Three new Chinese species of Palpifer Hampson from the collection of Franz Daniel (Lepidoptera: Hepialidae).

New Chinese Palpifer species are described from Yunnan and Fujian provinces. The male of Palpifer nielseni sp. n. is described from specimens housed at the Witt Museum Weiden and the Zoologisches Forschungsmuseum Alexander Koenig, while a male of P. chui sp. n. and a male and female of Palpifer climoi sp. n., are described from specimens in the latter collection only. Specimens were originally part of the Franz Daniel collection, collected in 1934-1935 from elevations of 2,300 and 3,000 m. The new species are diagnosed primarily by differences in the male genitalia. The female genitalia of P. climoi sp. n. represent the second published description for Palpifer. Four unique features of the forewing supporting monophyly of Palpifer are discussed.

Biogeographic-tectonic calibration of 14 nodes in a butterfly timetree.

The butterfly subtribe Coenonymphina (Nymphalidae: Satyrinae) comprises four main clades found, respectively, in (1) the Solomon Islands, (2) Australasia, (3) NW South America and (4) Laurasia, with a phylogeny: 1 (2 (3 + 4)). In assessing biogeographic evolution in the group we rejected the conversion of fossil-calibrated clade ages to likely maximum clade ages by the imposition of arbitrary priors. Instead, we used biogeographic-tectonic calibration, with fossil-calibrated ages accepted as minima. Previous studies have used this approach to date single nodes (phylogenetic-biogeographic breaks) in a group, but we extended the methodology to date multiple nodes. Within the Coenonymphina as a whole, 14 nodes coincide spatially with ten major tectonic events. In addition, the phylogenetic sequence of these nodes conforms to the chronological sequence of the tectonic events, consistent with a vicariance origin of the clades. Dating of the spatially coincident tectonic features provides a timescale for the vicariance events. The tectonic events are: pre-drift intracontinental rifting between India and Australia (150 Ma); seafloor spreading at the margins of the growing Pacific plate, and between North and South America (140 Ma); magmatism flare-up along the SW Pacific Whitsunday Volcanic Province-Median Batholith (130 Ma); a change from extension in the Clarence basin, eastern Australia, to uplift of the Great Dividing Range (114 Ma); Pamir Mountains uplift, foreland basin dynamics and high eustatic sea-levels leading to marine transgression of the proto-Paratethys Ocean eastward to Central Asia and Xinjiang (100 Ma); predrift rifting and seafloor spreading west of New Caledonia (100-50 Ma); sinistral strike-slip displacement along the proto-Alpine fault, New Zealand (100-80 Ma); thrust faulting in the Longmen Shan and foreland basin dynamics around the Sichuan Basin (85 Ma); pre-drift rifting in the Coral Sea basin (85 Ma); and dextral displacement on the Alpine fault (20 Ma).

Descriptions of two new genera and six new species of ghost-moths (Lepidoptera: Hepialoidea: Hepialidae) from south-eastern and southern Brazil.

Agripialus gen. n. and Mutipialus gen. n. are described to accommodate the following species: A. campos sp. n., A. variabilis sp. n., A. itatiaia sp. n., A. caparao sp. n., M. dilatus sp. n., and M. monticolus sp. n. All are from southeastern and southern Brazil. Diagnostic characters suggest a closer phylogenetic relationship between these two genera, than to any other described genus.

Taxonomic composition and monophyly of the genus Magnificus (Lepidoptera: Hepialoidea: Hepialidae).

The genus Magnificus Yan, 2000 was originally established to include M. jiuzhiensis Yan, 2000 and M. zhiduoensis Yan, 2000. Here we also include the species M. bouvieri (Oberthür, 1913) comb. n., M. dirschi (Bang-Haas, 1939) comb. n., M. miniatus (Chu Wang, 1985a) comb. n., M. regius (Staudinger, 1896) comb. n., and M. roseus (Oberthür, 1911) comb. n. Monophyly of Magnificus is supported by the shared presence of a unique shape of the tergosternal sclerite in the anterio-abdominal region and pseudoteguminal arm in the male genitalia. We also provide evidence for the inclusion of Magnificus and the northern Eurasian genera Hepialus Fabricius, Pharmacis Hübner, Triodia Hübner and Zenophassus Tindale as members of a larger monophyletic group supported by the shared presence in the male genitalia of a deep U-shaped medial notch in the posterior margin of the saccus and an expanded triangular flange at the lateral corners of the notch. The reciprocally allopatric distribution of Magnificus and its probable sister group is consistent with each originating locally by vicariance over different parts of a widespread ancestral range. This divergence may be the result of tectonic uplift in central Asia.

The Galápagos Islands: biogeographic patterns and geology.

In the traditional biogeographic model, the Galápagos Islands appeared a few million years ago in a sea where no other islands existed and were colonized from areas outside the region. However, recent work has shown that the Galápagos hotspot is 139 million years old (Early Cretaceous), and so groups are likely to have survived at the hotspot by dispersal of populations onto new islands from older ones. This process of metapopulation dynamics means that species can persist indefinitely in an oceanic region, as long as new islands are being produced. Metapopulations can also undergo vicariance into two metapopulations, for example at active island arcs that are rifted by transform faults. We reviewed the geographic relationships of Galápagos groups and found 10 biogeographic patterns that are shared by at least two groups. Each of the patterns coincides spatially with a major tectonic structure; these structures include: the East Pacific Rise; west Pacific and American subduction zones; large igneous plateaus in the Pacific; Alisitos terrane (Baja California), Guerrero terrane (western Mexico); rifting of North and South America; formation of the Caribbean Plateau by the Galápagos hotspot, and its eastward movement; accretion of Galápagos hotspot tracks; Andean uplift; and displacement on the Romeral fault system. All these geological features were active in the Cretaceous, suggesting that geological change at that time caused vicariance in widespread ancestors. The present distributions are explicable if ancestors survived as metapopulations occupying both the Galápagos hotspot and other regions before differentiating, more or less in situ.

Four new species of Splendid Ghost Moths (Lepidoptera: Hepialidae: Aenetus) from Australia and Papua New Guinea.

Four new Aenetus Herrich-Schäffer species are described from northern Australasia; Aenetus simonseni sp. nov. from the top-end of the Northern Territory, Australia, A. maiasinus sp. nov. from the Kimberley region of Western Australia, A. trigonogrammus sp. nov. from south-eastern Queensland, Australia, and A. albadamanteum sp. nov. from eastern Papua New Guinea. Aenetus simonseni sp. nov. and A. maiasinus sp. nov. appear to belong to the tegulatus-group of species (sensu Grehan et al. 2018), A. trigonogrammus sp. nov. is part of the splendens-group of species (sensu Simonsen 2018), while A. albadamanteum sp. nov. shares morphological similarities with A. hampsoni (Joicey Noakes, 1914), A. crameri Viette, 1956, and A. toxopeusi Viette, 1956, from New Guinea, and A. cohici Viette, 1961 from New Caledonia. The four new species are illustrated and compared with superficially similar species in morphology and, for two species, molecular (mtDNA COI gene) sequences.

Taxonomic revision of Schausiana Viette with two new species from Guatemala and notes on biogeography and correlated tectonics (Lepidoptera: Hepialidae).

We morphologically validate the genus Schausiana Viette, 1950 as a monophyletic group comprising five species-S. phalerus (Druce, 1887) comb. n. and S. trojesa (Schaus, 1901) from Mexico, S. chalciope sp. n. and S. maishei sp. n. from Guatemala and S. pharus (Druce, 1887) comb. n. from Mexico to Costa Rica. Potential morphological monophyly of the genus is supported by the unique shared presence of specialized scales spine-like, elongate spinous, or piliform, located along the forewing veins. We investigate possible close phylogenetic relationships of this genus with the Mexican and Central American genera Pallas C. Mielke Grehan, 2015, and Phassus Walker, 1856, and the southeastern Brazilian genus, Phthius C. Mielke Grehan, 2017. Also discussed are the shared presence of several characters supporting the morphological monophyly of Schausiana, Phassus, and Pallas as 'phassine' Hepialidae. The distribution range of Schausiana is bounded in its northwestern range by the Guerrero terrane. Other than the widespread S. pharus with a disjunct record from Costa Rica, the southern distribution boundary corresponds to the Motagua-Polochic Fracture Zone in Guatemala. We suggest that the ancestral range of Schausiana did not include the Guerrero terrane and was possibly also absent from the Chocos Block (Honduras, El Salvador, Nicaragua) and the Costa Rica-Panama arc of Central America. Further divergence of Schausiana is predicted to have been influenced by tectonic events during late Cretaceous or early Cenozoic time. Colour photos are presented for all species for the first time. Holotypes of S. maishei, sp. n., and S. chalciope, sp. n., are deposited in the Universidad del Valle de Guatemala, Guatemala.

Ghost-moths of Trinidad and Tobago with description of a new genus and a new species (Lepidoptera: Hepialidae).

The Hepialidae species of Trinidad and Tobago are documented and two species are recognized. The new and monotypic genus Wallacella, gen. n., (Lepidoptera, Hepialidae) is erected to include Phassus guianensis Schaus of Guyana (Wallacella guianensis, comb. n.). The male holotype and genitalia are also illustrated for the first time. Potential generic affinities with Druceiella (Viette) and Pfitzneriana (Viette) are considered. Gymelloxes juliusboosi sp. n. is described from Trinidad and Tobago and compared to its congeneric species. The sternite VIII and male genitalia are diagnostic for this new species. The male holotype is deposited in the collection of the Natural History Museum, London, UK.

Morphological notes on Gazoryctra sciophanes (Ferguson) and G. confusus (Edwards) (Lepidoptera: Hepialoidea: Hepialidae).

The genus Gazoryctra Hübner comprises 10 species in North America and four in northern Eurasia. The remaining diversity of North American Hepialidae is represented by four species of Sthenopis Packard, three species of Phymatopus Wallengren, and one species of Korscheltellus Börner (Nielsen et al. 2000; Grehan Knyazev 2019). The North American distribution of Gazoryctra extends between Alaska and southern Appalachians and southern Rocky Mountains (Grehan Mielke 2018). As with other North American Hepialidae, Gazoryctra is absent from much of the southern-central United States where there is ostensibly suitable habitat present as this genus is found in forested regions where it is believed to feed on roots or other organic matter (Schweitzer et al. 2011). This absence may be due to the lack of colonization following regression of inland seas that covered much of this region until the end of the Mesozoic (Grehan Mielke 2018).

Description of Walkeriella miraculosa, gen. n. et sp. n. from South-East Peru (Lepidoptera: Hepialoidea: Hepialidae).

Walkeriella miraculosa gen. n. et sp. n. (Lepidoptera: Hepialoidea: Hepialidae) is based on two specimens collected from rainforest southwest of Puerto Maldonado, Tambopata, Madre de Dios department, southeastern Peru. The distinct genitalia and wing venation separate this taxon from other genera of Hepialidae although four derived characters states are shared in common with the cibyrine clade. The holotype male is deposited in the collection of the Museo de Historia Natural, Lima, Peru.

New species of Endoclita (Lepidoptera: Hepialidae) and revived species status of E. kosemponis from Taiwan.

Two species of Endoclita C. . R. Felder, 1874 from Taiwan are evaluated for their taxonomic status: Endoclita meifenga Buchsbaum Grehan sp. n. is described as new for a unique male from the central mountains of Taiwan, and E. kosemponis (Strand, 1916), stat. rev. is elevated to species status. We show that the forewing markings and genitalic characteristics of E. meifenga sp. n. distinguish this species from all other Endoclita. The external appearance of E. kosemponis is similar to E. sinensis (Moore, 1877) but the two species show distinct differences in the male and female genitalia. Preliminary comparison of E. sinensis from Taiwan and Japan suggests they represent more than one species. A sample of the COI gene was sequenced for E. meifenga sp. n. and the primary types of E. meifenga sp. n. and E. kosemponis sp. n. are illustrated here for the first time. The taxonomic status of the non endemic Taiwanese E. davidi (Poujade, 1886) and E. sinensis needs future evaluation.

A new Endoclita (C. . R. Felder, 1874) from the High Mountains of Taiwan with notes about its evolutionary origins (Lepidoptera: Hepialidae).

A new species Endoclita atayala Buchsbaum Hsu sp. n. is based on a single female specimen collected at light in the high mountains of northern Taiwan. Comparisons with all other Endoclita (C. R. Felder, 1874) species confirms that E. atayala sp. n. exhibits distinctive and unique wing pattern characteristics and probably has a sister group relationship with E. crenilimbata (Le Cerf, 1919) of southeastern China and illustrated here for the first time, and E. nodus (Chu Wang, 1985) of northeastern China. The species diversity of Taiwanese Hepialidae is described and the biogeographic implications of the E. atayala sp. n. relationship for the origin of endemic species in Taiwan are reviewed.

Evolutionary biogeography and tectonic history of the ghost moth families Hepialidae, Mnesarchaeidae, and Palaeosetidae in the Southwest Pacific (Lepidoptera: Exoporia).

The biogeographic history of Exoporia (Lepidoptera) in the Southwest Pacific is reconstructed for genera and species that show distributional boundaries corresponding to tectonic structures in the region. Correlations with tectonic formations of Mesozoic origin such as the Whitsunday Volcanic Province and Otway-Bass-Gippsland Basin system in Australia, the Vitiaz Fracture Zone in northern Melanesia, and the Western Province-Eastern Province boundary, Waitaki Fault Zone, and Waihemo Fault Zone of New Zealand are presented as evidence of an East Gondwana origin for genera and species before the geological separation of Australia and New Zealand. The correlated boundaries also suggest that many extant species retain at least parts of their original East Gondwana distribution ranges. The presence of Exoporia on the northern Melanesian Arc, New Caledonia, and New Zealand is attributed to the tectonic isolation of these areas when East Gondwana expanded into the Pacific following retreat of the Pacific Plate subduction zone. Local endemism of Mnesarchaeidae in New Zealand is interpreted as the result of an original vicariance from a widespread ancestor ('Exoporia') resulting in two allopatric descendants -a narrowly distributed Mnesarchoidea and a widely distributed Hepialoidea. The current overlap of these two groups in New Zealand is explained as the result of subsequent range expansion by the Hepialoidea prior to geological fragmentation of East Gondwana. The potential impact of Cretaceous geography on modern distributions is also considered for Exoporia in southern Africa and northern America. Along with lateral displacement of Exoporia, tectonic processes also contributed to the origin of high elevation endemics through a process of passive tectonic uplift.

New species of Dugdaleiella, gen. nov., Kozloviella, gen. nov., and Pfitzneriella Viette from upper elevation Andes of Ecuador and Peru (Lepidoptera: Hepialidae).

Six new Hepialidae species are described from high elevation habitats in the Andes of Peru and Ecuador. One species is assigned to the genus Kozloviella gen. n., K. viazmenskyi sp. n., from Peru, and five to the genus Pfitzneriella Viette, 1951: P. antonkozlovi sp. n., P. olafi sp. n., P. titarenkoi, sp. n., P. yuliyakovalevae sp. n. from Peru and P. rawlinsi sp. n., from Ecuador. Dugdaleiella gen. n., is proposed to accommodate the Ecuadorian species Dugdaleiella monticola (Maassen, 1890) comb. n., formerly placed in Pfitzneriella. The taxonomic (but not the phylogenetic) status of P. lucicola (Maassen, 1890) is unaltered pending future opportunity to examine the type series. Preliminary phylogenetic analysis of the morphological characters supports Kozloviella gen. n., and Pfitzneriella as sister taxa, and separates P. titarenkoi sp. n., from the other species of Pfitzneriella. We suggest that the ancestor of these high elevation genera already occupied a pre-Andean upland habitat across Peru and Ecuador and this ancestor further diverged into the modern genera and species during the Andean orogeny. We consider it likely that there is a considerable diversity of endemic high elevation Hepialidae that remains to be discovered in the eastern Andes between Venezuela and Bolivia.

Re-characterization of Gymelloxes with a re-description of Gymelloxes terea from Central America (Lepidoptera: Hepialidae).

The hepialid genus Gymelloxes Viette, 1952 is characterized by, and differs from all other genera, by the male genitalia and a combination of characters. G. terea is redescribed due to the brevity of the original description. The male genitalia of Gymelloxes terea (Schaus, 1892) include two features that are unique within the Hepialidae - a posteriorly angled and digitiform tergal lobe, and two shallow, peg-like processes on the posterior margin of the saccus. The male phallus also exhibits minute sclerotized spicules or spots on the apex of an otherwise membranous tube. The external genitalia of the female includes a sclerotized antevaginalis with a broad, dorsally projecting, flat central margin. The species is included within the cibyrine clade of Hepialidae by the structure of the tergosternal connection and narrow spacing between Sc and R on the hindwing. The potential systematic significance of specialized similarities shared with other cibyrine Hepialidae is discussed. A lectotype of Dalaca terea Schaus, 1892 is here designated.

Mona Lisa smile: the morphological enigma of human and great ape evolution.

The science of human evolution is confronted with the popular chimpanzee theory and the earlier but largely ignored orangutan theory. The quality and scope of published documentation and verification of morphological features suggests there is very little in morphology to support a unique common ancestor for humans and chimpanzees. A close relationship between humans and African apes is currently supported by only eight unproblematic characters. The orangutan relationship is supported by about 28 well-supported characters, and it is also corroborated by the presence of orangutan-related features in early hominids. The uniquely shared morphology of humans and orangutans raises doubts about the almost universal belief that DNA sequence similarities necessarily demonstrate a closer evolutionary relationship between humans and chimpanzees. A new evolutionary reconstruction is proposed for the soft tissue anatomy, physiology, and behavioral biology of the first hominids that includes concealed ovulation, male beard and mustache, prolonged mating, extended pair-bonding, "house" construction, mechanical "genius," and artistic expression.