Exoskeletal structure in the Ordovician trilobite

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-
bites have been reviewed by Dalingwater (1973),
and therefore only studies after this date are
discussed below.
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
distinguished: an inner thick layer and an outer
thin layer. The inner layer was found to be
composed of calcite crystals which have their
c-axes oriented more or less perpendicular to the
exoskeletal surface. In six species this layer
showed fine lamellae. N o lamination was recog-
nized in the outer layer, which was found to be
prismatic in structure, pigmented, or composed
of apatite. In the remaining eight species, the
exoskeleton consisted of the inner layer only.
Dalingwater & Miller ( 1977) distinguished
three zones in the exoskeleton of the Lower
Ordovician (Kundan) trilobite Asaphus
(Asaphus) raniceps: (1) a narrow outer zone
comprising up to one-fifth of the total exoskelet-
al thickness and consisting of laminar units about
5 pm thick; (2) a wide central zone with four to
five laminar units 50-70 pm thick; and (3) a
narrow innermost zone with laminar units 5 pm
thick. The exoskeletal surface was found to be
covered occasionally by a very thin prismatic
layer. Three exoskeletal zones similar to those in
Asaphus were distinguished by St@rmer(1980)
in chasmopinid and phacopid trilobites. In this
material, the laminar units in the outer zone had
a thickness of about 5 pm, in the middle zone
20-30 pm, and in the inner zone about 10 pm.
Material and methods
In this paper the exoskeletal ultrastructure is
described in Flexicalymene sp. from the Upper
Ordovician Maquoketa Shale, Iowa, USA, and
compared with that in crustaceans and chelicer-
ates. In the material studied the state of pre-
servation of exoskeletal structures appears to
exceed that in any other trilobites in which these
structures have been studied previously. All
specimens described here were collected from
the lowermost Elgin coquina beds of the Ma-
quoketa Shale, at Graf, Iowa, USA. The coquina
consists largely of specimens of the cephalopod
Isorthoceras sociule (e.g. Tasch 1955; Flower
1962), but about 50 fragments of trilobite exo-
skeletons belonging to Flexicalymene sp. were
found by crushing about 5 kg of rock samples
into small pieces (0.5-1.0 cm3).
The exoskeleton was fractured vertically and
studied either unetched, or etched for three to
four hours with a chromium sulphate solution
(Mutvei 1972). The specimens were mounted on
specimen stubs, coated with evaporated gold,
and studied with the scanning electron micro-
226 Harry Mutvei
t
, , d show that the exocuticle has a horizontally
Fig. I . Fkxicdymrne sp. Block diagram of the exoskeleton.
Note difference in mineralization between exocuticle ( e m )and
endocuticle ( e n d o ) .b, endocuticular bulb; d , ducts; f , tuber-
cles.
scope. The original mineral composition of the
trilobite exoskeleton was mostly calcitic (Daling-
water 1973; Teigler & Towe 1975). However, in
the trilobites studied here the calcite has been
replaced by a hydrated phosphate (dahllite
according to Grandjean et al. 1964579) and
other minerals, as in the cephalopod shells from
the same locality, and this has facilitated pre-
servation of structures down to ultrastructural
level. There are clear indications that this re-
placement took place at an early diagenetic
stage.
Exoskeleton in Flexicalymene
Two layers can usually be distinguished on ver-
tical fracture planes of the exoskeleton. These
layers are here termed the exocuticle and the
endocuticle (exo, endo, Fig. l ) , in agreement
with the terminology adopted for structurally
similar layers in the exoskeleton of extant de-
capod crustaceans (e.g. Mutvei 1974). The ex-
ocuticle was probably covered by a thin epicuti-
cle which is not preserved.
Exocuticle. - The exocuticle forms the outer
layer of the exoskeleton, comprising about 20%
of the total exoskeletal thickness in most prepa-
rations (exo, Fig. 1). It is occasionally thinner,
probably because its outermost part has been
worn off.
Studies of untreated vertical fracture planes
LETHAIA 14(1981)
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-
connected by parabolically arched sheets of
fibres forming the inter-laminae (indicated by
arrows in Fig. 2B). In a preparation etched for
three hours with a chromium sulphate solution,
the organic fibres became wholly or partially
demineralized, and their diameter decreased
considerably to less than 0.1 pm (exo, Fig. 3, and
the separate fragment in the lower right corner of
the same Fig.). In most preparations, however,
the mineralized fibre-bundles have been dia-
genetically altered and form granulae, usually
less than 0.5 pm in diameter. The occurrence of
the horizontal laminar units is still indicated by a
more or less distinct horizontal arrangement of
the granulae.
The exocuticle of Flexicalyrnene sp. has the
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
the laminar units seem to be much thinner in
Flexicalymene. The calcified organic fibres and
their arrangement into laminae and inter-laminae
have not been observed in previous studies.
Endocuticle. - The boundary between the en-
docuticle and the exocuticle is distinct because
the former is always considerably more mineral-
ized than the latter. The endocuticle constitutes
the principal inner layer of the exoskeleton,
comprising about 80% of the total exoskeletal
thickness (endo, Fig. 1). On unetched vertical
fracture planes its structure appears to be clearly
different from that of the exocuticle. It is com-
posed of crystal blocks with a distinct cleavage
oriented obliquely to the exoskeletal surface.
These blocks have no distinct boundaries, and
their size and shape are highly variable. Their
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
organic laminae.
At the boundary with the exocuticle, the en-
docuticle forms numerous bulbs which protrude
outwards into the exocuticle (Fig. 1, 6, Fig. 2A).
Each bulb seems to contain a large duct which
opens into a tubercle on the exoskeletal surface
(Fig. 1 ) . Similar endocuticular bulbs occur in the
extant decapod crustacean Cancer pagurus
(Hegdahl et al. 1977a, Figs. 1-3).
O n the vertical fracture planes of the endocuti-
cle, etched for four hours with a chromium
sulphate solution, distinct horizontal organic
laminae are often observed (Figs. 4A, B). These
laminae extend through the crystal blocks. In a
preparation shown in Fig. 4A (endo), the hori-
zontal laminae seem to attain a maximum thick-
ness of 2 pm. These thick laminae are found in
the endocuticular bulbs around the large ducts,
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
are arranged regularly around smaller ducts (in-
dicated by arrows in Fig. 4B). These laminae,
which have a thickness less than 0.5 pm, usually
disappear about halfway between adjacent
ducts. Remains of thin horizontal laminae were
also found attached to the pore canal walls (Fig.
5B). In none of the endocuticular laminae can
the organic fibres be distinguished.
The endocuticle is traversed vertically by
ducts of various diameters. Ducts’of about 3-7
pm (Fig. I , indicated by arrows in Fig. 4B) and
15-20 pm (Fig. 1) in diameter were observed
most frequently. In the cephalon, the larger of
these ducts seem to extend through prominent
tubercles to the exoskeletal surface (Fig. 1; see
also Evitt & Whittington 1953; Stqirmer 1980).
In previous studies (e.g. Osmolska 1975; Teig-
ler & Towe 1975; Dalingwater & Miller 1977;
Stqirmer 1980), the pore canals in trilobites have
been observed only in light microscopic prepara-
tions, where their diameters have been estimated
to be 1-2 pm. In Flexicalymene sp., the pore
canak could not be observed on untreated ver-
tical fracture planes of the exoskeleton. Howev-
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
solution, pore canals with well preserved walls
were seen occasionally (Fig. SA, B). These ca-
nals have a diameter of about 0.3 p,m, and their
walls are not helical or twisted as those in some
extant crustaceans. A series of disc-like exten-
sions project horizontally from the wall of each
canal (Fig. 5B). These extensions are situated
about 0.2 pm from each other, and represent
remains of horizontal organic laminae of the
endocuticle (see above).
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)
exocuticle in being horizontally laminate (exo,
endo, Fig. 1). The organic laminar units are
mostly very thin and the endocuticle is consider-
ably more mineralized than the exocuticle.
Moulting. - Prior to moulting in arthropods,
much of the old exoskeleton is broken down
enzymatically. The products of digestion are
resorbed and re-used in the formation of the new
exoskeleton. Among crustaceans, only ostra-
codes seem not to resorb their heavily mineral-
ized carapaces before moulting (Turpen &
Angel1 1971). In extant decapod crustaceans, a
resorption from exuvia occurs generally, but its
total amount is variable (Passano 1960). For
example, in Palinurus the resorption before
moulting amounts to 23% of the organic matter
and 13% of the mineral matter, and in Carcinus
to 79% of the organic matter and 18% of the
mineral matter (Passano 1960). However, the
resorption seems not to effect the thickness of
the exuvia to any greater extent. The ultra-
structural changes due to resorption are not
known.
Among chelicerates, the extant Lirnulus and
probably also fossil eurypterids resorb practical-
ly all the endocuticle. The exuvia of Limulus are
therefore thin, forming about 20% of the original
exoskeletal thickness (Mutvei 1977). A similar
magnitude of resorption takes place in insect
exoskeletons before moulting (Wigglesworth
1959).
It is generally believed that most trilobite
exoskeletons in the fossil record represent ex-
uvia, and it is reasonable to assume that the
majority of Flexicalymene exoskeletons also
represent such moult stages. If resorption has
taken place, then the endocuticle in the trilobite
exuvia would be characterized by ( 1 ) a reduction
in thickness, and/or (2) a formation of pores and
cavities, particularly around ducts and pore can-
als, as a result of digestion of the organic matter
and dissolution of the mineral matter. However,
none of these characteristics could be observed
in the trilobite exoskeletons, including those of
Flexicalymene. It is therefore probable that be-
fore moulting, no resorption took place from the
heavily calcified exoskeleton, other than a
softening along the sutures where the exo-
skeleton split open. As in extant ostracodes,
trilobites were probably incapable of reusing
organic and mineral matter from the old exo-
skeleton during the formation of a new exo-
skeleton. With regard to its structure, a trilobite
E.xoskeleta1 structure in FLEXICAL
YMENE 23 1
exuvium cannot be distinguished therefore from
the exoskeleton of a dead animal (see also Hen-
ningsmoen 1975:181 ).
Comparisons with crustaceans and
chelicerates
Towe (1978:459) expressed doubts as to whether
‘trilobites have a typical arthropod cuticle’. He
pointed out that fine horizontal laminae,
observed sporadically in a limited number of
trilobites, ‘lack any evidence of microfibrils or
parabolic structures’, the latter structures refer-
ring to the inter-laminae.
As demonstrated here, both the exo- and
endocuticle in Flexicalymene sp. have a horizon-
tal, fine-laminate organic structure. Typical in-
ter-laminae composed of parabolic arcs of orga-
nic fibres occur in the exocuticle. These organic
fibres probably consisted of protein and chitin,
as similar fibres in extant arthropods (e.g. Hack-
man 1971). During diagenesis, the fibres lost
their chemical characteristics (Teigler & Towe
1975:139) but in places retained their shape and
arrangement. Consequently, the trilobite exo-
skeleton possesses a typical arthropod struc-
ture.
The exoskeleton in trilobites agrees with that
in extant crustaceans in that it is composed of
horizontal organic laminar units, and it is miner-
alized. In podocope ostracodes, the organic
fibres in the heavily mineralized carapaces
usually form a reticulate structure (Jgrgensen
1970; Bate & East 1975). However, a typical
organic lamination has been observed in less
mineralized juvenile carapaces of Podocopina
(Rosenfeld 1979, Figs. 8, 9) and in non-mineral-
ized carapaces of Myodocopina (Bate & East
1975).
Among chelicerates, adult Limulus (Xipho-
sura, Merostomata) has a thin, horizontally
laminate exocuticle and a thick, vertically lamin-
ate endocuticle. During ontogeny, the organic
endocuticular structure seems to change con-
siderably. Thus, in juvenile Limulus only the
marginal parts of the carapace are vertically
laminate (‘non-laminate’), whereas the remain-
ing parts are horizontally laminate (Dalingwater
1980). The exoskeleton in the fossil eurypterids
studied so far (Dalingwater 1973, 1980) has a
similar structure to that in adult Lirnufus: the
thin exocuticle is horizontally laminate and the
thick endocuticle vertically laminate (Mutvei
232 Harry Mutvei
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
pedipalps have a layer with a vertically laminate
structure (Mutvei 1977) corresponding to the
‘non-laminate zone’ of Dalingwater (1980). The
rest of the scorpion exoskeleton is horizontally
laminate but the inter-laminae are often conspi-
cuously thick (Mutvei 1977). In extant spiders
the exoskeleton is horizontally laminate (Barth
1969).
As seen in Fig. 6, a layer with vertically
laminate organic structure (vf) also occurs in the
distal end of the chela of the extant decapod
crustaceans Homarus gammarus and Nephrops
norvegicus. This layer replaces the outer part of
the endocuticle, whereas the inner part (endo)
has retained its normal, horizontally laminate
structure.
It is obvious that the development of the verti-
cally laminate organic structure does not have a
great taxonomic and phylogenetic significance
LETHAIA 14 (1981)
among arthropods. As pointed out previously
(Mutvei 1977), there is no fundamental differ-
ence between the vertically and horizontally
laminate structures, because the vertical laminae
seem to correspond to the parabolic fibrous
sheets in the inter-laminae. This condition is also
clearly indicated in the right lower corner of Fig.
6. In trilobite exoskeletons a vertically laminate
organic structure has not yet been observed.
As in crustaceans, the exoskeleton in trilobites
is mineralized. In the endocuticle, which is con-
siderably more mineralized than the exocuticle,
the c-axes of the calcite crystals are oriented
essentially perpendicular to the exoskeletal sur-
face (Teigler & Towe 1975). A similar orientation
of the c-axes is found in the heavily mineralized
exo- and endocuticles of ostracode carapaces
(e.g., Jgrgensen 1970). The high degree of miner-
alization of the exoskeletons in ostracodes and
trilobites explains their resistance against dia-
LETHAIA 14 (1981)
genetic changes (Bergstram 1979:29). ln the
less mineralized trilobite exocuticle, the original
crystallographic orientation is still unknown.
-
The exocuticle seems to aeree in its deeree of
mineralization to the exo- and endocuticles of
- Dalinewater, J. E. 1973: Trilobite cuticle microstructure and
extant decapod crustaceans. For example, in
Cancer pagurus small calcite crystals in the
exocuticle have a variable and irregular
arrangement, whereas in the endocuticle they
are Oriented main’y to the long Of
the protein-chitin fibres (Hegdahl et al. 1977a,
b). Travis (1970) did not find any-~ Dreferred
crystallographic orientation of small calcite crys-
tals in the exo- and endocuticles of the freshwa-
ter crayfish Orconectes ririlis.
The following two factors explain some of the
difficulties encountered in the study of trilobite
exoskeletons at an ultrastructural level: ( 1 ) Orga-
nic fibres and their arrangement into horizontal
laminae and inter-laminae can be expected to be
observed practically only in the exocuticle,
where mineralization was less extensive.
However, the exocuticle is comparatively thin
and is therefore easily worn off. In the heavily
mineralized endocuticle the organic laminae are
partially absent, and the organic fibres cannot be
distinguished. (2) The horizontal organic laminar
units in trilobite exo- and endocuticles are most-
ly extremely thin (0.2-0.5 pm). In extant arthro-
pods with equally thin laminar units, e.g., many
insects, the organic fibres and their arrangement
cannot be observed distinctly with SEM. The
latter condition can also be expected in trilo-
bites. The mineralized organic fibres of the lami-
nae and inter-laminae in Flexicalymene sp. could
therefore be observed only in some of the thick
laminar units which occur close to the endocu-
ticular boundary.
Acknowledgements. - This study was financially supported by
Grants 287-102 and 103 from the Swedish Natural Science
Research Council. I wish to thank Professor W. M . Furnish
and Professor 9. F. Glenister for supplying me with the
Maquoketa material and for guidance in field.
References
Barth, F. G. 1969: Die Feinstruktur des Spinneninteguments.
I. Die Cuticula des Laufbeins adulter hautungsferner Tiere
(Cupiennius s a k i Keys.). 2. Zellforsch. mikrosk. Anat. 97,
137- 159.
Bate, R. H. & East, 9. H . 1975: The ultrastructure of the
ostracode (Crustacea) integument. Bulls. A m . Puleont. 65,
529-547.
16 Lethaia 3/81
~
Exoskeletal structure in FLEXICAL YMENE 233
Bergstrom, J . 1979: Morphology of fossil arthropods as a guide
to phylogenetic relationships. I n Gupta, A.P. (ed.): Arthro-
pod Phvlogenv, 3-53, Van Nostrand Reinhold Co,, New
Yo&.
I
composition. Palaeontcjlogy 16, 827-839.
Dalingwater, J . E. 1975: Further observations on eurypterid
cuticles. Fossils and Struta 4 , 271-279.
Dalingwater, J . E. 1980: SEM observations on the cuticles of
some chelicerates. 8 . fnternationer Arachnologen-Kon-
gress, W i m 285-289.
Dalingwater. J . E. & Miller, J . 1977: The laminae and cuticular
organization of the trilobite Asaphus raniceps. Palaeontolo-
<pv20,
,* 21-32,
Evitt, W.R. & Whittington, H. B. 1953: The exoskeleton of
Nexiculymene (Trilobita). J . Paleontol. 27, 49-55.
Flower, R . H. 1962: Notes on the Michelinoceratida. N e w
Mexico Bureau ojMines, Mem. 10, 2 W 2 .
Grandjean, J . , Gregoire, Ch. & Lutts. A. 1964: On the mineral
components and the remnants of organic structures in shells
of fossil molluscs. Bull. Acad. Roy. Belg., Cl. Sci. 50,
562-595.
Hackman, R. H. 1971: The integument of Arthropoda. I n
Florkin, M. & Scheer, 9. T. (eds.): Chemical Zoology 4 ,
1-53. Academic Press, New York and London.
Hegdahl, T., Silness. J. & Gustavsen, F. 1977a:‘The structure
and mineralization of the carapace of the crab (Cuncer
pagurus L.). 1. The endocuticle. Zoologica Scripta 6, 89-99.
Hegdahl, T.. Gustavsen, F. & Silness. J . 1977b: The structure
and mineralization of the carapace of the crab (Cancer
pagurus L.). 2. The exocuticle. Zoologicu Scripta 6, 101-
105.
Henningsmoen, G. 1975: Moulting in trilobites. Fossils and
Struta 4 , 179-200.
Jdrgensen, N. 0. 1970: Ultrastructure of some ostracods. Bull.
Geoi. Soc. Denmurk 20, 79-92.
Mutvei, H. 1972: Ultrastructural studies on cephalopod shells.
Part 1: The septa and riphonal tube in Nautilus. Bull. Ceol.
Instn. Univ. Uppsalu N . S . 3 , 237-261.
Mutvei, H. 1974: SEM studies on arthropod exoskeletons.
Part 1: Decapod crustaceans, Homarus gammarus L. and
Curcinus maenas (L.). Bull. Geol. Instn. Univ. Uppsaln N .
S . 4, 73-80.
Mutvei. H. 1977: SEM studies on arthropod exoskeletons. 2.
Horseshoe crab Limulus polvphemus (L.) in comparison
with extinct eurypterids and recent scorpions. Zoologica
Scripta 6, 203-2 13.
Osmolska. Halszka. 1975: Fine morphological characters in
some Upper Palaeozoic trilobites. Fossils and Srratu 4 ,
201-207.
Passano, L. M. 1960: Molting and its control. I n Waterman, T.
H. (ed.): The Physiology ofcrustarea I , 473-536. Academic
Press, New York and London.
Rosenfeld, A. 1979: Structure and secretion of the carapace in
some living ostracodes. Lefhaia 12, 353-360.
Stbrmer. L. 1980: Sculpture and microstructure of the exo-
skeleton in chasmopinid and phacopid trilobites. Palaeon-
tology 23, 237-271.
Tasch, P. 1955: Paleoecologic observations on the orthoceratid
coquina beds of the Maquoketa at Graf, Iowa. J . Paleontol.
29, 51%518.
Teigler, D. J. & Towe, K. M. 1975: Microstructure and
composition of the trilobite exoskeleton. Fossils and Strata
4 , 137-149.
Towe, K . M. 1978: Do trilobites have a typical arthropod
cuticle? Palaeontology 21, 459461.
Travis, D. F. 1970: The comparative ultrastructure and orga-
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.