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1. Ejaculate Allocation and Sperm Characteristics Differ among Alternative Male Types in a Species of Fish with Cooperation and Competition among Unrelated Males

   https://doi.org/10.3390/cells10102612  

   Alonzo, Suzanne H.; Stiver, Kelly A.; Kindsvater, Holly K.; Marsh-Rollo, Susan E.; Nugent, Bridget; Kazancıoğlu, Erem (October 2021, Cells)

   Sexual selection arising from sperm competition has driven the evolution of immense variation in ejaculate allocation and sperm characteristics not only among species, but also among males within a species. One question that has received little attention is how cooperation among males affects these patterns. Here we ask how male alternative reproductive types differ in testes size, ejaculate production, and sperm morphology in the ocellated wrasse, a marine fish in which unrelated males cooperate and compete during reproduction. Nesting males build nests, court females and provide care. Sneaker males only “sneak” spawn, while satellite males sneak, but also help by chasing away sneakers. We found that satellite males have larger absolute testes than either sneakers or nesting males, despite their cooperative role. Nesting males invested relatively less in testes than either sneakers or satellites. Though sneakers produced smaller ejaculates than either satellite or nesting males, we found no difference among male types in either sperm cell concentration or sperm number, implying sneakers may produce less seminal fluid. Sperm tail length did not differ significantly among male types, but sneaker sperm cells had significantly larger heads than either satellite or nesting male sperm, consistent with past research showing sneakers produce slower sperm. Our results highlight that social interactions among males can influence sperm and ejaculate production. 

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2. Experimentally induced variation in neuroendocrine processes affects male reproductive behaviour, sperm characteristics and social interactions

   https://doi.org/10.1111/mec.14999  

   Nugent, Bridget M.; Stiver, Kelly A.; Hofmann, Hans A.; Alonzo, Suzanne H. (February 2019, Molecular Ecology)

   Abstract While extensive research has focused on how social interactions evolve, the fitness consequences of the neuroendocrine mechanisms underlying these interactions have rarely been documented, especially in the wild. Here, we measure how the neuroendocrine mechanisms underlying male behaviour affect mating success and sperm competition in the ocellated wrasse (Symphodus ocellatus). In this species, males exhibit three alternative reproductive types. “Nesting males” provide parental care, defend territories and form cooperative associations with unrelated “satellites,” who cheat by sneaking fertilizations but help by reducing sperm competition from “sneakers” who do not cooperate or provide care. To measure the fitness consequences of the mechanisms underlying these social interactions, we used “phenotypic engineering” that involved administering an androgen receptor antagonist (flutamide) to wild, free‐living fish. Nesting males treated with flutamide shifted their aggression from sneakers to satellite males and experienced decreased submissiveness by sneaker males (which correlated with decreased nesting male mating success). The preoptic area (POA), a region controlling male reproductive behaviours, exhibited dramatic down‐regulation of androgen receptor (AR) and vasotocin 1a receptor (V1aR) mRNA following experimental manipulation of androgen signalling. We did not find a direct effect of the manipulation on male mating success, paternity or larval production. However, variation in neuroendocrine mechanisms generated by the experimental manipulation was significantly correlated with changes in behaviour and mating success: V1aR expression was negatively correlated with satellite‐directed aggression, and expression of its ligand arginine vasotocin (AVT) was positively correlated with courtship and mating success, thus revealing the potential for sexual selection on these mechanisms. 

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3. Sperm and alternative reproductive tactics: a review of existing theory and empirical data

   https://doi.org/10.1098/rstb.2020.0075  

   Kustra, Matthew C.; Alonzo, Suzanne H. (December 2020, Philosophical Transactions of the Royal Society B: Biological Sciences)

   null (Ed.)

   Males that exhibit alternative reproductive tactics (ARTs) often differ in the risk of sperm competition and the energetic trade-offs they experience. The resulting patterns of selection could lead to between-tactic differences in ejaculate traits. Despite extensive research on male ARTs, there is no comprehensive review of whether and what differences in sperm traits exist between male ARTs. We review existing theory on ejaculate evolution relevant to ARTs and then conduct a comprehensive vote-counting review of the empirical data comparing sperm traits between males adopting ARTs. Despite the general expectation that sneaker males should produce sperm that are more competitive (e.g. higher quality or performance), we find that existing theory does not predict explicitly how males adopting ARTs should differ in sperm traits. The majority of studies find no significant difference in sperm performance traits between dominant and sneaker males. However, when there is a difference, sneaker males tend to have higher sperm performance trait values than dominant males. We propose ways that future theoretical and empirical research can improve our understanding of the evolution of ejaculate traits in ARTs. We then highlight how studying ejaculate traits in species with ARTs will improve our broader knowledge of ejaculate evolution. This article is part of the theme issue ‘Fifty years of sperm competition’. 

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4. Defense against outside competition is linked to cooperation in male–male partnerships

   https://doi.org/10.1093/beheco/arz206  

   Hellmann, Jennifer K.; Stiver, Kelly A.; Marsh-Rollo, Susan; Alonzo, Suzanne H.; Taborsky, ed., Michael (December 2019, Behavioral Ecology)

   Abstract Male–male competition is a well-known driver of reproductive success and sexually selected traits in many species. However, in some species, males work together to court females or defend territories against male competitors. Dominant (nesting) males sire most offspring, but subordinate (satellite) males are better able to obtain fertilizations relative to unpartnered males. Because satellites only gain reproductive success by sneaking, there has been much interest in identifying the mechanisms enforcing satellite cooperation (defense) and reducing satellite sneaking. One such potential mechanism is outside competition: unpartnered satellites can destabilize established male partnerships and may force partnered satellites to restrain from cheating to prevent the dominant male from replacing them with an unpartnered satellite. Here, we manipulated perceived competition in the Mediterranean fish Symphodus ocellatus by presenting an “intruding” satellite male to established nesting and satellite male pairs. Focal satellite aggression to the intruder was higher when focal satellites were less cooperative, suggesting that satellites increase aggression to outside competitors when their social position is less stable. In contrast, nesting male aggression to the intruder satellite increased as spawning activity increased, suggesting that nesting males increase their defense toward outside competitors when their current relationship is productive. We found no evidence of altered spawning activity or nesting/satellite male interactions before and after the presentation. These results collectively suggest that response to outside competition is directly linked to behavioral dynamics between unrelated male partners and may be linked to conflict and cooperation in ways that are similar to group-living species. 

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5. Program of the 87th Annual Meeting of the American Association of Physical Anthropologists

   https://doi.org/10.1002/ajpa.23489  

   DeCasien, A.R. (April 2018, American Journal of Physical Anthropology)

   Across taxa, sexually selected traits are more variable in the target sex than 1) the same trait in the opposite sex or 2) non-sexually selected traits, likely due to their condition-dependent expression. In humans, males show greater variability in certain cognitive abilities and brain structures that 1) may facilitate intra- or intersexual competition and 2) are greater/larger in males on average, suggesting these traits may also have been subject to sexual selection. This study investigates sex differences in brain structure variability in chimpanzees. Although male chimpanzees exhibit strong intrasexual competition, reproductive skew is reduced by female mate choice and male coercion. In vivo MRI scans were collected from 226 (135F/91M) individuals and surface areas were calculated for 25 cortical sulci. Outliers for each sex and sulcus were removed prior to analysis. We measured sex differences in variability by calculating the ratio of male-to-female standard deviations of MCMCglmm residuals, controlling for age, rearing condition, scanner type, and kinship. We tested for significant sex differences through permutation. We find that males are significantly more variable at the cingulate (ratio=1.18;p=0.043), middle-frontal (ratio=1.36;p=0.001), occipital-lateral (ratio=1.20;p=0.029), occipital- temporal-marginal (ratio=1.8;p=0.006), superior-temporal (ratio=1.36;p<0.001), subcentral-posterior (ratio=1.62;p=0.033), and superior-parietal (ratio=1.21;p=0.028) sulci. These regions are associated with social perception, face recognition, and motion prediction. Females are more variable at the medio-parietal-occipital sulcus (ratio=0.78;p=0.009), a region associated with planning. This is the first study to demonstrate greater male variability in brain structure in a nonhuman primate species, and suggests sexual selection may lead to greater variability in male cognition across taxa. 

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