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Article

Out of Pikermi: The Occurrence of Bohlinia in the Late Miocene of the Central Mediterranean

by
Antonella Cinzia Marra
1,2
1
Department MIFT, University of Messina, Viale Stagno D’Alcontres 31, I-98166 Messina, Italy
2
Museal System of the University of Calabria (SiMU), Section of Palaeontology, Via Pietro Bucci, cubo 14/b, I-87036 Arcavacata di Rende, Italy
Geosciences 2025, 15(2), 44; https://doi.org/10.3390/geosciences15020044
Submission received: 5 January 2025 / Revised: 26 January 2025 / Accepted: 28 January 2025 / Published: 1 February 2025
(This article belongs to the Section Biogeosciences)
Browse Figures
Versions Notes

Abstract

:
During the Late Miocene, Giraffidae of the genus Bohlinia were widespread in the Greco-Iranian bioprovince (Pikermian biome), while the occurrence in Africa (Chad) was disproven. The fossils of Bohlinia cf. attica described and compared here come from the Late Miocene of Cessaniti (southern Italy), associated with another giraffid, Samotherium cf. boissieri, and the large mammals Stegotetrabelodon syrticus, Tragoportax cf. rugosifrons, cf. Ceratotherium advenientis, and an undetermined Anthracotherid. In terms of paleogeography, the association should be related to the expansion of Pikermian species out of the Greco-Iranian bioprovince and calls for more in-depth considerations. Pikermian giraffids, and Bohlinia in particular, play an important role in understanding the paleoenvironmental and paleogeographical contexts in the Central Mediterranean, an area undergoing major geological changes.

1. Introduction

The Late Miocene outcrops of Cessaniti (the municipality of Cessaniti, the Province of Vibo Valentia, Calabria, Southern Italy; Figure 1a) produced an interesting terrestrial mammal assemblage [1,2,3,4,5,6,7]. The fossils come from the “Clypeaster sandstones”, as informally named by Nicotera (“arenarie a Clypeaster” [8]) for the impressive abundance of echinoids. Neogene deposits, including the “Clypeaster sandstones”, lie on a Hercynian crystalline basement and outcrop with different thicknesses throughout an area spanning from the coast near Briatico–Tropea–Cape Vaticano and the inland Mount Poro (Figure 1b) [4,5,6,7,8,9,10].
The most significant stratigraphic section is exposed in a quarry near Cessaniti, named Gentile, after the owner’s surname. The succession includes, from the bottom to the top [4,8,9,10,11], the following units:
  • Unit 1: “dark clayish sandstone with Ostrea and Cerithium”, alternating with coarse sandstone, interpreted as deposited in a lagoonal environment;
  • Unit 2: “Clypeaster sandstone”, interpreted as deposited in a shallow marine environment;
  • Unit 3: “Heterostegina yellow sandstone”, interpreted as deposited in open marine conditions;
  • Unit 4: “Orbulina marl”, interpreted as deposited in a hemipelagic environment.
The succession is interpreted as a marine transgression from coastal lagoon to open sea environments [4,9,10,11,12,13,14,15]. From a depositional point of view, it is interesting to point out that the “Clypeaster sandstones” are interrupted by periodical subaerial expositions and fluvial deposition, probably occurring during brief interruptions in the major transgressive event or for tectonics [4].
Marra et al. [4] bracketed the stratigraphic succession from Unit 1 to the base of Unit 4 between 8.1 and 7.2 Ma.
Besides the invertebrate fossil fauna particularly rich in echinoids, Unit 2 released remains of marine and land mammals (Table 1).
The Sirenian Metaxytherium serresii is the most abundant marine mammal, easily recoverable and identifiable for its pachyostotic bones; the species is also present in Unit 3 [16,17,18,19,20]. Cetaceans rarely occur and are represented by Odontocetes (Physiteroidea indet.) and Mysticetes (a few fossils of Heterocetus cf. guiscardii) [19,20,21].
The land mammals found at Gentile’s quarry are as follows: Stegotetrabelodon syrticus, Tragoportax cf. rugosifrons, cf. Ceratotherium advenientis, an undetermined Anthracotherid, Samotherium boissieri [2,3,4,5,6], and another giraffid previously attributed to Bohlinia cf. attica [3,22], here fully described, compared, and attributed to Bohlinia attica (Table 1). Only Stegotetrabelodon syrticus is also present in Unit 1, where one deciduous tooth has been found [2,23]. Tragoportax cf. rugosifrons and Samotherium boissieri also come from neighbor sites correlatable to Unit 2 of Gentile’s quarry succession [5].
The land mammal remains probably reached the marine depositional environment by being transported by rivers or floods. The fossils are, for the most part, in a good state of preservation. Some of them suffered intense fragmentation post-fossilization due to the quarry works and/or weathering [3,4,24].
The mammal assemblage represents a bioprovince defined by the association of Afro-Arabian and Greco-Iranian elements [1,3], clearly distinct from the Tusco-Sardinian (including terrains of actual Tuscany and Sardinia (Figure 1a; TSB in Figure 2)) and Apulo-Abruzzi (including terrains of actual Apulia and Abruzzi (Figure 1a; AAB in Figure 2)) bioprovinces, characterized by endemic taxa evolved in insular conditions (not fully reported in the available paleogeographic reconstructions) [25,26]. The possible location of the Cessaniti land is still not determined (Figure 2), and a possible spreading through North Africa has been hypothesized [1,3].
The Cessaniti fossils referrable to Bohlinia are significant for the chronological and geographical distribution of the species, as well as for the paleogeographical reconstruction of the Central Mediterranean, an area subject to major geological events during the Late Miocene (Figure 2).

2. Materials and Methods

2.1. Materials

All the fossils described here come from Gentile’s quarry, often referred to as Cessaniti from the name of the municipality in which is located.
The fossils are in a state of preservation ranging from good to fragmented, as indicated in the description. The studied fossils are now stored in two museums, for each of which the abbreviation used in this text and the inventory is given as follows:
  • Museal System of the University of Calabria, Section of Palaeontology, Via Pietro Bucci 87036 Rende, Cosenza, Italy; abbreviation: SiMU.
  • City Museum of Ricadi, Via Roma, 12, 89866 Santa Domenica, Vibo Valentia, Italy; abbreviation: MU.RI
The fossils described here are detailed as follows (each fossil is indicated with the museum’s abbreviation followed by the inventory number):
  • Upper left toothrow (P4-M3) with fragmentary alveolar portions of the maxillary bone; SiMU-CMA01.
  • Left distal radius; MU.RI-874.
  • Right metacarpal; MU.RI-972.
  • Left astragalus; SiMU-CMA02.
  • Fragmentary astragalus; MU.RI-877.
  • Left cubonavicular; SiMU-CMA04.
  • Right cubonavicular; SiMU-CMA05.
  • The fossils come from the upper part of Unit 2, indicated as levels 6-7 by Carone and Domning [19] and as CG.SH2-CG.SH3 by Marra et al. [4].

2.2. Methods

The fossils studied here come from Gentile’s quarry and were dispersed in the Clypeaster sandstones. Their recovery was possible thanks to the periodical field activities of amateur paleontologists during suspensions of quarrying works authorized by the superintendence [27,28]. Due to this collection method, it was not possible to obtain all the recovery data. However, the amateur paleontologists documented the findings, allowing them to identify the levels of provenance with a good approximation [27,28].
The fossil teeth and the bones were, in some cases, consolidated with glue and then studied and described using anatomy textbooks and specific papers. Measurements were taken using a caliber, according to Von Der Driech [29], and referring to the main papers on Giraffids, cited when needed. The measurements were included in the morphological description of each specimen.
The bone descriptions followed the standard anatomical terminology. The terminology adopted for describing molars and premolars is in Figure 3d.

3. Systematic Paleontology

Class: Mammalia Linnaeus, 1758;
Order: Artiodactyla Owen, 1848;
Suborder: Ruminantia Scopoli, 1777;
Family: Giraffidae Gray, 1821;
Subfamily: Bohlininae Solounias, 2007;
Genus: Bohlinia Matthew, 1929;
Type species: Bohlinia attica (Gaudry et Lartet, 1856).
The studied materials: an upper left toothrow (P3-M3) with fragmentary alveolar portions of the maxillary bone (SiMU-CMA01); a left distal radius (MU.RI-874); a right metacarpal broken into two parts (MU.RI-972); a complete (SiMU-CMA02) and a fragmentary astragalus (MU.RI-877); and a left (SiMU-CMA04) and a right cubonavicular (SiMU-CMA05).
The stratigraphic occurrence: Late Miocene (Upper Tortonian; European Land Mammal Age: Turolian) of Cessaniti (Vibo Valentia, Italy), Clypeaster sandstones from levels 6 and 7 according to Carone and Domning [19], corresponding to CG.SH2-CG.SH3 in Marra et al.’s study [4].

3.1. Descriptions

3.1.1. Toothrow

The left toothrow (SiMU-CMA01) is preserved from P4 to M3 and is held with a small portion of the alveolar portion of the maxilla (Figure 3). A P3, still under restoration, was also found and probably belonged to the same toothrow. The molars and premolars are brachyodont and large in an initial/medium stage of wear. The enamel is finely crenulated. A cingulum is present, more robust, and evident on the buccal side.
P3: A worn P3 possibly belonging to the toothrow (SiMU-CMA01) is under restoration. The P3 is worn and has a well-developed metastyle and parastyle. The external tubercle is prominent. Measurements (in mm): breadth = 22.4; length = 21.5.
P4: The metastyle and the parastyle are highly evident. The parastyle is more developed than the metastyle. The external tubercle is robust and high. It is furrowed, and this makes it look divided into two columns. Measurements (in mm): breadth = 26.8; lingual length = 18.0; labial length = 17.5.
M1: The metacone is wide, very prominent, and directed toward the lingual side. The central part of the metacone is not very prominent. The mesostyle is prominent and robust, with a U-shaped section in the occlusal view. The hypocone is arcuate and not prominent. The paracone has weakly arcuate ribs; its pillar is wide and prominent. The parastyle is well-developed. The protocone is more lingually projected than the hypocone and is arcuated, with a prominent anteriorly directed point. The entostyle is absent. Measurements (in mm): anterior breadth = 26.9; posterior breadth = 26.7; lingual length = 22.5; labial length = 23.7.
M2: The metastyle is slightly prominent. In the occlusal view, the metacone is weakly arcuate and slightly prominent, with straight ribs. The mesostyle has a U-shaped section. In the labial view, the metacone shows an almost flat wall, where the metastyle is slightly evident, while the mesostyle is prominent and robust. The hypocone is arcuate and wide. The posterior rib has a very weak fold. The anterior rib has a fold on the external enamel. The paracone has slightly arcuate ribs and is wide and highly prominent on the labial side. The parastyle is very robust and prominent. The protocone is arcuated and anteriorly pointed toward the lingual side. The protocone is more protruding toward the lingual side than the hypocone. The entostyle is absent. Measurements (in mm): anterior breadth = 31.5; posterior breadth = 29.4; lingual length = 26.4; labial length = 28.6.
M3: A small posterior enamel ridge is present, almost in the center on the posterior side. The metastyle is well-developed and pointed toward the lingual side. The anterior and posterior ribs of the metacone are almost straight. The metacone is very slightly prominent. In the labial view, the wall of the metacone is almost flat and is rearward with respect to the highly prominent metastyle and mesostyle. The latter is elevated, robust, and very prominent. In the occlusal section, it is elongated and pointed toward the labial side. The mesostyle is very wide at its base. The hypocone is well-developed and prominent. The entostyle is small, short, and not worn. The paracone is wide and prominent and pointed toward the labial side. The anterior and posterior ribs of the paracone are highly arcuate. The parastyle is prominent, with a wide base. The protocone is wide and arcuate, with a wide point and lingually directed. An entostyle between the protocone and paracone is present. The anterior rib of the protocone is more arcuate than the posterior one, being almost straight. Measurements (in mm): anterior breadth = 31.6; posterior breadth = 28.5; lingual length = 28.0; labial length = 29.0.
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Figure 3. Left toothrow (SiMU-60-CMA01) in labial (a), occlusal (b), and lingual (c) views; (d) dental nomenclature used for descriptions.
Figure 3. Left toothrow (SiMU-60-CMA01) in labial (a), occlusal (b), and lingual (c) views; (d) dental nomenclature used for descriptions.
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3.1.2. Fore Limb Bones

The left radius (MU.RI-874) is represented by a distal epiphysis (Figure 4d). Despite the strong fragmentation of the specimen, the articular surface is well-preserved. The scaphoid’s facet is oval, concave in the anterior part, and convex posteriorly. Its major axis is oblique. The facet of the intermedium is oblique and has a wide anterior portion, where it is concave and is then directed posteriorly with an elongated convex surface. The facet of the ulnare is anteriorly concave and posteriorly convex. The lateral styloid process is pronounced and robust. The distal breadth of the distal epiphysis is 102 mm.
The right metacarpal (MU.RI 972) is broken into two parts, and the central portion of the diaphysis was lost because it was very fragmented (Figure 4a–c). The epiphyses are preserved well enough to permit descriptions and measurements. In the proximal epiphysis, articular view, the facet of the trapezoid–capitatum is extended laterally and anteroposteriorly. A step separates the facet of the uncinatum, which lies on a slightly lower plane. The facet of the uncinatum is flat and less wide on the lateral side. The synovial fossa is wide on the articular face. It seems unopened on the palmar face, but the specimen is intensively fragmented. In the anterior and posterior views, the proximal end appears flared toward the medial and lateral sides. The proximal portion of the shaft, although fragmented, shows a deep central groove delimited by a wide medial ridge, more pronounced than the lateral one. The distal condyles are pronounced. The distal epiphysis is flared on the lateral side. Measurements (in mm): proximal transverse diameter = 90.8; proximal anteroposterior diameter =53.1; distal transverse diameter = 88.0; distal anteroposterior diameter = 49.9.

3.1.3. Tarsal Bones

Two astragali are present in Cessaniti’s collection. The best-preserved one is the left astragalus labeled SiMU-CMA02 (Figure 4g–i), while the other is fragmentary (MU.RI-877). The astragali are slender. In the anterior view, the neck is wide; the lateral ridge of the trochlea has a vertical major axis and is elevated and massive; the collum tali is massive on the medial side and wide on the lateral side; and in the head, the lateral articular facet is extended, and the medial one is small. In the posterior view, the lateral ridge of the trochlea has an elevated and massive posterior notch; the medial ridge of the trochlea protrudes toward the medial side and is rounded and massive; the proximal triangular fossa is deep; the articular surface of the calcaneus is slightly convex, with a medial border uniformly concave over to the medial side, a circular concavity near the mediodistal border, and a straight lateral border; and the lateral notch of the trochlea is elevated. The measurements of SiMU-CMA02 in mm: medial length = 78.6; lateral length = 89.0; medial anteroposterior diameter = 55.6; lateral anteroposterior diameter = 49.6; distal transverse diameter = 49.
The cubonaviculars include a right one (SiMU-CMA05; Figure 4e,f) and a left one (SiMU-CMA04). In the proximal view, the astragal facets are concave and separated by a smooth ridge. The calcaneum facet is flat and laterally oriented. In the distal view, the metatarsal facet is flat, quadrangular in shape, and has a central depression; the anterior cuneal facet (of the cuneiform) has a semilunar shape and is flat; and a pronounced step separates the anterior cuneal facet from the small posterior cuneal facet, which lies on a lower plane. Measurements (in mm): SiMU-CMA04 length = 82; transverse diameter = 91.2; SiMU-CMA04 length = 91.5; transverse diameter = 94.6.
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Figure 4. (a) Right metacarpal (MU.RI-972) in proximal articular view; (b) proximal portion of right metacarpal (MU.RI-972) in posterior (palmar) view; (c) distal portion of right metacarpal (MU.RI-972) in posterior (palmar) view; (d) left radius (MU.RI-874) in distal articular view; (e) right cubonavicular (SiMU-CMA05) in distal articular view; (f) right cubonavicular (SiMU-CMA05) in proximal articular view; (g) left astragalus (SiMU-CMA02) in dorsal view; (h) left astragalus (SiMU-CMA02) in plantar view; (i) left astragalus (SiMU-CMA02) in lateral (external) view.
Figure 4. (a) Right metacarpal (MU.RI-972) in proximal articular view; (b) proximal portion of right metacarpal (MU.RI-972) in posterior (palmar) view; (c) distal portion of right metacarpal (MU.RI-972) in posterior (palmar) view; (d) left radius (MU.RI-874) in distal articular view; (e) right cubonavicular (SiMU-CMA05) in distal articular view; (f) right cubonavicular (SiMU-CMA05) in proximal articular view; (g) left astragalus (SiMU-CMA02) in dorsal view; (h) left astragalus (SiMU-CMA02) in plantar view; (i) left astragalus (SiMU-CMA02) in lateral (external) view.
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4. Discussion

The morphology and biometry of Cessaniti’s specimens addressed comparisons with Bohlinia attica, but Samotherium boissieri, the other giraffid found at Cessaniti, and Palaeotragus roueni, having similar dental morphologies, were also considered.
The specimens of Cessaniti attributable to Bohlinia attica represent the westernmost occurrence of the species, typically occurring in the Late Miocene of the Greco-Iranian bioprovince (Greece, Turkey, Bulgaria, the Republic of North Macedonia, Iraq, and Iran) from the Vallesian (European Neogene Mammal Age MN9/10) to the Late Turolian (MN13) [30,31,32,33,34,35,36,37,38].
The expansion of Bohlinia to the Central Mediterranean is still not fully understood and concerns the whole mammal assemblage of Cessaniti, where African (Ceratotherium advenientis), (Stegotetrabelodon syrticus) and Greco-Iranian (Tragoportax cf. rugosifrons, Samotherium boissieri, and Bohlinia attica) taxa occur, suggesting intriguing inferences in the biochronologic, paleogeographic, and paleoclimatic reconstructions of the Central Mediterranean.

4.1. Comparisons

The most significant bone is the astragalus, indubitably attributable to Bohlinia. The bones have also been compared with Samotherium boissieri, the other Giraffid species present in Cessaniti’s mammal assemblage. The dental morphologies are not highly distinctive for the Giraffid species due to the limited specimens and the variability in characteristics. The similarities in some morphological characteristics among Giraffid species suggest comparing the measurements of the tooth row with Palaeotragus rouenii.
The tooth row (SiMU-CMA01) is comparable in size and morphology to that of Bohlinia attica (Table 2). Unfortunately, the incompleteness of the specimen does not permit us to calculate the indexes for comparisons, such as 100 × P2-P4/M1-M3. The length of the molar row (M1-M3= 90 mm on the labial side; 87.1 mm on the lingual side) is shorter than Samotherium boissieri, longer than Palaeotragus rouenii, and comparable to the largest specimens of Bohlinia attica (Table 2) [33,34,35,36,38,39,40,41,42,43]. The molars seem proportionally larger than the available sample of Bohlinia attica (Table 2).
The teeth from Cessaniti are finely crenulated, large, and brachyodont, similar to Bohlinia. Moreover, as in Bohlinia, the premolars are large with respect to the molars. The molars are broader than in Palaeotragus. Despite the similarities between Palaeotragus rouenii and Bohlinia attica, the size suggests that it should be ruled out that the tooth row belongs to Palaeotragus (Table 2 and Table 3).
In the premolars of the studied specimen, the external tubercles of premolars are well developed, and the styles are pointed and progressively larger from the top to the base, as in Bohlinia [39]. In SiMU-CMA01, the bifurcation of the parastyle, considered primitive for Bohlinia and occurring in some specimens [39], is not observed, as well as the oblique orientation of the external tubercles. The specimen from Samos (CM458) described and figured by Parizad et al. [38] and the specimen from Cessaniti have P4 with a robust and prominent metastyle, parastyle, and external tubercle, separated by furrows. In both specimens, the external tubercle is furrowed. The premolars of the toothrow from Cessaniti are not molarized, unlike some specimens of Bohlinia Kostopoulos [39].
The molars of SiMU-CMA01 have the following characteristics in common with Bohlinia [33,35,36,38,39,44]: strong development of the styles; an angular protocone, which is lingually projected; a well-developed parastyle and mesostyle; the parastyle and paracone connected at the base; a well-developed paracone; a quite flat metacone, progressively less developed than the paracone from M1 to M3; a very weak metastyle; and a basal cingulum that is stronger in the external wall. The parastyle is stronger than the mesostyle, and this latter is prominent in the specimens described by Kostopoulos [40] and Parizad et al. [38]. In SiMU-CMA01, the mesostyle is prominent, weak in M3, and robust in M2 and M1, while the parastyle is inclined to the front. These characteristics appear like those of the specimen CM458 from Samos, Greece (Figure 5 in Parizad et al.’s study [38]) The teeth of Samotherium boissieri have variable strength of styles and cones, and some characteristics are not constant. The specimen from Cessaniti has the labial groove of the P4 external tubercle and the lingually projected protocone in common with S. boissieri. The latter characteristic is also common in Bohlinia (Bohlin [38,39,40,45]). The toothrow of SiMU-CMA01 does not present some characteristics observed in S. boissieri [40]: a posterior rib of the paracone that is less steep than the anterior one; a parastyle and paracone of equal robustness; a flat and oblique lingual wall, observed in some P4; a basal tubercle in the posterior lobe of the molars, which is often observed; and a narrow protocone of the molars.
The radius of MU.RI 874 is fragmentary, but the dimensions and the oblique ridge between the facets for the scaphoid and the ulnare permit us to assign it to Bohlinia.
The metacarpal of MU.RI 972 falls within the morphometric variability of Bohlinia attica and clearly differs from Samotherium boissieri, accordingly with the distinctive characteristics of metapodials and their variability in Giraffidae [33,35,39,40,46]. Two metacarpals attributed to Samotherium boissieri (MU.RI 864 and MU.RI 865) come from the “Clypeaster sandstones” outcropping at Contrada Malopara, a site near Gentile’s quarry [3,4,28] and have been considered in comparisons. In the proximal epiphysis, the articular facets lie in the same plane in Bohlinia and MU.RI 972, while the articular facets are separated by a steep step in Samotherium boissieri. The proximal epiphysis of MU.RI 972 is semi-oval like Bohlinia and unlike Samotherium boissieri, where it is semicircular. Although fragmentary, the most proximal portion of the diaphysis of MU.RI 972 presents a deep central groove, delimitated by a large and elevated medial ridge and a less elevated and sharp ridge, like Bohlinia and unlike S. boissieri, where the central trough is less deep and the ridges are less elevated and weaker. The distal epiphysis is sensibly larger than the shaft, like in Bohlinia, while in S. boissieri, it is less large.
The astragal morphometry falls within the lower values of variability known for Bohlinia (Table 4) [3,34,47]. The astragalus is a very distinctive bone among Giraffids.
At Cessaniti, one astragalus of Samotherium boissieri comes from the “Clypeaster sandstones” at Gentile’s quarry. The SiMU-CMA02 and MU.RI-877 specimens share the following characteristics with Bohlinia: general slenderness; a wide neck; massive collum tali; uniformly convex articular facets in the plantar view; and an elevated posterior notch of the lateral trochlea. On the contrary, the astragalus of Samotherium boissieri appears more massive, and the medial and lateral trochlea have a small difference in height. The measurements are within the range of Bohlinia and fit with smaller specimens (Table 4). The Palaeotragus astragali are smaller, while the Samotherium boissieri ones are similar to smaller Bohlinia specimens but show a different morphology.
The cubonaviculars can be ascribed to Bohlinia for the perfect coincidence with the morphology of the astragalus of the proximal articulation and the separation of the facets of the metatarsal by a deep groove [34].

4.2. Chronology, Distribution, and Paleoecology

The genus Bohlinia only includes B. attica: the validity of the species Bohlinia nikitiae (Nikiti, Greece) is still pending for the poor preservation status and the attribution of some specimens to Palaeotraginae [36,37,39]. The specimens attributed to the species Bohlinia adoumi (Toros-Menalla Chad), were later attributed to Palaeotraginae [36,38,49].
Bohlinia attica is a typical species of the Pikermian mammal association, spread in the Greco-Iranian bioprovince between 8.3 and 7.0 Ma, and dominated by Gomphotheriidae, Rhinocerotidae, Giraffidae, Bovidae, Equidae, Felidae, and Hyaenidae, while Cervidae was remarkably absent [31,50,51,52,53,54].
The available field data on the Cessaniti record, recently verified, report that the fossils of B. attica and associated mammals were collected in the upper part of Unit 2 at Gentile’s quarry, more precisely, in levels 6 and 7, according to Carone and Doming [19], and the two sedimentary units of the shoreface sands CG.SH2 and CG.SH3 were recognized by Marra et al. [4]. The remains of Tragoportax cf. rugosifrons and Samotherium boissieri have also been found at correlatable levels in the surrounding area (Table 1). Stegotetrabelodon is also present in Unit 1 (Table 1). The mammal assemblage falls within the time interval from 8.1 of Unit 1 to 7.2 Ma at the base of Unit 4.
At Cessaniti, Bohlinia attica occurs with another giraffid typical of the Pikermian biome, Samotherium boissieri [3,22], the Eurasian mid-sized bovid Tragoportax cf. rugosifrons [5], the Rhinoceronthid cf. Ceratotherium advenientis, which is tentatively ascribed to the Rhinocerotina African lineage [7], and the North African Prodoscidean Stegotetrabelodon syrticus [1,2]. Bohlinia attica co-occurs with Samotherium boissieri at Corakyerler (Turkey) and Injana and with Tragoportax rugosifrons at Hadjidimovo, Strumyani (Bulgaria), and Karaslari (the Republic of Macedonia) (Table 5). These three taxa co-occur only at Cessaniti.
The proboscidean sample of Cessaniti has been attributed to Stegotetrabelodon syrticus with plesiomorphic characteristics [1,2]. Stegotetrabelodon syrticus is known from As Sahabi (Lybia), where it is dated to 6.8–5.3 Ma, while the fossils from the Baynunah Formation (Abu Dhabi, United Arab Emirates), previously attributed to S. syrticus, have been re-attributed to the new subspecies Stegotetrabelodon syrticus emiratus, dated to 8.0–6.0 Ma, more likely 7.7–7.0 Ma [55,56]. That being so, the plesiomorphic characteristics recognized in the Cessaniti specimens [1,2] are undergoing revision.
The new species cf. Ceratotherium advenientis is not strongly indicative, being based on only one specimen and dubiously attributed to the genus [7]. The species has been ascribed to Rhinocerotina and related to the African genera Diceros and Ceratotherium by cladistic analysis [7].
The land mammals might have reached the lands of Cessaniti at the time of the westward spreading of the Pikermian biome [3,4,6]. The expansion of the Arabian desert during the Late Tortonian (7.9–7.4 Ma ago) should have probably caused the wave to Africa of the Pikermian biome [57]. The hypothesis of the spreading of the Cessaniti mammal precursors through North Africa and then Calabria [3,4,6] needs stronger data. In the current state of knowledge, the possible position of the lands of Cessaniti in the Late Miocene is not known (Figure 2).
In the Cessaniti assemblage, mammals common in the Greco-Iranian bioprovince prevailed, while African elements were not undoubtedly present. Moreover, the records in Africa are too discontinuous to reach more convincing conclusions.
The occurrence of Bohlinia adoumi in Chad has been disproven [36,38,49,58]. Samotherium boissieri has not been recorded in Africa, and few remains are attributed to the genus at As Sahabi (Libya), Bou Hanifia (Algeria), and Nakali (Kenya; [49]). The genus Traportax had a wide distribution in Eurasia and Africa during the Late Miocene, while the species Tragoportax rugosifrons was spread in the Greco-Iranian bioprovince [32,59] (Table 5).
The mammal assemblage of Cessaniti is not comparable to the other ones occurring in Italy, in the northmost area of the Central Mediterranean. The Tusco-Sardinian and Apulo-Abruzzi bioprovinces are characterized by endemic mammals that evolved under insular conditions [25].
The possible relationship with the mammal assemblage from the Sicilian site of Gravitelli (Messina, Sicily; Figure 1a) is still pending. The sites have been tentatively considered as part of the same bioprovince for the non-endemic characteristics of their faunas and their proximity [3,4,22,25,60], but they possibly have only the genus Ceratotherium in common. The faunal assemblage, updated by modern studies carried out on museum casts and figures by Seguenza [61,62] due to the loss of original fossils, consisted of Mesopithecus sp., Metailurini indet., Hyaenictitherium hyaenoides, Plioviverrops orbignyi, Zygolophodon turicensis, Ceratotherium sp., Bovidae indet., Hexaprotodon? siculus, and Propotamochoerus provincialis [25,60,63,64,65,66]. The mammal assemblage of Gravitelli has prevalent European affinity with maybe two African elements, the new species Hexaprotodon? siculus and Ceratotherium sp.; moreover, a recent revision moved the original Messinian age to a time older than 7 million years [66]. In the state of the art, no firm conclusions can be drawn.
From a paleoecological point of view, data from the mesowear and anatomy suggest that Bohlinia attica and Samotherium boissieri were browsers [67]. Tragoportax rugosifrons was a grazer occupying an ecological niche comparable to that of the extant Hippotragus, a dweller of forest–savannah alternating with open spaces [68].
These dietary categories are consistent with a mosaic environment with wooded areas alternating with an open environment. This reconstruction is also coherent with the presence of Stegotetrabelodon, a ground-dwelling grazer–browser, and Ceratotherium, a ground-dwelling grazer.

5. Conclusions

  • The specimens from Cessaniti enlarge the available data on Bohlinia attica.
  • The occurrence of Bohlinia attica at Cessaniti, associated with Samotherium boissieri, confirms the westward expansion of the Pikermian biome.
  • The mammal assemblage from Cessaniti includes taxa with Greco-Iranian and Afro-Arabian affinities.
  • The dispersal is still not known for the geological complexity of the Central Mediterranean area.
  • The North African fossil record is too discontinuous to define the pathway and timing of dispersal to Cessaniti.
  • Nonetheless, the evidence of the expansion of the Pikermian biome in the Late Miocene is well-documented and probably linked to the increasing aridity.
  • The significance of the Cessaniti assemblage in an area as complex as the Central Mediterranean deserves further research.

Funding

This research was funded by the University of Messina, FFABR 2022/23, assigned to A.C. Marra.

Data Availability Statement

The original contributions presented in this study are included in the article. Further inquiries can be directed to the corresponding author.

Acknowledgments

The author is deeply indebted to: Fabrizio Sudano and Michele Mazza, the past and actual superintendents, respectively (SABAP of Reggio Calabria, Italy; sabap-rc.cultura.gov.it) for allowing this research; to Maurizio Paoletti, president of the SiMU, and Adriano Guido, responsible for the SiMU Section of Paleontology, the University of Calabria, for their support; the Geosciences staff for the kindness in every step of the process; last but not least, the four anonymous reviewers for their helpful suggestions which improved the paper.

Conflicts of Interest

The author declares no conflicts of interest.

Abbreviations

The following abbreviations were used in this manuscript:
SiMUMuseal System of the University of Calabria, Section of Palaeontology, Via Pietro Bucci 87036 Rende, Cosenza, Italy; abbreviation: SiMU
MU.RICity Museum of Ricadi, Via Roma, 12, 89866 Santa Domenica, Vibo Valentia, Italy: abbreviation: MU.RI Directory of open access journals

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Figure 1. (a) Geographic locations of the sites cited in this text: (b) simplified geological map of the area of Cessaniti–Cape Vaticano–Mount Poro (square in (a); modified from Gramigna et al.’s study [11]).
Figure 1. (a) Geographic locations of the sites cited in this text: (b) simplified geological map of the area of Cessaniti–Cape Vaticano–Mount Poro (square in (a); modified from Gramigna et al.’s study [11]).
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Figure 2. Paleogeography of the Peri-Tethys (modified from Meulenkamp and Sissingh’s study [26]); black lines indicate faults; red lines partially delineate the modern geography.
Figure 2. Paleogeography of the Peri-Tethys (modified from Meulenkamp and Sissingh’s study [26]); black lines indicate faults; red lines partially delineate the modern geography.
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Table 1. List of fossils of land mammals of Cessaniti, with the exception of Bohlinia, described in this paper. Fossils are from Gentile’s quarry and correlatable deposits: * Papaglionti; ** Malopara.
Table 1. List of fossils of land mammals of Cessaniti, with the exception of Bohlinia, described in this paper. Fossils are from Gentile’s quarry and correlatable deposits: * Papaglionti; ** Malopara.
Stegotetrabelodon syrticus [1,2]cf. Ceratotherium advenientis [7]Tragoportax cf.
rugosifrons [5]		Samotherium boissieri [3,4]
Unit 2Mandible
Incisor
Fragmentary molar
Two fragmentary humeri
Right II metacarpal
Incomplete femur		Fragmentary skull
Two fragmentary teeth
Few postcranial bones		Hemimandible
radius
Astragalus
Anterior phalanx
humerus *
Anterior phalanx *
metatarsal *
Posterior phalanx *		Two metacarpals
Two metatarsals
Complete manus **
Fragmentary ulna **
Distal radius **
Fragmentary humerus **
Unit 1One worn DP4
Table 2. Measurements of M1-M3 length of compared Giraffidae.
Table 2. Measurements of M1-M3 length of compared Giraffidae.
Bohlinia atticaPalaeotragus
rouenii		Palaeotragus
quadricornis		Samotherium
boissieri
76 [36]73 [42]91 [43]111[40]
83[36]58 [43]74 [43]104.5[40]
79[36]67 [39] 104.3[40]
ca. 76[36]67.1[39] 105.5[40]
ca. 75[36]68.3[39] 105.9[40]
77[39]67.6[39] 91.8[40]
85.5[39]63.3[39] 103.8[40]
92.7[38] 11.13[40]
83.5[38] 110[40]
115[43]
105[43]
Table 3. Dental measurements of Bohlinia attica from Cessaniti and sites of Greece (Nikiti [39], Pikermi [36], Samos [38], and Ditiko [33]), the Republic of Macedonia (Kirokuçuk [36]), and Iran (Maragheh [38]). Legends: Lg—lingual side; Lb—labial side; A—anterior side; P—posterior side; mean—mean value; min.—minimum value; max.—maximum value; spec.—specimen’s inventory number.
Table 3. Dental measurements of Bohlinia attica from Cessaniti and sites of Greece (Nikiti [39], Pikermi [36], Samos [38], and Ditiko [33]), the Republic of Macedonia (Kirokuçuk [36]), and Iran (Maragheh [38]). Legends: Lg—lingual side; Lb—labial side; A—anterior side; P—posterior side; mean—mean value; min.—minimum value; max.—maximum value; spec.—specimen’s inventory number.
MeasureNikitiPikermi SamosKirokuçukMaraghehDitikoCessaniti
Mean Min. Max.Spec. Spec. Spec. Spec.
P3 length18.5517.819.2 CM45821.9 M35223.2 21.5
M35320.8
P3 breadth19.501723.4 CM45820.1 M35224.2 22.4
M35319.1
P4 length18.2716.520.6 19.8 M35224.8 Lg18.0
M35322.5 Lb17.5
P4breadth22.3519.426.0122 23.2A22.8M35224.8 26.4
223 B22.7M35319.2
M1 length25.1223.826.3 26.2 M35226.827Lg22.5
M35324.726.4Lb23.7
M1 breadth25.1524.027.4127 25A26.0M35227.7 A26.9
226 B26.7M35327.0 P26.7
M2 length26.7422.830.2 29.4 M35235.2 Lg26.4
M353- Lb28.6
M2 breadth28.1526.533.2 28.3 M35230 A31.5
M353- P29.4
M3 length27.525.731.0 31 M35231.4 Lg28.0
M353- Lb29.0
M3 breadth27.5626.030.0 28.8 M35228 A31.6
M353- P28.5
Table 4. Measurements of astragali compared with Giraffidae.
Table 4. Measurements of astragali compared with Giraffidae.
SpeciesRef.Lateral LengthMedial LengthDistal BreadthAnte_Post Distal Depth
Cessaniti specimen 8978.65955.6
Bohlinia attica[33] 8159
[48]87.7–103.4 (min.–max.)76.5–89.6 (min.–max.)55–68 (min.–max.)46.5–61.1 (min.–max.)
[47]1039976.6
[47]1059470
[47]1059171
[47]1089473
Palaeotragus rouenii[40]76.6 47.2
[40]73.56547
Samotherium boissieri[40]90.0878.557.6
[40]96.582.359.7
[40]97.585.158.4
[40]89.278.659.3
[40]89.079.258.0
[40]88.478.059
[40]86.873.756.4
[40]85.675.056.8
[40]89.376.261.0
[40]84.476.553.5
[40]88.877.655.0
[40]89.075.859.0
[40]86.077.255.7
[40]90.076.656.0
[40]92.281.062.3
Table 5. Distribution of the artiodactyls taxa belonging to the Cessaniti assemblage; data (https://nowdatabase.org/now/database/, accessed on 26 January 2025) by The NOW Community/CC BY 4.0, January 2025; *: several outcrops in the area.
Table 5. Distribution of the artiodactyls taxa belonging to the Cessaniti assemblage; data (https://nowdatabase.org/now/database/, accessed on 26 January 2025) by The NOW Community/CC BY 4.0, January 2025; *: several outcrops in the area.
LocalityCountryAge (Ma)Bohlinia
attica		Samotherium boissieriTragoportax
rugosifrons
AzmakaBulgaria7.2X
Gorna Susica Bulgaria8.3–7.285X
HadjidimovoBulgaria7.6–7.1X X
KalimantsiBulgaria7.6–7.1X
KrodimovoBulgaria7.6–7.1X
StrumyaniBulgaria7.6–7.1X X
KirokuçukN Macedonia8.9–5.3X
Dolni DisanN Macedonia8.9–5.3X
KaraslariN Macedonia7.6–7.1X X
DitikoGreece7.1–5.3X
KerassiaGreece8.9–7.1X
NikitiGreece9.9–8.9 X
Pikermi *Greece7.454–7.1X
Pyrgos VassilissisGreece7.212–7.14X
Ravin de la PluieGreece9.426–9.311X
Ravin Zouaves *Greece9.9–8.1X
Samos *Greece8.9–5.3 XX
Vathylakkos *Greece7.489–7.454X
Corakyerler Turkey8.9–7.6XX
Esendere Turkey8.9–7.6X
KüçükçekmeceTurkey8.9–5.3X
Gülpinar Turkey7.6–7.1 X
KarainTurkey8.9–7.1 X
KavakdereTurkey8.254–8.108 X
Sinap *Turkey11.2–7.1X
InjanaIraq8.9–7.6XX
Maragheh *Iran8.9–7.1X X
Novo-ElizavetovkaUkraine8-9–7.1 X
TarakliaMoldova7.6–7.1 XX
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Marra, A.C. Out of Pikermi: The Occurrence of Bohlinia in the Late Miocene of the Central Mediterranean. Geosciences 2025, 15, 44. https://doi.org/10.3390/geosciences15020044

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Marra AC. Out of Pikermi: The Occurrence of Bohlinia in the Late Miocene of the Central Mediterranean. Geosciences. 2025; 15(2):44. https://doi.org/10.3390/geosciences15020044

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Marra, Antonella Cinzia. 2025. "Out of Pikermi: The Occurrence of Bohlinia in the Late Miocene of the Central Mediterranean" Geosciences 15, no. 2: 44. https://doi.org/10.3390/geosciences15020044

APA Style

Marra, A. C. (2025). Out of Pikermi: The Occurrence of Bohlinia in the Late Miocene of the Central Mediterranean. Geosciences, 15(2), 44. https://doi.org/10.3390/geosciences15020044

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