L.
I(V
Hydrobiologia 440: 249-260, 2000.
M.B. Jones, J.M.N. Azevedo, A.I Neto, A.C. Costa d; A.M. Frias Marrins (eds), Island, Ocean and Deep-Sea Biology.
249
O 2000 Kluwer Academic Publishers. Printed in the Netherlands.
Resting stages in a submarine canyon: a component of
shallow-deep-sea coupling?
Luigi Della ~omrnasa',Genuario ~elrnontel,Alberto palanques2, Pere puig2 &
Ferdinand0 B oero
'
' Dipartimento di Biologia, Stazione di Biologia Marina, CoNISMa, Universita degli Studi di Lecce,
73100 Lecce, Italy
E-mail: boero @unile.it
2~nstitutde Ci2ncies del Mar (CSIC), Plaqa del Mar s/n, 08039 Barcelona, Spain
Key words: submarine canyon, resting stages, shallow-deep-sea coupling, life cycle
Abstract
The ecological importance of resting stages in shallow waters is well known, but their presence in the deep sea is
practically unrecorded. Samples of sinking particles were collected from April 1993 to May 1994 in and around the
Foix Canyon (northwest Mediterranean Sea) using PPS3 sediment traps located between -600 m and -1 180 m.
Dead and viable organisms were collected, and inorganic empty shells constituted most of the biologically-derived
matter. Resting stages, considered as POM, had a flux of up to 70000 items m-2 d-l. They were the second
most abundant fraction of total POM after tintinnids (mainly represented by empty, chitinous loricas), and first
of the viable POM fraction. Most remained unidentified, but 58 morphotypes were referable to coastal species of
Dinophyta, Tintinnina and Calanoida. Resting stages were rare in samples collected from the open slope adjacent
to the canyon. These preliminary data suggest an important role of submarine canyons in concentrating POM and
transferring it from shallow to deep-sea habitats. Due to their resistance to degradation processes, resting stages are
probably the only POM component that can return to shallow areas by upwelling currents occurring in the canyon.
Introduction
The ecological importance of resting stages in neritic
areas is widely investigated (Matsuoka, 1985; Ishikawa & Taniguchi, 1994; Belmonte et al., 1995; Sonnemann & Hill, 1997; Montresor et al., 1998). Lindley
(1990) and Marcus (1995) stated that the presence of
resting eggs of Calanoida, Cladocera and Rotifera in
marine shelf sediments was inversely correlated with
distance from the coastline andlor depth. The importance of cysts in deep waters is mostly unknown. Only
Dale & Dale (1992) have recorded a flux of organicwalled and calcareous dinoflagellate cysts into the
deep sea at a rate of up to several thousands items
m-2 d-I. Lack of interest in the patterns and processes
related to cyst-mediated plankton-benthos dynamics is
due probably to the tendency to consider plankton ecology as limited to the water column (Boero et al., 1996;
Marcus & Boero, 1998). This is particularly true for
deep waters, where only biogeochemical cycles are re-
cognised as important components of benthic-pelagic
coupling.
In marine canyons, physical oceanographic models
suggest that water circulation is characterised by both
downwelling and upwelling events (Hickey, 1995;
Puig et al., 2000). Both hydrographic an satellite data
indicate a role of submarine canyons of! the northeast
Spanish coast in the modification of the surface flow
(Mas6 et al., 1990). Moreover, submarine canyons
play an important role in biological events and are
often underestimated areas of high species diversity.
They attract bottlenose whales and small odontocetes
(Gowans &Whitehead, 1995; Whitehead et al., 1997),
and host many species of decapods (Cartes & Sarda,
1993; Maynou et al., 1996), nematodes (Soetaert &
Heip, 1995), rmcronektonic crustaceans (MacquartMoulin & Patriti, 1996), fish assemblages (Stefanescu
et al., 1994) and hydromedusae (Gili et al., 1998,
1999).
2'25.
w
Figure 1. Foix Canyon topography and sediment trap locations
This work investigates the flow of resting stages
in a marine canyon, and discusses its possible role in
benthic-pelagic (and deep-shallow sea) coupling.
Materials and methods
camera. Resting stage (or 'cyst', according to Belmonte et a]., 1995) abundance was compared with that
of remaining biogenic particles (grouped according to
nine categories: silicoflagellates, dinoflagellates, foraminiferans, radiolarians, tintinnids, Oligotrichina,
metazoans, faecal pellets, others). The number of observed items was related with both collecting area
and time, obtaining the estimated flux (items mP2
d-'). All the biogenic material was separated qualitatively into two fractions: inorganic empty shells (of
silicoflagellates, radiolarians and foraminiferans) and
particulate organic matter (POM), to which cysts were
assigned.
Cyst description was based on morphological traits
(shape, size, surface features and content colour)
which were compared with available literature information. Unidentified morphotypes were roughly
separated, according to their size, into 'protistan' types
(diameter up to 40 pm), incertae sedis (diameter
from 41 to 70 pm) and 'metazoan' types (minimum
diameter 70 pm).
Sinking particles were collected in and around the
Foix submarine Canyon off the Catalan coast near
Results
Barcelona (Spain) (Fig. l ) , by a PPS3 sediment trap
system with a 0.125 m2collecting area for eac,h trap.
Collected particles were preserved automatically in
Among the five samples collected at the deepest
5% formalin solution to avoid organic matter degradacanyon station (Site l), the total flux of biogenic
tion. A rotatory collector, electronically controlled by
particles with a diameter >20 p m reached a maxa timerlmotor assembly, alternated six cups at intervals
imum (1 779 490 items mP2 d-') in spring 1993
of 15/16 days (Heussner et al., 1990). ~ r o m ~ ~ r i l 1 9 9 3 (sample STOlC11) and a minimum (632 990 items
to May 1994, sainples were collected at four sites,
m-2 d-') in summer (sample ST06C11) (Fig. 2) when
three inside the canyon and one in the adjacent open
the highest flux of cysts was recorded. In summer,
slope, at different depths and heights from the bottom
all four sampling sites combined (samples ST06C11,
(see Puig & Palanques, 1998 for further details).
ST04C12, ST06C13, ST06C14), the total flux of bioEight samples were chosen for this study (Table
genic particles (632 990 items mP2 d-') was highest
1): five (one for each season from April 1993 to May
at the deepest canyon site (sample ST06Cll) and min1994) from the deepest station in the canyon (Station
imum (211 258 items m-2 d-') at 500 m above the
canyon bottom (sample ST06C14).
1) and three (one from each other site) collected in
the summer, when resting stages were most abundant
The most abundant biogenic particles were silicoat Station 1. Samples were sonicated for 3 mins by
flagellates (recorded only as inorganic shells in all
an ultrasound device (Branson Sonifier 2200), sieved
samples: up to 1 528 050 items m-2 dK1 in STOlCll
sample, see Fig. 2). The shells of the silicoflagelat four different mesh sizes (250 p m , 125 p m , 63
p m and 20 pm), and diluted in 30 ml of 4% formlate DictyochaJibula were always the most abundant
items. Tintinnids were the second more abundant
alin solution. Two whole-sieved fractions (>250 p m
group, with a maximum (511 500 items mP2 d-')
and 250-125 p m ) and 10% aliquots of the others
recorded in winter (sample ST05C21). Resting stages
(125-63 p m and 63-20 p m ) were analysed for each
sample. Aliquot size was achieved by a preliminary
ranked third, with a peak (73 810 items m p 2 d-')
statistical analysis optimising subsampling according
in the summer (sample ST06C11) and a minimum
to Bros & Cowell (1987). Resting stage separation and
(2857 items mp2 d-') in the same season in the open
identification were camed out with an inverted micmslope adjacent to the canyon (sample ST04C12). The
remaining categories (dinoflagellates, foraminiferans,
scope (Zeiss Axiovert S 100) connected with a photo-
Tnble 1. General features of observed samples
Sample
Station
Sampling depth (m)
radiolarians, Oligotrichina, metazoans, faecal pellets,
and 'others')were a minority o f the isolated particles
(Fig. 2). As a whole, they ranged between 7536 items
m P 2 d t l in the open slope adjacent to the canyon
(summer: sample ST04C12) and 165 990 items m P 2
d-' (autumn: sample ST12C11), when a peak o f
faecal pellets occurred.
Most biogenic particles were inorganic empty
shells o f silicoflagellates, radiolarians and foraminiferans. In the canyon-adjacent open slope (sample
ST04C12), the organic component (POM) reached its
highest value (32.63% o f total flux) during autumn
(sample ST05C21), and lowest (2.6% o f total flux) in
summer (Fig. 3). In general, tintinnids (only 0.3% o f
this category in the form o f viable items, the remaining
as empty loricas) were the most abundant POM fraction, followed by cysts (only 23% o f them in the form
o f empty shells, the remaining as viable forms) (Fig.
4).
Cysts
1
Cysts were represented by 58 morphotypes: 35 protists, 12 Metazoa and 11 incertae sedis due to taxonomic uncertainty. Most protistan cysts were unidentified (19 morphotypes); 10 morphotypes (nine Dinophyta and one Tintinnina) were identified to genus
level and six to class level (Polyhymenophorea and
Chrysophyceae). Among the metazoan cysts, seven
morphotypes were calanoid copepods (the remaining
ones were unidentified).
Due to difficulties in identification o f the dinoflagellate Scrippsiella at species level on the basis
o f cyst morphology (Montresor et al., 1998), different morphotypes were considered under the group
Scrippsiella-like. All recognised resting stages, ex-
Distance from
the bottom (m)
Sampling period
cept the pelagic dinoflagellate Thoracosphaera albatrosiana, were coastal species and most have been
recorded from recent sediment samples (Matsuoka,
1988; Lewis, 1991; Sonneman & Hill, 1997; Belmonte, 1998;Montresor et al., 1998). Thoracosphaera
albatrosiana is known only as a cyst and is recorded
in great numbers from deep-sea sediment traps both
in the Pacific and the Atlantic Oceans (Dale & Dale,
1992).
The total flux o f protistan and incertae sedis cysts
was seven times higher than the total flux o f metazoan resting stages. The flux o f empty protistan cysts
was a seventh o f that o f viable ones, while the flux
o f empty metazoan cysts was eight times higher than
that o f viable ones. The most abundant cyst types were
T albatrosiana among protists, and 'Centropagiidae
egg l ' , among metazoa (see Appendix 1 for detailed
description of cyst morphologies).
Discussion
Puig & Palanques (1998)stated that the Foix Canyon
receives particles not only from the overlaying water
column, but also from the shelf waters, whereas on
the adjacent open slope, sinking particles come mainly
from the overlying water column, with a small contribution o f particles transferredfrom the shelf. Our data,
obtained from the flux o f biogenic sediment particles,
support this model (Puig & Palanques, 1998). The flux
o f POM recorded in the open slope (10 557 items m-2
d-I) was many times lower than the average (185 775
items m P 2d p l )o f all samples from inside the canyon.
This suggests that the Foix Canyon accumulates POM
enriched by organic biogenic sediment coming from
the continental shelf.
STOlCll ST06Cll STl2Cll ST05C21 ST12C21 S T W 1 2 ST06C13 ST06C14
Sample
Figure 2. Estimated flux (no items mP2 dC1) of the most abundant categories (see Table 1 for sample description).
POM is the main source of biomass and energy for
the deep sea; it includes large food falls, consisting of
animal carcasses, along with terrigenous and coastal
plant debris, as well as fine particulate organic matter mostly from planktonic animals, including faecal
pellets, moults, and phytoplankton (Gage & Tyler,
1991). Resting stages have not been considered previously as a POM component. In the Foix Canyon,
resting stages represent the second fine-POM fraction
(even the most abundant fraction in sample STO6Cl1)
and could play an important role in shallow-deep-sea
coupling. Boero (1994), Boero et al. (1996), Belmonte
& Rossi (1998) and Marcus & Boero (1998) suggested that cyst-mediated dynamics call for a re-appraisal
of the classical separation of plankton and benthos.
This has been stressed also by Pati et al. (1999) who
showed that cysts (never considered by meiobenthologists) constituted 52% of the meiobenthos of a coastal
lagoon in terms of number of individuals and 30% in
terms of biomass!
Marcus & Boero (1998) hypothesised that cysts
should represent a fundamental biological link, via
submarine canyons, in shelf-slope and shallow-deep
sea coupling. The preliminary data from the Foix
Canyon corroborate their suggestion. All the 23 identified morphotypes in the present study, with th;
exception of Thoracosphaera albatrosiana (see Dale
& Dale, 1992), belonged to coast3 taxa. Scrippsiella spp. and Calciodinellum operosum, recorded from
deep sea by Dale & Dale (1992), are dinoflagellate
cysts, typically recorded in coastal areas by several authors (Lewis, 1991; Belmonte et al., 1995; Montresor
et al., 1997, 1998). The occurrence of cysts of coastal
species in the canyon suggests transport from neritic
areas by downwelling phenomena. The flux of cysts
inside the canyon varied from 4050 items mP2 d-' to
73 810 items mP2 d-'. The lowest flux was recorded
in the canyon-adjacent slope (2857 items m-2 d-l).
In the Atlantic and Pacific Oceans, Dale & Dale
(1992) reported a flux of dinoflagellate cysts ranging
between 401 cysts m-2 d-' and 14 584 cysts m-2 dC1
from -389 m to -5582 m. In coastal areas, the sedimentation flux of resting stages is very variable. In the
Mar Piccolo of Taranto, Rubino et al. (1996) recorded a sinking rate of about 23 000 resting stages mP2
d-' (March 1996). In the Gulf of Naples, Montrtsor
et al. (1998) found an average cyst flux of 280000
cysts m-2 d-' . Therefore, the resting stage flux recorded in the Foix Canyon appears more similar to fluxes
recorded in coastal areas than to those of the deep sea.
The Foix Canyon, thus, acts as an accumulator
of resting stages. Further investigations are needed to
Inorganic
empty shells
POM
STOlCll STMCll STl2Cll STOSC2l ST12C21 ST04C12 ST06C13 ST06C14
Sample
Figure 3. Composition of biogenic seaments (see Table 1 for sample description).
Q Others
Bl Faecal pellets
Oligotrichina
klMetazoans
Fora@niferans
E l Dinoflagellates
Tintinnid shells
HViable tintinnids
Empty resting stages
Full resting stages
STOlCll ST06Cll STl2Cll ST05C21 ST12C21 ST04C12 ST06C13 ST06C14
Sample
Figure 4. Relative abundance of POM components (sce Table 1 for sample description).
Plore I. Observed resting stase morphotypes (see Appendix 1 for morphological descr.ip~ions).Scale har 20 prn, except 14 (50 p m )
P l o ~ e2. Ohserved resting stage n~orpho~ypcs
(see Appendix I for morphological descl-iptions). Scale bar: 20 /ImFigures 1-14: 50 /In1 Figures
15-10,
Plntr S. Observed resting stage morphotypes (see Appendix I for morphological descriptions). Scale bar: 50 W r n Figures 1-8, 16-19: 20 prn
Figurcs 9-15.
verify whether it refuels coastal planktonic populations via upwelling. During upwelling events, significant quantities of water coming from the deep bottom
are pumped out of the canyon walls toward the coasts
(Peffley & O'Brien, 1976; Klinck, 1995; Allen, 1996).
If the canyon head is located sufficiently close to the
coast, the canyon-upwelled water reaches the euphotic
zone, becoming readily available to the biota (Denman & Powell, 1984; Hickey, 1995). As suggested by
t
Marcus & Boero (1998), upwelled waters m ~ g haffect
coastal planktonic populations by not only supplying
dissolved nutrients, but also recruiting propagules in
the form of cysts. If this biological coupling between
shallow and deep sea is confirmed, the functioning
of coastal waters would be intimately linked with
that of offshore ones through canyon-driven propagule
circulation.
Many authors invoked the need to establish a complete model to better understand the functioning of
marine systems (Denman & Powell, 1984). Hickey
(1995) called for a synergy between physical and biological oceanography to li'nk topographical features or
physical mechanism to biological effects. On the other
hand, Boero et al. (1996) and Marcus & Boero (1998)
proposed biological cycles as a necessary complement
to biogeochemical cycles. The study of resting stage
dynamics in submarine canyons might be a further
step in this direction.
Acknowledgements
We thank J. M. Gili (Institut de Cikncies del Mar,
CSIC, Barcelona) for his stimulating encouragement
to start the study of submarine canyon sediments. This
work partly fulfils the requirements for the PhD training of L. Della Tommasa. Funds from the programme
"Biodiversid e interazioni nelle cornunit8 marine"
(COFIN) of the Ministero dell'Universit8 e Ricerca
Scientifica e Tecnologica. This research was supported by project AMB92-0251-C02-01 funded by the
Comisidn Interministerial de Ciencia y Tecnologia and
by projects MAS2-CT93-0053 andMAS3-CT95-0037
funded by the EEC.
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Appendix I. Cyst morphological descriptions
Plate 1
Dinophyta cysts
1.1.
Calciodinellum operosum Deflandre. Diameter 40 gm. spherical, wall calcareous, surface crested with a clear
paratabulation on the outer layer, brown. References: Dale & Dale (1992: 25, P1. 1.1 Figure 13); Montresor et al.
(1997); Montresor et al. (1998: 2297, Figure 20.
1.2.
cf. Diplopelfa parva (AbC, 1941) Matsuoka. Synonym: Dissodium parvum AbC. Diameter 32 pm,
processes 6 pm, spherical, with many short spines, brownish. References: Bolch & Hallegraeff (1990: 183, Figures 3 1 a x ) ;
Matsuoka (1988: 102, P1. 1 Figures G-K).
1.3.
Lingulodinium polyedrum (Stein) Dodge. Paleontological taxon: Lingulodinium machaerophorum (Deflandre
& Cookson) Wall. Synonym: Gonyaulaxpolyedra Stein. Diameter 26 pm, processes &8 pm, spherical, with
both bent and upright long conical spines, brown. References: Wall &Dale (1968: 294, P1. 1 Figure 18); Kokinos
& Anderson (1995); Lewis & Hallett (1997).
1.4.
Penfapharsodinium tyrrhenicum (Balech) Montresor et al. Paleontological taxon: Calcicarpinum bivalvum
Versteegh. Synonym: Peridinium tyrrhenicum Balech. Diameter 30 pm, wall calcareous, with distinctive ridges
conferring a broadly triangular-shape, brown. References: Montresor et al. (1993: 223, Figures 1 and 4); Sonneman &
Hi11 (1997: 157, Figures 12a-b); Montresor al. (1.898: 2296, Figure 2d).
Appendix 1. contd.
1.5.
1.8.
1.9
Protoperidinium compressum (AbC) Balech. Paleontological taxon: Stelladinium reidii Bradford; Stelladinium
stellatum (Wall) Reid. Synonym: Peridinium compressum Abt; fl stellaturn Wall & Dale. Length 20 p m , width
16 pm, processes 1&14 pm, stellate, with one apical, two antapical and two lateral processes, light brown.
References: Bolch & Hallegraeff (1990: 181, Figure 22); Sonneman & Hi11 (1997: 163, Figure 19).
Protoperidinium cf. nudum (Meunier) Balech. Paleontological taxon: Selenopemphix quanta (Bradford)
Matsuoka. Synonym: Peridinium cf. nudum (Meunier) Wall & Dale. Length 32 pm, width 28 pm, spines 6-8
pm, oval, with many conical spines, brown. References: Wall & Dale (1968: 301, PI. 4 Figure 5); Sonneman &
Hi11 (1997: 163-164, Figures 22a-b).
Protoperidinium oblongum (Aurillius) Parke & Dodge. Paleontological taxon: Votadinium calvum Reid.
Synonym: Peridinium divergens Ehrenb. var. oblongum Aurivillius; P oblongum (Aurivillius) Lebours;
P. oceanicum Schiller. Length 60 pm, width 40 pm, peridinoid, dorso-ventrally compressed, with rounded apices
and without spines, light brown. References: Wall & Dale (1968: 295, P1. 1 Figure 23); Bolch & Hallegraeff
(1990: 181, Figure 19a); Sonnneman & Hi11 (1997: 164, Figure 26a).
Scrippsiella-like. Length 25 pm-38 pm, width 22-31 pm, spherical or oval, calcareous, thickly provided with
short squat or conical spines, body red to brown. References: Lewis, (1991); Montresor et al. (1998).
Thoracosphaera albatrosiana Kampaner. Diameter 25 pm, spherical, calcareous, surface smooth, archeopyl
shaped as described by Dale & Dale (1992), brown. References: Dale & Dale (1992: 25, P1. 1.1 Figure 14).
Chrysophycean cysts
1.10.
Stomatocyst type I . Diameter 20 pm, spherical, surface smooth, wall thick, apical pore with cylindrical collar.
Similar to stomatocyst 15 1 Zeeb & Smol. Reference: Duff et al. (1995: 56, Figure 43).
Tintinnid cysts
1.11.
cf. Fusopsis sp.. Length 20 p m , width 18 pm, flask-shaped, surface smooth, apically-capped, antapically-tailed,
greenish. Reference: Reid &John (1978: 552, Figure If).
Polyhymenophorea (Protozoa, Ciliophora) flask-shaped cysts
1.12.
Type; Cil unl. Length 28 pm, width 25 p m , envelope thickness 3 pm, flask-shaped, surface smooth, external
envelope gelatinous, neck conical with a large pore, light brown.
1.13
Type: Cil un2. Length 44 pm, width 30 pm, flask-shaped, surface smooth, protoplasm granulous, apical process,
brownish.
1.14.
Type: Cil un3. Length 125 p m , width 78 pm, flask-shaped, surface smooth, wall thick, brown.
1.15.
Type: Cil un4. Length 38-40 pm, width 25 pm, flask-shaped, surface smooth, neck conical with a large pore,
greenish.
1.16.
Type: Cil un5. Length 50 pm, width 34 p m , flask-shaped, surface smooth, wall double, apical prominence,
brownish.
Unidentified protistan cysts
Type: Prot unl. Diameter 28-47 pm, spherical, surface smooth, wall thin, protoplasm granulous, greenish or
1.17.
.
.
brownish.
1.18
Type: Prot un2. Diapeter 25-34 p m , spherical, surface smooth, wall thin, with gelatinous crown, yellowish.
1.19.
Type: Prot un3, Diameter 14 pm, enyelope 8 p m thick, spherical, surface smooth, wall thin, with thick
gelatinous crown, greenish.
1.20.
Type: Prot un4. Diameter 22 pm, envelope 2-10 p m thick, spherical, surface smooth, wall thin, with gelatinous
crown and apical pore, greenish.
Plate 2
2.1.
2.2.
2.3.
2.4.
2.5
Type: Prot un5. Diameter 33 pm, spherical surface smooth, wall thin, with irregular mucilaginous layer, brown.
Type: Prot un6. Diameter 33 pm, spherical, surface smooth, wall thin, protoplasm with lipidic masses, greenish
Type: Prot un7. Diameter 18 p m , spherical, surface smooth, wall thin, with mucilaginous crown, protoplasm
with lipidic masses, yellowish.
Type: Prot un8. Diameter 20 pm, envelope 14 p m thick, spherical, surface smooth, wall thin, wrapped by
mucilaginous crown, brownish.
Type: Prot un9. Diameter 22 pm, spherical, surface smooth, wall thick, protoplasm granulous, light brown.
Appendix 1. contd
Type: Prot unlO. Diameter 25-38 p m , spherical, surface smooth, wall thick, with apical pore, brownish.
Type: Prot un 1 1. Diameter 32 p m , spherical, surface smooth, wall thick, protoplasm granulous, brown.
Type: Prot un12. Diameter 25-37 p m , spherical, surface smooth, wall thick, with apical pore, yellowish.
Type: Prot un13. Diamcter 32-38 p m , spherical, surface smooth, wall thick, black.
Type: Prot un14. Diameter 28 p m , gelatinous envelope 5 p m thick, spherical, surface smooth, wall thick, with
gelatinous crown, outer mucilage, yellowish.
Type: h o t un15. Diameter 35 p m , spherical, with many short and pointed processes, wall thick, red body in the
protoplasm, brownish.
Type: Prot un16. Diameter 25 p m ; spines 3 p m , spherical, with many short and thin spines, yellowish.
Type: Prot un17. Length 34 -55 p m , width 3 W 8 p m , oval, cyst, surface smooth, wall thin, with apical duct and
lipid assemblages in the protoplasm, yellowish.
Type: Prot un18. Length 32-52 pm, width 25-34 p m , oval, surface smooth, wall thick, with apical prominence,
yellowish.
Type: Prot un19. Length 38 p m , width 27 pm, envelope 32 p m thick, elliptical, surface smooth, wall thin,
wrapped by mucilaginous crown, brownish.
Metazoan cysts
2.16.
2.17.
2.18.
2.19.
Calanoida
Calanoida
Calanoida
Calanoida
eggl. Diameter 78-88 p m , spherical, with many short spines, empty.
egg2. Diameter 125 pm, spines 1 6 2 0 p m , spherical, with many conical spines, black.
egg3. Diameter 125 p m , spines 5 p m , spherical, with many short spines, brown.
egg4. Diameter 160 p m , spines 5-8 p m , spherical, with many short and thin spines, brownish.
Plate 3
3.1.
3.2.
3.3.
Acartiidae eggl. Diameter 58 p n , spines 6-10 p m , spherical, with many pointed and branched conical spines,
light brown.
Centropagiidae egg 1. Diameter 48 p m , spines 15-20 pm, spherical, with long, pointed and branched conical
spines, light brown.
Centropagiidae egg 2. Diameter 60 p m , processes 16 pm, spherical, with long, thin and bent spines, light brown
Unidentified metazoan resting stages
3.4.
Type: Egg unl. Diameter 90 pm, cap-shaped, surface smooth, protoplasm with lipidic masses, brown.
3.5.
Type: Egg un3. Length 125 pm, width 88 p m , oval, surface smooth, wall thin, with gelatinous crown and apical
pore, brown.
3.6.
Type: Egg un4. Diameter 82 pm, spherical, surface rough, with many squat processes, light brown.
3.7.
Type: Egg un5. length 90 p m , width 65 pm, oval, surface smooth, wall thin, with apical pore, brown.
3.8.
Type: Egg un6. Length 85 pm, width 58 p m , oval, surface smooth, wall thin, with apical duct, brownish.
Incertae sedis
3.9.
3.10.
3.1 1
Type: Inc unl. Diameter 45 p m , spherical, surface smooth, wall thick, yellowish.
Type: Inc un2. Diameter 55 pm, spherical, surface smooth, wall thick, with amphipolar prominences, brown.
Type: Inc un3. Diameter 52 p m , spherical, surface smooth, wall double, protoplasm with lipidic masses,
brownish.
Type: Inc un4. Diameter 55 p m , spherical, surface irregular, wall double, light brown.
Type: Inc un5. Diameter 47 pm, spherical, with short pointed processes, greenish.
Type: Inc un6. Diameter 52 p m , spherical, surface irregular, wall double, brown.
Type: Inc un7. Length 40-65 p m , width 34-56 p m , round-oval, surface smooth, wall thin, with apical pore, light
brown.
Type: Inc un8. Length 70 p m , width 48 p m , oval, surface smooth, wall double, with apical pore, protoplasm
with lipidic masses, brown.
Type: Inc un9. Length 55 p m , width 36 p m , elliptical, surface smooth, wall thin, protoplasm granulous,
greenish.
Type: Inc unlO. Diameter 65 p m , spherical, surface smooth, wall thin, with irregular gelatinous crown, brown.
Type: Inc unl 1. Diameter 48 p m , spherical, surface smooth, wall double, with septcd gelatinous crown,
protoplasm with lipidic masses, brownish.