New Fossil Scorpion from the Chiapas Amber Lagerst?tte

A new species of scorpion is described based on a rare entire adult male preserved in a cloudy amber from Miocene rocks in the Chiapas Highlands, south of Mexico. The amber-bearing beds in Chiapas constitute a Conservation Lagerstätte with outstanding organic preservation inside plant resin. The new species is diagnosed as having putative characters that largely correspond with the genus Tityus Koch, 1836 (Scorpiones, Buthidae). Accordingly, it is now referred to as Tityus apozonalli sp. nov. Its previously unclear phylogenetic relationship among fossil taxa of the family Buthidae from both Dominican and Mexican amber is also examined herein. Preliminarily results indicate a basal condition of T. apozonalli regarding to Tityus geratus Santiago-Blay and Poinar, 1988, Tityus (Brazilotityus) hartkorni Lourenço, 2009, and Tityus azari Lourenço, 2013 from Dominican amber, as was Tityus (Brazilotityus) knodeli Lourenço, 2014 from Mexican amber. Its close relationships with extant Neotropic Tityus-like subclades such as ‘Tityus clathratus’ and the subgenus Tityus (Archaeotityus) are also discussed. This new taxon adds to the knowledge of New World scorpions from the Miocene that are rarely found trapped in amber.     

A remarkable new fossil species of Amplectister with peculiar hindleg modifications (Coleoptera: Histeridae): further evidence for myrmecophily in Cretaceous clown beetles

Myrmecophily is a phenomenon of the symbiosis of organisms that depend on various ant (Formicidae) societies. Such interspecies associations are found in several unrelated lineages within the clown beetle family Histeridae. Recent studies have suggested that the origin of myrmecophily can be traced back to mid-Cretaceous based on a few fossil records from Kachin amber from northern Myanmar. Here, we describe a remarkable new species, Amplectister terapoides n. sp., from Kachin amber. This is the second species of the extinct genus Amplectister Caterino and Maddison, which has been found from the same amber deposit and has also been considered to be myrmecophilous. The new species here described has the most heavily modified hindlegs in any fossil histerids or even beetles discovered until now, indicating further evidence for ant colony association. Our discovery demonstrates that significant and diverse morphological adaptations to myrmecophily had already occurred during the Cretaceous.     

How long can insect species exist? Evidence from extant and fossil Micromalthus beetles (Insecta: Coleoptera)

Six new fossils of Micromalthus (Coleoptera: Archostemata) from Dominican amber are compared with extant and previously described fossil micromalthid beetles. The amber inclusions are well preserved and all important morphological features are visible. Taking into account the morphological variability of the extant species Micromalthus debilis LeConte, 1878 , it is not possible to find any morphological features that distinguish the fossils from the extant species. This also applies to the Dominican amber inclusion described as Micromalthus anasi Perkovsky, 2008, which therefore is considered a junior synonym of M. debilis. The lack of morphological change in M. debilis over time might possibly be explained by unusually stable environmental conditions, as this species occupies a very specialized ecological niche in decaying timber. A general survey of fossil insects indistinguishable from extant species is presented. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 300–311.     

The earliest fossil record of the extant rove beetle genus Phloeocharis from mid-Cretaceous Kachin amber of northern Myanmar and its biogeographic implications (Coleoptera: Staphylinidae: Phloeocharinae)

Phloeocharis Mannerheim is the largest genus within the problematic rove beetle subfamily Phloeocharinae, with a single extinct and 44 recent species recorded from the Holarctic Region. Until now, the oldest fossil record of Phloeocharis was known from Late Cretaceous (Turonian) amber from New Jersey, USA. Here we describe 2Phloeocharis burmana n. sp. from mid-Cretaceous (Albian–Cenomanian) Kachin amber from northern Myanmar, as the earliest extinct species of this genus. Our finding also sheds light on the biogeography of Phloeocharis, since no recent or extinct species have so far been recorded from the Oriental Region. Furthermore, the discovery of 2P. burmana n. sp. extends the Mesozoic diversity of the phloeocharine rove beetles both taxonomically and morphologically, particularly from Kachin amber.     

Phylogeny of the North American catfish family Ictaluridae (Teleostei: Siluriformes) combining morphology,genes and fossils

We performed the first combined‐data phylogenetic analysis of ictalurids including most living and fossil species. We sampled 56 extant species and 16 fossil species representing outgroups, the seven living genera, and the extinct genus ?Astephus long thought to be an ictalurid. In total, 209 morphological characters were curated and illustrated in MorphoBank from published and original work, and standardized using reductive coding. Molecular sequences harvested from GenBank for one nuclear and four mitochondrial genes were combined with the morphological data for total evidence analysis. Parsimony analysis recovers a crown clade Ictaluridae composed of seven living genera and numerous extinct species. The oldest ictalurid fossils are the Late Eocene members of Ameiurus and Ictalurus. The fossil clade ?Astephus placed outside of Ictaluridae and not as its sister taxon. Previous morphological phylogenetic studies of Ictaluridae hypothesized convergent evolution of troglobitic features among the subterranean species. In contrast, we found morphological evidence to support a single clade of the four troglobitic species, the sister taxon of all ictalurids. This result holds whether fossils are included or not. Some previously published clock‐based age estimates closely approximate our minimum ages of clades.     

Lobule shape evolution in Radula (Jungermanniopsida): one rate fits all?

The quantification of lobule shape for Radula spp. shows that there is overlap in lobule shape space occupied by subgenera, such that lobule shape does not always reflect relationships. Morphological convergence caused by lineages repeatedly traversing shared regions of morphospace appears commonplace in Radula, and means that many pairs of relatively unrelated species have similar lobule shapes. When observed over time, as in comparisons between fossil and extant species, this may give the impression of stasis if fossil species resemble modern species by chance, independent of their relatedness. This poses a challenge to relating fossils of known age to extant lineages, particularly when fossils are sterile. Significant rate variation between lineages was identified by Adams' Q‐mode analysis, with the fastest subgenus evolving 23 times more quickly than the slowest. Species of subgenus Volutoradula and subgenus Metaradula are apparently over‐dispersed throughout lobule morphospace according to Sidlauskas' method; morphometric branch lengths and hypervolumes in other subgenera can be explained by a stochastic process. In contrast, Bayesian analysis of macroevolutionary mixtures (BAMM) identified a single evolutionary rate as having the highest posterior probability. Consideration of the three independent accessions into auriculate lobule morphospace by Cladoradula and Radula, wherein convergent lobule shapes result from convergent lobule ontogenies and are correlated with bipinnately branched shoot systems and robust primary stems, leads to an ontogenetic hypothesis driven by structural requirements for light interception, under which auriculate lobules are a spandrel. It is speculated that lobules themselves, however, may be a key innovation facilitating radiation into microsites devoid of or depauperate in fungal endophytes. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 178 , 222–242.     

On the systematic position of the Early Cretaceous fern genus “Athyrium”

The oldest fossils assigned to Athyrium (mostly based on the sorus morphology) comprise fronds and spores from the Lower Cretaceous of Northeast Asia. However, most molecular dating suggests that extant Athyrium diverged from its sister genus during the Eocene or later, implying that the Cretaceous fossils probably belong to another polypodiaceous taxon. By examining the sorus morphology of extant genera related to the family Athyriaceae, we found that the primary diagnostic feature for assigning the Cretaceous fossils to Athyrium, i.e., the sorus shape, is common to the entire extant family, or plesiomorphic for the genus. As the fronds are more commonly preserved than the reproductive parts, we compared the fossil frond morphology with those of living taxa of the family that is divided into two types. The Cretaceous fossil we examined here bears the frond’s costal groove characters on adaxial side, which is more closely related to that of the Deparia-clade instead of the clade including Athyrium and other genera of the family. The observation is further confirmed by the cladistic analysis using morphological characters. The systematic position of the Early Cretaceous “Athyrium” was resolved as a stem member of the total Athyriaceae using a tip-dating approach with the Fossilized Birth-Death model in a Bayesian framework. Our study suggests that Early Cretaceous fossils previously assigned to Athyrium require taxonomic revision.     

Middle Miocene Cephalotaxus (Taxaceae) from Yunnan,Southwest China,and its implications to taxonomy and evolution of the genus

The infrageneric taxonomy of the genus Cephalotaxus Siebold et Zuccarini ex Endlicher (Taxaceae) is controversial as morphological characters have been doubted to change coherently among species. Although the epidermal features are considered as important characters in the taxonomy of conifers, they have not been systematically studied in this genus. Neogene foliage shoots with well-preserved epidermal structure not only can provide systematic links to the extant species, but also have implications for the taxonomical definition of the extant species by their epidermal features. In this paper, the morphologies of leafy shoots combining epidermal structure of a Neogene Cephalotaxus, which was collected from the middle Miocene of Southwest China, and ten extant species (variants) were studied. Morphology as well as epidermal structures was compared among species (variants) in Cephalotaxus. Based on the comparisons, the taxonomical definition of extant species in genus Cephalotaxus are discussed. Our results indicate that leaf morphological characters in combination with the features of epidermal structure can distinguish species in genus Cephalotaxus. The fossil foliage shoots from the middle Miocene of Southwest China were assigned to a new species, C. maguanensis n. sp., which shows close affinities to the extant C. hainanensis. On the basis of leaf and epidermal information, we suggest there are seven extant species in Cephalotaxus. With global cooling and the uplift of the Qinghai-Tibet Plateau since the middle Miocene, the cooling climate induced the differentiation and southward expansions of C. maguanensis, and its extant offspring, C. hainanensis, is distributed to Hainan Island and the Indo-Chinese peninsula.     

白垩纪中期克钦琥珀毛蚊科(双翅目:毛蚊总科)昆虫一新属新种

毛蚊科是双翅目长角亚目的一类昆虫,其种类丰富,广布世界各地.毛蚊化石记录较为丰富,但往往都是保存于湖相沉积地层中,琥珀中的化石较为罕见.本文根据缅甸北部克钦地区产出的白垩纪中期琥珀中保存的昆虫标本建立了 1新属新种——缅甸白垩叉毛蚊(Cretpenthetria burmensis gen.et sp.nov.),归入...     

The oldest known chironomids of the tribe Tanytarsini (Diptera: Chironomidae) indicate plesiomorphic character states

Taxonomic studies on chironomids of the tribe Tanytarsini recorded in Eocene Baltic amber from the Rovno region have revealed two new species: Archistempellina perkovskyi and Stempellinella ivanovae. Both species display interesting characters supporting evolutionary trends in the tribe. A. perkovskyi nov. sp. combines plesiomorphic characters, thus consolidating the hypothesis that the genus Archistempellina is one of the basal lineages of the tribe Tanytarsini. The antennal flagellum composed of 13 discernible segments found in S. ivanovae nov. sp. is defined as a plesiomorphic character state opposite to a reduced number of flagellomeres in extant species; the nipple-like process on the apex of the gonostylus is recognised as a unique character in the Tanytarsini and a possible homologue of the horn-like tip of the gonostylus known from several extant species of the genus Stempellinella Brundin. A key to determination of all genera and species of the Tanytarsini recorded in the Eocene is presented. Notes on evolutionary trends in the tribe are also provided.     

?Kenyaichthyidae fam. nov. and ?Kenyaichthys gen. nov. – First Record of a Fossil Aplocheiloid Killifish (Teleostei,Cyprinodontiformes)

The extant Cyprinodontiformes (killifishes) with their two suborders Cyprinodontoidei and Aplocheiloidei represent a diverse and well-studied group of fishes. However, their fossil record is comparatively sparse and has so far yielded members of the Cyprinodontoidei only. Here we report on cyprinodontiform fossils from the upper Miocene Lukeino Formation in the Tugen Hills of the Central Rift Valley of Kenya, which represent the first fossil record of an aplocheiloid killifish. A total of 169 specimens - mostly extraordinarily well preserved - and a sample of ten extant cyprinodontiform species were studied on the basis of morphometrics, meristics and osteology. A phylogenetic analysis using PAUP was also conducted for the fossils. Both the osteological data and the phylogenetic analysis provide strong evidence for the assignment of the fossils to the Aplocheiloidei, and justify the definition of the new family †Kenyaichthyidae, the new genus †Kenyaichthys and the new species †K. kipkechi sp. nov. The phylogenetic analysis unexpectedly places †Kenyaichthys gen. nov. in a sister relationship to the Rivulidae (a purely Neotropical group), a probable explanation might be lack of available synapomorphies for the Rivulidae, Nothobranchiidae and Aplocheilidae. The specimens of †K. kipkechi sp. nov. show several polymorphic characters and large overlap in meristic traits, which justifies their interpretation as a species flock in statu nascendi. Patterns of variation in neural and haemal spine dimensions in the caudal vertebrae of †Kenyaichthys gen. nov. and the extant species studied indicate that some previously suggested synapomorphies of the Cyprinodontoidei and Aplocheiloidei need to be revised.     

A new microinvertebrate with features of mites and tardigrades in Dominican amber

From time to time, small, fragile, previously unknown fossil invertebrates are found in specialized habitats. Occasionally, as in the present case, a fragment of the original habitat that existed millions of years ago is also preserved. The present article describes a previously unknown microinvertebrate in Dominican amber that cannot be placed in any group of extant invertebrates. Placed in a new family, genus, and species, the fossil shares characters with both tardigrades and mites, but clearly belongs to neither group. The several hundred fossil individuals preserved in the amber shared their moist, warm habitat with pseudoscorpions, nematodes, fungi, and protozoa. The large number of fossils provided additional evidence of their biology, including their reproductive behavior, developmental stages, and food. While there is no extant group that can accommodate these fossils, and we have no knowledge of any extant descendants, this discovery shows that unique lineages of minute invertebrates were surviving in the mid‐Tertiary.     

How diverse were ferns in the Baltic amber forest?

Diverse temperate forest types and a high atmospheric humidity have recently been suggested for the Eocene source area of Baltic amber. However, ferns are astonishingly rare as inclusions in this amber, which is in contrast to other seed‐free land plants, fungi, and lichens. Moreover, the identities of some of the few described putative fern taxa are dubious, and some fossils were even assigned to the Paleozoic seed fern genera Alethopteris, Pecopteris and to the form genus Sphenopteris containing Paleozoic and Mesozoic fern‐like leaf fossils. Here, we review previously described fern inclusions from Baltic amber and identify further fern‐like leaf inclusions as belonging to the extant angiosperm genus Comptonia (sweet ferns, Myricaceae). We conclude that only one taxon, Matonia striata (Matoniaceae), can with confidence be identified as a Polypodiopsida representative. Although “Pecopteris” humboldtiana is so far only known as sterile foliage, its leaf morphology strongly suggests that also this taxon belongs to the Polypodiopsida rather than to any other tracheophyte lineage. We propose accommodating “Pecopteris” humboldtiana in the new genus Berendtiopteris. “Alethopteris” serrata and “Sphenopteris” phyllocladoides are not to be regarded as evidence of ferns from Baltic amber. Reinvestigation of the holotypes of these two taxa did not reveal to which tracheophyte lineages these fossils belong. We suggest that the scarcity of fern remains from Baltic amber may reflect both a relatively low fern diversity in the source area of the fossil resin, and an absence or rarity of epiphytic and climbing ferns as observed in modern temperate forest ecosystems.     

The phylogeny of Strepsiptera (Hexapoda)

Previous phylogenetic analyses of Strepsiptera have been limited to characters from only males or first instar larvae, and by poor taxonomic sampling. This investigation is the first cladistic analysis to use more than fourfold as many characters as any prior study, and a broader sampling of taxa. The analysis of 189 morphological characters of all stages of representatives of all extant strepsipteran families and characters of adult males of amber fossils results in the following branching pattern: (?Protoxenos+ (?Cretostylops + (?Mengea + (Mengenillidae + (Corioxenidae + (Bohartillidae + (Halictophagidae + (Elenchidae + (?Protelencholax + (Myrmecolacidae + (Callipharixenidae + (Xenidae + Stylopidae)))))))))))). The basal placement of the Baltic amber fossil ?Protoxenos and the Burmese amber fossil ?Cretostylops is well founded. Even though ?Cretostylops is older than ?Protoxenos it is almost certainly not the most basal strepsipteran group but the sister group of a clade comprising the Baltic amber fossil ?Mengea + Strepsiptera s. str. (excl. stemlineage). Monophyly of Mengenillidae, Stylopidia, Stylopiformia s.l., Corioxenidae, Xenidae, and Stylopidae is confirmed. Mengenillidia is paraphyletic (with respect to ?Mengea (Mengeidae)), Elenchidae (with respect to ?Protelencholax) and the genus Stichotrema (with respect to the Baltic amber fossils). Thus Protelencholacidae fam. n. is described, and S. weitschati and S. triangulum are transferred to Palaeomyrmecolax. A ground plan of adult male Strepsiptera is provided and evolutionary interpretations are presented based on the obtained cladograms. © The Willi Hennig Society 2005.     

Early evolution of Cupedidae revealed by a mid‐Cretaceous reticulated beetle from Myanmar (Coleoptera: Archostemata)

Cupedidae, the most species‐rich family of the archaic suborder Archostemata, were abundant, diverse and widespread in the Mesozoic, yet little is known about the early evolution and biogeography. This stems, in part, from a lack of exceptionally preserved fossils from the Mesozoic and of formal phylogenetic study of both extant and extinct taxa. Here we describe and illustrate a new fossil from mid‐Cretaceous Burmese amber, and provide a phylogeny combining both fossils and all known extant genera of Archostemata. A dataset of 43 ingroup taxa and four outgroup taxa based on 110 morphological characters was analysed under parsimony. The results indicate that Priacma LeConte and Paracupes Kolbe, as well as the Cretaceous genera Barbaticupes Jarzembowski et al. and Mallecupes Jarzembowski et al., together form a sister clade to the rest of Cupedidae. Priacma megapuncta sp.n. is attributed to the relict North American Priacma by the presence of distinct subtruncate elytral apices, lateral elytral margins with two rows of sharp teeth, and peculiar fixing epipleural folds near the elytral apices. Our discovery of the first fossil species of Priacma in Burmese amber reveals the antiquity and wider distribution of the genus in the late Mesozoic. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:313565C2‐4F42‐48BD‐8720‐F379DE202868 .     

A Review of the Fossil Record of the Genus Itea (Iteaceae, Saxifragales) with Comments on its Historical Biogeography

Itea is a genus of about 20 species of trees and shrubs that are today native to southeastern North America, eastern Asia, and eastern Africa. In this paper, I review the fossil record of Itea, which is based on four types of fossils: diporate, psilate pollen attributed to Itea or the dispersed pollen genus Iteapollis; carpofossils representing fruits and seeds attributed to Itea europaea; flowers preserved in amber and assigned to Adenanthemum iteoides; and leaf impressions attributed to Itea. The distributions of these fossils indicate that Itea was present in western North America from the early Eocene to Miocene, in eastern North America beginning no later than the early Miocene, and in western Eurasia from the late Eocene to Pliocene. Only one datapoint is known from eastern Asia; it is early Miocene in age. Based on the fossil record, it can be inferred that Itea crossed between continents over both the Bering Land Bridge and North American Land Bridge, and that it reached Africa from Europe via Anatolia. Thus, it is predicted that the sole extant North American species, I. virginica, may be most closely related to the sole extant African species, I. rhamnoides. The potential application of Itea fossils to calibrating phylogenetic trees generated from molecular sequence data is also discussed.     

A new species and subgenus of Desmanthodium (Compositae,Helianheae) from southern Mexico

Desmanthodium congestum, a new species from Chiapas, Mexico, is described and illustrated. Its morphological features are clearly distinguishable from other species of the genus, so much so that this taxon forms a new subgenus withinDesmanthodium, subgenusMultiaggregata.     

Early lineages of Vespidae (Hymenoptera) in Cretaceous amber

Three wasp (Hymenoptera: Vespidae) fossils in Cretaceous amber (Late Albian) of northern Myanmar are described. Two are new species of the Mesozoic genus Curiosivespa (Rasnitsyn): C. zigrasi sp.n. and C. striata sp.n. The third species, Protovespa haxairei gen.n. et sp.n. , has a combination of features unique among Mesozoic Priorvespinae and the extant subfamilies. These well preserved fossils provide new morphological data for a cladistic analysis of the basal lineages of Vespidae. Results suggest that Euparagiinae is the sister group of all other Vespidae. The new genus Protovespa appears more closely related to extant Masarinae, Eumeninae and social wasps than to Priorvespinae. We assign it to a new subfamily: Protovespinae. Finally, fossil information combined with a phylogenetic tree shows that the main groups of Vespidae probably evolved during the Early Cretaceous. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:1E7E4796‐6E70‐4D81‐BB34‐0FEEA765DC25 .