**海巨龍屬**（學名：*Thalassotitan*，意為"海洋巨人"）是一屬已滅絕的大型滄龍，生活於約6600萬年前晚白堊紀馬斯垂克期的摩洛哥。該屬下有且僅有**殘暴海巨龍**（*T. atrox*）一個物種，該物種描述自一組發現於奥勒德阿卜杜恩盆地（Ouled Abdoun Basin）的化石。海巨龍屬於**傾齒龍族**（學名：Prognathodontini），該類群包括傾齒龍等其他滄龍。 

該屬的發現表明滄龍類在白堊紀晚期已經演化出了占據顶级掠食者生態位的物種。嚴重磨損的海巨龙牙齿和發現於正模标本附近的、酸蝕過的化石表明，海巨龙可能會捕食小型沧龙科、蛇颈龙、大型鱼类和海龟。

語源學
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海巨龙的属名*Thalassotitan*組合自古希腊語"θìλασσα"（*thálassa*，意為"海"）和"τιτìν"（*tītā́n*，"巨人"），指代海巨龙的巨大体型。種加詞*atrox*則來自拉丁語，意為"残忍的"或"无情的"，指代该物种的顶级捕食者地位和化石上因种內搏鬥產生的大量咬痕。

描述
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復原後的海巨龙與成年人的大小对比
海巨龙是最大的沧龙之一，其头骨可达1.3米，身长 可達9-10米。

### 頭骨

沧龙头骨結構示意图
像所有的倾齿龙一样，海巨龙的头骨又钝又结实，前颌骨是承载头骨顶端的骨头，从侧面看非常短，但从背面看则宽而凸。前颌骨的身体包含许多被称为神经血管孔的凹坑，这些凹坑被认为容纳了对触摸非常敏感的触觉神经。内腭杆是前颌骨的一个很长的延伸部分，一直延伸到额骨，它在上颌骨(主要承载牙齿的上颌骨)和外鼻孔(容纳鼻孔的开口)之间穿过时很宽，但当它与额骨接触时变窄成一个细长的杆。在上颌骨 之间，内鼻骨形成一个明显的低而短的龙骨。上颌骨短而结实，而且很深。它的表面是平的，除了在牙齿上方有一个低而宽的脊衬。

Neurovascular foramina line this margin, increasing in size as they progress towards the back of the skull. The texture of the maxilla's surface is rough, which is especially apparent in larger individual, caused by a network of veined grooves to house blood vessels. The external nares extend from and to above the fourth and twelfth maxillary teeth. The jugal bone, which is located just below the eye, is broad and robust. The frontal bone is short and broad, shaped almost like an isosceles triangle, with large neurovascular foramina at the center. The pineal foramen, which contains the parietal eye, is small and long. The supratemporal fenestrae, large openings between the eyes and the back end of the skull, take up nearly a quarter of the entire skull length and are somewhat triangular. The dentary, the tooth-bearing bone of the lower jaw, is short, wide, robust, and curved concave towards the upper jaws. Many bones of the upper jaw are tightly sutured together; its two tooth-bearing bones the premaxilla and maxilla were connected through interlocking joints containing an unusual series of flanges and grooves while an interdigitating set of tongue-and-groove joints secure the maxilla and prefrontal bone.

### 牙齒

Skull with teeth shown
*Thalassotitan* teeth are roughly conical in shape, lightly curved, large in size, and robust in build. They are most similar to the teeth of *P. saturator* except in being slightly shorter and stockier. Tooth crowns are slightly swollen around its base next to the root, but they do not form a round circumference. The surfaces of the crown are generally smooth but may sometimes have faint ridges depending on individual or ontogenetic variation. The enamel at the tip contain veinous ridges and coarse bumps. Cutting edges are well-developed and finely serrated. Each tooth has two cutting edges, but their positions differ depending on the tooth's position in the jaw. Towards the front of the jaw, the front-facing cutting edges are more pronounced than the diminished back-facing edges. In the middle and near the end of the jaw, both edges are of equal development and located diametrically opposite to each other. At the end of the jaw, the back-facing edges become more pronounced. The tooth roots are massive and barrel-shaped. Deep pits occur within the roots, from which new replacement teeth are formed.

Like all mosasaurs, *Thalassotitan* had four types of teeth, corresponding to the jaw bones they are located on. On the upper jaw were the premaxillary teeth, maxillary teeth, and pterygoid teeth (located separate from the main jawline near the rear of the skull); while on the lower jaw only the dentary teeth were present. *Thalassotitan* had in each jaw row from front to back: two premaxillary teeth, twelve maxillary teeth, at least six pterygoid teeth (the pterygoids were not fully preserved), and fourteen dentary teeth. The dentary teeth are generally flatter by the side than the maxillary teeth. Heterodonty is present, meaning that tooth shape changes down the jawline. The first four or five teeth are tall, narrow, and slightly curved, which become stockier, erect, and more robust around the middle of the jawline, then become shorter (as broad as they are tall), hooked, and flatter by the side. The pterygoid teeth are strongly hooked but are also large and robust, nearly approaching the size of the teeth on the main jawlines.

### 顱後骨骼

The postcranial skeleton is not fully known, only fossils representing a little more than the front half of the body have been found.

The general shape of the vertebrae are typical for mosasaurines. They are procoelous, meaning that the front side is deeply cupped concavely and the back side is bulged convexly. The cervical (neck) vertebrae are slightly wider than long. Its atlas holds rectangular or triangular neural arches; another single tall neural arch is also present at the top of the vertebra. The articular surfaces, which atach to the cartilage that connect vertebrae together, is initially heart-shaped but becomes rounded at the rearmost cervicals. The dorsal (back) vertebrae are slightly longer than wide with tall neural arches, rounded articular surfaces, and large rectangular transverse processes. The ribs are short and robust.

The pectoral girdle is robust and most similar with that in *P. overtoni* and *Mosasaurus conodon*, albeit more square-shaped than the latter. The two bones making up the girdle, the scapula and coracoid, are similar in sizes. They loosely contact with each other, but their contact point is nevertheless wider than the glenoid fossa. The scapula is shaped like a square, being as long as it is wide. It lacks a defined scapular neck but expands from front to back, forming a fan-like convex blade. The coracoid is also somewhat squarish and lacks a well-defined neck. Its margins are weakly concave in the front and back but very convex at the bottom.

The forelimbs formed long paddles that resembled mosasaurin mosasaurs like *Mosasaurus* and *Plotosaurus* but more primitive in possessing longer but fewer phalanges. The humerus is very stocky and resemble that in *P. overtoni* except in the expansion of the glenoid condyle beyond the postglenoid process. The radius is unusually shaped for a mosasaur. It is as large as the humerus and much larger than the ulna and takes on a crescent-like or subrectangular form, unlike smaller hourglass-shaped radii in typical mosasaurs.

分類
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*T. atrox* is most closely related to *Prognathodon saturator* (left) and *P. currii* (right), and the three species may be congeneric.
*Thalassotitan* is a member of the Prognathodontini tribe of the Mosasaurinae subfamily, with other members including *Prognathodon* and *Gnathomortis*. Morphologically, it is most similar to the giant mosasaurs *P. currii* and *P. saturator*, and a phylogenetic analysis by Longrich et al. (2022) recovered *Thalassotitan* in a clade between the two. This creates an unnatural paraphyletic relationship that reflects a wider issue with the genus *Prognathodon* as a whole. Several studies over the past decade found that *Prognathodon* is in general not monophyletic and in need of revision. Longrich et al. (2022) suggested such a revision may include an expansion of the *Thalassotitan* genus to include *P. currii* and *P. saturator*. However, due to a high degree of convergent evolution in the relationship-determining traits among many mosasaurs (especially among prognathodontines), phylogenetic results between each study are seldom consistent, mystifying exactly which species must be revised to stabilize the Prognathodontini. For example, some studies recovered *P. currii* and *P. saturator* as phylogenetically unrelated species with either falling outside a monophyletic *Prognathodon*, while other studies yield variable placements for the type species *P. solvayi* as either outside a monophyletic *P. currii*-*P. saturator* clade in support of Longrich et al. (2022) or within it, which would in theory invalidate *Thalassotitan* as a junior synonym under the principle of priority.

The following cladogram is modified from Longrich et al. (2022).


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| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
| Mosasaurinae | 

|  |  |
| --- | --- |
|   | *Kourisodon* |
|   |
|   | 

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| --- | --- |
|   | *Clidastes* |
|   |
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| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
|   | 

|  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
| Globidensini | 

|  |  |  |  |  |  |  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
|   | 

|  |  |
| --- | --- |
|   | *Globidens simplex* |
|   |
|   | *Globidens schumani* |
|   |
|   | *Globidens phosphaticus* |
|   |
|   | *Prognathodon rapax* (=*Ancylocentrum hungerfordi*) |
|   |

 |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Globidens alambamensis* |
|   |
|   | *Globidens dakotensis* |
|   |

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|   |

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|   |
| Prognathodontini | 

|  |  |
| --- | --- |
|   | *Gnathomortis* |
|   |
|   | 

|  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
|   | 

|  |  |
| --- | --- |
|   | *Prognathodon overtoni* |
|   |
|   | 

|  |  |  |  |  |  |  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
|   | 

|  |  |
| --- | --- |
|   | *Prognathodon saturator* |
|   |
|   | 

|  |  |
| --- | --- |
|   | ***Thalassotitan atrox*** |
|   |
|   | *Prognathodon currii* |
|   |

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|   |

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|   |
|   | *Prognathodon giganteus* |
|   |

 |
|   |

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|   |
|   | 

|  |  |
| --- | --- |
|   | *Prognathodon lutugini* |
|   |
|   | *Prognathodon solvayi* |
|   |

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|   |

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|   |

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|   |

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|   |
| Mosasaurini | 

|  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- |
|   | 

|  |  |
| --- | --- |
|   | *Moanasaurus* |
|   |
|   | *Mosasaurus mokoroa* |
|   |

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|   |
|   | 

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| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
|   | 

|  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
|   | 

|  |  |  |  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
|   | 

|  |  |
| --- | --- |
|   | *Mosasaurus conodon* |
|   |
|   | *Plesiotylosaurus* |
|   |
|   | *Plotosaurus* |
|   |

 |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Mosasaurus missouriensis* |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Mosasaurus lemonnieri* |
|   |
|   | 

|  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- |
|   | 

|  |  |
| --- | --- |
|   | *Mosasaurus hoffmannii* |
|   |
|   | *Mosasaurus beaugei* |
|   |

 |
|   |
|   | *Mosasaurus maximus* |
|   |

 |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Liodon* |
|   |
|   | *Mosasaurus sp.* (MGGC 21876) |
|   |

 |
|   |

 |
|   |

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|   |

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|   |
|   | "Magahouanga mosasaurine" |
|   |

 |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Carinodens* |
|   |
|   | *Xenodens* |
|   |

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|   |

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|   |

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|   |

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|   |

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|   |

 |
|   |

古環境學
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Life restoration of *Thalassotitan* (left) and other animals of the Ouled Abdoun Basin
The phosphate deposits of Morocco have revealed an extremely diverse environment of late maastrichtian age. The oceans of the area were full of an abundance of fish, from bony fish like *Enchodus* and *Stratodus* to cartilaginous fish like *Cretalamna, Squalicorax* and *Rhombodus.* There was also an abundance of marine reptiles, most notably the mosasaurs, with more than 10 genera alone known from this single site. This possibly suggests that niche partitioning took place here, in which predators take on different niches to avoid competition with one another (for example, mosasaurs like *Carniodens* and *Globidens* had blunt teeth for crushing shellfish while *Thalassotitan* and *Mosasaurus* hunted much larger food). Other marine reptiles include the elasmosaurid plesiosaur *Zarafasaura*, the sea turtle *Alienochelys* and the gavialoid crocodilian *Ocepesuchus* .

It seems that *Thalassotitan* was an apex predator in its ecosystem, with evidence being digestive damage found on some of the fossils in the nearby vicinity including those of plesiosaurs, turtles, and large fish. In the skies flew multiple species of pterosaurs including the azhdarchid *Phosphatodraco*, the nyctosauromorph *Alcione* and *Simurghia*, the nyctosaurid *Barbaridactylus*, and the possible pteranodontid *Tethydraco*. On land three species of dinosaurs are known, these being the abelisaur *Chenanisaurus,* the small lambeosaurine hadrosaur *Ajnabia* and a so far unnamed titanosaur. *Thalassotitan* lived alongside other giant mosasaurs like *Prognathodon*, *Mosasaurus* and *Gavialimimus*, as well as smaller mosasaurs like *Xenodens*, *Halisaurus* and *Pluridens*.

參考文獻
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1. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18  Longrich, Nicholas R.; Jalil, Nour-Eddine; Khaldoune, Fatima; Yazami, Oussama Khadiri; Pereda-Suberbiola, Xabier; Bardet, Nathalie.. Cretaceous Research. 2022-12-01, **140**. ISSN 0195-6671. doi:10.1016/j.cretres.2022.105315 （英语）.
2. ↑  Johan Yans; M'Barek Amaghzaz; Baadi Bouya; Henri Cappetta; Paola Iacumin; László Kocsis; Mustapha Mouflih; Omar Selloum; Sevket Sen; Jean-Yves Storme; Emmanuel Gheerbrant.. Gondwana Research. 2014, **25** (1): 257–269. Bibcode:2014GondR..25..257Y. S2CID 129475565. doi:10.1016/j.gr.2013.04.004.
3. ↑  Catherine R. C. Strong; Michael W. Caldwell; Takuya Konishi; Alessandro Palci.. Journal of Systematic Palaeontology. 2020, **18** (21): 1769–1804. ISSN 1477-2019. S2CID 224978215. doi:10.1080/14772019.2020.1818322.
4. ↑  Dale A. Russell.  **23**. New Haven: Bulletin of the Peabody Museum of Natural History. 1967: 240  [2023-05-15]. OCLC 205385. （原始内容存档于2023-07-10）.
5. ↑  Nathalie Bardet; Nour-Eddine Jalil; France de Lapparent de Broin; Damien Germain; Olivier Lambert; Mbarek Amaghzaz.. PLOS ONE. 2013, **8** (7): e63586. Bibcode:2013PLoSO...863586B. PMC 3708935 可免费查阅. PMID 23874378. doi:10.1371/journal.pone.0063586 可免费查阅.
6. ↑  Peggy Vincent; Nathalie Bardet; Xabier Pereda Suberbiola; Baâdi Bouya; Mbarek Amaghzaz; Saïd Meslouh.. Gondwana Research. 2011, **19** (4): 1062–1073. Bibcode:2011GondR..19.1062V. S2CID 129404886. doi:10.1016/j.gr.2010.10.005.
7. ↑  Nicholas R. Longrich; David M. Martill; Brian Andres.. PLOS Biology. 2018, **16** (3): e2001663. PMC 5849296 可免费查阅. PMID 29534059. doi:10.1371/journal.pbio.2001663 可免费查阅.
8. ↑  Fernandes, Alexandra E.; Mateus, Octávio; Andres, Brian; Polcyn, Michael J.; Schulp, Anne S.; Gonçalves, António Olímpio; Jacobs, Louis L.. Diversity. 2022, **14** (9). 741. doi:10.3390/d14090741 可免费查阅.
9. ↑  Nicholas R. Longrich; Xabier Pereda Suberbiola; R. Alexander Pyron; Nour-Eddine Jalil.. Cretaceous Research. 2020, **120**: 104678  [2023-05-15]. S2CID 228807024. doi:10.1016/j.cretres.2020.104678. （原始内容存档于2021-04-04）.
10. ↑  Nicholas R. Longrich; Xabier Pereda-Suberbiola; Nour-Eddine Jalil; Fatima Khaldoune; Essaid Jourani.. Cretaceous Research. 2017, **76**: 40–52. S2CID 133063691. doi:10.1016/j.cretres.2017.03.021.
11. ↑  Trevor Rempert; Alexander Vinkeles Melchers; Ashley Rempert; Muhammad Haque; Andrew Armstrong.  (PDF). The Journal of Paleontological Sciences. 2022, **22**: 1–22  [2023-05-15]. （原始内容存档 (PDF)于2022-01-15）.