Bulletin of the American Museum of Natural History

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The Bulletin, published continuously since 1881, consists of longer monographic volumes in the field of natural sciences relating to zoology, paleontology, and geology. Current numbers are published at irregular intervals. The Bulletin was originally a place to publish short papers, while longer works appeared in the Memoirs. However, in the 1920s, the Memoirs ceased and the Bulletin series began publishing longer papers. A new series, the Novitates, published short papers describing new forms.

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    The Trachylepis (Squamata: Scincidae) of Angola : an integrative taxonomic review with the description of seven new species (Bulletin of the American Museum of Natural History, no. 465)
    (American Museum of Natural History., 2024-02-20) Ceríaco, Luis M. P.; Marques, Mariana P.; Parrinha, Diogo.; Tiutenko, Arthur.; Weinell, Jeffrey L.; Butler, Brett O.; Bauer, Aaron M.
    The genus Trachylepis is currently represented by 21 species in Angola, most of them part of nomenclaturally and taxonomically challenging species complexes. In this study we present an integrative taxonomic revision of the genus in Angola and describe seven new species: Trachylepis attenboroughi, sp. nov., Trachylepis bouri, sp. nov., Trachylepis hilariae, sp. nov., Trachylepis ovahelelo, sp. nov., Trachylepis suzanae, sp. nov., Trachylepis vunongue, sp. nov., and Trachylepis wilsoni, sp. nov. As result of our taxonomic revisions, 25 valid Trachylepis species are now confirmed from Angola. A phylogenetic analysis using a combination of mitochondrial (16S, ND2) and nuclear (RAG1) markers, as well as morphological data, supports the recognition of the new species. In addition, data support the revalidation of Trachylepis albopunctata (Bocage, 1867), Trachylepis notabilis (Peters, 1879), and Trachylepis ansorgii (Boulenger, 1907). We also provide a redefinition of Euprepes anchietae Bocage, 1866, which we synonymize with Trachylepis maculilabris (Gray, 1845). Given that the type material for Trachylepis albopunctata, T. angolensis, and T. anchietae has been lost or destroyed, we designate neotypes for the purpose of nomenclatural stability. The description of the new species and the revision and revalidation of previously described Angolan species contribute to a better understanding of the taxonomy and biogeography of the genus, as well as to the general biogeographic patterns and evolution of the Angolan fauna.
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    Jormungandr walhallaensis : a new mosasaurine (Squamata: Mosasauroidea) from the Pierre Shale Formation (Pembina Member: Middle Campanian) of North Dakota (Bulletin of the American Museum of Natural History, no. 464)
    (American Museum of Natural History., 2023-10-30) Zietlow, Amelia R.; Boyd, Clint A.; Van Vranken, Nathan E.
    Mosasaurs are large, carnivorous aquatic lizards with a global distribution that lived during the Late Cretaceous. After 200 years of scientific study, new mosasaur species are still being discovered as new localities are explored and specimens collected long ago are reevaluated using modern standards of species delimitation. Even so, the phylogenetic positions of many key taxa are unresolved and therefore our understanding of mosasaur macroevolution is muddled. Here, we describe a new genus and species of mosasaurine mosasaur comprising a partial skull and skeleton from the Pembina Member of the Pierre Shale Formation in Cavalier County, North Dakota. The lower bound on the age of the specimen is 80.04 ±0.11 Ma, provided by the underlying bentonite bed. Its skull and jaws are nearly complete, and the postcranial skeleton preserves seven cervical vertebrae with three hypapophyseal peduncles, 11 ribs, and five anterior dorsal vertebrae. The new specimen was scored into a modified version of an existing phylogenetic matrix of Mosasauroidea and was recovered in a polytomy with Clidastes; however, given that its morphology is significantly different from that of Clidastes, we refer it to a new genus and species, Jormungandr walhallaensis. Notably, this new taxon shares a mosaic of features seen in both basal (e.g., Clidastes; high dental counts) and derived (e.g., Mosasaurus; subrectangular quadrate) mosasaurines, in addition to possessing its own unique suite of autapomorphies. Given that it possesses morphology intermediate between Clidastes and Plotosaurini, we suspect that future analyses of mosasaur phylogeny, following the addition of new characters and taxa, will recover Jormungandr as transitional between them. Its occurrence increases the known diversity of mosasaurs from the Pembina Member and is the earliest mosasaur to possess autapomorphies of Plotosaurini. Finally, we also analyzed the matrix using different outgroups to test their effect on tree topology and resolution, and found that including multiple nonmosasauroid anguimorphs increased resolution, but not support, of mosasaurid ingroup relationships.
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    Cranial and postcranial morphology of the insectivoran-grade mammals Hsiangolestes and Naranius (Mammalia, Eutheria) : with analyses of their phylogenetic relationships (Bulletin of the American Museum of Natural History, no. 463)
    (American Museum of Natural History., 2023-06-26) Ting, Suyin; Wang , Xiaoming (Paleontologist); Meng, Jin (Paleontologist)
    Early Cenozoic “insectivorans” possess some of the most primitive morphologies among eutherian mammals. Studies of these archaic mammals offer insights into the early diversifications of basal eutherians. Despite such importance, early fossil “insectivorans” from Asia are poorly known due to a scarcity of fossil remains, which often consist only of fragmentary jaws and teeth. Discoveries of remarkably well-preserved fossil “insectivorans”, including complete skulls and articulated postcranial skeletons, from the early Eocene Hengyang Basin in south-central Hunan Province, China, offer a rare opportunity to thoroughly study two taxa belonging to different families. Fine-grained red beds from Hengyang Basin preserve extraordinary fossils with morphological structures rarely seen elsewhere. Thin sections of a skull of Hsiangolestes youngi Zheng and Huang, 1984, for example, reveal the extremely delicate nasal and maxillary turbinates, which, as far as we are aware, are the first known from fossils of this age. We thus take this opportunity to document in detail the cranial and dental morphology, as well as postcranial skeletons, of the Hengyang “insectivorans”. In this monograph, we describe several complete skulls and serial sections of a skull, as well as many partial skulls, mandibles, and postcranial skeletons of Hsiangolestes youngi, an Asian early Eocene insectivoran-grade mammal. We also report a new species of Naranius Russell and Dashzeveg, 1986—N. hengdongensis—an Asian early Eocene cimolestid and describe its well-preserved skulls and mandibles. Hsiangolestes is endemic to Asia. It is currently known only from the earliest Eocene Lingcha Formation, Hengyang Basin, Hunan Province, China. Naranius closely resembles Cimolestes Marsh, 1889, the type genus of the family Cimolestidae. It is mainly distributed in Asia and known from the earliest Eocene deposits in the Bumban Member of the Naran Bulak Formation, Nemegt Basin, of Mongolia, and the Lingcha Formation, Hengyang Basin, Hunan Province, China. The only record of Naranius reported outside of Asia is N. americanus from the early Wasatchian Red Hot Local Fauna, Mississippi, United States. Using PAUP and TNT search algorithms, we place these Hengyang taxa within phylogenetic context of other fossil “insectivorans” from the Mesozoic and early Cenozoic of Asia together with some well-known Holarctic taxa. A phylogenetic analysis of 290 cranial and dental characters from 36 fossil and modern insectivoran-grade taxa is presented, focusing on new materials of Hsiangolestes youngi and Naranius hengdongensis. Based on the results of our phylogenetic analyses, we propose that (1) Hsiangolestes, Prosarcodon, Sarcodon, and Sinosinopa, form a monophyletic group, for which we propose the family name Sarcodontidae; (2) the family Cimolestidae should be restricted to Naranius and Cimolestes, which are sister taxa; (3) the systematic position of Naranius americanus is uncertain; and (4) the family Micropternodontidae should be restricted to Micropternodus and its allies in North America.
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    Transverse canal foramen and pericarotid venous network in Metatheria and other mammals (Bulletin of the American Museum of Natural History, no. 462)
    (American Museum of Natural History., 2023-06-21) MacPhee, R. D. E.; Gaillard, Charlène; Forasiepi, Analía M.; Sulser, R. Benjamin
    Although few nondental features of the osteocranium consistently discriminate marsupials from placentals, the transverse canal foramen (TCF) has been repeatedly offered as a potential synapomorphy of crown-group Marsupialia and their closest allies. To explore this contention appropriately, the TCF needs to be evaluated in relation to the morphofunctional complex of which it is a part, something never previously undertaken in a systematic fashion. This complex, here defined as the pericarotid venous network (PCVN), is assessed using osteological, histological, and ontogenetic information. Although the TCF is usually thought of as a marsupial attribute, some living placentals also express it. What do these clades actually share in regard to this feature, and how do they differ? Our leading hypothesis is that the chief components of the PCVN begin development in the same way in both Marsupialia and Placentalia, but they follow different ontogenetic trajectories in terms of persistence, size, and connections with other elements of the cephalic venous vasculature. Similarities include shared presence of specific emissary and emissarylike veins in the mesocranial region that connect part of the endocranial dural vasculature (cavernous sinus or CS) to the systemic circulation (external and internal jugular veins plus the cerebrospinal venous system). In marsupials the principal pericarotid vessels are the transverse canal vein (TCV) and internal carotid vein (ICV). These veins almost always attain relatively large size during marsupial ontogeny. By contrast, in most placentals their apparent homologs (among others, emissary vein of the sphenoidal foramen and internal carotid venous plexus) evidently slow down or terminate their growth relatively early, and for this reason they play only a proportionally minor role in cephalic drainage in later life. In both clades, these vessels (informally grouped with others in the same region as pericarotid mesocranial distributaries, or PMDs) play a variable role in draining the CS in conjunction with the much larger petrosal sinuses. A pneumatic space within the basisphenoid—called the sphenoid sinus in placentals, transverse basisphenoid sinus (TBS) in marsupials—communicates with PCVN vasculature and should be considered an integral part of the network. The TBS contains red marrow tissues that are active centers of extramedullary hematopoiesis in young stages of some species, although how widespread this function may be in marsupial clades is not yet known. Previous explorations of the marsupial PCVN have been largely limited to determining whether, in any given taxon, a continuous passageway linking the right and left TCFs could be demonstrated running through the basisphenoid (“intramural” condition). It has long been known that a number of species apparently lack this particular passageway, and that the TCFs instead open into the braincase (“endocranial” condition). Puzzlingly, some species appear to have both passageways, others one or the other, and a few none at all, thus inviting questions about their equivalency and the circumstances under which the CS is actually drained by the TCV. Morphologically, these uncertainties can be resolved by viewing the full TCV as a tripartite entity, consisting of a trunk and rostral and caudal branches. The trunk, or the part that leaves the TCF for the external jugular system, receives the rostral and caudal branches, if both are present, within the body of the basisphenoid. The rostral or intramural branch has little or no direct communication with the endocranium in most investigated species. By contrast, the caudal or endocranial branch is an ordinary emissarium, in that it connects a part of the endocranial system of dural veins with the extracranial circulation. Determining branch routing alone does not adequately capture the scale of morphological variety and function encountered in marsupial PCVN organization. We distinguish five patterns of association between TCVs and other PCVN components. These patterns, based on both histological and osteological criteria, are defined as follows: (1) Simple: only rostral passageway present, caudal passageway absent or reduced to a thread; rostral branch veins form midline confluence within TBS in advance of hypophysis; minimal interaction with CS and its distributaries; rostral and caudal portions of TBS discontinuous. (2) Complex: mostly as in (1), except both rostral and caudal branches present and functional; caudal branches communicate with CS/ICV and do not form a confluence; TBS more extensive. (3) Compound: mostly as in (2), except TBS greatly expanded, incorporating most of rostral branch canals, which are correspondingly short. (4) Hybrid: differs from others in that only the pathways for enlarged caudal branches are significant; they originate from the CS/ICV caudal to the position of the hypophysis; rostral branches absent or highly reduced. (5) Indeterminate: transverse foramina, canals, and branches absent or unidentifiable as such, presumably due to vascular involution early in ontogeny. In light of TCV composition, the trunk of the TCV can be considered a mixed-origin vein, maximally receiving both a quasisystemic or emissarylike vessel (rostral branch) that does not originate from endocranial dural vessels, and a true emissarial vessel (caudal branch) that does. Some extant geomyoid rodents and strepsirrhine primates exhibit enlarged venous structures in the mesocranial region; these are briefly surveyed for comparative purposes, but resemblances to conditions in marsupials are superficial and unmistakably interpretable as convergences. Members of the extinct marsupial sister group Sparassodonta sometimes lack detectable TCFs, as do other non-marsupial metatherians in the fossil record. Evidence for the transverse canal and other PCVN components in other therians is briefly outlined. In summary, the development of mesocranial vasculature as outlined in this paper is hypothesized to be basal for therians, but Marsupialia and Placentalia radically differ in the end expression of PMDs in the adult stage. In prenatal stages of both clades, initial differentiation of these distributaries is presumably similar, but, compared to marsupials, in almost all placental groups these vessels are retained in an undeveloped or neotenic state. By contrast, enhanced expression of the TCV trunk and its branches seems to be a genuine novelty characterizing Marsupialia, although one probably present in some other metatherian groups. Accordingly, the transverse foramen, canal, and related features are probably best regarded as an innovation occurring in the marsupial stem, not a synapomorphy of the crown group as previously suggested by some authors.
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    Systematic revision of Thomasomys cinereus (Rodentia: Cricetidae: Sigmodontinae) from northern Peru and southern Ecuador, with descriptions of three new species (Bulletin of the American Museum of Natural History, no. 461)
    (American Museum of Natural History., 2023-06-21) Pacheco Torres, Víctor R. (Víctor Rául); Ruelas, Dennisse
    Thomasomys cinereus is the type species of Thomasomys, type genus of the sigmodontine tribe Thomasomyini. As currently recognized, Thomasomys includes 48 species, all of which are endemic to humid montane or premontane forests in the tropical Andes. Although it has been suggested that T. cinereus is a species complex, this hypothesis has yet to be critically evaluated. Herein we provide a revision of the species based on a qualitative assessment of external, craniodental, and soft morphological traits; morphometric analyses; a phylogenetic analysis based on cytochrome b gene sequences; species delimitation methods; and first-hand examination of type material. Our analyses of genetic data recovered four distinct clades within T. cinereus, one corresponding to T. cinereus sensu stricto (restricted to the montane forests delimited by the Río Marañón, Río Huancabamba, and Río Tablachaca in Cajamarca department, Peru) and three new species: Thomasomys lojapiuranus, sp. nov., from the montane forests of Piura department, Peru, and Loja province, Ecuador; T. shallqukucha, sp. nov., restricted to the Kañaris montane forests in the Peruvian department of Lambayeque; and T. pagaibambensis, sp. nov., restricted to the montane forests of Pagaibamba in Cajamarca department, Peru. These species can be distinguished by several discrete morphological traits of the skull, dentition, mandible, stomach, palatal rugae, and glans penis. Genetic distances among these taxa range from 5.06%–7.65% at the cytochrome b locus, and delimitation analyses based on cytochrome b sequence data support their recognition as distinct species. Our results suggest the existence of previously unsuspected dispersal barriers in the Andes of northern Peru, and they confirm that the Río Marañón is a formidable barrier that limits the distribution of species of Thomasomys as well as other sigmodontine rodents.
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    Generic revisions of the Scopaeina and the Sphaeronina (Coleoptera: Staphylinidae: Paederinae: Lathrobiini) (Bulletin of the American Museum of Natural History, no. 460)
    (American Museum of Natural History., 2023-06-21) Herman, Lee H.
    The generic classifications of the paederine subtribes Scopaeina Mulsant and Rey, 1878, and Sphaeronina Casey, 1905, are revised. Sphaeronina, revised status, is resurrected from synonymy. Keys to the included genera of both subtribes are included. Newly discovered characters in both subtribes are discussed and illustrated. The Scopaeina now includes Scopaeus, Hyperscopaeus, Micranops, Orus, and Trisunius. The account for each genus includes its diagnostic characters, a description, summary of the general distribution, and list of the included species and specimens examined. Scopaeus Erichson, 1839, has a revised definition and is now restricted to species that have not only a constricted neck and a trichobothrium adjacent to and at about the middorsal margin of the eye, but also a metathoracic/mesofemoral stridulum comprised of a lateral, metaventral file and mesofemoral plectral ridges, slender, apically acute, metakatepisternal processes, and a middorsally fused median lobe of the aedeagus. The stridulum, redefined herein as a file and plectrum that when rubbed together produce stridulation in insects. The metaventral file and mesofemoral plectral ridges of Scopaeus, is, heretofore, unknown in the Staphylinidae or perhaps, even the Coleoptera. Variations of the stridulum and metakatepisternal processes are illustrated and described for each species group. Five genus-group names in the Western Hemisphere, Scopaeomerus Sharp, 1886, and Euscopaeus Sharp, 1886, are new synonyms of Scopaeus; Scopaeodera Casey, 1886, Scopaeoma Casey, 1905, and Scopaeopsis Casey, 1905, are revised status junior synonyms of Scopaeus. The species in those generic groups are now included in species groups of Scopaeus. Hyperscopaeus Coiffait, 1984, new status, is elevated to genus from subgeneric status in Scopaeus. Trisunius Assing, 2011, new subtribal assignment, is moved from the Medonina to the Scopaeina. Typhloscopaeus Jarrige, 1951, incertae sedis, formerly a subgenus of Scopaeus, is of unknown placement, but the species and generic names are retained in Scopaeus awaiting study of the type. Orus cervicula Casey, 1905, revised combination, is returned to Orus from Scopaeus. Orus femoralis (Sharp, 1887), new combination, is transferred from Scopaeus. There are now three named species of Orus with narrow necks. Scopaeus chiriquensis (Sharp, 1886), S. guatemalensis (Sharp, 1886), S. obscurus (Sharp, 1886), and S. palmatus (Sharp, 1886), new combinations, are transferred to Scopaeus from Scopaeomerus. Medon mexicanus (Bernhauer, 1910), new combination, is transferred to Medon from Scopaeomerus. Scopaeus crassitarsis (Sharp, 1886), S. gracilicornis (Sharp, 1886), S. impar (Bierig, 1935), new combinations, are transferred to Scopaeus from Euscopaeus.The following names are transferred from Scopaeus to Hyperscopaeus as new combinations: Hyperscopaeus admixtus (Fagel, 1973), H. albertvillensis (Fagel, 1973), H. allardianus (Fagel, 1973), H. andrewesi (Cameron, 1931), H. angolanus (Fagel, 1973), H. bamaniaensis (Fagel, 1973), H. borneensis (Cameron, 1941), H. bredoanus (Fagel, 1973), H. calidus (Bernhauer, 1932), H. confusoides (Fagel, 1973), H. confusus (Fagel, 1973), H. consimilis (Fagel, 1973), H. convexiceps (Bernhauer, 1932), H. corpulentus (Fagel, 1973), H. decelleanus (Fagel, 1973), H. dolosus (Fagel, 1973), H. endrodyanus (Fagel, 1973), H. errans (Fagel, 1973), H. erraticus (Fagel, 1973), H. fageli (Levasseur, 1981), H. fallaciosus (Fagel, 1973), H. filicornis (Fagel, 1973), H. flavidulus (Fagel, 1973), H. flavocastaneus (Lea, 1923), H. fluviatilis (Fagel, 1973), H. fossiceps (Eppelsheim, 1885), H. fuliginosus (Fagel, 1973), H. fulvescens (Motschulsky, 1858), H. fusculus (Motschulsky, 1858), H. gigantulus (Bernhauer, 1929), H. girardianus (Fagel, 1973), H. hova (Fauvel, 1905), H. hulstaertianus (Fagel, 1973), H. intermixtus (Fagel, 1973), H. kaszabianus (Fagel, 1973), H. katanganus (Fagel, 1973), H. kivuanus (Fagel, 1973), H. lamtoensis (Fagel, 1973), H. leleupianus (Fagel, 1973), H. leopoldvillensis (Fagel, 1973), H. lescuyeri (Delaunay, Coache, and Rainon, 2019), H. levasseuri (Lundgren, 1982), H. longiusculus (Fagel, 1973), H. machadoanus (Fagel, 1973), H. major (Eppelsheim, 1885), H. methneri (Bernhauer, 1932), H. minutulus (Fagel, 1973), H. mulongoensis (Fagel, 1973), H. nitidiceps (Fagel, 1973), H. nitidicollis (Fagel, 1973), H. opacicollis (Bernhauer, 1942), H. overlaetianus (Fagel, 1973), H. parvicornis (Fauvel, 1900), H. procerus (Kraatz, 1859), H. pruinosulus (Eppelsheim, 1885), H. pseudomethneri (Fagel, 1973), H. puberulus (Kraatz, 1859), H. reduncus (Fagel, 1973), H. ripicola (Fagel, 1973), H. rubricollis (Fagel, 1973), H. rubrotestaceus (Kraatz, 1859), H. ruguliceps (Fagel, 1973), H. ruziziensis (Fagel, 1973), H. semifuscus (Kraatz, 1859), H. senegalensis (Fagel, 1973), H. seydeli (Cameron, 1952), H. simillimus (Fagel, 1973), H. simulator (Fagel, 1973), H. spathiferus (Coiffait, 1970), H. spinosophallatus (Frisch, 2012), H. subconfusus (Fagel, 1973), H. subprocerus (Coiffait, 1978), H. surdus (Fagel, 1973), H. suspectus (Fauvel, 1907), H. tchapembanus (Fagel, 1973), H. thoracicus (Motschulsky, 1858), H. tristis (Bernhauer, 1929), H. vagans (Fagel, 1973, and H. voltae (Fagel, 1973). Sphaeronina, revised status, is resurrected from synonymy and now includes Sphaeronum Sharp, 1876, Tripectenopus Lea, 1918, Typhloleleupius Fagel, 1964, and Coecoscopaeus Coiffait, 1982; the last three genera are new assignments to the subtribe. Sphaeronina is redefined by the presence of a hypopharyngeal peg, an enlarged protibial concavity with combs, a ventral denticle on the left mandible, and a groove on the outer edge of the mandibles; additional possible diagnostic characters are discussed. Sphaeronum, Tripectenopus, Typhloleleupius, and Coecoscopaeus are redescribed; the genera are found, respectively, in the American tropical and subtropical regions, Australia, southern Africa and perhaps Madagascar, and Tunisia. Few African and Australian were available for study. Scopaeodracus Scheerpeltz, 1935, is a new synonym of Tripectenopus. Tripectenopus handschini (Scheerpeltz, 1935), new combination, is transferred from Scopaeodracus; Tripectenopus australiae (Fauvel, 1878), T. microps (Lea, 1923), T. pectinatrix (Lea, 1923), and T. torrensensis (Blackburn, 1891), new combinations, are transferred from Domene.
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    Comparative anatomy of the insect tracheal system, part 1. Introduction, apterygotes, Paleoptera, Polyneoptera (Bulletin of the American Museum of Natural History, no. 459)
    (American Museum of Natural History., 2023-03-10) Herhold, Hollister W.; Davis, Steven R., 1983-; DeGrey, Samuel P.; Grimaldi, David A.
    A broad comparative study of insect respiratory morphology is presented. Tracheae, epidermal invaginations extending into the body in branching networks of tubes, supply tissues with direct access to air for gas exchange. While previous tracheal studies focused on a handful of taxa and lacked in consistency, here a unified system of tracheal nomenclature is established using visualizations from micro-CT scanning of representatives from apterygotes, Paleoptera, and Polyneoptera, totaling 29 species, 29 genera, and 26 families in 13 insect orders. Three-dimensional visualizations of named tracheal branches establish robust assessments of homology and provide a framework for further studies across class Insecta. Patterns in respiratory architecture are presented along with a discussion of future investigations into phylogenetic and physiological questions.
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    Revision of the Nearctic species of the genus Amiota Loew (Diptera: Drosophilidae) (Bulletin of the American Museum of Natural History, no. 458)
    (American Museum of Natural History., 2022-09-15) Jones, Lance Eric; Grimaldi, David A.
    Thorough biotic inventories are still needed even in families with paradigm organisms like Drosophilidae, including well-studied areas such as North America. This work presents a taxonomic revision of the species of the genus Amiota Loew in North America and the Nearctic portion of Mexico. Amiota steganoptera Malloch is currently excluded from the Nearctic and Amiota setigera Malloch is synonymized under Amiota humeralis Loew. Specimens of Amiota subtusradiata Duda were not encountered during this study along with its synonym Amiota quadrata Takada and Toda; however, based on previous descriptions we include A. subtusradiata in the Nearctic fauna. All other previously described species from the Nearctic are redescribed. Thirty-six species are described as new: Amiota amputata, A. antitormentum, A. avipes, A. biacuminis, A. brayi, A. byersi, A. cervites, A. cruciatum, A. didens, A. durangoensis, A. elsaltoensis, A. floridiensis, A. forceps, A. fulvitibia, A. hyalou, A. imperator, A. incurva, A. laevifurca, A. latilabrum, A. mcalpinei, A. multiplex, A. nanonigrescens, A. occidentalis, A. onyx, A. oviraptor, A. pseudominor, A. raripennis, A. sinaloensis, A. subnebojsa, A. tessae, A. texas, A. tibialis, A. tormentum, A. uniacuminis, A. wheeleri, and A. zaliskoi. This increases the total species known in the Nearctic from 13 to 49. All species in the Nearctic are illustrated, adult diagnostic features are discussed, and distributions are provided. A cladogram based on parsimony analysis of 46 morphological characters established species groups in the genus. Most of the Nearctic species were accommodated into 10 species groups. Three species groups were previously erected for species in China and Europe. Seven species groups are newly established: the avipes, cervites, hsui, mariae, nebojsa, nigrescens, and subtusradiata groups.Diversity in Amiota appears to be partially dependent on elevation and latitude in the Nearctic, with high diversity found in southern Ontario, the Appalachians, the Ozarks, mountain forests of Arizona and New Mexico, and the Sierra Madre of central Mexico. The taxonomic history of the Nearctic species is reviewed, and various aspects of their biology is presented. Males of species in the A. mariae species group are polymorphic for mirror-image, asymmetric genitalia, called chiral variants. Besides morphology, larval saproxyly, adult lachryphagy, and biogeography are reviewed. Challenges to the study of Amiota and future prospects are discussed.
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    Craniodental morphology and phylogeny of marsupials (Bulletin of the American Museum of Natural History, no. 457)
    (American Museum of Natural History., 2022-06-28) Beck, Robin M. D.; Voss, Robert S.; Jansa, Sharon A.
    The current literature on marsupial phylogenetics includes numerous studies based on analyses of morphological data with limited sampling of Recent and fossil taxa, and many studies based on analyses of molecular data with dense sampling of Recent taxa, but few studies have combined both data types. Another dichotomy in the marsupial phylogenetic literature is between studies focused on New World taxa and those focused on Sahulian taxa. To date, there has been no attempt to assess the phylogenetic relationships of the global marsupial fauna based on combined analyses of morphology and molecular sequences for a dense sampling of Recent and fossil taxa. For this report, we compiled morphological and molecular data from an unprecedented number of Recent and fossil marsupials. Our morphological data consist of 180 craniodental characters that we scored for 97 terminals representing every currently recognized Recent genus, 42 additional ingroup (crown-clade marsupial) terminals represented by well-preserved fossils, and 5 outgroups (nonmarsupial metatherians).Our molecular data comprise 24.5 kb of DNA sequences from whole-mitochondrial genomes and six nuclear loci (APOB, BRCA1, GHR, RAG1, RBP3 and VWF) for 97 marsupial terminals (the same Recent taxa scored for craniodental morphology) and several placental and monotreme outgroups. The results of separate and combined analyses of these data using a wide range of phylogenetic methods support many currently accepted hypotheses of ingroup (marsupial) relationships, but they also underscore the difficulty of placing fossils with key missing data (e.g., †Evolestes), and the unique difficulty of placing others that exhibit mosaics of plesiomorphic and autapomorphic traits (e.g., †Yalkaparidon). Unique contributions of our study are (1) critical discussions and illustrations of marsupial craniodental morphology including features never previously coded for phylogenetic analysis; (2) critical assessments of relative support for many suprageneric clades; (3) estimates of divergence times derived from tip-and-node dating based on uniquely taxon-dense analyses; and (4) a revised, higher-order classification of marsupials accompanied by lists of supporting craniodental synapomorphies. Far from the last word on these topics, this report lays the foundation for future research that may be enabled by the discovery of new fossil taxa, better-preserved material of previously described taxa, novel morphological characters (e.g., from the postcranium), and improved methods of phylogenetic analysis.
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    Spatial and temporal distribution of the island-dwelling Kogaionidae (Mammalia, Multituberculata) in the uppermost Cretaceous of Transylvania (Western Romania) (Bulletin of the American Museum of Natural History, no. 456)
    (American Museum of Natural History., 2022-06-14) Csiki-Sava, Zoltán; Vremir, Mátyás; Meng, Jin (Paleontologist); Vasile, Ștefan; Brusatte, Stephen; Norell, Mark
    The latest Cretaceous kogaionid multituberculates from Transylvania (western Romania) were part of an endemic European clade of mammals that underwent an insular radiation at the end of the Cretaceous and then survived the end-Cretaceous mass extinction that extinguished many groups of contemporary therians. Transylvanian kogaionids lived on what was an island during the latest Cretaceous—“Hațeg Island”—and their fossils are found in the uppermost Campanian to upper Maastrichtian deposits of the Hațeg, Rusca Montană, and southwestern Transylvanian basins. This fossil record has improved dramatically over the past several decades, in part resulting from our decade-long joint Romanian-American-Scottish fieldwork, and comprises one of the most impressive and complete archives of Mesozoic mammals, including not only jaws and teeth but several incomplete skulls and partial skeletons.We here review the fossil record of kogaionids from Transylvania. We report four new occurrences from the Hațeg Basin, update information on previously described ones, and use our database to reassess the chronostratigraphical and geographical distribution of kogaionids and their evolutionary patterns. Although it was previously suggested that large and small kogaionids had largely mutually exclusive spatial distributions, we recognize the cooccurrence of small and large taxa in various units, suggesting a sympatric distribution across their entire chronostratigraphic range. We also identify a novel pattern: small kogaionids appear somewhat earlier than their larger relatives in all well-sampled sedimentary successions, suggesting that kogaionid colonizations of Hațeg Island and component regions took place at small body size and that body size increased only later through local evolution. We find correlations between body size, preservation style, and sedimentary context, which give insight into kogaionid paleobiology and diversity. Larger kogaionids are represented more often by partial skulls and occasionally skeletons compared with small kogaionids, which are usually represented only by isolated teeth, regardless of provenance. Larger kogaionids currently have a higher recognized local taxic diversity than their smaller relatives. We hypothesize that this may be in part a consequence of preservational bias related to body size, as more complete specimens may be more easily diagnosed as distinct taxa than those that are represented by more fragmentary and/or incomplete fossils. If true, the taxic diversity of smaller kogaionids may currently be underestimated. Finally, we identify correspondence between sedimentary facies and preservation style. Red-colored fine-grained rocks, suggestive of well-drained, oxidized floodplain paleoenvironments, yield more complete specimens than drab, greenish or grayish sediments deposited in more poorly drained parts of the floodplain. This pattern may suggest habitat preferences for better-drained floodplain environments and a semifossorial lifestyle for some taxa. As the kogaionid fossil record improves, we can further test the hypotheses and patterns outlined above. The pace of new kogaionid discoveries by our team and others indicates that a more complete picture of kogaionid distribution, paleobiology, and evolution will emerge in the coming years, contributing to a more profound understanding of this peculiar group of island-dwelling Mesozoic mammals.
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    An annotated checklist of Recent opossums (Mammalia: Didelphidae) (Bulletin of the American Museum of Natural History, no. 455)
    (American Museum of Natural History., 2022-04-04) Voss, Robert S.
    Living opossums (Didelphidae) comprise 125 species in 18 genera and 4 subfamilies. This synopsis lists all the didelphid taxa (subfamilies, tribes, genera, subgenera, and species) currently recognised as valid, summarizes information about typification, synonyms, and geographic distributions, remarks noteworthy recent changes in usage, and comments on still outstanding problems. A concluding discussion rejects the notion that the almost twofold increase in opossum species from 1993 to the present is “taxonomic inflation” and considers the impact of new kinds of data and new methods of data analysis on species delimitation.
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    Fourteen new, endemic species of shrew (genus Crocidura) from Sulawesi reveal a spectacular island radiation (Bulletin of the American Museum of Natural History, no. 454)
    (American Museum of Natural History., 2021-12-15) Esselstyn, Jacob A.; Setiawan Achmadi, Anang; Handika, Heru; Swanson, Mark T.; Giarla, Thomas C.; Rowe, Kevin C.
    After nearly a decade of field inventories in which we preserved voucher specimens of the small terrestrial mammals of Sulawesi, we combined qualitative and quantitative analyses of morphological traits with molecular phylogenetics to better understand the diversity of shrews (Soricidae: Crocidura) on the island. We examined the morphology of 1368 specimens and obtained extensive molecular data from many of them, including mitochondrial DNA sequences from 851 specimens, up to five nuclear exons from 657 specimens, and thousands of ultraconserved elements from 90 specimens. By iteratively testing species limits using distinct character datasets and appropriate taxon sampling, we found clear, mostly consistent evidence for the existence of 21 species of shrews on Sulawesi, only seven of which were previously recognized. We divide these 21 species into five morphogroups, provide emended diagnoses of the seven previously named species, and describe 14 new species. The Long-Tailed Group contains Crocidura caudipilosa, C. elongata, C. microelongata, new species, and C. quasielongata, new species; the Rhoditis Group contains C. rhoditis, C. pseudorhoditis, new species, C. australis, new species, and C. pallida, new species; the Small-Bodied Group contains C. lea, C. levicula, C. baletei, new species, C. mediocris, new species, C. parva, new species, and C. tenebrosa, new species; the Thick-Tailed Group contains C. brevicauda, new species and C. caudicrassa, new species; and the Ordinary Group contains C. musseri, C. nigripes, C. normalis, new species, C. ordinaria, new species, and C. solita, new species. Documenting these endemic species reveals a local radiation (20 of the 21 species are members of an endemic clade) in which elevational gradients played a prominent role in either promoting speciation, or at a minimum, fostering the cooccurrence of phenotypically similar species. As now understood, the species-level diversity of Crocidura on Sulawesi is nearly three times the known diversity of any other insular shrew fauna. This study highlights the fact that if we wish to understand the true extent of biodiversity on Earth, large-scale, vouchered organismal inventories followed up with thorough examinations of genetic, morphological, and geographic traits are sorely needed in montane tropical regions, even for purportedly well-studied groups such as mammals.
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    Systematics of the relictual Asian scorpion family Pseudochactidae Gromov, 1998, with a review of cavernicolous, troglobitic, and troglomorphic scorpions (Bulletin of the American Museum of Natural History, no. 453)
    (American Museum of Natural History., 2021-09-30) Prendini, Lorenzo; Ehrenthal, Valentin L.; Loria, Stephanie F.
    The first integrative systematic revision of the relictual Asian scorpion family Pseudochactidae Gromov, 1998, making use of an unprecedented collection of material acquired during several expeditions to most of the type localities, is presented. The subfamilies, genera and species of Pseudochactidae are revised based on a phylogenetic analysis of 140 morphological characters and 8608 nucleotide base pairs of concatenated DNA sequence from two nuclear and three mitochondrial gene loci, and a multivariate statistical analysis of 22 ratios and 8 counts for 60 specimens. Three subfamilies, four genera and six species are recognized in the family. Troglokhammouaninae, subfam. nov., is created to restore the monophyly of the nominotypical subfamily Pseudochactinae Gromov, 1998. Aemngvantom, gen. nov., is created to accommodate Aemngvantom lao (Lourenço, 2012), comb. nov., and Aemngvantom thamnongpaseuam gen. et sp. nov. Four new synonyms are presented: Troglokhammouanus louisanneorum Lourenço, 2017 = Troglokhammouanus steineri Lourenço, 2007, syn. nov.; Vietbocap thienduongensis Lourenço and Pham, 2012 = Vietbocap canhi Lourenço and Pham, 2010, syn. nov.; Vietbocap aurantiacus Lourenço et al., 2018 = V. canhi, syn. nov.; Vietbocap quinquemilia Lourenço et al., 2018 = V. canhi, syn. nov. Revised diagnoses of the subfamilies, genera and species, with comparative images, a key and distribution maps are provided, along with a summary of available data on ecology and conservation status, where applicable. Among the Southeast Asian pseudochactids, all of which appear to be obligately cavernicolous, the three species of Vietbocapinae Lourenço, 2012, are highly troglomorphic whereas the sole species of Troglokhammouaninae is barely so. Applying recently revised definitions of the Schiner-Racovitza system for the classification of subterranean organisms, only Vietbocapinae can be considered troglobitic. The global diversity of cavernicolous, troglomorphic and troglobitic scorpions is similarly revisited and a key to ecological classification of cavernicolous and troglomorphic scorpions presented. The world totals of troglomorphic vs. troglobitic scorpions are currently 58 vs. 28 species, in 29 vs. 17 genera and 15 vs. 13 families, respectively.
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    Systematic revision of the arboreal Neotropical “thorellii” clade of Centruroides Marx, 1890, bark scorpions (Buthidae C.L. Koch, 1837) with descriptions of six new species (Bulletin of the American Museum of Natural History, no. 452)
    (American Museum of Natural History., 2021-09-16) Goodman, Aaron M.; Prendini, Lorenzo; Francke, Oscar F.; Esposito, Lauren A.
    The arboreal Neotropical “thorellii” clade of Centruroides Marx, 1890, bark scorpions (Buthidae C.L. Koch, 1837) is revised, using a novel approach to species delimitation. A phylogenetic analysis, based on 112 morphological characters and 1078 aligned DNA nucleotides from the mitochondrial Cytochrome c Oxidase Subunit I (COI) gene, provided the framework for placing singletons from geographically disparate localities (and often with suboptimal preservation) using COI minibarcodes, thereby enlarging the taxon sample for diagnosis and delimitation of morphological species. Six new species are described, tripling the known diversity in the clade to nine: Centruroides berstoni, sp. nov.; Centruroides catemacoensis, sp. nov.; Centruroides chanae, sp. nov.; Centruroides cuauhmapan, sp. nov.; Centruroides hamadryas, sp. nov.; Centruroides yucatanensis, sp. nov. Revised diagnoses are presented for Centruroides hoffmanni Armas, 1996, Centruroides rileyi Sissom, 1995, and Centruroides schmidti Sissom, 1995. Comparative images, a key and distribution maps for all species of the clade are provided, along with a summary of available data for their ecology.
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    Mammalian diversity and Matses ethnomammalogy in Amazonian Peru. Part 4, Bats (Bulletin of the American Museum of Natural History, no. 451)
    (American Museum of Natural History., 2021-08-27) Velazco, Paúl M.; Voss, Robert S.; Fleck, David W. (David William), 1969-; Simmons, Nancy B.
    In this report, the fourth of our monographic series on mammalian diversity and Matses ethnomammalogyin the Yavarí-Ucayali interfluvial region of northeastern Peru, we document the occurrence of 98 species of bats, including 11 emballonurids, 2 noctilionids, 66 phyllostomids, 1 furipterid, 4 thyropterids, 7 vespertilionids, and 7 molossids. New species based on specimens collected in this region (Peropteryx pallidoptera, Micronycteris matses, Hsunycteris dashe, Sturnira giannae, and Thyropterawynneae) have already been described elsewhere, but noteworthy distributional and taxonomicresults newly reported here include the first specimen of Diclidurus isabella from Peru and the diagnosis of Glossophaga bakeri as a species distinct from G. commissarisi. Lists of examined voucher specimens, identification criteria, essential taxonomic references, and summaries of natural history observations are provided for all species. Original natural history information reported herein includes numerous observations of roosting behavior obtained by indigenous Matses collaborators. We assess the Yavarí-Ucayali bat inventory for completeness and conclude that more species remain to be discovered in the region, where as many as 116 species might be expected. Most of the “missing” species (those expected based on geographic criteria but not actually observed) are aerial insectivores, a guild that is notoriously difficult to sample by mistnetting. Of the 98 species in the observed regional fauna, only 71 are known to occur sympatrically at Jenaro Herrera, by far the best-sampled locality between the Yavarí and Ucayali rivers. Faunal comparisons with extralimital inventories (e.g., from Brazil, Ecuador, and French Guiana) suggest that frugivorous bats are substantially more speciose in western Amazonia than in eastern Amazonia, a result that is consistent with previous suggestions of an east-to-west gradient in the trophic structure of Amazonian mammal faunas. As previously reported, the Matses have only a single name for “bat,” but they recognize the existence of many unnamed local species, which they distinguish on the basis of morphology and behavior. However, by contrast with the well-documented accuracy of Matses observations about primates and other game species, recorded Matses monologs about bat natural history contain numerous factual errors and ambiguities. Linguistic underdifferentiation of bat diversity and inaccurate natural history knowledge are both explained by cultural inattention to small, inedible, and inoffensive nocturnal fauna.
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    Systematic revision of the sand scorpions, genus Buthacus Birula, 1908 (Buthidae C.L. Koch, 1837) of the Levant, with redescription of Buthacus arenicola (Simon, 1885) from Algeria and Tunisia (Bulletin of the American Museum of Natural History, no. 450)
    (American Museum of Natural History., 2021-04-27) Cain, Shlomo; Gefen, Eran; Prendini, Lorenzo
    Scorpions of the genus Buthacus Birula, 1908 (Buthidae C.L. Koch, 1837), commonly known as "sand scorpions," are widespread in the sandy deserts of the Palearctic, from West Africa to India. Although many new species of Buthacus were described in recent years, no modern revision exists for the genus and the limits of many infrageneric taxa remain unclear. The present contribution addresses the species of Buthacus recorded from the Levant, defined here as the region of the Middle East including Syria, Lebanon, Jordan, Israel, the Palestinian territories, and the Sinai Peninsula (Egypt). Prior to this study, five species and subspecies, including several synonyms, were recognized from the region. Based on extensive new collections, a reassessment of the morphology (including multivariate statistical analysis), and a phylogenetic analysis of morphological and DNA sequence data, published elsewhere, seven species of Buthacus are now recognized from the Levant, raising the number of species in the genus to 30. Three new species are described: Buthacus amitaii, sp. nov., endemic to Israel; Buthacus arava, sp. nov., endemic to Israel and Jordan; and Buthacus levyi, sp. nov., endemic to Egypt, Israel, and perhaps Libya. Buthacus arenicola (Simon, 1885) is redescribed and restricted to northeastern Algeria and central Tunisia, and Buthacus leptochelys (Ehrenberg, 1829) redescribed and restricted to Egypt, Sudan, and perhaps Libya. Buthacus armasi Lourenço, 2013, stat. rev., from southern Algeria, and Buthacus spatzi (Birula, 1911), stat. rev., from southern Tunisia and western Libya, are revalidated, and Buthacus fuscata Pallary, 1929, stat. nov. et stat. rev., from southern Algeria, revalidated and elevated to the rank of species. Buthacus nitzani Levy et al., 1973, stat. nov., currently restricted to Israel but probably present in the Sinai Peninsula (Egypt), is elevated to the rank of species. Buthacus tadmorensis (Simon, 1892), stat. rev., recorded from Iran, Iraq, Jordan, Syria, and Turkey, and Buthacus yotvatensis Levy et al., 1973, stat. rev., endemic to Israel and Jordan, are redescribed and revalidated. Three new synonyms are presented: Androctonus (Leiurus) macrocentrus Ehrenberg, 1828 = Buthacus leptochelys (Ehrenberg, 1829), syn. nov.; Buthus pietschmanni Penther, 1912 = Buthacus tadmorensis (Simon, 1892), syn. nov.; Buthacus granosus Borelli, 1929 = Buthacus leptochelys (Ehrenberg, 1829), syn. nov. Buthacus arenicola and the seven species of Buthacus occurring in the Levant are diagnosed and illustrated to modern standards, with updated distribution maps. A list of the currently recognized species of Buthacus, and a key to identification of the species occurring in the Levant are also presented.
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    Cranial morphology and phylogenetic relationships of Trigonostylops wortmani, an Eocene South American native ungulate (Bulletin of the American Museum of Natural History, no. 449)
    (American Museum of Natural History., 2021-04-19) MacPhee, R. D. E.; Hernández Del Pino, Santiago; Bond, Mariano; Kramarz, Alejandro; Forasiepi, Analía M.; Sulser, R. Benjamin
    In 1933 George G. Simpson described a remarkably complete skull of Trigonostylops, an Eocene South American native ungulate (SANU) whose relationships were, in his mind, quite uncertain. Although some authorities, such as Florentino Ameghino and William B. Scott, thought that a case could be made for regarding Trigonostylops as an astrapothere, Simpson took a different position, emphasizing what would now be regarded as autapomorphies. He pointed out a number of features of the skull of Trigonostylops that he thought were not represented in other major clades of SANUs, and regarded these as evidence of its phyletic uniqueness. Arguing that the lineage that Trigonostylops represented must have departed at an early point from lineages that gave rise to other SANU orders, Simpson reserved the possibility that Astrapotheriidae might still qualify (in modern terms) as its sister group. Even so, he argued that the next logical step was to place Trigonostylops and its few known allies in a separate order, Trigonostylopoidea, coordinate with Astrapotheria, Notoungulata, Litopterna, and Pyrotheria. Simpson's classification was not favored by most later authors, and in recent decades trigonostylopids have been almost universally assigned to Astrapotheria. However, his evaluation of the allegedly unique characters of Trigonostylops and its allies has never been systematically treated, which is the objective of this paper. Using computed tomography, the skull of Trigonostylops is compared, structure by structure, to a variety of representative SANUs as well as extant perissodactylans (which together comprise the clade Panperissodactyla) and the "condylarthran" Meniscotherium. In addition to placing Simpson's character evaluations in a comparative context, we also provide detailed assessments of many vascular and pneumatization-related feature of panperissodactylans never previously explored. Overall, we found that this new assessment strengthened the placement of Trigonostylops within a monophyletic group that includes Astrapotherium and Astraponotus, to the exclusion of other SANU clades. Although Trigonostylops cannot be considered as morphologically distinct or unusual as Simpson thought, our comparative and phylogenetic analyses have helped to generate a number of hypotheses about character evolution and function in SANUs that may now be fruitfully tested using other taxon combinations.
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    Systematics of the "giant" Ricinulei (Ricinoididae: Ricinoides) of West Africa : with descriptions of five new species and comparative morphology of the male copulatory apparatus. (Bulletin of the American Museum of Natural History, no. 448)
    (New York, NY : American Museum of Natural History, 2021-04-05) Botero-Trujillo, Ricardo; Sain, Colby E.; Prendini, Lorenzo
    The Ricinulei Thorell, 1876, or "hooded tick-spiders," are among the rarest and least studied arachnid orders. Ricinoides Ewing, 1929, the only Old World genus of extant ricinuleids, with 11 species described from tropical West Africa, is the most neglected of the three genera currently recognized. A lack of attention to the systematics of Ricinoides has created a disparity between its taxonomic diversity and that of the New World genera, Cryptocellus Westwood, 1874, and Pseudocellus Platnick, 1980, in which many new species have been described in recent decades. The present contribution provides a revised diagnosis of Ricinoides, which includes two new, putative synapomorphies for the genus and addresses the systematics and morphology of a group of West African species, which includes the world's largest ricinuleids and the type species of the genus. This group of nine species, referred to as the "giant" Ricinulei, shares a unique combination of characters, many of which appear to be unique to the group, and appears to be monophyletic. Four species of this group are redescribed, with revised diagnoses, based on reexamination of the type material: Ricinoi des afzelii (Thorell, 1892), from Sierra Leone; Ricinoides atewa Naskrecki, 2008, from Ghana; Ric inoides feae (Hansen, 1921), from Guinea-Bissau; and Ricinoides westermannii (Guérin-Méneville, 1838), from Togo. Five new species are described, raising the number of species in the genus to 16: Ricinoides eburneus, sp. nov., and Ricinoides taii, sp. nov., from Côte d'Ivoire; Ricinoides iita, sp. nov., from Nigeria; Ricinoides kakum, sp. nov., from Ghana; and Ricinoides nzerekorensis, sp. nov., from Guinea. Comparative illustrations of the adult morphology are presented for all nine species. The male copulatory apparatus is described and illustrated in detail, and new terminology and characters presented. The female spermathecae are described and illustrated for six species in which the females are known, representing the first illustrated comparison of these structures in African ricinuleids. Geographical distribution records are revised and updated for the different species, and their distributions mapped.
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    Evolution in the genus Rhinella : a total evidence phylogenetic analysis of neotropical true toads (Anura: Bufonidae). (Bulletin of the American Museum of Natural History, no. 447)
    (New York, NY : American Museum of Natural History, 2021-03-31) Pereyra, Martín O.; Blotto, Boris L.; Baldo, Diego; Chaparro, Juan C.; Ron, Santiago R.; Elias-Costa, Agustín J.; Iglesias, Patricia P.; Venegas, Pablo J.; Thomeí, Maria Tereza C.; Ospina-Sarria, Jhon Jairo; Maciel, Natan M.; Rada, Marco; Kolenc, Francisco; Borteiro, Claudio; Rivera-Correa, Mauricio; Rojas Runjaic, Fernando J.M.; Moravec, Jiri; Riva, Ignacio de la; Wheeler, Ward C.; Castroviejo-Fisher, Santiago; Grant, Taran; Haddad, Celio F.B.; Faivovich, Julian
    True toads of the genus Rhinella are among the most common and diverse group of Neotropical anurans. These toads are widely distributed throughout South America, inhabiting a great diversity of environments and ecoregions. Currently, however, the genus is defined solely on the basis of molecular characters, and it lacks a proper diagnosis. Although some phenetic species groups have traditionally been recognized within Rhinella, the monophyly of some of them have been rejected in previous phylogenetic analyses, and many species remain unassigned to these poorly defined groups. Additionally, the identity and taxonomy of several species are problematic and hinder the specific recognition and description of undescribed taxa. In this work, we first perform phylogenetic analyses of separate mitochondrial and nuclear datasets to test the possible occurrence of hybridization and/or genetic introgression in the genus. The comparative analysis of both datasets revealed unidirectional mitochondrial introgressions of an unknown parental species into R. horribilis ("ghost introgression") and of R. dorbignyi into R. bernardoi; therefore, the mitochondrial and nuclear datasets of these species were considered separately in subsequent analyses. We performed total-evidence phylogenetic analyses that included revised molecular (four mitochondrial and five nuclear genes) and phenotypic (90 characters) datasets for 83 nominal species of Rhinella, plus several undescribed and problematic species and multiple outgroups. Results demonstrate that Rhinella was nonmonophyletic due to the position of R. ceratophrys, which was recovered as the sister taxon of Rhaebo nasicus with strong support. Among our outgroups, the strongly supported Anaxyrus + Incilius is the sister clade of all other species of Rhinella. Once R. ceratophrys is excluded, the genus Rhinella is monophyletic, well supported, and composed of two major clades. One of these is moderately supported and includes species of the former R. spinulosa Group (including R. gallardoi); the monophyletic R. granulosa, R. crucifer, and R. marina Groups; and a clade composed of the mitochondrial sequences of R. horribilis. The other major clade is strongly supported and composed of all the species from the non-monophyletic R. veraguensis and R. margaritifera Groups, the former R. acrolopha Group, and R. sternosignata. Consistent with these results, we define eight species groups of Rhinella that are mostly diagnosed by phenotypic synapomorphies in addition to a combination of morphological character states. Rhinella sternosignata is the only species that remains unassigned to any group. We also synonymize nine species, treat three former subspecies as full species, and suggest that 15 lineages represent putative undescribed species. Lastly, we discuss the apparently frequent occurrence of hybridization, deep mitochondrial divergence, and "ghost introgression"; the incomplete phenotypic evidence (including putative character systems that could be used for future phylogenetic analyses); and the validity of the known fossil record of Rhinella as a source of calibration points for divergence dating analyses.
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    Phylogeny, classification, Mesozoic fossils, and biogeography of the Leiinae (Diptera: Mycetophilidae). (Bulletin of the American Museum of Natural History, no. 446)
    (American Museum of Natural History., 2021-03-16) Oliveira, Sarah Siqueira; Amorim, Dalton de Souza
    The relationships among the genera of fungus gnats in the mycetophilid subfamily Leiinae are unclear, and the monophyly of this group is questionable. This monograph provides an extensive phylogenetic study of theLeiinae based on morphological data from a large taxonomic sample, including all genera that have ever been assigned to the subfamily and a wide outgroup sampling to properly test subfamilial monophyly. A data matrix with 128 morphological features of 117 terminal taxa was carried out under parsimony using different implicit weight schemes. All recovered topologies support a monophyletic Leiinae that is more restricted than the usual delimitation of the subfamily. We found no consistent evidence that a clade with Docosia Winnertz, Novakia Strobl, Ectrepesthoneura Enderlein, and Tetragoneura Winnertz group together with the remaining genera of Leiinae. A name with subfamily rank--Tetragoneurinae, already present in the literature--is used here to refer to this group. The allactoneurine genera Sticholeia Søli and Allactoneurade Meijere form a clade with the leiine genus Leiella and the genera of Manotinae, which is deeply nested within the Leiinae. The male terminalia patterns found within the subfamily are analyzed and illustrated. A classification for the Leiinae is proposed grouping 33 genera in seven clades ranked as tribes: Selkirkiini Enderlein, Megophthalmidiini, trib. nov., Rondaniellini, trib. nov., Cycloneurini Shaw and Shaw, Manotini Edwards, Anomalomyiini, trib. nov., and Leiini Edwards. A key for the world genera of Leiinae is also provided. The Cretaceous mycetophilid fossil record is revisited and the biogeographic evolution of the Leiinae is discussed.