Evidence for Carboniferous origin of the order Mantodea (Insecta: Dictyoptera) gained from forewing morphology

Homologies of the forewing venation pattern of the order Mantodea (Insecta: Dictyoptera) consistent with the accepted insect wing venation groundplan are proposed. A comparative morphological analysis was carried out based on a broad taxonomic sample of extant taxa. Besides macromorphological aspects, focus is given to the pattern of the tracheal system as a basis for establishing primary homologies. All extant praying mantids exhibit a composite stem composed of the posterior radius (RP) and the media (M) and most praying mantids exhibit a fusion of the anterior branch of RP + M with the anterior radius (RA). The wing venation of the species ?Mesoptilus dolloi, previously assigned to the polyphyletic fossil assemblage ‘Protorthoptera’, is re‐interpreted in the light of the new homology statement. Our interpretation suggests that it is a putative stem‐Mantodea, as are some other ‘protorthopterous’ taxa. This hypothesis implies that the total‐group Mantodea arose as soon as the Late Carboniferous, i.e. about 175 million years earlier than previously estimated. This analysis contributes to the view that most of the Late Carboniferous ‘Protorthoptera’ are stem‐representatives of the major polyneopteran clades (e.g. cockroaches, grasshoppers and crickets, rock‐crawlers), suggesting a survivorship of several main Pterygota lineages at the end‐Permian extinction event higher than previously expected. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 79–113.     

Fossil lice (Insecta: Phthiraptera) reconsidered

Abstract.  Five papers describing fossils considered to be Phthiraptera (lice) by their authors are reviewed. We place the specimens described in three papers in the Acari (mites) and regard them as probably not fossils. A fourth paper describes what appears to be a fossil insect. However, we do not consider it to be a louse. The fifth paper describes a more recent fossil that differs little from extant lice.     

Afrotropical and Nearctic genera of Odonata in the French Oligocene: biogeographic and paleoclimatic implications (Insecta: Calopterygidae,Aeshnidae)

New species of the genera Sapho and Epiaeschna are recorded in the Oligocene of Aix- en-Provence, Bouches-du-Rhône, France. Sapho legrandi n. sp. is the third fossil representative of this recent African genus and Epiaeschna pseudoheros n. sp. is the fifth fossil species of this recent North American genus. The fossil species Triaeschna gossi from the Eocene of England, Epacantha magnifica from the Late Oligocene of Kazakhstan, and Mediaeschna matutina from the Oligocene of China, are considered species of Epiaeschna and the three fossil genera Triaeschna Campion 1916, Mediaeschna Zhang 1989, and Epacantha Martynov 1929 are synonymized with Epiaeschna. The closely related genera Umma and Sapho inhabit warm humid forests of Western Africa. Their presence in two Oligocene deposits of France supports the hypothesis of a warm humid palaeoenvironment for Armissan (Aude, France), and Aix-en-Provence.     

Systematic methods,fossils, and relationships within Heteroptera (Insecta)

Three recent papers dealing with phylogenetic relationships within the Heteroptera are discussed and analysed. A character set representing 43 taxa and 78 characters is used to test theories presented in those papers. The conclusions of Grimaldi and Engel concerning the placement of the Cretaceous fossil taxon Cretopiesma in the Piesmatidae are rejected in favour of placement in the Aradidae. The placement by Nel et al. of Protodoris from Eocene amber of the Paris Basin in the Thaumastocoridae is considered ambiguous because it has none of the diagnostic characters of that family group and is therefore regarded as incertae sedis. The arguments of Sweet concerning the elevation of the Aradoidea to infraordinal status on the basis of autapomorphies are also treated as invalid. General arguments against the use of phenetic methods in palaeontology, and ad hoc approaches under the guise of cladistics, are offered, with the conclusion that rigorous cladistic analyses are a prerequisite to testable conclusions concerning the placement of fossil and Recent taxa. © The Willi Hennig Society 2009.     

The insect trace fossil Tonganoxichnus from the middle Pennsylvanian of Indiana: Paleobiologic and paleoenvironmental implications

The ichnogenus Tonganoxichnus, produced by one or more monuran insect taxa, is now recorded from the Middle Pennsylvanian Mansfield Formation of Indiana. Tonganoxichnus is a resting trace that has three important implications. First, it represents a recurrent behavioral pattern in Upper Carboniferous to Lower Permian marginal marine environments of North America. Second, it provides finely resolved anatomical information for axial and appendicular body structures and behaviors that are difficult to determine from body‐fossil material alone. Third, integrated sedimentologic and ichnologic observations indicate that the Tonganoxichnus assemblage, inclusive of other ichnotaxa, is common in tidal rhythmites that were developed under freshwater conditions, probably in the innermost part of estuarine systems, close to or at the fluvioestuarine transition.     

New and poorly known Cenozoic clown beetle compressions from France (Insecta: Coleoptera: Histeridae)

Summary

Two French fossil Histeridae are described or revised, viz. Hister cerestensis n. sp. from the Oligocene of Céreste (Lubéron) and Plegaderus pitoni (Theobald, 1935), n. comb. from the Pliocene of Chambon Lake (Puy-de-Dôme). Morphometric data are used to compare fossil and extant taxa, with multivariate statistics. A list of the fossil Histeridae is given.     

Saccoglossus testa from the Mazon Creek fauna (Pennsylvanian of Illinois) and the evolution of acorn worms (Enteropneusta: Hemichordata)

The limited fossil record of enteropneust hemichordates (acorn worms) and the few external features that distinguish the four families have provided a challenge to our understanding of the evolution of the group and their various feeding adaptations. The middle Pennsylvanian Saccoglossus testa sp. nov. from the Mazon Creek, Westfalian D Carbonate Formation, Francis Creek Shale of northern Illinois provides evidence for the exploitation of surface sediments. Saccoglossus testa has a long proboscis characteristic of the extant genus Saccoglossus, a specialist in surface deposit feeding. The collar is as long as it is wide. The anterior trunk lacks a distinctively wide branchial region. These three features distinguish it from its sympatric enteropneust species Mazoglossus ramsdelli Bardack that has a proboscis characteristic of an infaunal deposit feeder. It is the seventh known species of fossil enteropneust, including a resting trace of a Lower Triassic fossil that has collar lips that characterize the extant deep‐sea family Torquaratoridae, and which represents a second parallel evolution of surface deposit feeding. An analysis of the seven fossils shows that the earliest Enteropneusta had a relatively simple harrimaniid‐like body plan, and that the spengelid, the torquaratorid and lastly the most complex ptychoderid body plan appeared in that chronological order.     

Recent genus Notonecta (Insecta: Heteroptera: Notonectidae) in the Lower Cretaceous of San Luis,Argentina: Palaeoecological implications

We describe a new species of the genus Notonecta, i.e. Notonecta mazzoniae n. sp. from the Lower Cretaceous of Argentina. It is the first fossil representative of the family described from South America, demonstrating the great antiquity of the genus, bearing identical ventral chaetal system correlated with respiratory for such a long period. The new species is recorded only by immature stages from the second to the fifth instars. Specimens are considered to be exuviae and not body fossils. Absence of body fossils may be due to taphonomic bias instead of palaeoecological constrains. In addition we found some specimens of Anisopinae, also considered as nymph exuviae, which represent the oldest record of the taxon.     

A case of mistaken identification of an amber fossil male strepsipteran (Insecta: Strepsiptera)

A well-preserved specimen of amber Strepsiptera was erroneously described as Stylops neotropicallis Kogan and Poinar, 2010. The taxonomic position of the species was based on a count of six antennomeres (typical of the Stylopidae); however, further observations showed that there are actually seven antennomeres, which places the fossil in the family Myrmecolacidae, and it is herein redescribed in the genus Palaeomyrmecolax Kulicka, 2001. Furthermore, nuclear magnetic resonance spectroscopy analysis of a sample of the amber piece containing the fossil revealed that it was Baltic rather than Dominican amber as originally thought. The fossil was compared with another specimen from Baltic amber in the Poinar collection, which shows close affinity to Palaeomyrmecolax succineus Kulicka, 2001, the type species of the genus. Palaeomyrmecolax neotropicallis (n.comb.) differs from that specimen and seems to differ also from the other four species in the genus Palaeomyrmecolax.     

Mesozoic Trachypachidae (Insecta: Coleoptera) from China

Abstract: Four well‐preserved beetles, attributed to Eodromeinae within Trachypachidae, are described from the Mesozoic of Inner Mongolia, China. Eodromeus robustus sp. nov., E. daohugouensis sp. nov. and Unda chifengensis sp. nov. are from the Middle Jurassic of Daohugou. The diagnostic characters for the two genera are revised, and all species of these genera are keyed. A new genus and species, Sinodromeus liutiaogouensis gen. nov., sp. nov., is described from the Lower Cretaceous Yixian Formation of Liutiaogou. The current fossil records of Trachypachidae from China are reviewed, and only five species (four described here) can be convincingly attributed to this family. The morphological disparity of Mesozoic Eodromeinae suggests that they evolved a broad spectrum of locomotory lifestyles.     

STUDIES OF CARBONIFEROUS FUNGI. II. THE STRUCTURE AND ORGANIZATION OF MYCOCARPON,SPOROCARPON, DUBIOCARPON,AND COLEOCARPON (ASCOMYCOTINA)

The problematic organism, Sporocarpon, was among the first fossil fungi to be described. The genus was actually a heterogeneous assemblage of fungal forms, and was eventually divided into several taxa including Sporocarpon, Dubiocarpon, and Mycocarpon. In the present paper these organisms, as well as a similar, previously undescribed genus, Coleocarpon; are described from Lower, Middle, and Upper Pennsylvanian strata in North America, and from Upper Carboniferous sediments in England. Although initially considered to be radiolarian-like protozoa, these structures show far more similarity to ascomycetous cleistothecia. All forms are ornamented, spherical bodies with a complex wall which encloses scattered asci and ascospores. Taxa are distinguished primarily by differences in the organization of the cleistothecial wall. Structurally similar organisms are found today among the Eurotiales.     

Phylogeny of ensign scale insects (Hemiptera: Coccoidea: Ortheziidae) based on the morphology of Recent and fossil females

The Ortheziidae (ensign scale insects) is a morphologically well‐defined family. The morphology and occurrence in the fossil record suggests a probable early origin of the family in scale insect evolution. The present phylogenetic analysis – based on 69 morphological characters of female ortheziids, using 39 exemplar Recent species – provides the first analytical assessment of relationships among living genera of the family, as well as the relationships of eight fossil species, based on complete, well‐preserved specimens in amber. Monophyly of the subfamilies Newsteadiinae, Ortheziinae and Ortheziolinae is supported, but Nipponortheziinae is found to be paraphyletic by inclusion of the Ortheziolinae. Thus, the subfamily Ortheziolinae is reduced in rank to tribe Ortheziolini stat.n. , which now includes Matileortheziola Kozár & Foldi, Ortheziolacoccus Kozár, Ortheziolamameti Kozár and Ortheziola?ulc. Consequently, the tribes Matileortheziolini, Ortheziolacoccini and Ortheziolamametini are synonymized ( syn.n. ) here under Ortheziolini. Five new species and one new genus of fossil ensign scales are described from three amber deposits: Burmorthezia gen.n. with type species Burmorthezia kotejai sp.n. and also B. insolita sp.n ., both in mid‐Cretaceous Burmese amber (98 Ma) and Arctorthezia baltica sp.n. in Eocene Baltic amber (c. 43 Ma) based on second‐instar nymphs; Mixorthezia kozari sp.n . and M. dominicana sp.n . in Miocene Dominican amber (c. 17 Ma) based on adult females. Fossil placements are unambiguous, with Burmorthezia forming a stem to crown‐group (Recent and Tertiary) Ortheziidae. A summary of described fossil ortheziids is provided.     

Reiterative growth in the complex adaptive architecture of the Paleozoic (Pennsylvanian) filicalean fern Kaplanopteris clavata

Reiteration is a widespread component of plant growth whose evolutionary importance in ferns is not recognized widely. We introduce and discuss the growth architecture of Kaplanopteris clavata, a fossil filicalean fern from the Pennsylvanian (ca. 305 million yeas old), focusing on types of reiteration exhibited by this species and on the adaptive and phylogenetic significance of reiteration for ferns in general. Kaplanopteris clavata combines two types of reiterative growth where growth modules are borne on fronds: (1) entire fronds derived from primary pinnae, and (2) epiphyllous plantlets. This combination of reiterative pathways, unique among fossil and living ferns, allowed K. clavata to explore ecospace through an opportunistic combination of scrambling, climbing and epiphytic growth. Kaplanopteris clavata underscores the organographic importance of fronds (as opposed to stems) in the adaptive architecture of ferns, emphasizing functional convergences between the different Baupla̋ne of ferns and angiosperms. This unique combination of reiterative pathways is interpreted as a derived condition illustrating the structural and developmental complexity achieved by some filicaleans during the first major evolutionary radiation of leptosporangiate ferns.     

The phylogeny of Ephemeroptera in Pterygota revealed by the mitochondrial genome of Siphluriscus chinensis (Hexapoda: Insecta)

The mayfly species Siphluriscus chinensis (Siphluriscidae) has valuable structures useful for phylogeny reconstruction, given its putative basal position within the Ephemeroptera. Here its nearly complete mitochondrial genome is sequenced. We built phylogenetic trees through multiple analytical strategies with some other insect mitogenomes. Structurally, the obtained mitochondrial genome of S. chinensis is 16,616 bp in length, ¹ containing 37 genes and an extra trnK-like (trnK2 (AAA)) gene. The 12 PCGs start with typical ATN codons, except the nad1 gene which starts with an unnormalized TTG. Like other known mayfly mitogenomes, the strand bias has negative AT-skew and negative GC-skew. Phylogenetically, our topologies suggest that Odonata is the basally diverged clade in Pterygota; Ephemeroptera is the sister group of the Neoptera; and S. chinensis is indeed the most basal mayfly branch.     

Mesoderm formation in a springtail,Tomocerus ishibashii Yosii (Collembola : Tomoceridae)

The mesoderm formation of Tomocerus ishibashii (Collembola : Tomoceridae) is described. Mesodermal cells are formed after the beginning of the formation of the primary dorsal organ, and originate from the entire region of the embryonic area. After completion of the blastodermic cuticles, cells of mesoderm and ectoderm concentrate towards a ventral midline and form a well-defined 2-layered germ band. The manner of mesoderm formation in the Collembola is similar to that in Diplura and Myriapods, except for the Chilopoda; the mesoderm of the Thysanura s. lat. and Pterygota originates from a localized zone of the embryo. Within the Hexapoda, mesoderm formation is categorized into 2 types: Type 1—unlocalized origin, in the Collembola and Diplura, and Type 2—localized origin, in the Thysanura s. lat. and Pterygota. Types 1 and 2 are thought to be plesiomorphic and apomorphic, respectively.     

The fossil record of Glypturus Stimpson, 1866 (Crustacea,Decapoda, Axiidea,Callianassidae) revisited,with notes on palaeoecology and palaeobiogeography

Abstract: The fossil record of the callianassid genus Glypturus (Decapoda, Axiidea) is re‐evaluated. Our systematic revision, both of extant and fossil taxa, is based on major cheliped morphology only, thus providing an important impetus for palaeontological studies. Both spination and tuberculation of chelipeds are herein considered of great taxonomic importance. Presence of spines on the upper margins of the merus and propodus and the lower margin of the carpus are significant for generic assignment, whereas the extent of tuberculation on lateral surfaces of the propodus is important for assignment at the species level. Altogether, four extant and six exclusively fossil species of Glypturus are recognized. Several extinct callianassid taxa are now transferred to the genus, namely Callianassa berryi, Callianassa fraasi, Callianassa munieri, Callianassa pugnax and Callianassaspinosa; Callianassa pseudofraasi is considered a junior synonym of C. fraasi. Based on a comparison of ecological preferences of extant representatives, the presence of Glypturus in the fossil record is considered to be linked with tropical to subtropical, nearshore carbonate environments of normal salinity. We argue that Glypturus is of Tethyan origin, with a stratigraphical range going as far back as the Eocene. Since then, the genus migrated both westwards and eastwards, establishing present‐day communities in the western Atlantic and Indo‐West Pacific which both comprise several distinct species. In the presumed area of origin, the genus does no longer occur today. The exlusively fossil (middle Eocene) genus Eoglypturus from Italy is considered closely related to Glypturus and is thus assigned to the subfamily Callichirinae as well.     

STUDIES OF PALEOZOIC MARATTIALEANS: AN EARLY PENNSYLVANIAN SPECIES OF THE FERTILE FERN SCOLECOPTERIS

Remains of the fossil Marattiales are very rare in Lower Pennsylvanian sediments. The present report describes a new species of the fertile fern foliage Scolecopteris from the Lewis Creek, Kentucky locality (Lower or lower Middle Pennsylvanian). Scolecopteris conicaulis n. sp. has radial synangia composed of a ring of 4–7 elongate, exannulate sporangia. Most features of the synangia of S. conicaulis were previously hypothesized to be primitive in Scolecopteris based on geologically younger species. Supposed primitive characters include the large synangium pedicel with fiber core, an outer-facing sporangial wall lacking differentiation or zonation, and large spores. The anatomy of the sporangium walls, pinnule morphology, and general spore type support an association with the Minor group of Scolecopteris. The new species is similar in several important features to Scolecopteris (Cyathotrachus) altus, the only other anatomically preserved fertile marattialean known from this early time, and indicates a considerably earlier origin for fertile foliage of this type.     

New griffenfly,Bohemiatupus elegans from the Late Carboniferous of western Bohemia in the Czech Republic (Odonatoptera: Meganisoptera: Meganeuridae)

Abstract

A new griffenfly, Bohemiatupus elegans n. gen., n. sp. (Meganeuridae) is described from the Upper Carboniferous (Bolsovian) deposits of the Ov?ín near Radnice in western Bohemia (Czech Republic). The new taxon based on fore- and hindwing venation is compared with the other meganeurid genera. It is the first record of a large griffenfly from the continental basins of the Bohemian Massif supplementing the other giant insects such as Bojophlebia prokopi Kukalová-Peck 1985 or Carbotriplura kukalovae Kluge 1996 from the same strata.     

A new Serpukhovian (Mississippian) fossil flora from western Argentina: Paleoclimatic, paleobiogeographic and stratigraphic implications

An uppermost Mississippian (Serpukhovian) fossil flora from western Argentina is described for the first time. Its particular composition includes both, warm climate plants typical from the Pennsylvanian of Gondwana, namely pteridosperms, sphenophytes as well as arboreous lycopsids; and small lycopsids characteristic of cooler climates of pre-Serpukhovian times. A warm–temperate climate is inferred for this association, which fills a time gap between those Pennsylvanian and Mississippian classic floras previously known. The presence of Tomiodendron, Nothorhacopteris kellaybelenensis and Paracalamites allow the recognition of the Paraca Floral Realm in Argentina, supporting the existence of a nonglacial paleoclimate period, developed just before of the Bashkirian–Moscovian Gondwanan glaciation and after a latest Visean glaciation. It likewise would indicate the paleogeographic position of south west Gondwana, which would have acceded to a warm temperature latitudinal belt during Serpukhovian times. The recognition of this floral assemblage has relevant biostratigraphic and lithostratigraphic implications for the western margin of Gondwana. It defines a new zone for western Argentina, herein named Frenguellia eximia–Nothorhacopteris kellaybelenensis–Cordaicarpus cesarii (FNC) zone, which is recorded in the oldest known lithostratigraphic unit from the Paganzo Basin, the Loma de los Piojos Formation nom. nov.