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Cuticular microstructure of some Silurian Homalonotid trilobites from Sweden

Dalingwater, John E

ABSTRACT-Etched slices of the cuticle of the Silurian trilobite Homalonotus rhinotropis Angelin, 1854, from two localities in Skane, southern
Sweden, have been examined with the scanning electron microscope. The two major subdivisions of the cuticle are a "prismatic" (originally
laminated) outer layer, about 20-30 (mu)m thick, and a principal layer, 200-300 ,(mu)m thick, which may be foliated or roughly laminated.
The original finely-laminated aspect of the outer layer may sometimes be seen as a continuous layer, but is more frequently detected as "shaped
inclusions" set in a prismatic background. These "shaped inclusions" are considered to have been produced by accumulation of minerals around
the apices of 3-6 (mu)m perpendicular canals early in calcification and preferentially preserved because of subtle differences from the products of
later stages of the process. It is possible that calcification started before ecdysis. The difficulties involved in ascertaining the original structure of
the principal layer are outlined, and possible reasons for differences in its preservation from the outer layer are proposed.

INTRODUCTION

EXTANT ARTHROPODS with mineralized exoskeletons are in the minority. A few rare examples can be found in the insects and in the
chelicerates; amongst the myriapods only the millipedes have calcified cuticles. Crustaceans are generally characterized as having heavily
mineralized exoskeletons, but while this is true of ostracodes and cirripedes, other groupssuch as copepods-have entirely organic cuticles. Even
within the malacostracans (the "higher Crustacea," which includes the crabs and lobsters) examples can be found with unmineralized cuticles.
Trilobites are therefore exceptional in that, for the whole of their history, almost all representatives of the group had heavily mineralized dorsal
exoskeletons. This type of hardening is almost certainly a derived condition, because early members of the group (or perhaps a sister group) had
unmineralized cuticles. It is perhaps relevant to mention here that all arthropod cuticles show some basic similarities in their organic structures
and that hardening by incorporating minerals within the organic framework is just one of several developmental options employed by the group.

Trilobites have a long history, are often extremely abundantly represented, and have a world-wide distribution; their exoskeletons have high
fossilization potential. However, until recently, almost nothing was known about the fine structures of their cuticles, despite the best endeavors of
workers (e.g., McAllister and Brand, 1989) over the previous hundred years or so. Because the mineralized fabric of a trilobite cuticle is probably
a reflection of a delicate organic framework, only in exceptional conditions are sufficiently fine details preserved. Furthermore, special polishing,
etching, and examination techniques are needed to investigate such material. We were fortunate in recognizing the special preservation of the
cuticle of Ellipsocephalus from the Cambrian of Sweden and being able to apply the correct preparative and investigative techniques to reveal
exceptional preservation in a laminated outer layer (LOL) of its cuticle (see Dalingwater et al., 1991). One of us (D. J. S.) had long suspected that
Tapinocalymene from the Silurian of the Welsh Borderland would reveal fine cuticular structure (Dalingwater et al., 1993a), because of his
awareness of the sedimentary conditions in which they were preserved (and we found similar preservation in the Silurian of Gotland). Despite
searching through many major collections in Britain and abroad, the only additional material we have discovered with fine structure preserved
in the LOL is a series of specimens of Homalonotus rhinotropis Angelin, 1854, from the upper Silurian of Skane, Sweden. We have already given a
brief description of this material and partially discussed its significance (Dalingwater et al., 1996); here we take the opportunity to add to our
description and to refine our discussion.

DISCUSSION

Subdivisions of the cuticle.-A prismatic outer layer has for some time been regarded as a significant original subdivision of the trilobite cuticle
along with a much thicker principal layer. However, we have here, and in previous papers, presented strong evidence that originally the outer
layer was finely laminate in many species [i.e., those few species from which we have described a laminated outer layer (LOL) and a much
greater number in which a prismatic outer layer, which must now be regarded as secondary, has probably replaced the LOL]. The LOL is
comparable with the pre-ecdysial exocuticle of extant decapod crustaceans; it was probably formed underneath the old cuticle prior to ecdysis and
calcified shortly after ecdysis.

"Shaped inclusions. "-The variety of laminated "shaped inclusions" described from the outer layer of the homalonotid cuticle are all considered to
have been produced by the amalgamation of spherical accumulations of material around the apices of the 3-6 (mu)m diameter perpendicular
canals. We have explained these spheroids as representing sites of early crystallization with a subtly different mineralogy to later phases and
therefore preferentially preserved or, in other examples (Ellipsocephalus, Calymene spp. from Gotland, and many of the structures termed
Osmolska cavities), preferentially destroyed. For a more detailed discussion, see Dalingwater et al. (1993a). We have not previously considered
the significance of the spheroidal shape. Early mineralization in the exocuticle of crabs takes the form of thin horizontal plaques, rather than
spherical accumulations, around the apices of bristle ducts. However "bun-shaped deposits" are formed in the outer endocuticle, again centered on
bristle ducts (Digby, 1967). Mineralization of cuticle comprises transporting mineralizing materials, essentially calcium and carbonate and/or
phosphate ions, from absorption sites, through the epidermis and into a preformed organic framework or template; initiating crystallization,
perhaps involving seeding and enzymatic catalysis; and completion of the process by amalgamation of the accumulations centered around the
initial sites of crystallization. If the 3-6 (mu)m canals in homalonotid cuticles, and similar-sized canals in other trilobite cuticles we have studied,
were involved not only as crystallization initiation centers but also in transporting one or more of the materials of the process, then there must
necessarily have been points of exit for these materials from the canals into the cuticle. If these exit points were situated more centrally in the

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LOL, then this might result in the spherical aspect of the initial areas of crystallization centered on the canals.

Principal layer.-The original structure of the principal layer remains elusive. There are at least four possibilities: nonlaminate, finely laminate,
coarsely laminate, and a mixture of laminate and nonlaminate.

The only arthropods to have regions of calcified cuticle with an almost entirely nonlaminate aspect are some ostracodes (carapaces) and some
cirripedes (plates and valves) (see Dalingwater and Mutvei (1990) for some details). But these structures are essentially rigid shells and the
trilobite cuticle, though relatively thick and heavily calcified, was a true exoskeleton, not a shell.

A combination of laminate and nonlaminate cuticle is found in some chelicerates-scorpions, limulids, and possibly eurypterids-and in some
decapod crustaceans (Mutvei, 1974, 1977). However, regions with nonlaminate, preferred orientation zones interspersed with laminate
nonpreferred orientation zones are usually only found in specialized parts of the exoskeleton subject to directional forces such as the tail-spine of
limulids.

Fine lamina units are characteristically found in the exocuticle of calcified arthropod cuticles, with relatively wide units towards the center of
the endocuticle and with these wide units flanked by somewhat narrower units. All units of uncalcified cuticles are generally very finely
laminated. Wide, rather coarse lamina units have been described from the center of the principal layer of a number of trilobite species; two
Devonian species show very well-defined units when viewed under reflected light, but did not reveal detail when examined with the SEM
(Dalingwater et al., 1993b). Some of the light micrographs figured by Teigler and Towe (1975) also show well-defined lamination, again
particularly in Devonian species.

Present evidence suggests, therefore, that the principal layer was laminate, with a central zone of wide lamina units flanked by outer and inner
zones of narrower lamina units. This tripartite zonation may produce some differentiation on recrystallization, which results in the slight
differences between the central and the outer and inner regions of the principal layer mentioned earlier in this paper.

Differences in the preservation of the principal layer and the LOL may be the result of a range of factors: differences in composition of the
mineralizing materials (perhaps there was originally more phosphate in the LOL); differences in the extent of mineralization, with the LOL less
heavily mineralized; and differences in the organic template with possibly, by analogy with decapod crustacean cuticle (see Dalingwater and
Mutvei, 1990), a reticulate arrangement of microfibres in the LOL and a microfibrillar arrangement in the principal layer. Such a difference in
fiber arrangement would affect the disposition of the mineral phase of the mineralized cuticle.

Notes on calcification.-The simulation of the early stages of mineralization in the LOL and possible roles for the pore canals and interprismatic
septa have already been discussed in our preliminary account of the homalonotid material (Dalingwater et al., 1996). We would like to add to
our concluding remarks in that paper the idea that one can examine mineralization on a number of levels of scale: from the intimate associations
of minerals with the cuticular microfibrils that can only be glimpsed at the highest magnifications of the transmission electron microscope to
aspects of crystallization that can be viewed with a hand-lens. Difficulties persist, even using Recent material, in reconciling observations made
at various levels of this spectrum and at different stages of the molt cycle.

Our final suggestion concerns the spheroids and other shapes described from the outer layer. In some crustaceans, mineralization of the new
cuticle actually starts before exuviation (see Dalingwater and Mutvei, 1990), though clearly this must take place in a way that does not hinder
the necessary expansion of the cuticle immediately after molting. The advantages to a trilobite of emerging with a partly mineralized exocuticle
(effectively studded with "mini-spicules")offering some degree of protection and speeding up the urgent progression to a fully formed, fully
mineralized exoskeleton-seem clear.

ACCEPTED 18 AUGUST 1998

REFERENCES

DALINGWATER, J. E., AND H. MUTVEI. 1990. Arthropod exoskeletons, p. 83-96. In J. G. Carter (ed.), Skeletal Biomineralization: Patterns,
Processes and Evolutionary Trends, Volume I. Van Nostrand Rheinhold, New York.

-, S. J. HUTCHINSON, H. MUTVEL, AND D. J. SIVETER. 1991. Cuticular ultrastucture of the trilobite Ellipsocephalus polytomus from the Middle
Cambrian of Oland, Sweden. Palaeontology, 34:205-217. AND -. 1993a. Cuticular ultrastructure of some Silurian calymenid trilobites from the
Welsh Borderland and Gotland. Palaeontographica, Abteilung A, 229:37-49, 10 pl. AND -. 1993b. Trilobite cuticle calcification, p. 13-20. In I.
Kobayashi, H. Mutvei, and A. Sahni (eds.), Structure, Formation and Evolution of Fossil Hard Tissues. Tokai University Press, Tokyo.

-, AND -. 1996. Mineralization of some homalonotid trilobite cuticles. Bulletin de l'Institut oceanographique, Monaco, nombre special, 14,
4:343-351.

DIGBY, P. S. B. 1967. Calcification and its mechanism in the shore crab, Carcinus maenus (L.). Proceedings of the Linnaean Society of London,
178:129-146.

MCALLISTER, J. E., AND U. BRAND. 1989. Primary and diagenetic microstructures in trilobites. Lethaia, 22:101-111.

MUTVEI, H. 1974. SEM studies on arthropod exoskeletons, Pt. 1, Decapod crustaceans, Homarus gammarus L. and Carcinus maenas (L.). Bulletin

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of the Geological Institutions of the University of Uppsala, New Series, 4:73-80.

-. 1977. SEM studies on arthropod exoskeletons, Pt. 2, Horseshoe crab Limulus polyphemus (L.) in comparison with extinct eurypterids and
Recent scorpions. Zoologica Scripta, 6:203-213.

TEIGLER, D. J., AND K. M. TowE. 1975. Microstructure and composition of the trilobite exoskeleton. Fossils and Strata, 4:137-149, 9 pl.
WHITTINGTON, H. B. 1993. Morphology, anatomy and habits of the Silurian homalonotid trilobite Trimerus. Memoirs of the Association of
Australasian Palaeontologists, 15:69-83.

JOHN E. DALINGWATER, DEREK J. SIVETER, AND HARRY MUTVEI 50 Westgate, Hale, Cheshire WA15 9AZ, U.K.,

Geological Collections, University Museum of Natural History, Parks Road, Oxford OX1 3PW, U.K., and Sektionen for Paleozoologi,
Naturhistoriska Riksmuseet, S-10005 Stockholm, Sweden

Copyright Paleontological Society Mar 1999

Provided by ProQuest Information and Learning Company. All rights Reserved

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