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We understand very little about the timing and origins of bioluminescence, particularly as a predator avoidance strategy. Understanding the timing of its origins, however, can help elucidate the evolution of this ecologically important signal. Using fireflies, a prevalent bioluminescent group where bioluminescence primarily functions as aposematic and sexual signals, we explore the origins of this signal in the context of their potential predators. Divergence time estimations were performed using genomic-scale datasets providing a robust estimate for the origin of firefly bioluminescence as both a terrestrial and as an aerial signal. Our results recover the origin of terrestrial beetle bioluminescence at 141.17 (122.63–161.17) Ma and firefly aerial bioluminescence at 133.18 (117.86–152.47) Ma using a large dataset focused on Lampyridae; and terrestrial bioluminescence at 148.03 (130.12–166.80) Ma, with the age of aerial bioluminescence at 104.97 (99.00–120.90) Ma using a complementary Elateroidea dataset. These ages pre-date the origins of all known extant aerial predators (i.e. bats and birds) and support much older terrestrial predators (assassin bugs, frogs, ground beetles, lizards, snakes, hunting spiders and harvestmen) as the drivers of terrestrial bioluminescence in beetles. These ages also support the hypothesis that sexual signalling was probably the original function of this signal in aerial fireflies. 

Oxygen is an important and often limiting reagent of a firefly’s bioluminescent chemical reaction. Therefore, the development of the tracheal system and its subsequent modification to support the function of firefly light organs are key to understanding this process. We employ micro-CT scanning, 3D rendering, and confocal microscopy to assess the abdominal tracheal system in Photinus pyralis from the external spiracles to the light organ’s internal tracheal brush, a feature named here for the first time. The abdominal spiracles in firefly larvae and pupae are of the biforous type, with a filter apparatus and appear to have an occlusor muscle to restrict airflow. The first abdominal spiracle in the adult firefly is enlarged and bears an occlusor muscle, and abdominal spiracles two through eight are small, with a small atrium and bilobed closing apparatus. Internal tracheal system features, including various branches, trunks, and viscerals, were homologized across life stages. In adults, the sexually dimorphic elaboration and increase in volume associated with tracheal features of luminous segments emphasizes the importance of gas exchange during the bioluminescent process. 

Oxygen is an important and often limiting reagent of a firefly’s bioluminescent chemical reaction. Therefore, the development of the tracheal system and its subsequent modification to support the function of firefly light organs are key to understanding this process. We employ micro-CT scanning, 3D rendering, and confocal microscopy to assess the abdominal tracheal system in Photinus pyralis from the external spiracles to the light organ’s internal tracheal brush, a feature named here for the first time. The abdominal spiracles in firefly larvae and pupae are of the biforous type, with a filter apparatus and appear to have an occlusor muscle to restrict airflow. The first abdominal spiracle in the adult firefly is enlarged and bears an occlusor muscle, and abdominal spiracles two through eight are small, with a small atrium and bilobed closing apparatus. Internal tracheal system features, including various branches, trunks, and viscerals, were homologized across life stages. In adults, the sexually dimorphic elaboration and increase in volume associated with tracheal features of luminous segments emphasizes the importance of gas exchange during the bioluminescent process. 

Firefly (Coleoptera: Lampyridae) taxonomy has undergone numerous changes over the past 100 years. In order to help provide stability to the group, types for several of the Lampyridae of the Biologia Centrali Americana were determined or designated in early 2019. Here we provide treatments for the remaining Lampyridae and determine the holotype specimens for four species and designate lectotype specimens for 33 species. 

The Biologia Centrali Americana (B.C.A.) is comprised of eight volumes that deal specifically with Coleoptera. These volumes were split into 18 parts and were published between 1879 and 1911. The family Lampyridae was treated in two parts, the main text (1881) with a supplement (1884). Within volume three, part 2, Gorham lists ~90 species in 14 genera, not including the Phengodini subfamily. Of these, Gorham provided original descriptions for 37 species. During recent research visits (2018 and 2020) the authors were able to study material pertinent to the B.C.A. We were able to confidently designate holotypes, lectotypes, and paralectotypes following ICZN articles 73.1 and 74.1 within these species. Two species described by Gorham (1881) are not treated here. Phaenolis nirgricollis was located with a single specimen, already designate as the holotype. Two female syntypes of Photinus consanguineous were located, however Oliver (1907) synonymized these females with Photinus pyralis. These designations contribute to a larger taxonomic effort to stabilize the nomenclature of this group. The species described in the supplement will be treated in a future work. Subfamilies are listed according to Martin et al. (2019) and genera/species within each subfamily are listed according to the order in Gorham (1881).