© Author(s) 2020. CC Atribution 4.0 License
Well-preserved cuticle of Atherfieldastacus magnus  
(Decapoda, Glypheida) from the Aptian of Mexico 
Dobro ohranjena kutikula raka Atherfieldastacus magnus  
(Decapoda, Glypheida) iz aptijskih plasti v Mehiki
Oscar GONZÁLEZ-LEÓN1, Josep A. MORENO-BEDMAR2, Ricardo BARRAGÁN-MANZO2  
& Francisco J. VEGA2
1Posgrado en Ciencias de la Tierra, Universidad Nacional Autónoma de México, CDMX 04510, México,  
and Facultad de Estudios Superiores Iztacala, Universidad Nacional Autónoma de México, 54070 México;  
e-mail: oscar.gonzalez@unam.mx
2Instituto de Geología, Universidad Nacional Autónoma de México, Ciudad Universitaria, Coyoacán, CDMX 04510, 
Mexico; e-mails: josepamb@geologia.unam.mx, ricardor@geologia.unam.mx, vegver@unam.mx
Prejeto / Received 9. 12. 2019; Sprejeto / Accepted 2. 4. 2020; Objavljeno na spletu / Published online 22. 4. 2020
Key words: Pleocyemata, Mecochiridae, cuticular structure, Lower Cretaceous, Chihuahua, Mexico
Ključne besede: Pleocyemata, Mecochiridae, kutikula, spodnja kreda, Chihuahua, Mehika
Abstract
The cuticle structure of fossil decapod crustaceans is an important tool, not only for palaeocological and 
taphonomic interpretations, but also as a potential way to characterise systematically genera and even species 
the cuticle of which has not been severely altered by diagenetic processes. Localities with abundant decapod 
crustacean remains can be interpreted either as reflecting mass mortality events or just simple accumulations 
of exuviae, on the basis of completeness and comparison of cuticle structures between specimens of the same 
species from different localities. Association with anoxic events by microfacies analyses can offer clues to 
explain the unusual abundance of decapod crustacean remains. This is the case for the Early Cretaceous lobster 
Atherfieldastacus magnus (M’Coy, 1849), which is found in large numbers in different Lower Cretaceous (mainly 
Aptian) lithostratigraphic units across the globe. In this case, we document the well-preserved cuticle structure of 
specimens from the upper Aptian of Chihuahua (Mexico), preserved three-dimensionally, mainly in concretions, 
which were studied in different transverse sections showing the cuticle in diverse portions of the lobster body. 
Thin cuticle layers show the typical crustacean cuticular structure that suggest these are corpses preserved in 
an anoxic environment.
Izvleček
Analiza strukture kutikule fosilnih deseteronožcev je pomembno orodje ne le za paleoekološke in tafonomske 
interpretacije, ampak tudi kot možen način za sistematsko opredelitev rodov in celo vrst, v kolikor kutikula ni 
diagenetsko spremenjena. Na podlagi ohranjenosti in primerjave strukture kutikule med primerki iste vrste z 
različnih nahajališč razlikujemo nahajališča s pogostimi ostanki rakov. Ta lahko kažejo na množičen pogin ali 
zgolj na akumulacije levov deseteronožcev. V povezavi s prepoznanimi anoksičnimi dogodki v mikrofacialnih 
analizah nam lahko metoda služi za razlago množičnih nakopičenj fosilnih deseteronožcev na nekaterih lokacijah. 
Tak primer je zgodnjekredni jastog Atherfieldastacus magnus (M'Coy, 1849), katerega številne ostanke najdemo 
v različnih litostratigrafskih enotah spodnje krede (predvsem v aptiju) po vsem svetu. V prispevku predstavljamo 
dobro ohranjeno strukturo kutikule osebkov iz zgornjega aptija iz nahajališča Chihuahua (Mehika). Vzorce 
tridimenzionalno ohranjene kutikule primerkov iz konkrecij smo pregledali na različnih prečnih presekih z 
različnih delov telesa jastoga. Tanke plasti kutikule z značilno strukturo kažejo, da gre v našem primeru za 
trupla, ki so se ohranila v anoksičnem okolju.
GEOLOGIJA 63/1, 39-46, Ljubljana 2020
https://doi.org/10.5474/geologija.2020.004
40 Oscar GONZÁLEZ-LEÓN, Josep A. MORENO-BEDMAR, Ricardo BARRAGÁN-MANZO, & Francisco J. VEGA
Introduction
An interesting factor of the study of decapod 
crustaceans is the review and examination of 
their cuticule structure. At most localities, cuticle 
structure is obscured by mineral replacement of 
the original carbonate, but modified by diagenet-
ic processes as well (Vega et al., 2005). In previous 
studies (e.g., Dennell, 1960; Hegdahl et al., 1977a, 
b; Roer & Dillaman, 1984), cuticle structure of 
Recent taxa has been studied, while other authors 
have demonstrated the presence of cuticle in the 
fossil record (e.g., Neville & Berg, 1971; Feldmann 
& Tshudy, 1987; Vega et al., 1994, 2005; Feld-
mann & Gaździcki, 1998; Guinot & Breton; 2006; 
González-León et al., 2016, 2018, among others). 
Studies of the functional morphology and tapho-
nomic implications have been addressed by vari-
ous authors (Schäfer, 1951; Guinot, 1979; Plotnick 
et al., 1988; Savazzi, 1988; Haj & Feldmann, 2002; 
Waugh et al., 2004). The use of this structure for 
taxonomic purposes is complicated because there 
are only few well-established characters. With 
this in mind, Waugh et al. (2009) analysed the 
morphological characters of some decapod crus-
taceans for possible future phylogenetic analysis.
Decapod crustaceans rank amongst the most 
common animals inhabiting a number of differ-
ent environments, both at the present day (Abele, 
1974) and in the past (Klompmaker et al., 2013; 
Schweitzer & Feldmann, 2014). The calcified cu-
ticle of decapod crustaceans comprises the hard 
exoskeleton of the animal and is composed of 
three layers (Haj & Feldmann, 2002); these lay-
ers have been documented in some fossil decapod 
crustaceans as well (Neville & Berg, 1971; Taylor, 
1973; Dalingwater, 1977; Vega et al., 1994, 1998; 
Feldmann & Gaździcki, 1998; Haj & Feldmann, 
2002; Waugh & Feldmann, 2003; Vega et al., 2005; 
Waugh et al., 2006; Amato et al., 2008; Waugh et 
al., 2009; González-León et al., 2016, 2018). The 
decapod cuticle has a very distinctive structure 
when observed in cross section. In spite of the fact 
that decapod crustacean cuticle is frequently pre-
served in material from Mesozoic and Cenozoic 
shelf deposits (Vega et al., 2005), very few efforts 
have been made as to how to distinguish corpses 
from exuviae. For this reason, it is important to 
recognise and characterise the microstructure as 
a potential tool in preliminary identification of, 
at least, major decapod crustacean groups and 
taphonomic interpretations (Feldmann & Tshudy, 
1987; Vega et al., 1994; Klompmaker et al., 2015). 
The present paper analyses and complements 
information on cuticule structure of numerous 
specimens of Atherfieldastacus magnus that are 
preserved in concretions from the upper Aptian 
La Peña Formation in Chihuahua State (northern 
Mexico).
Locality and stratigraphy
The main locality is in the Cerro Chino region 
(Chihuahua State), close to the towns of Coyame 
del Sotol and Cuchillo Parado (Fig. 1). Specimens 
were collected from upper Aptian strata assigned 
to the La Peña Formation (Fig. 2); for details on 
these localities and local stratigraphy, reference 
is made to Ovando-Figueroa et al. (2017) and 
González-León et al. (2018).
Fig. 1. Locality map showing the fossil site in northern Mexico (Chihuahua State) (modified from González-León et al., 2018).
41Well-preserved cuticle of Atherfieldastacus magnus (Decapoda, Glypheoidea) from the Aptian of Mexico
Material and methods
About 20 calcareous concretions were collect-
ed near Abuja Colorada, in a fossiliferous section 
dominated by shale. Specimens recorded herein 
were recovered from concretions of varying size, 
between 3 and 12 cm in length (Fig. 3) and were 
prepared with a Paleotools ME-9100 pneumatic 
percutor and subsequently sectioned transversely 
with a diamond saw blade and glued to microscop-
ic slides with resin, which were then polished by 
hand, using Kemet polishing abrasive. A Zeiss po-
larising microscope, with an adapted Canon EOS 
Mark I camera, was used to take numerous images 
of cuticule structure. Thin sections and complete 
specimens are deposited in the Colección Nacion-
al de Paleontología “María del Carmen Perrilli-
at”, Instituto de Geología, Universidad Nacional 
Autónoma México (abbreviation: IGM).
Systematic palaeontology
Order Decapoda Latreille, 1802
Suborder Pleocyemata Burkenroad, 1963
Infraorder Glypheida Zittel, 1885
Superfamily Glypheoidea von Zittel, 1885
Family Mecochiridae Van Straelen, 1925
Genus Atherfieldastacus Simpson in Robin, 
Charbonnier, Merle, Simpson, Petit & Fernandez, 
2016
Fig. 2. Stratigraphical sections of outcrops of upper Aptian 
strata in Chihuahua State, showing fossiliferous beds (modi-
fied from González-León et al., 2018).
Fig. 3. Atherfieldastacus magnus (M’Coy, 1849), Abuja Colorada Canyon section (locality 1), Chihuahua State, northern 
Mexico; a near-complete specimen (IGM 9478) preserved in a calcareous nodule. Anatomical abbreviations are as follows: a = 
branchiocardiac groove; ac =antennal carina; b = antennal groove; c = post-cervical groove; cd = cardiac groove; e1e = cervical 
groove; gc = gastro-orbital carina; hr = hepatic ridge; i = inferior groove; p1-2 = pereiopods; r1-r3 = branchial ridges; s2-6 = 
pleonal somites; t = telson. Scale bar in cm. Photograph: Josep A. Moreno-Bedmar (modified from González-León et al., 2018).
42 Oscar GONZÁLEZ-LEÓN, Josep A. MORENO-BEDMAR, Ricardo BARRAGÁN-MANZO, & Francisco J. VEGA
Fig. 4. A-G, Several views of thin sections of Atherfieldastacus magnus (M’Coy, 1849) from the upper Aptian of Chihuahua 
State, northern Mexico. Abbreviations: Endo = endocuticle; Epi = epicuticle; Exo = exocuticle.
43Well-preserved cuticle of Atherfieldastacus magnus (Decapoda, Glypheoidea) from the Aptian of Mexico
Fig. 5. Several views of thin sections of Atherfieldastacus magnus (M’Coy, 1849) from the upper Aptian of Chihuahua State, 
northern Mexico. Abbreviations: Biot = bioturbation; Endo = endocuticle; Epi = epicuticle; Exo = exocuticle; Pca = pore 
channels; Pyr = pyrite; Qz = quartz. Scale bars in µm.
44 Oscar GONZÁLEZ-LEÓN, Josep A. MORENO-BEDMAR, Ricardo BARRAGÁN-MANZO, & Francisco J. VEGA
Type species: Meyeria magna M’Coy, 1849, by 
original designation.
Other included species: Atherfieldastacus ra-
pax (Harbort, 1905) and A. schwartzi (Kitchin, 
1908).
Atherfieldastacus magnus (M’Coy, 1849)
(Fig. 3)
Diagnosis: See González-León et al. (2018).
Material examined: Specimens in 22 calcare-
ous concretions, of which eight were sectioned for 
analysis of cuticular structure; in total, 30 thin 
sections of different portions of the lobster body 
were obtained.
Cuticle structure
Analysis and discussion: In our analysis of cut-
icule structure, it was possible to recognise clear-
ly the three cuticle layers. In some cases, only a 
single layer was discernible. Elements of cuticule 
microstructure, such as pore canals, were also 
observed (Figs. 4, 5). Previously, such features 
had been recorded by Feldmann & Tshudy (1987), 
Vega et al. (1994) and González-León et al. (2016, 
2018), both for other species and for Atherfieldas-
tacus magnus, but recrystallised cuticles do not 
show clear layers (González-León et al., 2016).
The newly collected specimens clearly pres-
ent three discrete layers of cuticle. The first layer 
observed is the epicuticle (epi), which normally 
has a thin bilaminar structure; this could not be 
observed. Below the epicuticle is the second layer 
or exocuticle (exo), composed of chitin protein fi-
bres, stacked in layers with variable orientations 
(Green & Neff, 1972; Haj & Feldmann, 2002). This 
layer is altered, but still discernible in almost all 
specimens studied (Fig. 4). The microstructure 
is replaced by sparry calcite, as seen in Figures 
4A-C and G, although some fibres can still be not-
ed (Fig. 4E). The most strongly calcified layer is 
the third one; this is the endocuticle (endo) which 
presents broad lamellae in the outer portion and 
thin laminations on the inner part (Feldmann & 
Tshudy, 1987). Vertical laminations within the 
endocuticle were noted in specimens from Chi-
huahua and interpreted as pore channels (Figs. 
4A, E; 5H, K). A pigmented layer at the top of the 
endocuticle could also be observed (Fig. 4D-F). 
This might be associated with the original pig-
ment (quinona), as previously recognised by Tay-
lor (1973) and Vega et al. (1994). An example of 
how the microstructure and boundaries between 
layers can be altered by diagenetic processes 
was observed as well (Fig. 4G). The epicuticle 
can be clearly recognised (Figs. 4B, E; 5F, G, I, 
L), but only as a single layer, not as a double lay-
er, which is typical. The membranous layer was 
not preserved, similar to what has been recorded 
for other extinct species (Roer & Dillaman, 1984; 
Vega et al., 1994, 2005; Haj & Feldmann, 2002).
Conclusions
The completeness of cuticule structure (espe-
cially the basis of the endocuticle) and the 3-D 
preservation and articulation of carapaces with 
appendages suggest that the Chihuahua spec-
imens represent corpses that were accumulat-
ed during anoxic events. The presence of small 
pyrite crystals in the matrix and larger ones in 
appendages (Fig. 5B, C, J) supports such an in-
terpretation, along with bioturbations observed 
in some thin sections; these were possibly caused 
by scavengers that were feeding on cuticle re-
mains and other organic matter (Fig. 5A). Abun-
dant pyrite has also been observed in specimens 
of Atherfieldastacus magnus from the Aptian of 
Colombia (González-León et al., 2016). This sug-
gests that localities around the world, where A. 
magnus is abundant, may represent anoxic events 
that either killed the lobster populations and/or 
preserved the remains of this globally distribut-
ed species during the Early Cretaceous.
Acknowledgements
We are grateful for support from the Universidad 
Nacional Autónoma de México, through Dirección 
General de Asuntos del Personal Académico, with 
project PAPIIT-IN107617. Our sincere thanks go to 
Rok Gašparič for kindly invitating us to participate 
in this special volume, to Torrey Nyborg (Loma Linda 
University, California, USA) and Guenter Schweigert 
(Staatliches Museum für Naturkunde, Stuttgart, 
Germany) and John Jagt (Natuurhistorisch Museum 
Maastricht, The Netherlands) for suggestions on how 
to improve an earlier version of the typescript and to 
Marco A. Argáez Martínez (Invertebrate Paleontology 
Laboratory, Instituto de Geología, Universidad 
Nacional Autónoma de México) who produced all thin 
sections.
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