Evolution of weaponry in female bovids
Open accessAbstract
Weaponry is ubiquitous in male ungulates and is driven by intrasexual selection, but the mystery surrounding its sporadic presence in females remains unsolved. Female horns are often smaller and shaped differently to male horns, suggesting a different function; indeed, hypotheses explaining the presence of female horns include competition for food, male mollification and defence against predators. Here we use comparative phylogenetic analyses to show that females are significantly more likely to bear horns in bovids that are conspicuous due to large body size and living in open habitats than inconspicuous species living in closed habitats or that are small. An inability to rely on crypsis or take refuge in deep vegetation has apparently driven the evolution of horns for defence against predators in female bovids, a finding supported by many field observations. Typically, exceptions are small species where females are territorial (e.g. duikers) and use horns in intrasexual contests. Furthermore, we suggest that conspicuousness and territoriality hypotheses may explain other instances of female cranial weaponry (i.e. antlers and ossicones) in other horned ruminants. Our phylogenetic reconstruction indicates that the primary function of horns in females is linked to antipredator defence in most clades, but occasionally to intrasexual competition in others.
AI snapshot
Written from the paper’s full text.
TL;DR
Comparative phylogenetic analyses of 117 bovid species show that female horns are most strongly predicted by conspicuousness—large body size and open habitat—and by female territoriality, with 80 of 82 horned female species explained by these factors. The authors conclude that female horns primarily function in antipredator defence in most clades, and occasionally in intrasexual competition.
Key findings
- In phylogenetically corrected stepwise regressions (n = 116), habitat openness was the strongest predictor of female horn presence in Analysis I (t = 4.592, p < 0.001, r = 0.397), followed by female territoriality (t = 2.930, p = 0.004, r = 0.266) and body mass (t = 2.218, p = 0.029, r = 0.204); shoulder height and group size were not significant.
- In Analysis II, the composite exposure metric outcompeted openness (t = 4.604, p < 0.001, r = 0.397), while territoriality (t = 2.958, p = 0.004, r = 0.268) and body mass (t = 2.108, p = 0.037, r = 0.194) remained significant; the final model had adjusted R² = 0.208.
- Categorical tests found that 92.3% (60/65) of exposed species had female horns (χ² = 32.919, d.f. = 1, p < 0.001), and 88.4% (23/26) of territorial species had female horns (χ² = 5.384, d.f. = 1, p = 0.020); combined exposure/territoriality accounted for 80 of 82 horned species (χ² = 73.430, d.f. = 1, p < 0.001).
- The main exceptions were bongo (Tragelaphus eurycerus) and mountain anoa (Bubalus quarlesi); the authors also propose that horn loss in female chiru (Pantholops hodgsonii) may relate to their use of shallow dug depressions to hide from predators.
Methods
- Comparative study of 117 bovid species; female horn presence/absence was scored from Caro et al. (2003), with species scored as horned if any female members possessed horns.
- Predictors included log-transformed shoulder height, log-transformed body mass, habitat openness score, female territoriality, group size, and a composite exposure metric (log10 shoulder height × mean openness of primary habitats).
- Analyses used a published Cetartiodactyla supertree with additional resolution, standardized independent contrasts, multiple linear regressions forced through the origin with forward stepwise model selection, and χ² tests for categorical exposure and territoriality.
Limitations
- The authors state that they did not explicitly test population-level differences in female territoriality and horn presence, that extensions to other female ruminants are speculative and cannot be easily tested systematically due to small sample sizes, and that the horn–crypsis association could not be formally analyzed because the sample was too small.
- Evident from the text: horn scoring ignored intraspecific variation by classifying a species as horned if any female possessed horns, and the comparative regressions are correlational, so they cannot establish causation.
Model-written summary. Check it against the paper before relying on it.
Integrity signals
Registry
Cites
Statistics
Claims the paper makes
0 of 7 cited sources checked · 0 supported · 0 flagged
- Findingown data§ 4. Discussion
In all models, measures of conspicuousness explained the greatest amount of variance in the presence of horns, followed by female territoriality.
“In all models, measures of conspicuousness (openness and exposure) explained the greatest amount of variance in the presence of horns, followed by female territoriality.”
Quote in the paperSubstantial claim - Findingown data§ 3. Results
88.4 per cent (23/26) of species where females are territorial also exhibit horns in females.
“Similarly, 88.4 per cent (23/26) of species where females are territorial also exhibit horns in females (figure 1; χ2 = 5.384, d.f. = 1, p = 0.020).”
Quote in the paperSubstantial claim - Findingown data§ 3. Results
With body mass included in the model, shoulder height did not have a significant effect on female horn presence.
“With body mass included in the final model, shoulder height did not have a significant effect on female horn presence (t = − 0.164, n = 116, p = 0.870, r = − 0.015).”
Quote in the paperNot a substantial claim - Findingown data§ 4. Discussion
The most parsimonious reconstruction of the evolution of female horns based on the hypothesized phylogenetic tree indicates one potential case of horn loss.
“The most parsimonious reconstruction of the evolution of female horns based on our hypothesized phylogenetic tree (figure 1) indicates one potential case of horn loss.”
Quote in the paperNot a substantial claim - Findingown data§ 3. Results
When territoriality and conspicuousness hypotheses were considered simultaneously, 80 out of 82 species where females carry horns could be accounted for.
“Importantly, when the territoriality and conspicuousness hypotheses were considered simultaneously, 80 out of the 82 species where female carry horns could be accounted for (figure 1; χ2 = 73.430, d.f. = 1, p < 0.001, table 1).”
Quote in the paperSubstantial claim
- Findingown data§ 4. Discussion
Only two bovid species fail to conform to the predictions that female horns evolved in conspicuous species as weapons against predators, or in territorial species for use in conspecific aggression.
“Only two bovid species (see above) fail to conform to our predictions that female horns evolved in conspicuous species as weapons against predators, or in territorial species for use in conspecific aggression (table 1).”
Quote in the paperSubstantial claim - Findingown data§ 3. Results
Females of heavier bovid species were more likely to have horns.
“Females of heavier species were also more likely to have horns (t = 2.218, n = 116, p = 0.029, r = 0.204).”
- Also stated in: § 3. Results · plain
“Similar to the first test, species in which females defend territories and large species were also more likely to bear horns (territoriality: t = 2.958, n = 116, p = 0.004, r = 0.268; body mass: t = 2.108, n = 116, p = 0.037, r = 0.194).”
Quote in the paperSubstantial claim - Also stated in: § 3. Results · plain
- Study design and conductown data§ 2. Material and methods
The tracing of female horn evolution on the composite tree has strong phylogenetic signal (p < 0.001).
“We determined that the tracing of female horn evolution on this composite tree has strong phylogenetic signal (Blomberg et al. 2003; p < 0.001) using PDAP 6.0 software (Garland et al. 1993); therefore, related species tend to resemble each other in their tendency for females to have horns.”
Quote in the paperSubstantial claim0 of 2 sources checked - Interpretationreasoning§ 4. Discussion
Large size must be a benefit in self-defence for conspicuous, open country species.
“Conspicuous, open country species that are tall or large in size cannot hide, may have slower escape speeds, and are driven to carry horns to defend themselves and offspring; in addition, their large size must be a benefit in self-defence.”
Quote in the paperNot a substantial claim - Interpretationreasoning§ 4. Discussion
Inconspicuous, small or closed habitat species are cryptic and relieved of the burden of evolving antipredator weaponry.
“Conversely, inconspicuous, small or closed habitat species are cryptic and relieved of the burden of evolving antipredator weaponry.”
Quote in the paperSubstantial claim - Interpretationown data§ 4. Discussion
Nearly all other cases of horns are found in territorial taxa where females use horns to defend their territories against conspecific females.
“Nearly all other cases of horns are found in territorial taxa where females use horns to defend their territories against conspecific females.”
Quote in the paperSubstantial claim - Interpretationreasoning§ 4. Discussion
Conspicuous, open country species that are tall or large are driven to carry horns to defend themselves and offspring.
“Conspicuous, open country species that are tall or large in size cannot hide, may have slower escape speeds, and are driven to carry horns to defend themselves and offspring; in addition, their large size must be a benefit in self-defence.”
Quote in the paperSubstantial claim - Interpretationreasoning§ 4. Discussion
The chiru's novel means of reducing exposure may have tipped the balance in favour of crypsis and relaxed the need to evolve or maintain weaponry in females of this species.
“This novel means of reducing exposure may have tipped the balance in favour of crypsis and relaxed the need to evolve or maintain weaponry in females of this species.”
Quote in the paperSubstantial claim- Rests on an assumption: Chiru lack horns, have large bodies, live in open deserts and grasslands, and are highly exposed to predators.
- Rests on an assumption: Chiru dig shallow depressions in the dirt to lie down in and hide from predators, reducing their exposure to predators from afar.
- Interpretationreasoning§ 4. Discussion
Female horns and cryptic body pelage might be evolutionarily dissociated in bovids.
“This suggests that female horns and cryptic body pelage might be evolutionarily dissociated in bovids.”
Quote in the paperSubstantial claim - Interpretationown data§ 4. Discussion
The results suggest that competition between females over food in large groups has had little effect on the evolution of horns in females.
“Our results suggest that competition between females over food in large groups has had little effect on the evolution of horns in females.”
Quote in the paperSubstantial claim
- May combine two claims. 11
- The quote only partly states this claim. 1
- Classified as probably not a substantial claim. 4
May combine two claims.
Findingown data§ 3. ResultsGroup size had no effect on the presence of horns in females and was left out of the final regression model.
“Similarly, group size had no effect on the presence of horns in females (t = 1.255, n = 116, p = 0.212, r = 0.118) and was left out of the final regression model (F3,113 = 11.103, p < 0.001, adjusted R2 = 0.207).”
Quote in the paperNot a substantial claimThe quote only partly states this claim.
Findingown data§ 3. Results92.3 per cent (60/65) of exposed bovid species had horns in females.
“we found that 92.3 per cent (60/65) of exposed species had horns (figure 1; χ2 = 32.919, d.f. = 1, p < 0.001).”
Quote in the paperSubstantial claimMay combine two claims.
Findingown data§ 3. ResultsFemale territoriality had a significant effect on the presence of horns in female bovids; females that actively mark or defend territories have horns.
“We also found that territoriality had a significant effect on the presence of horns in females (t = 2.930, n = 116, p = 0.004, r = 0.266) in which females that actively mark or defend territories have horns.”
- Also stated in: § 3. Results · hedged
“Similar to the first test, species in which females defend territories and large species were also more likely to bear horns (territoriality: t = 2.958, n = 116, p = 0.004, r = 0.268; body mass: t = 2.108, n = 116, p = 0.037, r = 0.194).”
Quote in the paperSubstantial claim- Also stated in: § 3. Results · hedged
Probably not a substantial claim: may not pass S3 (load-bearing).
Findingown data§ 3. ResultsWhen exposure was included in the model, habitat openness was not included in the final model.
“Because exposure was included in the model, habitat openness was not included in the final model (t = −0.037, n = 116, p = 0.970, r = − 0.004).”
Quote in the paperNot a substantial claimMay combine two claims.
Findingown data§ 3. ResultsIn female bovids, openness of habitat had the greatest effect on the presence of horns, and females of species living in more open environments were more likely to bear horns.
“We found that openness of habitat had the greatest effect on the presence of horns (t = 4.592, n = 116, p < 0.001, r = 0.397), where females of species living in more open environments were more likely to bear horns.”
Quote in the paperSubstantial claimMay combine two claims.
Findingown data§ 3. ResultsExposure outcompeted openness as a predictor of horns in females; females of highly exposed species were much more likely to have horns than less-exposed species.
“we found that exposure outcompeted openness as a predictor of horns in females: females of highly exposed species were much more likely to have horns than less-exposed species (t = 4.604, n = 116, p < 0.001, r = 0.397).”
Quote in the paperSubstantial claimMay combine two claims.
Interpretationown data§ 4. DiscussionHorns probably evolved several times in female bovids for defence (bovines, antilopines, caprines, hippotragines and alcelaphines) and once for intrasexual combat (cephalophines).
“Figure 1 shows that horns probably evolved several times in female bovids for defence (bovines, antilopines, caprines, hippotragines and alcelaphines) and once for intrasexual combat (cephalophines).”
Quote in the paperSubstantial claimMay combine two claims.
Premisecitation§ 4. DiscussionFemale bongos are non-territorial, large, and live in dense forest, but use their horns in aggressive interactions in female groups to establish a rank order; smaller-horned cows tend to defer to individuals with longer horns.
“Female bongos (T. eurycerus) are non-territorial, large, and live in dense forest, but use their horns in aggressive interactions in female groups to establish a rank order (Estes 1991) with smaller-horned cows tending to defer to individuals with longer horns (Kingdon 1982).”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Estes 1991
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Kingdon 1982
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
Classified as probably not a substantial claim. May combine two claims.
Study design and conductown data§ 2. Material and methodsForward stepwise model selection procedures were used to avoid collinearity and determine which predictors best explained the variance in the presence of horns in females.
“To avoid collinearity issues and to determine which of these predictors best explained the variance in the presence of horns in females, we used forward stepwise model selection procedures: all five factors were simultaneously tested against each other with the most significant factor entering the model first.”
Quote in the paperSubstantial claimMay combine two claims.
Interpretationnone given§ 4. DiscussionConspicuous, open country species that are tall or large cannot hide and may have slower escape speeds.
“Conspicuous, open country species that are tall or large in size cannot hide, may have slower escape speeds, and are driven to carry horns to defend themselves and offspring; in addition, their large size must be a benefit in self-defence.”
Quote in the paperNot a substantial claimMay combine two claims.
Interpretationreasoning§ 4. DiscussionWeaponry in female bovids is principally driven by natural selection, whereas weaponry in males in all taxa and in females in certain taxa is driven by intrasexual selection.
“Weaponry in female bovids is thus principally driven by natural selection, whereas weaponry in males in all taxa and in females in certain taxa is driven by intrasexual selection, both processes driving evolution of the same morphological structure in one taxonomic group.”
Quote in the paperSubstantial claimMay combine two claims.
Interpretationown data§ AbstractThe primary function of horns in females is linked to antipredator defence in most clades, but occasionally to intrasexual competition in others.
“Our phylogenetic reconstruction indicates that the primary function of horns in females is linked to antipredator defence in most clades, but occasionally to intrasexual competition in others.”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S2 (checkable), S3 (load-bearing).
Interpretationnone given§ 4. DiscussionThe paper's arguments are also relevant to other female ruminants.
“We believe that our arguments are also relevant to other female ruminants; although, they cannot be easily tested systematically due to small sample sizes.”
Quote in the paperNot a substantial claimClassified as probably not a substantial claim.
Study design and conductcitation§ 2. Material and methodsStandardized independent contrasts were calculated for each factor to account for shared ancestry.
“We calculated standardized independent contrasts (FIC: Felsenstein 1985) for each of the above factors (territoriality, shoulder height, body mass, habitat openness and group size) using the PDTREE module (Midford et al. 2005) in Mesquite 2.5 (Maddison & Maddison 2008) to account for shared ancestry.”
Quote in the paperSubstantial claim0 of 3 sources checked- cites Felsenstein 1985 source
- cites Midford et al. 2005
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Maddison & Maddison 2008
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
May combine two claims.
Interpretationown data§ 4. DiscussionGroup size was less effective at explaining the variance in female horn presence, and its significant role in past studies may be a byproduct of its significant correlation with the other predictors in the model.
“Group size was less effective at explaining the variance in female horn presence and its significant role in past studies may be a byproduct of its significant correlation with the other predictors in the model.”
Quote in the paperSubstantial claim
The paper relies on this without support.
Chiru lack horns, have large bodies, live in open deserts and grasslands, and are highly exposed to predators.
“Female Tibetan antelope or ‘chiru’ (Pantholops hodgsonii) lack horns, have large bodies, live in open deserts and grasslands, and would appear to be highly exposed to predators in this landscape.”
The paper relies on this without support.
Chiru dig shallow depressions in the dirt to lie down in and hide from predators, reducing their exposure to predators from afar.
“However, chiru are unique in that they dig shallow (approx. 30 cm) depressions in the dirt to lie down in and hide from predators, probably reducing the degree to which they are exposed to predators from afar.”
Self-check
Percentages, sums and reported differences re-computed from the paper’s own numbers.
Each contradiction between two passages listed here was confirmed by an independent check.
Territorial classification conflicts with its stated definition
MinorThe methods say females were scored as territorial if they participated in territorial marking and defence. The results count three hornless species as territorial while saying they mark territories but do not defend them.
“Females were scored as ‘territorial’ if they actively participated in territorial marking and defence against conspecifics; male territoriality was disregarded.”
“Females of the remaining three territorial hornless species (Madoqua spp.) mark territories but do not defend them.”
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Self-citation
- Authors
- Stankowich, T., & Caro, T.
- Affiliations
- University of Massachusetts Amherst (US)
- University of California, Davis (US)
- Corresponding author
- Stankowich, Theodore
- Venue
- Proceedings of the Royal Society B: Biological Sciences
- Publisher
- The Royal Society
- Published
- 2009-09-16
- Volume
- 276
- Issue
- 1677
- Pages
- 4329-4334
- ISSN
- 0080-4649 (ISSN-L)
- 0962-8452
- 1471-2954
- PMC
- PMC2817105
- OpenAlex
- W2146089548
- Cited by
- 108
- References
- 40
- Citations by year
- 2026: 5 · 2025: 9 · 2024: 4 · 2023: 6 · 2022: 11 · 2021: 8 · 2020: 8 · 2019: 6 · 2018: 4 · 2017: 5 · 2016: 6 · 2015: 5 · 2014: 7 · 2013: 8 · 2012: 5
- FWCI
- 5.55
- Type
- journal article
- Language
- en
- Open access
- green
- www.ncbi.nlm.nih.gov
- Locations
- Proceedings of the Royal Society B Biological Sciences (publishedVersion)
- PubMed (publishedVersion)
- PubMed Central (submittedVersion)
- Funders
- University of Massachusetts Amherst
- Topics
- Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Animal Behavior and Reproduction
- Physical Sciences › Earth and Planetary Sciences › Paleontology › Evolution and Paleontology Studies
- Physical Sciences › Mathematics › Geometry and Topology › Morphological variations and asymmetry
- Keywords
- Bovidae, predator defense, intrasexual competition, female territoriality, conspicuity, habitat openness, body size, duikers, antler, ossicones, horn evolution, sexual selection
- MeSH terms
- Adaptation, Biological, Animals, Body Weights and Measures, Biological Evolution, Female, Horns, Phylogeny, Ruminants, Species Specificity, Territoriality, Ecosystem, Body Size
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Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
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