian)

Decapoda Shrimp-like decapod Chapada do Araripe, Ceara, Santana Fm, ? Phosphatization Y mid-Cretaceous (Aptian- Thoracic segment Wilby and Martill (1992); age indet. Brazil Romualdo Mbr Cenomanian) from Martill (2007)

Stomatopoda Nodosculda fisherorum Fort Worth area, Texas, USA Pawpaw Fm 3D Not mid-Cretaceous (late Pleon Franţescu (2012) Franţescu, 2012 mentioned Albian)

Decapoda Decapods Las Hoyas, Spain La Huérguia Fm Compressed Phosphatization Y Early Cretaceous – Briggs and Wilby (1996) indet. (Barremian)

Penaeoidea §Antrimpos Southern Germany Solnhofen Compressed Phosphatization Y Late Jurassic – Briggs and Kear (1994) Limestone (Tithonian)

Penaeoidea Antrimpos speciosus Southern Germany Solnhofen Compressed Y Late Jurassic Pleon MNHN.F.A33519; herein Münster, 1839 Limestone (Tithonian)

Penaeoidea Aeger tipularius (von Southern Germany Solnhofen Compressed Phosphatization Y Late Jurassic – Wilby and Briggs (1997) 312 Schlotheim, 1822) Limestone (Tithonian)

Penaeoidea Penaeid crustacean Southern Germany Solnhofen Compressed Phosphatization Y Late Jurassic – Wilby and Briggs (1997) Limestone (Tithonian)

Polychelida Cycleryon bourseaui Audo Canjuers Lagerstätte, Var Calcaires blancs de Compressed Y Late Jurassic (early Pleopods Audo et al. (2014) et al., 2014 Department, France Provence Fm Tithonian)

Glypheidea Gigacerina saemanni Cerin, France Cerin Lithographic Compressed Y Late Jurassic Pleon herein (Oppel, 1861) Limestone Fm (Kimmeridgian- Tithonian)

Decapoda Decapod Nusplingen, Germany Nusplingen Compressed Phosphatization Y Late Jurassic (late – Briggs et al. (2005) indet. Plattenkalk Kimmeridgian)

Decapoda Shrimp Cerin, France Cerin Lithographic Compressed Phosphatization Y Late Jurassic – Wilby and Briggs (1997) indet. Limestone Fm (Kimmeridgian- Tithonian) Earth-Science Reviews194(2019)306–326

⁎ Glypheidea Glypheopsis voultensis La Voulte-sur-Rhône, France La Voulte-sur- 3D Y Middle Jurassic Carapace and pleon Charbonnier et al. (2013) Charbonnier et al., 2013 Rhône Lagerstätte (Callovian)

Polychelida Voulteryon parvulus Audo La Voulte-sur-Rhône, France La Voulte-sur- 3D Y Middle Jurassic Carapace Jauvion et al. (2016) et al., 2014 Rhône Lagerstätte (Callovian)

⁎ Polychelida Cycleryon romani Audo La Voulte-sur-Rhône, France La Voulte-sur- Compressed and 3D Y Middle Jurassic Carapace and pleon Audo et al. (2014) et al., 2014 Rhône Lagerstätte (Callovian)

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Clade Taxon Locality Stratigraphic unit Preservation Preservation Konservat- Age (young to old) Part with muscle Source specimen: compressed process Lagerstätte? preserved (C) or three- dimensional (3D)

⁎ Polychelida Eryon ellipticus Van La Voulte-sur-Rhône, France La Voulte-sur- 3D Y Middle Jurassic Carapace and pleon Charbonnier (2009) Straelen, 1923 Rhône Lagerstätte (Callovian)

⁎ Polychelida Proeryon giganteus (Van La Voulte-sur-Rhône, France La Voulte-sur- Compressed and 3D Y Middle Jurassic Carapace and pleon Audo et al. (2014); SC pers. Straelen, 1923) Rhône Lagerstätte (Callovian) obs.

⁎ Polychelida Willemoesiocaris ovalis La Voulte-sur-Rhône, France La Voulte-sur- 3D Y Middle Jurassic Carapace and pleon Audo et al. (2014); SC pers. (Van Straelen, 1923) Rhône Lagerstätte (Callovian) obs.

⁎ Polychelida Hellerocaris falloti (Van La Voulte-sur-Rhône, France La Voulte-sur- Compressed and 3D Y Middle Jurassic Carapace and pleon Audo et al. (2014); SC pers. Straelen, 1923) Rhône Lagerstätte (Callovian) obs.

Penaeoidea Archeosolenocera straeleni La Voulte-sur-Rhône, France La Voulte-sur- Compressed and 3D Y Middle Jurassic Carapace and pleon herein; Jauvion et al. (2017) Carriol and Riou, 1991 Rhône Lagerstätte (Callovian)

⁎ Penaeoidea Aeger brevirostris Van La Voulte-sur-Rhône, France La Voulte-sur- Compressed and 3D Y Middle Jurassic Carapace and pleon Charbonnier (2009) Straelen, 1923 Rhône Lagerstätte (Callovian)

⁎ Erymida Eryma ventrosum (von La Voulte-sur-Rhône, France La Voulte-sur- 3D Y Middle Jurassic Carapace and pleon Charbonnier et al. (2010); SC Meyer, 1835) Rhône Lagerstätte (Callovian) pers. obs.

Polychelida Proeryon laticaudatus Holzmaden, Germany Posidonia Shale Compressed Y Early Jurassic Cheliped and pleon Beurlen (1930) 313 Beurlen, 1928 (as P. Fm (Toarcian) giganteus Beurlen, 1930)

Decapoda shrimp Osteno, Italy Lombardische Compressed Phosphatization Y Early Jurassic – Wilby and Briggs (1997) indet. Kieselkalk Fm (Sinemurian)

Peracarida Tealliocaris robusta Peach, Bearsden, Glasgow, Limestone Coal Fm Compressed ?Phosphatization Not Carboniferous Tergite Clark (2013) 1908 Scotland mentioned (Serpukhovian- Bashkirian)

Peracarida Tealliocaris woodwardi Gullane, Scotland Gullane shrimp- Compressed Phosphatization Not Carboniferous (Visean) Possible traces of Briggs and Clarkson (1985) (Etheridge, 1877) bed, Gullane Fm mentioned muscles in pleon

Aeschronect- Joanellia lundi Schram Fergus County, Montana, Heath Shale Compressed Not Carboniferous Mandible and Schram and Schram (1979) ida and Schram, 1979 USA mentioned (Mississippian) pleopod muscles

Penaeoidea Aciculopoda mapesi Ryan Quarry, Oklahoma, Woodford Shale Compressed Phosphatization Not Late Devonian Pleon Feldmann and Schweitzer Feldmann and USA mentioned (Famennian) (2010) Earth-Science Reviews194(2019)306–326 Schweitzer, 2010

Phyllocarida Montecaris gogoensis Locality 92, Canning Basin, Gogo Fm Compressed Phosphatization Not Middle-Late Devonian ?carapace Briggs et al. (2011) Briggs et al., 2011 Western Australia mentioned (Givetian-Frasnian)

Phyllocarida Nahecaris stuertzi Hunsrück and Taunus Hunsrück Slate Compressed Pyritization Y Early Devonian Carapace adductor Bergström et al. (1987) Bergström et al., 1987 regions, Germany muscle scar

Phyllocarida Ceratiocaris Site on the Upper Iowa Winneshiek Compressed Phosphatization Y Middle Ordovician Posterior carapace Briggs et al. (2015) winneshiekensis Briggs River, Iowa, USA Lagerstätte (Darriwilian) and anterior pleon et al., 2015 A.A. Klompmaker, et al. Earth-Science Reviews 194 (2019) 306–326

(e.g., Charbonnier et al., 2014), the Late Cretaceous of Sahel Alma in Lebanon (e.g., Charbonnier et al., 2017), and the Late Jurassic of Solnhofen in Germany (e.g., Garassino and Schweigert, 2006). Due to this Lagerstätten-effect, trends through time in muscle occurrences do not reflect diversity patterns. For example, decapod crustaceans first became diverse in the Late Jurassic (Klompmaker et al., 2013), but their diversity continued to rise into the Cenozoic (Schweitzer and Feldmann, 2015; Sepkoski, 2000), from which relatively few occurrences of muscle preservation are known. Marine, malacostracan-bearing Lagerstätten are primarily known from the Mesozoic. Most malacostracans with muscles preserved are Decapoda, but muscles are also known from phyllocarids and hoplocarids (including stomatopods and aeschronectids). Within decapods, brachyurans or true crabs became the dominant clade beginning in the latest Cretaceous (Klompmaker et al., 2013; Schweitzer and Feldmann, 2015), but they are scarcely represented in our list of muscle occurrences (Fig. 5; Table 1). Brachyurans first became abundant in the Late Jur- assic, predominantly found in coral- and sponge-associated habitats Fig. 6. The degree of compaction of muscles preserved in fossil malacostracan occurrences. 3D = muscles preserved in three dimensions; com- (Klompmaker et al., 2013; Krobicki and Zatoń, 2008). However, they pressed = muscles preserved in two dimensions. See Table 1 for all data. are rarely found in the Tithonian Solnhofen Lagerstätte in southern Germany (Feldmann et al., 2016), close to coral- and sponge-associated habitats inhabited by many brachyurans (e.g., Schweitzer and 3.2. Muscle preservation through time per malacostracan clade Feldmann, 2009; Wehner, 1988). Likewise, brachyurans are also absent in the Callovian La Voulte-sur-Rhône Lagerstätte in France After a comprehensive search of the literature for muscles preserved (Charbonnier et al., 2014: Table 1), but time-equivalent crabs are found in malacostracans supplemented with new occurrences, we report a in oolitic limestones in France (Crônier and Boursicot, 2009). Although total of 47 occurrences. Most of them originate in the Jurassic and brachyurans are more common in Cretaceous Lagerstätten (e.g., Cretaceous (Fig. 5; Table 1). Most occurrences are from Konservat-La- Charbonnier et al., 2017; Klompmaker, 2013: Appendix A; Larghi, gerstätten, which is not surprising because such Lagerstätten are de- 2004), they still do not dominate decapod assemblages. fined based on the presence of soft tissues and well-articulated speci- mens (Seilacher, 1970). Most occur in the three malacostracan-rich Lagerstätten: the Middle Jurassic of La Voulte-sur-Rhône in France

Fig. 7. Best-preserved previously known muscles in fossil malacostracans. A–B. The Devonian shrimp Aciculopoda mapesi Feldmann and Schweitzer, 2010, with pleonal muscles, from the USA (Oklahoma) (from Feldmann and Schweitzer, 2010: Figs. 3C, 4). C–D. Muscles preserved near the telson of the mud shrimp Upogebia aronae Haug et al., 2013, from the Eocene of Mexico, with arrows indicating muscle fibers (from Haug et al., 2013: Fig. 6A–B). E. Likely muscles in the thoracic region of the crab Pinnixa sp. (width = 9.2 mm) from the Pleistocene of the USA (Oregon) (from Zullo and Chivers, 1969: Pl. 1.1).

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Fig. 8. Muscles preserved in major cheliped propodi of the mid-Holocene ghost shrimp Sergio sp. from the Pacific side of Panama (Farfan/Amador beaches). A–C. Outer side of propodi with increasingly exposed and well-preserved muscles. D. Bundles attached to the inner side of the cuticle. E. Backscatter SEM image of strings of muscles. F. Light photography image of lowermost muscle bundle in E. G. SEM image of sheath (left) around muscle bundle. H–I. SEM images of different magnifications of the muscle fibers. Estimated width for D = ~2 mm. A: UF 126596; B: UF 126598; C–I:UF251887.

3.3. Types of muscle preservation proportionally and volumetrically. Therefore, under the assumption of equal preservational potential across muscle groups, muscles preserved Most muscles are preserved in compressed specimens rather than in in pleons may be found more frequently. However, preservation po- 3-dimensionally (3D) preserved specimens (Fig. 6). Most of the 3D- tential may not be equal for each muscle type as suggested for a preserved occurrences are from the La Voulte-sur-Rhône Lagerstätte, phosphatized Cretaceous polychaete (Wilson et al., 2016), but this as- whereas other occurrences of that Lagerstätte were represented by both pect has not been investigated for malacostracans thus far. Moreover, compressed and 3D-preserved specimens. Other 3D-preserved speci- gut-derived microbes appear to play a key role in the preservation of mens with muscle are found primarily in the Cenozoic (Zullo and internal anatomy by consuming tissue but also mediating authigenic Chivers, 1969; herein). The muscles in the compressed specimens mineralization in the branchiopod brine shrimp Artemia Leach, 1819, consist predominantly of re-mineralized muscles and sometimes out- after rupture at the mid- to hindgut junction (Butler et al., 2015). Dif- lines of muscles (e.g., Feldmann et al., 2011). Muscles in malacos- ferent guts are found in the carapace to the pleonal region in mala- tracans are primarily preserved through phosphatization (Table 1). costracans (McGaw and Curtis, 2013). Thus, the position of gut wall rupture may be critical for differential preservation of muscles within an individual in a closed system. However, external microbes may also 3.4. Location of preserved muscles enter the body cavity in different manners (e.g., mouth, at appendage joints after disarticulation) and could affect muscle preservation. Preserved muscles are most frequently found in the pleon (27/47 occurrences), whereas they occur less often in appendages (9/47) and in the carapace/cephalothorax (13/47). The pleons of malacostracans 3.5. Known and new preserved muscles in fossil malacostracans including Penaeoidea, Polychelida, and Stomatopoda contain primarily muscles for swimming and other movement purposes (e.g., Hertzler and Most occurrences of muscles in malacostracans have either been Freas, 2009; Vogt, 2002; Fig. 2). Conversely, the carapace of malacos- figured and mentioned as part of papers focusing primarily onsys- tracans contains a variety of other tissues as well (Davie et al., 2015; tematics (e.g., Audo and Charbonnier, 2013; Haug et al., 2013; Vega Warner, 1977; Fig. 1), implying that the pleon contains more muscles et al., 2007) or are part of papers about a variety of aspects of

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Fig. 9. Monocrystal x-ray diffraction and SEM energy-dispersive x-ray spectroscopy analyses of the muscle and cuticle of the cheliped major propodi of themid-Holocene ghost shrimp Sergio sp. from the Pacific side of Panama (Farfan/Amador beaches). A–B. Monocrystal XRD graphs of the muscle fibers (A) and cuticle (B) ofspecimenUF 251889. A. Diffractogram of a muscle fiber microsampling, attributed to hydroxyapatite and/or fluorapatite (peaks noted with a). B. Diffractogram ofacuticlemicro- sampling, attributed to magnesian calcite (peaks noted with c). C–D. Elemental map produced by an SEM-EDS (energy-dispersive x-ray spectroscopy) showing elevated concentrations of phosphorous indicated by purple dots where mineralized muscle is present, but not on the cuticle present on the very bottom part (specimen UF 251887). The electrons emitted were unable to reach and/or were reflected/emitted so that they were not detected for some deeper parts of the specimen, including the‘hole’in center-right. (For interpretation of the references to color in this figure legend, the reader is referred to the web version of this article.) preservation (e.g., Briggs and Kear, 1994; Jauvion et al., 2016). Other papers merely mention but do not show the presence of muscles (Franţescu, 2012). Some of the best figured examples of muscles in fossil malacostracans are shown in Fig. 7. A survey of material in the collection of the Florida Museum of Natural History (AAK), part of collection of the Muséum National d'Histoire Naturelle (SC, CJ), and part of the collection of the Natural History Museum of Denmark (SLJ) has yielded 18 new taxon occur- rences with muscles preserved. While all new occurrences can be found in Table 1, we here highlight a number of particularly well-preserved specimens. One example are major cheliped propodi from ghost shrimp Sergio sp. from the Pacific side of Panama. These specimens originate from the mid-Holocene (5015 ± 50 years BP based on AMS C-14 analysis of the co-occurring bivalve Lirophora mariae (d'Orbigny, 1846) [UF 233967]). Multiple specimens show evidence of remineralized muscles of varying quality (Fig. 8A–C). One specimen in particular shows superbly pre- served muscles (Fig. 8C–I), one of the best examples in the malacos- tracan fossil record. Muscles are attached to the inner side of the cuticle (Fig. 8D) and, at the largest scale, are composed of bundles that merge at the point of attachment to the cuticle. These bundles consist of fibers (Fig. 8F) and the individual fibers, in turn, consist of smaller fibers (Fig. 8H). No more details can be seen at even higher magnification (Fig. 8I). Some fibers appear to be surrounded by a sheath-like structure (Fig. 8F, middle left part; Fig. 8G). A microCT scan (Appendix 1) sug- gests that additional muscles are preserved inside this specimen un- derneath the cuticle. The muscle remains are phosphatized to hydro- xyapatite and/or fluorapatite, whereas the cuticle is composed of magnesian calcite as suggested by a combination of monocrystal x-ray diffraction (Fig. 9A–B) and energy-dispersive x-ray spectroscopy Fig. 10. Muscles preserved in the fossil dendrobranchiate shrimp Archeosolenocera (Fig. 9C–D). The muscle bundles are part of the adductor (closer) straeleni Carriol and Riou, 1991, from the Middle Jurassic (Callovian) of the La muscle, whereas obvious muscles belonging to the opener muscle are Voulte-sur-Rhône Lagerstätte in France (MNHN.F.A58641). A sagittal polished not visible (Fig. 8C). However, a plate is located at the position where section was prepared from the pleon to show muscle fibers. A. Entire specimen opener muscles would have been (Fig. 8C, top part). This plate may (photograph by Peter Massicard). B–C. SEM in lens detector image of muscle fibers, represent an apodeme that served as an attachment surface for these muscle fibers in light grey (exact location within pleon not known). C. Magnified muscles (Fig. 4A), as suggested previously for a Late Jurassic axiid image of lower middle part of B.

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Fig. 11. The fossil glypheid lobster Gigacerina saemanni (Oppel, 1861) from the Late Jurassic (Kimmeridgian/Tithonian) lithographic limestones of the Cerin La- gerstätte, France, and muscles from the extant glypheid lobster Neoglyphea inopinata Forest and Saint Laurent, 1975, for comparison. A. Complete fossil specimen MHNL.20015625 in dorso-ventral compression, under UV light. B–C. Close-up of pleonal somites 4 and 5 with strings of muscle fibers. D–E. Close-up of tail fan with strings of muscle fibers in both the basipodite and the proximal region of telson. Abbreviations: bap = basipodite, di = diaeresis, en = uropodal endopod, ex = uropodal exopod, mf = muscle fibers (in blue), pl = pleopods, s4–s6 = pleonal somites 4 to 6, t = telson. F. For comparison, elongated strings ofmusclefibers from pleonal somite s6 from the extant glypheid lobster MNHN-IU-2008-14,762 collected by Mission MUSORSTOM 1 (Philippines, N. Lubang Islands). Photographs: C. Lemzaouda, line drawings: S. Charbonnier. (For interpretation of the references to color in this figure legend, the reader is referred to the web version ofthis article.)

(Carpentier et al., 2006). Jurassic (Callovian) of the La Voulte-sur-Rhône Lagerstätte in France Other new examples include the dendrobranchiate shrimp (Fig. 10); the glypheid lobster Gigacerina saemanni (Oppel, 1861) from Archeosolenocera straeleni Carriol and Riou, 1991, from the Middle the Late Jurassic (Kimmeridgian/Tithonian) lithographic limestones of

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Fig. 12. Pleon of an undetermined fossil stomatopod (MGUH 32025) from the Eocene (Ypresian-Lutetian) Lillebælt Clay Formation of Trelde Næs near Fredericia, Denmark, with muscles preserved. A. Entire specimen. B–C. Closeups of one of the pleonal segments with muscle fibers. the Cerin Lagerstätte in France (Fig. 11); and an undetermined fossil a species, which may be primarily caused by variable preservation (e.g., stomatopod from the Eocene (Ypresian-Lutetian) Lillebælt Clay For- Fraaije, 2014: Figs. 7, 11). Muscle scars appear more difficult to spot on mation of Trelde Næs near Fredericia in Denmark (Fig. 12). In all cases, smaller specimens. muscles were found in the pleon, but also in the telson and adjacent Scars have been used for the taxonomic placement of species. For basipodite for the glypheid. The longitudinal orientation of the muscles example, for the Jurassic crab Tanidromites Schweitzer and Feldmann, in the glypheid suggest that they represent deep extensor muscles or 2008, distinguishing between species is done partly based on muscle dorsal pleonal muscles, which may apply too for the dendrobranchiate scars (Starzyk, 2013, 2015a, 2016); for the Jurassic–Cretaceous crab shrimp, although the precise position of the muscles is not known (see Eodromites Patrulius, 1959, hepatic pits (=scars of antennar extensor Hertzler and Freas, 2009). The similarly oriented muscles in the sto- muscle mostly likely, see Glaessner, 1969) were mentioned in the di- matopod represent dorsoventral trunk muscles (Fig. 3). The long- agnosis of genus (Starzyk, 2015b); and muscle scars in the mesogastric itudinally oriented muscles in the telson and basipodite of the glypheid region are part of diagnosis of the Middle Jurassic crab Homolus Eudes- may represent the telson flexor or extensor and the uropod rotator or Deslongchamps, 1835 (Schweitzer and Feldmann, 2010). remotor (sensu Vogt, 2002, for a crayfish). Despite the presence of muscle scars in many malacostracan taxa, they have not been studied systematically. Glaessner (1969: R407–R409) in the Treatise of Invertebrate Paleontology on Decapoda 4. Muscle scars indicated that muscle scars deserve more detailed comparative study, but no such studies have been performed thus far. In addition to muscles, muscle scars can give additional information To explore first-order patterns of muscle scars in fossil malacos- about the presence, shape, and size of the muscles. Muscle scars in the tracans, we surveyed the published literature from 2004 to 2006 and malacostracan fossil record are particularly visible on the inner side of 2014 to 2016 (357 articles in total) to obtain a reasonable sample size the cuticle where the muscles are attached in a reticulate pattern for this century. We searched manuscript texts containing fossil spe- (Figs. 13–14). However, recurring discoloration of the cuticle in a re- cimen images for referrals to muscle scars and we checked for scars in ticulate pattern may also signify muscle attachment spots on the inside the provided figures. Posterior gastric pits as an attachment forthe of the cuticle (e.g., Carpentier et al., 2006: Fig. 5C; Hyžný and stomach muscle (Glaessner, 1969; McLaughlin, 1980) and other pits as Klompmaker, 2015: Fig. 8; Rumsey et al., 2016: Fig. 3). Muscle scars possible attachment places of muscles are not included in this search have previously been noted primarily from internal molds lacking the because they cannot be observed consistently in the provided figures. cuticle (e.g., Berglund and Feldmann, 1989; Bishop, 1978; Crônier and We found that muscle scars are present from the Devonian onward and Boursicot, 2009; Glaessner, 1933, 1960; Schweigert and Koppka, 2011). become more common starting in the Jurassic (Fig. 15; Appendix 2), We noted that muscle scars have occasionally been incorrectly referred coinciding with the first major species-level diversification of decapods to as ornamentation because true ornamentation is found only on the (Klompmaker et al., 2013). The 162 occurrences of malacostracans cuticle surface, which is usually not the case for muscle scars. These showing muscle scars is a minimum estimate because we did not scars are not always present or equally obvious on each internal mold of

318 A.A. Klompmaker, et al. Earth-Science Reviews 194 (2019) 306–326 include ambiguous muscle scars and not all images were of adequate region of paguroid anomurans, and lateral to the cardiac region for quality to check for muscle scars. There is no preferred rock type in Brachyura (Appendix 2). These scars represent attachments for the which muscle scars are found; when malacostracans are preserved as closer or opener muscle, the posterior gastric muscle, the anterior internal molds, muscle scars can be found. gastric muscle, and the attractor epimeralis muscle, respectively Muscle scars are predominantly found on the inner and outer sides (Figs. 1–4). of primarily cheliped (first pereopod) propodi of Axiidea, at the baseof Brachyura, Axiidea, and Anomura comprise most of the muscle scar the mesogastric region of primarily Brachyura, on the anterior gastric occurrences (Fig. 16A; Appendix 2). Except for one phyllocarid

(caption on next page)

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Fig. 13. Muscle scars in fossil decapod crustaceans. A, B. Outer and inner sides of a partial major cheliped propodus of Callianassidae from the late Pleistocene–Holocene of Falmouth 01, Trelawny Parish, Jamaica (UF 239327). C. Inner side of the cuticle of a carapace of the crab Mithrax cf. M. pleuracanthus Stimpson, 1871, from the late Pliocene–early Pleistocene of the MacAsphalt Shell Pit, Sarasota County, Florida, USA (UF 29057; see Klompmaker et al., 2015, for other side; Windsor and Felder, 2017, for genus name). D. Internal mold of the major cheliped propodus of the ghost shrimp Calliax michelottii (A. Milne Edwards, 1860) from the early Miocene of Slovenia (UMJGPA 77874) (from Hyžný and Klompmaker, 2015: Fig. 8C). E. Internal mold of a carapace of the crab Dromiopsis elegans Reuss, 1858, from the Danian of Faxe in Denmark (MGUH 24379). F. Internal mold of the anterior part of the carapace of the hermit crab Pilgrimcheles vonmeyeri Fraaije, 2014, from the Kimmeridgian of Westerberg, Germany (MAB k.3335) (from Fraaije, 2014: Fig. 7A). G. Internal mold (and some of the cuticle) of a carapace of the crab Harpactocarcinus punctulatus (Desmarest, 1817), from the Eocene of Laverda at Salcedo, Vicenza, Italy (GBA 2010/151/0058/01). H. Internal mold of a carapace of the squat lobster Hispanigalathea tithonia Robins et al., 2016, from the Tithonian of Ernstbrunn, Austria (NHMW 2007z0149/0438) (from Robins et al., 2016: Fig. 13). I. Internal mold of a carapace of the lectotype of the crab Goniodromites complanatus Reuss, 1858, from the Kimmeridgian–lower Berriasian of Ignaziberg, Czech Republic (GBA2007/096/0008). J. A carapace of the crab Lophoranina bittneri (Lőrenthey, 1902), from the Eocene of Fumane di Polesella at Verona, Italy (GBA 2010/243/0004), with erosion of the cuticle exposing the muscle scars in the mid-part of the carapace. K. Internal mold of a carapace of the squat lobster Vasconilia ruizi (Van Straelen, 1940), from the late Albian of Koskobilo, Spain (MAB k.2598) (from Klompmaker et al., 2012: Fig. 7A). L. Internal mold of a carapace of the crab Navarradromites pedroartali Klompmaker et al., 2012, from the late Albian of Koskobilo, Spain (MAB k.3182) (from Klompmaker et al., 2012: Fig. 8B). occurrence, muscle scars were absent in non-decapod malacostracans figured herein, many of the muscles resemble those found inextant such as Isopoda and Stomatopoda, but the number of published articles representatives. Of the two best preserved individuals, the position and on fossil representatives of clades was also limited. Whether muscle the pattern of cylindrical bundles of the closer muscle in the mid- scars are less common in these clades requires more research, but Holocene Sergio sp. (Fig. 8) is very reminiscent of those of modern muscle scars have been mentioned from a Miocene stomatopod (Hof callianassid ghost shrimp (Fig. 4C). Much more remarkable is the strong and Schram, 1998: p. 328). Muscle attachments can be seen through the similarity of the pleonal muscles of the Late Devonian penaeoid shrimp cuticle in some recent isopods (Bruce and Bussarawit, 2004: Fig. 5; to a modern shrimp from the same superfamily (Feldmann and Poore and Storey, 2009: Fig. 1A). Therefore, fossil isopods with muscle Schweitzer, 2010: Fig. 4); many muscle groups in the modern specimen scars may be expected with targeted research, but we have not been could be identified on the fossil as well. Similarly, pleonal muscles ina able to find such instances thus far. Late Jurassic glypheid lobster and an Eocene stomatopod do not seem Although muscle scars in fossil malacostracans have been known for to differ much from modern representatives. Thus, morphologically a long time (e.g., Glaessner, 1933), they are not often recognized to the similar specimens contain comparable muscles, which we refer to as present day. Of the 162 malacostracan occurrences with muscle scars, “muscle conservatism”. Muscles in another marine arthropod, a only 30 (19%) were specifically mentioned in the text (Fig. 16B; Ap- horseshoe crab from the Late Jurassic of Germany, are also remarkably pendix 2). Most of such occurrences are found in systematic pa- comparable to those in modern horseshoe crabs (Briggs et al., 2005). leontology articles, for which the description and interpretation of all Muscle attachment scars provide additional evidence for the pre- morphological features including muscle scars is one of the main goals. sence, size, shape, and position of muscles. For ghost shrimps, the po- However, muscle scars were not mentioned for 132 of the occurrences sition and approximate size of closer muscle scars does not appear to (81%), including 67 occurrences (41%) for which the respective taxa have changed much through time (Hyžný and Klompmaker, 2015: were described, which implies that muscle scars were not recognized. Fig. 8). Unlike penaeoid and ghost shrimps, brachyurans have under- We strongly recommend including muscle scars in taxon descriptions. gone a remarkable change in shape from longer than wide in the Jur- In contrast, 65 muscle scar occurrences (40%) were part of papers that assic, as in a lobster ancestor, to primarily wider than long in the did not describe the respective taxa. It is no surprise that the muscle Cenozoic combined with tucking of the pleon under the carapace (e.g., scars were not mentioned in these cases. Glaessner, 1960; Guinot et al., 2013). This trend has led to a large re- To evaluate the commonness of muscle scars in the fossil malacos- duction of muscles in the pleon and some reorganization of muscles in tracan literature, we surveyed the same 357 articles for the presence of the carapace (e.g., Glaessner, 1960; Guinot et al., 2013). However, little muscle scars. Muscle scars were present in 24% (87) of the articles is known about the evolution of muscle scars in the carapace of Bra- (Fig. 16C; Appendix 3), indicating that muscle scars are common. This chyura throughout their evolutionary history. The same applies to percentage is a minimum estimate because not all images were of suf- several clades of Anomura that have evolved a crab-like morphology ficient quality to determine whether muscle scars were present. The (Keiler et al., 2015, 2017; Scholtz, 2014). number of muscle scar occurrences and the percentage of papers with muscle scars jointly suggest that evidence for muscle scars is ubiquitous 6. Conclusions and future directions and provides the opportunity to study large-scale patterns of muscle evolution and attachment. Our search into the literature has yielded 29 occurrences of muscles The patterns of skeletal calcification affecting the likelihood of preserved in fossil malacostracans to which we add 18 new occurrences muscle scar preservation are unexplored. It may be possible that very in the first overview of muscles in the malacostracan fossil record. Most prominent muscle scars on fossil cheliped propodi of ghost shrimps are of these occurrences are found in known Konservat-Lagerstätten. As our associated with a rather thin cuticle, whereas the absence of muscle survey of malacostracan taxa with muscles preserved in Konservat- scars in the fossil lobster chelae may be related to their strong calcifi- Lagerstätten has not been exhaustive, additional study of museum cation and ornamentation. Alternatively, the depth of the muscle scars collections and new collecting efforts are expected to result in addi- on the inside of the cuticlar layers may play a role in the preservation of tional species with muscles preserved. More late Cenozoic species muscle scars. In durophagous brachyuran crabs, the muscle scars are should be checked for the presence of muscles. It is possible that some very prominent also in strongly calcified chelae, which may be related specimens were overlooked because of a bias in research towards to their extreme claw strength (Kosloski and Allmon, 2015; Schenk and Mesozoic Konservat-Lagerstätten. Lithification and compaction should Wainwright, 2001; Taylor, 2000). not have impacted late Cenozoic taxa as much as older specimens. The muscles in the major cheliped propodi of a mid-Holocene ghost 5. Brief comparison of fossil and modern muscles and muscle shrimp and the muscles in the pleon of a Late Devonian penaeoid scars shrimp are comparable to those of related modern specimens. Thus, for similarly-shaped body plans, we hypothesize conservatism in the evo- Although not all muscle groups are preserved in the specimens lution of musculature. Most changes in muscle structure may be

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Fig. 14. Muscle scars in fossil decapod crustaceans as in Fig. 13, but with muscles marked. Inferred muscles: red = closer (adductor) muscles; light green = posterior gastric muscles; light blue = anterior gastric muscles; blue = attractor epimeralis muscles; yellow = antennar extensor muscles; pink = posterior gastric pits for stomach muscle; purple = internal mandible adductor muscles; orange = branchial muscles. (For interpretation of the references to color in this figure legend, the reader is referred to the web version of this article.)

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50 Malacostracan clade

Anomura 40 Astacidea Axiidea Brachyura 30 Gebiidea Glypheidea Phyllocarida 20

Occurrences with muscle scars muscle Occurrences with 10

0

Silurian Permian Triassic Jurassic Cambrian Devonian Neogene Ordovician Cretaceous Paleogene Quaternary Carboniferous

Fig. 15. Raw taxon occurrences with muscle scars per Meso- and Cenozoic period by malacostracan crustacean infraorder. Results are based on a survey from the literature published from 2004 to 2006 and 2014 to 2016 (see Appendix 2). Data are not normalized for the unequal duration of periods. n = 162.

Fig. 16. Muscle scar occurrences by clade (A), the recognition of these muscle scar occurrences (B), and the proportion of papers containing evidence of muscle scars (C). N(D): muscle scars were not mentioned, even though the taxa were described; N(N): muscle scars were not mentioned, but the taxa were not described. The results are based on a survey from the malacostracan literature published from 2004 to 2006 and 2014 to 2016 (see Appendices 2–3). expected for Brachyura, having undergone dramatic changes in car- can complement studies on muscle evolution through time, and may be apace and pleonal shape since the Jurassic. Additional collecting of particularly revealing for the shape changes associated with Brachyura. Brachyura from Konservat-Lagerstätten or re-examination of existing collections may yield specimens that can give clues about muscle de- Supplementary data to this articles (Appendices 1 to 3) can be found velopment in true crabs. Future research could also focus on char- online at https://doi.org/10.1016/j.earscirev.2019.04.012. acterizing the preservation of muscles within and across species using many individuals to more quantitatively assess the relative preservation Acknowledgments potential of muscles across the malacostracan body. In the first overview of muscle scars in fossil malacostracans, our We thank Darrell Kaufman (Northern Arizona University) for C-14 survey of 357 articles published from 2004 to 2006 and 2014 to 2016 dating of a bivalve associated with a ghost shrimp, Ann Heatherington yielded 162 occurrences with muscle scars on primarily internal molds. and Michael Kesler (both University of Florida) for help with SEM They are found most often on brachyuran carapaces and axiidean photography and EDS analyses, and Sylvain Pont (IMPMC, MNHN) for cheliped propodi. Nearly all occurrences are found from the Jurassic help with XRD analyses. Rodney Feldmann (Kent State University) onwards in a variety of sedimentary settings and typically not in provided high-resolution images of the Devonian shrimp, Carolin Haug Lagerstätten. These scars are recognized by authors in only 19% (30) of (LMU Munich) confirmed the presence of muscles in a Miocene sto- the occurrences. Muscle scars are found in 24% (87) of the articles, matopod, and Nathan Jud (William Jewell College) and PCP PIRE in- suggesting that muscle scars are very common. This rich source of data terns helped with making the microCT movie. We thank publishers and can be used for many more studies on muscle scar presence, muscle editors for permissions to reuse previously published images. Useful identification, preservation, and evolution through time. Muscle scars comments by two anonymous reviewers helped to improve this

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