David Penney

The University of Manchester · Faculty of Life Sciences

The previous issue of Palaeoentomology brought the first set of papers honoring David A. Grimaldi on the occasion of his 65th birthday. With the current one, it is continued, in recognition of his impact on the fields of amber studies, palaeontology, palaeo- and neoentomology, and evolutionary biology. After the success of Jurassic Park (both the M...

With 557 named spider species in 202 genera and 58 families, Baltic amber represents the richest source of fossil spiders anywhere on the planet, with research describing this remarkable fossil fauna dating back more than 200 years. The identification of numerous species is problematic because many of the older descriptions are inadequate by curren...

Spiders are the most diverse and important terrestrial predators in modern ecosystems. Therefore, fossil spiders are fundamental to understanding past terrestrial ecosystems, especially coevolution with their principal prey, the insects. Being generally soft bodied, spiders have a poor fossil record, but where they do occur, it is in the exceptiona...

Fossils in amber provide a unique insight into the evolutionary history of arthropods (insects, spiders and their relatives), the most diverse and ecologically important group of animals on the planet today. The fossils provide direct evidence of many groups familiar to us now, living alongside extinct forms such as the dinosaurs. Furthermore, the...

to ascertain, through analytical means, the impact of microCT scanning and confocal microscopy (CLSM) on different amber and copal types.

Palaeontology is a rather unique career choice, in that at one point or another most kids (and some adults) want to be a palaeontologist, but very few of them are ever able to realize their dream. However, there are more avenues into palaeontology related work today than there ever have been and the main purpose of this book is to highlight some of...

Within the last few decades technological advances such as the Internet and new imaging techniques (e.g., computed tomography and synchrotron scanning) have revolutionized how we do research in the 21st century. High-quality publications resulting from international, interdisciplinary collaborations are now commonplace, though there is still much s...

Keroplatid fungus gnats (Diptera: Keroplatidae) were hitherto known from the Caribbean island of Hispaniola from a single named species in Miocene amber from the Dominican Republic. Here we describe the new fossil species Proceroplatus preziosii Evenhuis & Penney based on a mature male specimen, which differs markedly from both fossil and extant ta...

A new specimen of Lacinius Thorell, 1876; (Opiliones: Phalangiidae) from Eocene Baltic amber is described. We interpret it as conspecific with a slightly younger record from the German Bitterfeld amber, originally referred to as the extant species L. erinaceus Staręga, 1966. Our new specimen reveals pedipalpal apophyses on both the patella and the...

Fossil insects in amber are often preserved with life-like fidelity and provide a unique insight to forest ecosystems of the geological past. Baltic amber has been studied for more than 300 years but despite the large number of described fossil species (ca. 3500 arthropods) and abundance of fossil material, few attempts have been made to try and qu...

Palaeoentomology represents the interface between two huge scientific disciplines: palaeontology – the study of fossils, and entomology – the study of insects. However, fossils rarely feature extensively in books on insects, and likewise, insects rarely feature in books about fossils. Similarly, college or university palaeontology courses rarely ha...

Although µ-CT and CLSM are exceptionally valuable for the investigation of amber inclusions, they have the potential to physio-chemically alter amber. Thus, access to collections for investigations utilising novel methodologies is currently hampered by a lack of understanding of their potential harmful effects on amber samples, which may consist of...

Insects preserved in copal, the sub-fossilized resin precursor of amber, have potential value in molecular ecological studies of recently-extinct species and of extant species that have never been collected as living specimens. The objective of the work reported in this paper was therefore to determine if ancient DNA is present in insects preserved...

Over the last few decades the internet, increased computing capabilities, molecular techniques and new imaging technologies have altered the way we do research in the 21st century. The result is a great increase in multi-disciplinary research and novel avenues incorporating spiders as model organisms. In short, we are in a new era of research, bein...

The development of second generation sequencing technology has resulted in the rapid production of large volumes of sequence data for relatively little cost, thereby substantially increasing the quantity of data available for phylogenetic studies. Despite these technological advances, assembling longer sequences, such as that of entire mitochondria...

Keroplatid fungus gnats (Diptera: Keroplatidae) were hitherto known from the Caribbean island of Hispaniola from a single named species in Miocene amber from the Dominican Republic. Here we describe the new fossil species Proceropla-tus preziosii Evenhuis & Penney based on a mature male specimen, which differs markedly from both fossil and extant t...

Inclusions in amber and copal provide us with a unique insight into terrestrial palaeocommunities because they represent a palaeobiocoenosis: a naturally co-occurring group of organisms that perished at they same point in time and in the same place. We report the first (sub)fossilized example of a spider population, preserved in Colombian copal, wh...

Amber and copal are renowned for preserving insects and other inclusions with lifelike fidelity. However, due to their frozen-in-time nature, they are taxonomically subequal to Recent insects, which commonly require dissection in order to identify them to species level. This can be overcome to a degree through digital dissection using computed tomo...

The Gambia, located on the coast of tropical West Africa supports a diverse range of habitat types, from forests to mangrove swamps, savannah grasslands and coastal ecosystems. It has a rich and diverse flora and fauna, which for some groups such as birds and mammals is relatively well known. However, for groups such as arachnids and most insect or...

A new spider species of the extant genus Craspedisia Simon (Araneae: Theridiidae) is described from a fossil in Miocene amber from the Dominican Republic. X-ray computed tomography (CT) was used to reveal important features that were impossible to view using traditional microscopy, because of the position in which the spider is preserved. Craspedis...

Summary A fossil pirate spider (Araneae, Mimetidae) is documented in amber from the Eocene Cambay Formation of India. It represents the first described spider from this deposit and the oldest fossil record of the family, extending its known geological range by approximately 3–8 million years from the previously oldest fossils in Eocene Baltic amber...

AbstractA new species of the cricket genus Proanaxipha Vickery & Poinar (Orthoptera: Gryllidae: Pentacentrinae) from Early Miocene Dominican amber is described and illustrated. Proanaxipha madgesuttonae sp. n. is distinguished from congeners by: (1) head capsule bearing a distinctive posteriorly bilobed colour spot on the vertex; (2) presence of cr...

X-ray computed tomography is used to identify a unique example of fossilized phoresy in 16 million-year-old Miocene Dominican amber involving a springtail being transported by a mayfly. It represents the first evidence (fossil or extant) of phoresy in adult Ephemeroptera and only the second record in Collembola (the first is also preserved in amber...

Spiders (Araneae) are one of the most species-rich orders on Earth today, and also have one of the longest geological records of any terrestrial animal groups, as demonstrated by their extensive fossil record. There are currently around 1150 described fossil spider species, representing 2.6% of all described spiders (i.e. extinct and extant). Data...

Fossil arachnids date back more than 400 million years to the Silurian period, making them one of the first animal groups to appear in terrestrial ecosystems. However, despite significant progress in the discovery and description of fossil arachnids in recent years there has been no basic overview of the arachnid fossil record published in a broadl...

For the kingdom Animalia, 1,552,319 species have been described in 40 phyla in a new evolutionary classification. Among these, the phylum Arthropoda alone represents 1,242,040 species, or about 80% of the total. The most successful group, the Insecta (1,020,007 species), accounts for about 66% of all animals. The most successful insect order, Coleo...

High-resolution phase-contrast X-ray computed tomography (CT) reveals the phoretic deutonymph of a fossil astigmatid mite (Acariformes: Astigmata) attached to a spider's carapace (Araneae: Dysderidae) in Eocene (44-49 Myr ago) Baltic amber. Details of appendages and a sucker plate were resolved, and the resulting three-dimensional model demonstrate...

Compared to insects, fossil spiders have received only scant attention in the literature. Previously, the only works available were numerous scientific papers, many published in foreign languages. Most of these are basic descriptive taxonomic works, with very few considering broader biological concepts. Despite a significant increase in the discove...

A new species of fossil lace bug, Phatnoma mattijoae sp. nov., is described from a single specimen in Miocene amber from the Dominican Republic. It represents the first fossil record of the genus, extending its known range by approximately 16 million years. The new fossil appears to be most closely related to the extant P. ovatum Champion, 1897, fr...

The new fossil family Grandoculidae fam. n. is erected for a Cretaceous (Campanian) Canadian amber spider Grandoculus chemahawinensis formerly placed tentatively in the extinct family Lagonomegopidae. Following the descriptions of several new lagonomegopid genera from widespread fossil deposits, it is now clear that their overall somatic morphology...

Computed tomography (CT) methods were applied to a problematic fossil spider (Arachnida: Araneae) from the historical Berendt collection of Eocene (ca. 44-49 Ma) Baltic amber. The original specimens of Ocypete crassipes Koch and Berendt 1854 are in dark, oxidised amber and the published descriptions lack detail. Despite this, they were subsequently...

Fossils preserved in amber provide us with a unique insight into the biodiversity and ecology of long extinct tropical forests. Today, similar habitats contain a major proportion of the planet’s biodiversity, although they are disappearing at an alarmingly rapid rate, primarily as a result of human activities such as deforestation. The hot, humid e...

Genetic differences among tree species, their hybrids and within tree species are known to influence associated ecological communities and ecosystem processes in areas of limited species diversity. The extent to which this same phenomenon occurs based on genetic variation within a single tree species, in a diverse complex ecosystem such as a tropic...

A new species of the extant spider family Anapidae is described from a fossil mature male in Eocene amber from the Baltic region and tentatively assigned to the genus Balticoroma Wunderlich, 2004. Phase contrast X-ray computed micro-tomography was used to reveal important features that were impossible to view using traditional microscopy. Balticoro...

A new species of brown lacewing (Insecta: Neuroptera: Hemerobiidae) is described from Eocene Baltic amber. Sympherobius siriae sp. nov. is the second fossil species of the genus so far described. The other, Sympherobius completus Makarkin et Wedmann is also from Baltic amber. The fossil record of Hemerobiidae is reviewed.

Kinchloe Roberts et al. (2008) proposed a technique, using outline-morphometric and linear analyses to permit identification of spider compression fossils to family level. This work focussed on fossil spiders from the Florissant Formation, Colorado, but the aim was to investigate a technique that would be broadly applicable to any compression fossi...

Propagation-based phase contrast imaging allows low absorbing materials to be studied non-destructively using hard xrays, generating additional edge contrast through interference effects. Here we will present recent work undertaken at Manchester University's "Henry Moseley X-ray Imaging Facility" studying the applicability of lab-based phase contra...

The beauty and mysteries of amber have fascinated humankind since its first discovery. Specimens with fossil inclusions inside were particularly revered and still are today, although for very different reasons. Fossils in amber represent a unique insight into subtropical and tropical forest ecosystems of the past. Today, such habitats contain a maj...

The first fossil Molinaranea is described, from middle Miocene Dominican amber. This record extends the known range of the genus back 16 million years; it also extends the geographical range of the genus through time, with extant species known only from Chile, Argentina, the Falkland Islands, and Juan Fernandez Island. A parsimony-based phylogeneti...

Conflicting opinions regarding the relative importance of dispersal versus vicariance in understanding Caribbean biogeography continue to stimulate lively debate, despite recent advances in regional tectonics, geology, palaeogeography and palaeontology. The Greater Antilles are young geographical features, probably mid-Miocene and Hispaniola has a...

Over the last three decades, the fossil record of spiders has increased from being previously biased towards Tertiary ambers and a few dubious earlier records, to one which reveals a much greater diversity in the Mesozoic, with many of the modern families present in that era, and with clearer evidence of the evolutionary history of the group. We he...

The spiders and other arachnids of West Africa are poorly known and this is particularly true of the Gambian fauna. At least nine different arachnid orders are now known to occur in the country and all of them are included in this introductory work. They include spiders, harvestmen, mites and ticks, camel spiders, scorpions, pseudoscorpions, taille...

The butterflies of West Africa are generally well known, but there are no country-by-country field guides for the interested amateur naturalist. The butterfly fauna of The Gambia is not particularly diverse in a West African context, but it does contain interesting elements of both Sahelian, Sudanian and Guinean Savannah species, because the countr...

The Plectreuridae is a relatively small, ecribellate, haplogyne spider family consisting of only two extant genera. Kibramoa Chamberlin has seven described species restricted to the USA and Mexico (Gertsch 1958) and Plectreurys Simon is known from 22 species, with similar distributions (Gertsch 1958, Jiménez 2006), but also including Cuba and Costa...

We provide a brief report on the feeding behavior of two salticid species in Bijilo Forest, The Gambia: Holcolaetis vellerea Simon 1909 and Menemerus bivittatus (Dufour 1831). The former was observed consuming a giant huntsman spider ?Heteropoda sp. (Sparassidae), which was much larger than itself and represents the first published evidence of aran...

. The purpose of this application, under Article 23.9.3 of the Code, is to conserve the generic name Myrmarachne MacLeay, 1839 for a well-known and widespread group of extant, ant-mimicking jumping spiders (Araneae, salticidae) by suppression of a little-used senior subjective synonym, Entomocephalus Holl, 1829, introduced for a fossil spider prese...

Ricinulei is an arachnid order of great antiquity with fossils of the extinct families Curculioididae and Poliocheridae known from the Pennsylvanian (c. 300Ma) of Europe and North America (Selden 1992). However, extant species (Ricinoididae) are known only from Africa (Ricinoides) and South and Central America (Cryptocellus and Pseudocellus), with...

A new fossil Linyphiidae: Linyphiinae is described from 125–135 Ma old (Upper Neocomian–basal Lower Aptian) Cretaceous amber from the Kdeirji/Hammana outcrop, Lebanon. This is the oldest known linyphiid as well as the oldest described amber spider. The first major radiation of the linyphiid subfamilies occurred in the early Cretaceous, if not befor...

The spider family Lagonomegopidae was described a decade ago from two specimens in Upper Cretaceous Siberian amber from the Taimyr Peninsula, and placed in the superfamily Palpimanoidea. Lagonomegopidae is known only from Cretaceous amber. Undiscovered extant species are considered unlikely because of their frequent occurrence in Cretaceous ambers...

Afrarchaea grimaldii new species (Archaeidae, Archaeinae) from 88–95 Ma (Cenomanian–Turonian) Upper Cretaceous amber (Burmite) from Myanmar (Burma) is described. This is the first spider to be described from this deposit and is the oldest known Archaeidae sensu stricto extending the known range of the family by approximately 50 Ma from the previous...

A new fossil spider, Palaeoperenethis thaleri gen. nov. and sp. nov., from Middle Eocene (44–52 Ma) strata from Horsefly, British Columbia, Canada, is described in the family Pisauridae. The localities in the Okanagan Highlands are well known for their insect fauna preserved together with fish and plants in varved diatomaceous lacustrine sediments....

The species-level diversity of fossil Chelicerata is summarized for each order. 1952 valid species of fossil chelicerates are currently recognized. of which 1593 are arachnids. In order of abundance they are: Araneae (979 fossil species). Actinotrichida (283). Eurypterida (241), Scorpiones (111), Xiphosura (96), Trigonotarbida (71), Pseudoscorpione...

The author is a Visiting Research Fellow at the University of Manchester, UK and the leading world expert on fossil spiders preserved in amber and in interpreting what they can reveal about the ecology of the extinct forests in which they lived. In this book Dr Penney uses a unique, integrated approach, which combines and compares information deriv...

A fossil tarantula (Araneae: Mygalomorphae: Theraphosidae) is described from an exuvium in Tertiary (Miocene) Chiapas amber, Simojovel region, Chiapas State, Mexico. It is difficult to assign it further taxonomically, but it is the first mygalomorph recorded from Chiapas amber and only the second unequivocal record of a fossil theraphosid. With a c...

There are many influences on how assessors grade themselves and others. Oral presentations are useful for exploring such factors in peer, self‐ and tutor marked assessments, being rapidly completed and assessed, commonly used in HE and very difficult to anonymize. This opportunistic study examined the effects of gender and level of attainment on th...

When most people hear the word ‘fossil’ they tend to conjure up images of giant dinosaurs such as Tyrannosaurus rex or shelled marine invertebrates. Prior to the Hollywood blockbuster movie Jurassic Park, which was based on recreating dinosaurs through extracting their DNA from fossil mosquitoes preserved in amber, few non-palaeontologists would en...

We apply Very-High-Resolution X-Ray Computed Tomography (VHR-CT) to a minute fossil spider (~1 mm long) from Eocene amber of the Paris Basin, France. We demonstrate that the newly described genus and species of Micropholcom-matidae, Cenotextricella simoni, retains excellent details of the somatic and male pedipalpal morphology that allows unqualifi...

We apply Very-High-Resolution X-Ray Computed Tomography (VHR-CT) to a minute fossil spider (~1 mm long) from Eocene amber of the Paris Basin, France. We demonstrate that the newly described genus and species of Micropholcommatidae, Cenotextricella simoni, retains excellent details of the somatic and male pedipalpal morphology that allows unqualifie...

The Wealden Group of southern England is a succession of non-marine sediments from the lower Cretaceous (c. 141–119 Ma), which also outcrop in Belgium and northwest Germany. Most insect orders are present as fossils, including Neuroptera, one of the most basal holometabolous orders. The fossil record of Neuroptera is diverse, spanning Permian–Recen...

A new spider species from the family Oonopidae is described from lowermost Eocene amber from Le Quesnoy, Oise department, Paris Basin, France. Orchestina parisiensis sp. n. is the first spider to be described from French amber. Preliminary observations on the fossil spider assemblage support a warm climate in this region during the lowermost Eocene...

Two new spiders, Quamtana huberi new species (Pholcidae) and Selenops sp. indet. (Selenopidae), are described from Lowermost Eocene (Ypresian) amber from Le Quesnoy, Oise department, Paris Basin, France. Both specimens represent the oldest known fossils of their respective families. This is the first fossil record of the extant genus Quamtana, exte...

Fossils preserved in amber provide a unique insight into terrestrial forest palaeoenvironments and also to micro- and macro-evolutionary processes. They may also prove useful in predicting the consequences of current global climate change. In order to appreciate the potential these amazing fossils have to offer, we need to understand the processes...

The aerial orb web woven by spiders of the family Araneidae typifies these organisms to laypersons and scientists alike. Here we describe the oldest fossil species of this family, which is preserved in amber from Alava, Spain and represents the first record of Araneidae from the Lower Cretaceous. The fossils provide direct evidence that all three m...

The new species Burlagonomegops alavensis (Araneae: Lagonomegopidae) is described from Lower Cretaceous (Aptian) amber from Álava (Basque Country), Spain. This is the first fossil spider to be described from this deposit and extends the known geological range of this family by approximately 15¿20 Ma, from the previously oldest described lagonomegop...

The first description of a Mermithidae-Linyphiidae parasite-host association is presented. The nematode is preserved exiting the abdomen of the host, which is a juvenile Tenuiphantes species (Araneae, Linyphiidae), collected from the Isle of Mull, UK. An updated taxonomic list of known mermithid spider hosts is provided. The ecology of known spider...

  The Neotropical fossil amber Hersiliidae are revised. Prototama Petrunkevitch is a senior synonym of both Priscotama Petrunkevitch and Neotama Baehr and Baehr. Fictotama Petrunkevitch and Perturbator Petrunkevitch are removed from synonymy and are considered nomina dubia because the fossils are poorly preserved. The following new combinations are...

To justify faunistic comparisons of ambers that differ botanically, geographically and by age, we need to determine that resins sampled uniformly. Our pluralistic approach, analysing size distributions of 671 fossilized spider species from different behavioural guilds, demonstrates that ecological information about the communities of two well-studi...

The first fossils of the extant New Zealand spider family Huttoniidae are described from Cretaceous (Campanian) amber from Alberta and Manitoba, Canada. The specimens are juveniles and poorly preserved, but the following combination of characters permits identification as huttoniids: general habitus, carapace without a raised cephalic region or fov...

  The spider family Oonopidae is described from Cretaceous ambers from Myanmar and Canada for the first time. Orchestina albertenis sp. nov. is the first spider to be described from Canadian Grassy Lake amber and only the second spider to be described from Canadian amber. The specimen in amber from Myanmar extends the known range of the extant genu...

The project Assembling the Tree of Life (AToL)—Phylogeny of Spiders is an ambitious, collab-orative, six-year project, which aims to construct a robust cladogram for all spider families. The resulting phylogeny will be based on morphological, molecular, behavioural and palaeontological data. Fossil spiders are not considered in current systematic c...

  Two spiders (Filistatidae) in Miocene Dominican Republic amber, one newly identified and only the second known fossil of this family, have autospasized legs (detached at a predetermined locus of weakness when restrained by a non-self-induced source) at the patella-tibia joint. In both specimens, droplets of haemolymph (blood) are preserved exitin...

Fossils preserved in amber represent only a small fraction of the biota which was alive in the amber forest. In order to make accurate palaeoenvironmental reconstructions, it is necessary to determine the biases which affect preservation. Only the Baltic and Dominican Republic amber deposits have had inclusions described in sufficient quantities su...

Misionella didicostae new species is described from 15-20 Ma Miocene amber from the Dominican Republic as the first fossil record of the family Filistatidae. The biogeography of the extant (Brazil and Argentina) and the new fossil species supports the hypothesis that the developing northern Greater Antilles and northwestern South America were brief...

Peer assessment provides a useful mechanism to develop many positive qualities in students studying in higher education (HE). Potential influences on peer‐awarded marks include student qualities such as gender, HE background (e.g. university affiliation) and participation in the development of the assessment criteria. Many studies that have investi...

The new species Floricomus fossilis (Araneae: Linyphiidae) is described from Miocene Dominican Republic amber. This is the first fossil record of Floricomus, extending its known geological range by 15¿20 Ma, and is the first record of the genus outside North America and Canada. Extant species may exist on Hispaniola, given the similarities between...

Penney, D. 2004. Cretaceous Canadian amber spider and the palpimanoidean nature of lagonomegopids. Acta Palaeonto− logica Polonica 49 (4): 579–584. The first formally described spider from mid−Campanian (76.5–79.5 Ma), Upper Cretaceous amber from Cedar Lake, Manitoba, Canada is named as Grandoculus chemahawinensis new genus and species. It belongs...

The oldest pisaurid spider Palaeohygropoda myanmarensis gen. et sp. nov. (Araneae: Pisauridae) is described from 100–107 Mya (Albian) Cretaceous amber (Burmite) from Myanmar (Burma). This specimen extends the known range of the family by approximately 60 My from the previously oldest record in Baltic amber. It predicts the presence of the extant sp...

The oldest described fossils of the extant spider family Araneidae (Araneinae; gen. et sp. indet.), the extant genus Orchestina (Oonopidae; O. sp. indet.) and the new fossil genus Palaeosegestria (Segestriidae; P. lutzzii gen. et sp. nov.) are presented from Upper Cretaceous amber of New Jersey. The known fossil range of the extant family Araneidae...