Dorateuthis syriaca Woodward, 1883

Dorateuthis syriaca Woodward, 1883

Figs. 2–4 View Fig View Fig , 8–19 View Fig View Fig .

1878 Sepialites ; Fraas 1878: 346. 1883 Dorateuthis syriaca n. sp.; Woodward 1883: 1–5, pl. 1. 1888 Curculionites senonicus View in CoL ; Kolbe 1888: 135, pl. 11, fig. 8. 1896 Sepialites ; Woodward 1896: 231. 1922 Dorateuthis syriaca Woodward, 1883 ; Naef 1922: 118. 1922 “ Plesioteuthis fraasi ” Woodward, 1896 ; Naef 1922: 133, fig. 50. 1922 “ Sepialites sahil –almae ” O. Fraas, 1878; Naef 1922: 134, fig.

49c. 1943 Sepialites sahil almae ; Klinghardt 1943: 12, fig. 8. 1946 Leptoteuthis syriaca Woodward, 1883 ; Roger 1946: 14, pl. 4: 5,

6, pl. 9: 1, 2. 1986 Dorateuthis sahilalmae Naef, 1922 ; Engeser and Reitner 1986:

3, fig. 1, pl. 1: 1. 1986? Dorateuthis sp. ; Engeser and Reitner 1986: 4, pl. 1: 2. 1987 Sepialites sahilalmae (O. Fraas) Naef, 1922 ; Riegraf 1987: 97. 1988 Dorateuthis sahilalmae Naef, 1922 ; Engeser 1988: 43. 2006 a Dorateuthis cf. syriaca Woodward, 1883 ; Fuchs 2006b: 7, fig. 4,

pls. 1–3. 2006 b Dorateuthis sahilalmae ; Fuchs 2006a: pls. 17h, 22f. 2007b Dorateuthis syriaca Woodward, 1883 ; Fuchs et al. 2007b: 246. 2007b Dorateuthis sahilalmae Naef, 1922 ; Fuchs et al. 2007b: 246. 2007 Dorateuthis syriaca Woodward, 1883 ; Fuchs 2007: 64, fig. 4. 2007? Dorateuthis sahilalmae Naef, 1922 ; Fuchs 2007: 64. 2011 a Dorateuthis syriaca Woodward, 1883 ; Fuchs and Larson 2011a:

237, figs. 2–6, 7.5, 8. 2015 Dorateuthis syriaca Woodward, 1883 ; Nixon 2015: 8, fig. 6a, c,

d.

→ Fig. 3. A. The holotype BMNH C5017 of plesioteuthid coleoid Dorateuthis syriaca Woodward, 1883 , from the upper Santonian of Sahel Aalma, Lebanon, as illustrated by multiple non-destructive imaging techniques. A 1, µXRF overlay of copper (blue), yttrium (green) and phosphorous (pink) distributions over the entire individual; A 2, rotated close-up of the posterior section with the fin outline visible on the right; A 3 –A 5, individual images from the RTI stack showing the anterior part of the body, head, and arm crown (A 3), the posterior part of the body (A 4), and the central part of the body (A 5); A 6, A 7, UV photographs of the entire individual (A 6) and the anterior part of the body, head, and arm crown (A 7). B. Hypothesized illustration of the holotype (Jessica Gardner). The Y-enriched elongate feature in A 1 is not considered part of the individual, but part of a contemporaneous organism. Scale bars 10 mm .

2015 Dorateuthis syriaca Woodward, 1883 ; Jattiot et al. 2015: 152, figs. 4–6, 10, 11, 12.1–3, 13.2–3, 14.

2016 Dorateuthis syriaca Woodward, 1883 ; Donovan and Fuchs 2016: 20, figs. 12–15.

2016b Dorateuthis syriaca ; Fuchs et al. 2016a: 438, figs. 1, 9.

2018 Dorateuthis syriaca Woodward, 1883 ; Gueriau et al. 2018: 985, fig. 5.

2020 Dorateuthis syriaca Woodward, 1883 ; Fuchs 2020: 11, fig. 3.

2021c Dorateuthis syriaca Woodward, 1883 ; Klug et al. 2021c: 7, fig. 5.

Holotype: BMNH C5017 , original of Woodward (1883: pl. 1).

Type locality: Sahel Aalma, Lebanon.

Type horizon: Upper Santonian, Upper Cretaceous.

Description.— Redescription of the holotype. — It is preserved in ventral view (Fig. 3). The body length (the posterior mantle margin to the anterior tip of the arms) is ~ 68 mm. The length of the gladius is ~ 40 mm and corresponds with the length of the mantle. It has a triangular median field and convex (Fig. 3A 3, A 6) anterior margin (~ 4 mm wide). The gladius has well defined lateral reinforcements (keels) and a central bipartite ridge. All are continuous from the anterior to the posterior margin (Fig. 3A 6, B). The apical angle is 6.3°. There is no evidence of a central median field encompassing the individual’s bipartite ridge. There is no evidence of lateral fields, hyperbolar zones or a conus.

µXRF elemental mapping and UV photography enables the most precise assessment of arm length to date. The holotype has eight tapered arms (Fig. 3A 1, A 7, B). Six appear complete and range from 18–21 mm in length. Two lack their distal sections and have a preserved length of 15 and 17 mm. Using the index (arm length max to gladius length) outlined in Fuchs (2020) the arms are “moderate” in size (a ratio of 0.53). The preserved diameter of seven of the arm bases varied between 0.5 and 1.4 mm. The arm with the smallest diameter ( 0.5 mm) is partially obscured by another arm, and therefore the original diameter was likely larger. There is no evidence of an additional tentacular pair, or any associated hooks that were noted in the original description by Woodward (1883). No suckers or cirri were observed.

µXRF elemental mapping reveals the cephalic cartilage (Fig. 3A 1), which is enriched in phosphorus and observed as two concave, semi-circular traces that flank the oesophagus in an anterior-posterior orientation. The cartilage is about 9 mm in length and is positioned between the posterior margin of the arm crown and the anterior lateral keels. The structures have a larger medial separation than is normally observed in dorsal view, and do not have the “pear-shape” associated with ventral view ( Fuchs and Larson 2011a: fig. 4), therefore we interpret what is preserved to be a central slice (Fig. 3A 1). The length of the cephalic cartilage was used as a proxy for eye diameter for the holotype. The ratio between the eyes and the gladius length ( diametermax to gladius length) is ~0.23. The cephalic cartilage flanks a mass of soft tissue that corresponds with the oesophagus and possibly the brain or optic lobes. The position of the cephalic cartilage is inconsistent with the eye position supposed by Woodward (1883). Indeed, imaging reveals their interpretation reflects degraded tissue at the base of the arm crown (Fig. 3A 1, A 7). Small protrusions, visible in the UV photographs and elemental maps, are positioned anteriorly to the lateral keels and likely represent soft tissue (Fig. 3A 1, A 3, A 6, A 7). It is possible that these represent posterior salivary glands or are the remnants of retractor musculature. Two areas of digestive contents are visible (Fig. 3A 4, A 6, B). The anterior-most area was originally described as an ink sac ( Woodward 1883: pl. 1), however, our multimodal imaging reveals these structures are in fact digestive remnants. A random sampling of fragment sizes in these two digestive assemblages shows that the anterior remains are roughly 0.2–0.4 mm in length, while those in the posterior section have larger components (approximately 0.2–1.2 mm). A small ink sac is more posteriorly located (Fig. 3A 4, B). RTI and µXRF mapping also reveal faint imprints of what we interpret to be elongate gill lamellae and a posterior fin Fig. 3A 1 to A 5). The contours of the fin revealed by elemental mapping (Fig. 3A 2) resemble the “oar-shaped” characteristic described for Dorateuthis ( Fuchs 2020) . As fins are other-

wise unknown in individuals from Sahel Aalma ( Fuchs and Larson 2011a), this represents the only known coleoid fin tissue preserved from this locality for this species.

Redescription of Dorateuthis syriaca .— Size: The diagnostic mantle length for Dorateuthis is 201–400 mm ( Fuchs 2020), which is categorised as “medium” in size. These sizes were defined by Fuchs (2020: 7) as “very small” (< 50 mm) small ( 50–200 mm), “medium” ( 201–400 mm), “large” ( 401–1500 mm), and “very large” (> 1500 mm). The mean mantle length in this study is smaller (~ 112 mm, Tables 3 and 4), however, one of our measured samples, MNHN.F.A50402 ( Jattiot et al. 2015: fig. 4.3), did fall within the range for the “medium” sized mantle length ( 241 mm). The rest are either “small” (~93%, n = 50), or “very small” (6%, n = 3). The body length was able to be measured for 43% (n = 23) of the individuals in the whole sample, and ranges from 68 mm (the holotype) to 456 mm ( MNHN.F.A50402, Jattiot et al. 2015: fig. 4.3).

Gladius size and shape: All the measured individuals have a triangular-shaped median field ( Figs. 2 View Fig and 4A–C View Fig ) with the widest section located at the anterior-most margin of the gladius. The shape of the anterior margin is slightly convex. This varies from the current diagnosis ( Fuchs 2020), though it supports a previous, tentative interpretation ( Fuchs 2006a: fig. 4). An anteriorly rounded imprint or contour on the ventro-medial section of the gladius is observed in four individuals (e.g., Fig. 4D View Fig ). It is located just posterior to the anterior margin and bears a resemblance to the medial component of the head retractors seen in Vampyroteuthis infernalis Chun, 1903 ( Bizikov and Toll 2016: fig. 12). As such, we suggest this feature corresponds with an attachment site for muscle tissue.

87% of the individuals have the slender (gladius widthmax to gladius length) gladius proportions of Dorateuthis ( Table 3, SOM: table 3), though 13% show a very slender

0.20 0.18 300

250 0.16 0.14)

mm 200 index 0.12 (Gladius 0.10

0.08

length 150

0.06

Gladius 100 0.04

50 0.02 0.00 0

Hjoula Haqel Sahel Lebanon Hjoula Haqel Sahel Lebanon

Aalma unde- Aalma unde-

termined termined

A B C

gladius. All the individuals exhibit the requisite apical (opening) angle (<12°, mean of 6.9°). We found no statistical differences between gladius size, shape, apical angle, or body size, and any of the three specific localities ( Fig. 5 View Fig ). However, Sahel Aalma shows the most size disparity as the smallest (the holotype, with a mantle size of 40 mm) and largest ( MNHN.F.A 50402, 241 mm) individuals are from this outcrop, which was not expected given the small sample size.

Gladius reinforcements, lateral fields, and median field area: Each of the 54 studied individuals has characteristic lateral reinforcements (keels) that are pronounced, and continuous from the anterior to the posterior margins of the gladius ( Figs. 2 View Fig , 4A–C View Fig ). More than half of the sample (63%) exhibited some form of median reinforcements ( Fig. 4A–C View Fig , SOM: table 4), longitudinally bisecting the gladius. These median reinforcements vary and take the form of either a central medial line ( Figs. 2 View Fig , 4B View Fig ) or ridge ( Figs. 2 View Fig , 4C View Fig ), which may be uni- or bi-partite. A central median field, an elongate area in the central section of the median field ( Figs. 2 View Fig , 4A View Fig ) may also be visible and span the line or ridge; they are also observed without a line or ridge. So far, there is no evidence to explain the type of median reinforcement exhibited. Longitudinal striations on the gladius ( Fig. 4A, B View Fig ) were commonly observed in the central median field in both dorsal and ventral view. These have previously been illustrated in plesioteuthids, such as Plesioteuthis prisca Rüppell, 1829 ) ( Naef 1922: fig. 42b) and noted to be “fine longitudinal lines” (translated from German).

The gladius has three observed morphological variations Fig. 6 View Fig ). The most common type of gladius (78% of the sample) shows a complete absence of lateral fields ( Fig. 6A View Fig ). Seven individuals have anteriorly projected lateral fields Fig. 6B View Fig ), which extend approximately 80% to 90% along the length of the gladius. Four have elongated posterior lateral fields (30% of the gladius length) with no evidence of anterior lateral fields ( Fig. 6C View Fig ). These gladius morphotypes could not be associated with other soft tissue characters e.g., arm length, total body size, or locality) in the study sample. It is possible that the variation observed is due to post mortem processes (e.g., mineral replacement across the gladii, disarticulation, or non-uniform compression),

ontogenetic stage, dimorphism, or polymorphism enabling potential variation in muscular attachment (e.g., Toll 1998; Fuchs et al. 2016a; Bizikov and Toll 2016).

Where lateral fields were observed, the area of the median field was calculated following the formula given in Fuchs (2020) (median field area to gladius areatotal), who notes this is “very large” (>0.8) in Dorateuthis . All the individuals measured had a median field area>0.8 consistent with the diagnosis ( SOM: table 3).

Conus: The conus is preserved in only two individuals, one in ventral view ( Fig. 4E View Fig ), the other in lateral. The length represents 3–5% of the gladius (conus lengthmax to gladius lengthmax), and 80% of the width (conus widthmax to gladius widthmax). No diagnostic indices currently exist for this character.

Arms: Twenty-two individuals (41%) provided the basis for arm measurements in this sample. Each of these individuals preserve at least one arm; nineteen preserve two or more, and only two show a complete arm crown. Both the latter individuals show the diagnostic differentiation in arm length noted for the species: a more elongated dorsal arm pair, a relatively short ventral pair, and two intermediatesized pairs in lateral position ( Fuchs and Larson 2011a: fig. 3). The three smallest individuals (gladius < 50 mm) in the sample, including the holotype, show very little variation in arm length ( SOM: table 2).

Comparisons between arm and gladius lengths were conducted ( Fig. 7 View Fig ). These results indicated a general relative increase in relative arm length to mantle length as would be expected, supporting previous analyses that linked differentiation in the arm crown with allometry ( Fuchs 2006a; Fuchs and Larson 2011a; Jattiot et al. 2015).

Chemical investigation reveals that the muscular mantle tissue of D. syriaca is commonly enriched in yttrium and strontium ( Fig. 8 View Fig ) the presence of which is indicative of substitution for calcium in calcium phosphate (apatite) during diagenesis ( Gueriau et al. 2018). This is consistent with phosphatization, a mode of preservation common in Cretaceous Lebanese coleoid fossil soft tissues ( Clements et al. 2017; Klug et al. 2021c). Donovan and Fuchs (2016) supposed that arms were also preserved in apatite though they lacked mineralogical analyses. XRF data shows an enrichment in yttrium and strontium and supports their initial assumption ( Fig. 8A View Fig ). The substitution of these elements in varying soft tissues within a specimen has not been fully investigated, and this variation could be a diagenetic artifact, or could possibly reflect a bias of phosphatization, which is known to be highly selective of particular tissues (as outlined in Clements et al. 2022).

Armature: Eight individuals (15%) exhibit faint circular structures (~ 1–2 mm in diameter) on the arms that are interpreted here as suckers ( Fig. 9A View Fig , SOM: fig. 1). These appear uniserial and radially symmetrical when observed under UV light. Although no individual has an entire row observable, we infer they were present along the entire length since they are visible on proximal, medial, and distal portions of the arms in the separate specimens. There is no indication of attachment types, and no evidence of sucker linings, hooks, or cirri. Three individuals show remnants of the axial nervous system in the arm crown ( Fig. 9B View Fig ). These axial nerve elements ( Klug et al. 2023; Rowe et al. 2023) are visible in UV light, and observed as delicate paired filaments, or dotted lines that follow the contours of the arms. Though one of these individuals have preserved the structure along the entire length of the arm, remnants are observed both proximally and medially ( Fig. 9A View Fig , SOM: fig. 1).

Head: The cephalic cartilage ( Fig. 9 View Fig ) is evident in 31% of the individuals. Using the methodology for determining orientation of the cephalic cartilage as outlined by Fuchs and Larson (2011a), we determine that in our samples it is preserved both dorsoventrally and laterally. It is most clearly visible in the UV photographs and elemental maps where the phosphatized cartilage fluoresces ( Fig. 9A View Fig 1, B 1 View Fig ). Structures interpreted as statocysts (e.g., Figs. 9B View Fig , 10A) are present on 4 individuals, just posterior to the cephalic cartilage. Internal calcareous statoliths ( Boyle and Rodhouse 2008) are not observable.

Eye measurements were obtained on 15 specimens (28% of the total individuals). As we had more precise measurements for the gladius, the gladius length was used to identify a ratio for maximum eye diameter (gladius length:eye diameter 0.13–0.25) rather than follow the index of Young and Vecchione (1996), which compares the eye radius to head width. Five individuals also preserve evidence of a central eye lens, which can be observed in Fig. 10A by the positioning of the capillary system in the eye ( Donovan and Fuchs 2016; Fuchs et al. 2016b; Fuchs and Larson 2011a). This appears in natural light as delicate filaments of yellow staining in a reticulated pattern. This pattern appears dark under UV light ( Fig. 10A).

Preservation of the buccal area is common and observed in ~72% of the individuals. The rostral tips, darker in colour than the rest of the beak chitin, are often visible under natural light. The muscular tissues of the buccal mass fluoresce under UV light ( Figs. 9B View Fig 1 View Fig , 10A, 11A View Fig 3). µXRF analysis demonstrates that the muscular tissue has elevated strontium traces compared with the matrix, indicating replacement of the original organic material by calcium phosphate ( Gueriau et al. 2018). The length of the buccal mass was measured in 31 individuals. The ratio of buccal mass length/gladius length ranges from 0.08–0.19. More detailed measurements of the buccal mass were taken on individuals NPL 52121 ( Fig. 11C View Fig ) and BHI 2203 to capture the 2D shape, which were scaled to provide a reconstruction, the first of its kind for D. syriaca (see Fig. 12 View Fig ).

Posterior salivary glands and other paired structures: Previous descriptive works on D. syriaca have identified a pair of posterior salivary glands located next to the anterior-most parts of the lateral margins of the gladius ( Lukeneder and Harzhauser 2004: pl. 2.2; Jattiot et al. 2015: figs. 12.1, 12.2). Our study includes two individuals figured in Jattiot et al. (2015) and Lukeneder and Harzhauser (2004) ( Fig. 11B and D View Fig respectively), that provide a comparator for this feature. The paired structures observed in the holotype closely resemble these posterior salivary glands.

Four individuals show paired structures that differ visually from the previously described posterior salivary glands ( Figs. 11A, C View Fig , 13 View Fig ). Two exhibit ovoid-shaped imprints in the region of the anterior lateral keels ( Fig. 11A, C View Fig ). In natural light, one of these structures is visible laterally to the anterior gladius margin in both individuals (shown here in NPL 52121, Fig. 11C View Fig ). The XRF and MSI images reveal a corresponding ovoid area, which is visible on the opposing side of the gladius margin. But for the shape, there is no evidence to suggest that this is anything other than mantle tissue. As such, it is possible that this ovoid structure reflects the anterior section of a funnel. The other two individuals possess paired structures with a rope-like morphology ( Fig. 13 View Fig ). These are either visible as hemispherical projections anteriorly angled towards the lateral keels ( Fig. 13A View Fig ), or medially angled inwards from the lateral keels ( Fig. 13B View Fig ) and are interpreted here as remnants of funnel retractor muscles ( Bizikov and Toll 2016; Fuchs et al. 2016a).

Digestive system: Preserved digestive contents are present in 19 individuals. Typically, the remains are ingested, and their relative position in the body enables comparison with the digestive organs in extant coleoids ( Mangold and Young 1998; Wells 2011). One individual ( MNHN.F.A88589) preserves remains in 3 areas ( Fig. 14A View Fig 1 –A View Fig 4 View Fig ): two are positioned in the posterior section of the mantle ( Fig. 14A View Fig 1 View Fig , A 3, A 4) and likely reflect the stomach and caecum. The composition of these remains varies which supports the different functions of these two digestive organs. The mass of digestive remains interpreted here to be stomach contents ( Fig. 14A View Fig 3), contains an articulated ray fin and a pelvic girdle, likely from a teleost fish; the posterior-most mass ( Fig. 14A View Fig 4 View Fig ) has a collection of less distinct, individual bones surrounded by a soft mass and likely represents caecal contents. The third mass of digestive remains ( Fig. 14A View Fig 2 View Fig ) is positioned more anteriorly in the mantle, is tubular in shape, and also retains evidence of fish bones. The position and contents are consistent with the presence of a crop.

The individuals examined in the study enabled observation of almost all elements of the digestive system of D. syriaca . While the preservation of ingested food matter is not uncommon in organisms that are phosphatically preserved stomach tip buccal salivary digestive mass gland gland

(see Clements et al. 2022), phosphatization appears to be highly selective in terms of which tissues are replaced by apatite—and while ingested organics appear to preferentially phosphatise, internal digestive integument is comparatively rare in the fossil record (Clements et al. 2022). This seems to be the case in D. syriaca , however, the relative positions of the ingested organics within the organism allows the determination of an Octobrachia-like gut configuration. One key character identified by this study is the crop of D. syriaca ; the presence of a crop is known in Recent Vampyromorpha ( Fig. 14A View Fig 5 View Fig ) and most Octopoda , but it is absent in modern Decabrachia ( Mangold and Young 1998: fig. 1). As such, this configuration of the digestive system is consistent with the assignment of Dorateuthis to Octobrachia .

Funnel: A funnel can be inferred in BHI 2213, preserved in dorso-lateral view ( Fig. 15B View Fig ). The feature was not observed directly, though its presence was determined by a stained elongated mass within the boundaries of the mantle. The staining is interpreted here to delineate the internal part of the funnel. The anterior margin of this is located just posteriorly and ventrally to the head, consistent with the placement of the funnel in both fossil and modern coleoids.

This stained impression shows evidence of diagenetic circular structures ( Fig. 15B View Fig ) associated with the presence of ink ( Klug et al. 2021c). These circular patterns are also present in individual MNHN.F. R 06746 on an unresolved structure ( Fig. 15A View Fig 3). Roger (1946) suggested it was a rectal bulb (translated from French), and Jattiot et al. (2015) indicated it may be a buccal mass. Given its location ( Fig. 15A View Fig ) and the presence of the diagenetic circular patterns, we interpret this structure to represent an accumulation of the long strings of pigmented faeces (coprolite) released from the mantle via the funnel ( Boyle and Rodhouse 2008).

Gills: Remains of gills are present in 12 individuals (22%). They are preserved either as impressions ( Fig. 16A View Fig ), orange/ yellow ( Fig. 16B View Fig ), or black stains ( Fig. 16C View Fig ). The orange staining of the gill relics represents pyritization, rather than phosphatization ( Donovan and Fuchs 2016). The most complete gills ( Fig. 16C View Fig ) show the branchial heart (~ 2 mm length max), the main efferent vessel (~ 14 mm), and a minimum of 17 gill blades. Combined, this represents ~21% of the gladius length. The associated lamellae are not preserved, though the filamentous gill blades taper distally (~4– 1.3 mm). Lamellae (n = minimum of 9) and gill blades are present in MNHN.F. A88589 ( Fig. 16A View Fig ). Under natural light, the blades are visible as orange filaments, and the lamellae are preserved as contoured relief. No main efferent vessel is observed.

Fins: The “oar-shaped” fin described for D. syriaca ( Fuchs 2020: fig. 3, 1e) is observed in the holotype from the Santonian of Sahel Aalma (Fig. 3A 1, A 2, B), as well as six (~11%) other individuals ( Fig. 17 View Fig ) from the Cenomanian outcrops. It is preserved as a mineralized structure ( Fig. 17A View Fig ), an impression ( Fig. 17B View Fig ) or an orange-coloured stain ( Fig. 17E View Fig ), making it generally distinguishable from the matrix under natural light. Its visualization can be enhanced using µXFR elemental mapping ( Figs. 8B, C View Fig and 17D View Fig ) and/ or UV photography ( Fig. 17C, F View Fig ) .

Reproductive system: Reproductive organs were tentatively included in a summary of known soft tissues in D. syriaca ( Donovan and Fuchs 2016) . This was based on a description by Roger (1946: fig. 7) of spermatophores in individual MNHN.F. R 06746. Observations on this individual show that what Roger (1946) interpreted to be spermatophores are, instead, digestive contents ( Fig. 15A View Fig 1 View Fig ). However, in MNHN.F.A88588 there are impressions present which possibly represent oviductal glands ( Fig. 15C View Fig ). These correspond with a titanium-enriched area that is observable using µXRF mapping ( Fig. 8B View Fig ).

Novel anatomic characters of Dorateuthis syriaca : The multimodal imaging approach and quantitative analyses performed in this work has enabled a revision of the summary of known soft tissues in D. syriaca ( Fig. 18 View Fig , SOM: table 5) that were previously synthesized in Donovan and Fuchs (2016). Furthermore, we have identified morphological characters, which were not previously described for the genus including paired retractor muscles, axial nerves in the arms, and an Octobrachia-type digestive system. For the first time, we also provide clear evidence of a funnel, circulatory system, and excretory system. Lastly, we discount the presence of tentacles, tentacular pockets, and hooks within the arm crown, and confirm that D. syriaca possessed suckers.

Stratigraphic and geographic range.—Lower upper Cenomanian–upper Santonian of Lebanon.