Five new species of Hoplitomeryx from the Neogene of Abruzzo and Apulia (central and southern Italy) with revision of the genus and of Hoplitomeryx matthei Leinders, 1983 Author Mazza, Paul P. A. Author Rustioni, Marco text Zoological Journal of the Linnean Society 2011 2011-11-25 163 4 1304 1333 http://dx.doi.org/10.1111/j.1096-3642.2011.00737.x journal article 10.1111/j.1096-3642.2011.00737.x 0024-4082 5442183 HOPLITOMERYX MINUTUS SP. NOV. ( TABLE 1 , FIGS 1 , 5 ) Holotype : Right hemimandible SCT 77. Paratypes : Mandibles SCT 60, SCT 73, SCT 78, SCT 155, RGM 261.147 , RGM 261.136 . Type locality and horizon: Holotype and paratypes tagged SCT from the Tortonian Scontrone Member of the Lithothamnium Limestone ( Patacca et al ., 2008 ; 41°45′15.54″N , 14°2′13.14″E ), outskirts of Scontrone , southern border of the National Park of Abruzzi , L’Aquila , central Italy . Paratype RGM 261.147 from the Messinian karstic fissure filling called Chiro 29 in the limestone quarry Chiro , and paratype RGM 261.136 from the Messinian karstic fissure filling called Nazario 4 in the limestone quarry Nazario. Both quarries between Apricena and Poggio Imperiale, Foggia, Gargano promontory, south-eastern Italy ( 41°48′12″N , 15°23′04″E ) . Preservation and deposition of type specimens: Scontrone specimens: Soprintendenza Archeologica dell’Abruzzo ( Chieti , central Italy ); Gargano specimens: Museum Naturalis , Leiden (the Netherlands) . Etymology: The name refers to the small dimensions of the species. Diagnosis: Apomorphies of species: mandible with horizontal ramus slender and sinuous, and with uniformly convex ventral profile; lower cheek dental formula: 3– 3 in the Scontrone specimens, 2–3 in the Gargano specimens; brachyodont lateral dentition ( HI = 0.67); lower premolars very small with no paraconid, robust and salient metaconid and strong entoconid; lower molars with widely spaced, triangular labial conids, flattened mesial enamel walls, isolated hypoconid, even in heavily worn teeth, strong metastylids, postentocristids, and ectostylids; distal margin of hypoconulid separated from entoconulid towards the occlusal edge of the crown and fused with it towards the collar; lingual enamel wall smooth, labial enamel wall moderately corrugated; molars with cingulum on the mesiolabial base of protoconids. Differential diagnosis: See Table 1 . Description of holotype : Fragment of horizontal ramus, fairly low and slender, with sharp ventral border. Mandibular scissure barely marked. The specimen preserves the three lower molars. Labial conids markedly triangular, inclined outwards and backwards and fairly widely separated from one another. Mesial walls flat. Preprotocristid abruptly bent lingually forming a marked angle. Mesial fossette opened lingually. Postprotocristid never fused to other structures keeping free all the way down to base of crown. Hypoconid isolated from other conids. Robust wear-resistant lingual ribs and stylids, which give teeth high occlusal relieves. Ectostylids in all molars. Well-developed mesiolabial cingula at the base of protoconids. In m3 distal margin of hypoconulid occlusally separated from entoconulid, rapidly fused to it towards base of tooth. Labial enamel moderately corrugated, lingual enamel smooth. Additional characters shown by paratypes : Horizontal ramus tapers gradually rostrally. Lower premolar to molar ratio: p2-p4/m1-m3: 0.50; p3-p4/ m1-m3: 0.38 in the Scontrone specimens; not assessable in the available Gargano specimens. Very thin film of cementum-like veneer occasionally present in labial valleys of lower molars. p2. Tooth stout, somewhat compressed mesiodistally and enlarged labiolingually. Parastylid well developed, paraconid absent. Metaconid the highest cuspid, it protrudes considerably lingually. Entoconid well developed, entostylid absent. Lingual enamel wall smooth. p3–p4. Morphologically similar to p2, but larger and proportionally more elongated mesiodistally. Parastylid well developed, paraconid rudimentary and Table 1. Differential diagnoses of dental characters Hoplitomeryx (genus characters) H. minutus H. falcidens H. apulicus H. matthei H. apruthiensis H. magnus Check teeth brachyodont Brachyodont lateral Mesodont Brachyodont lateral Mesodont Mesodont or mesodont dentition dentition Lower cheek dental Lower cheek dental Lower cheek dental Lower dental formula: in Lower cheek dental formula: 3– 3 in the formula: 3– 3 in the formula: 3– 3 in the Gargano specimens: formula: 2–3 Scontrone specimens, Scontrone specimens, Scontrone specimens, 2–3, unknown in 2–3 in the Gargano 2–3 in the Gargano 2–3 in the Gargano Scontrone specimens specimens specimens representatives at the moment Variously developed parastyle and mesostyle in upper cheek teeth Sabre-like upper canines Strong, slender, and can Sometimes be sharp upper Canines present Premolars compressed mesiodistally and enlarged labiolingually Upper molars often Molars elongated wider than they are mesiodistally and long somewhat compressed labiolingually P3 with centrally situated and lingually directed protocone P4 and M3 with large metaconule Upper molars with Upper molars very triangular protocone similar to one another, and trapezoidal with trapezoidal, metaconule closely packed lingual crescents Lingual cusps very tightly spaced in M1, somewhat more separated in M2 and M3 Protocone somewhat smaller than metacone Metacone sometimes Weak external rib of with small pillars on metacone labial face of upper molars Labial wall of metacone almost flat Strong external paracone Paracone supported by rib prominent median rib Mesodont Lower cheek dental formula: unknown at the moment in the Scontrone representatives, 2–3 in Gargano specimens Presence of upper canines External rib of metacone absent Paracone supported by robust and prominent median rib Table 1. Continued Hoplitomeryx (genus characters) H. minutus H. falcidens H. apulicus H. matthei H. apruthiensis H. magnus Incipient entostyle Entostyle (in upper No entostyles (in upper Tiny entostyle on second occasionally from teeth absent) teeth) molars mesial wall of metaconule of first and second upper molars Cingula absent in upper Thin cementum cover cheek teeth Lower molars with In lower molars labial Labial conids closely widely spaced labial cuspids widely spaced conids, spaced and somewhat backward-verging Lower molars with In lower molars labial Labial conids triangular labial cuspids triangular markedly triangular conids, Flattened mesial Flattened mesial walls enamel walls Check teeth with globose cuspids and stylids not prominent lower molars with swollen lingual enamel walls and with cuspids that remain isolated even in worn teeth Check teeth with occlusal surfaces markedly inclined outwards because of strong differential wear between inner and outer cuspids M1 with very reduced entostyle Cementum absent and overall smooth enamel in upper cheek teeth Lower molars with lingual cuspids relatively less worn than labial cuspids Lower molars with widely spaced, triangular, backward-verging labial cuspids Flattened mesial enamel walls In worn molars protoconid connected mesially with metaconid and distally with entoconid, and hypoconid connected with postprotocristid and postentocristid p4 nonmolarized Elongated Mesiodistally Elongated Compressed Compressed compressed trigonid and small talonid Lingually open mesial fossette Small paraconid Lower premolars very Paraconid absent or Paraconid sometimes Paraconid present in Paraconid absent or occasionally present small with no present only in present in unworn unworn or poorly worn rudimentary paraconid, robust unworn lower lower premolars lower premolars and salient premolars entoconid well metaconid, and developed in lower strong entoconid premolars In p2 protoconid and metaconid of same height In p3 protoconid larger than metaconid, opposite in p4 Isolated hypoconid Hypoconid well developed in lower premolars Parastylid well developed in lower premolars Metastylid generally Strong metastylids Metastylids well Metastylids well Poorly developed stylids Metastylid strong well developed in developed and developed but not lower molars protruding lingually protruding lingually Ectostylid occasionally Strong ectostylids Ectostylid low and Ectostylid very small Ectostylid small in m1, Small extostylid on each present, sometimes robust in m1, low in m1, absent in absent in m2 and m3 lower molar quite developed, on and blunt in other other molars lower molars molars Complete postentocristid Strong postentocristids Postentocristids well Postentocristids well Postentocristids not Postentocristid strong in lower molars developed and developed but not protruding lingually protruding lingually protruding lingually (in m3 postentocristid hardly developed) Infrequent mesiolabial Molars with cingulum Weak cingulum at the Cingulum absent (in Cingula absent (in lower Cingula absent in lower cingulum on the mesiolabial base of the lower teeth) teeth) cheek teeth base of protoconids protoconid in m1 and m2 Third lower molars with wide, mesiolingually open mesial fossette Elongated In p3 and p4 paraconid barely developed or totally absent Protoconid and metaconid of same height protoconid isolated in unworn molars Hypoconid isolated in unworn molars Metastylid prominent, but blunt Ectostylids blunt and robust Postentocristid prominent, but blunt Protoconids with faint traces of cingulum at their base present only in barely worn teeth, otherwise totally absent; metaconid robust, very salient, exceeding in height the cupsids of all lower premolars, and in contact with protoconid. Entoconid well developed and protruding lingually. Entostylid also well developed. Lingual enamel wall smooth. Table 1. Continued

Hoplitomeryx
(genus characters)	H. minutus	H. falcidens	H. apulicus	H. matthei	H. apruthiensis	H. magnus
Distal lobe of m3	Distal margin of	Hypoconulid and	Distal margin of	Distal margin of	Distal margin of	Third lower molars with
constantly bilophed,	hypoconulid	entoconulid with	hypoconulid fused,	hypoconulid separated	hypoconulid separated	tear-shaped
normally constituted	separated from	distal margins	or not, to the	from entoconulid	from entoconulid	hypoconulid and
by crescent	entoconulid towards	separated to halfway	entoconulid	occlusalwards and	occlusalward, and	entoconulid not fused
hypoconulid and large,	the occlusal edge of	the crown height	fused to it towards the	fused to it towards	distally
tear-shaped	the crown and fused	collar	the collar
entoconulid, both with	with it towards the
distally extending	collar
cristids, not fused	Cementum occasionally
together along the	present
distal margin of the	Lingual enamel wall	Enamel smooth	Enamel smooth	Labial enamel wall	Lingual walls somewhat	Labial enamel wall
tooth or otherwise	smooth, labial	corrugated, lingual	more corrugated than	corrugated, lingual
fused, but only in	enamel wall	wall smoother	labial walls in upper	wall smooth in lower
fairly worn teeth	moderately	check teeth, opposite	check teeth
corrugated	in lower ones
Mandible slender or	Mandible with	Mandible slender, with	Horizontal ramus of	Mandibles with	Mandible with slightly	Mandible with
pachyostotic	horizontal ramus	moderately sinuous	mandible slender	horizontal ramus	sinuous horizontal	horizontal ramus
slender and sinuous,	horizontal ramus	and slightly sinuous,	pachyostotic	ramus	massive lateromedially
and with uniformly	and convex ventral	with straight or
convex ventral	profile under cheek	slightly convex
profile	teeth corpus	ventral profile
Long diastema
between lateral
dentition and incisor
corpus
Mandibular scissure	Marked mandibular
slightly marked	scissure

Figure 1. A, Hoplitomeryx minutus sp. nov. , holotype right mandible (SCT 77): lingual view. B, Hoplitomeryx minutus sp. nov. , holotype right mandible (SCT 77): labial view. C, Hoplitomeryx minutus sp. nov. , holotype right mandible (SCT 77): occlusal view. D, Hoplitomeryx falcidens sp. nov. , holotype left mandible (SCT 177): labial view. E, Hoplitomeryx falcidens sp. nov. , holotype left mandible (SCT 177): lingual view. F, Hoplitomeryx falcidens sp. nov. , holotype left mandible (SCT 177): occlusal view. G, Hoplitomeryx falcidens sp. nov. , right maxillary (RGM 260.941): labial view. H, Hoplitomeryx falcidens sp. nov. , right maxillary (RGM 260.941): lingual view. I, Hoplitomeryx falcidens sp. nov. , right maxillary (SCT 70): occlusal view. m1. Tooth tightly in contact with fourth premolar. External rib of metaconid and entoconid strong, metastylid and postentocristid also quite robust. Robust ectostylid. Mesiolabial cingulum variously developed and at times supporting tiny cuspule. m2. Labial cuspids widely separated from one another with low, blunt ectostylid in between. m3. Low, pillar-like enamel structure sometimes present where distal margins of hypo- and entoconulid meet. Measurements: See Table 2.