Pelage variation and morphometrics of closely related Callithrix marmoset species and their hybrids
Authors: Malukiewicz Joanna , Warren Kerryn , Boere Vanner , Bandeira Illaira L. C. , Curi Nelson H. A. , das Dores Fabio T. , Fitorra Lilian S. , Furuya Haroldo R. , Igayara Claudia S. , Milanelo Liliane , Moreira Silvia B. , Molina Camila V. , Nardi Marcello S. , Nicola Patricia A. , Passamani Marcelo , Pedro Valeria S. , Pereira Luiz C. M. , Petri Bruno , Pissinatti Alcides , Quirino Adriana Alves , Rogers Jeffrey , Ruiz-Miranda Carlos R. , Silva Daniel L. , Silva Ita O. , Silva Monique O. M. , Summa Juliana L. , Zwarg Ticiana , Ackermann Rebecca R. 
Published: September 2024

Abstract
Background
Hybrids are expected to show greater phenotypic variation than their parental species, yet how hybrid phenotype expression varies with genetic distances in closely-related parental species remains surprisingly understudied. Here, we investigate pelage and morphometric trait variation in anthropogenic hybrids between four species of Brazilian Callithrix marmosets, a relatively recent primate radiation. Marmoset species are distinguishable by pelage phenotype and morphological specializations for eating tree exudates. In this work, we (1) describe qualitative phenotypic pelage differences between parental species and hybrids; (2) test whether significant quantitative differences exist between parental and hybrid morphometric phenotypes; and (3) determine which hybrid morphometic traits show heterosis, dysgenesis, trangression, or intermediacy relative to the parental trait. We investigated cranial and post-cranial morphometric traits, as most hybrid morphological studies focus on the former instead of the latter. Finally, we estimate mitogenomic distances between marmoset species from previously published data.

Results
Marmoset hybrid facial and overall body pelage variation reflected novel combinations of coloration and patterns present in parental species. In morphometric traits, C. jacchus and C. penicillata were the most similar, while C. aurita was the most distinct, and C. geoffroyi trait measures fell between these species. Only three traits in C. jacchus x C. penicillata hybrids showed heterosis. We observed heterosis and dysgenesis in several traits of C. penicillata x C. geoffroyi hybrids. Transgressive segregation was observed in hybrids of C. aurita and the other species. These hybrids were also C. aurita-like for a number of traits, including body length. Genetic distance was closest between C. jacchus and C. penicillata and farthest between C. aurita and the other species.

Conclusion
We attributed significant morphometric differences between marmoset species to variable levels of morphological specialization for exudivory in these species. Our results suggest that intermediate or parental species-like hybrid traits relative to the parental trait values are more likely in crosses between species with relatively lesser genetic distance. More extreme phenotypic variation is more likely in parental species with greater genetic distance, with transgressive traits appearing in hybrids of the most genetically distant parental species. We further suggest that fewer developmental disturbances can be expected in hybrids of more recently diverged parental species, and that future studies of hybrid phenotypic variation should investigate selective pressures on Callithrix cranial and post-cranial morphological traits.