Fezouata Formation Explained

Fezouata Formation
Type:Geological formation
Age:[1]
Period:Floian
Prilithology:Shale
Region:Drâa-Tafilalet
Country: Morocco
Underlies:Zini Formation
Thickness:>1000m (3,000feet), combined for upper and lower formations, the two exceptionally preserved intervals ~25m (82feet) and 15m (49feet) respectively
Coordinates:30.6°N -5.7°W
Paleocoordinates:-73.9°N 108.1°W

The Fezouata Formation or Fezouata Shale is a geological formation in Morocco which dates to the Early Ordovician.[2] [3] [4] [5] It was deposited in a marine environment, and is known for its exceptionally preserved fossils, filling an important preservational window beyond the earlier and more common Cambrian Burgess shale-type deposits.[6] The fauna of this geological unit is often described as the Fezouata biota, and the particular strata within the formation which exhibit exceptional preservation are generally termed the Fezouata Lagerstätte.

Biota

Over 1,500 non-mineralized specimens, representing 50 distinct taxa that have a composition similar to earlier Burgess Shale type biotas, have been recovered from the formations in addition to a less abundant shelly fauna.[6] The make-up of the community varies significantly through the stratigraphic sequence, with both abundances and faunal composition changing as time progresses.[6] Major burrowing is not present, but there are small (1-3 mm wide) burrows in the sediment, which may indicate that there is not enough oxygen in the water or sediment.[6] Particularly notable is the presence of bryozoa and graptolites,[6] forms that are absent in the Cambrian period. Diverse echinoderms indicate a normal range of salinity, and the overall shelly assemblage is not significantly different from the normal shelly fauna expected in open Ordovician waters.[6] The non-mineralized cohort contains a range of forms familiar from the Burgess Shale: Demosponges,[7] lobopods, barnacles, annelids, radiodonts (e.g. Aegirocassis),[8] possible halkieriids, marrellomorphs, paleoscolecid worms, nektaspids, skaniids as well as the expected problematica. Other Ordovician oddballs are also present, including mitrates,[9] machaeridians, cheloniellids and xiphosurans in abundance.[6]

Depositional setting

The fossiliferous strata were deposited just above storm wave base (offshore to lower shoreface transition), at between 50mand150mm (160feetand490feetm) water depth. Organisms were likely buried in situ.[10] Because of its placement above storm wave base, storms would have mobilized sediment that could be quickly deposited, trapping animals and leading to their preservation.[6] [11] Consequently, the assemblage is dominated by benthic organisms.[6]

Preservation

Fossils of the Fezouata Formation, which are usually squashed flat (although some do retain some degree of their original three-dimensionality) are often coated with a dusting of pyrite, and tin; this aspect of the fossil preservation is very similar to that at Chengjiang.[6] Non-mineralized appendages are often preserved.[6] While the formation as a whole is over 1000m (3,000feet) thick, only two intervals, 25m (82feet) and 15m (49feet) thick, provide exceptional preservation.[12] [13] Both of these intervals are located near the top of the lower formation, corresponding to the Araneograptus murrayi and Hunnegraptus copiosus graptolite zones respectively.

Location and stratigraphy

The fossils occur within an area of 500km2, in southeast Morocco's Draa Valley, north of Zagora. Stratigraphically productive layers are found through a 1.1km (00.7miles)-thick column of rock that spans the Tremadocian and Floian epochs.[6] Two stratigraphic intervals of the formation are fossiliferous: the lower is Late Tremadocian and sits 260to above the base of the formation; the upper, at 570to, is mid-Floian in age.[14]

History

The Lagerstätten were first identified in the late 1990s when a local fossil collector, Ben Moula, showed some of the finds to a PhD student who was then working in the area.[15] [16]

IUGS geological heritage site

In respect of the 'exceptional fossil preservation bridging the Cambrian Explosion and the Great Ordovician Biodiversification', the International Union of Geological Sciences (IUGS) included the 'Ordovician Fezouata Shale Fossil Site at Jbeltizagzaouine' in its assemblage of 100 'geological heritage sites' around the world in a listing published in October 2022. The organisation defines an IUGS Geological Heritage Site as 'a key place with geological elements and/or processes of international scientific relevance, used as a reference, and/or with a substantial contribution to the development of geological sciences through history.'[17]

Paleobiota

After[18] and subsequent literature:

Radiodonts

Apart from the three named species of Fezouata radiodonts, three other unnamed species occur in the formation: a third species of Pseudoangustidontus, an aegirocassisine, and a predatory hurdiid formerly mistaken for a eurypterid.[19]

Radiodonts of the Fezouata Formation
GenusSpeciesStratigraphyNotesImages
Aegirocassis[20] A. benmoulaeAraneograptus murrayi zone,Baltograptus minutus zone?[21] A giant filter-feeding aegirocassisine hurdiid radiodont. The largest known radiodont and one of the largest known panarthropods, at 2m (07feet) in length.
Pseudoangustidontus[22] P. duplospineusSagenograptus (Araneograptus) murrayi zone and Upper FezouataA filter-feeding aegirocassisine hurdiid radiodont. Originally considered enigmatic until 2023, where it was redescribed as a radiodont. Like the contemporary Aegirocassis, this genus was most likely a filter feeder, but it could have preyed upon larger food.
P. izdiguaSagenograptus (Araneograptus) murrayi zone
P. sp.Sagenograptus (Araneograptus) murrayi zone

Trilobites

The largest trilobite individuals in the Fezouata Formation tend to inhabit deep oxygenated waters with minimal influence from storms or larger predators.[23]

Trilobites of the Fezouata Formation
GenusSpeciesStratigraphyNotesImages
AgerinaA. quadrata?Cymatograptus protobalticus zone,?Baltograptus jacksoni zoneA leiostegiid which is common at intermediate depths in the transitional zone between shoreface and offshore sediments.[24]
AmpyxA. priscusAraneograptus murrayi zone up to the ?Baltograptus jacksoni zone.[25] A long-spined eyeless raphiophorid which often occurs in distinctive linear clusters. These clusters are interpreted as collective behavior akin to modern Palinurus lobsters, either as biological aggregations (for spawning or moulting), or migrating groups due to environmental pressures such as storms. Other trilobites occasionally become entangled with the Ampyx clusters. This species is widely distributed in time and is most common in relatively deep (upper offshore) sediments.
AnacheirurusA. adseraiAraneograptus murrayi zone[26] [27] A small pilekiine cheirurid, up to 4.8cm (01.9inches) long, which is common in relatively deep (upper offshore) sediments. Fossils of this species are known to preserve appendages similar in form to those of Cambrian trilobites. Known from multiple life stages, including late meraspids. Some specimens may have previously been misidentified as Lehua or Parapilekia.
ApatokephalusA. cf. incisusAraneograptus murrayi zone,?Cymatograptus protobalticus zoneAn uncommon remopleuridid found in relatively shallow (lower nearshore) sediments.
A. sp.Lower Fezouata,Upper FezouataA remopleuridid.
AsaphellusA. fezouataensis?Baltograptus jacksoni zoneAn endemic asaphid. Associated body fossils and trace fossils help to identify this species as the tracemaker for a distinctive resting trace, Rusophycus carleyi.[28] Found at intermediate depths in the transitional zone between shoreface and offshore sediments.
A. aff. jujuanusAraneograptus murrayi zone,?Cymatograptus protobalticus zone,

?Baltograptus jacksoni zone

A small and common asaphid up to 6.2cm (02.4inches) long, without clear preferences for specific depths.
A. sp.Araneograptus murrayi zoneAn asaphid. Known from multiple life stages, including late meraspids.
A. stubbsi[29] Araneograptus murrayi zoneA large endemic asaphid up to 38cm (15inches) long. Found in relatively shallow (lower nearshore) sediments.
A. tataensisAraneograptus murrayi zone,[30] Upper FezouataAn uncommon medium-sized endemic asaphid. Some specimens have been fossilized with unusual flattened antennae, the only reliable attributions of these structures in asaphids. The antennae may have had a particular sensory function to assist in hunting or scavenging. Found in relatively shallow (lower nearshore) sediments.
AsaphopsisA. sp.Lower FezouataA Dikelokephalina-like trilobite.
BasilicusB. destombesiBaltograptus jacksoni zoneA large asaphine asaphid up to 33cm (13inches) long.[31]
B. sp.A very large asaphine asaphid up to 51cm (20inches) long.
B. vidaliA large asaphine asaphid up to 36cm (14inches) long.
BathycheilusB. gallicusUpper FezouataA bathycheilid.
BavarillaB. sp.Araneograptus murrayi zoneA small bavarillid up to 4.1cm (01.6inches) long.
B. zemmourensisAraneograptus murrayi zone,Upper FezouataAn endemic bavarillid common in relatively shallow (lower nearshore) sediments.
Ceraurinella?C.? sp.Upper FezouataA cheirurid.
ColpocorypheC. pradesensis?Baltograptus jacksoni zone,Baltograptus minutus zoneA rare calymenid.
C. thorali?Cymatograptus protobalticus zone,Baltograptus minutus zoneA rare calymenid.
C. sp.Araneograptus murrayi zone,?Cymatograptus protobalticus zoneA rare calymenid found in relatively deep (upper offshore) sediments.
DactylocephalusD. sp.A large asaphid up to 29cm (11inches) long.
DikelokephalinaD. brenchleyi[32] Araneograptus murrayi zoneA large endemic dikelokephalinid up to 33.7cm (13.3inches) long. Often found in mass death assemblages, possibly for spawning akin to modern horseshoe crabs.
D. sp.Lower FezouataA large dikelokephalinid up to 29.5cm (11.6inches) long.
Ectillaenus?E.? sp.Baltograptus minutus zoneAn illaenid.
EoharpesE. sp.A harpetid.
EulomaE. sp.Araneograptus murrayi zone,?Cymatograptus protobalticus zone,

?Baltograptus jacksoni zone

A small eulomid up to 3.8cm (01.5inches) long. Found at intermediate depths in the transitional zone between shoreface and offshore sediments.
FouloniaF. peregrinaAraneograptus murrayi zone,?Baltograptus jacksoni zoneA cheirurid.
GeragnostusG. sp.Araneograptus murrayi zone,?Baltograptus jacksoni zoneA very rare artiopodan arthropod in the order Agnostida, which is often considered a group of trilobites. Very small, at up to 0.9cm (00.4inches) long.
HarpidesH. sp.A harpidid of questionable occurrence.
IndiligensI. sp.Araneograptus murrayi zoneA ptychopariid known from multiple life stages, including mid-stage meraspids.
LeptoplastidesL. salteriAnisograptus matanensis zone,Rhabdinopora flabelliformis anglica zoneAn olenid, the only diagnostic member of the family to be found in Africa.[33] Previously misidentified as Beltella.
LichakephalusL. stubbsi[34] A large lichakephalid.
Megistaspis (Ekeraspsis)M. (E.) cf. filacoviAraneograptus murrayi zone,Upper FezouataA small isoteline asaphid up to 5.6cm (02.2inches) long. Common in relatively shallow (lower nearshore) sediments.
M. (E.) hammondiAraneograptus murrayi zone[35] A large isoteline asaphid up to 28.5cm (11.2inches) long, not counting a pronounced spine on the pygidium. Some specimens preserving gut traces and appendages showing new anatomical traits not seen in other trilobites. The cephalic appendages hosted long spines capable of combing through sediment for food. This species' feeding behavior may be responsible for Cruziana rugosa, a trace fossil common throughout Ordovician Gondwana. Known from multiple life stages, including mid to late meraspids.
NeseuretusN. cf. attenuatus?Cymatograptus protobalticus zone,Baltograptus minutus zoneA calymenid found in particularly shallow sediments.
NileusN. deynouxiAraneograptus murrayi zoneA nileid known from multiple life stages, including late meraspids.
Nobiliasaphus?N.? sp.A large asaphine asaphid up to 33cm (13inches) long.
OgyginusO. forteyi hammondiA very large ogygiocaridinine asaphid, up to 45cm (18inches) long.
O. sp.Araneograptus murrayi zoneA very large ogygiocaridinine asaphid, up to 49.1cm (19.3inches) long.
OrometopusO. sp.Araneograptus murrayi zoneKnown from multiple life stages, including early to late meraspids.
ParabathycheilusP. gallicusAraneograptus murrayi zone,?Cymatograptus protobalticus zone,

?Baltograptus jacksoni zone

A common bathycheilid without clear preferences for specific depths.
Parvilichas[36] P. marochiiUpper FezouataA lichid.
Pharostomina?P.? sp.Lower FezouataA pharostomatid.
PlatycorypheP. sp.Upper FezouataA homalonotid.
PlatypeltoidesP. magrebiensisAraneograptus murrayi zoneA large endemic nileid, up to 23cm (09inches) long. Typically considered a single species. Known from multiple life stages, including early to late meraspids. Found in relatively shallow (lower nearshore) sediments.
PradoellaP. tazzarinensis?Cymatograptus protobalticus zone,Baltograptus minutus zoneA calymenid.
PrionocheilusP. aff. languedocensisAraneograptus murrayi zone,?Baltograptus jacksoni zoneA pharostomatid found in relatively shallow (lower nearshore) sediments.
SelenopeltisAn odontopleurid.
SymphysurusS. angustatusAraneograptus murrayi zone,Upper FezouataA nileid found in relatively shallow (lower nearshore) sediments.
S. ebbestadiAraneograptus murrayi zoneA common nileid with gregarious habits, often associated with graptolite debris at multiple water depths. Some specimens have been fossilized in the process of a mass moulting event, including some of the few known fossils of post-moult trilobites.[37] Known from multiple life stages, including mid to late meraspids.
S. sicardi?A small nileid up to 4.5cm (01.8inches) long
S. sp.Araneograptus murrayi zoneSeveral additional Symphysurus species, one which reaches 7.5cm (03inches) in length, and another reaching 4.2cm (01.7inches).
ToletanaspisT. aff. borniAraneograptus murrayi zone,?Cymatograptus protobalticus zone,

?Baltograptus jacksoni zone

A dalmanitid without clear preferences for specific depths.

Other arthropods

Many arthropods of the Fezouata Biota remain unnamed and undescribed. These include synziphosurines, xiphosurans (horseshoe crabs), eurypterids, chasmataspidids, phyllocarids, ostracods, a canadaspidid, a leanchoiliid, a cheloniellid (Eoduslia?),[38] a possible retifaciid, and a lepadomorph barnacle.

Other arthropods of the Fezouata Formation
GenusSpeciesStratigraphyNotesImages
Brachyaglaspis[39] B. singularisUpper Fezouata (mid-Floian)An aglaspidid with a very large cephalon, a short trunk, and no eyes.
Enosiaspis[40] E. hrungnirAraneograptus murrayi zoneAn acercostracan marrellomorph with a shield-like dorsal carapace.
"Furca"Sagenograptus (Araneograptus) murrayi zoneA marrellid marrellomorph which has not been formally named or described in detail. It had three pairs of cephalic appendages, 13 pairs of trunk appendages, and a cephalic shield with six large setae-bearing spines. Both adult and juvenile specimens are known of this species,[41] including individuals caught in the act of moulting.[42] The name "Furca mauretanica" is an informal nomen nudum for this species, first introduced in a PhD thesis. Multiple marrellid species may be present in the formation.
Setapedites[43] S. abundantisAraneograptus murrayi zoneA small but abundant offacolid "synziphosurine" (early euchelicerate). Though most closely related to Dibasterium and Offacolus from the Silurian Herefordshire Lagerstätte, it also shows similarities to Habelia from the Cambian, and thus serves as a transitional form among Ordovician euchelicerate evolution.
TariccoiaT. tazagurtensis[44] Araneograptus murrayi zoneA liwiid nektaspid with a large cephalon and pygidium and sharply angled thorax tergites. One of the few liwiid fossils with soft tissue of the digestive system preserved.
ThelxiopeT. sp.Araneograptus murrayi zoneA mollisoniid, possibly related to chelicerates.[45]
TremaglaspisT. sp.An aglaspidid, one of the first expectionally-preserved invertebrates found in the formation.

Echinoderms

Many echinoderm species of the Fezouata Biota remain unnamed and undescribed. These include representatives of chauvelicystids, other cornutes, anomalacystitids, mitrocystitids, eocrinoids, rhenopyrgids, and somasteroids.[46] Fezouata stylophoran fossils include soft tissue preserved among the skeletal elements, helping to unravel controversial details of their anatomy and ecology.[47]

Specific echinoderm species may form dense fossil beds in some layers of the formation, a phenomenon which is particularly common in the mid-late Tremadocian (Araneograptus murrayi graptolite zone). Most echinoderm beds are dominated by only a few species, often representatives of stylophorans or the eocrinoid Rhopalocystis, with few other animal fossils. Through nearly the entire the formation, small Rhopalocystis species dominate "meadow"-like ecosystems in shallow waters impacted by storms. In the youngest layers of the formations, diploporites usurp this niche. Conversely, stylophorans are opportunistic colonizers of deeper low-oxygen seabeds in some intervals of the Araneograptus murrayi zone. Echinoderms are uncommon at intermediate depths, which have a higher proportion of brachiopod and trilobite fossils.

Nevertheless, there are quite a few exceptions which contradict these broad rules. Many sites record diverse deep-water ecosystems protected from both storms and insufficient oxygen. Numerous species of echinoderms and other invertebrates coexist at these localities, even if one echinoderm species outnumbers other fossils at any given time. Large Rhopalocystis species, Macrocystella, Plasiacystis, and Balantiocystis are common components of these assemblages.

Fezouata reconstructs the uneven nature of the Cambrian-Ordovician transition of echinoderm faunas. Cosmopolitan Late Cambrian hallmarks (such as cornute stylophorans) maintian their abundance in oxygen-poor areas, while newer groups (crinoids, diploporites, asterozoans) make inroads into more oxygenated waters. The rarity of carbonate platforms and hard substrates in the Gondwanan area delayed the diversification of crinoids and edrioasteroids in the region. This also provided more space for the establishment of a distinctive South Polar ecosystem dominated by eocrinoids, mitrates, solutans, and eventually diploporites.

Echinoderms of the Fezouata Formation
GenusSpeciesStratigraphyNotesImages
AethocrinusA. cf. mooreiAraneograptus murrayi zoneA crinoid.
AmpelocarpusA. sp.Araneograptus murrayi zoneA rare cornute.
AmygdalothecaA. griffeiAraneograptus murrayi zoneAn uncommon cornute.
AnatifopsisA. escandreiHunnegraptus copuosus zoneA rare mitrate.
A. trapeziiformisAraneograptus murrayi zone,Hunnegraptus copiosus zone,

?Baltograptus jacksoni zone

A common mitrate.
AnedriophusA. moroccoensis?Baltograptus jacksoni zoneAn uncommon and endemic edrioasteroid.
ArgodiscusA. espilezorum?Cymatograptus protobalticus zone,?Baltograptus jacksoni zoneAn uncommon edrioasteroid.
AristocystitesA. cf. sinuosusAraneograptus murrayi zoneA rare diploporite.
AspidocarpusA. sp.Araneograptus murrayi zoneA rare mitrate.
BalanocystitesB. primus?Baltograptus jacksoni zoneA common mitrate.
BalantiocystisB. regnelliAraneograptus murrayi zone,?Cymatograptus protobalticus zone,

?Baltograptus jacksoni zone

A common eocrinoid. The dominant echinoderm at sites Z-F26 and Z-F23, which preserve diverse fossil assemblages buried in calm-water fine-grained siltstone of the ?Cymatograptus protobalticus zone. Also abundant in higher-energy deposits of the ?Baltograptus jacksoni zone.
BohemiaecystisB. sp.Araneograptus murrayi zoneA locally abundant cornute with a large sample size, allowing for a reconstruction of decay pathways in stylophoran soft tissue and skeletal material.[48]
ChauvelicystisC. spinosaAraneograptus murrayi zoneA rare cornute.
C. ubaghsiAraneograptus murrayi zoneAn uncommon cornute.
C. vizcainoiAraneograptus murrayi zoneA rare cornute.
ChinianocarposC. sp."Azygograptus interval"An uncommon mitrate.
FlabellicarpusF. rushtoniAraneograptus murrayi zoneA rare cornute.
GalliaecystisG. ubaghsiAraneograptus murrayi zoneA rare cornute.
G. sp.Araneograptus murrayi zoneAn uncommon cornute.
HanusiaH. sp.Araneograptus murrayi zoneAn uncommon cornute.
IocrinusI. sp.?Cymatograptus protobalticus zoneA rare disparid crinoid.
LagnocystisL. pyramidalisAraneograptus murrayi zoneA rare mitrate.
LingulocystisL. aff. deani?Cymatograptus protobalticus zone,?Baltograptus jacksoni zoneA rare eocrinoid.
L. elongataAraneograptus murrayi zone,?Baltograptus jacksoni zoneA rare eocrinoid.
MacrocystellaM. bohemicaAraneograptus murrayi zone,?Baltograptus jacksoni zoneA common glyptocystitid rhombiferan. Crushed but complete skeletons of this species are abundant at Oued Beni Zoli locality Z-F5, a site preserving a diverse assemblage of storm-influenced siltstone in the Araneograptus murrayi zone.
NanocarpusN. cf. dolambiiAraneograptus murrayi zoneA rare cornute.
Nimchacystis[49] N. agterbosiAraneograptus murrayi zoneAn uncommon minervaecystid solutan.
Paleosphaeronites?P.? prokopiAraneograptus murrayi zone,?Baltograptus jacksoni zoneA rare diploporite.
cf. PareocrinusAraneograptus murrayi zone,?Cymatograptus protobalticus zoneAn uncommon eocrinoid.
PeltocystisP. cornutaAraneograptus murrayi zoneAn uncommon mitrate.
PlasiacystisP. mobilisAraneograptus murrayi zone up to the Baltograptus minutus zoneA common minervaecystid solutan. Paddles of this species are abundant at sites Z-F13c and Z-F24, a pair of diverse faunal assemblages emplaced in sandy storm deposits of the Hunnegraptus copuosus zone.
ProcothurnocystisP. sp.Araneograptus murrayi zoneA common cornute.
ProkopicystisP. sp.Araneograptus murrayi zoneA rare cornute.
RamseyocrinusR. sp.?Cymatograptus protobalticus zone,?Baltograptus jacksoni zoneA rare disparid crinoid.
RhopalocystisR. destombesiAorograptus victoriae zone,Araneograptus murrayi zone,

Hunnegraptus copiosus zone,

?Baltograptus jacksoni zone

A common and endemic eocrinoid which can be abundant in certain layers. It is a small but robust species often found in densely jumbled fossiliferous lenses. These lenses are interpreted as debris from shallow-water "meadows", uprooted and washed down into deeper areas by storms.[50]
R. fragaAraneograptus murrayi zoneA rare and endemic eocrinoid, similar to R. destombesi in form and preservation style.
R. grandisAraneograptus murrayi zoneA rare and endemic eocrinoid, similar to R. havliceki in form and preservation style.
R. havlicekiAraneograptus murrayi zoneA common and endemic eocrinoid. It is a large but fragile species which is often found disarticulated in siltstone beds. Unlike R. destombesi, it was likely a deep-water specialist buried in place.
R. zagoraensisAraneograptus murrayi zone,Hunnegraptus copiosus zoneA common and endemic eocrinoid, similar to R. destombesi in form and preservation style.
ThoralicystisT. sp.Araneograptus murrayi zoneA common cornute.
T. zagoraensis?Baltograptus jacksoni zoneA rare cornute.
VillebrunasterV. fezouatensis[51] Araneograptus murrayi zoneA chinianasterid somasteroid, a sea star-like echinoderm with a broad pentagonal body. Originally described in a new genus, Cantabrigiaster,[52] but subsequently referred to Villebrunaster.
VizcainocarpusV. sp.Araneograptus murrayi zoneA rare mitrate.

Molluscs

Molluscs of the Fezouata Formation
GenusSpeciesStratigraphyNotesImages
Alococoncha?A.? sp."Azygograptus interval"A rare ctenodontid protobranch bivalve. It was likely an infaunal deposit feeder.[53]
BabinkaB. prima?Cymatograptus protobalticus zone,?Baltograptus jacksoni zone,

"Azygograptus interval"

A common bivalve, tentatively identified as one of the oldest lucinids (hatchet shells). It was likely an infaunal filter-feeder capable of rapid vertical movement within sediment. Like modern lucinids, it may have been a specialist in oxygen-poor environments thanks to endosymbiotic sulfide-oxidizing bacteria.
BactrocerasB. sp."late early to mid Floian"A very rare baltoceratid "nautiloid" cephalopod in the order Orthocerida.[54]
BathmocerasB. australe"late early to mid Floian"A very rare bathmoceratid "nautiloid" cephalopod in the order Ellesmerocerida.
B. taichoutense"late early to mid Floian"
Calvapilosa[55] C. kroegeriAraneograptus murrayi zoneA mollusc in the group Sachitida (the proposed clade containing living aculiferans and their extinct stem group relatives such as halkieriids). It had a flattened body with an extensive radula, a mantle covered in bristle-like sclerites, and a single flat calcareous shell on the head.
CarcassonnellaC. courtessolei?Cymatograptus protobalticus zone,?Baltograptus jacksoni zoneA common tergomyan monoplacophoran.[56]
C. sp.Araneograptus murrayi zone,?Cymatograptus protobalticus zone?A common tergomyan monoplacophoran known from tiny juveniles which died in relatively deep, oxygen-poor waters.
C. vizcainoi?Cymatograptus protobalticus zoneA common tergomyan monoplacophoran.
Cardiolaria?"Azygograptus interval"A rare afghanodesmatid protobranch bivalve. It may have been an infaunal filter-feeder.
Cienagomya?C.? sp.?Cymatograptus protobalticus zone,A very rare actinodontid bivalve in the family Intihuarellidae. It was likely an infaunal filter-feeder.
CoxiconchiaC. guiraudi?Cymatograptus protobalticus zone,"Azygograptus interval"A common bivalve, tentatively identified as one of the oldest lucinids (hatchet shells). It was likely an infaunal filter-feeder capable of rapid vertical movement within sediment. Like modern lucinids, it may have been a specialist in oxygen-poor environments thanks to endosymbiotic sulfide-oxidizing bacteria.
DestombesicerasD. zagorense"late early to mid Floian"An apicrinoceratid "nautiloid" cephalopod in the order Discosorida. The least rare nautiloid found in the formation, though still uncommon.
EkaterodontaE. courtessolei?Cymatograptus protobalticus zone,?Baltograptus jacksoni zoneA rare afghanodesmatid protobranch bivalve. It was likely an infaunal deposit feeder.
GlyptarcaG. sp.?Cymatograptus protobalticus zone,A rare arcid bivalve. It was likely an infaunal filter-feeder.
LesueurillaL. prima?Cymatograptus protobalticus zone,?Baltograptus jacksoni zone,

Baltograptus minutus zone

A lesuerillid gastropod.
Pelecyogyra[57] P. fezouataensisAraneograptus murrayi zoneA common onychochilid paragastropod, an extinct group of gastropod-like molluscs with asymmetrically coiled shells. It was gregarious and primarily found in shallow oxygenated waters represented at the Oued Beni Zoli locality.
PolymeresP. sp."late early to mid Floian"A rare polymerid "nautiloid" cephalopod in the order Dissidocerida.
Praenucula?P.? sp.?Cymatograptus protobalticus zone,A very rare praenuculid protobranch bivalve known from a single specimen. It was likely an infaunal deposit feeder.
ProtocyptendocerasP. longicameratum"late early to mid Floian"A very rare protocameroceratid "nautiloid" cephalopod in the order Endocerida.
RedoniaR. michelae?Cymatograptus protobalticus zone,?Baltograptus jacksoni zone,

"Azygograptus interval"

A very common actinodontid bivalve in the family Redoniidae. It was likely a shallowly infaunal filter-feeder capable of rapid reburial, with the majority of the shell embedded in sediment.
RibeiriaR. sp.Upper FezouataA rostroconch.[58]
RiocerasR. sp."late early to mid Floian"A very rare rioceratid "nautiloid" cephalopod in the order Ellesmerocerida.
SinuitesS. sp.?Baltograptus jacksoni zoneA rare bellerophontoid gastropod.
ThoralispiraT. laevis?Baltograptus jacksoni zoneA common tergomyan monoplacophoran.

Conodonts

Conodonts from the Fezouata Formation are mostly coniform elements of Early Ordovician species.[59] Overall diversity is rather low, and species which were common in temperate and tropical seas are apparently absent. The Fezouata Formation appears to be an exemplar of the 'subpolar domain', an assemblage of cold-water coastal conodonts native to the South Polar region of the Early Ordovician. Similar conodont faunas are known from Early Ordovician deposits in Central Europe, which was also located near the South Pole. The 'subpolar domain' survived into the Middle Ordovician and expanded into areas now found in the Middle East.

Fezouata conodonts are difficult to correlate to biostratigraphic systems in nearby temperate regions such as Baltica. Most species appear to correspond to the time interval stretching from the Oelandodus elongatus-Acodus deltatus subzone of the Paroistodus proteus zone (uppermost Tremadocian) up through the Prioniodus oepiki zone (lower Floian).

Conodonts of the Fezouata Formation
GenusSpeciesNotesImages
AcodusA. deltatusAn acodontid prioniodontid
CornuodusC. longibasisA strachanognathid
DrepanodusD. arcuatusA drepanoistodontid or protopanderodontid
DrepanoistodusD. cf. forcepsA drepanoistodontid
PaltodusP. cf. inaequalisA drepanoistodontid
P. spp.
ParapaltodusP. cf. flexuosusAn acanthodontid
P. aff. simplicissimus
ParoistodusP. cf. parallellusA drepanoistodontid
P. proteus
PeriodonP. cf. primusA periodontid ozarkodinid
P. cf. selenopsis
PrioniodusP. cf. gilbertiA balognathid prioniodontid
ProtopanderodusP. cf. leonardiiA protopanderodontid
ScalpellodusS. aff. gracilisAn acanthodontid
ScolopodusS. aff. krummiA protopanderodontid
SemiacontiodusS. cf. cornuformisA protopanderodontid
S. sp.
StolodusS. cf. stolaA belodellid
TripodusT. aff. laevisAn acodontid
T. cf. sweeti

Graptolites

Graptolites in the Fezouata Formation are important for biostratigraphic correlation to other regions. Most species found in the formation are assigned to planktic (graptoloid) groups, though some mesh-shaped taxa (like Araneograptus and Rhabdinopora) may have been transitional between sessile (dendroid) graptolites and free-floating graptoloids.[60] Sessile forms, such as Didymograptus, Dictyonema, Webbyites, and rhabdopleurids, are also present but much more rare.[61]

The graptolites of the Fezouata Formation are distributed over 10 biozones. In order, these zones are: the Anisograptus matanensis zone (1), Rhabdinopora flabelliformis anglica zone (2), “Adelograptustenellus zone (3), Aorograptus victoriae zone (4), Araneograptus murrayi zone (5), Hunnegraptus copiosus zone (6), ?Cymatograptus protobalticus zone (7), ?Baltograptus jacksoni zone (8), Baltograptus minutus zone (9), and the “Azygograptus interval” (10). Not all of the index taxa which these zones are named for are known from the Fezouata Formation. The Tremadocian-Floian boundary is approximately at the level between the Hunnegraptus copiosus and ?Cymatograptus probalticus zones.

The most well-preserved fossils in the Fezouata Formation generally come from strata of the late Tremadocian (late Araneograptus murrayi zone and early Hunnegraptus copiosus zone) and the mid-late Floian (late Baltograptus minutus zone and early "Azygograptus interval").

Graptolites of the Fezouata Formation
GenusSpeciesGraptolite biozoneNotesImages
"Adelograptus""A." tenellusAdelograptustenellus zoneA multiramous anisograptid (basal graptoloid).
AncoragraptusA. bulmaniAorograptus victoriae zone A "psigraptid"-type anisograptid, with a small number of reclined stipes.
AnisograptusA. cf. matanensisAnisograptus matanensis zoneA multiramous anisograptid. Very rare, only a single fossil may indicate its presence.
AraneograptusA. murrayiAraneograptus murrayi zoneA common basal graptoloid with a mesh-like conical colony form. Potentially a species of Sagenograptus.
AzygograptusA. eivionicusAzygograptus interval”A rare "azygograptid", a unique type of early graptoloid with a single uniserial stipe, convergently similar to the Silurian-Devonian monograptids.
BaltograptusB. deflexusBaltograptus minutus zoneA didymograptid graptoloid, with two subhorizontal to pendent stipes.
?B. jacksoni?Baltograptus jacksoni zone
?B. kurckiBaltograptus minutus zone
B. minutusBaltograptus minutus zone
ChoristograptusC. louaiAorograptus victoriae zone A small and rare anisograptid.
ClonograptusC. multiplex?Cymatograptus protobalticus zone,?Baltograptus jacksoni zoneA large and common multiramous dichograptid.
C. rigidusAraneograptus murrayi zone,Hunnegraptus copiosus zone
?Cymatograptus?C. protobalticus?Cymatograptus protobalticus zoneA rare didymograptid.
"Didymograptus""D." sp.Araneograptus murrayi zone,Hunnegraptus copiosus zoneA rare didymograptid.
ExpansograptusE. sp.?Baltograptus jacksoni zoneA rare didymograptid with two horizontal stipes.
HolograptusH. sp.Baltograptus minutus zoneA multiramous dichograptid.
HunnegraptusH. copiosusHunnegraptus copiosus zoneA multiramous anisograptid.
"Kiaerograptus""K." supremusAraneograptus murrayi zone,Hunnegraptus copiosus zoneA "psigraptid"-type anisograptid, with a small number of reclined stipes.
ParadelograptusP. norvegicusAraneograptus murrayi zone,Hunnegraptus copiosus zoneA multiramous sigmagraptid graptoloid.
P. tenuisAraneograptus murrayi zone
ParatemnograptusP. magnificusAraneograptus murrayi zone,Hunnegraptus copiosus zoneA large and common multiramous anisograptid.
RhabdinoporaR. flabelliformis anglicaRhabdinopora flabelliformis anglica zoneA basal graptoloid with a mesh-like conical colony form transitional between dendroids (such as Dictyonema) and early graptoloids such as anisograptids.
R. flabelliformis canadensisAnisograptus matanensis zone
R. flabelliformis flabelliformisAnisograptus matanensis zone,Rhabdinopora flabelliformis anglica zone
R. flabelliformis socialisAnisograptus matanensis zone
indeterminate rhabdopleuridSagenopterus (Araneograptus) murrayi zoneTubes of Rhabdopleura-like pterobranchs have been found growing around the edge of a cephalopod shell, one of the few examples of hard substrate to be found in the formation. This association is similar to the ecology reported for Yuknessia from the Burgess Shale.
Schizograptus?S. sp.Baltograptus minutus zoneA common multiramous dichograptid, also sometimes classified as Holograptus.
Tetragraptus"T. bulmani"Araneograptus murrayi zoneA common tetragraptid graptoloid with two to four reclined to pendent stipes.
Other Tetragraptus speciesAraneograptus murrayi zone,?Baltograptus jacksoni zone
WebbyitesW. felix[62] "probable late Tremadocian"An enigmatic feather-shaped organism identified as a sessile graptolite.

Brachiopods

Brachiopods of the Fezouata Formation
GenusSpeciesStratigraphyNotesImages
AcrotheleA. sp.Lower FezouataAn acrothelid linguliform.
AngusticardiniaA. sp.Upper FezouataAn angusticardiniid orthid.
LingulaL. salteriUpper FezouataA lingulid linguliform.
OrbitheleO. vanaUpper FezouataAn acrothelid linguliform.
PaurorthisP. tadristensisUpper FezouataA paurorthid orthid.
PlectorthisP. simplexLower FezouataA plectorthid orthid.
RanorthisR. fasciataLower and Upper FezouataA ranorthid orthid.
TarfayaT. marocanaUpper FezouataA heterorthid orthid.

Sponges

Many sponges from the Fezouata biota remain unnamed, including protomonaxonid demosponges (leptomitids, "choiids", hamptoniids, piraniids), a hazeliid, reticulosan hexactinellids (asthenospongiids, etc.), and other indeterminate forms.[63] Many of the sponges have affinities with Cambrian taxa common in Burgess Shale-type faunas. Though at least 27 sponge species have been recorded in the biota, nearly all occurrences are monospecific death assemblages, with the exception of Pirania auraeum, which has a broader and less dense distribution in the formation. Periodic unstable seafloor conditions (potentially related to seasonal disruptions) would have favored species-poor colonization events over short time periods, rather than a stable and diverse equilibrium.

This is unusual relative to other Ordovician sponge ecosystems, such as the Builth Inlier of Wales. Another difference is how Fezouata's sponge fauna consists mostly of protomonaxonids, with a few reticulosans occupying course-grained shallow seabeds. Conversely, in Wales there is a clear succession of diverse and sturdy lithistids and thick-walled hexactinellids in shallow reefs and other energetic areas, with protomonaxonids at intermediate depths, and reticulosans in the deepest and calmest environments.

Sponges of the Fezouata Formation
GenusSpeciesStratigraphyNotesImages
ChoiaC. sp.Upper FezouataA "choiid".
ChoiaellaC. sp.A "choiid".
HamptoniaH. sp.A hamptoniid.
'Hamptonia'H.' christiUpper FezouataAn indeterminate protomonaxonid. Previously considered a species of Hamptonia, but likely belongs to a new genus.
PiraniaP. auraeumUpper FezouataA piraniid.
Valospongia?V.? sp.A reticulosan.

Other animals

Many animals from the Fezouata biota remain unnamed and undescribed, including bryozoans, chordate "dermal plates", annelid worms, priapulids, problematica, and at least five new species of lobopod, including an armored form.

Other animals of the Fezouata Formation
GenusSpeciesStratigraphyNotesImages
ArchaeoconulariaA. sp.Lower and Upper FezouataA common large conulariid, up to 50cm (20inches) tall.[64]
CavernolitesC. senexUpper FezouataA hyolith.
ElegantilitesE. sp.Upper FezouataA hyolith.
EoconulariaE. sp.Lower and Upper FezouataAn abundant small conulariid (less than 5cm (02inches) tall). Frequently found attached to brachiopods or even other conulariids.
Gamalites?G.? sp.Upper FezouataA hyolith.
GompholitesG. sp.Upper FezouataA hyolith.
NephrothecaN. sp.Upper FezouataA hyolith.
Palaeoscolex?P.? tenensisAraneograptus murrayi zoneA palaeoscolecid worm. Though most palaeoscolecids are inferred to be burrowing carnivores, remains from the Fezouata Formation are more consistent with an opportunistic epibenthic lifestyle, patrolling for organic matter on the surface of the seabed.[65] [66]
PauxillitesP. thaddei[67] Upper FezouataA medium-sized pauxillitid hyolith.
PlumulitesP. bengtsoni[68] Araneograptus murrayi zoneA machaeridian (armored annelid worm). This species is the only machaeridian known to preserve soft tissue and jaw material, helping to demonstrate their affinities with aphroditiform polychaetes (scaleworms and kin).[69]
SphenothallusS. spp.Lower and Upper FezouataA tubular organism likely closely related to conulariids.

Other organisms

Chitinozoans, acritarchs, and algae have been recorded from the formation.[70]

Notes and References

  1. 10.1111/let.12240. Age calibration of the Lower Ordovician Fezouata Lagerstätte, Morocco. Lethaia. 2017. Lefebvre. Bertrand. Gutiérrez-Marco. Juan C. Lehnert. Oliver. Martin. Emmanuel L. O. Nowak. Hendrik. Akodad. Mustapha. El Hariri. Khadija. Servais. Thomas. 51. 2. 296–311. free.
  2. http://www.fossilworks.org/cgi-bin/bridge.pl?action=displayStrata&geological_group=&formation=Fezouata&group_formation_member=Fezouata Fezouata Formation
  3. http://www.fossilworks.org/cgi-bin/bridge.pl?action=displayStrata&geological_group=&formation=Fezouata%20Shale&group_formation_member=Fezouata%20Shale Fezouata Shale
  4. http://www.fossilworks.org/cgi-bin/bridge.pl?action=displayStrata&geological_group=&formation=Lower%20Fezouata&group_formation_member=Lower%20Fezouata Lower Fezouata Formation
  5. http://www.fossilworks.org/cgi-bin/bridge.pl?action=displayStrata&geological_group=&formation=Upper%20Fezouata&group_formation_member=Upper%20Fezouata Upper Fezouata Formation
  6. Van Roy . P. . Orr . P. J. . Botting . J. P. . Muir . L. A. . Vinther . J. . Lefebvre . B. . Hariri . K. E. . Briggs . D. E. G. . Derek Briggs . 2010 . Ordovician faunas of Burgess Shale type . Nature . 465 . 7295 . 215–8 . 2010Natur.465..215V . 10.1038/nature09038 . 20463737 . 4313285.
  7. 737–748. 2007. 10.1016/j.geobios.2007.02.006. J.. 40. 'Cambrian' demosponges in the Ordovician of Morocco: Insights into the early evolutionary history of sponges. Geobios. Botting. 6 . 2007Geobi..40..737B.
  8. Van Roy . P. . Briggs . D. E. G. . Derek Briggs. 10.1038/nature09920 . A giant Ordovician anomalocaridid . Nature . 473 . 7348 . 510–513 . 2011 . 21614078 . 2011Natur.473..510V . 205224390 .
  9. 10.1016/j.annpal.2007.06.003. First report of the mitrate Peltocystis cornuta Thoral (Echinodermata, Stylophora) in the Lower Ordovician of central Anti-Atlas (Morocco). 2007. Lefebvre . B.. Botting . J.. Annales de Paléontologie. 93. 3. 183. 2007AnPal..93..183L.
  10. Martin. Emmanuel L.O.. Pittet. Bernard. Gutiérrez-Marco. Juan-Carlos. Vannier. Jean. El Hariri. Khadija. Lerosey-Aubril. Rudy. Masrour. Moussa. Nowak. Hendrik. Servais. Thomas. Vandenbroucke. Thijs R.A.. Van Roy. Peter. Vaucher. Romain. Lefebvre. Bertrand. The Lower Ordovician Fezouata Konservat-Lagerstätte from Morocco: Age, environment and evolutionary perspectives. Gondwana Research. 34. 2016. 274–283. 1342-937X. 10.1016/j.gr.2015.03.009. 2016GondR..34..274M. free.
  11. Web site: Saleh . Farid . Candela . Yves . Harper . David A. T. . Polechová . Marika . Lefebvre . Bertrand . Pittet . Bernard . Storm-Induced Community Dynamics in the Fezouata Biota (Lower Ordovician, Morocco) . (limited access to full article) geoscienceworld.org . Society for Sedimentary Geology PALAIOS, print edition: 21 December 2018, pp. 535-541 (Vol. 33, No. 12) . 1 March 2019.
  12. Web site: Saleh . Farid . Pittet . Bernard . Perrillat . Jean-Philippe . Lefebvre . Bertrand . Orbital Control on Exceptional Fossil Preservation (abstract only) . geoscienctworld.org (subscription w/ free abstract) . Geology (magazine published) 1 February 2019 . 1 March 2019.
  13. News: Shifts in Earth's Orbit Increase the Chances of Spectacular Fossils . The Economist . The Economist, print edition: January 26th 2019, p. 72 (1/2-page based on article in Geology--in prior footnote--1/Feb/2019). 1 March 2019.
  14. 10.1111/let.12240. Age calibration of the Lower Ordovician Fezouata Lagerstätte, Morocco. Lethaia. 2017. Lefebvre. Bertrand. Gutiérrez-Marco. Juan C. Lehnert. Oliver. Martin. Emmanuel L. O. Nowak. Hendrik. Akodad. Mustapha. El Hariri. Khadija. Servais. Thomas. 51. 2. 296–311. free.
  15. Exceptionally Preserved Faunas from the Lower Ordovician of the Anti-Atlas, Morocco. Van Roy, P. Lefebvre, B. El Hariri, K. Hafid, A. 2008. First international Conference and Exhibition, Marrakech, Morocco.
  16. 10.1038/news.2010.234. Jones . N.. Weird wonders lived past the Cambrian. 2010. Nature.
  17. Web site: The First 100 IUGS Geological Heritage Sites . IUGS International Commission on Geoheritage . IUGS . 13 November 2022.
  18. Van Roy . Peter . Briggs . Derek E. G. . Gaines . Robert R. . September 2015 . The Fezouata fossils of Morocco; an extraordinary record of marine life in the Early Ordovician . Journal of the Geological Society . en . 172 . 5 . 541–549 . 10.1144/jgs2015-017 . 2015JGSoc.172..541V . 129319753 . 0016-7649.
  19. Potin . Gaëtan J.-M. . Gueriau . Pierre . Daley . Allison C. . 2023 . Radiodont frontal appendages from the Fezouata Biota (Morocco) reveal high diversity and ecological adaptations to suspension-feeding during the Early Ordovician . Frontiers in Ecology and Evolution . 11 . 10.3389/fevo.2023.1214109 . free . 2296-701X.
  20. Van Roy . Peter . Daley . Allison C. . Briggs . Derek E. G. . 2015 . Anomalocaridid trunk limb homology revealed by a giant filter-feeder with paired flaps . Nature . en . 522 . 7554 . 77–80 . 10.1038/nature14256 . 25762145 . 2015Natur.522...77V . 1476-4687.
  21. Saleh . Farid . Vaucher . Romain . Vidal . Muriel . Hariri . Khadija El . Laibl . Lukáš . Daley . Allison C. . Gutiérrez-Marco . Juan Carlos . Candela . Yves . Harper . David A. T. . Ortega-Hernández . Javier . Ma . Xiaoya . Rida . Ariba . Vizcaïno . Daniel . Lefebvre . Bertrand . 2022-12-13 . New fossil assemblages from the Early Ordovician Fezouata Biota . Scientific Reports . en . 12 . 1 . 20773 . 10.1038/s41598-022-25000-z . 36513689 . 9747710 . 2022NatSR..1220773S . 2045-2322.
  22. Van Roy . Peter . Tetlie . O. Erik . 2006 . A spinose appendage fragment of a problematic arthropod from the Early Ordovician of Morocco . Acta Palaeontologica Polonica . 51 . 2 . 239–246.
  23. Saleh . Farid . Vidal . Muriel . Laibl . Lukáš . Sansjofre . Pierre . Gueriau . Pierre . Pérez-Peris . Francesc . Lustri . Lorenzo . Lucas . Victoire . Lefebvre . Bertrand . Pittet . Bernard . El Hariri . Khadija . Daley . Allison C. . 2021 . Large trilobites in a stress-free Early Ordovician environment . Geological Magazine . en . 158 . 2 . 261–270 . 10.1017/S0016756820000448 . 2021GeoM..158..261S . 0016-7568.
  24. Martin . Emmanuel L.O. . Vidal . Muriel . Vizcaïno . Daniel . Vaucher . Romain . Sansjofre . Pierre . Lefebvre . Bertrand . Destombes . Jacques . 2016 . Biostratigraphic and palaeoenvironmental controls on the trilobite associations from the Lower Ordovician Fezouata Shale of the central Anti-Atlas, Morocco . Palaeogeography, Palaeoclimatology, Palaeoecology . en . 460 . 142–154 . 10.1016/j.palaeo.2016.06.003. 2016PPP...460..142M .
  25. Vannier . Jean . Vidal . Muriel . Marchant . Robin . El Hariri . Khadija . Kouraiss . Khaoula . Pittet . Bernard . El Albani . Abderrazak . Mazurier . Arnaud . Martin . Emmanuel . 2019-10-17 . Collective behaviour in 480-million-year-old trilobite arthropods from Morocco . Scientific Reports . en . 9 . 1 . 14941 . 10.1038/s41598-019-51012-3 . 2045-2322 . 6797724 . 31624280. 2019NatSR...914941V .
  26. Pérez-Peris . Francesc . Laibl . Lukáš . Vidal . Muriel . Daley . Allison C. . 2021 . Systematics, morphology, and appendages of an Early Ordovician pilekiine trilobite Anacheirurus from Fezouata Shale and the early diversification of Cheiruridae . Acta Palaeontologica Polonica . 66 . 4 . 857–877 . 10.4202/app.00902.2021.
  27. Laibl . Lukáš . Drage . Harriet B. . Pérez-Peris . Francesc . Schöder . Sebastian . Saleh . Farid . Daley . Allison C. . 2023 . Babies from the Fezouata Biota: Early developmental trilobite stages and their adaptation to high latitudes . Geobios . en . 81 . 31–50 . 10.1016/j.geobios.2023.06.005. 2023Geobi..81...31L .
  28. Gibb . Stacey . Chatterton . Brian D. E. . Gingras . Murray K. . 2010-11-30 . Rusophycus carleyi (James, 1885), Trace Fossils from the Lower Ordovician of Southern Morocco, and the Trilobites that Made Them . Ichnos . en . 17 . 4 . 271–283 . 10.1080/10420940.2010.535452 . 2010Ichno..17..271G . 1042-0940.
  29. Fortey . Richard A. . 2009 . A New Giant Asaphid Trilobite from the Lower Ordovician of Morocco . Memoirs of the Association of Australasian Palaeontologists . 37 . 9–16.
  30. Edgecombe . Gregory D. . Fortey . Richard. A. . 2023 . A novel antennal form in trilobites . Journal of Paleontology . en . 97 . 1 . 152–157 . 10.1017/jpa.2022.59 . 2023JPal...97..152E . 0022-3360.
  31. Corbacho . Joan . Vela . Joan Antoni . 2010 . Giant Trilobites from Lower Ordovician of Morocco . Batalleria . 15 . 3–34.
  32. Fortey . Richard A. . 2011 . Trilobites of the genus Dikelokephalina from Ordovician Gondwana and Avalonia . Geological Journal . en . 46 . 5 . 405–415 . 10.1002/gj.1275 . 2011GeolJ..46..405F . 0072-1050.
  33. Hopkins . Melanie J. . Gutiérrez-Marco . Juan Carlos . Di Silvestro . Gianpaolo . 2024-01-15 . First occurrence of well-preserved Ordovician trilobites of the family Olenidae from Africa . Journal of Paleontology . en . 1–10 . 10.1017/jpa.2023.60 . 0022-3360. free .
  34. Fortey . Richard A. . 2012 . The first known complete Lichakephalid trilobite, lower Ordovician of Morocco . Memoirs of the Association of Australasian Palaeontologists . 42 . 1–7.
  35. Gutiérrez-Marco . Juan C. . García-Bellido . Diego C. . Rábano . Isabel . Sá . Artur A. . 2017-01-10 . Digestive and appendicular soft-parts, with behavioural implications, in a large Ordovician trilobite from the Fezouata Lagerstätte, Morocco . Scientific Reports . en . 7 . 1 . 39728 . 10.1038/srep39728 . 2045-2322 . 5223178 . 28071705. 2017NatSR...739728G .
  36. Corbacho . Joan . Vela . Joan A. . Parvilichas marochii: New genus and species of Lichidae from the Zagora region (Morocco); Early Ordovician (Floian) . Scripta Musei Geologici Seminarii Barcinonensis [Ser. palaeontologica] . 14 . 3–13.
  37. Drage . Harriet B. . Vandenbroucke . Thijs R.A. . Van Roy . Peter . Daley . Allison C. . 2019 . Sequence of post-moult exoskeleton hardening preserved in a trilobite mass moult assemblage from the Lower Ordovician Fezouata Konservat-Lagerstätte, Morocco . Acta Palaeontologica Polonica . 64 . 2 . 261–273 . 10.4202/app.00582.2018.
  38. Sumrall . Colin D. . Zamora . Samuel . 2011 . Ordovician edrioasteroids from Morocco: faunal exchanges across the Rheic Ocean . Journal of Systematic Palaeontology . 9 . 3 . 425–454 . 2011JSPal...9..425S . 10.1080/14772019.2010.499137 . 1477-2019.
  39. Ortega-Hernández . Javier . Van Roy . Peter . Lerosey-Aubril . Rudy . May 2016 . A new aglaspidid euarthropod with a six-segmented trunk from the Lower Ordovician Fezouata Konservat-Lagerstätte, Morocco . Geological Magazine . en . 153 . 3 . 524–536 . 2016GeoM..153..524O . 10.1017/S0016756815000710 . 0016-7568 . 130254788.
  40. Legg . David A. . 2016 . An acercostracan marrellomorph (Euarthropoda) from the Lower Ordovician of Morocco . The Science of Nature . en . 103 . 3–4 . 21 . 10.1007/s00114-016-1352-5 . 26922777 . 2016SciNa.103...21L . 0028-1042.
  41. Laibl . Lukáš . Gueriau . Pierre . Saleh . Farid . Pérez-Peris . Francesc . Lustri . Lorenzo . Drage . Harriet B. . Bath Enright . Orla G. . Potin . Gaëtan J.-M. . Daley . Allison C. . 2023 . Early developmental stages of a Lower Ordovician marrellid from Morocco suggest simple ontogenetic niche differentiation in early euarthropods . Frontiers in Ecology and Evolution . 11 . 10.3389/fevo.2023.1232612 . free . 2296-701X.
  42. Drage . Harriet B. . Legg . David A. . Daley . Allison C. . 2023 . Novel marrellomorph moulting behaviour preserved in the Lower Ordovician Fezouata Shale, Morocco . Frontiers in Ecology and Evolution . 11 . 10.3389/fevo.2023.1226924 . free . 2296-701X.
  43. Lustri . Lorenzo . Gueriau . Pierre . Daley . Allison C. . 2024-05-07 . Lower Ordovician synziphosurine reveals early euchelicerate diversity and evolution . Nature Communications . en . 15 . 1 . 3808 . 10.1038/s41467-024-48013-w . 38714651 . 2041-1723. 11076625 . 2024NatCo..15.3808L .
  44. Pérez-Peris . Francesc . Laibl . Lukáš . Lustri . Lorenzo . Gueriau . Pierre . Antcliffe . Jonathan B . Bath Enright . Orla G . Daley . Allison C . 2021 . A new nektaspid euarthropod from the Lower Ordovician strata of Morocco . Geological Magazine . en . 158 . 3 . 509–517 . 10.1017/S001675682000062X . 2021GeoM..158..509P . 0016-7568.
  45. Lerosey-Aubril . Rudy . Skabelund . Jacob . Ortega-Hernández . Javier . 2020-04-09 . Revision of the mollisoniid chelicerate(?) Thelxiope, with a new species from the middle Cambrian Wheeler Formation of Utah . PeerJ . en . 8 . e8879 . 10.7717/peerj.8879 . free . 32296605 . 7151752 . 2167-8359.
  46. Lefebvre . Bertrand . Allaire . Ninon . Guensburg . Thomas E. . Hunter . Aaron W. . Kouraïss . Khaoula . Martin . Emmanuel L.O. . Nardin . Elise . Noailles . Fleur . Pittet . Bernard . Sumrall . Colin D. . Zamora . Samuel . 2016 . Palaeoecological aspects of the diversification of echinoderms in the Lower Ordovician of central Anti-Atlas, Morocco . Palaeogeography, Palaeoclimatology, Palaeoecology . en . 460 . 97–121 . 10.1016/j.palaeo.2016.02.039. 2016PPP...460...97L .
  47. Lefebvre . Bertrand . Guensburg . Thomas E. . Martin . Emmanuel L.O. . Mooi . Rich . Nardin . Elise . Nohejlová . Martina . Saleh . Farid . Kouraïss . Khaoula . El Hariri . Khadija . David . Bruno . 2019 . Exceptionally preserved soft parts in fossils from the Lower Ordovician of Morocco clarify stylophoran affinities within basal deuterostomes . Geobios . en . 52 . 27–36 . 10.1016/j.geobios.2018.11.001. 2019Geobi..52...27L .
  48. Saleh . Farid . Lefebvre . Bertrand . Dupichaud . Christophe . Martin . Emmanuel L.O. . Nohejlová . Martina . Spaccesi . Léa . 2023 . Skeletal elements controlled soft-tissue preservation in echinoderms from the Early Ordovician Fezouata Biota . Geobios . en . 81 . 51–66 . 10.1016/j.geobios.2023.08.001. 2023Geobi..81...51S .
  49. Dupichaud . Christophe . Lefebvre . Bertrand . Milne . Claire H. . Mooi . Rich . Nohejlová . Martina . Roch . Renaud . Saleh . Farid . Zamora . Samuel . 2023 . Solutan echinoderms from the Fezouata Shale Lagerstätte (Lower Ordovician, Morocco): diversity, exceptional preservation, and palaeoecological implications . Frontiers in Ecology and Evolution . 11 . 10.3389/fevo.2023.1290063 . free . 2296-701X.
  50. Allaire . Ninon . Lefebvre . Bertrand . Nardin . Elise . Martin . Emmanuel L.O. . Vaucher . Romain . Escarguel . Gilles . 2017 . Morphological disparity and systematic revision of the eocrinoid genus Rhopalocystis (Echinodermata, Blastozoa) from the Lower Ordovician of the central Anti-Atlas (Morocco) . Journal of Paleontology . en . 91 . 4 . 685–714 . 10.1017/jpa.2017.6 . 2017JPal...91..685A . 0022-3360. 11336/44871 . free .
  51. Blake . Daniel B. . Hotchkiss . Frederick H. C. . 2022-08-01 . Origin of the subphylum Asterozoa and redescription of a Moroccan Ordovician somasteroid . Geobios . 72-73 . 22–36 . 10.1016/j.geobios.2022.07.002 . 2022Geobi..72...22B . 0016-6995. free .
  52. Hunter . Aaron W. . Ortega-Hernández . Javier . January 2021 . A new somasteroid from the Fezouata Lagerstätte in Morocco and the Early Ordovician origin of Asterozoa . Biology Letters . en . 17 . 1 . 20200809 . 10.1098/rsbl.2020.0809 . 1744-9561 . 7876607 . 33465330.
  53. Polechová . Marika . October 2016 . The bivalve fauna from the Fezouata Formation (Lower Ordovician) of Morocco and its significance for palaeobiogeography, palaeoecology and early diversification of bivalves . Palaeogeography, Palaeoclimatology, Palaeoecology . en . 460 . 155–169 . 2016PPP...460..155P . 10.1016/j.palaeo.2015.12.016.
  54. Kröger . Björn . Lefebvre . Bertrand . August 2012 . Palaeogeography and palaeoecology of early Floian (Early Ordovician) cephalopods from the Upper Fezouata Formation, Anti-Atlas, Morocco . Fossil Record . en . 15 . 2 . 61–75 . 10.1002/mmng.201200004 . free. 2012FossR..15...61K .
  55. Vinther . Jakob . Parry . Luke . Briggs . Derek E. G. . Van Roy . Peter . February 2017 . Ancestral morphology of crown-group molluscs revealed by a new Ordovician stem aculiferan . Nature . en . 542 . 7642 . 471–474 . 10.1038/nature21055 . 28166536 . 2017Natur.542..471V . 0028-0836. 1983/70368bb0-c2df-490a-8b45-26e7b4397596 . 205253410 . free .
  56. Ebbestad . Jan Ove R. . October 2016 . Gastropoda, Tergomya and Paragastropoda (Mollusca) from the Lower Ordovician Fezouata Formation, Morocco . Palaeogeography, Palaeoclimatology, Palaeoecology . en . 460 . 87–96 . 2016PPP...460...87E . 10.1016/j.palaeo.2016.01.003.
  57. Ebbestad . Jan Ove R. . Lefebvre . Bertrand . November 2015 . An unusual onychochilid mollusc from the Ordovician (Tremadocian) Fezouata Formation, Morocco . Geobios . en . 48 . 6 . 427–438 . 2015Geobi..48..427E . 10.1016/j.geobios.2015.09.004.
  58. Lefebvre . Bertrand . El Hariri . Khadija . Lerosey-Aubril . Rudy . Servais . Thomas . Van Roy . Peter . 2016 . The Fezouata Shale (Lower Ordovician, Anti-Atlas, Morocco): A historical review . Palaeogeography, Palaeoclimatology, Palaeoecology . en . 460 . 7–23 . 10.1016/j.palaeo.2015.10.048. 2016PPP...460....7L .
  59. Lehnert . Oliver . Nowak . Hendrik . Sarmiento . Graciela N. . Gutiérrez-Marco . Juan Carlos . Akodad . Mustapha . Servais . Thomas . October 2016 . Conodonts from the Lower Ordovician of Morocco — Contributions to age and faunal diversity of the Fezouata Lagerstätte and peri-Gondwana biogeography . Palaeogeography, Palaeoclimatology, Palaeoecology . en . 460 . 50–61 . 2016PPP...460...50L . 10.1016/j.palaeo.2016.03.023.
  60. Gutiérrez-Marco . Juan Carlos . Martin . Emmanuel L.O. . October 2016 . Biostratigraphy and palaeoecology of Lower Ordovician graptolites from the Fezouata Shale (Moroccan Anti-Atlas) . Palaeogeography, Palaeoclimatology, Palaeoecology . en . 460 . 35–49 . 2016PPP...460...35G . 10.1016/j.palaeo.2016.07.026.
  61. Nanglu . Karma . Waskom . Madeleine E. . Richards . Jared C. . Ortega-Hernández . Javier . 2023-10-11 . Rhabdopleurid epibionts from the Ordovician Fezouata Shale biota and the longevity of cross-phylum interactions . Communications Biology . en . 6 . 1 . 1002 . 10.1038/s42003-023-05377-x . 2399-3642 . 10567727 . 37821659.
  62. Muir . L A . Gutiérrez-Marco . J C . 2023 . https://kirj.ee/earth-publications/?filter[year=2023&filter[issue]=1468&filter[publication]=12591 A new species of the problematic organism Webbyites from the Early Ordovician Fezouata Biota of Morocco ]. Estonian Journal of Earth Sciences . en . 72 . 1 . 74 . 10.3176/earth.2023.24 . 1736-4728. free .
  63. Botting . Joseph P. . 2016 . Diversity and ecology of sponges in the Early Ordovician Fezouata Biota, Morocco . Palaeogeography, Palaeoclimatology, Palaeoecology . en . 460 . 75–86 . 10.1016/j.palaeo.2016.05.018. 2016PPP...460...75B .
  64. Van Iten . Heyo . Muir . Lucy . Simões . Marcello G. . Leme . Juliana M. . Marques . Antonio C. . Yoder . Naomi . October 2016 . Palaeobiogeography, palaeoecology and evolution of Lower Ordovician conulariids and Sphenothallus (Medusozoa, Cnidaria), with emphasis on the Fezouata Shale of southeastern Morocco . Palaeogeography, Palaeoclimatology, Palaeoecology . en . 460 . 170–178 . 2016PPP...460..170V . 10.1016/j.palaeo.2016.03.008 . free . 11449/161915.
  65. Martin . Emmanuel L.O. . Lerosey-Aubril . Rudy . Vannier . Jean . October 2016 . Palaeoscolecid worms from the Lower Ordovician Fezouata Lagerstätte, Morocco: Palaeoecological and palaeogeographical implications . Palaeogeography, Palaeoclimatology, Palaeoecology . en . 460 . 130–141 . 2016PPP...460..130M . 10.1016/j.palaeo.2016.04.009.
  66. Kouraiss . Khaoula . El Hariri . Khadija . El Albani . Abderrazak . Azizi . Abdelfattah . Mazurier . Arnaud . Vannier . Jean . 2018 . X-ray microtomography applied to fossils preserved in compression: Palaeoscolescid worms from the Lower Ordovician Fezouata Shale . Palaeogeography, Palaeoclimatology, Palaeoecology . en . 508 . 48–58 . 10.1016/j.palaeo.2018.07.012. 2018PPP...508...48K .
  67. Valent . Martin . 2015-10-31 . Pauxillites thaddei a new lower Ordovician hyolith from Morocco . Acta Musei Nationalis Pragae, Series B, Historia Naturalis . en . 51–54 . 10.14446/AMNP.2015.51 . 1804-6479.
  68. Vinther . Jakob . Van Roy . Peter . Briggs . Derek E. G. . 2008 . Machaeridians are Palaeozoic armoured annelids . Nature . en . 451 . 7175 . 185–188 . 10.1038/nature06474 . 18185586 . 2008Natur.451..185V . 1476-4687.
  69. Parry . Luke A. . Edgecombe . Gregory D. . Sykes . Dan . Vinther . Jakob . 2019-07-24 . Jaw elements in Plumulites bengtsoni confirm that machaeridians are extinct armoured scaleworms . Proceedings of the Royal Society B: Biological Sciences . en . 286 . 1907 . 20191247 . 10.1098/rspb.2019.1247 . 0962-8452 . 6661337 . 31337310.
  70. Nowak . Hendrik . Servais . Thomas . Pittet . Bernard . Vaucher . Romain . Akodad . Mustapha . Gaines . Robert R. . Vandenbroucke . Thijs R.A. . 2016 . Palynomorphs of the Fezouata Shale (Lower Ordovician, Morocco): Age and environmental constraints of the Fezouata Biota . Palaeogeography, Palaeoclimatology, Palaeoecology . en . 460 . 62–74 . 10.1016/j.palaeo.2016.03.007. 2016PPP...460...62N .