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Population dynamics of Equus conversidens(Perissodactyla, Equidae) from the late Pleistocene ofHidalgo (central Mexico): comparison with extant and

fossil equid populationsAlexis Pérez-Pérez, Victor Manuel Bravo-Cuevas, Philippe Fernandez

To cite this version:Alexis Pérez-Pérez, Victor Manuel Bravo-Cuevas, Philippe Fernandez. Population dynamics of Equusconversidens (Perissodactyla, Equidae) from the late Pleistocene of Hidalgo (central Mexico): com-parison with extant and fossil equid populations. Journal of South American Earth Sciences, Elsevier,2020, 106, pp.103100. �10.1016/j.jsames.2020.103100�. �halshs-03024467v2�

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15/12/2020 Population dynamics of Equus conversidens (Perissodactyla, Equidae) from the late Pleistocene of Hidalgo (central Mexico): Comp…

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Journal of South American Earth SciencesAvailable online 14 December 2020, 103100

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Population dynamics of Equus conversidens (Perissodactyla, Equidae) from thelate Pleistocene of Hidalgo (central Mexico): Comparison with extant and fossilequid populations

Alexis Pérez-Pérez , Victor Manuel Bravo-Cuevas , Philippe Fernandez

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https://doi.org/10.1016/j.jsames.2020.103100

HighlightsPaleodemography of Equus conversidens is explored by life-tables and Lewis-Lesliematrices.

Mortality pattern of E. conversidens is a prime-dominated profile.

Hunting pressure by non-human predators is related to the mortality pattern.

Predation and resource availability regulate population dynamics of E.conversidens.

AbstractThe Mexican horse Equus conversidens is a representative member of the Quaternary North American megafauna.Systematics, phylogeny, and stable isotope ecology of this species are known in some detail, although informationregarding population ecology is unexplored. Numerous fossil remains (isolated teeth, mandibles, skulls, and postcranialelements) of this horse have been collected from Pleistocene deposits that outcrop at southeastern Hidalgo, centralMexico. The available dental sample is suitable for recovering some aspects of its population dynamics. Our studycompares life history traits of cohorts of E. conversidens from the state of Hidalgo to extant and Pleistocene species ofequids from North America (United States and Canada), Europe (France), and Africa (Rwanda, Namibia). Life tables andage-structured models allow us to describe demographic parameters and mortality patterns. We highlight that E.conversidens was a typical K-strategists with high probability of surviving to adulthood with asymptotic growth rate(λ = 1.05) indicating a population close to balance. Considering the mammalian associations from different habitats insoutheastern Hidalgo and the flexible diet of E. conversidens, we suggest that ambush (Panthera atrox) and cursorialpredators (Canis dirus) played a major role in the prime-dominated mortality pattern of this species. The presence of awoodland-grassland ecosystem that sustained a high diversity of herbivores and the occurrence of large-sized predatorssuggest that the population dynamics were regulated by resource availability and selective predation, as it occurs inpopulations of the extant plains zebra, E. quagga.

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Keywords

Autoecology; Paleodemography; Equid cohorts; Mortality patterns; Leslie matrices

1. IntroductionLife tables and matrix population models are very efficient tools now widely applied in ecology to better understand thelife history traits of plants and animal species (Fujiwara and Diaz-Lopez, 2017). In both cases, multiple cohorts areparticularly well suited to interpret age structures through time (Fernandez et al., 2017).

Age-specific life tables were first applied in experimental biology to better understand invertebrate mortality patterns (e.g.Pearl and Miner, 1935). These models were then developed further, especially in many studies for wildlife management oflarge mammals (Caughley, 1966) or birds (Botkin and Miller, 1974). At the same time, matrix population models havebeen used to describe the evolution of multiple cohorts (projections in time, stationary versus unbalanced livingpopulations) (Bernardelli, 1941; Lewis, 1942; Leslie 1945, 1948), followed more recently by the very well-known andimpressive work of Caswell in the field of applied mathematical demography (2001).

Nevertheless, life table models have been poorly exploited in paleodemography except in a few studies involving largemammals (Kurtén, 1953; 1954a; 1954b; 1983; Van Valen, 1963; 1964; 1965; Voorhies, 1969; Reher 1970; 1973; 1974;Nimmo 1971; Hulbert, 1982; 1984; Koike and Oitaishi, 1985; 1987; Lyman, 1987; Turnbull and Martill, 1988; McDonald,1996; Fernandez and Legendre, 2003; Mihlbachler, 2003; 2007; O'Sullivan J., 2005; Fernandez et al., 2006; Fernandez2009; Prentiss et al., 2014), dinosaurs (Erickson et al. 2006, 2009; Steinsalz and Orzack, 2011), and small mammals(Korth and Evander, 1986; Bryant, 1991; Mihlbachler, 2012). In a similar way, age-structured Leslie models have receivedlittle interest in paleodemography although they have become a standard in population ecology (Fernandez and Boulbes,2010; Monchot et al., 2012; Rodríguez-Gómez et al., 2014, 2016; Martín-González et al., 2016).

In this study we highlight the remarkable properties of life table demographic parameters (see below i.e. l , s , R , r, Tb,etc.) with Leslie matrix model projections considering an outstanding sample of different horse species from differentecological contexts and mammalian associations. Our E. conversidens sample consists of at least 50 individuals from thelate Pleistocene of the state of Hidalgo. Here we compare it to the fossil cohorts of Equus lambei (Bluefish Caves, Canada)(Burke and Cinq-Mars, 1998), extant Equus quagga (Akagera, Rwanda) (Spinage, 1972), Equus zebra (Khomas Hocland,Namibia) (Joubert, 1974), Equus caballus (Pryor Mountain, Montana, United States) (Garrott and Taylor, 1990), Equushemionus (Southern and Dzungaria Gobi, Mongolia) (Lkhagvasuren et al., 2017), and Equus asinus (Grand Canyon,Arizona, United States) (Ruffner and Carothers, 1982). In addition, key paleobiological contexts in France were selectedfor inter-specific age structures comparisons, such as the hyena den of Fouvent with its accumulation of Equusgermanicus, and the Bau de l’Aubesier with its heavy anthropogenic hunting pressure on Equus mosbachensis(Fernandez and Legendre, 2003; Fernandez et al., 2006). Each of these populations offers an exceptional opportunity toprovide detailed information about recent and fossil equids whose causes of death and demographic parameters areknown.

In order to avoid biases resulting from different methodological approaches, individual ages for all fossil equids wereobtained from polynomial equations whose parameters were estimated by bootstrapping starting from the dental heightintervals of known age individuals (Fernandez and Legendre, 2003). Ternary diagrams and histograms are used toillustrate the age frequencies between equid populations in order to relate them to the two dominant theoretical types ofmortality: attritional vs. catastrophic, which are usually considered key components in describing fossil age structures inthe zooarchaeological literature.

Taken together, all of these aspects allow us to address issues that are central to our understanding of the Pleistocene E.conversidens: which demographic parameters are the most sensitive to the final state of the population from southeasternHidalgo, central Mexico? And which human or non-human predators have affected the assemblage of E. conversidens?

2. Study areaThe studied material comes from seven neighbor localities in southeastern Hidalgo known as Ventoquipa (HGO-9: 20°01′N - 98°20′ W, 2289 m.a.s.l.); Barranca Piedras Negras (HGO-23: 20º03′03″ N - 98º37′23″ W, 2460 m.a.sl.); Barranca delBerrendo (HGO-28: 20º01′20.7″ N - 98º37′37.9″ W, 2458 m.a.s.l.); Barranca San Agustín (HGO-29: 20°00′27.7″ N -98º37′59.9″ W, 2467 m.a.s.l.); Jagüey Viejo (HGO-42: 20°02.311″ N - 98°35.885″ W, 2528 m.a.s.l.); Arroyo las Cajas

x x 0

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(HGO-46: 20º07′31.7″ N - 98º58′07.7″ W, 2140 m.a.s.l.); and El Barrio (HGO-47: 20º07′41″ N - 98º56′02.7″ W,2184 m.a.s.l.). These localities are separated by a maximum distance of 67.55 km (between HGO-9 and HGO-47) and aminimum of 1.75 km (between HGO-28 and HGO-29) (Fig. 1). In this area, there are Quaternary deposits that consist ofclay, silt, and conglomeratic lenses set in a fluvial environment (Bravo-Cuevas, 2002) (Fig. 2). An important sample offossil remains has been recovered from these sedimentary deposits. The record includes rodents, felids, dire wolves,ground sloths, giant armadillos, horses, llamas, pronghorns, deer, bison, mammoths, gomphotheres, and mastodons(Bravo-Cuevas, 2001, 2002; Bravo-Cuevas et al., 2009a, 2009b, 2011, 2012, 2013, 2015, 2016, 2017, 2020). The presenceof material belonging to Bison and Panthera atrox is related to the Rancholabrean North American Land Mammal Age(NALMA) (Bell et al., 2004). The identification of Bison antiquus indicates that the maximum age of the fossil-bearingdeposits is of about 60 ka (Bravo-Cuevas and Vázquez-Cortes, 2019), corresponding to the Wisconsin glaciation in NorthAmerica.

Fig. 1

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Fig. 1. Index map of the study area in southeastern Hidalgo, central Mexico. The capital of the state, Pachuca city (largestar) and the fossil localities are indicated. Localities: HGO-9, Ventoquipa; HGO-23, Barranca Piedras Negras; HGO-28,Barranca del Berrendo; HGO-29, Barranca San Agustín; HGO-42, Jagüey Viejo; HGO-46, Arroyo las Cajas; HGO-47, ElBarrio.

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Fig. 2. (a) Lithostratigraphic correlation of the fossil localities and (b) Generalized lithostratigraphy of the fossil localities.The localities are labeled as in Fig. 1.

3. Material and methods

3.1. Dental fossil sample

The sample referable to E. conversidens considered here consists of 85 dental anatomical elements (Table 1). The fossilmaterial includes two skulls with right and left P2-M3, one partial skull with right P2-M3, one left maxillary fragmentwith P2-M1, one right mandible fragment with p3-m3, one right mandible fragment with p2-m2, two left mandiblefragments with p2-m3, five right mandible fragments with p2-m3, one right mandible fragment with p4-m3, twomandible fragments with right and left p2-m3, and 69 isolated cheek teeth (40 of the upper dentition and 29 of the lower

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dentition). The fossil specimens are cataloged and housed at the Museo de Paleontología, Universidad Autónoma delEstado de Hidalgo (UAHMP).

Table 1. Teeth specimens of Equus conversidens from Hidalgo. Upper/lower case for upper and lower dentitionrespectively (R, right; L, left).

Ventoquipa HGO-09 UAHMP-286 Rp4 58.1 7.43

Barranca Piedras Negras HGO-23 UAHMP-179 RP3 71.6 5.99

UAHMP-409 RM3 45.8 9.05

UAHMP-181 Lp4 53.6 7.94

Barranca del Berrendo HGO-28 UAHMP-941 RP2 38.7 9.42

UAHMP-509 RP4 57.5 8.06

UAHMP-905 RP3 67.0 6.74

UAHMP-314 RP4 63.7 6.83

UAHMP-319 LM1 43.3 9.97

UAHMP-594 LM1 13.1 23.85

UAHMP-901 LM1 34.7 12.47

UAHMP-909 RM1 34.7 12.47

UAHMP-316 LM2 66.6 6.69

UAHMP-942 LM2 65.0 6.92

UAHMP-356 LM3 61.3 7.36

UAHMP-317 RM3 20.8 18.67

UAHMP-939 RM3 60.8 7.40

UAHMP-943 RM3 67.9 6.75

UAHMP-215 Lp3 65.7 7.00

UAHMP-353 Lp4 64.7 6.75

UAHMP-4145 Rp4 50.2 8.40

UAHMP-945 Lm1 44.0 8.26

UAHMP-1916 Lp4 66.7 6.51

UAHMP-403 Rm1 28.9 12.52

UAHMP-508 Rm1 78.7 4.49

UAHMP-950 Rm1 19.4 17.51

UAHMP-1915 Rm1 54.3 6.96

UAHMP-4130 Rm1 20.0 17.14

UAHMP-944 Lm2 74.8 5.97

UAHMP-326 Rm3 38.7 11.16

UAHMP-390 Rm3 65.3 6.63

Barranca San Agustín HGO-29 UAHMP-417 LP4 33.9 13.40

UAHMP-415 LM1 47.4 9.06

Locality Locality ID Catalogue No. Dental element Crown height (mm) Estimated age (years)

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UAHMP-491 LM1 59.6 7.12

UAHMP-510 RM3 27.2 14.91

UAHMP-492 Lp2 53.8 5.33

UAHMP-512 Lp2 49.6 6.27

UAHMP-418 Lp4 65.3 6.68

UAHMP-932 Lm1 59.7 6.52

UAHMP-504 Lm1 49.6 7.46

UAHMP-387 Rm1 65.8 6.04

UAHMP-4145 Lm2 56.0 8.36

UAHMP-495 Rm2 41.3 10.62

UAHMP-416 Lm3 64.8 6.70

UAHMP-511 Lp3 Unworn 2.67

UAHMP-4170 Rm1 Unworn 0.92

Barranca Jagüey Viejo HGO-42 UAHMP-935 LP2 48.6 8.02

UAHMP-902 RM1 37.3 11.63

UAHMP-936 Lp4 28.3 15.47

UAHMP-949 Rm1 71.2 5.53

UAHMP903 Rm2 36.2 11.90

UAHMP-937 Rm2 59.9 7.94

Las Cajas HGO-46 UAHMP-1913 LP2 54.9 7.07

UAHMP-1911 LM2 51.9 8.95

UAHMP-1912 LM2 53.1 8.72

UAHMP-507 Lp4 63.5 6.88

El Barrio HGO-47 UAHMP-2706 RP2 37.3 9.65

UAHMP-1125 RP3 50.1 9.13

UAHMP-4014 RP3 44.8 10.03

UAHMP-900 LP4 63.6 6.84

UAHMP-1116 LP4 26.1 17.09

UAHMP-2709 LP4 57.9 7.61

UAHMP-2711 LP4 66.2 6.49

UAHMP-1123 RP4 39.7 11.39

UAHMP-1112 LM1 56.9 7.48

UAHMP-4017 LM1 43.7 9.87

UAHMP-1123 RM1 37.5 11.55

UAHMP-2712 RM1 12.8 24.05

UAHMP-4122 RM1 29.5 14.47

UAHMP-4123 RM1 46.8 9.19

UAHMP-4162 RM1 48.4 8.86

Locality Locality ID Catalogue No. Dental element Crown height (mm) Estimated age (years)Download




UAHMP-A RM1 24.3 16.93

UAHMP-1910 RM2 52.9 8.77

UAHMP-4019 RM2 53.7 8.63

UAHMP-2710 Lp3 18.6 18.26

UAHMP-4018 Lp4 30.9 14.10

UAHMP-2704 Rp4 60.6 7.18

UAHMP-B Rp4 49.7 8.95

UAHMP-2704 Lm1 72.6 5.37

UAHMP-4120 Lm1 50.8 7.32

UAHMP-957 Rm1 57.3 6.71

UAHMP-1109 Rm1 34.9 10.37

UAHMP-2708 Lm3 25.3 15.87

UAHMP-4016 Lm3 71.3 5.48

UAHMP-4215 Lm3 55.5 8.10

3.2. The fossil cohorts of E. conversidens from Hidalgo

An ancient population or paleopopulation consists of different cohorts through time and retains the basic traits of anextant population (e.g., genetic flow, age structure, and interbreeding) (MacFadden, 2008; Fernandez et al., 2017). Thetime interval of a deposit should be considered to characterize a paleopopulation (Hunter, 1998). The mammalianassociation from Hidalgo represents a within-habitat assemblage set in a fluvial environment, time-averaging estimatedlimits range from decades to 100 kyrs (Behrensmeyer et al., 2000; Bravo-Cuevas et al., 2017). This time interval has beenconsidered as a geological time duration that meets the assumption of multiple deposits corresponding to different cohortgenerations that interacted as an entire population (MacFadden, 2008).

MacFadden (1989) observed that intraspecific variation in dental characters in paleopopulations of fossil horses hascoefficients of variation (V) that range from about 4% to 10%, as it occurs in extant horse populations. In this regard, thecoefficients of variation related to the anteroposterior length (7.55%) and transverse width (5.59%) of the first uppermolars of E. conversidens from the state of Hidalgo are less than 10% (Table 2). Consequently, the fossil collection hasbeen treated as a natural assemblage of horses of the same species that shared the same area (southeastern Hidalgo) atthe same geological time-interval (late Pleistocene).

Table 2. Coefficient of variation (CV) in the anteroposterior length (APL) and transverse width (TW) of M1s of Equusconversidens from the late Pleistocene of southeastern Hidalgo, central Mexico (measurements in mm).

HGO-28 UAHMP-319 22.29 22.6

HGO-28 UAHMP-594 20.89 24.22

HGO-28 UAHMP-901 21.76 24.65

HGO-28 UAHMP-509 24.2 24.22

HGO-29 UAHMP-415 22.86 23.33

HGO-29 UAHMP-491 24.07 23.91

HGO-42 UAHMP-902 24.05 25.57

Locality Locality ID Catalogue No. Dental element Crown height (mm) Estimated age (years)

Locality ID APL TW

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HGO-47 UAHMP-1112 23.35 24.11

HGO-47 UAHMP-4017 22.18 22.98

HGO-47 UAHMP-1123 25.66 25.33

HGO-47 UAHMP-2712 20.35 27.44

HGO-47 UAHMP-4122 21.85 23.55

HGO-47 UAHMP-4123 24.92 25.18

HGO-47 UAHMP-4162 26.31 26.97

HGO-47 UAHMP-1116 21.7 23.87

CV 7.55 5.59

3.3. Estimation of the number of individuals

Teeth are often the best anatomical elements to estimate the number of individuals in fossil assemblages because they arebetter preserved than bones and represent diagnostic material for the identification of species (Lyman, 1994). The sampleof E. conversidens from Hidalgo consists of isolated teeth, mandibles, skulls, and postcranial remains, dental elementsbeing the most abundant (Bravo-Cuevas et al., 2011). For comparative purposes, we used two indices: Minimum Numberof Individuals (MNI) (Badgley, 1986) and the total Number of Identified Specimens (NISP) (Lyman, 2008). Assumingmultiples cohorts in the seven localities from central Mexico, the total MNI was summed between each one of them andcalculated considering the most abundant tooth from one side of the body (left or right). Partial or complete dental seriesset in skulls or mandibles were counted as one individual for NISP.

When fossil deposits are produced or modified by either humans, carnivores or natural processes such as those caused bymicro-organisms, we have to estimate the potential taphonomic biases. Here, we calculated the proportion between leftand right teeth according the method of Brain (1981) in order to evaluate dental preservation and potential disturbancesof the assemblage.

Finally, together the MNI, NISP and the dental preservation of E. conversidens were compared with middle and upperPleistocene Equus populations from France (Table 3).

Table 3. Comparison of the minimum number of individuals (MNI) and the total number of identified specimens (NISP),the frequencies of right and left teeth, and the degree of differential preservation (following Brain, 1981) of the horsepopulation of Equus conversidens from the late Pleistocene of southeastern Hidalgo, central Mexico, and selectedpopulations of Pleistocene Equus from France (data from Fernandez et al., 2006: Table 3, p. 180).

50 85 50 39 100 89.00%

49 353 186 167 372 94.89%

54 464 353 326 706 96.17%

67 396 173 218 436 89.67%

∗

Dp = (LT + RT) *100/Total expected teeth.

3.4. Individual age and age intervals estimation

We took into account isolated teeth and cheek teeth set in the maxilla or the mandible. We used the method of Fernandezand Legendre (2003), which uses crown height to assign each tooth (i.e., P2/p2, M3/m3 …) to a particular age interval.

Locality ID APL TW

MNI NISP Left teeth (LT) Right teeth (RT) Total expected Differential preservation (Dp∗)

Hidalgo

Bau del’Aubesier

Fouvent

Combe-Grenal

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The model is based on a regression analysis of curvilinear type and provides the individual age of teeth and its standarddeviation, mean prediction error whose parameters are estimated by bootstrap, as well as median values of regressioncoefficients.

The crown height (in mm) was measured using a digital caliper (range 0–150 mm, resolution 0.01 m, accuracy 0.003 mm,and was taken from the separation of the roots, straight to the occlusal surface along the mesostyle and the ectostylid inthe upper and lower cheek teeth respectively. We estimate the measurement uncertainty following the methodology ofJCGM (2008), recording a set of 30 measurements of crown height on the same tooth (UAHMP-941). The measurandvalue is ±0.0005.

The median (1), minimum and maximum (2) age (in years) for each tooth were estimated from the following equations(from Fernandez and Legendre, 2003: Table 5, pp. 1583; Fernandez et al., 2006: Table 2, pp. 180):

In these equations, a = slope; a , a , a = intercept; H = crown height, and E = mean prediction error. Estimation of the agedistribution of each tooth in appropriate age intervals is detailed in the step-by-step procedure of Fernandez (2009: Fig. 1,pp. 7). In this study, we also compared the age distributions derived from the MNI on the one hand, and the NISP on theother hand using a Kolmogorov-Smirnov non-parametric test and Spearman's rank-order correlation coefficient (Kleinand Cruz-Uribe, 1984; Vasileiadou et al., 2007) (Table 4).

Table 4. Comparison of age distribution derived from the minimum number of individuals (MNI) and the total number ofidentified specimens (NISP) indices, using a Kolmogorov-Smirnov test (K–S = 0.735, p > 0.10) and Spearman's rankcorrelation coefficient (r = 0.881, p > 0.01).

0–3 1.70 1.75

3–6 7.82 10.39

6–9 26.37 37.74

9–12 6.23 17.68

12–15 3.12 7.07

15–18 3.02 6.23

18–21 0.73 2.14

21–24 1.00 2.00

Number of individuals 50 85

Table 5. Life tables of E. conversidens (age intervals of three years) using the minimum number of individuals (MNI) andthe number of identified specimens (NISP) indices. Demographic parameters described as it is indicated in the method.

0 0.00 0.000 100.000 0.966

3 1.70 3.399 96.601 0.838

6 7.82 15.643 80.958 0.348

9 26.37 52.749 28.210 0.558

(1)

(2)

0 1 2 3

s

Age interval MNI NISP

x nx dx lx sx

MNI

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1)

2)

3)

12 6.23 12.469 15.740 0.604

15 3.12 6.235 9.505 0.364

18 3.02 6.043 3.462 0.578

21 0.73 1.462 2.000 0.000

24 1.00 2.000 0.000 0.000

0 0.00 0.000 100.000 0.979

3 1.75 2.059 97.941 0.875

6 10.39 12.225 85.717 0.482

9 37.74 44.406 41.311 0.497

12 17.68 20.798 20.512 0.595

15 7.07 8.312 12.200 0.399

18 6.23 7.330 4.870 0.483

21 2.14 2.518 2.353 0.000

24 2.00 2.353 0.000 0.000

3.5. Mortality patterns

We established the age structure of the population of E. conversidens with both histograms and ternary diagrams, as theyare the most common tools to compare mortality profiles (Stiner, 1991, 1994; Steele, 2005; Steele and Weaver, 2002;Discamps and Costamagno, 2015). MNI and NISP indices were used in histograms where each bar represents the numberof individuals that died in each age interval (Fig. 3a). To show the mortality patterns of equids proposed by Discamps andCostamagno, we rebuilt the limits of the areas of the ternary diagram. In the present study, three age categories wereretained as with the extant plains zebra E. quagga from the Serengeti (Grange et al., 2004: p. 526):

Juveniles (J) = non-reproductive individuals with an age of 0–3 years (year 3 ending at 36 months).

Prime adults (Pa) = individuals with an age of 3–15 years (year 3 beginning at 37 months), including potentiallyreproductive individuals like young adults and adults.

Old adults (Oa) = individuals with an age greater than 15 years.


x nx dx lx sx

NISP

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(1)

(2)

•

•

•

•

•

•

Fig. 3. (a) Mortality profiles of the population of Equus conversidens from the late Pleistocene of Hidalgo derived fromthe MNI and NISP indexes. (b) Ternary diagram and location of the population studied here and selected populations ofextinct and extant Equus. Horse populations: 1, E. conversidens (NISP); 1′, E. conversidens (MNI); 2, E. lambei; 3, E.mosbachensis; 4, E. germanicus; 5, E. quagga; 6, E. zebra; 7, E. caballus; 8, E. asinus; 9, E. hemionus.

The fraction of Potential Ecological Longevity (PEL) of the population of Equus conversidens from Hidalgo was estimatedassuming a PEL of about 24 years, corresponding to the oldest individuals (UAHMP-594: 23.85 ± 0.38 years andUAHMP-2712: 24.05 ± 0.389), which is comparable to that of the extant E. quagga (Spinage, 1972). The PEL was used forthe graphical demarcation between the zones in the ternary diagram, considering the following areas: Juveniles-Prime-Old (JPO), Juveniles-Old-Prime (JOP), Prime (P), and Old (O) (sensu nomenclature of Discamp and Costamagno, 2015).The fraction of PEL to each age category is as follows: Juveniles = 12.5%, Prime adults = 50%, and Old adults = 37.5%(Discamp and Costamagno, 2015).

Finally, the mortality pattern of E. conversidens from Hidalgo was compared with selected Pleistocene and extantpopulations of Equus.

3.6. Population dynamics: life tables and Leslie matrices

A static or time-specific life table model was constructed based on vertical or cross-sectional analysis (i.e., Caughley, 1977;Dodd and Stanton, 1990). This model is suitable for the analysis of the E. conversidens population because it considersindividuals of all ages belonging to different cohorts and multiple diachronic deposits over time (Fernandez et al., 2017).In this regard, two assumptions must be taken into account:

time-specific life tables are female-dominated models. However, it is widely accepted that male survival does not affectthe population growth rate of most natural mammalian populations (Caswell and Weeks, 1986; Lindström and Kokko,1998). This implies a balanced sex-ratio at birth (1:1) and a fecundity rate that is half the number of offspring producedalive at birth by a female of a given age.

it is assumed that the migration processes did not influence the age structure of the studied population (Caughley,1977).

The life table parameters used here are defined below (see also Fernandez et al., 2017).( ) refers indiscriminately either to “age interval” or “age class”. Nevertheless, strictly speaking the former begins at 0and the latter at 1 (Caswell, 2001). In this study the standardization of one-year age intervals was derived from the svalues following the procedure of Fernandez and Boulbes (2010: p. 801).

(n ) is the number of individuals in each age interval. It is derived either starting from the MNI or from the NISP (seecalculation above).

(d ) refers to the proportion of individuals of the whole cohort(s) dying in each age interval. It can be estimated asfollows:

(l ) is the cumulated survivorship and corresponds to the fraction of a cohort that survives to an age x. The first valuecan be expressed per 1, 100, or 1000 and subsequent values are estimated as:

In a standardized life table with regular pooled age-class intervals where d is not available, the algebraic relation with calculates the proportion:

( ) refers to the age-specific survival rate. The rate of individuals that survived from age x to x +1:

(m ) refers to the fecundity rate. This variable is defined as the average number of descendants per age interval.

x

x

x

x

x

x

x

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•

•

In extinct species, this demographic parameter could be inferred using the fecundity rate of extant species throughallometry and/or phylogenetic relationship (Martín-González et al., 2016). As mentioned previously, the fecundity rate ishalf the number of offspring alive at birth produced by a female in each age class, assuming a balanced sexual ratiobetween males and females of 1:1.

The fecundity of E. conversidens from Hidalgo is derived from the age-dependent birth and survival patterns observed infemales of the extant plains zebra E. quagga (Rubenstein and Nuñez, 2009: table 8.2, p. 206), considering their closephylogenetic relationships (Barrón-Ortiz et al., 2019). To obtain a more accurate approximation in the fecundity rate, weassumed that as in E. quagga, females with an age from 17 to 21 years decreased their fecundity by 6.49% (Grange et al.,2004), whereas females over 21 years show low survivorship expectancy by having commonly a null fecundity (Spinage,1972; Smuts, 1976).

(l m ) refers to the contribution of a given age interval to the population reproduction as the product of l and m .

(R ) refers to the net reproductive rate of a fossil cohort that is calculated as follows:

In this regard, if R < 1 indicates that population is declining whereas if R > 1 indicates an increasing population and R = 1means a perfect stationary population (Monchot et al., 2012; Martín-González et al., 2016).

Since the pioneer articles of Leslie (1945, 1948), the so-called Leslie matrices have become a standard in populationecology with the impressive book of Caswell (2001). A projection matrix reveals whether the age structure of the differentcohorts is related to a stationary population at any particular time. It gives both the picture of the initial fossil sample andthe projection in time of the entire population to characterize its viability of a population. Here we used the simplestmatrix projection model known as Lewis-Leslie:

Matrix A contains the age-specific fecundity rates (m , m …) as in the life tables. The diagonal contains the age-specificsurvival rates (s , s …) that corresponds to the values of s . The vector n (t) is obtained from the l , as the fraction of thetotal number of individuals in one-year intervals, whereas the vector n (t+1) is the new number of individuals in each ageinterval at the next succeeding census of the projected population [An(t) = n(t + 1)] (see details in Fernandez et al., 2017).The population projection of E. conversidens was performed using the software PopTools version 3.0.6 (Hood, 2008).

We first entered the age-specific survival (s ) and the fecundity data (m ) of the horse's population into our pre-breedingmatrix as it is the basic model used for mammals (see Caswell, 2001). The projection returns the asymptotic growth rate(λ), the exponential growth rate (r), and the mean generation time (T ) that corresponds to the mean age of a femalegiving birth in a reproductive event (Leslie, 1966). In this regard, decreasing populations are characterized by having λless than 1 and negative values for r. On the contrary, increasing populations have values greater than 1 for λ and positivevalues for r. In ideal conditions stationary populations have λ = 1 and r = 0 values (Lemos-Espinal et al., 2005; Fernandezet al., 2017).

Mammalian populations often show early fluctuations in the asymptotic growth rate, which is known as transient

dynamics. In this regard, the damping ratio describes how fast the transient dynamics is stabilized after

perturbation around the asymptotic growth rate regardless of population structure (Fernandez et al., 2017). We calculatedthe damping ratio using the software R Studio vr. 1.1.463, package popdemo (RStudio Team, 2015; Stott, 2018).

We also performed a disturbance analysis using the PopTool software (Hood, 2008), to explore the elasticity of theasymptotic growth rate related to changes in survival and fertility rates (see details in Caswell, 1978). The elasticity of theE. conversidens population from Hidalgo was compared and plotted with that of selected Pleistocene and extant Equuspopulations using ternary diagrams (Oli, 2004). We considered here: (1) elasticity of the survival of juveniles [e (Sj)],which is the sum of the elasticity values of individuals from 0 to 3 years; (2) elasticity of adult survival [e (Sa)], which is

x x x x

o

o o o

0 1

0 1 x x

x x

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the sum of the elasticity values of individuals of ≥4 years; and (3) elasticity of fertility [e (F)], which is the sum of allelasticity values corresponding to fertility.

The population dynamics and elasticities of E. conversidens were compared to the following equid populations: E. lambeifrom the Bluefish Caves, Wisconsinian of Yukon, Canada; in France E. germanicus from Fouvent, Oxygene Isotopic Stage3; E. mosbachensis from Bau de l'Aubesier, Oxygene Isotopic Stage 7–6; and the extant populations of the plains zebra E.quagga from Akagera, Rwanda; and the mountain zebra E. zebra from Khomas Hochland, Namibia (Spinage, 1972;Joubert, 1974; Burke and Cinq-Mars, 1998; Fernandez and Legendre, 2003; Fernandez et al., 2006; Fourvel et al., 2014,2015) (Supplementary Data).

4. Results and discussion

4.1. Population size

The estimated number of individuals in the sample of E. conversidens is 50 and 85 from MNI and NISP indicesrespectively. Classical works on population dynamics of ancient populations have shown that sample sizes between 30and 40 individuals are suitable for interpreting important demographic features (see Kurtén, 1953; 1954a; 1954b). Lyman(1987) also demonstrated that about 30 individuals are necessary if 10 age classes are used but this critical assumption isoften overlooked in zooarchaeological literature. In fact, optimal MNI depends on both the standard deviation ofindividual age and mean prediction error for each tooth category that will condition the width of the interval age classes,as well as their number (see details in Fernandez et al., 2017, Fig. 1, p. 480). The dental preservation of E. conversidenscalculated from Brain (1981) is comparable to that of Combe-Grenal and is somewhat lower in comparison to thePleistocene French sites of the Bau de l’Aubesier and Fouvent (Table 3). This suggests limited taphonomical biases fordental material between different deposits of our sample.

The Kolmogorov-Smirnov test indicates non-significant differences between MNI or NISP indices in the age distributionof E. conversidens (K–S = 0.74; p > 0.10) and the Spearman's rank correlation coefficient indicates a positive correlationbetween the estimated age distributions (rs = 0.881, p > 0.01) (Table 4, Fig. 3a). It is not surprising that NISP and MNIboth suggest the same age distribution, because it has been demonstrated that they were not independent of one another(see detailed discussions in Grayson, 1979, 1984; Lyman, 2008).

4.2. Mortality profiles

The mortality profiles of E. conversidens show a right asymmetry and a leptokurtic distribution. These profiles do not fitthe basic attritional (U-shaped) and catastrophic (L-shaped) mortality patterns. The observed patterns are more similarto the stalking-model proposed by Levine (1983: Fig. 4.8, p. 31) (Fig. 3a). Hunting mainly of prime adults with a bell-shaped distribution produces this pattern (Levine, 1983). Even if the dental preservation of our sample is quite good, thiskind of distribution has also often been documented in North American bison kill sites. Herds of animals were killed andtheir mortality pattern is usually attributed to poor preservation of the less ossified bones and teeth of youngsters (e.g.,Munson, 2000; Munson and Garniewicz, 2003; Munson and Marean, 2003).

The prime adults of the age interval 6–9 years represent 52.75% (MNI) and 44.41% (NISP), whereas the juveniles and oldadults of the age intervals 0–3 years and 21–24 years respectively represent each less than 10% of individuals for bothindices (Fig. 3a). The high percentage of prime adult individuals in comparison to individuals of other age classes may bethe result of an accumulation over time where predation played an important role (Levine, 1983). Other horse populationsshowing this pattern have been related to the influence of human and/or non-human predators, including the populationof E. lambei from Bluefish Caves, late Pleistocene of Canada (humans and large-sized predators, such as wolf, bear, andlion) (Burke and Cinq-Mars, 1998); the population of E. mosbachensis (Level I) from Bau de l'Aubesier, UpperPleistocene of southern France (selective hunting by Neanderthals) (Fernandez and Legendre, 2003); and thepaleopopulation of E. germanicus from Fouvent, Upper Pleistocene of eastern France (hyena den) (Fernandez et al.,2006).

The predominance of young adults and mature adults indicates a prime-dominated mortality pattern (Stiner, 1990;Kahlke and Gaudzinski, 2005; Bai et al., 2011). This pattern is consistent with the location of the population of E.conversidens in the ternary diagram, within the Prime adult area (P) (Fig. 3b). The comparison with selected populationsof fossil and extant Equus shows that E. lambei, E. mosbachensis, and E. germanicus also fall within the Prime adult area(P) whereas E. quagga, E. zebra, E. asinus, and E. hemionus are located within the Juveniles-Prime-Old area (JPO). Only

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E. caballus is distributed within the Juveniles-Old-Prime area (JOP) and none of the compared populations fall in the Oldadult area (O) (Fig. 3b).

A prime-dominated mortality pattern has been interpreted to reflect selective hunting strategies by humans (Stiner, 1990,1991, 1994; Driver and Maxwell, 2013); however, there is an ecologically complementary influence by non-humanpredators (Marín et al., 2017). This mortality pattern has been observed in large ungulates, including several Pleistocenehorse populations (e.g., Levine, 1983; Fernandez and Legendre, 2006; Marín et al., 2017).

The studied sample does not show anthropic traces, and direct (e.g., bone remains) or indirect evidence (e.g., stone toolsor bone surface modifications) of humans in southeastern Hidalgo during the late Pleistocene is not reported. However,the mammalian fauna from the late Pleistocene of southeastern Hidalgo includes large-sized predators, such as dire wolfCanis dirus and American lion Panthera atrox (Bravo-Cuevas et al., 2009, 2016). Equus conversidens represented asuitable prey for these predators (Bravo-Cuevas et al., 2016). Given this, the mortality profile of E. conversidens fromHidalgo probably was the result of the influence of these predators, providing high return rates to these carnivores(Stephens and Krebs, 1986).

4.3. Life tables and Leslie matrices

Table 5, Table 6 indicate the vital statics of the population of Equus conversidens from Hidalgo.

Table 6. Standardized life tables of E. conversidens (age intervals of one year) using the minimum number of individuals(MNI) and the number of identified specimens (NISP) indices. Demographic parameters described as it is indicated in themethod.

0 1.000 0.989 0.000 0.000

1 0.989 0.989 0.000 0.000

2 0.977 0.989 0.000 0.000

3 0.966 0.943 0.105 0.101

4 0.911 0.943 0.105 0.096

5 0.859 0.943 0.105 0.090

6 0.810 0.704 0.340 0.275

7 0.570 0.704 0.340 0.194

8 0.401 0.704 0.340 0.136

9 0.282 0.823 0.340 0.096

10 0.232 0.823 0.340 0.079

11 0.191 0.823 0.460 0.088

12 0.157 0.845 0.460 0.072

13 0.133 0.845 0.460 0.061

14 0.112 0.845 0.460 0.052

15 0.095 0.714 0.460 0.044

16 0.068 0.714 0.460 0.031

17 0.048 0.714 0.430 0.021

18 0.035 0.833 0.430 0.015

19 0.029 0.833 0.430 0.012

20 0.024 0.833 0.430 0.010

x lx sx mx l mx x

MNI

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21 0.020 0.833 0.430 0.009

22 0.017 0.833 0.000 0.000

23 0.014 0.833 0.000 0.000

24 0.012 0.833 0.000 0.000

R = 1.48

0 1.000 0.993 0.000 0.000

1 0.993 0.993 0.000 0.000

2 0.986 0.993 0.000 0.000

3 0.979 0.957 0.105 0.103

4 0.937 0.957 0.105 0.098

5 0.896 0.957 0.105 0.094

6 0.857 0.784 0.340 0.291

7 0.672 0.784 0.340 0.228

8 0.527 0.784 0.340 0.179

9 0.413 0.792 0.340 0.140

10 0.327 0.792 0.340 0.111

11 0.259 0.792 0.460 0.119

12 0.205 0.841 0.460 0.094

13 0.173 0.841 0.460 0.079

14 0.145 0.841 0.460 0.067

15 0.122 0.736 0.460 0.056

16 0.090 0.736 0.460 0.041

17 0.066 0.736 0.430 0.028

18 0.049 0.785 0.430 0.021

19 0.038 0.785 0.430 0.016

20 0.030 0.785 0.430 0.013

21 0.024 0.785 0.430 0.010

22 0.018 0.785 0.000 0.000

23 0.014 0.785 0.000 0.000

24 0.011 0.785 0.000 0.000

R = 1.79

The net reproductive rate (Ro) was 1.48 and 1.79 using MNI and NISP indices respectively (Table 6), suggesting anincreasing population. The observed Ro in all selected populations of Equus varies from 1.44 to 1.87, except for E. zebrathat showed a value of 0.40 (Table 7). Ro > 1 indicates an increasing population (Monchot et al., 2012; Martín-González etal., 2016).

Table 7. Comparison of the net reproductive rate (Ro), mean generational time (T ), exponential growth rate (r), growthrate (λ), and damping ratio (ρ) of the population of Equus conversidens from Hidalgo and selected populations of extinct

x lx sx mx l mx x

o

NISP

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b

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and extant Equus. A. E. conversidens (left and right value is for MNI and NISP indices respectively), B. E. lambei, C. E.mosbachensis, D. E. germanicus, E. E. quagga, F. E. zebra.

A 1.48, 1.79 9.21, 9.41 0.04, 0.06 1.04, 1.06 1.21, 1.24

B 1.66 9.08 0.07 1.07 1.21

C 1.44 6.86 0.05 1.05 1.36

D 1.87 7.53 0.08 1.08 1.29

E 1.78 10.31 0.06 1.05 1.21

F 0.40 8.40 −0.11 0.89 1.37

The mean generation time (Tb) of the population of E. conversidens is 9.21 and 9.41 using the MNI and NISP indicesrespectively, indicating that females produced an average of 1.63 descendants in about 9.31 years. This pattern of anincreasing population is also observed in E. lambei, E. mosbachensis, E. quagga, and E. germanicus, although the latterpopulation should be related to rapid population growth because females produced 1.87 descendants in about 7.53 years.By contrast, the population of E. zebra is related to a declining population, considering that females produced 0.40descendants in about 8.4 years (Table 7).

The exponential growth rate (r) related to the population growth of E. conversidens is 0.04 (MNI) and 0.06 (NISP) whichis very close to other selected species where r is about 0.06; only E. zebra has an r = – 0.11 indicating that the number ofindividuals in the population decreased by 11% each year (Table 7).

The asymptotic growth rate (λ) of E. conversidens and the other selected horse populations is > 1, except for E. zebrawhere λ is < 1 (Table 7). The average asymptotic growth rate for the species considered (excluding E. zebra) is 1.06. In thisregard, when λ = 1 suggests perfect stability and a non-impacted population (Fernandez et al., 2017). Ransom et al. (2016:pp.76) reported average λ of 1.12 for populations of wild and feral equids, Gaillard et al. (2000: pp. 370) indicated valuesof λ ranging from 1.25 to 1.35 related to maximum annual population growth rates in extant equids, and Fernandez et al.(2017: pp. 488) reported values ranging from 1.21 to 1.26 in extant populations of wild equids that all evolve in optimalconditions. The values of λ for the population of E. conversidens (MNI: 1.04, NISP: 1.06) from Hidalgo are somewhatlower than those observed in some populations of extant wild horses, although similar to the populations of E. lambei(λ = 1.07), E. mosbachensis (λ = 1.05), and E. germanicus (λ = 1.08) (Table 7).

Some populations of E. caballus and Equus ferus przewalskii inhabited areas without significant pressure frompredators, presenting somewhat high survival rates in all age class intervals (Ransom et al., 2016). By contrast, thepopulations of E. lambei, E. mosbachensis, and E. germanicus are related to the selective influence of diverse predatorswith a preference for adult individuals (Burke and Cinq-Mars, 1998; Fernandez and Legendre, 2003; Fernandez et al.,2006). This could have occurred in the population of Equus conversidens from Hidalgo, which is in agreement with theobserved prime-dominated mortality pattern.

The analysis of population dynamics of E. conversidens from Hidalgo suggests a relatively stable increasing populationwith slow growth (λ = 1.05, Tb = 9.31). Similar population dynamics are observed in stable populations of the modernplains zebra E. quagga from Serengeti, Tanzania (λ = 1.01, Tb = 8.24) (Grange et al., 2004), and different to that ofdeclining populations of the same species from Hwange, Zimbabwe related to intensive predation (λ = 0.094, Tb = 12.58)(Grange et al., 2015).

The damping ratio (ρ) of the population of E. conversidens is about 1.22 (Table 7). A damping ratio greater than 1indicates a population that stabilized around the λ somewhat rapidly after perturbations regardless of populationstructure (Fernandez et al., 2017). Here this parameter is slightly greater than 1 for all the species considered, suggestingthe convergence to stable size distribution was not so rapid. According to their values of ρ, a similar pattern should beexpected for the populations of E. conversidens, E. lambei, and E. quagga and to a lesser extent with E. germanicus(Table 7).

Population Ro Tb r λ ρ

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Fig. 4 shows the relative population size and fluctuations of the asymptotic growth rate over time for all the equids of thisstudy. The population size of E. conversidens derived from the NISP is higher than the one derived from the MNI, butboth indicate an increasing population; a similar pattern is observed in the populations of E. mosbachensis and E.quagga. Furthermore, E. germanicus has the fastest population growth, which is consistent with its highest values of netreproductive rate, exponential growth rate, and growth rate among the populations compared (Fig. 4, Table 7). Bycontrast, E. zebra is the only species that shows a negative growth curve reflecting a declining population (Fig. 4).


Fig. 4. (a) Projection of the population growth and (b) transient dynamics of Equus conversidens from the late Pleistoceneof Hidalgo and selected populations of extinct and extant Equus.

The fluctuations of the asymptotic growth rate (λ) suggest that the population of E. conversidens stabilized at theprojection time t and t derived from the MNI and NISP indices respectively. The projection from the MNI shows thatthe lambda values decreased from the time t (λ = 1.055) to t9 (λ = 1.042), indicating fluctuations in the age structureduring this time interval and the slower population growth; other minor fluctuations should occur before reachingstabilization at t . The projection from the NISP shows that the lambda values decreased from the time t (λ = 1.081) to t6(λ = 1.062), indicating major fluctuations during this time interval and reaching stabilization at t where the populationexperienced faster and constant growth (Fig. 4b). Both projections suggest fluctuations in the age structure related tojuvenile (0–3 years) and young-adults (3–6 years) age classes, occurring when the population experienced slower growth.Usually, these early fluctuations in the projection of a population are related to the initial conditions of the demographicparameters in the age structure (Fernandez et al., 2017).

The projection curve of E. conversidens is very similar to that of E. germanicus (t ), E. quagga (t ), and E. lambei (t ),whereas the population of E. mosbachensis converge to stable size more rapidly (t ) (Fig. 4a). All of these fluctuations areconsistent with the observed damping ratios for each species.

The elasticity values derived from all species are shown in the ternary diagram of Fig. 5. It indicates that survival,compared to fertility, is the most important demographic parameter in the population growth. It has been observed thatsurvival is an important demographic pattern in the population growth of ungulates (Gaillard et al., 2000; Oli, 2004;Funston and Mills, 2006). In the ternary diagram, the population of E. conversidens and the selected horse populationsoccupy a plot space with average elasticity values of fecundity lesser than 0.20 (e(F) = 0.12) and average elasticity values ofsurvival greater than 0.80 (e(Sj) = 0.46; e(Sa) = 0.42). These values are related to populations with slow life histories (Oli,2004).

27 31

0

27 0

31

26 29 31

17

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Fig. 5. Ternary diagram of elasticity values of Equus conversidens from the late Pleistocene of Hidalgo and selectedpopulations of extinct and extant Equus. e (F), fecundity; e (Sj), survivorship of juvenile individuals (0–3 years); e (Sa),survivorship of adults (≥4 years). Horse populations: 1, E. conversidens (NISP); 1′, E. conversidens (MNI); 2, E. lambei;3, E. mosbachensis; 4, E. germanicus; 5, E. quagga; 6, E. zebra.

The demographic models of E. conversidens derived from MNI and NISP indices suggest that the survival of primeindividuals (>3 years) is more important for population growth, as with E. quagga and E. zebra. The survival of juveniles(0–3 years of age) for E. lambei, E. mosbachensis and E. germanicus is somewhat more important for population growth.Finally, we observed that all the populations considered in this study fall within the “slow” area of the ternary diagram,which also reflects the theory of fast-slow continuum life stories (see review Oli, 2004). Our results are clearly related tothe life history of large mammals characterized by late maturity, long lifespan, high survival and fecundity rates as well ashigher λ in comparison with the “fast model”. By the same token, and despite the concerns regarding the adequacy of ther-K selection theory (Stearns, 1992 but see Boyce, 1984), the demographic parameters of all species studied here allow usto classify them as K-strategists in stable ecosystems (Pianka, 1970).

4.4. Ecological considerations

The extant plains zebra Equus quagga has a similar body size (ca. 300 kg) and is the closest relative species of E.conversidens (Alberdi et al., 1995; Barron-Ortiz et al., 2019), being a suitable model for comparison purposes. Severalstudies have shown that population dynamics of E. quagga is related to multiple factors, including seasonality, predation,environmental conditions, and/or resource availability (Mohelman, 2002; Georgiadis et al., 2003; Grange et al., 2004,2015; Funston and Mills, 2006; Owen-Smith, 2008; Rubenstein, 2010; Ransom et al., 2016; Kiffner et al., 2017).

Isotopic ecology and palynological studies indicate that savanna-like habitats with flood plains and channels were presentin southeastern Hidalgo during the late Pleistocene (Bravo-Cuevas et al., 2017; Pineda-Maldonado, 2017). This woodland-grassland ecosystem sustained a high diversity of large mammalian herbivores, providing a suitable environmentcharacterized by diverse food plant resources (Bravo-Cuevas et al., 2017). Considering their flexible feeding strategies,observed through mesowear and stable isotope analyses, individuals of E. conversidens were able to exploit the availablefood resources in different habitats (Bravo-Cuevas et al., 2011, 2017).

We propose that the influence of ambush (American lion) and cursorial social predators (dire wolf) are related to themortality profile of the population of E. conversidens. Other ungulates that represented potential prey of these largecarnivores have been reported at southeastern Hidalgo, including llamas, camels, deer, antelopes, and bison (Bravo-Cuevas et al., 2016). In this regard, populations of E. quagga experience different intensities of predation (Georgiadis etal., 2003; Owen-Smith, 2008; Funston and Mills, 2006), although it is not a limiting factor of population growth if thereis an abundance of large preys (Funston and Mills, 2006).

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Alberdi et al., 1995

Badgley, 1986

Bai et al., 2011

Given the above, the population dynamics of Equus conversidens from Hidalgo were regulated by the diversity of foodresources, as well as the selective predation of large carnivores, maintaining a stationary population that increasedslightly (Bowyer et al., 2014).

5. ConclusionThe current ecological standard tools such as life tables (time-specific model) and Lewis-Leslie matrices, are very efficientto derive paleodemographic parameters from the late Pleistocene cohorts of E. conversidens from Hidalgo. We comparedthis species to a sample of modern equids (E. asinus, E. zebra, E. hemionus, E. caballus) and fossil species (E.mosbachensis, E. germanicus, E. lambei) from different habitats. The mortality pattern of E. conversidens is a prime-dominated profile sometimes associated to hunting pressure or density-mediated destruction of faunal remains. Wepropose that the influence of natural predators produced this pattern, considering the association of large-sizedpredators, such as the dire wolf (Canis dirus) and the American lion (Panthera atrox), at southeastern Hidalgo during thelatter half of the Pleistocene. By the same token, it is proposed that population dynamics of E. conversidens from Hidalgowere regulated by predation and resource availability as it occurs in the extant plains zebra, E. quagga.

Authors contributionAlexis Pérez-Pérez: Conceptualization, Methodology, analysis, writing. Victor Manuel Bravo-Cuevas: Conceptualization,writing, editing. Philippe Fernandez: Conceptualization, Methodology, writing.

Declaration of competing interestThe authors declare that they have no known competing financial interests or personal relationships that could haveappeared to influence the work reported in this paper.

AcknowledgmentsThanks to Francisco J. Vega for handling and editing of the manuscript. We acknowledge the comments of theanonymous reviewers that improved the final version of the manuscript. We also appreciate the kind support of Nuria M.Morales-García (Bristol University, UK) for the linguistical review of the text.

Appendix A. Supplementary dataThe following is the Supplementary data to this article:

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ReferencesM.T. Alberdi, J.L. Prado, E. Ortiz-Jaureguizar

Patterns of body size changes in fossil and living Equini (Perissodactyla)Biol. J. Linn. Soc., 54 (1995), pp. 349-370Article Download PDF View Record in Scopus Google Scholar

C. BadgleyCounting individuals in mammalian fossil assemblages from fluvial environmentsPalaios, 1 (1986), pp. 328-338, 10.2307/3514695CrossRef View Record in Scopus Google Scholar

B. Bai, Y. Wang, J. Meng, X. Jin, Q. Li, P. Li

Download

https://ars-els-cdn-com.lama.univ-amu.fr/content/image/1-s2.0-S089598112030643X-mmc1.docx

https://ars-els-cdn-com.lama.univ-amu.fr/content/image/1-s2.0-S089598112030643X-mmc1.docx

https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/0024406695900152

https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/0024406695900152/pdf?md5=80876b3665fb689a46aa028a8907aa0e&pid=1-s2.0-0024406695900152-main.pdf

https://www.scopus.com/inward/record.url?eid=2-s2.0-0029109866&partnerID=10&rel=R3.0.0

https://scholar.google.com/scholar?q=Patterns%20of%20body%20size%20changes%20in%20fossil%20and%20living%20Equini

https://doi.org/10.2307/3514695

https://doi.org/10.2307/3514695


https://scholar.google.com/scholar_lookup?title=Counting%20individuals%20in%20mammalian%20fossil%20assemblages%20from%20fluvial%20environments&publication_year=1986&author=C.%20Badgley




Barrón-Ortiz et al., 2019

Bell et al., 2004

Behrensmeyer et al., 2000

Bernardelli, 1941

Botkin and Miller, 1974

Bowyer et al., 2014

Boyce, 1984

Brain, 1981

Bravo-Cuevas, 2001

Bravo-Cuevas, 2002

Taphonomic analyses of an early eocene litolophus (perissodactyla, chalicotherioidea) assemblagefrom the erlian basin, inner Mongolia, ChinaPalaios, 26 (2011), pp. 187-196, 10.2110/palo.2010.p10-079rCrossRef View Record in Scopus Google Scholar

C.I. Barrón-Ortiz, L.S. Avilla, C.N. Jass, V.M. Bravo-Cuevas, H. Machado, D. MothéWhat is Equus? Reconciling taxonomy and phylogenetic analysesFrontiers in Ecology and Evolution, 343 (2019), pp. 1-14, 10.3389/fevo.2019.00343Google Scholar

Ch J. Bell, E.L. Lundelius Jr., A.D. Barnosky, R.W. Graham, E.H. Lindsay, D.R. Ruez Jr., H.A. SemkenJr., H.A. Webb, R.J. ZakrewskiThe blancan, irvingtonian, and rancholabrean mammal agesM.O. Woodburne (Ed.), Late Cretaceous and Cenozoic Mammals of North America, Columbia University Press,United States (2004), pp. 232-314CrossRef Google Scholar

A.K. Behrensmeyer, S.M. Kidwell, R.A. GastaldoTaphonomy and paleobiologyPaleobiology, 26 (2000), pp. 103-147, 10.1017/S0094837300026907View Record in Scopus Google Scholar

H. BernardelliPopulation wavesJ. Bihar Res. Soc., 31 (I) (1941), pp. 3-18Google Scholar

D.B. Botkin, R.S. MillerMortality rates and survival of birdsAm. Nat., 108 (1974), pp. 181-192http://www.jstor.com/stable/2459849View Record in Scopus Google Scholar

R.T. Bowyer, V.C. Bleich, B.K.M. Stewart, J.C. Whiting, K.L. MonteithDensity dependence in ungulates: a review of causes, and concepts with some clarificationsCalif. Fish Game, 100 (2014), pp. 550-572View Record in Scopus Google Scholar

M.S. BoyceRestitution of r- and K-selection as a model of density-dependent natural selectionAnnu. Rev. Ecol. Evol. Syst., 15 (1984), pp. 427-447, 10.1146/annurev.es.15.110184.002235CrossRef View Record in Scopus Google Scholar

C.K. BrainThe Hunters or the Hunted? an Introduction to African Cave TaphonomyThe University of Chicago Press, Chicago, Illinois (1981), p. 365View Record in Scopus Google Scholar

V.M. Bravo-CuevasPleistocene faunal assemblage from south-central region of the state of Hidalgo, Central MexicoJ. Vertebr. Paleontol., 21 (2001), p. 35AGoogle Scholar

V.M. Bravo-CuevasDiferenciación geológica y bioestratigráfica de la Formación Tarango para el estado de Hidalgo,Centro de México, México, Hidalgo, Centro de Investigaciones en Ciencias de la TierraUniversidad Autónoma del Estado de Hidalgo, México, Technical Report (2002), p. 59View Record in Scopus Google Scholar

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https://doi.org/10.2110/palo.2010.p10-079r

https://doi.org/10.2110/palo.2010.p10-079r


https://scholar.google.com/scholar?q=Taphonomic%20analyses%20of%20an%20early%20eocene%20litolophus%20%20assemblage%20from%20the%20erlian%20basin,%20inner%20Mongolia,%20China

https://doi.org/10.3389/fevo.2019.00343

https://scholar.google.com/scholar?q=What%20is%20Equus%20Reconciling%20taxonomy%20and%20phylogenetic%20analyses

https://doi.org/10.1093/oseo/instance.00186936

https://scholar.google.com/scholar_lookup?title=The%20blancan%2C%20irvingtonian%2C%20and%20rancholabrean%20mammal%20ages&publication_year=2004&author=Ch%20J.%20Bell&author=E.L.%20Lundelius%20Jr.&author=A.D.%20Barnosky&author=R.W.%20Graham&author=E.H.%20Lindsay&author=D.R.%20Ruez%20Jr.&author=H.A.%20Semken%20Jr.&author=H.A.%20Webb&author=R.J.%20Zakrewski

https://doi.org/10.1017/S0094837300026907


https://scholar.google.com/scholar_lookup?title=Taphonomy%20and%20paleobiology&publication_year=2000&author=A.K.%20Behrensmeyer&author=S.M.%20Kidwell&author=R.A.%20Gastaldo

https://scholar.google.com/scholar_lookup?title=Population%20waves&publication_year=1941&author=H.%20Bernardelli

http://www.jstor.com/stable/2459849


https://scholar.google.com/scholar_lookup?title=Mortality%20rates%20and%20survival%20of%20birds&publication_year=1974&author=D.B.%20Botkin&author=R.S.%20Miller


https://scholar.google.com/scholar_lookup?title=Density%20dependence%20in%20ungulates%3A%20a%20review%20of%20causes%2C%20and%20concepts%20with%20some%20clarifications&publication_year=2014&author=R.T.%20Bowyer&author=V.C.%20Bleich&author=B.K.M.%20Stewart&author=J.C.%20Whiting&author=K.L.%20Monteith

https://doi.org/10.1146/annurev.es.15.110184.002235

https://doi.org/10.1146/annurev.es.15.110184.002235


https://scholar.google.com/scholar_lookup?title=Restitution%20of%20r-%20and%20K-selection%20as%20a%20model%20of%20density-dependent%20natural%20selection&publication_year=1984&author=M.S.%20Boyce


https://scholar.google.com/scholar?q=The%20Hunters%20or%20the%20Hunted%20an%20Introduction%20to%20African%20Cave%20Taphonomy

https://scholar.google.com/scholar_lookup?title=Pleistocene%20faunal%20assemblage%20from%20south-central%20region%20of%20the%20state%20of%20Hidalgo%2C%20Central%20Mexico&publication_year=2001&author=V.M.%20Bravo-Cuevas


https://scholar.google.com/scholar_lookup?title=Diferenciaci%C3%B3n%20geol%C3%B3gica%20y%20bioestratigr%C3%A1fica%20de%20la%20Formaci%C3%B3n%20Tarango%20para%20el%20estado%20de%20Hidalgo%2C%20Centro%20de%20M%C3%A9xico%2C%20M%C3%A9xico%2C%20Hidalgo%2C%20Centro%20de%20Investigaciones%20en%20Ciencias%20de%20la%20Tierra&publication_year=2002&author=V.M.%20Bravo-Cuevas




Bravo-Cuevas and Vázquez-Cortes, 2019

Bravo-Cuevas et al., 2009a

Bravo-Cuevas et al., 2009b

Bravo-Cuevas et al., 2011







V. Bravo-Cuevas, V. Vázquez-CortesTaxonomy, ecology, and biochronological implications of bison (artiodactyla, bovidae) from thelate Pleistocene of Hidalgo and puebla, central Mexico11th North American Paleontological Conference Program with Abstracts. PaleoBios, 36 (2019), p. 79Google Scholar

V.M. Bravo-Cuevas, M.A. Cabral-Perdomo, E. Ortiz-Caballero, Vargas J. PriegoLa Megafauna del PleistocenoK.A. González-Rodríguez, C. Cuevas-Cardona, J.M. Castillo-Cerón (Eds.), Los Fósiles del Estado de Hidalgo,Universidad Autónoma del Estado de Hidalgo, Pachuca, Hidalgo, México (2009), pp. 85-96View Record in Scopus Google Scholar

V.M. Bravo-Cuevas, E. Ortiz-Caballero, M.A. Cabral-PerdomoGliptodontes (Xenarthra, Glyptodontidae) del Pleistoceno Tardío (Rancholabreano) de Hidalgo,Centro de MéxicoBol. Soc. Geol. Mex., 61 (2009), pp. 267-276, 10.18268/BSGM2009v61n2a14CrossRef View Record in Scopus Google Scholar

V.M. Bravo-Cuevas, E. Jiménez-Hidalgo, J. Priego-VargasTaxonomía y hábito alimentario de Equus conversidens (Perissodactyla, Equidae) del PleistocenoTardío (Rancholabreano) de Hidalgo, centro de MéxicoRev. Mex. Ciencias Geol., 28 (2011), pp. 65-82View Record in Scopus Google Scholar

V.M. Bravo-Cuevas, E. Jiménez-Hidalgo, G. Cuevas-Ruíz, M.A. Cabral-PerdomoA small camelid Hemiauchenia from the late Pleistocene of Hidalgo, central MexicoActa Palaeontol. Pol., 57 (2012), pp. 497-508, 10.4202/app.2011.0005CrossRef View Record in Scopus Google Scholar

V.M. Bravo-Cuevas, E. Jiménez-Hidalgo, M.A. Cabral-Perdomo, J. Priego-VargasTaxonomy and notes on the paleobiology of the late Pleistocene (rancholabrean) antilocaprids(mammalia, artiodactyla, antilocapridae) from the state of Hidalgo, central MexicoRev. Mex. Ciencias Geol., 30 (2013), pp. 601-613View Record in Scopus Google Scholar

V.M. Bravo-Cuevas, N.M. Morales-García, M.A. Cabral-PerdomoDescription of mastodons (Mammut americanum) from the late Pleistocene of southeasternHidalgo, central MexicoBol. Soc. Geol. Mex., 67 (2015), pp. 337-347, 10.18268/BSGM2015v67n2a14CrossRef View Record in Scopus Google Scholar

V.M. Bravo-Cuevas, J. Priego-Vargas, M.A. Cabral-Perdomo, M.A. Pineda MaldonadoFirst occurrence of Panthera atrox (Felidae, Pantherinae) in the Mexican state of Hidalgo and areview of the record of felids from the Pleistocene of MexicoFossil Record, 19 (2016), pp. 131-141, 10.5194/fr-19-131-2016CrossRef View Record in Scopus Google Scholar

V.M. Bravo-Cuevas, F. Rivals, J. Priego-VargasPaleoecology (δ C and δ O stable isotopes analysis) of a mammalian assemblage from the latePleistocene of Hidalgo, central Mexico and implications on the better understanding ofenvironmental conditions in temperate North America (18º –36º N Lat.)Palaeogeogr. Palaeoclimatol. Palaeoecol., 485 (2017), pp. 632-643, 10.1016/j.palaeo.2017.07.018Article Download PDF View Record in Scopus Google Scholar

V.M. Bravo-Cuevas, E. Ortiz-Caballero, E. Jiménez-Hidalgo, C.I. Barrón-Ortiz, J.M. TheodorTaxonomía y hábito alimentario de ejemplares de Mammuthus columbi (Proboscidea:elephantidae) del centro y sur de MéxicoBol. Soc. Geol. Mex., 72 (2020), pp. 1-28, 10.18268/BSGM2018v72n1a#View Record in Scopus Google Scholar

13 18

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https://scholar.google.com/scholar?q=Taxonomy,%20ecology,%20and%20biochronological%20implications%20of%20bison%20%20from%20the%20late%20Pleistocene%20of%20Hidalgo%20and%20puebla,%20central%20Mexico


https://scholar.google.com/scholar_lookup?title=La%20Megafauna%20del%20Pleistoceno&publication_year=2009&author=V.M.%20Bravo-Cuevas&author=M.A.%20Cabral-Perdomo&author=E.%20Ortiz-Caballero&author=Vargas%20J.%20Priego

https://doi.org/10.18268/BSGM2009v61n2a14



https://scholar.google.com/scholar?q=Gliptodontes%20%20del%20Pleistoceno%20Tard%C3%ADo%20%20de%20Hidalgo,%20Centro%20de%20M%C3%A9xico


https://scholar.google.com/scholar?q=Taxonom%C3%ADa%20y%20h%C3%A1bito%20alimentario%20de%20Equus%20conversidens%20%20del%20Pleistoceno%20Tard%C3%ADo%20%20de%20Hidalgo,%20centro%20de%20M%C3%A9xico

https://doi.org/10.4202/app.2011.0005

https://doi.org/10.4202/app.2011.0005


https://scholar.google.com/scholar_lookup?title=A%20small%20camelid%20Hemiauchenia%20from%20the%20late%20Pleistocene%20of%20Hidalgo%2C%20central%20Mexico&publication_year=2012&author=V.M.%20Bravo-Cuevas&author=E.%20Jim%C3%A9nez-Hidalgo&author=G.%20Cuevas-Ru%C3%ADz&author=M.A.%20Cabral-Perdomo


https://scholar.google.com/scholar?q=Taxonomy%20and%20notes%20on%20the%20paleobiology%20of%20the%20late%20Pleistocene%20%20antilocaprids%20%20from%20the%20state%20of%20Hidalgo,%20central%20Mexico




https://scholar.google.com/scholar?q=Description%20of%20mastodons%20%20from%20the%20late%20Pleistocene%20of%20southeastern%20Hidalgo,%20central%20Mexico

https://doi.org/10.5194/fr-19-131-2016

https://doi.org/10.5194/fr-19-131-2016


https://scholar.google.com/scholar?q=First%20occurrence%20of%20Panthera%20atrox%20%20in%20the%20Mexican%20state%20of%20Hidalgo%20and%20a%20review%20of%20the%20record%20of%20felids%20from%20the%20Pleistocene%20of%20Mexico

https://doi.org/10.1016/j.palaeo.2017.07.018

https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/S003101821730216X

https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/S003101821730216X/pdfft?md5=71c8fba65f0189cf44ffa44c75f903cc&pid=1-s2.0-S003101821730216X-main.pdf


https://scholar.google.com/scholar?q=Paleoecology%20%20of%20a%20mammalian%20assemblage%20from%20the%20late%20Pleistocene%20of%20Hidalgo,%20central%20Mexico%20and%20implications%20on%20the%20better%20understanding%20of%20environmental%20conditions%20in%20temperate%20North%20America

https://doi.org/10.18268/BSGM2018v72n1a#


https://scholar.google.com/scholar?q=Taxonom%C3%ADa%20y%20h%C3%A1bito%20alimentario%20de%20ejemplares%20de%20Mammuthus%20columbi%20%20del%20centro%20y%20sur%20de%20M%C3%A9xico




Bryant, 1991

Burke, 1998

Caswell, 1978

Caswell, 2001

Caswell and Weeks, 1986

Caughley, 1966

Caughley, 1977

Discamps and Costamagno, 2015

Dodd and Stanton, 1990

Driver and Maxwell, 2013

Erickson et al., 2006

J.D. BryantAge-frequency profiles of micromammals and population dynamics of Proheteromys

floridanus (Rodentia) from the early Miocene Thomas Farm site, Florida (USA)Palaeogeogr. Palaeoclimatol. Palaeoecol., 85 (1991), pp. 1-14, 10.1016/0031-0182(91)90022-JView Record in Scopus Google Scholar

A. Burke, J. Cinq-MarsPaleoethological reconstruction and taphonomy of Equus lambei from the bluefish Caves, Yukonterritory, CanadaArtic, 51 (1998), pp. 105-115, 10.14430/arctic1052View Record in Scopus Google Scholar

H. CaswellA general formula for the sensitivity of population growth rate to changes in life historyparametersTheor. Popul. Biol., 14 (1978), pp. 215-230, 10.1016/0040-5809(78)90025-4Article Download PDF View Record in Scopus Google Scholar

H. CaswellMatrix Population Models: Construction, Analysis and Interpretation(second ed.), Sinauer Associates, Sunderland, Massachusetts (2001), p. 722Google Scholar

H. Caswell, D.E. WeeksTwo-sex models: chaos, extinction, and other dynamic consequences of sexAm. Nat., 128 (1986), pp. 707-735, 10.1086/284598View Record in Scopus Google Scholar

G. CaughleyMortality patterns in mammalsEcology, 47 (1966), pp. 906-918CrossRef Google Scholar

G. CaughleyAnalysis of Vertebrate PopulationsJohn Wiley and Sons, London (1977), p. 234View Record in Scopus Google Scholar

E. Discamps, S. CostamagnoImproving mortality profile analysis in zooarchaeology: a revised zoning for ternary diagramsJ. Archaeol. Sci., 58 (2015), pp. 62-76, 10.1016/j.jas.2015.03.021Article Download PDF View Record in Scopus Google Scholar

J.R. Dodd, R.J. StantonPopulations in PaleoecologyJ.R. Dodd, R.J. Stanton (Eds.), Paleoecology: Concepts and Applications, John Wiley and Sons, New York (1990),pp. 277-322Google Scholar

J.C. Driver, D. MaxwellBison death assemblages and the interpretation of human hunting behaviorQuat. Int., 297 (2013), pp. 100-109, 10.1016/j.quaint.2012.12.038Article Download PDF View Record in Scopus Google Scholar

G.M. Erickson, P.J. Currie, B.D. Inouye, A.A. WinnTyrannosaur life tables: an example of nonavian dinosaur population biologyScience, 313 (2006), pp. 213-217, 10.1126/science.1125721CrossRef View Record in Scopus Google Scholar

Download

https://doi.org/10.1016/0031-0182(91)90022-J


https://scholar.google.com/scholar?q=Age-frequency%20profiles%20of%20micromammals%20and%20population%20dynamics%20of%20Proheteromys%20floridanus%20%20from%20the%20early%20Miocene%20Thomas%20Farm%20site,%20Florida

https://doi.org/10.14430/arctic1052


https://scholar.google.com/scholar_lookup?title=Paleoethological%20reconstruction%20and%20taphonomy%20of%20Equus%20lambei%20from%20the%20bluefish%20Caves%2C%20Yukon%20territory%2C%20Canada&publication_year=1998&author=A.%20Burke&author=J.%20Cinq-Mars

https://doi.org/10.1016/0040-5809(78)90025-4


https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/0040580978900254/pdf?md5=5d2132ca70c15e3624c4caef0f71a1c4&pid=1-s2.0-0040580978900254-main.pdf


https://scholar.google.com/scholar_lookup?title=A%20general%20formula%20for%20the%20sensitivity%20of%20population%20growth%20rate%20to%20changes%20in%20life%20history%20parameters&publication_year=1978&author=H.%20Caswell

https://scholar.google.com/scholar_lookup?title=Matrix%20Population%20Models%3A%20Construction%2C%20Analysis%20and%20Interpretation&publication_year=2001&author=H.%20Caswell

https://doi.org/10.1086/284598


https://scholar.google.com/scholar_lookup?title=Two-sex%20models%3A%20chaos%2C%20extinction%2C%20and%20other%20dynamic%20consequences%20of%20sex&publication_year=1986&author=H.%20Caswell&author=D.E.%20Weeks

https://doi.org/10.2307/1935638

https://scholar.google.com/scholar_lookup?title=Mortality%20patterns%20in%20mammals&publication_year=1966&author=G.%20Caughley


https://scholar.google.com/scholar_lookup?title=Analysis%20of%20Vertebrate%20Populations&publication_year=1977&author=G.%20Caughley

https://doi.org/10.1016/j.jas.2015.03.021

https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/S0305440315001144

https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/S0305440315001144/pdfft?md5=432c2121c9d867452bb9902f357b095c&pid=1-s2.0-S0305440315001144-main.pdf


https://scholar.google.com/scholar_lookup?title=Improving%20mortality%20profile%20analysis%20in%20zooarchaeology%3A%20a%20revised%20zoning%20for%20ternary%20diagrams&publication_year=2015&author=E.%20Discamps&author=S.%20Costamagno

https://scholar.google.com/scholar_lookup?title=Populations%20in%20Paleoecology&publication_year=1990&author=J.R.%20Dodd&author=R.J.%20Stanton

https://doi.org/10.1016/j.quaint.2012.12.038


https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/S1040618213000037/pdfft?md5=084fe6e2838f1d2d64740613f40f5a28&pid=1-s2.0-S1040618213000037-main.pdf


https://scholar.google.com/scholar_lookup?title=Bison%20death%20assemblages%20and%20the%20interpretation%20of%20human%20hunting%20behavior&publication_year=2013&author=J.C.%20Driver&author=D.%20Maxwell

https://doi.org/10.1126/science.1125721

https://doi.org/10.1126/science.1125721


https://scholar.google.com/scholar_lookup?title=Tyrannosaur%20life%20tables%3A%20an%20example%20of%20nonavian%20dinosaur%20population%20biology&publication_year=2006&author=G.M.%20Erickson&author=P.J.%20Currie&author=B.D.%20Inouye&author=A.A.%20Winn




Erickson et al., 2009

Fernandez, 2009

Fernandez and Legendre, 2003

Fernandez and Boulbes, 2010

Fernandez et al., 2006

Fernandez et al., 2017

Fourvel et al., 2014

Fourvel et al., 2015

Funston and Mills, 2006

Fujiwara and Diaz-Lopez, 2017

G.M. Erickson, P.J. Makovicky, B.D. Inouye, C. Zhou, K. GaoA life table for Psittacosaurus lujiatunensis: initial insights into ornithischian dinosaurpopulation biologyAnat. Rec., 292 (2009), pp. 1514-1521, 10.1002/ar.20992CrossRef View Record in Scopus Google Scholar

P. FernandezDe l’estimation de l’âge individuel dentaire au modèle descriptif des structures d’âge des cohortesfossiles: l’exemple des Equidae et du time-specific model en contextes paléobiologiquespléistocènesBull. Soc. Prehist. Fr., 106 (2009), pp. 5-14, 10.3406/bspf.2009.13826CrossRef View Record in Scopus Google Scholar

P. Fernandez, S. LegendreMortality curves for horses from the Middle Palaeolithic site of Bau de l'Aubesier (Vaucluse,France): methodological, palaeo-ethnological, and palaeo-ecological approachesJ. Archaeol. Sci., 30 (2003), pp. 1577-1598, 10.1016/S0305-4403(03)00054-2Article Download PDF View Record in Scopus Google Scholar

P. Fernandez, N. BoulbesAnalyse démographique des cohortes du cheval pléistocène moyen de Romain-la-Roche (Doubs,France)Rev. Paleobiol., 29 (2010), pp. 771-801View Record in Scopus Google Scholar

P. Fernandez, J.-L. Guadelli, P. FosseApplying dynamics and comparing life tables for Pleistocene Equidae in anthropic (Bau del'Aubesier, Combe-Grenal) and carnivore (Fouvent) contexts with modern feral horse populations(Akagera, Pryor Mountain)J. Archaeol. Sci., 33 (2006), pp. 176-184, 10.1016/j.jas.2005.07.005Article Download PDF View Record in Scopus Google Scholar

P. Fernandez, C. Bonenfant, J.M. Gaillard, S. Legendre, H. MonchotLife tables and Leslie matrices for mammalian cohorts in different paleobiological contexts duringthe PleistoceneJ.P. Brugal (Ed.), Taphonomies, Editions des Archives Contemporaines, Paris (2017), pp. 477-497View Record in Scopus Google Scholar

J.B. Fourvel, P. Fosse, P. Fernandez, P.O. AntoineLa grotte de Fouvent, dit l'Abri Cuvier (Fouvent-le-Bas, Haute-Saône, France): analysetaphonomique d’un repaire d’hyènes du Pléistocène supérieur (OIS 3)Paléo, 25 (2014), pp. 79-99, 10.4000/paleo.2747CrossRef Google Scholar

J.B. Fourvel, P. Fosse, P. Fernandez, P.O. AntoineLarge mammals of fouvent saint-andoche (Haute-Saône, France): a glimpse into a late Pleistocenehyena denGeodiversitas, 37 (2015), pp. 237-266, 10.5252/g2015n2a5CrossRef View Record in Scopus Google Scholar

P.J. Funston, M.G.L. MillsThe influence of lion predation on the population dynamics of common large ungulates in theKruger National ParkS. Afr. J. Wildl. Res., 36 (2006), pp. 9-22View Record in Scopus Google Scholar

M. Fujiwara, J. Diaz-LopezConstructing stage-structured matrix population models from life tables: comparison of methodsPeerJ, 5 (2017), Article e3971

Download

https://doi.org/10.1002/ar.20992

https://doi.org/10.1002/ar.20992


https://scholar.google.com/scholar_lookup?title=A%20life%20table%20for%20Psittacosaurus%20lujiatunensis%3A%20initial%20insights%20into%20ornithischian%20dinosaur%20population%20biology&publication_year=2009&author=G.M.%20Erickson&author=P.J.%20Makovicky&author=B.D.%20Inouye&author=C.%20Zhou&author=K.%20Gao

https://doi.org/10.3406/bspf.2009.13826

https://doi.org/10.3406/bspf.2009.13826


https://scholar.google.com/scholar?q=De%20lestimation%20de%20l%C3%A2ge%20individuel%20dentaire%20au%20mod%C3%A8le%20descriptif%20des%20structures%20d%C3%A2ge%20des%20cohortes%20fossiles:%20lexemple%20des%20Equidae%20et%20du%20time-specific%20model%20en%20contextes%20pal%C3%A9obiologiques%20pl%C3%A9istoc%C3%A8nes

https://doi.org/10.1016/S0305-4403(03)00054-2


https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/S0305440303000542/pdfft?md5=628b53d968d34319b03b4ad74ab0378a&pid=1-s2.0-S0305440303000542-main.pdf


https://scholar.google.com/scholar?q=Mortality%20curves%20for%20horses%20from%20the%20Middle%20Palaeolithic%20site%20of%20Bau%20de%20lAubesier%20:%20methodological,%20palaeo-ethnological,%20and%20palaeo-ecological%20approaches


https://scholar.google.com/scholar?q=Analyse%20d%C3%A9mographique%20des%20cohortes%20du%20cheval%20pl%C3%A9istoc%C3%A8ne%20moyen%20de%20Romain-la-Roche



https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/S0305440305001548/pdfft?md5=eba0b87282f414141e8b745482907f50&pid=1-s2.0-S0305440305001548-main.pdf


https://scholar.google.com/scholar?q=Applying%20dynamics%20and%20comparing%20life%20tables%20for%20Pleistocene%20Equidae%20in%20anthropic%20%20and%20carnivore%20%20contexts%20with%20modern%20feral%20horse%20populations


https://scholar.google.com/scholar_lookup?title=Life%20tables%20and%20Leslie%20matrices%20for%20mammalian%20cohorts%20in%20different%20paleobiological%20contexts%20during%20the%20Pleistocene&publication_year=2017&author=P.%20Fernandez&author=C.%20Bonenfant&author=J.M.%20Gaillard&author=S.%20Legendre&author=H.%20Monchot

https://doi.org/10.4000/paleo.2747

https://doi.org/10.4000/paleo.2747

https://scholar.google.com/scholar?q=La%20grotte%20de%20Fouvent,%20dit%20lAbri%20Cuvier%20:%20analyse%20taphonomique%20dun%20repaire%20dhy%C3%A8nes%20du%20Pl%C3%A9istoc%C3%A8ne%20sup%C3%A9rieur

https://doi.org/10.5252/g2015n2a5

https://doi.org/10.5252/g2015n2a5


https://scholar.google.com/scholar?q=Large%20mammals%20of%20fouvent%20saint-andoche%20:%20a%20glimpse%20into%20a%20late%20Pleistocene%20hyena%20den


https://scholar.google.com/scholar_lookup?title=The%20influence%20of%20lion%20predation%20on%20the%20population%20dynamics%20of%20common%20large%20ungulates%20in%20the%20Kruger%20National%20Park&publication_year=2006&author=P.J.%20Funston&author=M.G.L.%20Mills




Gaillard et al., 2000

Garrott and Taylor, 1990

Georgiadis et al., 2003

Grange et al., 2004

Grange et al., 2015

Grayson, 1979

Grayson, 1984

Hood, 2008

Hulbert, 1982

Hulbert, 1984

Hunter, 1998

CrossRef Google Scholar

J.M. Gaillard, M. Festa-Bianchet, N.G. Yoccoz, A. Loison, C. ToïgoTemporal variation in fitness components and population dynamics of large herbivoresAnnu. Rev. Ecol. Evol. Syst., 31 (2000), pp. 367-393, 10.1146/annurev.ecolsys.31.1.367CrossRef View Record in Scopus Google Scholar

R.A. Garrott, L. TaylorDynamics of a feral horse population in MontanaJ. Wildl. Manag., 54 (1990), pp. 603-612, 10.2307/3809357CrossRef View Record in Scopus Google Scholar

N. Georgiadis, M. Hack, K. TurpinThe influence of rainfall on zebra population dynamics: implications for managementJ. Appl. Ecol., 40 (2003), pp. 125-136, 10.1046/j.1365-2664.2003.00796.xView Record in Scopus Google Scholar

S. Grange, P. Duncan, J.-M. Gaillard, A.R.E. Sinclair, P.J.P. Gogan, C. Packer, H. Hofer, M. EastWhat limits the Serengeti zebra population?Oecologia, 140 (2004), pp. 523-532, 10.1007/s00442-004-1567-6View Record in Scopus Google Scholar

S. Grange, F. Barnier, P. Duncan, J.-M. Gaillard, M. Valeix, H. Ncube, S. Périquet, H. FritzDemography of plains zebras (Equus quagga) under heavy predationPopul. Ecol., 57 (2015), pp. 201-214, 10.1007/s10144-014-0469-7CrossRef View Record in Scopus Google Scholar

D.K. GraysonOn the quantification of vertebrate archaeofaunasM.B. Schiffer (Ed.), Advances in Archaeological Method and Theory, vol. 2, Academic Press, New York (1979), pp.199-237Google Scholar

D.K. GraysonQuantitative Zooarchaeology: Topics in the Analysis of Archaeological FaunasAcademic Press, New York (1984), p. 202View Record in Scopus Google Scholar

G.M. HoodPopTools version 3.2.5Available online at:http://www.cse.csiro.au/poptools (2008)Google Scholar

R.C. Hulbert Jr.Dynamics of the three-toed horse neohipparion from the late miocene of FloridaPaleobiology, 8 (1982), pp. 159-167, 10.1017/S0094837300004504View Record in Scopus Google Scholar

R.C. Hulbert Jr.Paleoecology and population dynamics of the early Miocene (Hemingfordian) horse Parahippusleonensis from the Thomas Farm site, FloridaJ. Vertebr. Paleontol., 4 (1984), pp. 547-558, 10.1080/02724634.1984.10012030CrossRef View Record in Scopus Google Scholar

J.P. HunterPaleoecology meets ecology on questions of scaleTrends Ecol. Evol., 13 (1998), pp. 478-479Article Download PDF View Record in Scopus Google Scholar

Download

https://doi.org/10.7717/peerj.3971

https://scholar.google.com/scholar_lookup?title=Constructing%20stage-structured%20matrix%20population%20models%20from%20life%20tables%3A%20comparison%20of%20methods&publication_year=2017&author=M.%20Fujiwara&author=J.%20Diaz-Lopez

https://doi.org/10.1146/annurev.ecolsys.31.1.367

https://doi.org/10.1146/annurev.ecolsys.31.1.367


https://scholar.google.com/scholar_lookup?title=Temporal%20variation%20in%20fitness%20components%20and%20population%20dynamics%20of%20large%20herbivores&publication_year=2000&author=J.M.%20Gaillard&author=M.%20Festa-Bianchet&author=N.G.%20Yoccoz&author=A.%20Loison&author=C.%20To%C3%AFgo

https://doi.org/10.2307/3809357

https://doi.org/10.2307/3809357


https://scholar.google.com/scholar_lookup?title=Dynamics%20of%20a%20feral%20horse%20population%20in%20Montana&publication_year=1990&author=R.A.%20Garrott&author=L.%20Taylor

https://doi.org/10.1046/j.1365-2664.2003.00796.x


https://scholar.google.com/scholar_lookup?title=The%20influence%20of%20rainfall%20on%20zebra%20population%20dynamics%3A%20implications%20for%20management&publication_year=2003&author=N.%20Georgiadis&author=M.%20Hack&author=K.%20Turpin

https://doi.org/10.1007/s00442-004-1567-6


https://scholar.google.com/scholar?q=What%20limits%20the%20Serengeti%20zebra%20population

https://doi.org/10.1007/s10144-014-0469-7

https://doi.org/10.1007/s10144-014-0469-7


https://scholar.google.com/scholar?q=Demography%20of%20plains%20zebras%20%20under%20heavy%20predation

https://scholar.google.com/scholar_lookup?title=On%20the%20quantification%20of%20vertebrate%20archaeofaunas&publication_year=1979&author=D.K.%20Grayson


https://scholar.google.com/scholar_lookup?title=Quantitative%20Zooarchaeology%3A%20Topics%20in%20the%20Analysis%20of%20Archaeological%20Faunas&publication_year=1984&author=D.K.%20Grayson

http://www.cse.csiro.au/poptools

https://scholar.google.com/scholar_lookup?title=PopTools%20version%203.2.5&publication_year=2008&author=G.M.%20Hood

https://doi.org/10.1017/S0094837300004504


https://scholar.google.com/scholar_lookup?title=Dynamics%20of%20the%20three-toed%20horse%20neohipparion%20from%20the%20late%20miocene%20of%20Florida&publication_year=1982&author=R.C.%20Hulbert%20Jr.

https://doi.org/10.1080/02724634.1984.10012030

https://doi.org/10.1080/02724634.1984.10012030


https://scholar.google.com/scholar?q=Paleoecology%20and%20population%20dynamics%20of%20the%20early%20Miocene%20%20horse%20Parahippus%20leonensis%20from%20the%20Thomas%20Farm%20site,%20Florida


https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/S0169534798014888/pdfft?md5=895d92ea32bc21b0107ac6dadc2ec385&pid=1-s2.0-S0169534798014888-main.pdf


https://scholar.google.com/scholar_lookup?title=Paleoecology%20meets%20ecology%20on%20questions%20of%20scale&publication_year=1998&author=J.P.%20Hunter




JCGM, 2008

Joubert, 1974

Kahlke and Gaudzinski, 2005

Kiffner et al., 2017

Klein and Cruz-Uribe, 1984

Koike and Ohtaishi, 1985

Koike and Ohtaishi, 1987

Korth and Evander, 1986

Kurtén, 1953

Kurtén, 1954a

Kurtén, 1954b

JCGMEvaluation of measurement data - guide to the expression of uncertainty in measurementDocument produced by Working Group 1 of the Joint Committee for Guides in Metrology (JCGM/WG 1) (2008), p.134https://www.bipm.org/fr/publications/guides/Google Scholar

E. JoubertComposition and limiting factors of a Khoma Hochland population of Hartmann zebra Equuszebra hartmannaeMadoqua, 1 (1974), pp. 49-53View Record in Scopus Google Scholar

R.D. Kahlke, S. GaudzinskiThe blessing of a great flood: differentiation of mortality patterns in the large mammal record ofthe Lower Pleistocene fluvial site of Untermassfeld (Germany) and its relevance for theinterpretation of faunal assemblages from archaeological sitesJ. Archaeol. Sci., 32 (2005), pp. 1202-1222, 10.1016/j.jas.2005.03.004Article Download PDF View Record in Scopus Google Scholar

C. Kiffner, H. Rheault, E. Miller, T. Scheetz, V. Enriquez, R. Swafford, J. Kioko, H.H.T. PrinsLongterm population dynamics in a multi-species assemblage of large herbivores in East AfricaEcosphere, 8 (2017), Article e02027, 10.1002/ecs2.2027CrossRef Google Scholar

R.G. Klein, K. Cruz-UribeThe Analysis of Animal Bones from Archeological SitesUniversity of Chicago Press, Chicago, Illinois (1984), p. 273View Record in Scopus Google Scholar

H. Koike, N. OhtaishiPrehistoric hunting pressure estimated by the age composition of excavated Sika deer (Cervusnippon) using the annual layer of tooth cementJ. Archaeol. Sci., 12 (1985), pp. 443-456Article Download PDF View Record in Scopus Google Scholar

H. Koike, N. OhtaishiEstimation of prehistoric hunting rates based on the age composition of (Cervus nippon)J. Archaeol. Sci., 14 (1987), pp. 251-269Article Download PDF View Record in Scopus Google Scholar

W.W. Korth, R.L. EvanderThe use of age-frequency profiles of micromammals in the determination of attritional andcatastrophic mortality of fossil assemblagesPalaeogeogr. Palaeoclimatol. Palaeoecol., 52 (1986), pp. 227-236, 10.1016/0031-0182(86)90048-9Article Download PDF View Record in Scopus Google Scholar

B. KurténOn the variation and population dynamics of fossil and recent mammal populationsActa Zool. Fennica, 76 (1953), pp. 1-112View Record in Scopus Google Scholar

B. KurténPopulation dynamics and evolutionEvolution, 8 (1954), pp. 75-81, 10.1111/j.1558-5646.1954.tb00110.xCrossRef Google Scholar

B. KurténPopulation dynamics: a new method in paleontology

Download

https://www.bipm.org/fr/publications/guides/

https://scholar.google.com/scholar_lookup?title=Evaluation%20of%20measurement%20data%20-%20guide%20to%20the%20expression%20of%20uncertainty%20in%20measurement&publication_year=2008&author=JCGM


https://scholar.google.com/scholar_lookup?title=Composition%20and%20limiting%20factors%20of%20a%20Khoma%20Hochland%20population%20of%20Hartmann%20zebra%20Equus%20zebra%20hartmannae&publication_year=1974&author=E.%20Joubert



https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/S0305440305000634/pdfft?md5=a014776668c5187fa78ca47b74d99452&pid=1-s2.0-S0305440305000634-main.pdf


https://scholar.google.com/scholar?q=The%20blessing%20of%20a%20great%20flood:%20differentiation%20of%20mortality%20patterns%20in%20the%20large%20mammal%20record%20of%20the%20Lower%20Pleistocene%20fluvial%20site%20of%20Untermassfeld%20%20and%20its%20relevance%20for%20the%20interpretation%20of%20faunal%20assemblages%20from%20archaeological%20sites

https://doi.org/10.1002/ecs2.2027

https://doi.org/10.1002/ecs2.2027

https://scholar.google.com/scholar_lookup?title=Longterm%20population%20dynamics%20in%20a%20multi-species%20assemblage%20of%20large%20herbivores%20in%20East%20Africa&publication_year=2017&author=C.%20Kiffner&author=H.%20Rheault&author=E.%20Miller&author=T.%20Scheetz&author=V.%20Enriquez&author=R.%20Swafford&author=J.%20Kioko&author=H.H.T.%20Prins


https://scholar.google.com/scholar_lookup?title=The%20Analysis%20of%20Animal%20Bones%20from%20Archeological%20Sites&publication_year=1984&author=R.G.%20Klein&author=K.%20Cruz-Uribe


https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/0305440385900044/pdf?md5=8c1481f8dd638f028cba0a7c7f4c6673&pid=1-s2.0-0305440385900044-main.pdf


https://scholar.google.com/scholar?q=Prehistoric%20hunting%20pressure%20estimated%20by%20the%20age%20composition%20of%20excavated%20Sika%20deer%20%20using%20the%20annual%20layer%20of%20tooth%20cement


https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/0305440387900148/pdf?md5=e0ba0d0b534a964eb04b4c230a0099b0&pid=1-s2.0-0305440387900148-main.pdf


https://scholar.google.com/scholar?q=Estimation%20of%20prehistoric%20hunting%20rates%20based%20on%20the%20age%20composition%20of

https://doi.org/10.1016/0031-0182(86)90048-9


https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/0031018286900489/pdf?md5=8b36d0f77710746a60c07edb13677a42&pid=1-s2.0-0031018286900489-main.pdf


https://scholar.google.com/scholar_lookup?title=The%20use%20of%20age-frequency%20profiles%20of%20micromammals%20in%20the%20determination%20of%20attritional%20and%20catastrophic%20mortality%20of%20fossil%20assemblages&publication_year=1986&author=W.W.%20Korth&author=R.L.%20Evander


https://scholar.google.com/scholar_lookup?title=On%20the%20variation%20and%20population%20dynamics%20of%20fossil%20and%20recent%20mammal%20populations&publication_year=1953&author=B.%20Kurt%C3%A9n

https://doi.org/10.1111/j.1558-5646.1954.tb00110.x

https://doi.org/10.1111/j.1558-5646.1954.tb00110.x

https://scholar.google.com/scholar_lookup?title=Population%20dynamics%20and%20evolution&publication_year=1954&author=B.%20Kurt%C3%A9n




Kurtén, 1983

Lemos-Espinal et al., 2005

Leslie, 1945

Leslie, 1948

Leslie, 1966

Levine, 1983

Lewis, 1942

Lindström and Kokko, 1998

Lkhagvasuren et al., 2017

Lyman, 1987

Lyman, 1994

J. Paleontol., 28 (1954), pp. 286-292View Record in Scopus Google Scholar

B. KurténVariation and dynamics of a fossil antelope populationPaleobiology, 9 (1983), pp. 62-69View Record in Scopus Google Scholar

J.A. Lemos-Espinal, R.I. Rojas-González, J.J. Zúñiga-VegaTécnicas para el Estudio de Poblaciones de Fauna SilvestreCONABIO, Universidad Nacional Autónoma de México (2005), p. 157ppGoogle Scholar

P.H. LeslieOn the use of matrices in certain population mathematicsBiometrika, 33 (1945), pp. 183-212, 10.1093/biomet/33.3.183CrossRef View Record in Scopus Google Scholar

P.H. LeslieSome further notes on the use of matrices in population mathematicsBiometrika, 35 (1948), pp. 213-245, 10.2307/2332342CrossRef Google Scholar

P.H. LeslieThe intrinsic rate of increase and the overlap of successive generations in a population ofGuillemot (Uria aalge Pont.)J. Anim. Ecol., 35 (1966), pp. 291-301https//doi.org/10.2307/2396CrossRef View Record in Scopus Google Scholar

M.A. LevineMortality models and the interpretation of horse population structureG. Bailey (Ed.), Hunter–Gatherer Economy in Prehistory: A European Perspective, Cambridge University Press,Cambridge (1983), pp. 23-46View Record in Scopus Google Scholar

E.G. LewisOn the generation and growth of a populationSankhya, 6 (1942), pp. 93-96View Record in Scopus Google Scholar

J. Lindström, H. KokkoSexual reproduction and population dynamics: the role of polygyny and demographic sexdifferencesProc. Roy. Soc. Lond. B, 265 (1998), pp. 438-488, 10.1098/rspb.1998.0320Google Scholar

D. Lkhagvasuren, N. Batsaikhan, W.F. Fagan, E.C. Ghandakly, P. Kaczensky, T. Müller, R.Samiya, R. Schafberg, A. Stubbe, M. Stubbe, H. AnsorgeFirst assessment of the population structure of the Asiatic wild ass in MongoliaEur. J. Wildl. Res., 64 (2017), p. 3, 10.1007/s10344-017-1162-xGoogle Scholar

R.L. LymanOn the analysis of vertebrate mortality profiles: sample Size, mortality type, and hunting pressureAm. Antiq., 52 (1987), pp. 125-142, 10.2307/281064CrossRef View Record in Scopus Google Scholar

R.L. Lyman

Download


https://scholar.google.com/scholar_lookup?title=Population%20dynamics%3A%20a%20new%20method%20in%20paleontology&publication_year=1954&author=B.%20Kurt%C3%A9n


https://scholar.google.com/scholar_lookup?title=Variation%20and%20dynamics%20of%20a%20fossil%20antelope%20population&publication_year=1983&author=B.%20Kurt%C3%A9n

https://scholar.google.com/scholar_lookup?title=T%C3%A9cnicas%20para%20el%20Estudio%20de%20Poblaciones%20de%20Fauna%20Silvestre&publication_year=2005&author=J.A.%20Lemos-Espinal&author=R.I.%20Rojas-Gonz%C3%A1lez&author=J.J.%20Z%C3%BA%C3%B1iga-Vega

https://doi.org/10.1093/biomet/33.3.183

https://doi.org/10.1093/biomet/33.3.183


https://scholar.google.com/scholar_lookup?title=On%20the%20use%20of%20matrices%20in%20certain%20population%20mathematics&publication_year=1945&author=P.H.%20Leslie

https://doi.org/10.2307/2332342

https://doi.org/10.1093/biomet/35.3-4.213

https://scholar.google.com/scholar_lookup?title=Some%20further%20notes%20on%20the%20use%20of%20matrices%20in%20population%20mathematics&publication_year=1948&author=P.H.%20Leslie

https://doi.org/10.2307/2396


https://scholar.google.com/scholar?q=The%20intrinsic%20rate%20of%20increase%20and%20the%20overlap%20of%20successive%20generations%20in%20a%20population%20of%20Guillemot


https://scholar.google.com/scholar_lookup?title=Mortality%20models%20and%20the%20interpretation%20of%20horse%20population%20structure&publication_year=1983&author=M.A.%20Levine


https://scholar.google.com/scholar_lookup?title=On%20the%20generation%20and%20growth%20of%20a%20population&publication_year=1942&author=E.G.%20Lewis

https://doi.org/10.1098/rspb.1998.0320

https://scholar.google.com/scholar_lookup?title=Sexual%20reproduction%20and%20population%20dynamics%3A%20the%20role%20of%20polygyny%20and%20demographic%20sex%20differences&publication_year=1998&author=J.%20Lindstr%C3%B6m&author=H.%20Kokko

https://doi.org/10.1007/s10344-017-1162-x

https://scholar.google.com/scholar_lookup?title=First%20assessment%20of%20the%20population%20structure%20of%20the%20Asiatic%20wild%20ass%20in%20Mongolia&publication_year=2017&author=D.%20Lkhagvasuren&author=N.%20Batsaikhan&author=W.F.%20Fagan&author=E.C.%20Ghandakly&author=P.%20Kaczensky&author=T.%20M%C3%BCller&author=R.%20Samiya&author=R.%20Schafberg&author=A.%20Stubbe&author=M.%20Stubbe&author=H.%20Ansorge

https://doi.org/10.2307/281064

https://doi.org/10.2307/281064


https://scholar.google.com/scholar_lookup?title=On%20the%20analysis%20of%20vertebrate%20mortality%20profiles%3A%20sample%20Size%2C%20mortality%20type%2C%20and%20hunting%20pressure&publication_year=1987&author=R.L.%20Lyman




Lyman, 2008

MacFadden, 1989

MacFadden, 2008

Marín et al., 2017

Martín-González et al., 2016

McDonald, 1996

Mihlbachler, 2003

Mihlbachler, 2007

Mihlbachler, 2012

Vertebrate TaphonomyCambridge University Press, Cambridge (1994), p. 524View Record in Scopus Google Scholar

R.L. LymanQuantitative PaleozoologyCambridge University Press, Cambridge (2008), p. 348Google Scholar

B.J. MacFaddenDental character variation in paleopopulations and morphospecies of fossil horses and extantanalogsD.R. Prothero, R.M. Schoch (Eds.), The Evolution of Perissodactyls, Oxford University Press, New York (1989), pp.128-141Google Scholar

B.J. MacFaddenGeographic variation in diets of ancient populations of 5-million-year-old (early Pliocene) horsesfrom southern North AmericaPalaeogeogr. Palaeoclimatol. Palaeoecol., 226 (2008), pp. 83-94, 10.1016/j.palaeo.2008.03.019Article Download PDF View Record in Scopus Google Scholar

J. Marín, P. Saladié, A. Rodríguez-Hidalgo, E. CarbonellNeanderthal hunting strategies inferred from mortality profiles within the Abric Romaní sequencePloS One, 12 (2017), Article e0186970, 10.1371/journal.pone.0186970CrossRef Google Scholar

J.A. Martín-González, A. Mateos, G. Rodríguez-GómezA parametrical model to describe a stable and stationary age structure for fossil populationsQuat. Int., 413B (2016), pp. 69-77, 10.1016/j.quaint.2016.01.038Article Download PDF View Record in Scopus Google Scholar

H.G. McDonaldPopulation structure of the late pliocene (blancan) zebra Equus simplicidens (perissodactyla:equidae) from the hagerman horse quarry, IdahoK.M. Stewart, K.L. Seymour (Eds.), Palaeoecology and Palaeoenvironments of Late Cenozoic Mammals, Universityof Toronto, Canada (1996), pp. 134-155CrossRef Google Scholar

M.C. MihlbachlerDemography of late Miocene rhinoceroses (Teleoceras proterum and Aphelops malacorhinus)from Florida: linking mortality and sociality in fossil assemblagesPaleobiology, 29 (2003), pp. 412-428https://doi:10.1666/0094-8373(2003)0292.0.COView Record in Scopus Google Scholar

M.C. MihlbachlerSexual dimorphism and mortality bias in a small Miocene North American rhino, Menocerasarikarense: insights into the coevolution of sexual dimorphism and sociality in rhinosJ. Mamm. Evol., 14 (2007), pp. 217-238https://doi:10.1007/s10914-007-9048-4CrossRef View Record in Scopus Google Scholar

M.C. MihlbachlerPalaeodemographics of small-bodied mammals in Pleistocene environments: a case study ofmuskrats (Ondatra zibethicus: rodentia: Muridae) from north FloridaBiol. J. Linn. Soc., 106 (2012), pp. 41-56CrossRef View Record in Scopus Google Scholar

Download


https://scholar.google.com/scholar_lookup?title=Vertebrate%20Taphonomy&publication_year=1994&author=R.L.%20Lyman

https://scholar.google.com/scholar_lookup?title=Quantitative%20Paleozoology&publication_year=2008&author=R.L.%20Lyman

https://scholar.google.com/scholar_lookup?title=Dental%20character%20variation%20in%20paleopopulations%20and%20morphospecies%20of%20fossil%20horses%20and%20extant%20analogs&publication_year=1989&author=B.J.%20MacFadden

https://doi.org/10.1016/j.palaeo.2008.03.019


https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/S0031018208002319/pdfft?md5=f6ffcfa6cc8b1989c25e6aabf30a05a7&pid=1-s2.0-S0031018208002319-main.pdf


https://scholar.google.com/scholar?q=Geographic%20variation%20in%20diets%20of%20ancient%20populations%20of%205-million-year-old%20%20horses%20from%20southern%20North%20America

https://doi.org/10.1371/journal.pone.0186970


https://scholar.google.com/scholar_lookup?title=Neanderthal%20hunting%20strategies%20inferred%20from%20mortality%20profiles%20within%20the%20Abric%20Roman%C3%AD%20sequence&publication_year=2017&author=J.%20Mar%C3%ADn&author=P.%20Saladi%C3%A9&author=A.%20Rodr%C3%ADguez-Hidalgo&author=E.%20Carbonell



https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/S1040618216000707/pdfft?md5=c114480ea4d3ba75e4ca3ba9eb77f8ff&pid=1-s2.0-S1040618216000707-main.pdf


https://scholar.google.com/scholar_lookup?title=A%20parametrical%20model%20to%20describe%20a%20stable%20and%20stationary%20age%20structure%20for%20fossil%20populations&publication_year=2016&author=J.A.%20Mart%C3%ADn-Gonz%C3%A1lez&author=A.%20Mateos&author=G.%20Rodr%C3%ADguez-G%C3%B3mez

https://doi.org/10.3138/9781487574154-010

https://scholar.google.com/scholar?q=Population%20structure%20of%20the%20late%20pliocene%20%20zebra%20Equus%20simplicidens%20%20from%20the%20hagerman%20horse%20quarry,%20Idaho


https://scholar.google.com/scholar?q=Demography%20of%20late%20Miocene%20rhinoceroses%20%20from%20Florida:%20linking%20mortality%20and%20sociality%20in%20fossil%20assemblages

https://doi.org/10.1007/s10914-007-9048-4


https://scholar.google.com/scholar_lookup?title=Sexual%20dimorphism%20and%20mortality%20bias%20in%20a%20small%20Miocene%20North%20American%20rhino%2C%20Menoceras%20arikarense%3A%20insights%20into%20the%20coevolution%20of%20sexual%20dimorphism%20and%20sociality%20in%20rhinos&publication_year=2007&author=M.C.%20Mihlbachler

https://doi.org/10.1111/j.1095-8312.2011.01849.x


https://scholar.google.com/scholar?q=Palaeodemographics%20of%20small-bodied%20mammals%20in%20Pleistocene%20environments:%20a%20case%20study%20of%20muskrats%20%20from%20north%20Florida




Monchot et al., 2012

Munson, 2000

Munson and Marean, 2003

Munson and Garniewicz, 2003

Nimmo, 1971

Oli, 2004

Owen-Smith, 2008

Pearl and Miner, 1935

Pianka, 1970

Pineda-Maldonado, 2017

Prentiss et al., 2014

H. Monchot, P. Fernandez, J.-M. GaillardPaleodemographic analysis of a fossil porcupine (Hystrix refossa gervais, 1852) population fromthe upper Pleistocene site of geula cave (mount carmel, Israel)J. Archaeol. Sci., 39 (2012), pp. 3027-3038Article Download PDF View Record in Scopus Google Scholar

P.J. MunsonAge-correlated differential destruction of bones and its effects on archaeological mortality profileof domestic sheep and goatsJ. Archaeol. Sci., 27 (2000), pp. 391-407, 10.1006/jasc.1999.0463Article Download PDF View Record in Scopus Google Scholar

P.J. Munson, C.W. MareanAdults only? A reconsideration of Middle Paleolithic 'prime-dominated' reindeer hunting atSalzgitter LebenstedetJ. Hum. Evol., 44 (2003), pp. 263-273, 10.1016/S0047-2484(02)00163-XView Record in Scopus Google Scholar

P.J. Munson, R.C. GarniewiczAge-mediated survivorship of ungulate mandibles and teeth in canid-ravaged faunal assemblagesJ. Archaeol. Sci., 30 (2003), pp. 405-416, 10.1006/jasc.2002.0850Article Download PDF View Record in Scopus Google Scholar

B.W. NimmoPopulation dynamics of a Wyoming pronghorn cohort from the Eden-Farson site, 48SW304Plains Anthropol., 16 (1971), pp. 285-288View Record in Scopus Google Scholar

M.K. OliThe fast–slow continuum and mammalian life-history patterns: an empirical evaluationBasic Appl. Ecol., 5 (2004), pp. 449-463, 10.1016/j.baae.2004.06.002Article Download PDF CrossRef View Record in Scopus Google Scholar

N. Owen-SmithThe comparative population dynamics of browsing and grazing ungulatesI.J. Gordon, H.T. Prins (Eds.), The Ecology of Browsing and Grazing, Springer-Verlag, Berlin (2008), pp. 149-178CrossRef View Record in Scopus Google Scholar

R. Pearl, J.R. MinerExperimental studies on the duration of life. XIV. The comparative mortality of certain lowerorganismsQRB (Q. Rev. Biol.), 10 (1935), pp. 60-79View Record in Scopus Google Scholar

E.R. PiankaOn r- and K-SelectionAm. Nat., 104 (1970), pp. 592-597, 10.1086/282697View Record in Scopus Google Scholar

M.A. Pineda-MaldonadoInterpretación paleoambiental de un ecosistema del Pleistoceno tardío en Epazoyucan, sureste deHidalgo mediante palinología y un análisis ecomorfológicoMSc. Dissertation. Universidad Autónoma del Estado de Hidalgo, Mexico (2017)Google Scholar

A.M. Prentiss, H.S. Cail, L.M. SmithAt the malthusian ceiling: subsistence and inequality at bridge river, British columbiaJ. Anthropol. Archaeol., 33 (2014), pp. 34-48, 10.1016/j.jaa.2013.11.003Article Download PDF View Record in Scopus Google Scholar

Download


https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/S0305440312001707/pdfft?md5=738b718e4a9fef014a9897228fbb2289&pid=1-s2.0-S0305440312001707-main.pdf


https://scholar.google.com/scholar?q=Paleodemographic%20analysis%20of%20a%20fossil%20porcupine%20%20population%20from%20the%20upper%20Pleistocene%20site%20of%20geula%20cave

https://doi.org/10.1006/jasc.1999.0463


https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/S0305440399904636/pdf?md5=186f4b51794d42cd4c988215cfac26fa&pid=1-s2.0-S0305440399904636-main.pdf


https://scholar.google.com/scholar_lookup?title=Age-correlated%20differential%20destruction%20of%20bones%20and%20its%20effects%20on%20archaeological%20mortality%20profile%20of%20domestic%20sheep%20and%20goats&publication_year=2000&author=P.J.%20Munson

https://doi.org/10.1016/S0047-2484(02)00163-X


https://scholar.google.com/scholar?q=Adults%20only%20A%20reconsideration%20of%20Middle%20Paleolithic%20prime-dominated%20reindeer%20hunting%20at%20Salzgitter%20Lebenstedet



https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/S0305440302908502/pdf?md5=7920df7f672b634e92b332ccd092f71f&pid=1-s2.0-S0305440302908502-main.pdf


https://scholar.google.com/scholar_lookup?title=Age-mediated%20survivorship%20of%20ungulate%20mandibles%20and%20teeth%20in%20canid-ravaged%20faunal%20assemblages&publication_year=2003&author=P.J.%20Munson&author=R.C.%20Garniewicz


https://scholar.google.com/scholar_lookup?title=Population%20dynamics%20of%20a%20Wyoming%20pronghorn%20cohort%20from%20the%20Eden-Farson%20site%2C%2048SW304&publication_year=1971&author=B.W.%20Nimmo

https://doi.org/10.1016/j.baae.2004.06.002


https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/S1439179104000398/pdfft?md5=98638cce57b9ed927fdc1b5b1adc4888&pid=1-s2.0-S1439179104000398-main.pdf

https://doi.org/10.1016/j.baae.2004.06.002


https://scholar.google.com/scholar?q=The%20fastslow%20continuum%20and%20mammalian%20life-history%20patterns:%20an%20empirical%20evaluation

https://doi.org/10.1007/978-3-540-72422-3_6


https://scholar.google.com/scholar_lookup?title=The%20comparative%20population%20dynamics%20of%20browsing%20and%20grazing%20ungulates&publication_year=2008&author=N.%20Owen-Smith


https://scholar.google.com/scholar_lookup?title=Experimental%20studies%20on%20the%20duration%20of%20life.%20XIV.%20The%20comparative%20mortality%20of%20certain%20lower%20organisms&publication_year=1935&author=R.%20Pearl&author=J.R.%20Miner

https://doi.org/10.1086/282697


https://scholar.google.com/scholar_lookup?title=On%20r-%20and%20K-Selection&publication_year=1970&author=E.R.%20Pianka

https://scholar.google.com/scholar_lookup?title=Interpretaci%C3%B3n%20paleoambiental%20de%20un%20ecosistema%20del%20Pleistoceno%20tard%C3%ADo%20en%20Epazoyucan%2C%20sureste%20de%20Hidalgo%20mediante%20palinolog%C3%ADa%20y%20un%20an%C3%A1lisis%20ecomorfol%C3%B3gico&publication_year=2017&author=M.A.%20Pineda-Maldonado

https://doi.org/10.1016/j.jaa.2013.11.003


https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/S0278416513000597/pdfft?md5=b1c9b0aa4e3af9f19547b7e47a5d9faf&pid=1-s2.0-S0278416513000597-main.pdf


https://scholar.google.com/scholar_lookup?title=At%20the%20malthusian%20ceiling%3A%20subsistence%20and%20inequality%20at%20bridge%20river%2C%20British%20columbia&publication_year=2014&author=A.M.%20Prentiss&author=H.S.%20Cail&author=L.M.%20Smith




Ransom et al., 2016

Reher, 1970

Reher, 1973

Reher, 1974

Rodríguez-Gómez et al., 2014

Rodríguez-Gómez et al., 2016

RStudio Team, 2015

Rubenstein, 2010

Rubenstein and Nuñez, 2009

Ruffner and Carothers, 1982

J.I. Ransom, L. Lagos, H. Hrabar, H. Nowzari, D. Usukhjargal, N. SpasskayaWild and feral equid population dynamics

J.I. Ransom, P. Kaczensky (Eds.), Wild Equids. Ecology, Management, and Conservation, Johns HopkinsUniversity Press, Baltimore, Maryland (2016), pp. 68-83View Record in Scopus Google Scholar

C.A. ReherPopulation dynamics of the Glenrock Bison bison populationG.C. Frison (Ed.), The Glenrock Buffalo Jump, 48CO34: Late Prehistoric Period Buffalo Procurement andButchering on the Northwestern Plains, vol. 7, Plains Anthropologist Memoir (1970), pp. 51-55View Record in Scopus Google Scholar

C.A. ReherThe Wardell Bison bison sample: population dynamics and archaeological InterpretationG.C. Frison (Ed.), The Wardell Buffalo Trap, 48SH301: Communal Procurement in the Upper Green River Basin,Wyoming, University of Michigan Anthropology Papers No. 48, Ann Arbor (1973), pp. 89-105View Record in Scopus Google Scholar

C.A. ReherPopulation study of the Casper site BisonG.C. Frison (Ed.), The Casper Site: A Hell Gap Bison Kill on the High Plains, Academic Press, New York (1974), pp.113-124Google Scholar

G. Rodríguez-Gómez, A. Mateos, J.A. Martín-González, R. Blasco, J. Rosell, J. RodríguezDiscontinuity of human presence at atapuerca during the early middle Pleistocene: a matter ofecological competition?PloS One, 9 (2014), Article e101938, 10.1371/journal.pone.0101938CrossRef View Record in Scopus Google Scholar

G. Rodríguez-Gómez, A. Mateos, J.A. Martín-González, J. RodríguezMeasuring intraguild competition from faunal assemblages to compare environmental conditionsamong paleocommunitiesQuat. Int., 413B (2016), pp. 55-68, 10.1016/j.quaint.2015.11.087Article Download PDF View Record in Scopus Google Scholar

RStudio TeamRStudio: integrated development for R. RStudio, PBC, boston, MassachusettsAvailable online at:http://www.rstudio.com/ (2015)Google Scholar

D.I. RubensteinChapter 7 - ecology, social behavior, and conservation in zebrasAdv. Stud. Behav., 42 (2010), pp. 231-258, 10.1016/S0065-3454(10)42007-0Article Download PDF View Record in Scopus Google Scholar

D.I. Rubenstein, C.M. NuñezSociality and reproductive skew in horses and zebrasR. Hager, C.B. Jones (Eds.), Reproductive Skew in Vertebrates-Proximate and Ultimate Causes, CambridgeUniversity Press, Cambridge (2009), pp. 196-226CrossRef View Record in Scopus Google Scholar

G.A. Ruffner, S.W. CarothersAge structure, condition and reproduction of two Equus asinus (equidae) populations from grandCanyon national park, ArizonaSW. Nat., 27 (1982), pp. 403-411, 10.2307/3670715CrossRef Google Scholar

Download


https://scholar.google.com/scholar_lookup?title=Wild%20and%20feral%20equid%20population%20dynamics&publication_year=2016&author=J.I.%20Ransom&author=L.%20Lagos&author=H.%20Hrabar&author=H.%20Nowzari&author=D.%20Usukhjargal&author=N.%20Spasskaya


https://scholar.google.com/scholar_lookup?title=Population%20dynamics%20of%20the%20Glenrock%20Bison%20bison%20population&publication_year=1970&author=C.A.%20Reher


https://scholar.google.com/scholar_lookup?title=The%20Wardell%20Bison%20bison%20sample%3A%20population%20dynamics%20and%20archaeological%20Interpretation&publication_year=1973&author=C.A.%20Reher

https://scholar.google.com/scholar_lookup?title=Population%20study%20of%20the%20Casper%20site%20Bison&publication_year=1974&author=C.A.%20Reher




https://scholar.google.com/scholar?q=Discontinuity%20of%20human%20presence%20at%20atapuerca%20during%20the%20early%20middle%20Pleistocene:%20a%20matter%20of%20ecological%20competition



https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/S1040618215012136/pdfft?md5=7153af67b33b3711266dd572b8b79f7b&pid=1-s2.0-S1040618215012136-main.pdf


https://scholar.google.com/scholar_lookup?title=Measuring%20intraguild%20competition%20from%20faunal%20assemblages%20to%20compare%20environmental%20conditions%20among%20paleocommunities&publication_year=2016&author=G.%20Rodr%C3%ADguez-G%C3%B3mez&author=A.%20Mateos&author=J.A.%20Mart%C3%ADn-Gonz%C3%A1lez&author=J.%20Rodr%C3%ADguez

http://www.rstudio.com/

https://scholar.google.com/scholar_lookup?title=RStudio%3A%20integrated%20development%20for%20R.%20RStudio%2C%20PBC%2C%20boston%2C%20Massachusetts&publication_year=2015&author=RStudio%20Team

https://doi.org/10.1016/S0065-3454(10)42007-0


https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/S0065345410420070/pdfft?md5=65dc7bd6020ca847d66af5ddf85721b6&pid=1-s2.0-S0065345410420070-main.pdf


https://scholar.google.com/scholar_lookup?title=Chapter%207%20-%20ecology%2C%20social%20behavior%2C%20and%20conservation%20in%20zebras&publication_year=2010&author=D.I.%20Rubenstein

https://doi.org/10.1017/CBO9780511641954.010


https://scholar.google.com/scholar_lookup?title=Sociality%20and%20reproductive%20skew%20in%20horses%20and%20zebras&publication_year=2009&author=D.I.%20Rubenstein&author=C.M.%20Nu%C3%B1ez

https://doi.org/10.2307/3670715

https://doi.org/10.2307/3670715

https://scholar.google.com/scholar?q=Age%20structure,%20condition%20and%20reproduction%20of%20two%20Equus%20asinus%20%20populations%20from%20grand%20Canyon%20national%20park,%20Arizona




Smuts, 1976

Spinage, 1972

Stearns, 1992

Steele, 2005

Steele and Weaver, 2002

Steinsalz and Orzack, 2011

Stephens and Krebs, 1986

Stiner, 1990

Stiner, 1991

Stiner, 1994

Stott, 2018

G.L. SmutsPopulation characteristics of burchell's zebra (Equus burchelli antiquorum. H. Smith, 1841) in thekruger national parkS. Afr. J. Wildl. Res., 6 (1976), pp. 99-112Google Scholar

C.A. SpinageAfrican ungulate life tablesEcology, 53 (1972), pp. 645-652, 10.2307/1934778CrossRef View Record in Scopus Google Scholar

S.C. StearnsThe Evolution of Life HistoriesOxford University Press, Oxford, United Kingdom (1992), p. 249View Record in Scopus Google Scholar

T.E. SteeleComparing methods for analysing mortality profiles in zooarchaeological and paleontologicalsamplesInt. J. Osteoarchaeol., 15 (2005), pp. 1-17View Record in Scopus Google Scholar

T.E. Steele, T.D. WeaverThe modified triangular graph: a refined method for comparing mortality profiles inarchaeological samplesJ. Archaeol. Sci., 29 (2002), pp. 317-322, 10.1006/jasc.2001.0733Article Download PDF View Record in Scopus Google Scholar

D. Steinsalz, S.H. OrzackStatistical methods for paleodemography on fossil assemblages having small numbers ofspecimens: an investigation of dinosaur survival ratesPaleobiology, 37 (2011), pp. 113-125Google Scholar

D.W. Stephens, J.R. KrebsForaging TheoryPrinceton University Press, Princeton (1986), p. 247View Record in Scopus Google Scholar

M.C. StinerThe use of mortality patterns in archaeological studies of hominid predatory adaptationsJ. Anthropol. Archaeol., 9 (1990), pp. 305-351, 10.1016/0278-4165(90)90010-BArticle Download PDF View Record in Scopus Google Scholar

M.C. StinerHuman Predator and Prey MortalityWestview Press, Boulder (1991), p. 276Google Scholar

M.C. StinerHonor Among Thieves: a Zooarchaeological Study of Neandertal EcologyPrinceton University Press, Princeton (1994), p. 447Google Scholar

I. StottPopdemo: demographic modelling using projection matricesAvailable online at:https://cran.r-project.org/web/packages/popdemo/vignettes/popdemo.html (2018)Google Scholar

Download

https://scholar.google.com/scholar?q=Population%20characteristics%20of%20burchells%20zebra%20%20in%20the%20kruger%20national%20park

https://doi.org/10.2307/1934778

https://doi.org/10.2307/1934778


https://scholar.google.com/scholar_lookup?title=African%20ungulate%20life%20tables&publication_year=1972&author=C.A.%20Spinage


https://scholar.google.com/scholar_lookup?title=The%20Evolution%20of%20Life%20Histories&publication_year=1992&author=S.C.%20Stearns


https://scholar.google.com/scholar_lookup?title=Comparing%20methods%20for%20analysing%20mortality%20profiles%20in%20zooarchaeological%20and%20paleontological%20samples&publication_year=2005&author=T.E.%20Steele



https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/S0305440301907332/pdf?md5=2dd08008c10b5c91655a8338e203550a&pid=1-s2.0-S0305440301907332-main.pdf


https://scholar.google.com/scholar_lookup?title=The%20modified%20triangular%20graph%3A%20a%20refined%20method%20for%20comparing%20mortality%20profiles%20in%20archaeological%20samples&publication_year=2002&author=T.E.%20Steele&author=T.D.%20Weaver

https://scholar.google.com/scholar_lookup?title=Statistical%20methods%20for%20paleodemography%20on%20fossil%20assemblages%20having%20small%20numbers%20of%20specimens%3A%20an%20investigation%20of%20dinosaur%20survival%20rates&publication_year=2011&author=D.%20Steinsalz&author=S.H.%20Orzack


https://scholar.google.com/scholar_lookup?title=Foraging%20Theory&publication_year=1986&author=D.W.%20Stephens&author=J.R.%20Krebs

https://doi.org/10.1016/0278-4165(90)90010-B

https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/027841659090010B

https://www-sciencedirect-com.lama.univ-amu.fr/science/article/pii/027841659090010B/pdf?md5=90ebb18c364a9303e331c781c666dbad&pid=1-s2.0-027841659090010B-main.pdf


https://scholar.google.com/scholar_lookup?title=The%20use%20of%20mortality%20patterns%20in%20archaeological%20studies%20of%20hominid%20predatory%20adaptations&publication_year=1990&author=M.C.%20Stiner

https://scholar.google.com/scholar_lookup?title=Human%20Predator%20and%20Prey%20Mortality&publication_year=1991&author=M.C.%20Stiner

https://scholar.google.com/scholar_lookup?title=Honor%20Among%20Thieves%3A%20a%20Zooarchaeological%20Study%20of%20Neandertal%20Ecology&publication_year=1994&author=M.C.%20Stiner

https://cran.r-project.org/web/packages/popdemo/vignettes/popdemo.html

https://scholar.google.com/scholar_lookup?title=Popdemo%3A%20demographic%20modelling%20using%20projection%20matrices&publication_year=2018&author=I.%20Stott




Turnbull and Martill, 1988

Van Valen, 1963

Van Valen, 1964

Van Valen, 1965

Vasileiadou et al., 2007

Voorhies, 1969

W.D. Turnbull, D.M. MartillTaphonomy and preservation of a monospecific titanothere assemblage from

the Washakie formation (Late Eocene), Southern Wyoming: an ecological accident in the fossilrecordPalaeogeogr. Palaeoclimatol. Palaeoecol., 63 (1988), pp. 91-108Article Download PDF View Record in Scopus Google Scholar

L. Van ValenSelection in natural populations: Merychippus primus, a fossil horseNature, 197 (1963), pp. 1181-1183CrossRef View Record in Scopus Google Scholar

L. Van ValenAge in two fossil horse populationsActa Zool., 45 (1964), pp. 93-110View Record in Scopus Google Scholar

L. Van ValenSelection in natural populations. III. Measurement and estimationEvolution, 19 (1965), pp. 514-528View Record in Scopus Google Scholar

K. Vasileiadou, J.J. Hooker, M.E. CollinsonQuantification and age structure of semi-hypsodont extinct rodent populationsJournal of Taphonomy, 5 (2007), pp. 15-41View Record in Scopus Google Scholar

M.R. VoorhiesTaphonomy and Population Dynamics of an Early Pliocene Vertebrate Fauna, Knox County,NebraskaUniversity of Wyoming, Contributions of Geology Special Paper (1969), pp. 1-69CrossRef View Record in Scopus Google Scholar

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https://scholar.google.com/scholar?q=Taphonomy%20and%20preservation%20of%20a%20monospecific%20titanothere%20assemblage%20from%20the%20Washakie%20formation%20,%20Southern%20Wyoming:%20an%20ecological%20accident%20in%20the%20fossil%20record

https://doi.org/10.1038/1971181a0


https://scholar.google.com/scholar_lookup?title=Selection%20in%20natural%20populations%3A%20Merychippus%20primus%2C%20a%20fossil%20horse&publication_year=1963&author=L.%20Van%20Valen


https://scholar.google.com/scholar_lookup?title=Age%20in%20two%20fossil%20horse%20populations&publication_year=1964&author=L.%20Van%20Valen


https://scholar.google.com/scholar_lookup?title=Selection%20in%20natural%20populations.%20III.%20Measurement%20and%20estimation&publication_year=1965&author=L.%20Van%20Valen


https://scholar.google.com/scholar_lookup?title=Quantification%20and%20age%20structure%20of%20semi-hypsodont%20extinct%20rodent%20populations&publication_year=2007&author=K.%20Vasileiadou&author=J.J.%20Hooker&author=M.E.%20Collinson

https://doi.org/10.2113/gsrocky.8.special_paper_1.1


https://scholar.google.com/scholar_lookup?title=Taphonomy%20and%20Population%20Dynamics%20of%20an%20Early%20Pliocene%20Vertebrate%20Fauna%2C%20Knox%20County%2C%20Nebraska&publication_year=1969&author=M.R.%20Voorhies

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