Beruflich Dokumente
Kultur Dokumente
Flexicalymene
HARRY MUTVEI
Mutvei. Harry 1981 10 01: Exoskeletal structure in the Ordovician trilobite Flexicalymene. Lethaia.
LETHAIA Vol. 14. pp. 225-234. Oslo. ISSN 0024-1164.
Scanning Electron Microscope study of the exoskeletal ultrastructure of secondarily phosphatized
material of Nexicalymene sp. from the Upper Ordovician Maquoketa Shale, Iowa, USA, shows that the
exocuticle, comprising 20% of the total exoskeletal thickness, is composed of horizontal laminar units
between 0.2 and I pm thick. These units consist of primarily mineralized organic fibres which form
horizontal laminae interconnected by inter-laminae. The endocuticle is considerably more mineralizer
than the exocuticle, and its original organic structure cannot be observed in untreated preparations.
Etching with chromium sulphate reveals: ( 1 ) horizontal organic laminar units, 0.2 to 2 pm thick, and (2)
pore canals with non-twisted walls about 0.3 pm in diameter. Exuvia cannot be distinguished from the
exoskeletons of dead animals. The exoskeletal ultrastructure in trilobites agrees essentially with that in
crustaceans. 0 Trilobita, Flexicalymene, exoskeleton, cuticle, ultrastructure, Ordovician.
Harry Mutvei, Department of Palueozoology, Swedish Museum of Natural History, Box50007, S-104 05
Stockholm. and Palaeontological Institute, Box 558, S-75 122 Uppsala. Sweden: 28th January, 1981.
Previous studies on exoskeletal structure in trilo- in chasmopinid and phacopid trilobites. In this
bites have been reviewed by Dalingwater (1973), material, the laminar units in the outer zone had
and therefore only studies after this date are a thickness of about 5 pm, in the middle zone
discussed below. 20-30 pm, and in the inner zone about 10 pm.
Teigler & Towe ( 1975) studied exoskeletal
structure and composition in twenty trilobite
species ranging in age from Cambrian through
Devonian. In twelve species two layers were
Material and methods
distinguished: an inner thick layer and an outer In this paper the exoskeletal ultrastructure is
thin layer. The inner layer was found to be described in Flexicalymene sp. from the Upper
composed of calcite crystals which have their Ordovician Maquoketa Shale, Iowa, USA, and
c-axes oriented more or less perpendicular to the compared with that in crustaceans and chelicer-
exoskeletal surface. In six species this layer ates. In the material studied the state of pre-
showed fine lamellae. N o lamination was recog- servation of exoskeletal structures appears to
nized in the outer layer, which was found to be exceed that in any other trilobites in which these
prismatic in structure, pigmented, or composed structures have been studied previously. All
of apatite. In the remaining eight species, the specimens described here were collected from
exoskeleton consisted of the inner layer only. the lowermost Elgin coquina beds of the Ma-
Dalingwater & Miller ( 1977) distinguished quoketa Shale, at Graf, Iowa, USA. The coquina
three zones in the exoskeleton of the Lower consists largely of specimens of the cephalopod
Ordovician (Kundan) trilobite Asaphus Isorthoceras sociule (e.g. Tasch 1955; Flower
(Asaphus) raniceps: (1) a narrow outer zone 1962), but about 50 fragments of trilobite exo-
comprising up to one-fifth of the total exoskelet- skeletons belonging to Flexicalymene sp. were
al thickness and consisting of laminar units about found by crushing about 5 kg of rock samples
5 pm thick; (2) a wide central zone with four to into small pieces (0.5-1.0 cm3).
five laminar units 50-70 pm thick; and (3) a The exoskeleton was fractured vertically and
narrow innermost zone with laminar units 5 pm studied either unetched, or etched for three to
thick. The exoskeletal surface was found to be four hours with a chromium sulphate solution
covered occasionally by a very thin prismatic (Mutvei 1972). The specimens were mounted on
layer. Three exoskeletal zones similar to those in specimen stubs, coated with evaporated gold,
Asaphus were distinguished by St@rmer(1980) and studied with the scanning electron micro-
226 Harry Mutvei LETHAIA 14(1981)
t
, , d show that the exocuticle has a horizontally
laminate organic structure (4x0, Figs. I , ?A, B).
The total number of laminar units seems to vary
between 30 and 40 (Fig. 2A); the individual units
are usually 0.2-0.5 pm thick. Close to the en-
docuticular boundary the laminar units may
attain a thickness of about 1 pm (Figs. 2A, B).
The laminar units consist of organic fibres
which are primarily mineralized and probably
aggregated to form bundles. These fibre-bundles
are distinctly visible only in some preparations
close to the endocuticular boundary, where the
laminar units increase in thickness. In untreated
preparations their diameter is about 0.2 pm (Fig.
2B). Sheets of these fibre-bundles form consecu-
tive horizontal laminae (Fig. 2B) which are inter-
Fig. I . Fkxicdymrne sp. Block diagram of the exoskeleton.
Note difference in mineralization between exocuticle ( e m )and connected by parabolically arched sheets of
endocuticle ( e n d o ) .b, endocuticular bulb; d , ducts; f , tuber- fibres forming the inter-laminae (indicated by
cles. arrows in Fig. 2B). In a preparation etched for
three hours with a chromium sulphate solution,
the organic fibres became wholly or partially
scope. The original mineral composition of the demineralized, and their diameter decreased
trilobite exoskeleton was mostly calcitic (Daling- considerably to less than 0.1 pm (exo, Fig. 3, and
water 1973; Teigler & Towe 1975). However, in the separate fragment in the lower right corner of
the trilobites studied here the calcite has been the same Fig.). In most preparations, however,
replaced by a hydrated phosphate (dahllite the mineralized fibre-bundles have been dia-
according to Grandjean et al. 1964579) and genetically altered and form granulae, usually
other minerals, as in the cephalopod shells from less than 0.5 pm in diameter. The occurrence of
the same locality, and this has facilitated pre- the horizontal laminar units is still indicated by a
servation of structures down to ultrastructural more or less distinct horizontal arrangement of
level. There are clear indications that this re- the granulae.
placement took place at an early diagenetic The exocuticle of Flexicalyrnene sp. has the
stage. same thickness as that in Asaphus raniceps
(Dalingwater & Miller 1977, PI. 9: 1.2 and Fig. 2).
In both trilobites it is horizontally laminate, but
Exoskeleton in Flexicalymene the laminar units seem to be much thinner in
Flexicalymene. The calcified organic fibres and
Two layers can usually be distinguished on ver- their arrangement into laminae and inter-laminae
tical fracture planes of the exoskeleton. These have not been observed in previous studies.
layers are here termed the exocuticle and the
endocuticle (exo, endo, Fig. l ) , in agreement Endocuticle. - The boundary between the en-
with the terminology adopted for structurally docuticle and the exocuticle is distinct because
similar layers in the exoskeleton of extant de- the former is always considerably more mineral-
capod crustaceans (e.g. Mutvei 1974). The ex- ized than the latter. The endocuticle constitutes
ocuticle was probably covered by a thin epicuti- the principal inner layer of the exoskeleton,
cle which is not preserved. comprising about 80% of the total exoskeletal
thickness (endo, Fig. 1). On unetched vertical
Exocuticle. - The exocuticle forms the outer fracture planes its structure appears to be clearly
layer of the exoskeleton, comprising about 20% different from that of the exocuticle. It is com-
of the total exoskeletal thickness in most prepa- posed of crystal blocks with a distinct cleavage
rations (exo, Fig. 1). It is occasionally thinner, oriented obliquely to the exoskeletal surface.
probably because its outermost part has been These blocks have no distinct boundaries, and
worn off. their size and shape are highly variable. Their
Studies of untreated vertical fracture planes diameter usually ranges between 50 and 100 pm.
LETHAIA 14 (1981) Exoskeletal structure in FLEXICALYMENE
227
Fig. 2. Flexiculymene sp. 0 A . Untreated exoskeletal surface and vertical fracture plane within the glabella. Note distinct
horizontal lamina of the exocuticle (exo) at the boundary with an endocuticular bulb ( b ) .0 B. Untreated vertical fracture plane of
the exocuticle, showing bundles of calcified organic fibres in the horizontal laminar units. Fibre-bundles forming parabolic arcs
in the inter-laminae are indicated by arrows.
Unetched endocuticle lacks distinct horizontal O n the vertical fracture planes of the endocuti-
organic laminae. cle, etched for four hours with a chromium
At the boundary with the exocuticle, the en- sulphate solution, distinct horizontal organic
docuticle forms numerous bulbs which protrude laminae are often observed (Figs. 4A, B). These
outwards into the exocuticle (Fig. 1, 6, Fig. 2A). laminae extend through the crystal blocks. In a
Each bulb seems to contain a large duct which preparation shown in Fig. 4A (endo), the hori-
opens into a tubercle on the exoskeletal surface zontal laminae seem to attain a maximum thick-
(Fig. 1 ) . Similar endocuticular bulbs occur in the ness of 2 pm. These thick laminae are found in
extant decapod crustacean Cancer pagurus the endocuticular bulbs around the large ducts,
(Hegdahl et al. 1977a, Figs. 1-3). where they curve strongly outwards. In another
228 Harry Mutvei LETHAIA 14 (1981)
Fig. 3 . Flexicalymene sp. Vertical fracture plane of the exocuticle (exo) etched for three hours with chromium sulphate solu-
tion. Note partially decalcified organic fibres in the inter-laminae, close to the endocuticular (endo)boundary, and on a separate
fragment in the lower right comer.
preparation (Fig. 4B), thin horizontal laminae most frequently. In the cephalon, the larger of
are arranged regularly around smaller ducts (in- these ducts seem to extend through prominent
dicated by arrows in Fig. 4B). These laminae, tubercles to the exoskeletal surface (Fig. 1; see
which have a thickness less than 0.5 pm, usually also Evitt & Whittington 1953; Stqirmer 1980).
disappear about halfway between adjacent In previous studies (e.g. Osmolska 1975; Teig-
ducts. Remains of thin horizontal laminae were ler & Towe 1975; Dalingwater & Miller 1977;
also found attached to the pore canal walls (Fig. Stqirmer 1980), the pore canals in trilobites have
5B). In none of the endocuticular laminae can been observed only in light microscopic prepara-
the organic fibres be distinguished. tions, where their diameters have been estimated
The endocuticle is traversed vertically by to be 1-2 pm. In Flexicalymene sp., the pore
ducts of various diameters. Ducts’of about 3-7 canak could not be observed on untreated ver-
pm (Fig. I , indicated by arrows in Fig. 4B) and tical fracture planes of the exoskeleton. Howev-
15-20 pm (Fig. 1) in diameter were observed er, on the vertical fracture planes that were
LETHAIA 14 (I98 I ) YMENE 229
Exoskeletal structure in FLEXICAL
Fig. 4 . Flexicalymene sp. Vertical fracture planes of the endocuticle (endo) etched for four hours with chromium sulphate
solution. 0 A. Horizontal organic lamellae curve strongly outwards in the bulbs. Between the bulbs they are absent. 0B. Thin
horizontal organic lamellae occur around the ducts.
etched for four hours with a chromium sulphate sions project horizontally from the wall of each
solution, pore canals with well preserved walls canal (Fig. 5B). These extensions are situated
were seen occasionally (Fig. SA, B). These ca- about 0.2 pm from each other, and represent
nals have a diameter of about 0.3 p,m, and their remains of horizontal organic laminae of the
walls are not helical or twisted as those in some endocuticle (see above).
extant crustaceans. A series of disc-like exten- In summary, the endocuticle is similar to the
230 Harry Mutvei LETHAIA 14(1981)
Fig. 5. Flexicalymene sp. Pore canals on a vertical fracture plane etched for four hours with chromium sulphate solution. Note the
horizontal disc-like extensions from the pdre canal walls.These extensions are remains of horizontal organic lamellae.
LETHAIA 14 (1981) E.xoskeleta1 structure in FLEXICAL
YMENE 23 1
exocuticle in being horizontally laminate (exo, exuvium cannot be distinguished therefore from
endo, Fig. 1). The organic laminar units are the exoskeleton of a dead animal (see also Hen-
mostly very thin and the endocuticle is consider- ningsmoen 1975:181 ).
ably more mineralized than the exocuticle.
Fig. 6 . Nephrops norvegicus. Untreated vertical fracture plane of the exoskeleton in distal end of the chela showing structure of
the exocuticle (em), vertically laminate layer ( v n , and endocuticle (endo).
1977). In extant scorpions, the distal ends of the among arthropods. As pointed out previously
pedipalps have a layer with a vertically laminate (Mutvei 1977), there is no fundamental differ-
structure (Mutvei 1977) corresponding to the ence between the vertically and horizontally
‘non-laminate zone’ of Dalingwater (1980). The laminate structures, because the vertical laminae
rest of the scorpion exoskeleton is horizontally seem to correspond to the parabolic fibrous
laminate but the inter-laminae are often conspi- sheets in the inter-laminae. This condition is also
cuously thick (Mutvei 1977). In extant spiders clearly indicated in the right lower corner of Fig.
the exoskeleton is horizontally laminate (Barth 6. In trilobite exoskeletons a vertically laminate
1969). organic structure has not yet been observed.
As seen in Fig. 6, a layer with vertically As in crustaceans, the exoskeleton in trilobites
laminate organic structure (vf) also occurs in the is mineralized. In the endocuticle, which is con-
distal end of the chela of the extant decapod siderably more mineralized than the exocuticle,
crustaceans Homarus gammarus and Nephrops the c-axes of the calcite crystals are oriented
norvegicus. This layer replaces the outer part of essentially perpendicular to the exoskeletal sur-
the endocuticle, whereas the inner part (endo) face (Teigler & Towe 1975). A similar orientation
has retained its normal, horizontally laminate of the c-axes is found in the heavily mineralized
structure. exo- and endocuticles of ostracode carapaces
It is obvious that the development of the verti- (e.g., Jgrgensen 1970). The high degree of miner-
cally laminate organic structure does not have a alization of the exoskeletons in ostracodes and
great taxonomic and phylogenetic significance trilobites explains their resistance against dia-
LETHAIA 14 (1981) Exoskeletal structure in FLEXICAL YMENE 233
genetic changes (Bergstram 1979:29). ln the Bergstrom, J . 1979: Morphology of fossil arthropods as a guide
to phylogenetic relationships. I n Gupta, A.P. (ed.): Arthro-
less mineralized trilobite exocuticle, the original pod Phvlogenv, 3-53, Van Nostrand Reinhold Co,, New
crystallographic orientation is still unknown. Yo&.
-
The exocuticle seems to aeree in its deeree of
mineralization to the exo- and endocuticles of
- Dalinewater, J. E. 1973: Trilobite cuticle microstructure and
I
16 Lethaia 3/81
~
234 Harry Mutvei LETHAIA 14 (198 I )
nization of five calcified tissues. I n Schraer, H . (ed.): Wigglesworth, V . B. 1959: The Control of Growrh undForm: a
Biological Calcificurion: Cellular and Molecular Aspects, Study of the Epidermal Cell in an Insect. 140 pp. Cornell
203-31 1. North-Holland Publishing Company, Amsterdam. University Press, Ithaca, New York.
Turpen, J. B. & Angell, R. W. 1971: Aspects of molting and
calcification in the ostracod Heterocypris. B i d . Bull. 140,
33 1-338.