Fig 1.
Historical material of Varanus marathonensis Weithofer, 1888.
(A) Lectotype IPUW 1888-001-001, left maxilla from the late Miocene, MN12, of Pikermi (Greece) as figured in its original description [15]. (B) Paralectotype, isolated supraorbital from Pikermi (currently lost). (C, D) Ventral and left lateral view of the vertebra MNHN Pikermi n. 31 from the same locality as illustrated by Gaudry [49]. (E) Lateral view of the lectotype IPUW 1888-001-001 as is preserved nowadays. (F) Dorsolateral view of the same maxilla. Note that the expanded surface on the maxilla (mefp) was already shown by the original drawing by Weithofer, who also described it in the accompanying text. Scale bars equal 10 mm; (A) and (B) not to scale.
Fig 2.
Topotypic vertebrae of Varanus marathonensis Weithofer, 1888.
(A–C) Caudal vertebrae, IPUW 5187, from Pikermi (Greece), in dorsal (A), ventral (B) and right lateral (C) views. Note the pedestal for the chevron bone located close to the condyle (but separated from it). Scale bar equals 10 mm.
Fig 3.
Maxillae of Varanus marathonensis Weithofer, 1888 from Abocador de Can Mata.
(A–D) Right maxilla IPS50119 from the middle Miocene, MN7+8, of ACM/C4-C2 in lateral (A), medial (B), dorsal (C), and ventral (D) views. (E–H) Same as above but right maxilla IPS50292 from ACM/C3-A6. Note the medially expanded ascending surface of the facial process and the evident lateral sulcus developed along the narial margin. Scale bars equal 10 mm.
Fig 4.
Vertebrae of Varanus marathonensis Weithofer, 1888 from Abocador de Can Mata.
(A, B) Fragmentary trunk vertebra IPS42353 from ACM/C4-C2 in dorsal (A) and ventral (B) views. (C, D) Fragmentary trunk vertebra IPS42362 from ACM/C4-C2 in dorsal (C) and anterior (D) views. (E, F) Fragmentary trunk vertebra IPS43719 from ACM/C4-C2 in dorsal (D) and ventral (F) views. (G, H) Caudal vertebra IPS42400 from ACM/C4-C2 in ventral (G) and left lateral (H) views. (I, J) Caudal vertebra IPS42368 from ACM/C4-C2 in ventral (I) and right lateral (J) views. The arrows indicate the precondylar constriction. Scale bars equal 5 mm.
Fig 5.
Isolated maxillae of Varanus marathonensis Weithofer, 1888 from Cerro de los Batallones.
(A–D) Right maxilla BAT-3 1088a from the late Miocene, MN10, of Cerro de los Batallones in lateral (A), medial (B), dorsal (C), and ventral (D) views. (E, F) Left maxilla BAT-3 1088b in lateral (E) and medial (F) views. Scale bar equals 10 mm.
Fig 6.
Isolated prefrontal of Varanus marathonensis Weithofer, 1888 from Cerro de los Batallones.
(A, B) Right prefrontal BAT-3 1088c from Cerro de los Batallones in external (A) and internal (B) views. Scale bar equals 5 mm.
Fig 7.
Cranial bones of the almost complete skeleton of Varanus marathonensis Weithofer, 1888 from Batallones-3.
(A, B) Premaxilla in anterior (A) and posterior (B) views. (C–E) Left maxilla in medial (C), lateral (D), and dorsal (E) views. (F, G) Left palatine in dorsal (F) and ventral (G) views. (H, I) Left pterygoid in dorsal (H) and ventral (I) views. (J, K) Right ectopterygoid in dorsal (J) and ventral (K) views. (L, M) Left lacrimal in medial (L) and lateral (M) views. (N–P) Left quadrate in anterior (N), posterior (O), and lateral (P) views. Scale bars equal 5 mm.
Fig 8.
Cranial bones of the almost complete skeleton of Varanus marathonensis Weithofer, 1888 from Batallones-3.
(A, B) Left prefrontal in lateral (A) and medial (B) views. (C, D) Right supraorbital in dorsal (C) and ventral (D) views. (E–H) Left frontal in dorsal (E), ventral (F), lateral (G), and medial (H) views. (I) Right frontal in ventral view. (J, K) Left postorbitofrontal in ventral (J) and dorsal (K) views. (L, M) Parietal in dorsal (L) and ventral (M) views. Scale bars equal 5 mm.
Fig 9.
Braincase of the almost complete specimen of Varanus marathonensis Weithofer, 1888 from Batallones-3.
(A–D) Isolated parabasisphenoid in dorsal (A), ventral (B), anterior (C), and right lateral (D) views. (E–I) Braincase in right lateral (E), anterior (F), ventral (G), posterior (H), and dorsal (I) views. (J, K) Isolated left paroccipital process in anterior (J) and posterior (K) views. Scale bars equal 5 mm.
Fig 10.
Lower jaw of the almost complete specimen of Varanus marathonensis Weithofer, 1888 from Batallones-3.
(A, B) Left dentary in lateral (A) and medial (B) views. (C) Right dentary in medial view. (D–G) Posterior part of the right mandible (surangular, prearticular, articular) in lateral (D), medial (E), ventral (F), and dorsal (G) views. (H, I) Left coronoid in lateral (H) and medial (I) views. Scale bars equal 5 mm.
Fig 11.
Presacral vertebrae of the almost complete specimen of Varanus marathonensis Weithofer, 1888 from Batallones-3.
(A–E) Axis and third cervical vertebra in dorsal (A), ventral (B), anterior (C), posterior (D), and left lateral (E) views. (F–J) Trunk vertebrae in left lateral (F), dorsal (G), ventral (H), anterior (I), and posterior (J) views. Scale bars equal 5 mm.
Fig 12.
Sacral region of the almost complete specimen of Varanus marathonensis Weithofer, 1888 from Batallones-3, including the last trunk vertebra, two sacral vertebrae, one cloacal vertebra (plus a fragment of a possible second one), and first caudal vertebra.
Scale bars equal 5 mm.
Fig 13.
Tail of the almost complete specimen of Varanus marathonensis Weithofer, 1888 from Batallones-3.
(A) Complete tail. (B) Detail of the caudal vertebrae in right lateral view. Scale bars equal 10 mm.
Fig 14.
Forelimb of the almost complete specimen of Varanus marathonensis Weithofer, 1888 from Batallones-3.
(A, B) Right coracoid in lateral (A) and medial (B) views. (C–F) Left humerus in anterior (C), posterior (D), dorsal (E), and ventral (F) views. Scale bars equal 5 mm.
Fig 15.
Hindlimb of the almost complete specimen of Varanus marathonensis Weithofer, 1888 from Batallones-3.
(A, B) Left ilium in lateral (A) and medial (B) views. (C–F) Right femur in ventral (C), dorsal (D), anterior (E), and posterior (F) views. (G, H) Left femur in anterior (G) and ventral (H) views. (I, J) Left tibia in ventral (I) and dorsal (J) views. (K) Proximal end of an undetermined fibula in ventral view. Scale bars equal 5 mm.
Fig 16.
Majority-rule consensus tree based on the results of analysis 1A.
The position of Varanus marathonensis (as scored from information provided by the specimen from Batallones and the type of Varanus amnhophilis; this OTU is named “Vmarathonensis Vamnhophilis” in figures of the supplementary information) inside the Indo-Asian A group and as sister species to the Indovaranus subgenus is supported by 85% of the MPTs. Numbers in the time scale represent million years ago.
Fig 17.
Snouts of extant Varanus species showing the maxillary morphology in dorsal and right lateral view.
(A, B) Varanus ornatus (Daudin, 1803) ZFMK 14889. (C, D) Varanus komodoensis Ouwens, 1912 ZFMK 64998. (E, F) Varanus griseus (Daudin, 1803) SMF 32911. (G, H) Varanus bengalensis (Daudin, 1802) SMF 63456. Note in all cases the absence of a lateral sulcus along the edge of the narial aperture. In V. ornatus and V. komodoensis there is no hint of medial expansion of the medial edge of the maxilla (no medial concavity along the medial edge of the maxilla in dorsal view), whereas in V. griseus and V. bengalensis a sort of expansion is mostly located in the area anterior to the facial process (presence of a variably developed concavity). Among the four species here depicted, V. ornatus is the only one that bears blunt, molariform teeth. Scale bars equal 10 mm.
Fig 18.
Partial skull (ΜΥΤ-130, including the anterior portion of the skull) of Varanus marathonensis Weithofer, 1888 from Samos in right dorsolateral view.
Scale bar equals 5 mm.