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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)

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    122 pages : illustrations (some color) ; 26 cm.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

    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)

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    71 pages : color illustrations, map ; 26 cm.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

    Supplemental Material for 'New fossil giant panda relatives (Ailuropodinae, Ursidae) : a basal lineage of gigantic Mio-Pliocene cursorial carnivores (American Museum novitates, no. 3996)'

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    Supplemental Material for 'New fossil giant panda relatives (Ailuropodinae, Ursidae) : a basal lineage of gigantic Mio-Pliocene cursorial carnivores (American Museum novitates, no. 3996)' -

    New fossil giant panda relatives (Ailuropodinae, Ursidae) : a basal lineage of gigantic Mio-Pliocene cursorial carnivores (American Museum novitates, no. 3996)

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    71 pages : illustrations (chiefly color), color maps ; 26 cm.Among the fossil members of the giant panda subfamily of ursid carnivorans, Ailuropodinae, one group of species is of giant size, those of Indarctos. Indarctos species have some dental resemblances to and may be closely related to Agriotherium, although there are other clear differences between these taxa, and no known species has definitive shared derived traits that could link these two genera. Here we describe a rich suite of fossil material from both North America and eastern Asia, all belonging to a new genus, Huracan, which possesses characters shared with both Agriotherium and Indarctos but also has diagnostic autapomorphies. The new taxon was distributed widely in the Holarctic during the latest Miocene, including at least four species: the type species Huracan schneideri (previously Agriotherium schneideri) from the latest Hemphillian (Hh4) and possibly early Blancan North American Land Mammal “Ages” (NALMAs), North America; H. coffeyi from the early Late Hemphillian (Hh3) NALMA, North America; H. qiui, sp. nov., from the Baodean Asian Land Mammal “Age” (ALMA), northern China; and H. roblesi from the MN13 zone (latest Miocene–earliest Pliocene) of Spain. Huracan is the nearest sister taxon to Agriotherium, the latter herein considered to be an ailuropodine (in the tribe Agriotheriini) rather than a hemicyonid, and the common ancestor of both genera evolved from Indarctos (with resultant paraphyly of that taxon) or another Indarctos-like ailuropodine bear, likely in eastern Asia. The dentitions of Huracan and Agriotherium both are more specialised for carnivory than most Indarctos species, indicating a radiation of diverse ecological carnivores earlier in the history of the later-diverging, highly specialized herbivores in the giant panda lineage. Their postcranial morphology suggests that species in both genera (Huracan and Agriotherium) were more cursorial than species assigned to Indarctos, and thus well adapted to more open habitats. These derived traits may explain the worldwide replacement of Indarctos species by Huracan and Agriotherium species during the latest Miocene, in response to significant global cooling and expansion of C4 grasslands that occurred at that time

    Comparative anatomy of the insect tracheal system, part 1. Introduction, apterygotes, Paleoptera, Polyneoptera (Bulletin of the American Museum of Natural History, no. 459)

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    184 pages : illustrations (chiefly color) ; 26 cm.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

    Description of two new Labeo (Labeoninae; Cyprinidae) endemic to the Lulua River in the Democratic Republic of Congo (Kasai ecoregion) : a hotspot of fish diversity in the Congo basin (American Museum novitates, no. 3999)

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    22 pages : illustrations (some color), color map ; 26 cm.Labeo mbimbii, n. sp., and Labeo manasseeae, n. sp., two small-bodied Labeo species, are described from the lower and middle reaches of the Lulua River (Kasai ecoregion, Congo basin) in the Democratic Republic of Congo. The two new species are members of the L. forskalii species group and are genetically distinct from all other species of that clade. Morphologically they can be distinguished from central African L. forskalii group congeners except L. dhonti, L. lukulae, L. luluae, L. parvus, L. quadribarbis, and L. simpsoni in the possession of 29 or fewer (vs. 30 or more) vertebrae and from those congeners by a wider interpectoral, among other features. The two new species are endemic to the Lulua River and, although overlapping in geographical range and most meristic and morphometric measures, are readily differentiated by differing numbers of fully developed supraneural bones, predorsal vertebrae, snout morphology, and additional osteological features. The description of these two species brings the total of Labeo species endemic to the Lulua basin to three. The third endemic species, L. luluae, was previously known only from the juvenile holotype, but numerous additional specimens have now been identified. The cooccurrence of 14 Labeo species in the Lulua River, three of which are endemic, highlights this system as a hotspot of Labeo diversity in the Congo basin and across the continent

    Supplemental Material for 'Transverse canal foramen and pericarotid venous network in Metatheria and other mammals (Bulletin of the American Museum of Natural History, no. 462)'

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    Supplemental Material for 'Transverse canal foramen and pericarotid venous network in Metatheria and other mammals (Bulletin of the American Museum of Natural History, no. 462)' -

    Systematic revision of the whip spider family Paracharontidae (Arachnida: Amblypygi) with description of a new troglobitic genus and species from Colombia (American Museum novitates, no. 4000)

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    36 pages : illustrations (some color), map ; 26 cm.The ancient, enigmatic whip spider family Paracharontidae Weygoldt, 1996, representing the basalmost lineage of the arachnid order Amblypygi Thorell, 1883, is revised. The monotypic West African genus Paracharon Hansen, 1921, from Guinea Bissau, is redescribed, based on a reexamination and reinterpretation of the newly designated lectotype. A new troglobitic whip spider, Jorottui ipuanai, gen. et sp. nov., is described from a cave system in the upper basin of the Camarones River in the La Guajira Department of northeastern Colombia. This new taxon is the second extant representative of Paracharontidae and the first outside Africa. It is unambiguously assigned to the family based on several characters shared with Paracharon caecus Hansen, 1921, notably a projection of the anterior carapace margin, the tritosternum not projecting anteriorly, similar pedipalp spination, a reduced number of trichobothria on the tibia of leg IV, and cushionlike female gonopods. A detailed examination confirmed the absence of ocelli in both genera and the presence of three (Paracharon) vs. four (Jorottui, gen. nov.) prolateral teeth on the basal segment of the chelicera, the dorsalmost tooth bicuspid in both genera. The male gonopods of Paracharontidae are described for the first time. Paracharonopsis cambayensis Engel and Grimaldi, 2014, is removed from Paracharontidae and placed incertae sedis in Euamblypygi Weygoldt, 1996; amended, comparative diagnoses are presented for Paracharontidae and Paracharon; and previous interpretations of various diagnostic characters for Paracharontidae are discussed

    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)

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    127 pages : illustrations (chiefly color), color map ; 26 cm.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

    Generic revisions of the Scopaeina and the Sphaeronina (Coleoptera: Staphylinidae: Paederinae: Lathrobiini) (Bulletin of the American Museum of Natural History, no. 460)

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    193 pages : illustrations (some color) ; 26 cm.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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