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A new record and cuticular structure of Meyeria magna (Decapoda, Mecochiridae) from the lower Albian (Lower Cretaceous) of Colombia
Cretaceous Research 57 (2016) 342e349
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Cretaceous Research journal homepage: www.elsevier.com/locate/CretRes
A new record and cuticular structure of Meyeria magna (Decapoda, Mecochiridae) from the lower Albian (Lower Cretaceous) of Colombia lez-Leo n a, b, *, Pedro Patarroyo c, Josep Anton Moreno-Bedmar d, Oscar Gonza Torrey Nyborg e, Francisco J. Vega d noma de M Posgrado en Ciencias de la Tierra, Universidad Nacional Auto exico, M exico D.F., C.P. 04510, Mexico Facultad de Estudios Superiores Iztacala, Universidad Nacional Autonoma de M exico, Estado de M exico, 54070, Mexico c , Colombia Departamento de Geociencias, Universidad Nacional de Colombia, Cr.30 N. 45e03, Bogota d noma de M Instituto de Geología, Universidad Nacional Auto exico, M exico, D.F., C.P. 04510, Mexico e Loma Linda University, Department of Earth and Biological Sciences, Loma Linda, CA 92350, USA a
b
a r t i c l e i n f o
a b s t r a c t
Article history: Received 9 March 2015 Received in revised form 13 August 2015 Accepted in revised form 14 August 2015 Available online 30 October 2015
Eleven specimens of the lobster Meyeria magna from the Simití Formation (Santander Province, Colombia) represent a new occurrence for this species in South America. Ammonites collected both below and above the bed that yielded M. magna allow dating of these specimens as early Albian. Morphological characteristics observed in these Colombian specimens were compared with those of other specimens from Mexico, the United Kingdom and Spain. Our interpretation of the cuticular structure in thin section does not unambiguously allow interpretation of the specimens studied to corpses or molts. © 2015 Elsevier Ltd. All rights reserved.
Keywords: Lobsters South America Albian Ammonites Comparisons
1. Introduction Apart from a doubtful occurrence in Asia (Wang, 1981), the widely distributed Early Cretaceous lobster Meyeria magna M'Coy, 1849 is well known in North America and Europe (e.g. Vía, 1951; Vía Boada, 1975; Simpson & Middleton, 1985; Feldmann, Vega, pez, 1995, 2007; CalGarcía-Barrera, Rico-Montiel, & Martínez-Lo zada & Urquiola, 1999; Vega, Feldmann, Etayo-Serna, Bermúdez mez, 2008; Lo pez-Horgue, 2009; Bover-Arnal et al., Aguirre, & Go n, Moreno-Bedmar, & Vega, 2010; Najarro et al., 2011; Gonz alez-Leo 2014). In the present study, a new occurrence of M. magna is noted and represents the second record of this species from Colombia and South America. The taxonomy of Meyeria magna is well established on the basis of specimens from the Isle of Wight, United Kingdom, from where it was originally described by M'Coy (1849), along with several other specimens collected from Spain, Colombia and
Mexico (Vía Boada, 1975; Simpson & Middleton, 1985; Vega et al., lez-Leo n et al., 2014). The diagnostic features of 2008; Gonza eleven Colombian specimens are compared to those of other ocurrences of M. magna. Since cuticular structure in decapods is vilavery distinctive in transverse section (Vega, Feldmann, & Da Alcocer, 1994, 2005), thin sections of several specimens of M. magna from the above-mentioned countries were made and studied in order to characterize this feature. Cuticular structure of other fossil decapod species has been previously reported (e.g. Feldmann & Tshudy, 1987; Vega et al., 1994; Feldmann & Ga zdzicki, 1998; Haj & Feldmann, 2002; Waugh & Feldmann, 2003; Waugh, Feldmann, Schroeder, & Mutel, 2006). The current work constitutes a first step toward characterizing the cuticular structure of M. magna. In addition, it contributes to the chronostratigraphic range of this widely distributed species.
2. Geological setting * Corresponding author. lez-Leo n), E-mail addresses: [email protected] (O. Gonza [email protected] (P. Patarroyo), [email protected] (J.A. Moreno-Bedmar), [email protected] (T. Nyborg), [email protected] (F.J. Vega). http://dx.doi.org/10.1016/j.cretres.2015.08.006 0195-6671/© 2015 Elsevier Ltd. All rights reserved.
The eleven specimens of Meyeria magna analyzed in the present paper were collected from the Simití Formation at co-ordinates lez area during fieldwork in the 6 030 2500 N/73 4301000 W in the Ve
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lez area is located in Eastern Cordillera of central Colombia. The Ve (Fig. 1). The area the province of Santander to the north of Bogota has been renowned for its fossil record ever since the Humboldt expeditions, and, as such, a vast literature on the region has been published, in some cases including fossil illustrations such as Karsten (1856, 1886), Gerhardt (1897), Collet (1924), Beurlen (1938), Riedel (1938), Basse (1949), Bürgl (1956), Kakabadze and Thieuloy (1991), Bogdanova and Hoedemaeker (2004), Kakabadze and Hoedemaeker (1997, 2004) and Sharikadze, Kakabadze, and Hoedemaeker (2004). The Simití Formation is a lithostratigraphic unit within the Middle Magdalena Valley, Colombia; it overlies the Tablazo Formation in the Cundinamarca Basin (TablazoeMagdalena subbasin). lez is the upper San Its equivalent in the Villa de Leyva area near Ve Gil Formation, which overlies the lower San Gil Formation. In the present study, the Tablazo and Simití formations are preferred over the lower and upper San Gil formations because this nomenclature lez area by Ulloa and was adopted for the geological map of the Ve Rodríguez (1978, 1984) and Mendoza-Parada, Moreno-Murillo, and Rodríguez-Orjuela (2009). The Simití Formation, with a thickness of 410 m, is composed mainly of black shales with intercalated biosparites and biomicrites. Some beds show evidence of transport, which led to fossil fragmentation, while other beds contain foraminifera and flattened ammonites, bivalves, gastropods and decapod crustaceans. Ammonites of the genera Carloscaceresiceras Etayo-Serna (1979), Douvilleiceras de Grossouvre (1894) and Glottoceras Hyatt (1875), were collected both below and above specimens of M. magna, and this association indicates an early Albian date. Meyeria magna has been previously recorded from Colombia (Vega et al., 2008), in association with early Aptian cephalopod genera such as Heminautilus Spath (1927), Kutatisites Kakabadze , (1970) and Cheloniceras Hyatt (1903), from southwest of Bogota in the Upper Magdalena Valley subbasin. A depositional environment associated with initial offshore deposits following the progressive marine advance to the south of the basin is suggested. 3. Ammonite biostratigraphy In the section studied, ammonites are quite rare; two specimens
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were recovered, while a third one was left at the outcrop. The specimen of Carloscacereciseras cf. caceresi Etayo-Serna (1979) illustrated here (Fig. 2A) is an internal mold of a volumetric individual with wide sinusoid constrictions on the flank that resemble the holotype illustrated by Etayo-Serna (1979, pl. 10, Fig. 1). However, poor preservation does not allow definite identification to species level. According to Wright (1996), the genus Carloscacereciseras is a junior synonym of Anadesmoceras Casey, 1954, but we follow Bogdanova & Hoedemaeker (2004) in considering Carloscacereciseras to be a valid genus, the age range of which is early Albian (Bogdanova & Hoedemaeker, 2004). The second ammonite specimen represents a crushed shell of Douvilleiceras sp. (Fig. 2B). Because of poor preservation a specific identification is impossible. The lower Albian is usually divided into two biozones, the younger of which is the globally recorded Douvilleiceras mammillatum Zone. The genus Douvilleiceras is restricted to this zone (e.g. Casey, 1961; Reboulet et al., 2014). The equivalent in Colombia is the Douvilleiceras solitaeeNeodeshayesites columbianus Assemblage Zone (Etayo-Serna, 1979). The third ammonite, Glottoceras cf. raimondii n, 1908) (Fig. 2C), is a slightly crushed specimen with pyritic (Lisso replacement. This shell shows clear lower, mid and ventrolateral tubercles on primary ribs and intercalated ventrolateral tubercles in the ventral region are also observed. The low primary ribs start in the umbilical seam, the ventral area being flat to slightly concave. The ribs of this specimen are not as well developed as in the n (1908, p. 4, Fig. 1a, as Knemiceras raimondii); illustration by Lisso however, tuberculation is the same. Today this species is assigned to genus Glottoceras (e.g. Robert & Bulot, 2004; Bujtor, 2010), its range corresponding approximately to the middle and upper part of the Douvilleiceras mammillatum Zone. Specimens of Meyeria magna used in the present study were collected above the specimen of Carloscacereciseras cf. caceresi and below Douvilleiceras sp. and Glottoceras cf. raimondii (Fig. 2); therefore, an early Albian date is firmly established. 4. Systematic paleontology Order Decapoda Latreille, 1802 Suborder Pleocyemata Burkenroad, 1963 Infraorder Glypheidea von Zittel, 1885
Fig. 1. Locality map showing localities from which specimens of Meyeria magna were collected and stratigraphic column of the Lower Cretaceous (Albian) Simití Formation.
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Fig. 2. A. Lateral view of a specimen of Carloscacereciseras cf. caceresi, embedded in rock; B. Lateral view of Douvilleiceras sp. (UN-DG-AM-241); C. Lateral view of Glottoceras cf. raimondii (UN-DG-AM-240). Scale bar equals 10 mm.
Superfamily Glypheoidea von Zittel, 1885 Family Mecochiridae Van Straelen, 1925 Genus: Meyeria M'Coy, 1849 Type species. Astacus ornatus Phillips, 1829, by original designation. Included species. Meyeria ornata (Phillips, 1829); Meyeria magna M'Coy, 1849; Meyeria harveyi (Woodward, 1900); Meyeria rapax (Harbort, 1905); Meyeria schwarzi (Kitchin, 1908); Meyeria bolivari (Van Straelen, 1927); Meyeria gracilis (Glaessner, 1932); Meyeria mexicana (Rathbun, 1935); Meyeria houdardi (Van Straelen, 1936) and Meyeria crofti (Ball, 1960). Meyeria magna M'Coy, 1849 Figs. 3e4 1849 Meyeria magna M'Coy, p. 334, Fig. 4. 1863 Meyeria vectensis Bell, p. 33, pl. 10, Figs. 1e5. 1863 Oncopareia granulosa Vilanova, p.98, pl. 3, Fig. 2. 1881 Meyeria pearcei Spence-Bate in Lee, p. 87, pl. 204, Fig. 14. 1927 Meyeria bolivari Van Straelen, p. 80, pl. 1, Figs. 1e2. 1951 Hoploparia granulosa Vía, p. 154, Fig. 10. 1975 Meyeria magna M'Coy, Vía Boada, p. 33, Figs. 1.1e1.9, 2.1e2.6. 1985 Meyeria magna M'Coy, Simpson and Middleton, pp. 204e211, Figs. 1A, 2AeM, 6E, H, 7H, 8AeB, 9AeL. 1995 Meyeria pueblaensis Feldmann, Vega, García-Barrera, Rico pez, p. 404, Figs. 2.1e2.4. Montiel and Martínez-Lo 2008 Meyeria magna M'Coy, Vega et al., p. 5, Figs. 5.1e5.7, 6.1e6.7, 7.1, 7.6. pez-Horgue, p. 27, Fig. 2AeJ. 2009 Meyeria magna M'Coy, Lo 2011 Meyeria magna M'Coy, Astrop, 2011, p. 116, Fig. 1B. 2013 Meyeria magna M'Coy, Klompmaker, appendices AeB. lez-Leo n et al., pp. 10e14, 2014 Meyeria magna M'Coy, Gonza Figs. 10AeQ, 11AeM, 12AeJ.
5. Material and methods 5.1. Description of specimens 5.1.1. Carapace Laterally compressed and subcylindrical; cephalic region with three longitudinal spiny carinae; orbital, gastro-orbital and antennal carina parallel; short distance between orbital and gastroorbital carinae, antennal carina separated twice the distance from the others; cephalic carinae raised, antennal carina more raised than others; cervical groove deep, two-thirds of height of carapace, inclined 53 toward lower anterion margin, cervical groove
ventrally connected to antennal groove; branchiocardiac groove shallow, inclined 15 from upper part of posterior margin to midheight of carapace; post-cervical groove shallow and parallel to branchiocardiac groove; hepatic groove slightly deep, convex ventrally at intersection with antennal groove; shallow and undeveloped inferior groove, connected to hepatic groove; cuticle on anterior cardiac region with a few small tubercles; cuticle on branchial region covered by small tubercles of uniform size, becoming more numerous toward ventral margin and margin hepatic lobe. 5.1.2. Pleon Pleonal somites with three segments of longitudinal rows of tubercles, anterior and lower margins rounded, posterior margin straight; second to fifth pleonal somites of similar shape and size; sixth pleural segment triangular, strong ridge at contact with tergum; endopodite and exopodite of telson triangular, both with median keel, diaeresis present. Appendages not preserved, only fragments of first pereiopod and coxae of pereiopods 1e5. 5.2. Discussion Specimens are well preserved and only slightly crushed. Morphological features observed include: cephalic carinae (cc ¼ oc orbital carina þ gc gastro-orbital carina þ antennal carina) (Fig. 3A); three thoracic ridges in the branchial region (Fig. 3 A1, A2 and B); cervical, (e1e), hepatic (b1), branchiocardiac (a) and postcervical (c) grooves (Fig. 3 A1, A2, B, E1 and E2); ornamentation of granules on carapace can be observed (Fig. 3 A2, specimens D and E1), as well as ornamentation of the pleon (Fig. 3 C, F1 and F2). In Fig. 3D the diaeresis in the exopodite of the telson can be seen. Such characteristics have been recognized previously allowing the unequivocal assignment of the Colombian material to Meyeria magna. Fig. 4 illustrates carapace features observed in the Colombian material. 5.3. Material Eleven specimens of Meyeria magna (UN-DG-CR-020eUN-DGlez area, CR-030) from the lower Albian (Simití Formation) in the Ve n province of Santander, Colombia are housed in the Coleccio gica del Departamento de Geociencias, Universidad Paleontolo Nacional de Colombia.
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Fig. 3. AeF. Meyeria magna from the lower Albian of Colombia. Carapace UN-DG-CR-020, in left lateral (A1) and right lateral views (A2); B. Partial carapace and pleon, UN-DG-CR023, in right lateral view; C. UN-DG-CR-021 showing five pleonal somites and a partial telson, left side; D. Carapace fragment, UN-DG-CR-025, showing pleon and telson; E1. UN-DGCR-022, showing ornamentation of carapace, left lateral view; E2. Same specimen, right lateral view; F1. Detail of UN-DG-CR-030, pleonal somites, right lateral view; F2. Same specimen, showing details of pleon, left lateral view. Scale bar equals 10 mm.
5.4. Cuticular structure The material studied represents 12 thin sections obtained from five specimens of Meyeria magna, one from Colombia, two from Mexico, one from United Kingdom and one from Spain. The thin sections are housed in the Museo María del Carmen Perrilliat M., n Nacional de Paleontología, Instituto de Geología, Coleccio
noma de Me xico, Me xico D.F., Mexico Universidad Nacional Auto with the numbers IGM 4685 to IGM 4696. 5.4.1. Colombia (V elez) Four thin sections (numbered IGM 4685, IGM 4686, IGM 4687 and IGM4688) obtained from two median sections of the carapace. The age of this material is early Albian. 5.4.2. Mexico (Puebla) Four thin sections (numbered IGM 4689, IGM 4690, IGM 4691 and IGM 4692) obtained from the median part of the sectioned carapace and pereiopods. The age of this material is late Barremianeearly Aptian. 5.4.3. Spain (Cuchía) Two thin sections (numbered IGM 4693 and IGM 4694) obtained from the branchial region of the carapace. The age of this material also is early Aptian.
Fig. 4. Meyeria magna, UN-DG-CR-020, cephalothorax. Morphological characteristics: e1e ¼ cervical groove, c ¼ postcervical groove, a ¼ branchiocardiac groove, b1 ¼ hepatic groove, cephalic carinae (oc ¼ orbital carina, gc ¼ gastro-orbital carina, ac ¼ antennal carina) and tr ¼ thoracic ridges. Scale bar equals 10 mm.
5.4.4. United Kingdom (Isle of Wight) Two thin sections (numbered IGM 4695 and IGM 4696) obtained from the branchial region of the carapace. The age of this material is early Aptian.
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6. Cuticular structure in decapod crustaceans The order Decapoda includes organisms commonly called shrimps, lobsters and crabs. Decapod crustaceans rank amongst the most common animals inhabiting a number of different environments, both at present times (e.g. Abele, 1974) and in the past (e.g. Klompmaker et al., 2013, Schweitzer, Feldmann, & Kowalewski, 2013; Schweitzer & Feldmann, 2014). They are commonly found in the fossil record despite their low preservation potential when compared to other marine invertebrates (Kidwell & Flessa, 1995). For many fossil decapod crustacean taxa, morphological features such as the shape and contour of the carapace, the number and placement of spines, nodes and grooves and the orbital design, are the basis for classification (Haj & Feldmann, 2002). Decapod cuticle has a very distinctive structure when observed in cross section. Although most references on petrography and sedimentology illustrate or describe how to recognize some arthropod bioclasts such as those of trilobites and ostracods (Adams, Mackenzie, & Guilford, 1984; Flügel, 2004), there is no such description for decapods, in spite of the fact that decapod cuticle is of potentially common occurrence in Mesozoic and vila-Alcocer, & Filkorn, 2005). The Cenozoic shelf deposits (Vega, Da cuticle of extant decapod crustaceans is commonly composed of four discrete layers, namely epicuticle, exocuticle, endocuticle and a membranous layer, which are all well documented in the literature (e.g., Neville & Berg, 1971; Taylor, 1973; Dalingwater, 1977; Vega et al., 1994, 1998; Feldmann & Ga zdzicki, 1998; Haj & Feldmann, 2002; Vega et al., 2005; Amato, Waugh, Feldmann, & Schweitzer, 2008). The outermost layer of the cuticle is the epicuticle, commonly a mineralized bilaminar layer. The exocuticle is composed of chitinprotein fibers stacked in layers of continuously changing orientation (Green & Neff, 1972; Haj & Feldmann, 2002). The endocuticle is the thicker and most heavily calcified layer (Roer & Dillaman, 1984). In some lobsters, such as nephropids, the outer part of the endocuticle consists of broad lamellae while the innermost portion is thinly laminated (Feldmann & Tshudy, 1987). The innermost layer is the membranous layer, an uncalcified layer (Roer & Dillaman, 1984; Haj & Feldmann, 2002), making the chances of preservation in the fossil record low. Characterization of the cuticular structure has been used as evidence to suggest the presence of both molts and corpses in the fossil record of decapod crustaceans. Feldmann and Tshudy (1987) analyzed 58 specimens of Hoploparia stokesi (Weller, 1903) and concluded that the presence of horizontal lamellae of the endocuticle could be used to differentiate between corpses and molts. Vega et al. (1994) recognized the endocuticular, exocuticular lamellae, probably tegumental ducts, cell imprints in the epicuticular layer and the membranous layer in many samples of Costacopluma mexicana Vega & Perrilliat, 1989, suggesting the presence of corpses in the record of this species. For this reason, the study of cuticular structure in these organisms can be a potential tool in preliminary identification of major decapod groups and taphonomic interpretations (Feldmann & Tshudy, 1987; Vega et al., 1994; Klompmaker, Hy zný, & Jakobsen 2015). In the present study, we analyze and describe the cuticular structure of Meyeria magna employing thin sections of five specimens from different countries. 6.1. Analysis and discussion Analysis of cuticular structure in M. magna thin sections did not fully allow assignment of these specimens to either corpses or molts. In some specimens three discrete layers (epicuticle, exocuticle and endocuticle) could be distinguished whereas in others only a
single layer was clearly observed, the epicuticle. These features have previously been recognized (Feldmann & Tshudy, 1987; Vega et al., 1994). In none of the studied thin sections could the microstructure (horizontal lamellae, pore canals, tegumental ducts and balls) be observed. These features may have either been altered during the diagenetic process or were never present in the first place; a process difficult to determine. Examination of the Colombian thin sections shows three discrete layers of cuticle. The epicuticle is approximately 13 mm thick (Fig. 5aeb); the exocuticle is 23 mm thick and shows no evidence of lamellar structure (Fig. 5a). In the thickest layer (57 mm), the endocuticle was partially replaced by sparite crystals during diagenesis. Below the endocuticle it is possible to discern two other undifferentiated layers (Fig 5a). Since the membranous layer is not ordinarily preserved in the fossil record, we infer that these layers may reflect the diagenetic process. Unfortunately, insufficient preservation of the microstructure of the different layers does not permit us determine whether the Colombian material represent corpses or molts. The Colombian specimens of Meyeria magna were preserved disarticulated, suggesting the specimens were molts; however this is not necessarily true. There are many destructive processes that can lead to fragmentation of corpses, such as physical disturbance, transport and destruction in the sedimentary environment, predation and scavenging, bacterial degradation of soft tissues and exoskeleton, as well as chemical breakdown of such remains (Plotnick, 1986). Examination of Mexican specimens has allowed characterization of the same layers as those seen in the Colombian material (Fig. 5cee). We observed a bilaminar epicuticle (Fig. 5c), 10 mm thick, as the first layer; the second layer was identified as the exocuticle, 26 mm thick, and the third layer was the endocuticle, measuring 140 mm (Fig. 5dee). The characteristic lamellae and pore canals are absent in the Mexican specimens. As in the Colombian material, we also noted two undifferentiated layers below the base of the endocuticule that could be interpreted as the membranous layer; however, this hypothesis cannot be confirmed. The boundaries between exocuticle and endocuticle were only inferred (Fig. 5dee). Despite the presence of discrete layers in the cuticle of the Mexican specimens, we cannot document this record as corpses because the microstructure (horizontal lamellae) in the endocuticle and exocuticle is not evident. It is possible that these features were lost during diagenesis, but this is difficult to determine. When epicuticle, exocuticle and endocuticle layers are present, cuticle structure does suggest a specimen represents a corpse, as well as, when there is only one layer present, cuticular structure suggests a specimen represents a molt. This criterion can be applied in thin sections, when it is seen that the diagenetic process altered the cuticle microstructure. Based upon these observations, specimens from the United Kingdom and Spain could be interpreted as molts. In both cases, we recognized the epicuticule, but the other two layers (exocuticle and endocuticle) were inferred, because boundaries between layers were unclear. The thin sections were obtained from fragmentary specimens preserving carapace cuticle. Our examination of the best-preserved UK specimen has demonstrated a bilaminar structure in the epicuticle (16 mm thick), while the possible exocuticle measures approximately 40 mm and lamellae are not preserved across this layer; the inferred endocuticule is around 150 mm thick (Fig. 5feg). In material from Spain, the epicuticle was 15 mm, the layer interpreted as the exocuticule 24 mm and the inferred endocuticle (last preserved layer) 94 mm (Fig. 5hei). The boundaries between layers (exocuticle and endocuticle) in the UK and Spain specimens were inferred on the basis of different colorations of
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Fig. 5. Thin sections of cuticle of Meyeria magna, aeb (IGM 4685 and IGM 4686 from Colombia) and cee (IGM 4689 and IGM 4690 from Mexico), showing the three layers of cuticle: epicuticle (Ep) (bilaminar); endocuticle (En); exocuticle (Ex); and two undifferentiated layers; feg (IGM 4695 and IGM 4696 from United Kingdom) showing the three layers of cuticle, layer boundaries not clear; h-i (IGM 4693 and IGM 4694 from Spain) showing the three layers of cuticle, layer boundaries not clear. Scale bars equal 200 mm, except for b in which it equals 100 mm.
each layer, with no preservation of the lamellae tegumental ducts or pore canals in this structures (Fig. 5feg, heI, respectively). The presence of one layer (epicuticle) and two inferred layers without microstructure cannot be used to determine whether this specimen represents a corpse or a molt. For this reason, we propose to analyze more specimens using additional methods in order to establish features to differentiate between corpses and molts more clearly, irrespective of the presence of complete specimens in the fossil record of M. magna. As noted by Glaessner (1969), the disarticulated position (Salter's position) does not always represent molts (compare Bishop, 1986); conversely, the occurrence of complete specimens does not necessarily imply that such represent corpses.
7. Conclusions Eleven fragmentary specimens of Meyeria magna are described from the Simití Formation in the Velez area, province of Santander, Colombia. The occurrence of ammonites collected above and below the bed that yielded M. magna allowed to establish an Albian date for these specimens. Examination of the cuticular structure of M. magna from Colombia and Mexico in thin section helped to characterize the different layers that constitute the cuticle. Cuticle structure in material from the United Kingdom and Spain did not completely preserve the three layers of the cuticule. The absence of horizontal lamellae, canal pore, tegumental ducts and ball in the epicuticle, exocuticle and endocuticle can possibly be attributed to diagenesis. Therefore, these specimens could not be assigned to
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either corpses or molts. However, in cases in which the record is highly altered by diagenetic processes, the recognition of different layers of cuticle could be an approximation so as to distinguish between corpses or molts. Additional specimens and studies of M. magna, along with other decapod crustaceans, are needed to substantiate or refute this hypothesis. Acknowledgments The present study was supported by funds from the project noma de Me xico, through granted by Universidad Nacional Auto €l Klompmaker PAPIIT IN103214. We thank our reviewers, Adie (Florida Museum of Natural History, Gainesville), Sylvain Charum national d'Histoire naturelle, Paris) and John W. bonnier (Muse M. Jagt (Natuurhistorisch Museum, Maastricht, Netherlands) for comments and suggestions, as well as the editor-in-chief, Eduardo Koutsoukos, for comments and support. Acknowledges the field collaboration of the student group of Paleontology (Universidad ) in the year 2008. Fernando Nún ~ ez Nacional de Colombia - Bogota Useche (Laboratorio de Microscopía, Instituto de Geología UNAM) is acknowledged for providing facilities in the study of thin sections. Finally I thank to the Posgrado en Ciencias de la Tierra, Instituto de Geología, UNAM and CONACYT for their support during the development of the postgraduate studies. References Abele, L. G. (1974). Species diversity of decapod crustaceans in marine habitats. Ecology, 55, 156e161. Adams, A. E., Mackenzie, W. S., & Guilford, C. (1984). Atlas of sedimentary rocks under the microscope. Harlow: Longman, 104 pp. Amato, C., Waugh, D. A., Feldmann, R. M., & Schweitzer, C. E. (2008). The effect of calcification patterns on cuticle density in crabs: a key to lifestyle. Journal of Crustacean Biology, 28(4), 587e595. Astrop, T. I. (2011). Phylogeny and evolution of Mecochiridae (Decapoda: Reptantia: Glypheoidea): an integrated morphometric and cladistic approach. Journal of Crustacean Biology, 31(1), 114e125. Ball, H. W. (1960). Upper cretaceous Decapoda and Serpulida from James Ross Island, Graham Land. Falkland islands Dependencies survey. Scientific Reports, 24, 1e30. tace de Colombie (Am. Sud.). Basse, E. (1949). Quelques ammonites nouvelles du Cre Bulletin de la Soci et e g eologique de France, 5(18), 691e698. siles creta cicos (vermes, equínidos, crust Beurlen, K. (1938). Algunos fo aceos) de la gicos y paleontolo gicos de la Cordillera Oriental Cordillera Oriental. Estudios geolo de Colombia, parte, 3, 128e136. Bishop, G. A. (1986). Taphonomy of the North American decapods. Journal of Crustacean Biology, 6, 326e355. Bogdanova, T., & Hoedemaeker, P. H. (2004). Barremian-early albian Deshayesitidae and Silesitidae of Colombia. Scripta Geologica, 128, 183e312. Bover-Arnal, T., Moreno-Bedmar, J. A., Salas, R., Skelton, P. W., Bitzer, K., & Gili, E. (2010). Sedimentary evolution of an Aptian syn-rift carbonate system (Maestrat Basin, E Spain): effects of accommodation and environmental change. Geologica Acta, 8(3), 249e280. Bujtor, L. (2010). Systematics, phylogeny and homeomorphy of the Engonoceratidae Hyatt, 1900 (Ammonoidea, Cretaceous) and revision of Engonoceras duboisi Latil, ologie/Notebooks on Geology, Article 2010/08 (CG2010_ 1989. Carnets de Ge A08) (pp. 1e31). logo de las amonitas de Colombia. Parte I. Pulchelliidae. Bürgl, H. (1956). Cata Instituto de Geologia. Boletin de Geologia, 4(1), 1e119. Burkenroad, M. D. (1963). The evolution of the Eucarida (Crustacea, Eumalacostraca), in relation to the fossil record. Tulane Studies in Geology, 2, 3e17. n de “Mytilus vilanovae” La nderer (Bivalvo Calzada, S., & Urquiola, M. (1999). Revisio cico). Revista Espan ~ ola de Paleontología, 14(2), 293e295. creta Casey, R. (1954). New genera and subgenera of Lower Cretaceous ammonites. Journal of the Washington Academy of Sciences, 44(4), 106e115. Casey, R. (1961). The stratigraphical palaeontology of the Lower Greensand. Palaeontology, 3(4), 487e621. mien de Colombie. Eclogae Collet, L. W. (1924). Sur quelques Ammonites du Barre Geologicae Helvetiae, 18(4), 485e493. Dalingwater, J. E. (1977). Cuticular ultrastructure of a Cretaceous decapod crustacean. Geological Journal, 12, 25e32. Etayo-Serna, F. (1979). Zonation of the Cretaceous of Central Colombia by amn Especial, 2, 1e186. monites. INGEOMINAS, Publicacio Feldmann, R. M., & Ga zdzicki, A. (1998). Cuticular ultrastructure of fossil and living homolodromiid crabs (Decapoda: Brachyura). Acta Palaeontologica Polonica, 43, 1e19.
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Histoire naturelle, general et particuli ere des crustac es et des insectes (vol. 3). . Paris: F. Dufart, 468 pp. n, C. (1908). Contribucio n al conocimiento sobre algunos ammonites del Perú. Lisso In 4 Congreso Científico Latino-Americano, 1 Pan-Americano, Santiago de Chile, 16ae16c. pez-Horgue, M. A. (2009). New occurrences of Meyeria magna M'Coy, 1849 Lo (Decapoda, mecochiridae) in the early aptian and early albian of the Basque Cantabrian Basin (North Spain). Geogaceta, 47, 25e28. Mendoza-Parada, J., Moreno-Murillo, J. M., & Rodríguez-Orjuela, G. (2009). Sistema rstico de la Formacio n Rosablanca Creta cico inferior, en la provincia santanca lez, Colombia. Geología Colombiana, 34, 35e44. dereana de Ve M'Coy, F. (1849). On the classification of some British fossil Crustacea, with notices
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A new record and cuticular structure of Meyeria magna (Decapoda, Mecochiridae) from the lower Albian (Lower Cretaceous) of Colombia lez-Leo n a, b, *, Pedro Patarroyo c, Josep Anton Moreno-Bedmar d, Oscar Gonza Torrey Nyborg e, Francisco J. Vega d noma de M Posgrado en Ciencias de la Tierra, Universidad Nacional Auto exico, M exico D.F., C.P. 04510, Mexico Facultad de Estudios Superiores Iztacala, Universidad Nacional Autonoma de M exico, Estado de M exico, 54070, Mexico c , Colombia Departamento de Geociencias, Universidad Nacional de Colombia, Cr.30 N. 45e03, Bogota d noma de M Instituto de Geología, Universidad Nacional Auto exico, M exico, D.F., C.P. 04510, Mexico e Loma Linda University, Department of Earth and Biological Sciences, Loma Linda, CA 92350, USA a
b
a r t i c l e i n f o
a b s t r a c t
Article history: Received 9 March 2015 Received in revised form 13 August 2015 Accepted in revised form 14 August 2015 Available online 30 October 2015
Eleven specimens of the lobster Meyeria magna from the Simití Formation (Santander Province, Colombia) represent a new occurrence for this species in South America. Ammonites collected both below and above the bed that yielded M. magna allow dating of these specimens as early Albian. Morphological characteristics observed in these Colombian specimens were compared with those of other specimens from Mexico, the United Kingdom and Spain. Our interpretation of the cuticular structure in thin section does not unambiguously allow interpretation of the specimens studied to corpses or molts. © 2015 Elsevier Ltd. All rights reserved.
Keywords: Lobsters South America Albian Ammonites Comparisons
1. Introduction Apart from a doubtful occurrence in Asia (Wang, 1981), the widely distributed Early Cretaceous lobster Meyeria magna M'Coy, 1849 is well known in North America and Europe (e.g. Vía, 1951; Vía Boada, 1975; Simpson & Middleton, 1985; Feldmann, Vega, pez, 1995, 2007; CalGarcía-Barrera, Rico-Montiel, & Martínez-Lo zada & Urquiola, 1999; Vega, Feldmann, Etayo-Serna, Bermúdez mez, 2008; Lo pez-Horgue, 2009; Bover-Arnal et al., Aguirre, & Go n, Moreno-Bedmar, & Vega, 2010; Najarro et al., 2011; Gonz alez-Leo 2014). In the present study, a new occurrence of M. magna is noted and represents the second record of this species from Colombia and South America. The taxonomy of Meyeria magna is well established on the basis of specimens from the Isle of Wight, United Kingdom, from where it was originally described by M'Coy (1849), along with several other specimens collected from Spain, Colombia and
Mexico (Vía Boada, 1975; Simpson & Middleton, 1985; Vega et al., lez-Leo n et al., 2014). The diagnostic features of 2008; Gonza eleven Colombian specimens are compared to those of other ocurrences of M. magna. Since cuticular structure in decapods is vilavery distinctive in transverse section (Vega, Feldmann, & Da Alcocer, 1994, 2005), thin sections of several specimens of M. magna from the above-mentioned countries were made and studied in order to characterize this feature. Cuticular structure of other fossil decapod species has been previously reported (e.g. Feldmann & Tshudy, 1987; Vega et al., 1994; Feldmann & Ga zdzicki, 1998; Haj & Feldmann, 2002; Waugh & Feldmann, 2003; Waugh, Feldmann, Schroeder, & Mutel, 2006). The current work constitutes a first step toward characterizing the cuticular structure of M. magna. In addition, it contributes to the chronostratigraphic range of this widely distributed species.
2. Geological setting * Corresponding author. lez-Leo n), E-mail addresses: [email protected] (O. Gonza [email protected] (P. Patarroyo), [email protected] (J.A. Moreno-Bedmar), [email protected] (T. Nyborg), [email protected] (F.J. Vega). http://dx.doi.org/10.1016/j.cretres.2015.08.006 0195-6671/© 2015 Elsevier Ltd. All rights reserved.
The eleven specimens of Meyeria magna analyzed in the present paper were collected from the Simití Formation at co-ordinates lez area during fieldwork in the 6 030 2500 N/73 4301000 W in the Ve
lez-Leo n et al. / Cretaceous Research 57 (2016) 342e349 O. Gonza
lez area is located in Eastern Cordillera of central Colombia. The Ve (Fig. 1). The area the province of Santander to the north of Bogota has been renowned for its fossil record ever since the Humboldt expeditions, and, as such, a vast literature on the region has been published, in some cases including fossil illustrations such as Karsten (1856, 1886), Gerhardt (1897), Collet (1924), Beurlen (1938), Riedel (1938), Basse (1949), Bürgl (1956), Kakabadze and Thieuloy (1991), Bogdanova and Hoedemaeker (2004), Kakabadze and Hoedemaeker (1997, 2004) and Sharikadze, Kakabadze, and Hoedemaeker (2004). The Simití Formation is a lithostratigraphic unit within the Middle Magdalena Valley, Colombia; it overlies the Tablazo Formation in the Cundinamarca Basin (TablazoeMagdalena subbasin). lez is the upper San Its equivalent in the Villa de Leyva area near Ve Gil Formation, which overlies the lower San Gil Formation. In the present study, the Tablazo and Simití formations are preferred over the lower and upper San Gil formations because this nomenclature lez area by Ulloa and was adopted for the geological map of the Ve Rodríguez (1978, 1984) and Mendoza-Parada, Moreno-Murillo, and Rodríguez-Orjuela (2009). The Simití Formation, with a thickness of 410 m, is composed mainly of black shales with intercalated biosparites and biomicrites. Some beds show evidence of transport, which led to fossil fragmentation, while other beds contain foraminifera and flattened ammonites, bivalves, gastropods and decapod crustaceans. Ammonites of the genera Carloscaceresiceras Etayo-Serna (1979), Douvilleiceras de Grossouvre (1894) and Glottoceras Hyatt (1875), were collected both below and above specimens of M. magna, and this association indicates an early Albian date. Meyeria magna has been previously recorded from Colombia (Vega et al., 2008), in association with early Aptian cephalopod genera such as Heminautilus Spath (1927), Kutatisites Kakabadze , (1970) and Cheloniceras Hyatt (1903), from southwest of Bogota in the Upper Magdalena Valley subbasin. A depositional environment associated with initial offshore deposits following the progressive marine advance to the south of the basin is suggested. 3. Ammonite biostratigraphy In the section studied, ammonites are quite rare; two specimens
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were recovered, while a third one was left at the outcrop. The specimen of Carloscacereciseras cf. caceresi Etayo-Serna (1979) illustrated here (Fig. 2A) is an internal mold of a volumetric individual with wide sinusoid constrictions on the flank that resemble the holotype illustrated by Etayo-Serna (1979, pl. 10, Fig. 1). However, poor preservation does not allow definite identification to species level. According to Wright (1996), the genus Carloscacereciseras is a junior synonym of Anadesmoceras Casey, 1954, but we follow Bogdanova & Hoedemaeker (2004) in considering Carloscacereciseras to be a valid genus, the age range of which is early Albian (Bogdanova & Hoedemaeker, 2004). The second ammonite specimen represents a crushed shell of Douvilleiceras sp. (Fig. 2B). Because of poor preservation a specific identification is impossible. The lower Albian is usually divided into two biozones, the younger of which is the globally recorded Douvilleiceras mammillatum Zone. The genus Douvilleiceras is restricted to this zone (e.g. Casey, 1961; Reboulet et al., 2014). The equivalent in Colombia is the Douvilleiceras solitaeeNeodeshayesites columbianus Assemblage Zone (Etayo-Serna, 1979). The third ammonite, Glottoceras cf. raimondii n, 1908) (Fig. 2C), is a slightly crushed specimen with pyritic (Lisso replacement. This shell shows clear lower, mid and ventrolateral tubercles on primary ribs and intercalated ventrolateral tubercles in the ventral region are also observed. The low primary ribs start in the umbilical seam, the ventral area being flat to slightly concave. The ribs of this specimen are not as well developed as in the n (1908, p. 4, Fig. 1a, as Knemiceras raimondii); illustration by Lisso however, tuberculation is the same. Today this species is assigned to genus Glottoceras (e.g. Robert & Bulot, 2004; Bujtor, 2010), its range corresponding approximately to the middle and upper part of the Douvilleiceras mammillatum Zone. Specimens of Meyeria magna used in the present study were collected above the specimen of Carloscacereciseras cf. caceresi and below Douvilleiceras sp. and Glottoceras cf. raimondii (Fig. 2); therefore, an early Albian date is firmly established. 4. Systematic paleontology Order Decapoda Latreille, 1802 Suborder Pleocyemata Burkenroad, 1963 Infraorder Glypheidea von Zittel, 1885
Fig. 1. Locality map showing localities from which specimens of Meyeria magna were collected and stratigraphic column of the Lower Cretaceous (Albian) Simití Formation.
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Fig. 2. A. Lateral view of a specimen of Carloscacereciseras cf. caceresi, embedded in rock; B. Lateral view of Douvilleiceras sp. (UN-DG-AM-241); C. Lateral view of Glottoceras cf. raimondii (UN-DG-AM-240). Scale bar equals 10 mm.
Superfamily Glypheoidea von Zittel, 1885 Family Mecochiridae Van Straelen, 1925 Genus: Meyeria M'Coy, 1849 Type species. Astacus ornatus Phillips, 1829, by original designation. Included species. Meyeria ornata (Phillips, 1829); Meyeria magna M'Coy, 1849; Meyeria harveyi (Woodward, 1900); Meyeria rapax (Harbort, 1905); Meyeria schwarzi (Kitchin, 1908); Meyeria bolivari (Van Straelen, 1927); Meyeria gracilis (Glaessner, 1932); Meyeria mexicana (Rathbun, 1935); Meyeria houdardi (Van Straelen, 1936) and Meyeria crofti (Ball, 1960). Meyeria magna M'Coy, 1849 Figs. 3e4 1849 Meyeria magna M'Coy, p. 334, Fig. 4. 1863 Meyeria vectensis Bell, p. 33, pl. 10, Figs. 1e5. 1863 Oncopareia granulosa Vilanova, p.98, pl. 3, Fig. 2. 1881 Meyeria pearcei Spence-Bate in Lee, p. 87, pl. 204, Fig. 14. 1927 Meyeria bolivari Van Straelen, p. 80, pl. 1, Figs. 1e2. 1951 Hoploparia granulosa Vía, p. 154, Fig. 10. 1975 Meyeria magna M'Coy, Vía Boada, p. 33, Figs. 1.1e1.9, 2.1e2.6. 1985 Meyeria magna M'Coy, Simpson and Middleton, pp. 204e211, Figs. 1A, 2AeM, 6E, H, 7H, 8AeB, 9AeL. 1995 Meyeria pueblaensis Feldmann, Vega, García-Barrera, Rico pez, p. 404, Figs. 2.1e2.4. Montiel and Martínez-Lo 2008 Meyeria magna M'Coy, Vega et al., p. 5, Figs. 5.1e5.7, 6.1e6.7, 7.1, 7.6. pez-Horgue, p. 27, Fig. 2AeJ. 2009 Meyeria magna M'Coy, Lo 2011 Meyeria magna M'Coy, Astrop, 2011, p. 116, Fig. 1B. 2013 Meyeria magna M'Coy, Klompmaker, appendices AeB. lez-Leo n et al., pp. 10e14, 2014 Meyeria magna M'Coy, Gonza Figs. 10AeQ, 11AeM, 12AeJ.
5. Material and methods 5.1. Description of specimens 5.1.1. Carapace Laterally compressed and subcylindrical; cephalic region with three longitudinal spiny carinae; orbital, gastro-orbital and antennal carina parallel; short distance between orbital and gastroorbital carinae, antennal carina separated twice the distance from the others; cephalic carinae raised, antennal carina more raised than others; cervical groove deep, two-thirds of height of carapace, inclined 53 toward lower anterion margin, cervical groove
ventrally connected to antennal groove; branchiocardiac groove shallow, inclined 15 from upper part of posterior margin to midheight of carapace; post-cervical groove shallow and parallel to branchiocardiac groove; hepatic groove slightly deep, convex ventrally at intersection with antennal groove; shallow and undeveloped inferior groove, connected to hepatic groove; cuticle on anterior cardiac region with a few small tubercles; cuticle on branchial region covered by small tubercles of uniform size, becoming more numerous toward ventral margin and margin hepatic lobe. 5.1.2. Pleon Pleonal somites with three segments of longitudinal rows of tubercles, anterior and lower margins rounded, posterior margin straight; second to fifth pleonal somites of similar shape and size; sixth pleural segment triangular, strong ridge at contact with tergum; endopodite and exopodite of telson triangular, both with median keel, diaeresis present. Appendages not preserved, only fragments of first pereiopod and coxae of pereiopods 1e5. 5.2. Discussion Specimens are well preserved and only slightly crushed. Morphological features observed include: cephalic carinae (cc ¼ oc orbital carina þ gc gastro-orbital carina þ antennal carina) (Fig. 3A); three thoracic ridges in the branchial region (Fig. 3 A1, A2 and B); cervical, (e1e), hepatic (b1), branchiocardiac (a) and postcervical (c) grooves (Fig. 3 A1, A2, B, E1 and E2); ornamentation of granules on carapace can be observed (Fig. 3 A2, specimens D and E1), as well as ornamentation of the pleon (Fig. 3 C, F1 and F2). In Fig. 3D the diaeresis in the exopodite of the telson can be seen. Such characteristics have been recognized previously allowing the unequivocal assignment of the Colombian material to Meyeria magna. Fig. 4 illustrates carapace features observed in the Colombian material. 5.3. Material Eleven specimens of Meyeria magna (UN-DG-CR-020eUN-DGlez area, CR-030) from the lower Albian (Simití Formation) in the Ve n province of Santander, Colombia are housed in the Coleccio gica del Departamento de Geociencias, Universidad Paleontolo Nacional de Colombia.
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Fig. 3. AeF. Meyeria magna from the lower Albian of Colombia. Carapace UN-DG-CR-020, in left lateral (A1) and right lateral views (A2); B. Partial carapace and pleon, UN-DG-CR023, in right lateral view; C. UN-DG-CR-021 showing five pleonal somites and a partial telson, left side; D. Carapace fragment, UN-DG-CR-025, showing pleon and telson; E1. UN-DGCR-022, showing ornamentation of carapace, left lateral view; E2. Same specimen, right lateral view; F1. Detail of UN-DG-CR-030, pleonal somites, right lateral view; F2. Same specimen, showing details of pleon, left lateral view. Scale bar equals 10 mm.
5.4. Cuticular structure The material studied represents 12 thin sections obtained from five specimens of Meyeria magna, one from Colombia, two from Mexico, one from United Kingdom and one from Spain. The thin sections are housed in the Museo María del Carmen Perrilliat M., n Nacional de Paleontología, Instituto de Geología, Coleccio
noma de Me xico, Me xico D.F., Mexico Universidad Nacional Auto with the numbers IGM 4685 to IGM 4696. 5.4.1. Colombia (V elez) Four thin sections (numbered IGM 4685, IGM 4686, IGM 4687 and IGM4688) obtained from two median sections of the carapace. The age of this material is early Albian. 5.4.2. Mexico (Puebla) Four thin sections (numbered IGM 4689, IGM 4690, IGM 4691 and IGM 4692) obtained from the median part of the sectioned carapace and pereiopods. The age of this material is late Barremianeearly Aptian. 5.4.3. Spain (Cuchía) Two thin sections (numbered IGM 4693 and IGM 4694) obtained from the branchial region of the carapace. The age of this material also is early Aptian.
Fig. 4. Meyeria magna, UN-DG-CR-020, cephalothorax. Morphological characteristics: e1e ¼ cervical groove, c ¼ postcervical groove, a ¼ branchiocardiac groove, b1 ¼ hepatic groove, cephalic carinae (oc ¼ orbital carina, gc ¼ gastro-orbital carina, ac ¼ antennal carina) and tr ¼ thoracic ridges. Scale bar equals 10 mm.
5.4.4. United Kingdom (Isle of Wight) Two thin sections (numbered IGM 4695 and IGM 4696) obtained from the branchial region of the carapace. The age of this material is early Aptian.
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6. Cuticular structure in decapod crustaceans The order Decapoda includes organisms commonly called shrimps, lobsters and crabs. Decapod crustaceans rank amongst the most common animals inhabiting a number of different environments, both at present times (e.g. Abele, 1974) and in the past (e.g. Klompmaker et al., 2013, Schweitzer, Feldmann, & Kowalewski, 2013; Schweitzer & Feldmann, 2014). They are commonly found in the fossil record despite their low preservation potential when compared to other marine invertebrates (Kidwell & Flessa, 1995). For many fossil decapod crustacean taxa, morphological features such as the shape and contour of the carapace, the number and placement of spines, nodes and grooves and the orbital design, are the basis for classification (Haj & Feldmann, 2002). Decapod cuticle has a very distinctive structure when observed in cross section. Although most references on petrography and sedimentology illustrate or describe how to recognize some arthropod bioclasts such as those of trilobites and ostracods (Adams, Mackenzie, & Guilford, 1984; Flügel, 2004), there is no such description for decapods, in spite of the fact that decapod cuticle is of potentially common occurrence in Mesozoic and vila-Alcocer, & Filkorn, 2005). The Cenozoic shelf deposits (Vega, Da cuticle of extant decapod crustaceans is commonly composed of four discrete layers, namely epicuticle, exocuticle, endocuticle and a membranous layer, which are all well documented in the literature (e.g., Neville & Berg, 1971; Taylor, 1973; Dalingwater, 1977; Vega et al., 1994, 1998; Feldmann & Ga zdzicki, 1998; Haj & Feldmann, 2002; Vega et al., 2005; Amato, Waugh, Feldmann, & Schweitzer, 2008). The outermost layer of the cuticle is the epicuticle, commonly a mineralized bilaminar layer. The exocuticle is composed of chitinprotein fibers stacked in layers of continuously changing orientation (Green & Neff, 1972; Haj & Feldmann, 2002). The endocuticle is the thicker and most heavily calcified layer (Roer & Dillaman, 1984). In some lobsters, such as nephropids, the outer part of the endocuticle consists of broad lamellae while the innermost portion is thinly laminated (Feldmann & Tshudy, 1987). The innermost layer is the membranous layer, an uncalcified layer (Roer & Dillaman, 1984; Haj & Feldmann, 2002), making the chances of preservation in the fossil record low. Characterization of the cuticular structure has been used as evidence to suggest the presence of both molts and corpses in the fossil record of decapod crustaceans. Feldmann and Tshudy (1987) analyzed 58 specimens of Hoploparia stokesi (Weller, 1903) and concluded that the presence of horizontal lamellae of the endocuticle could be used to differentiate between corpses and molts. Vega et al. (1994) recognized the endocuticular, exocuticular lamellae, probably tegumental ducts, cell imprints in the epicuticular layer and the membranous layer in many samples of Costacopluma mexicana Vega & Perrilliat, 1989, suggesting the presence of corpses in the record of this species. For this reason, the study of cuticular structure in these organisms can be a potential tool in preliminary identification of major decapod groups and taphonomic interpretations (Feldmann & Tshudy, 1987; Vega et al., 1994; Klompmaker, Hy zný, & Jakobsen 2015). In the present study, we analyze and describe the cuticular structure of Meyeria magna employing thin sections of five specimens from different countries. 6.1. Analysis and discussion Analysis of cuticular structure in M. magna thin sections did not fully allow assignment of these specimens to either corpses or molts. In some specimens three discrete layers (epicuticle, exocuticle and endocuticle) could be distinguished whereas in others only a
single layer was clearly observed, the epicuticle. These features have previously been recognized (Feldmann & Tshudy, 1987; Vega et al., 1994). In none of the studied thin sections could the microstructure (horizontal lamellae, pore canals, tegumental ducts and balls) be observed. These features may have either been altered during the diagenetic process or were never present in the first place; a process difficult to determine. Examination of the Colombian thin sections shows three discrete layers of cuticle. The epicuticle is approximately 13 mm thick (Fig. 5aeb); the exocuticle is 23 mm thick and shows no evidence of lamellar structure (Fig. 5a). In the thickest layer (57 mm), the endocuticle was partially replaced by sparite crystals during diagenesis. Below the endocuticle it is possible to discern two other undifferentiated layers (Fig 5a). Since the membranous layer is not ordinarily preserved in the fossil record, we infer that these layers may reflect the diagenetic process. Unfortunately, insufficient preservation of the microstructure of the different layers does not permit us determine whether the Colombian material represent corpses or molts. The Colombian specimens of Meyeria magna were preserved disarticulated, suggesting the specimens were molts; however this is not necessarily true. There are many destructive processes that can lead to fragmentation of corpses, such as physical disturbance, transport and destruction in the sedimentary environment, predation and scavenging, bacterial degradation of soft tissues and exoskeleton, as well as chemical breakdown of such remains (Plotnick, 1986). Examination of Mexican specimens has allowed characterization of the same layers as those seen in the Colombian material (Fig. 5cee). We observed a bilaminar epicuticle (Fig. 5c), 10 mm thick, as the first layer; the second layer was identified as the exocuticle, 26 mm thick, and the third layer was the endocuticle, measuring 140 mm (Fig. 5dee). The characteristic lamellae and pore canals are absent in the Mexican specimens. As in the Colombian material, we also noted two undifferentiated layers below the base of the endocuticule that could be interpreted as the membranous layer; however, this hypothesis cannot be confirmed. The boundaries between exocuticle and endocuticle were only inferred (Fig. 5dee). Despite the presence of discrete layers in the cuticle of the Mexican specimens, we cannot document this record as corpses because the microstructure (horizontal lamellae) in the endocuticle and exocuticle is not evident. It is possible that these features were lost during diagenesis, but this is difficult to determine. When epicuticle, exocuticle and endocuticle layers are present, cuticle structure does suggest a specimen represents a corpse, as well as, when there is only one layer present, cuticular structure suggests a specimen represents a molt. This criterion can be applied in thin sections, when it is seen that the diagenetic process altered the cuticle microstructure. Based upon these observations, specimens from the United Kingdom and Spain could be interpreted as molts. In both cases, we recognized the epicuticule, but the other two layers (exocuticle and endocuticle) were inferred, because boundaries between layers were unclear. The thin sections were obtained from fragmentary specimens preserving carapace cuticle. Our examination of the best-preserved UK specimen has demonstrated a bilaminar structure in the epicuticle (16 mm thick), while the possible exocuticle measures approximately 40 mm and lamellae are not preserved across this layer; the inferred endocuticule is around 150 mm thick (Fig. 5feg). In material from Spain, the epicuticle was 15 mm, the layer interpreted as the exocuticule 24 mm and the inferred endocuticle (last preserved layer) 94 mm (Fig. 5hei). The boundaries between layers (exocuticle and endocuticle) in the UK and Spain specimens were inferred on the basis of different colorations of
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Fig. 5. Thin sections of cuticle of Meyeria magna, aeb (IGM 4685 and IGM 4686 from Colombia) and cee (IGM 4689 and IGM 4690 from Mexico), showing the three layers of cuticle: epicuticle (Ep) (bilaminar); endocuticle (En); exocuticle (Ex); and two undifferentiated layers; feg (IGM 4695 and IGM 4696 from United Kingdom) showing the three layers of cuticle, layer boundaries not clear; h-i (IGM 4693 and IGM 4694 from Spain) showing the three layers of cuticle, layer boundaries not clear. Scale bars equal 200 mm, except for b in which it equals 100 mm.
each layer, with no preservation of the lamellae tegumental ducts or pore canals in this structures (Fig. 5feg, heI, respectively). The presence of one layer (epicuticle) and two inferred layers without microstructure cannot be used to determine whether this specimen represents a corpse or a molt. For this reason, we propose to analyze more specimens using additional methods in order to establish features to differentiate between corpses and molts more clearly, irrespective of the presence of complete specimens in the fossil record of M. magna. As noted by Glaessner (1969), the disarticulated position (Salter's position) does not always represent molts (compare Bishop, 1986); conversely, the occurrence of complete specimens does not necessarily imply that such represent corpses.
7. Conclusions Eleven fragmentary specimens of Meyeria magna are described from the Simití Formation in the Velez area, province of Santander, Colombia. The occurrence of ammonites collected above and below the bed that yielded M. magna allowed to establish an Albian date for these specimens. Examination of the cuticular structure of M. magna from Colombia and Mexico in thin section helped to characterize the different layers that constitute the cuticle. Cuticle structure in material from the United Kingdom and Spain did not completely preserve the three layers of the cuticule. The absence of horizontal lamellae, canal pore, tegumental ducts and ball in the epicuticle, exocuticle and endocuticle can possibly be attributed to diagenesis. Therefore, these specimens could not be assigned to
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either corpses or molts. However, in cases in which the record is highly altered by diagenetic processes, the recognition of different layers of cuticle could be an approximation so as to distinguish between corpses or molts. Additional specimens and studies of M. magna, along with other decapod crustaceans, are needed to substantiate or refute this hypothesis. Acknowledgments The present study was supported by funds from the project noma de Me xico, through granted by Universidad Nacional Auto €l Klompmaker PAPIIT IN103214. We thank our reviewers, Adie (Florida Museum of Natural History, Gainesville), Sylvain Charum national d'Histoire naturelle, Paris) and John W. bonnier (Muse M. Jagt (Natuurhistorisch Museum, Maastricht, Netherlands) for comments and suggestions, as well as the editor-in-chief, Eduardo Koutsoukos, for comments and support. Acknowledges the field collaboration of the student group of Paleontology (Universidad ) in the year 2008. Fernando Nún ~ ez Nacional de Colombia - Bogota Useche (Laboratorio de Microscopía, Instituto de Geología UNAM) is acknowledged for providing facilities in the study of thin sections. Finally I thank to the Posgrado en Ciencias de la Tierra, Instituto de Geología, UNAM and CONACYT for their support during the development of the postgraduate studies. References Abele, L. G. (1974). Species diversity of decapod crustaceans in marine habitats. Ecology, 55, 156e161. Adams, A. E., Mackenzie, W. S., & Guilford, C. (1984). Atlas of sedimentary rocks under the microscope. Harlow: Longman, 104 pp. Amato, C., Waugh, D. A., Feldmann, R. M., & Schweitzer, C. E. (2008). The effect of calcification patterns on cuticle density in crabs: a key to lifestyle. Journal of Crustacean Biology, 28(4), 587e595. Astrop, T. I. (2011). Phylogeny and evolution of Mecochiridae (Decapoda: Reptantia: Glypheoidea): an integrated morphometric and cladistic approach. Journal of Crustacean Biology, 31(1), 114e125. Ball, H. W. (1960). Upper cretaceous Decapoda and Serpulida from James Ross Island, Graham Land. Falkland islands Dependencies survey. Scientific Reports, 24, 1e30. tace de Colombie (Am. Sud.). Basse, E. (1949). Quelques ammonites nouvelles du Cre Bulletin de la Soci et e g eologique de France, 5(18), 691e698. siles creta cicos (vermes, equínidos, crust Beurlen, K. (1938). Algunos fo aceos) de la gicos y paleontolo gicos de la Cordillera Oriental Cordillera Oriental. Estudios geolo de Colombia, parte, 3, 128e136. Bishop, G. A. (1986). Taphonomy of the North American decapods. Journal of Crustacean Biology, 6, 326e355. Bogdanova, T., & Hoedemaeker, P. H. (2004). Barremian-early albian Deshayesitidae and Silesitidae of Colombia. Scripta Geologica, 128, 183e312. Bover-Arnal, T., Moreno-Bedmar, J. A., Salas, R., Skelton, P. W., Bitzer, K., & Gili, E. (2010). Sedimentary evolution of an Aptian syn-rift carbonate system (Maestrat Basin, E Spain): effects of accommodation and environmental change. Geologica Acta, 8(3), 249e280. Bujtor, L. (2010). Systematics, phylogeny and homeomorphy of the Engonoceratidae Hyatt, 1900 (Ammonoidea, Cretaceous) and revision of Engonoceras duboisi Latil, ologie/Notebooks on Geology, Article 2010/08 (CG2010_ 1989. Carnets de Ge A08) (pp. 1e31). logo de las amonitas de Colombia. Parte I. Pulchelliidae. Bürgl, H. (1956). Cata Instituto de Geologia. Boletin de Geologia, 4(1), 1e119. Burkenroad, M. D. (1963). The evolution of the Eucarida (Crustacea, Eumalacostraca), in relation to the fossil record. Tulane Studies in Geology, 2, 3e17. n de “Mytilus vilanovae” La nderer (Bivalvo Calzada, S., & Urquiola, M. (1999). Revisio cico). Revista Espan ~ ola de Paleontología, 14(2), 293e295. creta Casey, R. (1954). New genera and subgenera of Lower Cretaceous ammonites. Journal of the Washington Academy of Sciences, 44(4), 106e115. Casey, R. (1961). The stratigraphical palaeontology of the Lower Greensand. Palaeontology, 3(4), 487e621. mien de Colombie. Eclogae Collet, L. W. (1924). Sur quelques Ammonites du Barre Geologicae Helvetiae, 18(4), 485e493. Dalingwater, J. E. (1977). Cuticular ultrastructure of a Cretaceous decapod crustacean. Geological Journal, 12, 25e32. Etayo-Serna, F. (1979). Zonation of the Cretaceous of Central Colombia by amn Especial, 2, 1e186. monites. INGEOMINAS, Publicacio Feldmann, R. M., & Ga zdzicki, A. (1998). Cuticular ultrastructure of fossil and living homolodromiid crabs (Decapoda: Brachyura). Acta Palaeontologica Polonica, 43, 1e19.
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