Phylogenetic analyses of the subfamily Scarabaeinae (Coleoptera: Scarabaeidae) provide new insights into the Mexican Transition Zone theory.

Being areas of biotic overlap located between biogeographic regions, transition zones function as natural laboratories. The present study explores the phylogenetic history of the dung beetle subfamily Scarabaeinae, in order to present an evolutionary scenario that allows inference of the biogeographic history of the Mexican Transition Zone (MTZ) and integration of the distributional patterns of its biota. The species sampling included 94 New World taxa (93 species of Scarabaeinae and one species of Aphodiinae). The phylogenetic relationships of the main clades recovered in our study were supported with PP values 0.95. Based on the BAYAREALIKE model to reconstruct the ancestral distributional patterns of Scarabaeinae, we inferred a complex scenario with 19 dispersal events, 15 vicariance events, and three extinctions. We suggest that the Ancient Neotropical and Tropical Paleoamerican patterns represent the most likely ancestral distributional patterns for the Scarabaeinae of the MTZ, which probably settle there during the Eocene-Oligocene. The rest of the Scarabaeinae distributional patterns were assembled in subsequent periods. The results suggest that the MTZ had two separate formation stages: a Paleo-MTZ (Eocene-Miocene) and a current MTZ (Pliocene-Anthropocene). We conclude that the evolutionary history as well as the dispersal-vicariance scenario for the Scarabaeinae of the MTZ fits the out of the tropics model.

The genomes of Hercules beetles reveal putative adaptive loci and distinct demographic histories in pristine North American forests.

Beetles, despite their remarkable biodiversity and a long history of research, remain lacking in reference genomes annotated with structural variations in loci of adaptive significance. We sequenced and assembled high-quality chromosome-level genomes of four Hercules beetles which exhibit divergence in male horn size and shape and body colouration. The four Hercules beetle genomes were assembled to 11 pseudo-chromosomes, where the three genomes assembled using Nanopore data (Dynastes grantii, D. hyllus and D. tityus) were mapped to the genome assembled using PacBio + Hi-C data (D. maya). We demonstrated a striking similarity in genome structure among the four species. This conservative genome structure may be attributed to our use of the D. maya assembly as the reference; however, it is worth noting that such a conservative genome structure is a recurring phenomenon among scarab beetles. We further identified homologues of nine and three candidate-gene families that may be associated with the evolution of horn structure and body colouration respectively. Structural variations in Scr and Ebony2 were detected and discussed for their putative impacts on generating morphological diversity in beetles. We also reconstructed the demographic histories of the four Hercules beetles using heterozygosity information from the diploid genomes. We found that the demographic histories of the beetles closely recapitulated historical changes in suitable forest habitats driven by climate shifts.

Long-read Sequencing Data Reveals Dynamic Evolution of Mitochondrial Genome Size and the Phylogenetic Utility of Mitochondrial DNA in Hercules Beetles (Dynastes; Scarabaeidae).

The evolutionary dynamics and phylogenetic utility of mitochondrial genomes (mitogenomes) have been of particular interest to systematists and evolutionary biologists. However, certain mitochondrial features, such as the molecular evolution of the control region in insects, remain poorly explored due to technological constraints. Using a combination of long- and short-read sequencing data, we assembled ten complete mitogenomes from ten Hercules beetles. We found large-sized mitogenomes (from 24 to 28 kb), which are among the largest in insects. The variation in genome size can be attributed to copy-number evolution of tandem repeats in the control region. Furthermore, one type of tandem repeat was found flanking the conserved sequence block in the control region. Importantly, such variation, which made up around 30% of the size of the mitogenome, may only become detectable should long-read sequencing technology be applied. We also found that, although different mitochondrial loci often inferred different phylogenetic histories, none of the mitochondrial loci statistically reject a concatenated mitochondrial phylogeny, supporting the hypothesis that all mitochondrial loci share a single genealogical history. We on the other hand reported statistical support for mito-nuclear phylogenetic discordance in 50% of mitochondrial loci. We argue that long-read DNA sequencing should become a standard application in the rapidly growing field of mitogenome sequencing. Furthermore, mitochondrial gene trees may differ even though they share a common genealogical history, and ND loci could be better candidates for phylogenetics than the commonly used COX1.

Distribution, Regionalization, and Diversity of the dung beetle genus Phanaeus MacLeay (Coleoptera: Scarabaeidae) using Species Distribution Models.

The genus Phanaeus is a well-known group whose taxonomy has been described multiple times. Its distribution was previously classified into 11 ecogeographic groups that are equivalent to areas of endemism. Here we use Species Distribution Models to describe species richness patterns. We measured beta-diversity and regionalized its distribution into one region and one transition zone, both with three dominions: Mexican Transition Zone (North American, Mexican, and Mesoamerican dominions) and Neotropical region (Pacific, Brazilian, and Atlantic Forest dominions). We also present a species checklist and updated the distribution maps for 73 of 81 species described so far that reflects all the taxonomical updates. We include a list of all the recorded locations (by country, state, and province), list the recorded habitats and biomes, and describe the modelled environmental conditions for each species.

Species Redescriptions and New Species of the Onthophagus mexicanus Species Group (Coleoptera: Scarabaeidae), with Notes on Distribution and Rodent-dung Beetle Associations.

The Onthophagus mexicanus species group includes at least 18 species: O. anewtoni Howden and Génier, O. arnetti Howden and Cartwright, O. browni Howden, O. cartwrighti Howden, O. championi Bates, O. concinnus Castelnau, O. cynomysi Brown, O. eulophus Bates, O. guatemalensis Bates, O. hecate (Panzer), O. mcclevei Howden and Génier, O. medorensis Brown, O. mexicanus Bates, O. orpheus (Panzer), O. polyphemi Hubbard, O. pseudoguatemalensis sp. n., O. totonacus sp. n. and O. velutinus Howden and Cartwright. Onthophagus pseudoguatemalensis sp. n. and O. totonacus sp. n. are described from Mexico (Jalisco and Veracruz, respectively). Onthophagus cartwrighti, O. championi, O. eulophus and O. guatemalensis are redescribed, while lectotypes are designated herein for O. championi and O. eulophus. The distribution of O. cartwrighti is clarified; a new country record is provided for O. championi (Honduras); new state records are reported for O. championi (Oaxaca and Veracruz, Mexico) and O. guatemalensis (Oaxaca, Mexico). The accurate distribution of O. eulophus remains unknown since its original description. Updated distribution maps are included for all the species within the group. An updated determination key to species of the O. mexicanus species group is provided. The rarity of O. eulophus and O. totonacus in the entomological collections is thought to be a consequence of their trophic habits; both species are suggested to be inquilines of rodent nests or burrows.

Onthophagus acernorus (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini), a new dung beetle species from Jalisco, Mexico.

The Onthophagus chevrolati Harold, 1869 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini) species complex was recently studied, and a majority of its species were newly described or redescribed (see Moctezuma Halffter 2020). Onthophagus viridichevrolati Moctezuma Halffter, 2020 was the only species within the O. chevrolati species complex recognized to occur in the Mexican state of Jalisco (Moctezuma Halffter 2020). Recently, new material belonging to the O. chevrolati species complex was collected from the maple (Acer; Sapindaceae) forests of Talpa de Allende, Jalisco, Mexico. The examination of these specimens showed that they are morphologically similar to O. chevrolati Harold, 1869 and belong to an undescribed species. In this study, we provide photographs of the habitus (male and female) and male genitalia of O. acernorus new species; an identification key to separate it from O. chevrolati, and an updated distribution map of the O. chevrolati species complex.

The integration processes of the distributional patterns in the Mexican Transition Zone: Phyletic, paleogeographic and ecological factors of a case study.

In this study, we take the Onthophagus chevrolati species group, likely a monophyletic species group as an example to analyze the processes that led to the biogeographic integration of the Holarctic fauna in the mountains of the Mexican Transition Zone to test our biogeographic hypotheses. We propose a change from the status of subspecies to species for O. oaxacanus Zunino Halffter, 1988 new status; O. howdeni Zunino Halffter, 1988 new status; O. jaliscensis Zunino Halffter, 1988 new status; O. longecarinatus Zunino Halffter, 1988 new status; O. omiltemius Bates, 1889 revised status; and O. retusus Harold, 1869 revised status. Consequently, the O. chevrolati group of species is currently made up of 47 species belonging to four species lines: O. vespertilio, O. hippopotamus, O. cyanellus and O. chevrolati. The diversification of the Onthophagus chevrolati species group in this region resulted from three hypothetical stages of evolution. In the first, the penetration and expansion of the ancestor of the O. chevrolati species group occurred before the Miocene and the Trans-Mexican Volcanic Belt arose. During the second stage, the O. hippopotamus species line expanded and evolved, integrating with the paleogeographic changes and the formation of the Trans-Mexican Volcanic Belt, as a consequence of volcanism during the Miocene-Pliocene. In the third and most recent stage, the O. chevrolati species line used the existing mountain systems and interglacial climate fluctuations of the Pleistocene to expand and diversify. Thus, the mountains of the Mexican Transition Zone are not simply periglacial refugia. The entomofauna of Holarctic origin present in the region evolved while the Earth's geological processes were underway.

The Onthophagus fuscus (Coleoptera: Scarabaeidae) species complex: an update and the description of a new species.

The Onthophagus fuscus Boucomont, 1932 (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini) species complex has a total of eight species, one of which is a new species (Onthophagus orizabensis new species) and three are elevated from the subspecies level (O. canescens Zunino Halffter, 1988 new rank; O. mycetorum Zunino Halffter, 1988 new rank; O. parafuscus Zunino Halffter, 2005 new rank). Onthophagus navarretorum Delgado Capistan, 1996 is included, while Onthophagus semiopacus has been removed from the species complex. The distribution map and key for the identification of species in this species complex are updated. Habitat preferences, ecological tolerance, and the restriction of this species complex to the mountains of the Mexican Transition Zone are discussed.

Two new species of Ateuchus with remarks on ecology, distributions, and evolutionary relationships (Coleoptera, Scarabaeidae, Scarabaeinae).

Two new species of the genus Ateuchus Weber are described from the region of Los Chimalapas, Oaxaca, Mexico: A. benitojuarezisp. n. and A. colossussp. n. A diagnosis for distinguishing these new species from the other species of this genus in North America is included. This paper is illustrated with pictures of the dorsal habitus and the male genitalia of the new species. The evolutionary relationships of the species are discussed, as well as their distribution and ecology. It is considered that the species of the genus Ateuchus present in North and Central America correspond to the Typical Neotropical and Mountain Mesoamerican distribution patterns.

ResumenDescribimos dos especies nuevas del género Ateuchus Weber, de la región de Los Chimalapas, Oaxaca, México: A. benitojuarezisp. n. y A. colossussp. n. Incluimos diagnosis para distinguir a estas especies nuevas del resto de especies del género presentes en Norteamérica. Presentamos ilustraciones del hábito dorsal y la genitalia masculina de las nuevas especies. Discutimos las relaciones evolutivas de las especies, así como también su distribución y ecología. Consideramos que las especies del género Ateuchus corresponden con los patrones de distribución Neotropical Típico y Mesoamericano de Montaña.

Two new species of the Phanaeus endymion species group (Coleoptera, Scarabaeidae, Scarabaeinae).

Phanaeus bravoensissp. n. is described from the coniferous-oak forests in the state of Guerrero, and P. huicholsp. n. from coniferous-oak forests and cloud forests in Jalisco and Nayarit. The new species are closely related to P. halffterorum and P. zoque respectively. Morphological trait combination, geographic distribution, and trophic habits show important differences among the studied species. A distribution map and an updated key to separate the species are included.

ResumenDescribimos a Phanaeus bravoensissp. n. proveniente de los bosques de coníferas y encino del estado de Guerrero, y a P. huicholsp. n. de los bosques de coníferas y encino y bosques de niebla de Jalisco y Nayarit. Las nuevas especies están cercanamente relacionadas con P. halffterorum y P. zoque, respectivamente. Las combinaciones de caracteres morfológicos, áreas de distribución geográfica y los hábitos alimenticios muestran diferencias importantes entre las especies estudiadas. Incluimos un mapa de distribución y una clave actualizada para separar a las especies.

A contribution to the knowledge of the mountain entomofauna of Mexico with a description of two new species of Onthophagus Latreille, 1802 (Coleoptera, Scarabaeidae, Scarabaeinae).

Recent intensive samplings carried out across the mountainous regions of El Pinal (Puebla, Mexico) have provided new insights into the main environmental factors that affect the geographic distribution of the scarabaeinae beetles of the Trans-Mexican Volcanic Belt above 2500 m a.s.l. This study is part of an ongoing project investigating the diversity and biogeography of copro-necrophagous beetles (Scarabaeinae, Aphodiinae, Geotrupinae and Silphidae) in the easternmost areas of the Trans-Mexican Volcanic Belt. Previous experience allows us to propose a series of predictions that we expect will provide possible explanations for current distribution patterns observed in Scarabaeinae and other groups of insects found in the Trans-Mexican Volcanic Belt. This mountain range has a primarily biogeographic importance, limiting the Mexican High Plateau in the South and connecting the western and eastern Sierra Madre mountain chains, which are considered the most important routes for dispersal of mountain fauna of northern origin. The taxonomic and biogeographic study of the species collected so far in El Pinal (including Onthophagus clavijeroisp. n. and Onthophagus martinpieraisp. n. described here), along with their possible relationships with other known species, allows us to answer the preliminary assumptions proposed.

ResumenCon base en capturas intensivas realizadas en la montaña de El Pinal (estado de Puebla, México) se plantea un análisis de los factores que intervienen en la distribución geográfica de la fauna de montaña (en este caso Scarabaeinae) por encima de los 2500 m de altitud, en el Sistema Volcánico Transversal (SVT), principal cordillera dispuesta en sentido latitudinal de México y de toda América. Este artículo es parte de una serie de trabajos coordinados para estudiar los escarabajos coprófagos y necrófagos de la parte oriental del SVT.La experiencia generada nos permite plantear una serie de premisas (Introducción) que se espera cumpla la distribución de la fauna de Scarabaeinae y de otros grupos en el SVT, cordillera de primera importancia biogeográfica, pues limita por el sur el Altiplano Mexicano (una península de Norte América) y une las dos principales cordilleras que se desarrollan N-S: las Sierras Madres Occidental y Oriental, las más importantes vías de penetración de la fauna de montaña de origen septentrional. Un análisis de las afinidades taxonómicas y biogeográficas de las colectas de Scarabaeinae en El Pinal (incluyendo dos especies nuevas de Onthophagus que se describen) nos permite dar respuesta en la Discusión a las premisas planteadas en la Introducción.