Rhinoceros beetle horn development reveals deep parallels with dung beetles.

Beetle horns are attractive models for studying the evolution of novel traits, as they display diverse shapes, sizes, and numbers among closely related species within the family Scarabaeidae. Horns radiated prolifically and independently in two distant subfamilies of scarabs, the dung beetles (Scarabaeinae), and the rhinoceros beetles (Dynastinae). However, current knowledge of the mechanisms underlying horn diversification remains limited to a single genus of dung beetles, Onthophagus. Here we unveil 11 horn formation genes in a rhinoceros beetle, Trypoxylus dichotomus. These 11 genes are mostly categorized as larval head- and appendage-patterning genes that also are involved in Onthophagus horn formation, suggesting the same suite of genes was recruited in each lineage during horn evolution. Although our RNAi analyses reveal interesting differences in the functions of a few of these genes, the overwhelming conclusion is that both head and thoracic horns develop similarly in Trypoxylus and Onthophagus, originating in the same developmental regions and deploying similar portions of appendage patterning networks during their growth. Our findings highlight deep parallels in the development of rhinoceros and dung beetle horns, suggesting either that both horn types arose in the common ancestor of all scarabs, a surprising reconstruction of horn evolution that would mean the majority of scarab species (~35,000) actively repress horn growth, or that parallel origins of these extravagant structures resulted from repeated co-option of the same underlying developmental processes.

Novel Features of the Prenatal Horn Bud Development in Cattle (Bos taurus).

Whereas the genetic background of horn growth in cattle has been studied extensively, little is known about the morphological changes in the developing fetal horn bud. In this study we histologically analyzed the development of horn buds of bovine fetuses between ~70 and ~268 days of pregnancy and compared them with biopsies taken from the frontal skin of the same fetuses. In addition we compared the samples from the wild type (horned) fetuses with samples taken from the horn bud region of age-matched genetically hornless (polled) fetuses. In summary, the horn bud with multiple layers of vacuolated keratinocytes is histologically visible early in fetal life already at around day 70 of gestation and can be easily differentiated from the much thinner epidermis of the frontal skin. However, at the gestation day (gd) 212 the epidermis above the horn bud shows a similar morphology to the epidermis of the frontal skin and the outstanding layers of vacuolated keratinocytes have disappeared. Immature hair follicles are seen in the frontal skin at gd 115 whereas hair follicles below the horn bud are not present until gd 155. Interestingly, thick nerve bundles appear in the dermis below the horn bud at gd 115. These nerve fibers grow in size over time and are prominent shortly before birth. Prominent nerve bundles are not present in the frontal skin of wild type or in polled fetuses at any time, indicating that the horn bud is a very sensitive area. The samples from the horn bud region from polled fetuses are histologically equivalent to samples taken from the frontal skin in horned species. This is the first study that presents unique histological data on bovine prenatal horn bud differentiation at different developmental stages which creates knowledge for a better understanding of recent molecular findings.

Chapter 6. The origin and diversification of complex traits through micro- and macroevolution of development: insights from horned beetles.

Understanding how development and ecology shape organismal evolution is a central goal of evolutionary developmental biology. This chapter highlights a class of traits and organisms that are emerging as new models in evo-devo and eco-devo research: beetle horns and horned beetles. Horned beetles are morphologically diverse, ecologically rich, and developmentally and genetically increasingly accessible. Recent studies have begun to take advantage of these attributes and are starting to link the microevolution of horned beetle development to the macroevolution of novel features, and to identify the genetic, developmental, and ecological mechanisms, and the interactions between them, that mediate organismal innovation and diversification in natural populations. Here, I review the most significant recent findings and their contributions to current frontiers in evolutionary developmental biology.

Development of the giraffe horn and its blood supply.

The giraffe horn is an unusual cranial exostosis that lacks clear delineation and categorization as either horn or antler. The distinction between the two is made by contrasting their methods of development and basic composition. This process of development has not been detailed in the giraffe, a factor contributing to the difficulty in distinguishing the classification of these horns. In a chronological series of giraffe horns from prenatal and postnatal animals, we have observed unique morphologies that define their proper location on the skull, the blood supply to them, and the transitions involved in their histological development. While our observations have facilitated the classification of the giraffe horn, our interpretations were not always in accord with previous reports.