Journal of Paleolithic Archaeology

, Volume 2, Issue 4, pp 359–377 | Cite as

Paleobiology and Taphonomy of a Middle Paleolithic Neandertal Tooth from Ciemna Cave, Southern Poland

  • John C. WillmanEmail author
  • Bolesław Ginter
  • Raquel Hernando
  • Marina Lozano
  • Krzysztof Sobczyk
  • Damian Stefański
  • Anita Szczepanek
  • Krzysztof Wertz
  • Piotr Wojtal
  • Mirosław Zając
  • Katarzyna Zarzecka-Szubińska
  • Paweł Valde-Nowak


Middle Paleolithic human remains from more northern Central Europe are rare and fragmentary despite a relative abundance of archeological remains in this region from this time period (Fig. 1a). To date, Neandertal remains in Poland are limited to three isolated molars from Stajnia Cave (Urbanowski et al. 2010; Dąbrowski et al. 2013; Nowaczewska et al. 2013). Additional fragmentary Neandertal fossils from the upland zone of Central Europe come from Švédův stůl (Ochoz) (Vlček 1969) and Kůlna Cave (Jelínek 1988) in the Moravian Karst, and Šipka Cave near Štramberk, Czech Republic (Svoboda et al. 1996). Neandertal remains from the Carpathians are limited to Šal’a 1 (Sládek et al. 2002), Šal’a 2 (Jakab 2005), and Gànovce 1 (Vlček 1955) in Slovakia; Subalyuk 1 and 2 (Pap et al. 1996) in Hungary; and Bordul Mare 1 (von Gaál 1928) further east in Romania. Given the extensive occupation of at least southern Poland into neighboring Moravia and the Carpathians during the Middle Paleolithic, the dearth of human remains raises questions regarding the roles of humans in site formation processes in the region. A fragmentary human tooth from Ciemna Cave in southern Poland, a partial mandibular incisor (Ciemna 1), provides additional data relevant to these questions and expands our knowledge of Neandertal behavior in the region.
Fig. 1

a and b Location of Ciemna Cave in relation to other Middle Paleolithic sites in Central Europe where human remains have been discovered

The Archeological Context

Ciemna Cave

Ciemna Cave is located in the Kraków-Częstochowa Upland, in the Prądnik Valley near Ojców, at ≈ 375 m asl and 80 m above the valley bottom (Fig. 1b). It is an extensive cave system formed in Jurassic limestone (Gradziński et al. 2007) (Fig. 2). The karstic processes caused slope destruction and partly exposed its western portions. The complex filling of the cave reflects the changing depositional circumstances, including inner (clays) and outer eolian agents (loess) during the Late Pleistocene. The localization of favorable occupation areas, such as terraces or openings, influenced the patterns of human site use through time (Valde-Nowak et al. 2016a, b) (Fig. 2).
Fig. 2

Ciemna Cave chamber outline, trenches, and location of the Ciemna 1 incisor

The investigation of Ciemna Cave started at the beginning of the nineteenth century (Czarnowski 1924). It focused mostly on the western part of the cave system, especially the Ogrojec (sector CO)—a large, roofless chamber—which opened during the first Pleniglacial (MIS 4) and provided favorable conditions for settlement, as is indicated by a Micoquian occupation. Test excavations of Holocene layers by J. Czarnowski took place at the beginning of the twentieth century (Czarnowski 1924). Later excavations in 1918–19 (Krukowski 1939–1948) uncovered substantial evidence for an Ojców industry and Prądnician cycle characterized by the presence of Prądniks and the Prądnik technique which is now included within the Micoquian Middle Paleolithic. Test excavations of the Ogrojec occurred again in the 1960s by Kowalski (1967, 2006), providing archeological insights similar to the earlier test excavations. A dense and compacted Micoquian level was documented within the MIS 3 interpleniglacial member covered by a thick loess layer.

Ongoing research at Ciemna Cave has been carried out since 2007. A primary focus has been the first test excavation within the inner part of the system—primarily the main chamber (sector CK). Ground-penetrating radar was used to assess the depth of the fill before excavations (Szynkiewicz 2007) and during excavations (Karczewski and Ostrowski 2013). Over 42 m2 of adjoining trenches were excavated (Fig. 2). All excavated lithics and bones were coordinated with a total station and georeferenced with photogrammetry (Valde-Nowak et al. 2016b). Arbitrary 10 cm spits in 1 × 1 m grid units were used to collect sediments for wet sieving though 1 mm mesh. Plant and animal macrofossils were obtained by flotation and wet sieving (0.5 mm mesh) of samples from each unit (Valde-Nowak et al. 2016b). During recent excavations, a rich geological sequence and archeological levels (Table 1, Fig. 3) characterized by relatively stable horizontal and vertical deposition were uncovered and documented in the main chamber (Valde-Nowak et al. 2014, 2016a). The stratigraphy of sector C (the area where the main chamber and the one parallel to the hillside of the valley joined each other near the modern entrance to the cave) was published by Krukowski (1924) and Kowalski (2006) but not studied in detail. So far, seven archeological levels document Late Pleistocene cultural contexts (IX to III from the bottom) in the main chamber with several Middle Paleolithic (Mousterian, Levallois-Mousterian, Taubachian, and Micoquian) lithic traditions represented (Table 1).
Table 1

Archeological sequence and radiocarbon dating of Ciemna Cave main chamber (sector CK). Shading indicates context of human incisora

Fig. 3

Lithostratigraphic context of the Ciemna 1 incisor (star) in layer 3. Layers are numbered 1–19. Section and profile along the eastern wall of the excavation

In the faunal assemblage, a paucity of herbivore remains is contrasted by an exceptional number of carnivore remains in the main chamber (sector CK), as well as in the more exterior Ogrojec section (CO) (Valde-Nowak et al. 2016b; Wojtal 2007). The dearth of herbivore remains suggests little or no transport of herbivore carcasses to Ciemna Cave by humans or carnivores (Valde-Nowak et al. 2016a). Ursids dominate the sequence containing the human tooth (Table 2, Online Resource Fig. 13), and the majority of identifiable carnivore remains with signs of digestion are from bears (Table 3). Bears will cannibalize and gnaw on other bears in hibernation dens (Rabal-Garcés et al. 2012; Stiner 1999; Sala and Arsuaga 2013; Sala et al. 2014), and digested cave bear bones recovered from Ciemna Cave suggest that this occurred (Table 3). Sediment from the main chamber contained thousands of deciduous cave bear teeth (NISP = 5535) as well as limb bones from very young bears. Therefore, the overall density of bear remains, and especially the abundance of very young individuals, suggests that Ciemna Cave was frequently used as a hibernation den and only occupied by humans in the absence of carnivores (Wojtal 2007; Valde-Nowak et al. 2016a, b).
Table 2

Non-human carnivores recovered from the hominin-bearing lithostratigraphic layer 3–4 of the main chamber (sector CK) of Ciemna Cave


Layer 3–4



  Canis lupus


  Canis sp.


  Crocuta spelaea




  Ursus speleus


  Ursus sp.


  Vulpes vulpes


  Vulpes sp.


Table 3

Non-human fauna exhibiting signs of digestion in hominin-bearing lithostratigraphic layers 3–4 of the main chamber (sector CK) of Ciemna Cave


Layer 3–4 (NISP)


Total number of digested bones






cf. Equus ferus (NISP = 2)


Magnum, sesamoid

cf. Rupicapra rupicapra (NISP = 2)


Phalanx II, capitatotrapezoideum

Ursus spelaeus


Naviculare, phalanx II, phalanx I, phalanx I

Ursus sp.


Metacarpal/metatarsal, phalanx I, phalanx I, pisiforme, sesamoid, sesamoid, lower second incisor

The present data concerning the Middle Paleolithic human incisor comes exclusively from the recent excavations (trenches I and II) in the Main Chamber as the further finds are still under ongoing investigation.

Archeological Context for the Ciemna 1 Incisor

The tooth was found during routine sediment screening. It is from archeological level III, the most recent Micoquian level, within lithostratigraphic level 2.3 and 3 in geological series IV (layers 5–2.2), which consist of brown and grayish-brown loams, silty loams, and loess with limestone rubble (Table 1, Figs. 2 and 3) (for additional stratigraphic context see Valde-Nowak et al. 2014, 2016a, b). The limestone rubble is weathered and smoothed, suggesting a mild climate during deposition of these layers (Valde-Nowak et al. 2014).

Archeological level III exhibits the greatest concentration of artifacts (n = 150), that includes cores, blanks, and waste as well as numerous distinct tool types (bifacial and partly prepared knives, notched and denticulate sidescrapers, and retouched flakes). However, the density of the scattered cultural material is low, since the potential living area is relatively vast—stretching across the full extent of the chamber (Fig. 2). A relatively large number of flakes, a few cores with refitted elements, bifacial knives, and a bone retoucher (Ginter et al. 2015) suggest a processing area, probably at the periphery of more stable settlement areas. These finds are a valuable source of counterpoint to the palimpsest record in the Ogrojec, which seems to have offered the most favorable occupational location during that time (Valde-Nowak et al. 2016a).

Archeological level III of the main chamber was dated by several radiocarbon methods, but different sample materials (i.e., bone, charcoal, and unburned wood), and dating methods provided results that are difficult to interpret (Alex et al. 2017). All radiocarbon dates from bone samples from Ciemna archeological levels III, IV, and V produced minimum ages, or dates beyond the radiocarbon calibration curve, whereas a charcoal sample produced a significantly younger age than bone from the same stratigraphic sequence (Table 1) (Alex et al. 2017). However, an undisturbed process of sediment formation and a lack of subsequent disturbances in this part of the cave system are supported by detailed spatial analyses of flint artifact and faunal remains (Valde-Nowak et al. 2014, 2016a). Furthermore, refittings—some of relatively small pieces—of flint products from archeological level III (Valde-Nowak et al. 2016a), and ongoing analyses of animal bone refittings by one of us (K. Zarzecka-Szubińska) indicate homogeneity of layers and minimal postdepositional movement of materials within the archeological levels containing the human tooth. In this light, the recent date from unburned wood (Alex et al. 2017) is best considered an intrusion incurred during modern excavations.

The latest dates, based on screened bone collagen of long bones controlled with FTIR, suggest the Micoquian layers are older than 50 ka BP (Alex et al. 2017), and corresponds well with the dating of other Micoquian sequences at Kůlna (Neruda 2017), Stajnia (Urbanowski et al. 2010; Dąbrowski et al. 2013; Nowaczewska et al. 2013), and Obłazowa (Alex et al. 2017). Thus, it is relatively certain that the human incisor, found within lithostratigraphic layer 2.3 and 3, is dated to the Micoquian given that all the bone remains from the dated archeological levels III, IV, and V (lithostratigraphic layers 2.3–8) have been dated to the Micoquian and contain Micoquian artifacts (Table 1). Thus, given the associated material culture, dating, and stratigraphic integrity of the archeological layers, as well as presence of diagnostic Neandertal fossils of similar chronologies in the region, we can be reasonably assured that Ciemna 1 belongs to a Neandertal despite an absence of diagnostic dental morphology (see below).

Materials and Methods

The state of preservation of Ciemna 1 severely limits potential observations. Macroscopic observations and standard mesiodistal dimensions of the crown were made on the original specimen.

To facilitate microscopic examination using an environmental scanning electron microscopy (ESEM: FEI Quanta 600), the original tooth was cleaned with 96% alcohol, and a mold was made with President Plus Jet light body polyvinylsiloxane (Coltene Whaledent), and a positive cast was made using Epo-Tek 301 epoxy (Epoxy Technology, Inc). The cast was sputter-coated with ~ 20 nm of gold and observed using a secondary electron detector, under high vacuum, at 20 kV, and a working distance > 20 mm.


Morphological Description and Paleobiological Considerations

The Ciemna 1 incisor is heavily modified by postmortem taphonomic processes (Fig. 4), which makes the identification of tooth position and side difficult. Labiolingual breadth is not measurable as the entire lingual surface is fractured and missing postmortem. In labial view, an oblique, postmortem fracture traverses the root from approximately the mesial cervix to the distal side of the root, leaving only a few millimeters of root intact. Very little mesial enamel is present aside from that wrapping around from the labial surface. The fractures exposed the pulp chamber postmortem. The estimated labial crown height is 6.0 mm and the total height with the crown plus root fragment is 9.7 mm.
Fig. 4

The Ciemna 1 mandibular incisor in occlusal (Oc), distal (Di), labial (La), mesial (Me), and lingual (Li) views. Scale bar represents 1 mm increments

In occlusal view, the Ciemna 1 crown exhibits a degree of asymmetry that suggests that it is a right mandibular second incisor (Fig. 4). Despite breakage, the root fragment exhibits a distal curvature consistent with right I2 morphology. The Ciemna 1 mesiodistal crown breadth is small (5.3 mm). The small breadth is likely an artifact of postmortem damage of the crown and an advanced stage of occlusal wear. Neandertal mean I1 mesiodistal breadth is 5.5 mm (SD = 0.5 mm, range = 4.4–6.5 mm, n = 12), and Neandertal mean I2 mesiodistal breadth is 6.5 (SD = 0.4 mm, range = 5.8–7.4 mm, n = 25). The Ciemna 1 mesiodistal breadth is suggestive of a mandibular first incisor, but labial convexity, crown asymmetry, and root curvature are more consistent with a right mandibular second incisor designation. Ciemna 1 is reliably designated as a right mandibular permanent incisor, and it is most likely an I2 based on morphological attributes.

Although the lingual half of the occlusal surface is missing, the labial enamel rim is intact, and dentin exposure can be estimated as stage 4 or 5 (Smith 1984). Incisor wear is an unreliable indicator of age-at-death as it can vary extensively relative to wear on the postcanine dentition, the latter providing a more accurate assessment of age. However, the degree of incisor wear on Ciemna 1 is within the range of variation documented elsewhere for younger adult (≈ 20–40-year-old) Neandertal mandibles (see Trinkaus 2011: Table S1). Therefore, the Ciemna 1 individual was likely an adult aged between 20 and 40 years at the time of death.

The labio-occlusal enamel edge has a modest degree of antemortem rounding. A well-developed interproximal groove (stage 4: Estalrrich et al. 2017) is visible below the cervix of the lateral root surface (Fig. 5). The groove can be discerned from non-carious cervical lesions (Grippo et al. 2012; Towle et al. 2018; Michael et al. 2010), by the presence of parallel, and labiolingually oriented, striations within the borders of the groove (Bermúdez de Castro et al. 1997; Ungar et al. 2001; Hlusko 2003; Bouchneb and Maureille 2004; Estalrrich et al. 2017) (Fig. 5). Within the groove, there are numerous wide (~ 30–70 μm), subparallel striations, many of which preserve microstriations within the borders of each larger striation (Fig. 5). As observed elsewhere (Bermúdez de Castro et al. 1997), several striations in the labial portion of the groove deviate at low angles before becoming straight and subparallel as they traverse lingually (Fig. 5). Other striations trace the border of the inferior border of the enamel at the cementoenamel junction (Fig. 5). The microstriations within the larger striations suggest abrasion using a probe with an irregular tip. Striation morphology more closely resembling experimental grooves made with rigid, or semi-rigid, probes (e.g., reindeer antler with abrasive particles or wood and abrasive particles: Bouchneb and Maureille 2004) than those made with more flexible grass stalks (Hlusko 2003; Estalrrich et al. 2017) or calf sinew with abrasives (Bouchneb and Maureille 2004).
Fig. 5

Detail of distal interproximal groove on the cervix of Ciemna 1. a Dashed outline of interproximal groove in the same view as b. b ESEM micrograph of the distal surface. c ESEM focus-stacked micrograph from an oblique, labio-distally oriented view. Note that the semicircular cross-section of the groove is more accentuated labially than lingually. d Detail of microstriations in the labial portion of the groove using light microscopy. Note that some striations deflect at low angles at the base of the groove, some trace the border of the CEJ, but most are subparallel along the length of the groove. Scales are 1 mm (2 × 0.5 mm increments)

The length of the interproximal groove, although incomplete due to postmortem damage, is approximately 3.9 mm. The width of the groove at the labial-most portion of the groove is 1.3 mm and the distal-most portion—at the approximate midpoint of the tooth if it were unbroken—is 1.1 mm. The wider labial versus lingual width of the groove is common morphology for “toothpick” grooves (Frayer and Russell 1987; Bermúdez de Castro et al. 1997; Ungar et al. 2001; Estalrrich et al. 2017). The semicircular cross-section of the groove is most pronounced at maximum groove width. The length of the Ciemna 1 groove, despite postmortem truncation, is long compared with those from the anterior dentitions of Neandertals from El Sidrón and Hortus, but the width is similar to that of the Hortus Neandertals (Estalrrich et al. 2017) (Table 4).
Table 4

Ciemna 1 interproximal groove dimensions compared to those in the anterior teeth of Neandertals from El Sidrón and Hortus (Estalrrich et al. 2017)a


Mesial surface

Distal surface






Ciemna 1




El Sidrón


3.3 (0.9)

0.6 (0.2)

2.8 (0.7)

0.6 (0.2)



2.8 (0.7)

1.4 (0.3)

2.7 (0.2)

1.1 (0.2)

aMeasurements provided as “average (standard deviation)”

bLength is based on presence of only the labial half of the root surface

cWidth for Ciemna 1 taken just anterior to root fracture

Taphonomic Considerations

The edges of the broken root exhibit only subtle edge rounding which is probably a result of modest soil acidity or sediment abrasion (Fernández-Jalvo and Andrews 2016) (Fig. 6a). Sediment is still present in the root canal indicating that the breakage of the root was not a recent phenomenon (Fig. 4). The vertical splitting of incisors at midline is not uncommon in similarly and lesser-worn Neandertal mandibular incisors (e.g., Feldhofer NN67, Švédův Stůl, Spy 571a, Vindija VI-201, and Vi-286: Willman, personal observation) and could result from changes in humidity or temperature (Hughes and White 2009; Fernández-Jalvo and Andrews 2016). Similar to the incisal enamel surface, the fractured edges of the occlusal dentin also appear rounded in macroscopic observations (Fig. 4), but the ESEM images show that the edges of the occlusal dentin surface fractures are postmortem (i.e., not rounded or polished like the occlusal dentin). The dentin on the mesial half of the occlusal surface contains four superficial, postmortem cracks—again, a likely result of changing humidity or temperature (Fig. 6c).
Fig. 6

Scanning electron micrographs of postmortem damage to mesial and occlusal surfaces of the Ciemna 1 incisor. a Postmortem fracture of root and enamel surface. b Dashed yellow oval surrounding area of enamel erosion or abrasion. Yellow arrows point to enamel microcracking. c Yellow arrows indicate corroded pits, red arrows point to dentin and enamel splitting and cracking, blue oval and arrow indicate postmortem enamel and dentin chipping. d Red arrows point to parallel postmortem “trampling” features. e Yellow arrows point to corroded pits, blue arrows indicate postmortem enamel and dentin chipping. All scales are 1 mm

Short and shallow striations with “sharp” edges located near midline resemble “trampling”-induced sediment abrasion (Martínez and Pérez-Pérez 2004). This alteration is characterized by multiple parallel striations from the compression and sliding of the enamel surface across a hard and abrasive substrate (Fig. 6d). Patches of altered enamel exhibiting cracks and erosion also indicate some degree of postdepositional enamel alteration (Martínez and Pérez-Pérez 2004) (Fig. 6b). Despite postmortem enamel alterations, patches of microwear are still evident on the labial surface and bordering the margin of the interproximal groove.

The labial surface was examined for antemortem instrumental striations indicative of the “stuff-and-cut” behaviors typical of Neandertals and other Pleistocene humans (Lalueza-Fox and Frayer 1997; Lozano et al. 2008, 2017; Estalrrich and Rosas 2013, 2015; Krueger et al. 2017; Willman 2017a). No antemortem instrumental striations are present, but it should be noted that the labial surface is heavily worn, incomplete, and exhibits additional postmortem surface alterations. The incisal enamel surface exhibits a roughened or irregular surface (Fig. 4) that mimics antemortem enamel chipping (Bonfiglioli et al. 2004; Scott and Winn 2011) that is commonly documented on archaic human anterior teeth (Lalueza-Fox and Frayer 1997; Lozano et al. 2008; Belcastro et al. 2018). When viewed macroscopically, the roughened edge of the labio-occlusal margin of the Ciemna 1 incisor superficially mimics a “nibbling effect” produced by an accumulation of many small enamel chips (Estalrrich and Rosas 2013, 2015). However, ESEM imaging shows postmortem erosion of the occlusal enamel rim and postmortem enamel loss along the mesial and distal edges of the occlusal surfaces that border the extensive longitudinal fractures bisecting the tooth (Fig. 6c, e). Thus, if antemortem enamel chipping was present, it can no longer be definitively identified on Ciemna 1.

A final taphonomic consideration concerns the presence of only an isolated and fragmentary human tooth. Antemortem fracture and dislocation of Ciemna 1 from trauma, incising food, non-masticatory behaviors, or other factors (Lukacs 2007) are unlikely. If traumatic, Ciemna 1 would be classified as a “complicated crown-root fracture” since there is enamel, dentin, and cementum involvement with pulp exposure (Glendor et al. 2007). However, such fractures generally occur at or near the cementoenamel junction and leave the root largely intact (Glendor et al. 2007; Lukacs 2007). The Ciemna 1 tooth is not only longitudinally split, but a substantial portion of the root is still intact. Such fracture morphology is difficult to reconcile with a traumatic episode. Furthermore, enamel cracking and fracture is generally restricted to the enamel crown, terminates at the cementoenamel junction, or only occasionally penetrates the underlying dentin (Viciano et al. 2012, 2015; Bernardini et al. 2014; Belcastro et al. 2018). Thus, a scenario in which the Ciemna 1 tooth was lost through antemortem trauma and discarded in the cave is unlikely.

Isolated teeth are consistent with extreme cranial bone modification by carnivores (stage V: Sala et al. 2014), and the presence of a single skeletal element of low dietary-utility is suggestive of late-stage carnivore scavenging whereby the body is completely disarticulated, damaged, and scattered over a large area (Haglund et al. 1989). Carnivore modification is also consistent with the postmortem crown and root fractures of Ciemna 1. However, without clear signs of digestive corrosion (e.g., Maureille et al. 2017) or tooth marks and other indicators (e.g., Sala et al. 2014; Sala and Arsuaga 2018), carnivore modification of Ciemna 1 cannot be confirmed.

The mosaic of well-preserved antemortem features (e.g., the interproximal groove, microwear, and dentin exposure) and postmortem features (cracking/splitting of the root, dentin cracks, modest rounding of fractured root edges, and abrasion/erosion of enamel) indicate a complex taphonomic history for the Ciemna 1 tooth. However, factors contributing to the ultimate deposition of the tooth are difficult to reconstruct.

Discussion and Conclusions

The paleobiological assessment of the Ciemna 1 incisor is most interesting with respect to the well-developed interproximal groove on the cervical margin. Interproximal grooves are not exceptional among Neandertals (e.g., Siffre 1911; Martin 1923; de Lumley 1973; Frayer and Russell 1987; Bermúdez de Castro et al. 1997; Bouchneb and Maureille 2004; Lozano et al. 2013; Smith et al. 2006; Lalueza-Fox et al. 1993; Ungar et al. 2001; Schmitz et al. 2002; Estalrrich et al. 2017; Frayer et al. 2017), but it does confirm that the incisor belongs to a Middle Paleolithic human. Furthermore, an interproximal groove is also noted on a second maxillary molar (S5000) from Stajnia Cave (Urbanowski et al. 2010), making the Ciemna 1 tooth the second Neandertal from Poland with this wear feature, but the first to exhibit an interproximal groove on an anterior tooth.

The type of material or artifact, and the reason for engaging in behavior(s), that produce interproximal grooves has been extensively discussed in the literature (Eckhardt and Piermarini 1988; Formicola 1988, 1991; Lukacs and Pastor 1988; Puech and Cianfarani 1988; Turner 1988; Brown and Molnar 1990; Frayer 1991; Bermúdez de Castro et al. 1997; Ungar et al. 2001; Hlusko 2003; Bouchneb and Maureille 2004; Lozano et al. 2013; Estalrrich et al. 2017). While working sinew, cordage, plant fibers, and similar materials between adjacent teeth may create some interproximal grooves (e.g., Schulz 1977; Lukacs and Pastor 1988; Brown and Molnar 1990), the majority of researchers agree that abrasion from a rigid or semi-rigid probe, or “toothpick”, is responsible for the formation of most documented interproximal grooves (Formicola 1988; Frayer, 1991; Bermúdez de Castro et al. 1997; Ungar et al. 2001; Hlusko 2003; Bouchneb and Maureille 2004; Estalrrich et al. 2017). Furthermore, grooves experimentally created with calf sinew, and abrasive particles produce very fine microstriations and lack a gross morphology characteristic of the grooves produced by abrasion with bone, antler, wood (Bouchneb and Maureille 2004), or grass stalks (Hlusko 2003, Estalrrich et al. 2017). Grooves on the interproximal surfaces around the cervical region of herbivore anterior teeth have been documented as well (von Koenigswald 1972; Young and Marty 1986; Spinage, cited in Frayer and Russell 1987; Puech and Cianfarani 1988; Franz-Odendaal 2002). However, the morphology of these grooves tends to be deeper and/or more “V”-shaped in cross section (see images in Young and Marty 1986; Puech and Cianfarani 1988; Franz-Odendaal 2002) than those of hominins. The microscopic characteristics of herbivore grooves do not align with those observed in hominins either (Puech and Cianfarani 1988). Thus, we find it most likely that a rigid, or semi-rigid, probe with a non-uniform tip was used for habitual “toothpicking” of the interproximal surface of the Ciemna 1 tooth.

Estalrrich et al. (2017) found that interproximal groove widths tended to be higher in the anterior dentition of the Hortus individuals compared with those from El Sidrón, the former Neandertals also exhibiting greater average occlusal wear than the latter. Experimental replication of toothpick grooves highlights how the repetition of abrasive toothpicking actions over time contribute to the ultimate morphology and dimensions of each groove (Hlusko 2003; Bouchneb and Maureille 2004; Estalrrich et al. 2017). Thus, the wide groove on the Ciemna 1 incisor corresponds well with the relatively well-worn occlusal surface of an older adult Neandertal—such as those from Hortus (Estalrrich et al. 2017). The length of the Ciemna 1 interproximal groove is quite long despite postmortem truncation. However, it is within the range of those documented on other anterior—albeit mostly maxillary—teeth from El Sidrón and Hortus (Estalrrich et al. 2017). While no oral paleopathology was documented, the Ciemna 1 incisor is too fragmentary to decisively conclude that oral pathology was absent. Thus, it is unclear if the toothpick groove was related to palliative/therapeutic behaviors (Lozano et al. 2013; Willman 2016, 2017b; Frayer et al. 2017), or hygienic ones (Ungar et al. 2001; Estalrrich et al. 2017).

A final consideration of the Ciemna 1 incisor concerns the paucity of Neandertal fossils in Central Europe (other than the large Krapina sample) despite a rich Middle Paleolithic archeological record in the region. Ciemna Cave, like many other Middle Paleolithic cave sites, is represented by palimpsests of carnivore and human activities from the alternating occupation of the cave (Enloe et al. 2000; Costamagno et al. 2005; Miracle 2005; Wojtal 2007; Enloe 2012; Valde-Nowak et al. 2016b; Sanz et al. 2017; Sanchis et al. 2019). The presence of only an isolated tooth, of low dietary-utility, is suggestive of extreme bone modification (Sala et al. 2014) and late-stage carnivore scavenging (Haglund et al. 1989). Canids can produce extensive damage to a human mandible (Haglund 1996), as can hyenas (Horwitz and Smith 1988), and a high degree of bone breakage and consumption is typical to hyenas and wolves (Sala and Arsuaga 2018). However, there is no unequivocal evidence of carnivore modification of Ciemna 1. Nevertheless, the state of the Ciemna 1 fossil is characteristic of other Neandertal remains from the Middle Paleolithic of Central Europe in terms of its fragmentary and isolated nature, which raises questions regarding the roles of humans in site formation processes in the region.

Thus, the isolated tooth from Ciemna Cave can now be added to the record of similarly fragmentary Central European Neandertal remains from Strajnia, Subalyuk, Šipka, Švédův Stůl, Kůlna Cave, Bordul Mare, Gànovce, and Šal’a. The tooth occurs, as with many of the other sites from the upland and Carpathian zones of Central Europe, in the context of a shelter that was principally a carnivore (especially ursid) den with only occasional human occupation.



Dr. Erik Trinkaus provided extensive comments and comparative metrics for analyses. Initial ESEM analyses were carried out with the assistance of Dr. Katarzyna Berent at the AGH University of Science and Technology, Academic Center for Materials and Nanotechnology, Kraków, Poland. Final ESEM analyses conducted at the Scientific and Technical Resources Service of the University Rovira i Virgili, Tarragona, Spain. The associate editor and three anonymous reviewers provided feedback that greatly improved this manuscript. To all we are grateful.

Funding Information

This work was supported by a Marie Skłodowska-Curie Actions (H2020-MSCA-IF-2016 No. 749188); AGAUR (Ref. 2018SGR1040 and 2018PFR-URV-B2-91) and the Polish National Science Centre (No. UMO-2014/15/B/HS3/02219 – “The Last Neanderthals in the Ciemna Cave”).

Compliance with Ethical Standards

Conflict of Interest

The authors declare that they have no conflict of interest.

Supplementary material

41982_2019_26_MOESM1_ESM.docx (3.1 mb)
ESM 1 (DOCX 3203 kb)


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Copyright information

© Springer Nature Switzerland AG 2019
corrected publication 2019

Authors and Affiliations

  • John C. Willman
    • 1
    • 2
    Email author
  • Bolesław Ginter
    • 3
  • Raquel Hernando
    • 1
    • 2
  • Marina Lozano
    • 1
    • 2
  • Krzysztof Sobczyk
    • 3
  • Damian Stefański
    • 4
  • Anita Szczepanek
    • 5
  • Krzysztof Wertz
    • 6
  • Piotr Wojtal
    • 6
  • Mirosław Zając
    • 4
  • Katarzyna Zarzecka-Szubińska
    • 7
  • Paweł Valde-Nowak
    • 3
  1. 1.IPHES, Institut Català de Paleoecologia Humana i Evolució SocialTarragonaSpain
  2. 2.Àrea de PrehistòriaUniversitat Rovira i Virgili (URV)TarragonaSpain
  3. 3.Institute of ArcheologyJagiellonian UniversityKrakówPoland
  4. 4.Archeological Museum of KrakówKrakówPoland
  5. 5.Institute of Archaeology and EthnologyPolish Academy of SciencesKrakówPoland
  6. 6.Institute of Systematics and Evolution of AnimalsPolish Academy of SciencesKrakówPoland
  7. 7.Department of PaleozoologyWrocław UniversityWrocławPoland

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