Manuscrit accepté / Accepted manuscript
New thylacocephalans from the Cretaceous Lagerstätten of Lebanon
Sylvain CHARBONNIER, Giorgio TERUZZI, Denis AUDO, Maxime LASSERON, Carolin HAUG and Joachim T. HAUG
Reçu le /Received date: 06/10/16 Accepté le /Accepted date: 10/11/16
Prière de citer l’article de la façon suivante / Please cite this article as:
CHARBONNIER S., TERUZZI G., AUDO D., LASSERON M., HAUG C. and HAUG J.T. (2017). – New thylacocephalans from the Cretaceous Lagerstätten of Lebanon. – Bulletin de la Société géologique de France, 188, n° thématique (sous presse).
1 New thylacocephalans from the Cretaceous Lagerstätten of Lebanon
2
3 SYLVAIN CHARBONNIER1, GIORGIO TERUZZI2, DENIS AUDO3,
4 MAXIME LASSERON1, CAROLIN HAUG4, JOACHIM T. HAUG4
5
6 1Muséum national d’Histoire naturelle, Paris ; Centre de Recherche sur la Paléobiodiversité et les
7 Paléoenvironnements (CR2P, UMR 7207), Sorbonne Universités, MNHN, UPMC, CNRS, 57 rue Cuvier F-75005
8 Paris (France). [email protected]
9
10 2Museo di Storia Naturale di Milano, Sezione di Paleontologia degli Invertebrati, Corso Venezia 55, 20121 Milano
11 (Italy). [email protected]
12
13 3Université de Rennes 1, Géosciences Rennes UMR 6118, Campus de Beaulieu, 35042 Rennes (France).
15
16 4LMU Munich, Department of Biology II and GeoBio-Center, Großhaderner Str. 2, 82152 Martinsried-Planegg,
17 (Germany). [email protected] 18 BSGF
19
20 Key-words. – Euarthropoda, Thylacocephala, new genera, Cenomanian, Hakel, Hadjoula, Sahel Alma, Lebanon.
21
22 Abstract. – Thylacocephalans (Euarthropoda, Thylacocephala) are characterized by their “bivalved” carapace and
23 three anterior prehensile appendages. It is still not clear how they used to live, or what their evolutionary history
24 is. This study focuses on new thylacocephalans from the Late Cretaceous Konservat-Lagerstätten of Lebanon,
25 which yielded the youngest representatives of the group. Three new genera and species are described in the
26 Cenomanian sublithographic limestones of Hakel and Hadjoula, and two new genera and one new species are
27 described in the Santonian chalky limestones of Sahel Alma. Among the specimens from Hakel and Hadjoula,
28 Paradollocaris vannieri, Thylacocaris schrami and Globulacaris garassinoi are the first reports of
29 thylacocephalans in the Cenomanian of Lebanon. Paradollocaris and Thylacocaris are assigned to Dollocarididae
1 ACCEPTED MANUSCRIPT 1 based upon their large optic notches limited by rostral and antero-ventral processes, their hypertrophied eyes, and
2 their posterior notches with dorsal and ventral spines. Moreover, Thylacocaris presents a very peculiar character:
3 an optic notch with two strong optic spines protecting the eye. Globulocaris is assigned to Protozoeidae based
4 upon its small carapace with a distinct dorsal notch anterior to a strong postero-dorsal spine. Among the specimens
5 from Sahel Alma, Keelicaris deborae is a new form of thylacocephalans in the Santonian of Lebanon. It presents
6 a very unusual keel-shaped carapace with terraces and punctuations, and is assigned to Microcarididae. The new
7 genus Hamaticaris, presenting a very peculiar hooked rostrum, is also erected for Protozoea damesi Roger, 1946.
8 These two species add to the well-known thylacocephalans from Sahel Alma: Pseuderichtus cretaceus Dames,
9 1886, Protozoea hilgendorfi Dames, 1886 and Thylacocephalus cymolopos Lange, Hof, Schram & Steeman, 2001.
10 The occurrence of such diverse fauna of thylacocephalans markedly increases the diversity of the group during the
11 Late Cretaceous. The diversity and abundance of the Sahel Alma thylacocephalans pose also the problem of causes
12 of their disappearance from the fossil record after the Santonian.
13
14
15 Nouveaux thylacocéphales des Lagerstätten du Crétacé du Liban
16
17 Mots-clés. – Euarthropoda, Thylacocephala, nouveaux genres, Cénomanien, Hakel, Hadjoula, Sahel Alma, Liban 18 BSGF 19 Résumé. – Les thylacocéphales (Euarthropoda, Thylacocephala) sont des euarthropodes caractérisés par leur
20 carapace « bivalve » et leurs trois appendices préhensiles antérieurs. Leur mode de vie et leur histoire évolutive
21 sont encore loin d’être bien connus. La présente étude documente de nouveaux thylacocéphales issus des
22 Konservat-Lagerstätten du Crétacé supérieur du Liban qui ont livré les représentants les plus récents du groupe.
23 Trois nouveaux genres et espèces sont décrits dans les calcaires sublithographiques cénomaniens de Hakel et de
24 Hadjoula, et deux nouveaux genres et une nouvelle espèce sont décrits dans les calcaires crayeux santoniens de
25 Sahel Alma. Parmi les spécimens de Hakel et de Hadjoula, Paradollocaris vannieri, Thylacocaris schrami et
26 Globulacaris garassinoi sont les premières mentions de thylacocéphales dans le Cénomanien du Liban.
27 Paradollocaris et Thylacocaris sont assignés aux Dollocarididae sur la base de leurs larges encoches optiques
28 limitées par des processus rostraux et antéro-ventraux, de leurs yeux hypertrophiés, et de leurs encoches
29 postérieures avec des épines dorsales et ventrales. De plus, Thylacocaris présente un caractère très singulier : une
30 encoche optique avec deux grosses épines optiques protégeant l’œil. Globulocaris est assigné aux Protozoeidae
2 ACCEPTED MANUSCRIPT 1 sur la base de sa petite carapace avec une très nette encoche dorsale à l’avant de l’épine postéro-dorsale. Parmi les
2 spécimens de Sahel Alma, Keelicaris deborae est une nouvelle forme de thylacocéphales dans le Santonien du
3 Liban. Possédant une très inhabituelle carapace en forme de quille de navire avec des côtes et des ponctuations, il
4 est assigné aux Microcarididae. Le nouveau genre Hamaticaris, présentant un rostre crochu très particulier, est
5 aussi érigé pour Protozoea damesi Roger, 1946. Ces deux espèces s’ajoutent aux thylacocéphales bien connus de
6 Sahel Alma : Pseuderichtus cretaceus Dames, 1886, Protozoea hilgendorfi Dames, 1886 et Thylacocephalus
7 cymolopos Lange, Hof, Schram & Steeman, 2001. La présence d’une faune aussi variée de thylacocéphales
8 augmente la diversité du groupe durant le Crétacé supérieur. La diversité et l’abondance des thylacocéphales de
9 Sahel Alma posent aussi le problème des causes de leur disparition du registre fossile après le Santonien.
10
11
12 INTRODUCTION
13
14 Thylacocephalans are among the most intriguing euarthropods of the Palaeozoic and Mesozoic
15 eras. They have a long stratigraphic record ranging from the Silurian [Haug et al., 2014] and
16 possibly earlier [Vannier et al., 2006: Cambrian] to the Late Cretaceous [Dames, 1886; Schram
17 et al., 1999], when they probably became extinct. They have a high palaebiogeographic
18 distribution and are known from allBSGF continents except An tarctica and South America [Hegna et
19 al., 2014]. They are notably known from Scotland, Spain, France, Germany, Italy, Austria,
20 Slovenia, Lebanon, Madagascar, China, Australia, United States, Mexico, and more recently
21 from Japan [Ehiro et al., 2015].
22 Their anatomy, mode of life and phylogenetical affinities have remained largely unresolved
23 [Rolfe, 1985; Schram et al., 1999; Lange et al., 2001; Vannier et al., 2006; Schram, 2014;
24 Vannier et al., 2016]. Thylacocephans are characterized by a metamerized body protected by a
25 sclerotized “bivalved” carapace (in the wide sense), a pair of compound eyes, sometimes very
26 large, and three pairs of prominent prehensile appendages with spiny tips converging towards
27 the mouth. In the case of some Jurassic taxa such as Dollocaris Van Straelen, 1923, these
3 ACCEPTED MANUSCRIPT 1 appendages distantly recall the raptorial appendages of extant mantis shrimps (stomatopod
2 crustaceans) and suggest predatory and hunting habits [Charbonnier, 2009; Charbonnier et al.,
3 2010; Vannier et al., 2016].
4 Historically, thylacocephalans have been tentatively assigned to a great variety of crustacean
5 groups (e.g., stomatopods: Fraas [1878], Hilgendorf [1885], Roger [1946], decapods: Secrétan
6 [1985], cirripedes: Arduini et al. [1980]). More recently, Haug et al. [2014] have proposed a
7 sister-group relationship with Remipedia based on morphological similarities between
8 Thylacares from the Waukesha fauna (Silurian, Wisconsin) and extant remipedes. These
9 resemblances concern the multisegmented and undifferentiated nature of the trunk the number
10 and detailed morphology of the sub-chelate raptorial appendages. According to Vannier et al.
11 [2016], this tentative placement of Thylacocephala close to remipedes is questionable given
12 major differences between the two groups in terms of body organisation and exoskeletal
13 structure. As the time of writing, the exact affinities of Thylacocephala still remain an open
14 question.
15 This paper reports new occurrences of thylacocephalans in the Cretaceous Konservat- 16 Lagerstätten from Lebanon. It increasesBSGF the palaeobiodiversity of last representatives of the 17 group during the Cenomanian (Hakel, Hadjoula) and the Santonian (Sahel Alma) just before
18 their assumed extinction. In addition, the new species provide new insights on the morphology
19 of Thylacocephala that may prove useful to understand the affinities and ecology of these
20 euarthropods.
21
22
4 ACCEPTED MANUSCRIPT 1
2 GEOLOGICAL SETTING
3
4 The three classical Konservat-Lagerstätten of Lebanon, Hakel, Hadjoula and Sahel Alma
5 (fig. 1), contain a remarkably preserved marine fauna dominated by fishes (Actinopterygii,
6 Chondrichthyes: Forey et al. [2003], Gayet et al. [2003]) and euarthropods (Crustacea,
7 Merostomata, Thylacocephala), associated with numerous soft-bodied organisms (Coleoidea,
8 Polychaeta). Hakel and Hadjoula are both late Cenomanian in age [Hemleben, 1977; Wippich
9 & Lehmann, 2004]. Sahel Alma is late Santonian in age [Roger, 1946; Ejel & Dubertret, 1966],
10 and is currently inaccessible, being in private property covered by olive trees and possibly
11 entirely exhausted.
12 Hadjoula and Hakel. — These two Lagerstätten are located in northwest Lebanon, ca
13 10 km east of Byblos (Jbail). They correspond to sublithographic limestones probably deposited
14 in small shallow basins comprising intra-shelf depressions (see review in Audo & Charbonnier
15 [2012]). The sublithographic limestones are intensely excavated for fossils which are sold to 16 tourists and collectors but also forBSGF scientific studies. By far the most abundant animals found in 17 the Hadjoula and Hakel biota are the fishes (in the wide sense) with a prolific and varied
18 assemblage of chondrichthyans and osteichthyans, both found in large numbers on bedding
19 planes [Gayet et al., 2003]. Euarthropods are also common, represented by decapod crustaceans
20 [Glaessner, 1945; Roger, 1946; Garassino, 1994, 2001; Haug et al., 2016; Charbonnier et al.,
21 in press], stomatopods [Ahyong et al., 2007], isopods (unpublished) and thylacocephalans (this
22 work). The euarthropods are most frequently preserved as compressions on the surface of
23 sublithographic limestone.
24 Sahel Alma. — The Sahel Alma Lagerstätte is located in northwest Lebanon, ca 20 km
25 northeast of Beirut. The “fish-beds” of Sahel Alma correspond to chalky laminated limestones
5 ACCEPTED MANUSCRIPT 1 formed in much deeper environments than those recognized in Hadjoula or Hakel. They were
2 deposited during an acceleration of the subsidence of the Arabian craton. This acceleration of
3 subsidence is probably a consequence of the onset of the collisional trend, which was
4 responsible for the closing of the Tethys [Ferry et al., 2007]. During a field trip to Lebanon in
5 2011, survey in the vicinity of Sahel Alma did not result in the discovery of any traces of the
6 outcrop. Apparently, the outcrop is in a private property, where some years ago an olive grove
7 was planted. The outcrop is therefore not available for new excavations anymore.
8 The Sahel Alma fauna comprises numerous chondrichthyans and osteichthyans [Gayet et al.,
9 2003], cephalopods [Roger, 1946; Jattiot et al., 2015] and rare annelids [Bracchi &
10 Alessandrello, 2005]. Euarthropods are the most common fossils, represented by decapod
11 crustaceans [Brocchi, 1875; Roger, 1946; Garassino, 1994, 2001; Audo & Charbonnier, 2013;
12 Charbonnier et al., in press], cirripedes [Gale, 2016], isopods [Feldmann, 2009; Feldmann &
13 Charbonnier, 2011] and dominated by thylacocephalans [Dames, 1886; Schram et al., 1999;
14 Lange et al., 2001].
15 16 BSGF 17 MATERIAL AND METHODS
18
19 The examined thylacocephalans come from Hakel (1 specimen), Hadjoula (23 specimens), and
20 Sahel Alma (92 specimens). Most of the Cenomanian (Hakel, Hadjoula) specimens were
21 collected during a field trip in 2011 organized by the Paris Museum and conducted by one of
22 the authors (SC). Santonian (Sahel Alma) specimens were collected during the expedition of
23 the professor Camille Arambourg in Syria and Iran (1938-1939). They are all housed in the
24 palaeontological collections of the Muséum national d’Histoire naturelle, Paris, France
25 (acronym: MNHN). Other specimens are housed in the Museo di Storia Naturale in Milano,
6 ACCEPTED MANUSCRIPT 1 Italy (acronym: MSNM) and were collected by Alessandro Garassino or donated by private
2 collectors.
3 Thylacocephalans from Hakel and Hadjoula are all preserved flattened on sublithographic
4 limestones. The cuticle is relatively well preserved but is generally microfractured due to the
5 compaction. An important recrystallization of the fossil can also occur, however infrequently.
6 Their aspect varies from whitish on cream-coloured sediment to red-brownish, this latter case
7 possibly corresponding to weathering of the fossil. Thylacocephalans (and also crustaceans) are
8 distinctly autofluorescent under UV light (appear yellow) and green light (appear light orange).
9 The specimens were therefore documented under macro-fluorescence settings and also under
10 cross-polarised light; both methods enhance the contrast between fossil and surrounding matrix
11 [for details see Haug et al. 2009, 2011, 2012]. Some of the best-preserved specimens were
12 partially prepared manually with a fine needle to allow a more detailed description.
13 The specimens from Sahel Alma are preserved within a soft, chalky limestone. UV
14 autofluorescence of the specimens (and also crustaceans) is generally faint or absent and can
15 only be observed for some specimens, under the right conditions and with an adapted colour 16 processing of pictures. All the specimensBSGF were studied using a binocular and a camera lucida. 17 Imaging of specimens were variably realized in natural light or UV fluorescence (pictures by
18 L. Cazes, P. Loubry and GT), cross-polarized light (pictures by DA), or green fluorescence
19 (pictures by JTH and CH).
20 Natural light pictures give a good idea of the aspect of the specimen to the naked eye. They are
21 obtained with a diffuse white light source and without polarizing filter. Cross-polarized light
22 pictures are obtained by illuminating specimens with polarized light and cancelling most of the
23 reflected light with a polarizer on the camera lens. This set-up allows to reduce drastically glare
24 and enhance contrast of the specimen [Bengtson, 2000].
7 ACCEPTED MANUSCRIPT 1 UV fluorescence is obtained by illuminating the fossil by a “dark-light” (blue to UV light).
2 Under this kind of light, numerous fossil euarthropods, especially from limestone display a
3 strong yellow autofluorescence. Pictures are digitally white-balanced to ease the visualisation
4 of fluorescence.
5 Green fluorescence works on the same principle as UV fluorescence, in this case, the
6 autofluorescence is light orange [Haug et al., 2009; Haug & Haug, 2011], and the green light
7 has to be cancelled by an orange/red filter to allow the visualisation of the fluorescence. Once
8 again, pictures are colour equilibrated to ease the visualisation of fluorescence.
9
10
11
12 SYSTEMATIC PALAEONTOLOGY
13
14 Preliminary remarks. — In the early 1980s, three higher taxonomic levels were applied to
15 thylacocephalans: the class Thylacocephala Pinna, Arduini, Pesarini & Teruzzi, 1982, class 16 Conchyliocarida Secrétan, 1983, BSGFand order Concavicarida Briggs & Rolfe, 1983. Rolfe [1985] 17 adopted the class Thylacocephala for the entire group, comprising two orders: Conchyliocarida
18 and Concavicarida. This systematic classification was followed by Schram [1990], Schram et
19 al. [1999], and Lange et al. [2001], although the genera included within each order are different
20 in each paper. No family level designations have been made, except for the Austriocarididae
21 Glaessner, 1931. Recently, Schram [2014] attempted, for the first time, to classify the
22 thylacocephalans to the family level. However, Ehiro et al. [2015] considered that no clear
23 definition was given to these families and it seems that further examination is necessary for
24 family-level classification. Even if it is probably perfectible, we follow the classification
25 proposed by Schram [2014].
8 ACCEPTED MANUSCRIPT 1
2
3
4 THYLACOCEPHALA Pinna, Arduini, Pesarini & Teruzzi, 1982
5 Order CONCHYLIOCARIDA Secrétan, 1983
6 Family Dollocarididae Schram, 2014
7
8 Paradollocaris Charbonnier nov. gen.
9 figs 2-5
10
11 Type species. — Paradollocaris vannieri Charbonnier nov. sp.
12
13 Etymology. — A combination of the Latin par, paris (close to) and Dollocaris Van Straelen,
14 1923. The gender of the genus is feminine.
15 16 Diagnosis. — Carapace with largeBSGF optic notch limited by elongate, sharp rostrum and antero- 17 ventral process; staple-like optic notch with straight, strongly inclined margin; rostrum with
18 hook-like distal extremity; dorsal margin with crest extending above the rostrum and interrupted
19 posteriorly; regular, convex ventral margin; large muscle scar; posterior margin with narrow
20 notch limited by dorsal and ventral spines; hypertrophied eye; eight pairs of gills; robust
21 prominent anterior appendages; posterior trunk somites bearing paddle-like limbs.
22
23 Discussion. — Paradollocaris is assigned to Dollocarididae based upon the following
24 morphological characters: large optic notch limited by rostral and antero-ventral processes,
25 hypertrophied eye, posterior notch with dorsal and ventral spines. The general morphology of
9 ACCEPTED MANUSCRIPT 1 the new genus is close to Dollocaris from the La Voulte-sur-Rhône Lagerstätte but some
2 important differences can be reported. Paradollocaris differs from Dollocaris by its staple-like
3 optic notch (strongly concave in Dollocaris), its elongate rostrum (short in Dollocaris), its
4 regular convex ventral margin (more sinuous and with wide concavity before reaching antero-
5 ventral process in Dollocaris), its subcircular muscle scar (ellipsoidal and prominent in
6 Dollocaris), and the absence of longitudinal lateral carina (present in Dollocaris).
7 Paradollocaris differs from Paraostenia Secrétan, 1985 and Mayrocaris Polz, 1994 by the
8 general morphology of its carapace. For instance, Paraostenia possesses a sub-rectangular
9 carapace with a reduced rostrum and almost straight optic and posterior notches (staple-like
10 optic notch with elongate rostrum in Paradollocaris), and a lateral carina associated to an
11 undulate row of tubercles (absent in Paradollocaris). Mayrocaris Polz, 1994 shows a triangular
12 ventral margin with a pronounced concavity before reaching the antero-ventral process (regular,
13 convex ventral margin in Paradollocaris).
14 Paradollocaris differs also from Victoriacaris Hegna, Vega & González-Rodríguez, 2014 and
15 Polzia Hegna, Vega & González-Rodríguez, 2014 by its staple-like optic notch and relatively 16 narrow posterior notch (large andBSGF concave optical and posterior notches in Victoriacaris; 17 concave optic notch and reduced posterior notch in Polzia), by its rostrum with hook-like distal
18 extremity (beveled anterior spine in Victoriacaris; simple spiny rostrum in Polzia), and by its
19 postero-ventral spine (rounded postero-ventral corner in Polzia).
20
21 Paradollocaris vannieri Charbonnier nov. sp.
22 figs 2-5
23
24 Etymology. — The specific epithet honours Jean Vannier, palaeontologist at the University of
25 Lyon, France.
10 ACCEPTED MANUSCRIPT 1
2 Type material. — Holotype MNHN.F.A57231, 16 paratypes MNHN.F.A57232–A57234,
3 A57236–A57239, A57241, A57242, A57246, A57248, A57253, A57258 and MSNM i24983,
4 i27858, i27859.
5
6 Type locality. — Hadjoula, Lebanon, Middle East.
7
8 Type age. — Late Cretaceous, Cenomanian.
9
10 Description. — Bivalved carapace, subtrapezoidal (holotype: length: ca 17.5 mm; height: ca
11 7.5 mm), laterally compressed, with very large optic notch and narrower posterior notch;
12 convex dorsal margin separating carapace in two identical valves, with a crest prolonged
13 frontally by the rostrum and interrupted in the posterior first quarter of carapace; staple-like
14 optic notch, limited by elongate, sharp rostrum and pointed antero-ventral process; optic notch
15 with straight margin, inclined at about 50° angle to longitudinal axis of carapace; elongate 16 rostrum with hook-like distal extremity;BSGF regular, convex ventral margin divided into two 17 sections of almost equal length: antero-ventral section slightly convex and postero-ventral one
18 straight and inclined at about 130° angle to longitudinal axis of carapace; muscle scar appearing
19 as large subcircular protuberance (diameter: about one third of carapace height) in antero-
20 ventral margin; muscle scar clearly in connexion with the basal level of anterior gills; posterior
21 margin with relatively narrow, concave notch, limited by dorsal and ventral longitudinal spines;
22 hypertrophied eye protruding through orbital notch; eight pairs of lamellate or phyllobranchiate
23 gills; prominent anterior appendages (prehensile or raptorial appendages) poorly preserved but
24 apparently very robust; posterior trunk somites bearing about 16 paddle-like limbs;
11 ACCEPTED MANUSCRIPT 1 ornamentation of carapace with circular punctuations possibly arranged in a pattern, probably
2 irregular vertical rows (not clearly visible due to the preservation).
3
4 Discussion. — Examined specimens of Paradollocaris vannieri vary in proportion, some are
5 slightly longer (Fig. 2A-F), some shorter (Fig. 3A-F). Close observations of shorter specimens
6 reveal in some cases wrinkles on the surface of the fossil. These wrinkles seem to indicate that
7 the fossil was compressed along its longitudinal axis, resulting in a small deformation. Taking
8 into account that some of the variations in proportion within our sample are linked to
9 deformation, and without other characters to distinguish the various specimens, we consider
10 that our sample is monospecific. However, some of the observed variations may also have a
11 biological cause and correspond to intraspecific variations, for instance sexual dimorphism. It
12 is unfortunately not possible to test these ideas considering the preservation of the specimens
13 and our current poor understanding of the thylacocephalan biology.
14 BSGF
12 ACCEPTED MANUSCRIPT 1 Thylacocaris Audo & Charbonnier nov. gen.
2 figs 6-8
3
4 Type species. — Thylacocaris schrami Audo & Charbonnier nov. sp.
5
6 Etymology. — Derived from the name of the class Thylacocephala. The gender of the genus is
7 feminine.
8
9 Diagnosis. — Elongate carapace with lateral outline almost elliptical; large optic notch limited
10 by, elongate, sharp rostrum and short antero-ventral process; slightly concave optic notch with
11 straight, strongly inclined margin, bearing two strong optic spines; dorsal margin with crest
12 extending above the rostrum and interrupted posteriorly; regular, slightly convex ventral
13 margin; large muscle scar located medially of ventral margin; posterior margin with narrow
14 notch limited by dorsal and ventral spines; hypertrophied eye; robust prominent anterior
15 appendages; posterior trunk somites bearing numerous paddle-like limbs. 16 BSGF 17 Discussion. — Thylacocaris is assigned to Dollocarididae based upon the following
18 morphological characters: large optic notch limited by rostral and antero-ventral processes,
19 hypertrophied eye, posterior notch with dorsal and ventral spines. Thylacocaris differs from
20 Dollocaris and Paradollocaris by its longitudinally elongate carapace, its slightly concave optic
21 notch (strongly concave in Dollocaris and staple-like in Paradollocaris), its elongate rostrum
22 (short in Dollocaris), its regular, almost straight ventral margin (more sinuous and with wide
23 concavity before reaching antero-ventral process in Dollocaris and convex in Paradollocaris),
24 its large muscle scar (ellipsoidal and prominent in Dollocaris), and the absence of longitudinal
25 lateral carina (present in Dollocaris).
13 ACCEPTED MANUSCRIPT 1 Thylacocaris differs from Paraostenia Secrétan, 1985 and Mayrocaris Polz, 1994 by the
2 general morphology of its carapace (ellipsoidal in lateral outline). For instance, Paraostenia
3 exhibits sub-rectangular carapace bearing optic and posterior notches almost straight and
4 without developed rostrum (concave optic notch with elongate rostrum in Thylacocaris), and a
5 lateral carina associated to an undulate row of tubercles (absent in Thylacocaris). Mayrocaris
6 Polz, 1994 shows a triangular ventral margin with a pronounced concavity before reaching the
7 antero-ventral process (regular, almost straight ventral margin in Thylacocaris).
8 Thylacocaris differs also from Victoriacaris Hegna, Vega & González-Rodríguez, 2014 and
9 Polzia Hegna, Vega & González-Rodríguez, 2014, mainly by the elongate shape of its carapace
10 (subtrapezoidal in Victoriacaris and in Polzia), by its elongate rostrum (beveled anterior spine
11 in Victoriacaris; simple spiny rostrum in Polzia), and by its optic spines (absent in Victoriacaris
12 and in Polzia).
13
14
15 Thylacocaris schrami Audo & Charbonnier nov. sp. 16 BSGFfigs 6-8 17
18 Etymology. — The specific epithet honours Frederick Schram, palaeontologist at the Burke
19 Museum, University of Washington, USA.
20
21 Type material. — Holotype MNHN.F.A57240, five paratypes MNHN.F.A57235, A57244, and
22 MSNM i24785, i25125, i26785.
23
24 Type locality. — Hadjoula, Lebanon, Middle East.
25
14 ACCEPTED MANUSCRIPT 1 Type age. — Late Cretaceous, Cenomanian.
2
3 Description. — Bivalved carapace, laterally compressed, longitudinally elongate (holotype:
4 length: ca 20.8 mm; height: ca 5.6 mm), almost elliptical in lateral outline, with very large optic
5 notch and narrower posterior notch; convex dorsal margin separating carapace in two identical
6 valves, with a crest prolonged frontally by the rostrum and interrupted in the posterior first
7 quarter of carapace; concave optic notch, limited by elongate, sharp rostrum and pointed antero-
8 ventral process; optic notch with straight margin, inclined at about 40° angle to longitudinal
9 axis of carapace, bearing in the dorsal part two strong spines downward inclined (lower spine
10 longer than upper spine); elongate rostrum with pointed distal extremity; regular, convex
11 ventral margin divided into two sections of almost equal length: antero-ventral section almost
12 straight and horizontal, and postero-ventral one straight and inclined at about 120° angle to
13 longitudinal axis of carapace; muscle scar appearing as large subcircular protuberance
14 (diameter: about one third of carapace height) in antero-ventral margin; posterior margin with
15 narrow, staple-like notch, limited by dorsal and ventral longitudinal spines; hypertrophied eye 16 protruding through orbital notch;BSGF lamellate gills, poorly visible in all available specimens; 17 prominent anterior appendages (prehensile or raptorial appendages) poorly preserved but
18 apparently robust; posterior trunk somites bearing paddle-like limbs of unassessable number
19 (numerous, probably about 16?); ornamentation of carapace with circular punctuations
20 uniformly arranged.
21
22 Discussion. — In paratype MNHN.F.B18836 of Thylacocephalus cymolopos, Lange et al.
23 [2001] interpreted articulated appendages found in the optic notch adjacent to the rostrum, as
24 evidence of small antennulae and antennae. Careful examination of this specimen leads us to a
25 different conclusion: the articulation of these two structures is only apparent and linked to the
15 ACCEPTED MANUSCRIPT 1 preservation of the cuticle of the thin carapace. These structures are similar to those observed
2 in the optic notch of Thylacocaris schrami, and most probably correspond to optic spines
3 protecting the hypertrophied eyes. Based on the interpretation proposed by Lange et al. [2001],
4 Schram [2014] supposed that the compound eyes of Thylacocephalus might be not
5 hypertrophied. Our new interpretation, coupled with the evidence of large eyes in Thylacocaris,
6 makes the presence of large eyes in Thylacocephalus possible, and questions its familial
7 placement within Microcarididae Schram, 2014. Comparison with Thylacocaris suggests
8 Thylacocephalus could instead correspond to a Dollocarididae.
9
10
11 Order CONCAVICARIDA Briggs & Rolfe, 1983
12 Family Protozoeidae Schram, 2014
13
14 Globulocaris Teruzzi & Charbonnier nov. gen.
15 fig. 9 16 BSGF 17 Type species. — Globulocaris garassinoi Teruzzi & Charbonnier nov. sp.
18
19 Etymology. — Derived from the Latin globulosus (spherical, rounded), alluding to the globular
20 shape of the carapace. The gender of the genus is feminine.
21
22 Diagnosis. — Small globular carapace, about twice as long as deep; convex dorsal margin
23 prolonged anteriorly by the rostrum and interrupted posteriorly at level of very short concave
24 notch anterior to the postero-dorsal spine; slightly concave optic notch, limited by sharp and
25 short rostrum and rounded antero-ventral process; ventral margin rounded in lateral outline;
16 ACCEPTED MANUSCRIPT 1 posterior margin limited by strong postero-dorsal spine and rounded postero-ventral process;
2 one short posterior spine immediately under postero-dorsal spine, both forming narrow and
3 concave notch; eye relatively large.
4
5 Discussion. — Globulocaris is assigned to Protozoeidae based upon the following
6 morphological characters: small carapace with distinct dorsal notch of posterior portion of
7 carapace anterior to strong postero-dorsal spine, and optic notch not particularly concave with
8 rounded antero-ventral process. Globulocaris differs from Protozoea Dames, 1886,
9 Pseuderichthus Dames, 1886, and Hamaticaris nov. gen. by its globular carapace
10 (subtrapezoidal in Protozea, Pseuderichthus, and Hamaticaris), its short simple rostrum (very
11 elongate in Protozoea; elongate with one infra-rostral spine in Pseuderichthus and
12 Hamaticaris), its dorsal margin with very short concave notch anterior to the postero-dorsal
13 spine (smooth and regular in Protozoea; large notch limited by dorsal spine in Pseuderichthus),
14 and its narrow posterior notch limited by spines (absent in Protozoea and Hamaticaris).
15 Globulocaris differs also from Victoriacaris Hegna, Vega & González-Rodríguez, 2014 and 16 Polzia Hegna, Vega & González-BSGFRodríguez, 2014, mainly by the globular shape of its carapace 17 (subtrapezoidal in Victoriacaris and Polzia), by its short rostrum (beveled anterior spine in
18 Victoriacaris), and by its narrow posterior notch (very large in Victoriacaris; almost absent in
19 Polzia).
20
21
22 Globulocaris garassinoi Teruzzi & Charbonnier nov. sp.
23 fig. 9
24
17 ACCEPTED MANUSCRIPT 1 Etymology. — The specific epithet honours Alessandro Garassino, formerly a palaeontologist
2 at the Museo di Storia Naturale di Milano, Italy.
3
4 Type material. — Holotype by monotypy MSNM i20655.
5
6 Type locality. — Hakel, Lebanon, Middle East.
7
8 Type age. — Late Cretaceous, Cenomanian.
9
10 Description. — Bivalved carapace; small globular, about twice as long as deep (length: 12 mm;
11 height: 7 mm); convex dorsal margin prolonged anteriorly by the rostrum (split in the present
12 specimen: see line drawing) and interrupted posteriorly at level of very short concave notch
13 anterior to the postero-dorsal spine; dorsal margin almost horizontal between the short concave
14 notch and the postero-dorsal spine; optic notch with slightly concave margin, inclined at about
15 90° angle to longitudinal axis of carapace, limited by sharp and short rostrum and rounded 16 antero-ventral process; ventral marginBSGF rounded in lateral outline (split in the present specimen: 17 see line drawing); muscle scar appearing as large circular protuberance (diameter: 3 mm) in
18 antero-ventral margin; posterior margin limited by strong postero-dorsal spine and rounded
19 postero-ventral process; one short posterior spine immediately under postero-dorsal spine, both
20 forming narrow and concave posterior notch; eye poorly preserved but relatively large;
21 prominent anterior appendages not preserved; posterior trunk somites poorly preserved.
22
23
24 Hamaticaris Charbonnier nov. gen.
25 fig. 10
18 ACCEPTED MANUSCRIPT 1
2 Type species. — Protozoea damesi Roger, 1946.
3
4 Etymology. — Derived from the Latin hamatus (hooked), alluding to the hooked rostrum. The
5 gender of the genus is feminine.
6
7 Diagnosis. — Carapace, trapezoidal in lateral outline, with very elongate spiny rostrum and
8 postero-dorsal spine; rostral spine directed slightly upward, accompanied by small anteriorly
9 oriented hook ventral to its base; antero-ventral process with small rounded projection present
10 just below large optic notch; dorsal margin with very short concave notch anterior to the
11 postero-dorsal spine; bifurcate or Y-shaped arrangement of ridges on central aspect of carapace;
12 three parallel, longitudinal rows of punctuations on dorsal aspect of carapace surface; gracile
13 anterior appendages (prehensile appendages); eight posterior trunk somites bearing eight
14 paddle-like limbs.
15 16 Preliminary remarks. — Dames BSGF[1886] erected the genera Protozoea (with P. hilgendorfi as 17 type species) and Pseuderichthus (with Ps. cretaceus as type species). The same fossils had
18 already been described but not named by Hilgendorf [1885], who drew direct comparisons
19 between these fossils and several larvae of extant crustaceans. Dames [1886] also thought that
20 these fossils represented larval forms of unknown genera of stomatopods. This idea that
21 Protozoea and Pseuderichthus were stomatopod larvae was later strongly endorsed by Van
22 Straelen [1938]. Later, Roger [1946] recognized the new form Protozoea damesi and concluded
23 that the two species of Protozoea had to represent probably remains of adult branchiopods. The
24 possibility these genera may be thylacocephalans was first suggested by Arduini et al. [1980:
25 369] and seconded by Pinna et al. [1982: 481]. Schram et al. [1999] reviewed the
19 ACCEPTED MANUSCRIPT 1 thylacocephalan fauna from Lebanon providing very detailed descriptions. However, these
2 authors maintained Roger’s species in Protozoea whereas the respective carapaces of Protozoea
3 hilgendorfi and P. damesi are clearly different and lead us to propose the new genus
4 Hamaticaris.
5
6 Discussion. — Hamaticaris is assigned to Protozoeidae based upon the following
7 morphological characters: very elongate rostrum, carapace much longer than deep, with
8 elongate postero-dorsal spine, and gracile anterior prehensile appendages. Hamaticaris differs
9 from Protozoea Dames, 1886, Pseuderichthus Dames, 1886, and Globulocaris by its
10 trapezoidal carapace (globular in Globulocaris), its very elongate rostrum with hooked infra-
11 rostral spine (very elongate and smooth in Protozoea, short with simple infra-rostral spine in
12 Pseuderichthus, short and simple in Globulocaris), its dorsal margin with very short concave
13 notch anterior to the postero-dorsal spine (smooth and regular in Protozoea; large notch limited
14 by dorsal spine in Pseuderichthus), its antero-ventral process with small rounded projection
15 (absent in three other genera), its regular posterior margin (with narrow posterior notch limited 16 by spines in Pseuderichthus andBSGF Globulocaris), and its Y-shaped arrangement of ridges on 17 central aspect of carapace (absent in three other genera). Hamaticaris differs also from
18 Victoriacaris Hegna, Vega & González-Rodríguez, 2014 and Polzia Hegna, Vega & González-
19 Rodríguez, 2014, mainly by its very elongate rostrum and postero-dorsal spine (beveled anterior
20 spine in Victoriacaris, short rostrum in Polzia), and by its simple posterior margin (with large
21 posterior notch in Victoriacaris).
22
23
24 Hamaticaris damesi (Roger, 1946) nov. comb.
25 fig. 10
20 ACCEPTED MANUSCRIPT 1
2 Protozoe damesi Roger, 1946: 59-61, fig. 49, pl. 6, figs 7-9.
3 Protozoe damesi – Pinna et al. [1982: 481]. — Arduini & Pinna [1989: 5, 32, fig. p.5].
4 Protozoea damesi – Schram et al. [1999: 778-784, fig. 3, pl. 4 & 5].
5 Type material. — Lectotype MNHN.F.B18843 (figured by Roger 1946: pl. 6, fig. 7);
6 24 paralectotypes: MNHN.F.B18842 (figured by Roger 1946: pl. 6, fig. 8), B18844, B18885
7 (figured by Roger 1946: pl. 6, fig. 9), A30652-A30669, A30672, A50518, A50523.
8 The composition of the original type material has been modified and debated by several authors.
9 Schram et al. [1999] stated that Roger [1946] proposed an extensive description of the species
10 but “without referring to specimen numbers”. They also designated specimen
11 MNHN.F.B18842, illustrated by Roger [1946: pl. 6, fig. 8], to be the lectotype, which is then
12 considered to be “the best of the surviving specimens. Of the original three specimens, the
13 counterpart of the holotype is lost”. However, careful examination of the original text reveals
14 that the type material was composed of almost 60 specimens from Sahel Alma and that Roger 15 [1946] did not selected a holotype.BSGF Another confusion comes from Arduini & Pinna [1989: 5, 16 caption of right figure] who fixed specimen MNHN.F.B18843, illustrated by Roger [1946: pl.
17 6, fig. 7], as lectotype [ICZN 1999: art. 74.6]. Thus the lectotype proposed by Schram et al.
18 [1999] is not valid. In conclusion, only 25 specimens from the original type material are still
19 housed in the palaeontological collections of the MNHN, Paris.
20 Type locality. — Sahel Alma, Lebanon, Middle East.
21
22 Type age. — Late Cretaceous, Santonian.
23
24 Description. — Bivalved carapace, trapezoidal in lateral outline, elongate (lectotype: length: ca
25 12 mm; height: ca 4 mm); very elongate rostrum (length about one-quarter the length of the
21 ACCEPTED MANUSCRIPT 1 central part of the carapace); bending at acute angle from the slightly convex dorsal margin,
2 directed slightly upward, accompanied at its base by small hooked infra-rostral spine (“crochet”
3 in Roger [1946], or “hook” in Schram et al. [1999]) pointing anteriorly; very elongate postero-
4 dorsal spine (length about one-quarter the length of the central part of the carapace), prolonging
5 dorsal margin; slightly convex dorsal margin, with small concave notch anterior to the postero-
6 dorsal spine; large, concave optic notch limited by rostral hook and rounded antero-ventral
7 process; antero-ventral process with small rounded, ventrally directed projection (or “knob” in
8 Schram et al. [1999] of the marginal carina, followed directly by a very slight notch; ventral
9 margin parallel to dorsal margin and making a 45° angle upwards to meet the base of the
10 postero-dorsal spine at an abrupt angle; straight posterior margin; bifurcate or Y-shaped
11 arrangement of ridges on central part of carapace (after Schram et al., 1999: “the stem and
12 dorsal fork of the Y is a long, V-like ridge flanked by shallow grooves, and the ventral fork is
13 a much shorter and shallower ridge extending posteriorly towards the ventral margin. The pits
14 in this region also seem to form rows. One row of about four or five pits closely parallels the
15 ventral carina, and another pit row is roughly parallel with the ridges of the Y-shaped structure, 16 but at some small distance from BSGFit”); ornamentation of carapace composed of three parallel, 17 longitudinal rows of punctuations, lying on dorsal part, just above Y-shaped ridges; dorsal row
18 with about 40 punctuations; median and ventral row with about 27 punctuations; eye poorly
19 preserved but relatively large; about eight pairs of gills; three gracile anterior prehensile
20 appendages; eight posterior trunk somites bearing eight paddle-like limbs (description largely
21 inspired by Schram et al., 1999).
22
22 ACCEPTED MANUSCRIPT 1 Family Microcarididae Schram, 1999
2
3 Keelicaris Teruzzi & Charbonnier nov. gen.
4 fig. 11
5
6 Type species. — Keelicaris deborae Teruzzi & Charbonnier nov. sp.
7
8 Etymology. — Derived from the Middle English kele (keel: lowest and principal timber of a
9 ship), alluding to the peculiar shape of the carapace.
10
11 Diagnosis. — Keel-shaped carapace with very narrow ventral margin and convex dorsal
12 margin; short rostrum; longitudinal lateral carina in median part of caparace; muscle scar
13 inserted at base of restricted ventral margin; very elongate, arcuate posterior margin; large, sub-
14 rectilinear optical notch; ornamentation composed of subparallel vertical rugations or terraces
15 (“ribs”), regularly spaced from dorsal margin, and terminating in restricted ventral region, 16 leaving just smooth muscle scarBSGF region; one longitudinal row of punctuations under dorsal 17 margin.
18
19
20
21
22
23
24 Discussion. — The peculiar keel-shaped carapace of Keelicaris with its deep, narrow ventral
25 margin is quite unique with respect to any known thylacocephalan carapace. It can remind the
23 ACCEPTED MANUSCRIPT 1 outline of Pseuderichthus or Victoriacaris, which carapaces are narrower in the ventral area.
2 However, in Keelicaris, the ventral margin is one eighth less long than the dorsal margin, while
3 in Victoriacaris it is half less long than the dorsal margin, and in Pseuderichthus it is a third
4 less long than the dorsal margin (anterior and posterior spines included). Both Pseuderichthus
5 and Victoriacaris have no surface ornamentation except punctuations in Pseuderichthus.
6 After several comparisons, Keelicaris is assigned to Microcarididae based upon its carapace
7 surface with rugations or terraces shared with Microcaris Pinna, 1974, Atropicaris Arduini &
8 Brasca, 1984, Rugocaris Tintori, Bigi, Crugnola & Danini, 1986, Ferrecaris Calzada & Mañé,
9 1993, Thylacocephalus Lange, Hof, Schram & Steeman, 2001, and Kitakamicaris Ehiro,
10 Sasaki, Kano, Nemoto & Kato, 2015. Keelicaris differs from all these genera by the general
11 morphology of its carapace. Concerning the ornamentation, the rugations in Keelicaris are
12 strong and subparallel, deviated at level of the lateral median carina, convergent and interrupted
13 in the muscle scar area (sinuous, imbricated rugations present on all the carapace surface in
14 Atropicaris, Microcaris and Kitakamicaris; sinuous, subparallel rugations present on all the
15 carapace surface in Ferrecaris; rounded, subparallel, widely spaced rugations in Rugocaris; 16 alternating ribs and grooves associatedBSGF to rows of punctuations , interrupted in ventral region – 17 muscle scar area – in Thylacocephalus). The interruption of rugations in the muscle scar area is
18 shared only by Keelicaris and Thylacocephalus but the shape of the ventral margin is very
19 different.
20
21
22 Keelicaris deborae Teruzzi & Charbonnier nov. sp.
23 fig. 11
24
24 ACCEPTED MANUSCRIPT 1 Etymology. — The specific epithet honours Debora Affer who prepared more than
2 1,000 specimens of thylacocephalans from the Sahel Alma Lagerstätte housed at the Museo di
3 Storia Naturale di Milano, Italy.
4
5 Type material. — Holotype by monotypy MSNM i27356.
6
7 Type locality. — Sahel Alma, Lebanon, Middle East.
8
9 Type age. — Late Cretaceous, Santonian.
10
11 Description. — Keel-shaped carapace (length : ca 40 mm: height: 17.5 mm) characterized by
12 deep, subtrapezoidal, ventral margin; short rostrum; very short, narrow ventral margin of about
13 one eighth of the dorsal margin length and located in the anterior half of carapace; rounded
14 muscle scar, inserted at base of restricted ventral margin; convex dorsal margin (posterior end
15 damaged), anteriorly prolonged by short rostrum; one longitudinal row of punctuations under 16 dorsal margin; very elongate, concaveBSGF and arcuate posterior margin, inclined at about 140° 17 angle to longitudinal axis of carapace (posterior end damaged); large, sub-rectilinear optical
18 notch, limited by short rostrum and little-marked antero-ventral process, inclined at about 50°
19 angle to longitudinal axis of carapace; ornamentation composed of subparallel vertical
20 rugations or terraces (“ribs”), regularly spaced from dorsal margin, and terminating in restricted
21 ventral region, leaving just smooth muscle scar region; one longitudinal lateral carina in median
22 part of caparace, superimposed to subvertical rugations; subvertical rugations presenting small
23 deviation at level of dorsal punctuations, and stronger deviation at level of median lateral carina;
24 points of most anterior rugations in ventral region bent backward at level of muscle scar; two
25 ACCEPTED MANUSCRIPT 1 rugations with bifurcate ventral extremity at level of muscle scar; no traces of limbs or soft-
2 parts.
3
4 Taphonomy. — The almost complete carapace is damaged in the posterior region. The two
5 halves are displaced: the dorsal region of the left half is partially visible in connection from the
6 right one which is completely exposed. The two halves are clearly distinguishable thanks to a
7 median dorsal line. The left half is bent and partially lying under the right one; this displacement
8 is probably linked to a partial distortion due to a post mortem deformation.
9
10
11
12 CONCLUSIONS
13
14 The new forms herein described increase our knowledge of the thylacocephalans during the
15 Late Cretaceous (table 1, fig. 12). The Cenomanian genera Paradollocaris, Thylacocaris, and 16 Globulocaris are described for theBSGF first time in this work. It is surprising considering that 17 Hadjoula and Hakel have, for a long time, been particularly renowned for their exquisite
18 preservation of euarthropods – most notably the crustaceans (see Charbonnier et al. [in press]
19 and references therein). The hypertrophied eyes of Paradollocaris and Thylacocaris and the
20 probable large eyes of Globulocaris suggest relatively deep-water organisms. This was already
21 mentioned in other Mesozoic Lagerstätten such as La Voulte-sur-Rhône, France [Charbonnier
22 et al., 2010].
23 The thylacocephalans of Sahel Alma exhibit a remarkable diversity, known just in the Middle
24 Jurassic of La Voulte-sur-Rhône, France [Charbonnier, 2009]. Five genera are present
25 (Protozoea, Pseuderichthus, Thylacocephalus, Hamaticaris, Keelicaris). They are also quite
26 ACCEPTED MANUSCRIPT 1 abundant in term of specimen numbers, at least concerning Protozoea hilgendorfi and
2 Pseuderichtus cretaceus. Based on the chalky nature of the sediment from Sahel Alma and due
3 to biases in collecting specimens in the field, no reliable statistic data are available. But, in the
4 sampled rocks from which the bulk of the Milano collection of thylacocephalans from Sahel
5 Alma was recovered in recent years, they appear to be the most abundant among macro-
6 invertebrates, including far more than 1,000 individuals in roughly 1 m3 of sampled rock. The
7 diversity and abundance of the Sahel Alma thylacocephalans is surprising since they represent
8 the last known occurrence of thylacocephalans in the fossil record.
9 According to Schram et al. [1999], several features of the Sahel Alma thylacocephalans suggest
10 good swimming capacities. Indeed, the rostral and postero-dorsal and spines of Protozoea,
11 Hamaticaris and Thylacocephalus are commonly typical of pelagic and planktic forms amongst
12 extant crustaceans (e.g. spinose larvae or adults of dendrobranchiate shrimps and
13 gnathophausian mysids). Such spines serve to increase buoyancy and reduce turbulent flow
14 around the body. Moreover, the seemingly light and flexible carapaces of Protozoea and
15 Hamaticaris is a feature found in many other thylacocephalans and swimming crustaceans. The 16 new species of thylacocephalansBSGF from Lebanon increase the palaeobiodiversity of the last 17 representatives of the group during the Cenomanian (Hakel, Hadjoula) and the Santonian (Sahel
18 Alma) just before their assumed extinction. However, they do not offer any new insights into
19 the tempo of thylacocephalan extinction at the end of the Cretaceous.
20 Recently, Hegna et al. [2014] reported the first known occurrences of Mesozoic
21 thylacocephalans in the Americas in the Cretaceous Muhi Quarry Lagerstätte. These authors
22 indicated that throughout their entire evolutionary history, thylacocephalans have a high
23 incidence of endemism. This pattern seems to hold true until the end of the Mesozoic because,
24 once again, there are no thylacocephalan species or genera shared between Mexico and the new
27 ACCEPTED MANUSCRIPT 1 species described from Lebanon. It shall be noted that the punctual fossil thylacocephalans may
2 also explain much of this endemism.
3 The same authors also judiciously concluded that only more work on Lagerstätten or new
4 localities will help us understand if the apparent range contraction of thylacocephalans at the
5 end of the Mesozoic is a real phenomenon or an artefact of sampling. The new forms from
6 Lebanon seem to support them: only intensive research and collect will permit to
7 palaeontologists to find new thylacocephalans. In the present case, we sorted more than 50,000
8 slabs of lithographic limestone from Hakel and Hadjoula (Expedition Charbonnier 2011) to find
9 only 25 specimens of thylacocephalans, which are herein described.
10 Finally, the description of these new species of thylacocephalans is another step toward a better
11 understanding of this still enigmatic group.
BSGF
28 ACCEPTED MANUSCRIPT 1 ACKNOWLEDGMENTS
2
3 We thank Frederick R. Schram and an anonymous reviewer for their valuable reviews of this
4 work and for their useful advices. Philippe Loubry and Lilian Cazes (UMR 7207 CR2P)
5 provided excellent photographs of specimens housed in the Paris and Milano museums. We are
6 pleased to acknowledge Jean-Michel Pacaud (MNHN, Direction des collections) for access to
7 the palaeontological collections. Finally, we are indebted to Pierre Abi Saad and Albert Abi
8 Saad (Memory of Time, Byblos, Lebanon), who lead us to the outcrops in Lebanon, received
9 us and provided us most of the specimens studied herein. The field trips of SC and DA received
10 financial support from the Action Transversale du Muséum 2011: « Biodiversité actuelle et
11 fossile. Crises, stress, restaurations et panchronisme : le message systématique ». The visit of
12 JTH to the MNHN, Paris, was funded by a grant from the European Commission’s (FP 6)
13 Integrated Infrastructure Initiative programme SYNTHESYS (FR-TAF 2590).
14
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18 morphology and systematic affinities of the group. – BMC Evolutionary Biology, 14,
19 159. DOI: 10.1186/s12862-014-0159-2.
20 HAUG C, HAUG J.T., WALOSZEK D., MAAS A., FRATTIGIANI R. & LIEBAU S. (2009). – New
21 methods to document fossil from lithographic limestones of southern Germany and
22 Lebanon. – Palaeontologia Electronica 12 (3), 6T, 1-12.
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24 (2012). – A holomorph approach to xiphosuran evolution—a case study on the ontogeny
25 of Euproops. – Development Genes and Evolution, 222, 253-268.
32 ACCEPTED MANUSCRIPT 1 HAUG J.T. & HAUG C. (2011). – Fossilien unter langwelligem Licht: Grün-Orange-Fluoreszenz
2 an makroskopischen Objeckten. – Archaeopteryx 29, 20-23.
3 HAUG J.T., AUDO D., CHARBONNIER S., PALERO F., PETIT G., ABI SAAD P. & HAUG C. 2016. –
4 The evolution of a key character, or how to evolve a slipper lobster. Arthropod Structure
5 & Development, 45, 97-107.
6 HAUG J.T., HAUG C., KUTSCHERA V., MAYER G., MAAS A., LIEBAU S., CASTELLANI C.,
7 WOLFRAM U., CLARKSON E.N.K. & WALOSZEK D. (2011). – Autofluorescence imaging,
8 an excellent tool for comparative morphology. – Journal of Microscopy, 244, 259-272.
9 HEGNA T.A., VEGA F.J. & GONZÁLEZ-RODRÍGUEZ K.A. (2014). – First Mesozoic
10 thylacocephalans (Arthropoda, ?Crustacea; Cretaceous) in the Western Hemisphere,
11 new discoveries from the Muhi Quarry Lagerstätte. – Journal of Paleontology, 88 (3),
12 606-616.
13 HEMLEBEN C. VON (1977). – Rote Tiden und die oberkretazischen Plattenkalke im Libanon. –
14 Neues Jahrbuch für Geologie und Paläontologie, Monatshefte, 1977 (4), 239-255.
15 HILGENDORF F. (1885). – Ueber cretacische Squilliden-Larven vom Libanon. – Sitzungs- 16 Berichte der Gesellschaft BSGFnaturforschender Freunde zu Berlin, 1885 (10) 184-185. 17 ICZN (1999). – International Code of Zoological Nomenclature. International Trust for
18 Zoological Nomenclature. – The Natural Museum, London, v-xxix + 306 p.
19 JATTIOT R., BRAYARD A., FARA E. & CHARBONNIER S. (2015). – Gladius-bearing coleoids from
20 the Upper Cretaceous Lebanese Lagerstätten: diversity, morphology and phylogenetic
21 implications. Journal of Paleontology, 89 (1), 148-167.
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23 the Cretaceous of Lebanon links the Thylacocephala to the Crustacea. – Palaeontology,
24 44 (5), 905-912.
33 ACCEPTED MANUSCRIPT 1 PINNA G. (1974). – I crostacei della fauna triassica di Cene in Val Seriana. Memorie della
2 Società italiana di Scienze naturali e del Museo civico di Storia naturale di Milano, 21
3 (1), 5-34.
4 PINNA G., ARDUINI P., PESARINI C. & TERUZZI G. (1982). – Thylacocephala : una nuova classe
5 di crostacei fossili. – Atti della Società Italiana di Scienze Naturali e del Museo Civico
6 di Storia Naturale in Milano, 123, 469-482.
7 POLZ H. (1994). – Mayrocaris bucculata gen. nov. sp. nov. (Thylacocephala, Conchyliocarida)
8 aus den Solnhofener Plattenkalken. – Archaeopteryx, 12, 35-44.
9 ROGER J. (1946). – Invertébrés des couches à poissons du Crétacé supérieur du Liban. –
10 Mémoires de la Société géologique de France (Nouvelle série), 51, 5-92.
11 ROLFE W.D.I. (1985). – Form and function in Thylacocephala, Conchyliocarida and
12 Concavicarida (?Crustacea): a problem of interpretation. – Transactions of the Royal
13 Society of Edinburgh, 76, 391-399.
14 SCHRAM F.R. (1990). – On Mazon Creek Thylacocephala. – Proceedings of the San Diego
15 Society of Natural History, 3, 1-16. 16 SCHRAM F.R. (2014). – Family levelBSGF classification within Thylacocephala, with comments on 17 their evolution and possible relationships. – Crustaceana, 87 (3), 340-363.
18 SCHRAM F.R., HOF C.H.J. & STEEMAN F.A. (1999). – Thylacocephala (Arthropoda: Crustacea?)
19 from the Cretaceous of Lebanon and implications for thylacocephalan systematics. –
20 Palaeontology, 42 (5), 769-797.
21 SECRÉTAN S. (1983). – Une nouvelle classe fossile dans la super-classe des Crustacés :
22 Conchyliocarida. – Comptes Rendus de l’Académie des Sciences, Paris, 296 (2), 741-
23 743.
34 ACCEPTED MANUSCRIPT 1 SECRÉTAN S. (1985). – Conchyliocarida, a class of fossil crustaceans: relationship to
2 Malacostraca and postulated behaviour. – Transactions of the Royal Society of
3 Edinburgh, 76, 381-389.
4 TINTORI A., BIGI E., CRUGNOLA G. & DANINI G. (1986). – A new Jurassic Thylacocephala
5 Rugocaris indunensis gen. n. sp. n. and its paleoecological significance. – Rivista
6 Italiana di Paleontologia e Stratigrafia, 92 (2), 239-250.
7 VANNIER J., CHEN J.-Y., HUANG D.-Y., CHARBONNIER S. & WANG X.-Q. (2006). – The Early
8 Cambrian origin of thylacocephalan arthropods. – Acta Palaeontologica Polonica, 51
9 (2), 201-214.
10 VANNIER J., SCHOENEMANN B., GILLOT T., CHARBONNIER S. & CLARKSON E. (2016). –
11 Exceptional preservation of eye structure in arthropod visual predators from the Middle
12 Jurassic. – Nature Communications, 7, 10320. DOI: 10.1038/ncomms10320.
13 VAN STRAELEN V. (1923). – Les Mysidacés du Callovien de La Voulte-sur-Rhône (Ardèche).
14 – Bulletin de la Société Géologique de France, 23 (4), 431-439.
15 WIPPICH G. & LEHMANN J. (2004). – Allocrioceras from the Cenomanian (mid-Cretaceous) of 16 the Lebanon and its bearingBSGF on the palaeobiological interpretation of heteromorphic 17 ammonites. – Palaeontology, 47 (5), 1093-1109.
18
19
20
35 ACCEPTED MANUSCRIPT 1
2 CAPTIONS
3
4 TABLE 1. –List of the thylacocephalans from the Konservat-Lagerstätten of Lebanon, with the
5 number of studied specimens, the age and the distribution.
6
7
8 FIG. 1. – Map of Lebanon showing the main fossiliferous localities yielding exceptionally
9 preserved Cenomanian faunas (Hadjoula, Hakel, En Nammoura Lagerstätten) and Santonian
10 faunas (Sahel Alma Lagerstätte).
11
12 FIG. 2. – Paradollocaris vannieri Charbonnier nov. gen, nov. sp. from the Cenomanian of
13 Hadjoula, Lebanon. A-F. holotype MNHN.F.A57231 showing carapace with preserved soft-
14 parts, right lateral view: cross-polarized light (A), line drawing of carapace (B), green-orange
15 fluorescence (C), interpretative line drawing of soft-parts (D); same as C, desaturated (E), or 16 step wise desaturated (F). G. ParatypeBSGF MNHN.F.A57232, carapace, right lateral view, cross- 17 polarized light. H. Paratype MNHN.F.A57233, specimen showing the bivalved carapace, left
18 lateral view, green-orange fluorescence desaturated. I. Paratype MNHN.F.A58238, carapace
19 showing ornamentation composed of punctuations, right lateral view, cross-polarized light. J.
20 Paratype MNHN.F.A57237, fragment of carapace showing posterior part of dorsal carina, right
21 lateral view, green-orange fluorescence desaturated. Abbreviations: avp= antero-ventral
22 process, dc= dorsal carina, dm= dorsal margin, gi= lamellate gills, ms= muscle scar, on= optical
23 notch, pn= posterior notch, pu= punctuation, pds= postero-dorsal spine, pvs= postero-ventral
24 spine, r= rostrum, tl= paddle-like limbs or trunk limbs, ts= trunk somites, vm= ventral margin.
36 ACCEPTED MANUSCRIPT 1 Scale bars: 5 mm. Line drawings: S. Charbonnier. Photographs: D. Audo (A, G, I) and J. Haug
2 (C, E, F, H, J).
3
4 FIG. 3. – Paradollocaris vannieri Charbonnier nov. gen, nov. sp. from the Cenomanian of
5 Hadjoula, Lebanon. A-D. Paratype MNHN.F.A57234, carapace with slight longitudinal
6 compression, right lateral view: cross-polarized light (A), note the punctuations of carapace;
7 green-orange fluorescence desaturated (B-D), close-up of rostrum (C) and close-up of anterior
8 appendages (poorly preserved) (D). E-F. Paratype MNHN.F.A57241, right lateral view: green-
9 orange fluorescence (E), interpretative line-drawing (F). Abbreviations: avp= antero-ventral
10 process, dc= dorsal carina, dm= dorsal margin, gi= lamellate gills, ms= muscle scar, on= optical
11 notch, pn= posterior notch, pu= punctuation, r= rostrum, rap= prominent anterior appendages,
12 tl= paddle-like limbs or trunk limbs, vm= ventral margin. Scale bars: 5 mm (A, B, E, F) and 2
13 mm (C, D). Line drawing: S. Charbonnier. Photographs: D. Audo (A) and J. Haug (B-E).
14
15 FIG. 4. – Paradollocaris vannieri Charbonnier nov. gen, nov. sp. from the Cenomanian of 16 Hadjoula, Lebanon. A-D. ParatypeBSGF MNHN.F.A57236, showing carapace with preserved soft- 17 parts, left lateral view: cross-polarized light (A), interpretative line drawing of carapace and
18 soft-parts (B), green-orange fluorescence (C), green-orange fluorescence step-wise desaturated
19 (D). E. Paratype MSNM i24983, left lateral view, green-orange fluorescence step-wise
20 desaturated. F. Paratype MNHN.F.A57248 showing carapace and soft-parts (eye and lamellate
21 gills), left lateral view, cross-polarized light. G-H. Paratype MNHN.F.A57246, carapace, right
22 lateral view (G) and line drawing (H). Abbreviations: avp= antero-ventral process, dc= dorsal
23 carina, dm= dorsal margin, gi= lamellate gills, ms= muscle scar, on= optical notch, pn=
24 posterior notch, pu= punctuation, pds= postero-dorsal spine, pvs= postero-ventral spine, r=
37 ACCEPTED MANUSCRIPT 1 rostrum, tl= paddle-like limbs or trunk limbs, vm= ventral margin. Scale bars: 5 mm. Line
2 drawings: S. Charbonnier. Photographs: D. Audo (A, F, G) and J. Haug (C-E).
3
4 FIG. 5. – Paradollocaris vannieri Charbonnier nov. gen, nov. sp. from the Cenomanian of
5 Hadjoula, Lebanon. A. Paratype MNHN.F.A57241, close-up of carapace showing punctuations
6 and lamellate gills, green-orange fluorescence step-wise desaturated. B. Paratype
7 MNHN.F.A57239, posterior part of specimen showing fragments of robust prominent anterior
8 appendages, right lateral view, green-orange fluorescence desaturated. C-D. Paratype MSNM
9 i27859, left lateral view, carapace with fragments of prominent anterior appendages, UV light
10 (C), and close-up of prominent anterior appendages (D), green-orange fluorescence desaturated.
11 Abbreviations: avp= antero-ventral process, dc= dorsal carina, dm= dorsal margin, gi=
12 lamellate gills, ms= muscle scar, on= optical notch, pn= posterior notch, pu= punctuation, pds=
13 postero-dorsal spine, pvs= postero-ventral spine, rap= prominent anterior appendages, vm=
14 ventral margin. Scale bars: 2 mm (A, D) and 5 mm (B, C). Photographs: J. Haug (A, B, C) and
15 P. Loubry (C). 16 BSGF 17 FIG. 6. – Thylacocaris schrami Audo & Charbonnier nov. gen, nov. sp. from the Cenomanian
18 of Hadjoula, Lebanon. A-D. Holotype MNHN.F.A57240, left lateral view, cross-polarized light
19 (A), line-drawing of carapace, and and soft-parts (B), carapace and fragments of anterior
20 prehensile appendages, green-orange fluorescence desaturated (C), close-up of optical notch
21 showing broken rostrum and two optical spines, green-orange fluorescence desaturated (D). E-
22 F. Paratype MSNM i24785, carapace, right lateral view, natural light (E), UV light (F).
23 Abbreviations: avp= antero-ventral process, dc= dorsal carina, dm= dorsal margin, ms= muscle
24 scar, on= optical notch, os= optical spine, pn= posterior notch, pu= punctuation, rap= prominent
25 anterior appendages, tl= paddle-like limbs or trunk limbs, ts= trunk somites, vm= ventral
38 ACCEPTED MANUSCRIPT 1 margin. Scale bars: 5 mm (A-C, E, F) and 2 mm (D). Line drawing: S. Charbonnier.
2 Photographs: D. Audo (A), J. Haug (C, D) and L. Cazes (E-F).
3
4 FIG. 7. – Thylacocaris schrami Audo & Charbonnier nov. gen, nov. sp. from the Cenomanian
5 of Hadjoula, Lebanon. A-D. Paratype MNHN.F.A57235, right lateral view, green-orange light
6 desaturated, subcomplete specimen showing carapace, eye, and prominent anterior appendages
7 (A), interpretative line drawing (B), close-up of optical notch with well-preserved eye, and two
8 optic spines (C), close-up of anterior appendages (D). E-F. Paratype MSNM i25125, carapace,
9 eyes and fragments of anterior appendages, natural light (E), green-orange light desaturated (F).
10 Abbreviations: avp= antero-ventral process, dc= dorsal carina, dm= dorsal margin, gi=
11 lamellate gills, ms= muscle scar, os= optical spine, pn= posterior notch, pu= punctuation, pds=
12 postero-dorsal spine, pvs= postero-ventral spine, r= rostrum, rap= prominent anterior
13 appendages, tl= paddle-like limbs or trunk limbs, ts= trunk somites, vm= ventral margin. Scale
14 bars: 5 mm (A, B, E, F) and 2 mm (D). Line drawing: S. Charbonnier. Photographs: J. Haug
15 (A, C, D, F) and L. Cazes (E). 16 BSGF 17 FIG. 8. – Thylacocaris schrami Audo & Charbonnier nov. gen, nov. sp. from the Cenomanian
18 of Hadjoula, Lebanon. A-C. Paratype MNHN.F.A57244, carapace, right lateral view, natural
19 light (A), line drawing (B), green-orange light desaturated, before preparation (C). D-F.
20 Paratype MSNM i24785, carapace, right lateral view, UV light (D), green-orange light
21 desaturated (E), close-up of rostrum (F). Abbreviations: avp= antero-ventral process, dm=
22 dorsal margin, ms= muscle scar, on= optical notch, os= optical spine, pn= posterior notch, pu=
23 punctuation, r= rostrum, tl= paddle-like limbs or trunk limbs, ts= trunk somites, vm= ventral
24 margin. Scale bars: 5 mm (A-E) and 2 mm (F). Line drawing: M. Lasseron. Photographs: D.
25 Audo (A), J. Haug (C, E, F) and L. Cazes (D).
39 ACCEPTED MANUSCRIPT 1
2 FIG. 9. – Holotype MSNM i20655 of Globulocaris garassinoi Teruzzi & Charbonnier nov. gen,
3 nov. sp. from the Cenomanian of Hakel, Lebanon. A. Carapace, left lateral view, natural light.
4 B. Interpretative line drawing. C. Image with inverted colours. Abbreviations: avp= antero-
5 ventral process, dm= dorsal margin, dcn= dorsal concave notch, ms= muscle scar, pn= posterior
6 notch, pds= postero-dorsal spine, pvs= postero-ventral spine, r= rostrum, ts= trunk somites,
7 vm= ventral margin. Scale bars: 2 mm. Line drawing: S. Charbonnier. Photographs: G. Teruzzi.
8
9 FIG. 10. Hamaticaris damesi (Roger, 1946), nov. gen., nov. comb. the Santonian of Sahel Alma,
10 Lebanon. A-B. Paralectotype MNHN.F.B18842, carapace, right lateral view (A) and
11 interpretative line drawing (B). C. Lectotype MNHN.F.B18843, carapace, right lateral view,
12 note the anterior prehensile appendages. D. Paralectotype MNHN.F.A30668, carapace, left
13 lateral view. E-F. Paralectotype MNHN.F.A30652, carapace, right lateral view (E) and
14 interpretative line drawing (F). G. Paralectotype MNHN.F.A30655, fragment of carapace, left
15 lateral view. H. Paralectotype MNHN.F.A30669, carapace, left lateral view. I. Paralectotype 16 MNHN.F.A30659, carapace, rightBSGF lateral view, note the Y-shaped ridges. J. Paralectotype 17 MNHN.F.A30672, carapace, left lateral view. K-L. Paralectotype MNHN.F.A30656, carapace,
18 left lateral view (K) and interpretative line drawing (L). Abbreviations: avp= antero-ventral
19 process, dm= dorsal margin, gi= gills, irs= infra-rostral spine, kn= knob, ms= muscle scar, on=
20 optical notch, pu= punctuation, pds= postero-dorsal spine, r= rostrum, rap= prominent anterior
21 appendages, ts= trunk somites, tl= paddle-like limbs or trunk limbs, vm= ventral margin. Scale
22 bars: 5 mm. Line drawing: S. Charbonnier. Photographs all in cross-polarized light: J. Haug
23 and G. Doitteau (K).
24
40 ACCEPTED MANUSCRIPT 1 FIG. 11. – Holotype MSNM i27356 of Keelicaris deborae Teruzzi & Charbonnier nov. gen,
2 nov. sp. from the Santonian of Sahel Alma, Lebanon. A. Carapace, right lateral view, natural
3 light. B. Carapace, right lateral view, UV light. C. Interpretative line drawing. Abbreviations:
4 avp= antero-ventral process, dm= dorsal margin, lc= lateral carina, lv= left valve, ms= muscle
5 scar, on= optical notch, pm= posterior margin, pu= punctuation, r= rostrum, ru= rugations or
6 terraces, rv, right valve, vm= ventral margin. Scale bars: 1 cm. Line drawing: S. Charbonnier.
7 Photographs: G. Teruzzi.
8
9 FIG. 12. – Line drawings of the carapaces of the thylacocephalan genera from the Late
10 Cretaceous Konservat-Lagerstätten of Libanon (front part to the left). A. Paradollocaris nov.
11 gen. B. Thylacocaris nov. gen. C. Globulocaris nov. gen. D. Hamaticaris nov. gen. E.
12 Keelicaris nov. gen. F. Thylacocephalus Lange, Hof, Schram & Steeman, 2001. F. Protozoea
13 Dames, 1886. G. Pseuderichtus Dames, 1886. Line drawings: S. Charbonnier.
BSGF
41 ACCEPTED MANUSCRIPT Figure 1
N
10 km
Hakel
Byblos Hadjoula (Jbeil)
Mediterranean En Nammoura Sea
Sahel Alma
Santonian Lagerstätte Beirut BSGF Cenomanian Lagerstätten
ACCEPTED MANUSCRIPT Figure 2
A B dc dm
pds r pn pu pvs on ms
avp vm
C D
gi
ts
eye ms
tl
E F
G BSGFH
I J dc
ms
tl
ACCEPTED MANUSCRIPT Figure 3
A B
C pu D
ms
r tl
vm rap on rap
E F dm r
dc gi pu eye
BSGFpn ms tl avp
vm
ACCEPTED MANUSCRIPT Figure 4
A B dm dc pds r gi pn
pvs eye
avp vm
C D
E F
eye BSGF G H dm dc r
pn pvs eye
tl ms avp vm
ACCEPTED MANUSCRIPT Figure 5
A B dc dm
dm pds
pn
pu pvs pu
gi ms gi gi vm gi
rap
rap rap
C D ms BSGFon vm avp
rap rap
ACCEPTED MANUSCRIPT Figure 6
dc A B dm
r pn ts pu eye tl os ms
avp vm
C D dm
pu
r
rap on
rap os
E F
BSGF
ACCEPTED MANUSCRIPT Figure 7
dm A B dc r pds pu os pn os ts pvs eye ms
avpavp vm
rap
rap
C D
vm
rap
rap
rap
E F dc dm
BSGF r ts gi
eye ms eye tl vm rap
ACCEPTED MANUSCRIPT Figure 8
A D
B E dm
r pn ts tl pu on os ms avp vm F dm
C
r
BSGF os
ACCEPTED MANUSCRIPT Figure 9
A
B dm
r dcn pds pn pvs r ts eye pvp ms
avp vm
vm C BSGF
ACCEPTED MANUSCRIPT Figure 10
A B dm pds r pu
ms on irs avp vm kn
C D dm r pds irs on pu ts ms rap avp tl rap kn vm E G dm
r
irs on F pu dm pds r ms tl gi avp kn ts on irs ms vm avp vm kn rap H dm pds gi ts
tl on
avp kn vm I BSGFJ
K L dm r pu pds gi irs on ts ms avp kn vm
ACCEPTED MANUSCRIPT Figure 11
A
B
C dm
lv pu
r
pm rv lc on ru ms BSGF avp vm
ACCEPTED MANUSCRIPT Figure 12 A E
B F
C G
D H
BSGF
ACCEPTED MANUSCRIPT Table 1
Studied THYLACOCEPHALA Age Distribution specimens CONCHYLIOCARIDA Dollocarididae Paradollocaris vannieri nov. gen., nov. sp. 17 Cenomanian Hadjoula Thylacocaris schrami nov. gen., nov. sp. 6 Cenomanian Hadjoula CONCAVICARIDA Protozoeidae Globulocaris garassinoi nov. gen., nov. sp. 1 Cenomanian Hakel Hamaticaris damesi (Roger, 1946) nov. gen., nov. comb. 25 Santonian Sahel Alma Keelicaris deborae nov. gen., nov. sp. 1 Santonian Sahel Alma Protozoea hilgendorfi Dames, 1886 51 Santonian Sahel Alma Pseuderichtus cretaceus Dames, 1886 12 Santonian Sahel Alma Microcarididae Thylacocephalus cymolopos Lange, Hof, Schram & Steeman, 2003 3 Santonian Sahel Alma
BSGF
ACCEPTED MANUSCRIPT