Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e20264964, enero-diciembre 2026 (Publicado Oct. 09, 2026)

Comparative feeding strategies of two Neotropical cervids

(Artiodactyla: Cervidae) from Costa Rica based on dental wear proxies

María Fernanda Martínez-Polanco1,2,3* https://orcid.org/0000-0002-5540-8981

1. Departamento de Antropología, Facultad de Ciencias Humanas, Universidad Nacional de Colombia, Sede Bogotá,

Avenida Carrera 30 45-03, Ciudad Universitaria, Bogotá, Colombia; mfmartinezp@gmail.com

2. Institut Català de Paleoecologia Humana i Evolució Social (IPHES-CERCA), Universitat Rovira i Virgili (URV), Zona

Educacional 4, Campus Sescelades (Edifici W3), 43007 Tarragona, Tarragona, España.

3. Departament d’Història i Història de l’Art, Universitat Rovira i Virgili (URV), Avinguda de Catalunya 35, 43002

Tarragona, Tarragona, España.

Received 25-I-2026. Corrected 25-V-2026. Accepted 18-VIII-2026.

ABSTRACT

Introduction: Dental mesowear and microwear analyses provide complementary proxies for reconstructing

herbivore feeding ecology at different temporal scales. While mesowear reflects long-term dietary abrasion,

microwear captures short-term feeding behavior, allowing evaluation of dietary stability and variability in season-

ally dynamic tropical environments.

Objectives: To assess short- and long-term dietary patterns in two Neotropical cervids, the white-tailed deer

(Odocoileus virginianus) and the Central American red brocket deer (Mazama temama), and to evaluate the

ecological significance of microwear variability in relation to habitat, seasonality, and foraging behavior.

Methods: Dental mesowear and low magnification microwear analyses were conducted on lower molars of

modern cervid specimens from Costa Rica. Microwear variables, including pits, scratches, and scratch texture,

were recorded within standardized enamel areas. Scratch variability was assessed using standard deviation and

coefficient of variation, and results were interpreted through comparative and descriptive approaches.

Results: Odocoileus virginianus exhibited moderate mesowear values combined with low microwear dispersion,

indicating a stable feeding strategy. In contrast, M. temama showed greater variability in microwear features,

particularly in scratch counts, showing a broader diet.

Conclusions: Tropical cervids may maintain consistent long-term feeding strategies while exhibiting variable

short-term dietary responses driven by ecological context and resource availability. Integrating mesowear,

microwear, and microwear variability provides a robust framework for interpreting cervid feeding ecology

in both modern and archaeological contexts, with implications for future comparative studies across tropical

environments.

Key words: dental mesowear; dental microwear; Odocoileus virginianus; Mazama temama; feeding ecology.

RESUMEN

Estrategias tróficas de dos cérvidos neotropicales (Artiodactyla: Cervidae)

de Costa Rica a partir de indicadores de desgaste dental

Introducción: El mesodesgaste y microdesgaste dental son enfoques complementarios que permiten reconstruir

la ecología trófica de herbívoros en escalas temporales diferentes. El mesodesgaste refleja una temporalidad larga

mientras que el microdesgaste presenta el comportamiento alimenticio del animal antes de morir. Estos dos

enfoques permiten conocer la estabilidad o la variabilidad de la dieta de los animales en ambientes tropicales.

https://doi.org/110.15517/2qn2gs59

VERTEBRATE BIOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e20264964, enero-diciembre 2026 (Publicado Oct. 09, 2026)

INTRODUCTION

Understanding how closely related species

exploit food resources is central to the study

of herbivore ecology and niche differentiation.

In ungulates, dietary strategies are commonly

structured along a continuum ranging from

browsing to grazing, with intermediate mixed-

feeding behaviors reflecting varying degrees

of dietary flexibility (Hofmann & Stewart,

1972). Dental wear analyses have become a

widely used approach for reconstructing feed-

ing strategies in both extant and fossil her-

bivores, as they provide direct evidence of

food properties and ingestion-related abrasives

(Semprebon et al., 2016).

Dental micro- and mesowear proxies cap-

ture dietary signals at different temporal scales.

Dental microwear reflects short-term feeding

behavior by recording the physical proper-

ties of foods consumed during the days or

weeks prior to death, whereas dental mesowear

integrates longer-term dietary patterns over

months or years by documenting cumulative

tooth wear related to food abrasiveness and

toughness (Ackermans et al., 2020; DeSantis,

2016; Fortelius & Solounias, 2000; Solounias &

Semprebon, 2002). Mesowear analysis is based

on the macroscopic examination of molar

cusp morphology, particularly cusp relief and

occlusal sharpness, which reflect the balance

between attrition and abrasion during masti-

cation. Following the general framework pro-

posed by Hofmann and Stewart (1972), these

wear patterns allow herbivorous mammals to

be broadly classified into browsing, grazing,

or mixed-feeding dietary categories. Animals

dominated by attrition typically retain sharper

cusps, characteristic of browsers consuming

softer vegetation, whereas grazers feeding on

more abrasive resources tend to exhibit more

rounded or blunted cusps due to higher lev-

els of abrasion (Fortelius & Solounias, 2000).

The combined application of mesowear and

microwear proxies therefore provides a more

robust framework for reconstructing feeding

strategies by integrating both long-term dietary

trends and short-term feeding signals (Davis &

Pineda-Munoz, 2016).

In paleontological and archaeological

research, comparisons of dental wear patterns

in extant species with well-documented diets

with those in fossil or archaeological teeth

remains are widely used to infer the feeding

Objetivos: Estudiar los patrones dietarios a corto y largo plazo en dos especies de venados neotropicales: el

venado de cola blanca (Odocoileus virginianus) y el temazate rojo (Mazama temama) y evaluar el significado

ecológico de la variabilidad del microdesgaste en relación con el hábitat, la estacionalidad y los comportamientos

de forrajeo.

Métodos: El mesodesgaste y el microdesgaste dental de baja magnificación fueron realizados en molares actuales

de individuos de Costa Rica. Las variables del microdesgaste, incluyendo hoyos, estrías y texturas de estrías fue-

ron estudiadas en áreas estandarizadas del esmalte. La variabilidad de las estrías se obtuvo usando la desviación

estándar y los coeficientes de variación. Los resultados se interpretaron mediante un enfoque comparativo y

descriptivo.

Resultados: Odocoileus virginianus presenta valores de mesodesgaste moderados con una dispersión baja en las

variables del microdesgaste, indicando una estrategia de alimentación estable. En el caso de M. temama se evi-

dencia una mayor variabilidad en los rasgos de microdesgaste, particularmente en número de estrías, mostrando

una dieta más amplia.

Conclusiones: Los venados tropicales mantienen unas estrategias tróficas estables a largo plazo, aunque pueden

ser variables como una respuesta al contexto ecológico y a la disponibilidad de los recursos. Integrar el mesodes-

gaste, el microdesgaste y la variabilidad del microdesgaste provee un contexto potente para interpretar la ecología

trófica tanto en venados actuales como arqueológicos y paleontológicos, brindando una base para estudios futuros

en distintos ambientes tropicales.

Palabras clave: mesodesgaste dental; microdesgaste dental; Odocoileus virginianus; Mazama temama; ecología

trófica.

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habits of past mammal populations (Faith &

Lyman, 2019). For this reason, establishing reli-

able reference datasets based on modern taxa

is essential for the accurate interpretation of

dental wear signals in deep-time and archaeo-

logical contexts.

The taxonomy of Neotropical cervids is

currently under reevaluation. Recent genetic

analyses indicate that the traditionally rec-

ognized Mazama temama represents a com-

plex of at least three species: M. temama

(Mexico–Honduras), Mazama reperticia

(Nicaragua–Panama), and Mazama zetta

(Colombia–Ecuador) (Escobedo-Morales et

al., 2025). Taxonomic relationships within the

widely distributed white-tailed deer, Odocoi-

leus virginianus, are also the subject of ongoing

discussion, given its broad geographic range

and ecological variability across the Americas

(López-Arévalo et al., 2020).

To provide ecological context for the inter-

pretation of dental wear proxies, it is important

to briefly consider the feeding ecology of the

two species examined in this study. The white-

tailed deer (O. virginianus) is generally char-

acterized as a highly plastic and opportunistic

species, capable of exploiting a wide spectrum

of plant resources across open and heteroge-

neous landscapes throughout the Americas

(Eisenberg, 1989; Emmons, 1999; Gallina-Tes-

saro et al., 2010). In contrast, Central American

red brocket (M. reperticia) is typically associ-

ated with forested environments and has been

described as a more selective feeder with a

stronger reliance on browse (Bello-Gutiérrez et

al., 2010; Eisenberg, 1989; Gallina-Tessaro et al.,

2019; Sarria-Perea, 2012). Although these eco-

logical differences are well documented, direct

comparisons of dietary strategies between these

taxa, based on dental wear proxies, remain

limited, particularly in Central America (Mar-

tínez-Polanco & Rivals, 2023).

Accordingly, the aim of this study is to

compare the dietary strategies of 16 mod-

ern deer specimens—including 10 white-tailed

deer (O. virginianus) and six Central American

red brocket deer (M. reperticia)—from two

localities in Costa Rica: Guanacaste Province

and Volcán Irazú, using dental micro- and

mesowear analyses. By integrating dietary

proxies that operate at complementary tem-

poral scales, this study evaluates patterns of

ecological niche differentiation across environ-

mentally contrasting settings and provides new

insights into feeding specialization and ecologi-

cal flexibility in tropical cervids, contributing to

a better understanding of resource partitioning

and dietary adaptation in both modern and

archaeological contexts.

MATERIALS AND METHODS

Specimen selection and provenance: The

cervid specimens analyzed in this study are

housed in the Natural History Museum of

Vienna and were collected during the Austrian

scientific expedition to Costa Rica in 1930. The

expedition was organized by the mammalogist

Otto Porsch, and the specimens examined here

were collected by Otto Koller during system-

atic mammal surveys conducted in different

regions of the country (Díaz-Bolaños, 2008;

Weber, 2008).

In this study, a total of 16 deer specimens

were analyzed, including 10 individuals of O.

virginianus and six individuals of M. reperticia.

Only adult individuals with fully erupted per-

manent dentition were included in the present

analysis. Of seven O. virginianus adult males,

six were collected in the vicinity of Bebedero,

Guanacaste Province, and one was collected

in Carrillo, within the primary forests of Santa

Clara. All adult females were collected in Bebe-

dero (Koller, 1934) (Fig. 1).

According to Koller’s original field notes,

all white-tailed deer specimens were person-

ally collected within the same general area

over a period of approximately three weeks

between June and July (Koller, 1934). Because

the specimens derive from a geographically

restricted locality and a very short collection

interval, environmental variation among indi-

viduals is expected to be minimal. Moreover,

dental microwear reflects short-term dietary

signals, typically representing the foods con-

sumed during the last days or weeks prior

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e20264964, enero-diciembre 2026 (Publicado Oct. 09, 2026)

to death. Therefore, seasonal dietary differ-

ences within this sample are likely limited.

This tightly constrained temporal and spa-

tial framework provides a rare opportunity to

examine intra-population variability in den-

tal wear patterns under broadly comparable

environmental conditions.

The Central American red brocket deer

series consists of six skulls without associated

postcranial material, all collected in the forested

areas of Volcán Irazú (Koller, 1934) (Fig. 1).

All individuals in this sample are adult males.

Although the Central American red brocket

deer sample is limited in size and derives from a

different locality, the uniformity in sex reduces

intra-sample variability and strengthens the

comparison of dietary strategies inferred from

dental micro- and mesowear proxies.

Mesowear is inherently a population-based

proxy, and the original framework proposed

by Mikael Fortelius and Daniel Solounias rec-

ommended sample sizes of 20-30 individuals,

although reliable approximations can be

obtained from smaller samples of around 10

specimens (Fortelius & Solounias, 2000).

While larger datasets are desirable, assem-

bling extensive modern comparative samples

is often limited by the availability of curated

skeletal material from wild populations (Pat-

terson, 2002). In the present study, the sample

includes 10 individuals of O. virginianus and

six of M. reperticia. Although relatively mod-

est, these sample sizes are comparable to those

reported in several mesowear studies of both

modern and fossil ungulates and still provide

meaningful ecological information when inter-

preted cautiously (Ackermans et al., 2020; Wil-

son et al., 2025). Furthermore, the integration

of mesowear with dental microwear analysis

strengthens dietary interpretations by combin-

ing proxies that operate at different temporal

scales, thereby improving the robustness of the

ecological inferences derived from the available

material.

Fig. 1. Map of Costa Rica indicating the sampling localities of the studied specimens. Symbols represent the collection sites

of white-tailed deer (Odocoileus virginianus) and Central American red brocket deer (Mazama reperticia) from Guanacaste

Province and Volcán Irazú.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e20264964, enero-diciembre 2026 (Publicado Oct. 09, 2026)

Age and sex considerations: Due to the

limited availability of detailed age-at-death data

and well-established wear sequences for cryptic

Neotropical deer species such as M. reperticia,

individuals could not be assigned to discrete

chronological age classes. To minimize poten-

tial age-related bias, only adult specimens were

included in the analyses, defined as individuals

with complete eruption of the third molar (m3).

Juvenile specimens were excluded because den-

tal microwear patterns in young individuals

may reflect transient dietary signals associated

with nursing, weaning, or incomplete dental

eruption and occlusal function.

Dental wear stages were evaluated using

qualitative morphological criteria based on

the functional condition of the occlusal sur-

face. Teeth exhibiting advanced stages of wear-

characterized by substantial cusp obliteration,

strongly reduced occlusal relief, and extensive

dentine exposure —were excluded to avoid dis-

tortions associated with senescent wear surfac-

es (Severinghaus, 1949). Restricting the sample

to adult individuals with functional occlusal

surfaces ensures comparability with previously

published microwear datasets and provides a

more reliable representation of typical feeding

ecology. In contrast, mesowear has been shown

to be less sensitive to age effects and more

strongly related to long-term dietary patterns,

particularly in intermediate age groups that

commonly dominate fossil assemblages (Rivals

et al., 2007).

Both male and female individuals were

pooled in the analyses, as previous studies

have demonstrated that dental wear patterns

do not differ significantly between sexes (Mar-

tínez-Polanco et al., 2023; Martínez-Polanco

& Rivals, 2023). Microwear observations were

therefore interpreted as reflecting species-level

dietary strategies rather than sex-specific feed-

ing behavior.

Tooth selection and dental wear analysis:

The dental wear analysis focused exclusively

on lower second molars (m2), all of which

correspond to left elements. No other molar

positions were considered in the present study.

The second molar was selected because of its

intermediate position in the tooth row, which

makes it less susceptible to extreme wear than

the first or third molars, while still providing

a reliable signal of dietary abrasion (Fortelius

& Solounias, 2000). The use of a single side

was intended to maintain methodological con-

sistency and to minimize potential sampling

bias. Restricting the analysis to a single side is

a common strategy in dental wear studies to

ensure comparability among specimens when

sample sizes are limited (Martínez-Polanco

et al., 2022). Lower second molars were ana-

lyzed because mandibular elements are more

frequently preserved in archaeological assem-

blages from the study area, and the present

dataset was designed to provide a comparative

framework applicable to such materials (Mar-

tínez-Polanco et al., 2019).

Tooth mesowear analysis: Dental

mesowear analysis was used to infer long-

term dietary strategies from macroscopic

wear patterns on molar crowns. Following the

mesowear ruler method originally proposed

by Fortelius and Solounias (2000), mesowear

reflects the balance between attrition (tooth-

to-tooth contact) and abrasion (food-to-tooth

contact) accumulated over extended periods,

providing an estimate of average dietary abra-

siveness (Ackermans et al., 2020; Fortelius &

Solounias, 2000).

Mesowear was assessed by visual inspec-

tion of cusp morphology, with emphasis on

cusp relief and occlusal sharpness. These fea-

tures were combined into an ordinal mesowear

score ranging from low values, indicative of

sharp cusps and high relief typically associated

with browsing diets, to high values reflecting

rounded or blunt cusps, characteristic of more

abrasive feeding strategies. In this scale, these

features are scored on a scale from zero (sharp

and high relief) to six (very blunt), and an aver-

age mesowear score (MWS) is calculated for

each taxon (Mihlbachler et al., 2011; Rivals et

al., 2013; Rivals et al., 2017).

To aid interpretation, mesowear scores

obtained for O. virginianus and M. reperticia

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e20264964, enero-diciembre 2026 (Publicado Oct. 09, 2026)

were compared with published reference val-

ues for extant ungulates representing brows-

ing, mixed-feeding, and grazing dietary guilds,

generated using comparable scoring protocols

(Fortelius & Solounias, 2000; Rivals et al.,

2011; Rivals et al., 2013; Rivals et al., 2017).

These reference data were used exclusively as a

descriptive framework and were not included

in statistical analyses.

Tooth microwear analysis: Dental

microwear was analyzed using the traditional

low-magnification microwear method based

on the counting of pits and scratches on the

occlusal enamel surface under stereomicros-

copy. This approach follows the protocol widely

applied in ungulate dietary reconstructions

(Solounias & Semprebon, 2002; Semprebon

et al., 2004). Dental microwear analysis was

used to reconstruct short-term dietary behavior

based on microscopic wear features formed on

the enamel surface during feeding (Fortelius &

Solounias, 2000). Microwear patterns consist

primarily of pits and scratches generated by

the interaction between food items, exogenous

abrasives, and tooth enamel. Because these fea-

tures form rapidly and are continuously over-

written, they reflect dietary intake during the

last days or weeks prior to death and provide

a snapshot of recent feeding behavior (Davis &

Pineda-Munoz, 2016; DeSantis, 2016; Solou-

nias & Semprebon, 2002; Xafis et al., 2017).

Prior to analysis, all teeth were examined

under a stereomicroscope to evaluate surface

preservation. Specimens showing post-mortem

damage, surface alteration, contamination, or

conservation-related modifications that could

obscure or mimic dietary wear were excluded.

Experimental studies have shown that tapho-

nomic processes such as trampling, sediment

pressure, or thermal alteration can modify

enamel surfaces and potentially produce fea-

tures resembling dietary microwear (Micó,

Blasco, Muñoz del Pozo et al., 2024; Micó et

al., 2024). Only molars with well-preserved

occlusal surfaces allowing clear identification

of primary microwear features were retained

for analysis.

Microwear analysis followed established

low-magnification protocols widely applied to

ungulates. High-resolution dental impressions

were obtained using light-body polyvinylsilox-

ane, and epoxy resin casts were produced for

observation. Microwear features were examined

under a stereomicroscope at 35X magnification

using transmitted light. Counts of pits and

scratches were recorded within a standardized

0.4 × 0.4 mm area located on the enamel band

of the protoconid of the lower second molar

(m2). The number of scratches allows dietary

categorization: browsers (0-17 scratches), graz-

ers (17.5-29.5 scratches), and mixed feeders,

which fall in the overlapping range (Fortelius

& Solounias, 2000). Bivariate plots were gener-

ated using R code following Rivals (2019), with

packages including readxl, ggrepel, plyr, lattice,

Rmisc, and ggplot2 (RStudio Team, 2022).

In addition to the total counts of pits and

scratches, supplementary microwear variables

were recorded to refine dietary interpreta-

tion, including the presence of cross scratches

(% XS), large pits (% LP), and gouges (% G).

Scratch texture was assessed using a scratch

width score (SWS), ranging from fine to coarse

scratches, with average values calculated per

species (Solounias & Semprebon, 2002; Sem-

prebon et al., 2011). Microwear variables were

interpreted using published reference frame-

works for browsing, mixed-feeding, and graz-

ing strategies, which were applied solely for

descriptive purposes.

All microwear observations were made by

a single trained observer following standard-

ized protocols. Scoring was done with aware-

ness of species identity but without considering

geographic origin, which was only included

during the analysis.

Following the approach proposed by Rivals

et al. (2015), the standard deviation (SD) and

coefficient of variation (CV) were assessed

(Rivals et al., 2015). These metrics provide

insights into short-term dietary heterogene-

ity within taxa and allow evaluation of the

degree of dietary consistency or flexibility

reflected in the microwear signal. Variability

in scratch counts was examined exclusively for

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e20264964, enero-diciembre 2026 (Publicado Oct. 09, 2026)

O. virginianus. This approach was adopted

because previous microwear studies on white-

tailed deer provide a well-established com-

parative framework for interpreting dispersion

metrics such as standard deviation and coef-

ficient of variation (Martínez-Polanco et al.,

2019; Martínez-Polanco et al., 2023). In addi-

tion, the precise collection dates available for

the Costa Rican white-tailed deer specimens

allow the microwear signal to be evaluated

within a well-constrained temporal context,

facilitating direct comparison with previously

published datasets.

Recent studies have successfully applied

combined dental wear approaches to M.

temama and M. reperticia (Martínez-Polan-

co, 2022; Martínez-Polanco & Rivals, 2023;

Martínez-Polanco, 2026); however, interpreting

scratch variability remains challenging, par-

ticularly in the absence of detailed chronologi-

cal information for all specimens. As a result,

dispersion metrics for this taxon were not

evaluated within the same analytical framework

used for white-tailed deer. Instead, microwear

data for M. reperticia are discussed primarily in

descriptive terms, emphasizing general patterns

of dental wear rather than formal assessments

of short-term dietary variability.

RESULTS

Tooth mesowear results: Mesowear scores

differed between the two cervid species ana-

lyzed (Table 1). M. reperticia (n = 6) displayed

low average mesowear values (mean MWS =

0.17, SD = 0.41), indicating predominantly

sharp cusps and high occlusal relief across the

sample. In contrast, O. virginianus (n = 10)

exhibited higher mesowear scores (mean MWS

= 0.70, SD = 0.67), reflecting a greater degree

of cusp rounding and reduced occlusal relief.

M. reperticia shows relatively low variability, as

indicated by the standard deviation of scratch

counts. O. virginianus presents more homoge-

neous distribution of mesowear scores within

the sample.

Mesowear analysis reveals clear differences

in dental wear patterns between the two cer-

vid species. M. reperticia consistently displays

low MWS, with values clustering near zero

and showing minimal variability, indicating

limited exposure to abrasive dietary compo-

nents. In contrast, O. virginianus exhibits higher

and more variable MWS values, falling within

the lower-to-intermediate range of the dietary

reference framework. When compared with

extant dietary categories, M. reperticia overlaps

Tabl e 1

Mesowear and microwear data of Mazama reperticia (red brocket deer) and Odocoileus virginianus (white-tailed deer) from

Costa Rica.

Mesowear Microwear

nMWS n NP NS % LP % G % XS SWS

M. reperticia M 0.17 9.25 10.50 0 0 10 0

SD 6 0.41 6 5.05 5.73 0 0 10 0

CV - 0.55 0.55

O. virginianus M 0.70 8.10 9.90 0 0 0 0.1

SD 10 0.67 10 2.32 2.13 0 0 0 0.1

CV - 0.29 0.22

Abbreviations: n: sample size, MWS: mesowear score, NS: number of scratches, NP: number of pits, % LP: percentage of

specimens with large pits, % G: percentage of specimens with gouges, % XS: percentage of specimens with cross scratches,

SWS: scratches width score (from zero: fine scratches only to two: coarse scratches only), M: mean, SD: standard deviation,

CV: coefficient of variation.

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e20264964, enero-diciembre 2026 (Publicado Oct. 09, 2026)

closely with leaf browsers, whereas O. virginia-

nus shows partial overlap with both leaf brows-

ers and mixed feeders. These patterns suggest a

more specialized browsing strategy in M. rep-

erticia and a broader dietary niche in O. virgin-

ianus, consistent with greater trophic flexibility

across different environmental settings in Costa

Rica (Fig. 2) (Tab. 1).

Tooth microwear results: Microwear vari-

ables showed moderate differences between the

two cervid species analyzed (Table 1). M. reper-

ticia (n = 6) presented an average of 9.25 num-

ber of pits (NP) and 10.50 number of scratches

(NS), with relatively high standard deviations

(SD = 5.05 for pits; SD = 5.73 for scratches),

indicating substantial inter-individual variabil-

ity. The coefficients of variation were similar

for pits and scratches (CV = 0.55 for both vari-

ables). Large pits and gouges were absent in this

sample, while cross scratches were recorded in

10 % of the observations. The average scratch

width score (SWS) was zero, indicating pre-

dominantly fine scratches.

O. virginianus (n = 10) exhibited slightly

lower mean values for pits (NP = 8.10) and

scratches (NS = 9.90), with smaller standard

deviations (SD = 2.32 for pits; SD = 2.13 for

scratches) and lower coefficients of variation

(CV = 0.29 for pits; CV = 0.22 for scratches).

No large pits, gouges, or cross scratches were

observed in this sample. The average scratch

width score was low (SWS = 0.1), reflect-

ing mainly fine scratches with occasional

coarser features.

The microwear bivariate plot shows the

position of the analyzed cervid samples rela-

tive to reference dietary envelopes defined

by extant browsers and grazers. The Costa

Rican specimens cluster in the low-to-moderate

scratch range and intermediate pit frequen-

cies, plotting close to the browser dietary space

and clearly outside the grazer envelope. Mean

values for O. virginianus and M. reperticia

overlap broadly, with both taxa characterized

by relatively low scratch counts and moderate

pit frequencies. This pattern indicates limited

consumption of abrasive grasses and suggests

a diet dominated by browse or soft vegetation

shortly before death. The proximity of the

samples to the browser reference ellipse sup-

ports a predominantly browsing feeding signal,

Fig. 2. Mesowear results for the Central American red brocket deer (Mazama reperticia) and white-tailed deer (Odocoileus

virginianus) from Costa Rica in comparison with data on ungulates with known diets: Data from Fortelius and Solounias

(2000), Rivals et al. (2011), Rivals et al. (2013), Rivals et al. (2017). Error bars correspond to the standard error of the mean

(±1 SEM).

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e20264964, enero-diciembre 2026 (Publicado Oct. 09, 2026)

with no evidence of a strong grazing compo-

nent (Fig. 3A). Fig. 3B illustrates representative

microwear features on the enamel bands of the

lower second molar (m2) of M. reperticia and

O. virginianus, highlighting the fine and coarse

scratches recorded during analysis.

Scratch variation, quantified using the

standard deviation (SD = 2.13) and coefficient

of variation (CV = 0.22), shows low disper-

sion. The white-tailed deer sample from Gua-

nacaste plots within area A of the heat map

(Fig. 4) shows a consistent microwear signal

Fig. 3. A. Bivariate plot of the average numbers of pits and scratches of Central American red brocket deer (Mazama

reperticia) and white-tailed deer (Odocoileus virginianus) from Costa Rica. Error bars correspond to the standard error of

the mean (± 1 SEM). Ellipses correspond to the Gaussian confidence ellipses (p = 0.95) on the centroid for the extant leaf

browsers and grazers from Solounias and Semprebon (2002) White-tailed deer (Odocoileus virginianus): fine and coarse

scratches (specimen NMW-4154), collected during the rainy season. B. Examples of microwear features on the enamel bands

of the occlusal surface of the lower second molar (m2). Central American red brocket deer (Mazama reperticia): fine and

coarse scratches (specimen NMW-4178).

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e20264964, enero-diciembre 2026 (Publicado Oct. 09, 2026)

suggesting multiple individuals experienced

similar short-term dietary conditions during

the same season.

DISCUSSION

The combined analysis of mesowear and

microwear provides complementary insights

into the feeding ecology of M. reperticia and

O. virginianus from Costa Rica, revealing clear

interspecific differences structured across tem-

poral scales. Mesowear results indicate low to

moderate tooth abrasion in both taxa, con-

sistent with diets dominated by relatively soft

vegetation. However, M. reperticia exhibits

consistently lower mesowear values, support-

ing a predominantly browsing strategy over

the long term, whereas O. virginianus shows

higher average mesowear scores, indicating a

greater contribution of moderately abrasive

food resources. In contrast, microwear pat-

terns show substantial overlap between the two

species. Although M. temama displays slightly

greater dispersion in microwear variables and

O. virginianus shows a somewhat more clus-

tered signal, these differences are subtle and

should be interpreted cautiously.

These contrasting signals emphasize the

importance of integrating dental proxies with

different temporal resolutions. The combi-

nation of low mesowear values and variable

microwear patterns in M. reperticia suggests

long-term dietary stability, recorded over the

extended functional lifespan of the tooth,

coupled with short-term flexibility reflected

in microwear signatures that capture dietary

behavior during the last days or weeks prior

to death. As a forest-dwelling species forag-

ing primarily in the understory, M. reperticia

consumes plant items whose abrasiveness var-

ies with species, phenology, and proximity to

the ground (Flores-Vázquez, 2021). Occasional

intake of more abrasive items, such as fallen

fruits or low vegetation, may increase scratch

counts without producing sustained tooth abra-

sion detectable in mesowear. Although, the

Fig. 4. Heat map (boundary lines of the three regions with the error probability) and the sample of white-tailed deer

(Odocoileus virginianus) from Guanacaste, Costa Rica. A. Seasonal or shorter events. B. Longer than a season events. C.

Separate events (data from Rivals et al., 2015).

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e20264964, enero-diciembre 2026 (Publicado Oct. 09, 2026)

species remains strongly browse-oriented, the

specific plant taxa consumed may vary season-

ally and across habitats. As noted by Martínez-

Polanco (2026), the fruits consumed by M.

temama may be relatively soft and therefore

unlikely to produce conspicuous microwear

features such as pits or gouges, a pattern also

observed in other Neotropical cervids. In addi-

tion, fruit consumption may be highly seasonal;

if most dental wear is generated while animals

feed on leaves, buds, and stems, the frugivorous

component of the diet may be weakly expressed

in the dental record. Although ecological stud-

ies report the consumption of fruits and flow-

ers, particularly in montane cloud forests of

Mexico (Villarreal-Espino-Barros et al., 2008),

dental wear patterns remain dominated by fea-

tures associated with soft browse. This suggests

that the mechanical properties and seasonal

availability of fruits may lead to an underrepre-

sentation of frugivory in microwear signatures

(Martínez-Polanco & Rivals, 2023), highlight-

ing the need to consider food properties and

seasonal dietary contributions when interpret-

ing dental wear patterns.

In contrast, the dental wear patterns of O.

virginianus indicate a relatively stable feeding

strategy across both short- and long-term tem-

poral scales. The Guanacaste specimens origi-

nate from a highly seasonal tropical dry forest

environment (Jiménez et al., 2016), yet they

display moderate mesowear values combined

with low microwear dispersion. All individuals

were collected during the early rainy season, a

period characterized by increased food avail-

ability and the selective consumption of young,

nutritionally rich leaves (Arceo et al., 2005;

Bello et al., 2001; Sánchez-Rojas et al., 1997).

Such selective foraging behavior likely reduces

short-term dietary heterogeneity and contrib-

utes to the relatively homogeneous microwear

signal observed (Martínez-Polanco et al., 2023).

These patterns are consistent with the well-

known ecological flexibility of O. virginianus,

which combines broad dietary potential with

selective feeding when preferred resources are

abundant (Granados et al., 2014).

Similar patterns have been documented

in other dietary studies from tropical dry for-

est systems. Microhistological analyses of fecal

samples from white-tailed deer at the Chamela

Biological Station in Mexico have documented

marked seasonal variation in diet composition

(Arceo et al., 2005). During the rainy season,

deer tend to consume a smaller number of

plant species but rely heavily on young leaves,

which offer higher nutritional quality and lower

concentrations of secondary compounds. In

contrast, dietary diversity increases during the

dry season, when fruits become a more impor-

tant resource. Experimental and comparative

studies of dental microwear further indicate

that fruit consumption may produce distinc-

tive features such as increased pit frequencies,

large pits, or gouges in some taxa (Semprebon

et al., 2011). However, these signals may vary

depending on the mechanical properties of

consumed fruits and the degree of environ-

mental contamination by exogenous particles.

The absence of these distinctive microwear fea-

tures in the sample analyzed suggests that fruit

consumption was probably limited during the

period when the animals were collected.

Finally, although the present study focus-

es on modern ecological contexts, the low

microwear variability observed in O. virginia-

nus has important implications for archaeologi-

cal research. Previous studies have shown that

variation in scratch dispersion can inform on

the temporal clustering of hunting events when

collection dates are well constrained (Martínez-

Polanco et al., 2019; Martínez-Polanco et al.,

2023). The consistency of the microwear signal

documented here therefore provides a valu-

able modern analogue for future archaeologi-

cal and palaeoecological studies investigating

seasonal patterns of deer exploitation across

different environments.

This study provides new insights into

short-term dietary variability in Central Amer-

ican cervids by integrating dental microwear

and mesowear analyses in extant populations

of O. virginianus and M. reperticia. The results

reveal clear interspecific differences in the

interpretative resolution of microwear proxies,

12 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e20264964, enero-diciembre 2026 (Publicado Oct. 09, 2026)

closely linked to ecological niche, foraging

behavior, and the availability of comparative

reference data.

In white-tailed deer, low scratch variabil-

ity and the clustering of individuals within a

single microwear area indicate repeated feeding

events concentrated within the same season.

Because the exact collection dates of these

specimens are known and comparable datasets

are available, microwear variability in O. virgin-

ianus can be robustly interpreted and directly

compared with both modern and archaeologi-

cal populations. These results reinforce the

value of white-tailed deer as a reliable reference

taxon for evaluating short-term dietary pat-

terns and temporal aspects of resource use in

paleoecological and archaeological contexts.

However, the limited number of specimens

and the restricted representation of seasonal

and ontogenetic variation should be considered

when extrapolating these patterns to broader

populations.

In contrast, M. reperticia exhibits pro-

nounced variability in microwear features, par-

ticularly in scratch counts, despite consistently

low mesowear values indicative of a long-term

browsing strategy. This pattern likely reflects

short-term dietary flexibility within forested

environments, where the abrasiveness of con-

sumed plant items may vary due to plant part

selection, phenology, or exogenous contami-

nation, rather than sustained shifts in trophic

strategy. While recent studies have successfully

applied combined dental wear approaches to

M. reperticia, the present results underscore

the need for careful ecological contextualiza-

tion when interpreting microwear dispersion

in this taxon.

Overall, this study highlights the impor-

tance of interpreting microwear variability

within a species-specific ecological framework

and of grounding such interpretations in well-

constrained modern reference datasets. White-

tailed deer emerge as a particularly robust

model for future comparative and archaeologi-

cal applications, while expanded regional and

seasonal datasets for Central American red

brocket deer will further refine the use of dental

wear proxies for reconstructing cervid ecology

and human–environment interactions in tropi-

cal settings.

Ethics statement: The author declares that

they all agree with this publication and that

they have made contributions that justify their

authorship; that there is no conflict of interest

of any kind; and that they have complied with

all relevant ethical and legal requirements and

procedures. All sources of funding are fully and

clearly detailed in the acknowledgements sec-

tion. The respective signed legal document is in

the journal’s archives.

ACKNOWLEDGMENTS

I acknowledge Alexander Bibl and Megumi

Kiesel at the Naturhistorisches Museum Wien,

for their assistance. I am grateful to María del

Pilar Martínez Polanco for her assistance in the

preparation of the map included in this manu-

script. This research received support from the

SYNTHESYS Project (AT-TAF-TA4-23) http://

www.synthesys.info/ which is financed by

European Community Research Infrastructure

Action under the FP7 “Capacities” Program.

The Institut Català de Paleoecologia Humana i

Evolució Social (IPHES) has received financial

support from the Spanish Ministry of Science

and Innovation through the “María de Maeztu”

programme for Units of Excellence (CEX2019-

000945-M) and the Generalitat de Catalunya

and AGAUR research group 2021-SGR-01237.

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