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... Goldfuss, 1820 Cervinae Goldfuss, 1820 Megacerini Viret, 1961Megaloceros Brokes, 1828Including: Megaceros Owen, 1844Praedama Portis, 1920;Dolichodoryceros kahlke, 1956. Figure 2 The European giant deer of the Pleistocene (after Van der Made, submitted). 25 50 75 100 125 150 175 200 225 25 50 75 100 125 150 175 200 225 20 cm 1b 2a 1c 1a 2b 3a 3b 279 THE LATEST EARLY PLEISTOCENE GIANT DEER MEGALOCEROS NOVOCARTHAGINIENSIS N. SP. ...
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... antler fragment (Figure 3-2) has the basal part of the brow tine flattened and clearly dipping towards the median plane. This is one of the specimens which has been described before as a knapping tool (Carbonell et al., 1981;Gibert Clols, 1984, 1989). ...
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... a position behind this groove, a crest is directed towards the buccal side of the tooth. If seen in anterior or pos- terior view (Figures 12-1b), the lingual wall is not very upright (feature 3); and it is not completely convex, but convex at the base and concave a little further from the base. This is a common feature in Dama. ...
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... P 2 (Figures 11-1, 12-4, 12-8) is similar to the P 3-4 , but differs in the paraprestyle being not well developed (but see Figure 12-2), in the para ectostyle being much more dominant on the buccal wall, in being more elon- gate and in having a still more marked groove on the lingual wall. ...
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... D 2 ( Figure 12-3) is again similar to the P 2 , but differs in being narrower and in having a much better separa- tion in two lingual lobes. ...
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... M 3 ( Figure 20) has a moderate relieve on the lin- gual side. The metapost and entoprecristids overlap (as is common in Dama), but are not connected by a me- taendocristid (which is common in Cervus). ...
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... size of the M 3 (as indicated by the width of the first lobe -DTa) is compared to that of other Dama-like deer in Figure 20. The minimum, average and maximum val- ues of the small sample from Cueva Victoria are larger than in any of the Early Pleistocene samples, including the large samples of Untermassfeld and Vallonnet, but are comparable to the Middle Pleistocene samples. ...
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... M 1 and M 2 ( Figure 21-5) have morphologies that are similar to that of the first two lobes of the third molar. ...
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... P 4 ( Figure 21-4, 21-7) has a deep furrow at the buccal side separating the two lobes. The second lobe tends to be narrower than the first one. ...
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... a buccal height of 15.7 mm. The entoconid of this tooth is not connected (at the occlusal surface) to the hypoprecristid (as in Dama), but in a more worn spec- imen ( Figure 21-7) these structures are connected (as in Cervus) (feature 6 of Lister, 1996). The wear stage has to be taken into account in the evaluation of this feature. ...
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... wear stage has to be taken into account in the evaluation of this feature. There is a well developed metaprecristid clos- ing the anterior fossid in two of the specimens (they are "molarized"), but it is absent in the third specimen ( Figure 21-7). The molarization of the P 4 is variable in probably all of the Dama-like species, but the frequency of the molarized morphology seems to increase in the younger samples or species (Table 8). ...
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... P 3 ( Figure 21-9) has a moderately developed metaconid, that is placed more posteriorly than the pro- toconid. It is peculiar, that this cusp is not well connected to the protoconid. ...
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... P 2 ( Figure 21-8) has a relatively low and "inflated" main cusp as in Dama, and unlike in Cervus, where this cusp is higher with flatter lingual and buccal sides. A pro- toprecrista is directed anteriorly, it does not curve much lingually and the parastylid is not demarcated as a dis- tinct structure, nor is there a separate paraconid (feature 2 of Lister, 1996). ...
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... M 1-3 (P3 (Figures 21-1, 21-3, 21-10) have moder- ately developed buccal styles and the lingual columns are not particularly strong (features 1 and 3 of Lister, 1996). ...
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... P 4 ( Figure 21-2) has no groove on the lingual wall and a moderate cingulum where the lingual wall curves postero-buccally. If seen in anterior view, the lingual wall is convex near to the crown base and straight or faint- ly concave higher up. ...
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... P 2-3 ( Figure 21-6) have a deep groove on the lin- gual side, forming two lingual lobes. ...
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... possible phylogenetic relationships are tentative. Dama rhenana (with a maximum of three tines per ant- ler), D. nestii (four tines), D. vallonnetensis, D. clactoni- ana (the tines fuse in a narrow palmation) and D. dama (with a wider palmation) seemed to fit in gradual trends towards increasing antler complexity and a lower bifur- cation of the brow tine and main beam ( Figure 18) and in gradual size changes (Figure 20), which was taken as an indication that they may belong to a single line- age (e.g. Van der Made, 1999). ...
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... deer from Cueva Victoria has the bifurcation of the brow tine and main beam lower than in the earliest species of this group (Figure 18). It is larger than most Early Pleistocene species, save for the sample from Atapuerca TD4-5, and is in the ranges of the Middle Pleistocene species (Figure 20). The long main beam without the beginning of the base of the second tine, is a resemblance to the Early Pleistocene species. ...
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... age indicated by M. novocarthageniensis de- pends on its phylogenetic position and three posibilities have been suggested: 1) If M. novocarthaginiensis would be intermediate be- tween M. savini and M. giganteus (e.g. Van der Made, 2001Made, , 2004), the age of Cueva Victoria (and of Cueva Negra) should be between some 500-600 and 400 ka 2) If M. novocarthaginiensis is a descendant of the species from Libakos and ancestral to M. savini (one of the variants in Figure 2; Van der Made & Tong, 2008; Van der Made, in prep.), the age of Cueva Victoria (and Cueva Negra) should be between some 1.2 Ma (the age of Libakos; Steensma, 1988) and 0.7-0.8 Ma (the appearance of M. savini). ...
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... this scenario, M. giganteus is either another descendant of M. novocarthaginiensis, or evolved as a parallel branch (in eastern Europe or western Asia). (MURCIA, SPAIN) 3) If M. novocarthaginiensis is a descendant of the spe- cies form Libakos and ancestral to M. giganteus (evolv- ing in parallel to the M. savini lineage; the other variant in Figure 2; Van der Made & Tong, 2008; Van der Made, in prep.), the age of Cueva Victoria (and Cueva Negra) would be between some 1.2 and 0.4 Ma (the entry of M. giganteus). ...
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... though it has not been explicitly stated, the fos- sils of Dama have played a role in the age assigned to Cueva Victoria (e.g. Van der Made, 2001, fig. 2). Their large size and the idea that the antler may have been palmate played a role. As discussed, above, the size is indeed more indicative of the Middle Pleistocene, but the sample from Atapuerca TD4+5 also reaches such large sizes and is latest Early Pleistocene in age. The units TD4+5 are below a polarity event, which has been ...
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Citations
... The attribution of Dama-like deer to the genus Dama remains controversial, although some researchers include them in this genus (e.g., Pfeiffer, 1999Pfeiffer, , 2005Van der Made, 2015;Pfeiffer-Deml, 2016Athanassiou, 2022;Van der Made et al., 2023). These species are assigned to other genera, including Cervus (s.l.) and "Pseudodama" Azzaroli, 1992(see Heintz, 1970Azzaroli, 1992;Kahlke, 1997Kahlke, , 2001Petronio, 1998, 2002;Breda and Lister, 2013;Breda et al., 2020;Cherin et al., 2022;Mecozzi et al., 2024). ...
... 5644/3367 is similar to the antler of a deer from Libakos (MNQ19) in Greece, which is assigned to Megaloceros sp. (Van der Made and Tong, 2008;Van der Made, 2015) or Praedama sp. (P. ...
... It was first identified as Megaloceros sp. (Kostopoulos, 1997), later as Eucladoceros giulii Kahlke, 1997(Van der Made, 1998 and as Arvernoceros cf. verestchagini (Croitor and Kostopoulos, 2004), and then distinguished as a new species, Rucervus gigans (Croitor, 2018b). ...
... Before these finds, the giant deer from Libakos (MNQ19, ca. 1.3 Ma) from Greece was considered the oldest representative of the genus Megaloceros (s.l.) in Europe (Van der Made and Tong, 2008;Van der Made, 2015. According to J. van der Made, the deer from Libakos might be at the base of the divergence of two lineages: one led to M. savini (=Praedama savini) and the other to M. giganteus (Van der Made, 2015, text-fig. ...
... Tempo and mode of the early dispersal of this taxon in Europe are still unknown and would require the taxonomical revision of the fallow deer remains from late Early to early Middle Pleistocene localities in western Europe. Examples are the remains from the Sierra de Atapuerca (Gran Dolina, Penal, Sima del Elefante, Trinchera Galerìa, van der Made 1998Made , 1999Made , 2001Made , 2013Made , 2015van der Made et al. 2003van der Made et al. , 2017, the complex moyen II at the Caune de l'Arago (France; Magniez et al. 2013) or the Italian remains of Cava Redicicoli , Pagliare di Sassa (Palombo et al. 2001) and Slivia (Ambrosetti et al. 1979). ...
The taxonomy of Quaternary medium-sized deer from Europe rests mainly on antler morphology, while adequate dental and postcranial diagnostic features are lacking. When complete antlers are not available, the taxonomic identifications are often attempted on chronological ground. A considerable number of mostly unpublished craniodental and postcranial remains of fallow deer from selected Italian sites from the late Early Pleistocene to the late Middle Pleistocene is here presented and discussed. The aim of this work is to test the validity of the diagnostic characters proposed in literature and to explore the variability of the fallow deer taxa. In addition, the analysis of the two reference samples from Riano and Ponte Molle allows to refine the features of Dama clactoniana. Finally, biometric comparison has been performed in order to investigate possible oscillations across time and/or differences among taxa.
... Van der Made and Mazo (2014), Van der Made (2015), and Van der Made et al. (2017) considered Dama vallonnetensis and Dama roberti to be on a lineage that reduced the tines and that the palmate species evolved from Dama nestii or Dama nestii farnetensis. Van der Made (2015) and Van der Made et al. (2016 recognized the palmate species Dama peloponesiaca and D. aff. ...
... Van der Made and Mazo (2014), Van der Made (2015), and Van der Made et al. (2017) considered Dama vallonnetensis and Dama roberti to be on a lineage that reduced the tines and that the palmate species evolved from Dama nestii or Dama nestii farnetensis. Van der Made (2015) and Van der Made et al. (2016 recognized the palmate species Dama peloponesiaca and D. aff. peloponesicaca, not recognized by the other authors ( Fig. 1). ...
... We can observe that there is a tendency for the successive species to have the brow tine in progressively lower positions. This is a common evolutionary tendency in the Cervinae and has been documented for Dama (Van der Made 1999; Van der Made et al. 2016Made et al. , 2017, Megaloceros, and Sinomegaceros (Van der Made & Tong 2008;Van der Made 2015 and is also known from Megaceroides (Praemegaceros), while in Eucladoceros there is no clear tendency, but the different species differ in this feature (Van der Made & Dimitrijevic 2015). Though there are fewer data on the position of the middle tine (B) than on the brow tine (A), in successive samples or species, it also tends to originate in a progressively lower position. ...
We describe fossils of a new species of fallow deer, Dama celiae. It is the end member of the lineage Dama farnetensis–D. vallonnetensis–D. roberti–D. celiae, which reduced the number of points of the antler from four to two, while the parallel lineage leading to the living fallow deer evolved more complex and palmate antlers. The fossils are from localities Pedro Jaro I and Orcasitas in the + 25–30-m terrace of the Manzanares river, which is correlated to MIS9 (337–300 ka) and which also yielded fossils of Megaloceros matritensis, a recently named species, end member of a lineage that survived longer than previously believed. A younger terrace of the Manzanares yielded remains of Haploidoceros, a rare deer known from two older localities in southern France and one younger locality in Spain. So many rare deer species in this valley indicates either endemism and a very special environment or that the record of fossil deer is much less known than generally assumed. Until recently, the European Middle Pleistocene record of deer had only one middle-sized species at a time. Now, it appears that there were up to three contemporaneous species of the size of a fallow deer. Acheulean lithic assemblages have been documented from the same sites as Dama celiae. This species was contemporaneous to Neanderthals with Acheulean culture. Cut marks suggest that it was consumed by them and probably was hunted
... In the case of the artiodactyls, the nomenclature of the dentition follows Bärmann & Rössner (2011) and Made (1996); the latter is also applied to the nomenclature and measurement of suids. For cervids and bovids the measures were taken according to specifications by Made & Tong (2008) and Made (1989Made ( , 2012, except for Ld for the distal phalanx when pertinent (Von Den Driesch 1976). Apart from the specified nomenclature in Table 1, equid phalanges are as 2FIII, with the Arabic number indicating the number of the phalanx and the roman numeral indicating the digit (also for metapodials). ...
... Graphic: Bivariate analysis with both posterior transverse diameter and antero-posterior diameter on the base of the crown of the left upper third molar. Data extracted fromMade (1999Made ( , 2012,Breda & Lister (2013) andMade et al. (2017). ...
The Lower Pleistocene site of Quibas, in Sierra de Quibas (Murcia, Spain) was discovered in 1994 and has since then provided abundant material of typical Epivillafranchian taxa. This biochron belongs to the Early-Middle Pleistocene transition (1.2 – 0.78 Ma), characterised by a change in orbital cyclicity from a 41 kyr cycle to 100 kyr that intensified the climate and culminated in the most important faunal turnover of the Pleistocene regarding large mammals. The Group of Palaeoanthropology of the National Museum of Natural Sciences (CSIC, Spain) and the Institut Català de Paleoecologia Humana i Evolució Social (IPHES-CERCA, Spain) carried out four field seasons from 2015 to 2018. Here we present the large herbivorous mammals recovered from the field, including the first citation of two taxa new to the locality: Stephanorhinus cf. etruscus and Bison cf. voigtstedtensis. We also provide the first description of previously mentioned taxa: Dama cf. vallonnetensis and Sus sp. Together with the remaining herbivores, the faunal community shows a strong European affinity with some regionalism. Compared with other Iberian localities, the site of Quibas stands out for the lack of hominin fossils or any evidence supporting their presence in the area, a peculiar scenario given that the Early-Middle Pleistocene transition broadly speaking sees the arrival of humans into Europe.
... Its classification in the separate genus Praedama is based on differences in antler morphology. Dolichodoryceros sues-senbornensis Kahlke, 1956, is a junior synonym (Croitor 2014;van der Made 2015). The recently described species Megaloceros novocarthaginiensis van der Made, 2015, has also Praedama morphology (van der Made 2015; Croitor 2018b). ...
The paleontological research during the last 160 years in Greece has recovered continental deer fossils from at least 100 localities, which span geochronologically from the late Miocene to the end of the Pleistocene, while scanty dental remains date from the middle Miocene. The following species are documented: Procapreolus pentelici (late Miocene), Procapreolus cusanus, Croizetoceros ramosus, Metacervocerus rhenanus, Eucladoceros cf. ctenoides, Rucervus gigans, Praedama aff. savini, Dama vallonnetensis (late Pliocene and Early Pleistocene), Praemegaceros pliotarantoides, Praemegaceros verticornis, Alces latifrons, Megaloceros giganteus, Cervus elaphus, Dama dama, Capreolus capreolus, “Cervus” peloponnesiacus (Middle and Late Pleistocene), while several other samples are incomplete and not identifiable to the species level. Despite their comparatively scarce representation in the fossil faunas, the cervids became more common in time periods with cooler climate, like during the Late Pleistocene.
... Cervus megaceros Hart, 1825 is based on the specimen from Rathcannon (Ireland) exposed in the Royal Dublin Society [40]. Cervus euryceros irlandicus Fischer, 1834 [43] is based on the specimen described by Hibbert [42] from the Isle of Man that is designated here as the lectotype of Megaloceros giganteus irlandicus (Fischer, 1834 [27,55,56], but is regarded by Vislobokova [26] as a junior synonym of M. giganteus giganteus. Hibbert [57] proposed to use the species name Cervus euryceros taken from Aldrovandi's interpretations of antique texts [58]. ...
... Cervus euryceros irlandicus Fischer, 1834 [43] is based on the specimen described by Hibbert [42] from the Isle of Man that is designated here as the lectotype of Megaloceros giganteus irlandicus (Fischer, 1834). The subspecies name M. giganteus irlandicus has been applied by van der Made [27,55,56], but is regarded by Vislobokova [26] as a junior synonym of M. giganteus giganteus. Hibbert [57] proposed to use the species name Cervus euryceros taken from Aldrovandi's interpretations of antique texts [58]. ...
... The giant deer from Sapozhok is characterized by the extremely short upper premolars (the premolar to molar series length ratio is 62.8%) and lower premolars (53.4%), thus achieving the most advanced condition of those characters among the giant deer forms ( Figure 5A). The data on giant deer from Ireland are adapted from Croitor et al. [29]; the data on giant deer from Rhine Valley are adapted from van der Made [27]; the data on giant deer metacarpals from Zhana-Aul are adapted from Kozhamkulova [67]; the data on Praedama giulii are adapted from Kahlke [68]; the data on P. novocarthaginiensis, P. matritensis and "Elaphurus" eleonorae are adapted from van der Made [56,69] and Vislobokova [70] The specimen from Sapozhok is characterized by the long robust type of metacarpals that are grouped on the scatter diagram together with the long metacarpals from Western Europe ( Figure 5B). Therefore, one can assume that the long robust metacarpals from Ireland should be ascribed to M. giganteus giganteus. ...
The article presents a preliminary morphological description of the holotype of Megaloceros giganteus (Blumenbach, 1799) that serves for the description of the species. The article proposes a taxonomical and morphological revision of the nominotypical subspecies M. giganteus giganteus and morphological comparison with other subspecies of M. giganteus. The cluster analysis of diagnostic craniodental and antler characters revealed the systematic position and phylogenetic relationships of M. giganteus with other cervid groups. The genus Praedama is regarded as a closely related phylogenetic branch that linked to the direct cursorial forerunner of Megaloceros that evolved in the middle latitudes of Western Siberia and northern Kazakhstan. The genus Dama has a distant relationship with Megaloceros and represents an earlier phylogenetic branch that evolved in the Ponto-Mediterranean area. The article discusses the secondary adaptations of M. giganteus forms to forest and woodland habitats in Europe and general paleobiogeographic features of the Megaloceros lineage.
... This study presented a phylogeny of Megaloceros with five species, including M. giganteus, M. savini and three species which were not named. Since this time, more data became available on Megaloceros and related forms and one of these species has been named (Van der Made, 2015). Moreover, the stratigraphy of the terraces of the Manzanares and nearby Jarama valleys became much better known and were dated (e.g. ...
... Van der Made and Tong (2008) noted that there were several species, recorded from Libakos, Cueva Victoria and the Madrid area, which resemble M. savini, but differing in some features. The material from Libakos has been described by Steensma (1988), but was not assigned to a species, the material from Cueva Victoria was recently named M. novocarthaginiensis (Van der Made, 2015) and the material from Madrid is described in this paper. Some recognize two genera Megaloceros for M. giganteus with antlers with a distal palmation and Praedama (= Dolichodoryceros) for M. savini with branching antlers (e.g. ...
... A phylogenetic model of the evolution of Megaloceros was proposed by Van der Made and Tong (2008). Since that time, one of the species was named (Van der Made, 2015) and further research lead to the reevaluation of some features. Titov and Shvyreva (2016) named the species Megaloceros stavropolensis based on an antler from the Georgievsk sand pit, correlated to MN17 (which corresponds to about 2.5-1.8 ...
The Irish elk or Megaloceros giganteus is an emblematic species of the Pleistocene, but its relatives are much less known and were believed to have gone extinct when the Irish elk dispersed into Europe. The species Megaloceros matritensis n. sp. is described here on the basis of material from ten localities and levels in a terrace of the Manzanares river, South of Madrid. It must have been a common species when it lived there some 300–400 ka ago, being contemporary of M. giganteus. This species acquired features, such as enlarged premolars, very thick molar enamel, and a low mandibular condyle, which are masticatory adaptations to an, as yet, unknown diet. The species itself formed part of the diet of people which lived in the area. Megaloceros matritensis fossils are found associated to stone tools of late Acheulean and early Mousterian type. If found in other areas, this species could be indicative for this transitional period. Giant deer might be expected to be good examples of Cope's rule, which holds that species tend to evolve larger body sizes. However, M. matritensis is the last member of a lineage which gradually decreased in size during the Middle Pleistocene. Contrary to what one might expect, size decrease is not un-common in the giant deer.
... According to Rössner & Rummel (2001), the sample from Cetina de Aragón should be included in Pomelomeryx boulangeri (Pomel, 1853). The small sized ruminant of Laerru is probably a new insular taxon and it can be associated to the same forms of Oschiri ( " Amphitragulus " sp.) (Van der Made 2008) and Sardara ( " Amphitragulus boulangeri " ) (Comaschi Caria 1953 ). The poorly preserved remains of Laerru are temporarily reported to Pecora indet. ...
... The " Oschiri fauna " is one of the oldest known endemic insular fauna of the Mediterranean (Bruijn & Rümke 1974; Van der Made 2008 ). In literature , this fauna is reported in two Sardinian localities: Oschiri and Sardara (Comaschi Caria 1953; Bruijn & Rümke 1974; Van der Made 2008). ...
... The " Oschiri fauna " is one of the oldest known endemic insular fauna of the Mediterranean (Bruijn & Rümke 1974; Van der Made 2008 ). In literature , this fauna is reported in two Sardinian localities: Oschiri and Sardara (Comaschi Caria 1953; Bruijn & Rümke 1974; Van der Made 2008). In the early 20 th century, the first vertebrate fossils were found in Oschiri, such as two maxilla fragments of ruminant. ...
A new vertebrate assemblage was discovered in an Early Miocene lacustrine deposit near the village of Laerru (northern Sardinia, Italy). The assemblage is composed by mammals, reptiles and a bird. The mammals are represented by three ruminants (cf. Sardomeryx oschiriensis, Pecora indet. small size and Pecora indet. very small size) and one dormouse (Peridyromys aff. murinus) while reptiles are represented by turtles (Trionychidae?) and crocodiles (cf. Diplocynodon sp.). A bird bone fragment is also reported and referred to Palaeortyx cf. brevipes (Galliformes). The assemblage can be related to the " Oschiri fauna " , one of the oldest endemic insular fauna known in the Medi-terranean. The age of the Laerru vertebrates is early-middle Burdigalian, between 18.8 and 18.3 Ma, corresponding to the mammal unit of the main land MN3. The predominance of ruminants confirms the good capacity of these mammals to colonize insular environments.
... Ecologically opportunistic cervids are most successful in young ecosystems with large amplitude of environmental fluctuations (Geist, 1998). Indeed, the paleontological record and modern fauna give only two examples of successful evolutionary survival of cervids on the African continent: Megaceroides algericus (Lydekker, 1890) and Cervus elaphus barbarus Bennet, 1833(Gentry, 2010. ...
... The first description of the species belongs to Lydekker (1890). He described a maxilla with an upper tooth series comprising P 4 -M 3 of a medium-sized deer from Hammam Mescoutine (Algeria) as Cervus algericus, noting a strongly developed cingulum, and assumed a possible phylogenetic relationship of the new species with the giant deer Megaloceros giganteus. ...
... : distance between M 3 and posterior edge of occipital condyle; M 1 -M 3 : length of upper molar series; P 2 -P 4 : length of upper premolar series; P 2 -pr.: distance between P 2 and prosthion; or-pr., distance between orbit and prosthion; Dor.: horizontal diameter of orbit; oroc.: distance between orbit and posterior edge of occipital condyle. (Lydekker, 1890: figured on p. 602), P 2 being completely destroyed and M 2 and M 3 being damaged; the cast is stored at the Natural History Museum of London (Lydekker, 1890: p. 604), collection number M10647 (Gentry, 2010). The length of the upper molar series M 1 -M 3 amounts to 58.5 mm (measured from the figure). ...
The unusual cranial morphology of the endemic extinct African deer Megaceroides algericus (Lydekker,
1890) is described. Some details of cranial and dental morphology suggest that M. algericus is closely
related to the Eurasian giant deer Megaloceros giganteus (Blumenbach, 1799). The paper presents also a
discussion on paleoecology and functional morphology of M. algericus, as well as its origin, phylogenetic
and taxonomic position. Megaloceros mugarensis (Di Stefano, 1996) from the middle Pleistocene of
Levant is regarded as a probable forerunner of M. algericus.
