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Introduction
I am an evolutionary biologist who studies evolutionary patterns and processes in arachnids (spiders and their kin). My research focuses on the diversity of spiders in the Rocky Mountain/Great Plains ecoregion (the Colorado Spider Survey). I am also involved with projects exploring the taxonomy, systematics, biogeography, and natural history of spiders, scorpions, and solifuges. My lab page is at http://www.dmns.org/science/museum-scientists/paula-cushing/
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February 1998 - present
Publications
Publications (127)
Background/Objectives: Arachnids are a megadiverse arthropod group. The present study investigated the chromosomes of pedipalpid tetrapulmonates (orders Amblypygi, Thelyphonida, Schizomida) and two arachnid orders of uncertain phylogenetic placement, Ricinulei and Solifugae, to reconstruct their karyotype evolution. Except for amblypygids, the cyto...
Cobweb spiders, or Theridiidae, with just over 2500 species, are the fourth largest spider family. They are usually small to medium-sized species that can occur worldwide in all terrestrial habitats. The common name cobweb spider refers to their web type. Occasionally, they are also called tangle-web spiders, but this is a rather vague term (who kn...
The yellow sac spider family has a turbulent history, as the classic yellow sac spiders, which are primarily discussed here, have changed their family affiliation three times in the last 30 years. Today, they form their own family, as was already proposed back in 1887. Almost 400 species are currently counted among 14 genera of Cheiracanthiidae.
The family of tube-dwelling spiders, Segestriidae, comprises around 180 species worldwide. They are predominantly small to medium-sized spiders that have only six eyes (which you might only see at a second glance, Fig. 4.9), have a cylindrical body structure, and betray themselves through a characteristic leg position. Three pairs of legs are direc...
Nearly 13% of all spider species are jumping spiders (Salticidae). With over 6500 species, they are the most species-rich family of all. Jumping spiders are found throughout the world and are abundant in all terrestrial habitats. They do not build capture webs but stalk their prey before jumping on it to overpower it (hence their name). This is fol...
Ground spiders are generally poorly known, as they are nocturnal and usually live quite cryptically. They are a group of spiders distributed worldwide and, with almost 2500 species, represent the sixth largest family. They are small to medium-sized spiders, most of which are colored in brown and black tones, often with parts of the body covered in...
With nearly 300 species, Uloboridae is a species-poor family of spiders, predominantly found in the subtropics and tropics, with only a few species reaching into the temperate zone. These predominantly small animals of 3–10 mm body length have, as a very special feature among spiders, completely reduced their venom glands. The reasons for this are...
Linyphiidae comprise nearly 5000 species worldwide, making them the second most common spider family after the jumping spiders. They are web-building spiders with a body length of 1–8 mm (although most species are only 1–3 mm), which are recognizable by their horizontal sheet webs. Ground-dwelling linyphiids build their tiny sheet webs in crevices...
Dictynidae, with around 500 species, is a globally distributed family of predominantly small spiders. Mesh-weavers belong to the cribellate spiders, which incorporate mesh threads (Chap. 1) into their small webs. These webs usually appear unstructured and are mostly deployed on plants, often around the spider’s retreat.
Funnel-web spiders are a family of medium to large (up to 18 mm) spiders, with almost 1400 species in 100 genera worldwide. They are web-building animals, and their web form also gives them their English name. Their web consists of a retreat open on both sides, with its edge widening on one side to form a catching sheet, so that the whole structure...
When we, as spider researchers, talk to laypeople about our favorite animals, two completely different reactions are normal: (1) Ah, these are the animals that build those beautiful webs (they always mean orb webs); or (2) Oh my goodness, these are the big, ugly, hairy creatures that lurk in our bathrooms and are even venomous. We, therefore, regul...
In this chapter, we offer various identification aids to determine the house spiders discussed here. We will not go to the species level but will be able to resolve their family affiliation. Such identification keys differ in their difficulty depending on the spider species and your prior knowledge. For this reason, in addition to a classic dichoto...
Apartments and houses have special climatic conditions, which allow a small group of spider species to live in our buildings. Through global goods transport, we have spread many of our house spiders throughout the world, so we often encounter the same species everywhere. There is no reason to be afraid of them, yet we describe how to remove spiders...
Hackled mesh-weavers, Amaurobiidae, are a small spider family of just under 300 species. These rather small to medium-sized spiders are predominantly dark in color and prefer moist habitats. Hackled mesh-weavers belong to the cribellate spiders, meaning that they produce ultrafine crimped threads (Chap. 1). These cribellate threads give the web a b...
Goblin spiders are a family of tiny spiders that grow to 1–2 (rarely 3) mm in size. Currently, almost 2000 species in 115 genera are known. These short-legged spiders are predominantly nocturnal and do not build capture webs but actively hunt small insects. Oonopids are found almost worldwide but have a clear focus in the tropics and subtropics of...
The small family includes about 200 species of six-eyed spiders. Included in this family is the genus Loxosceles, commonly referred to as recluse spiders or violin spiders (because of the marking on their forebody). These spiders are small to medium-sized, and their six eyes are arranged in three groups (Fig. 4.10). Recluse spiders naturally occur...
Giant crab spiders, also called huntsman spiders, with almost 1500 species, are a diverse family of predominantly large, free-living spiders. The similarity to crab spiders, which is reflected in their common name, is only superficial. The two families are not closely related. Their names just refer to similar movement patterns. Giant crab spiders...
One could argue that spitting spiders are a rather insignificant family. Currently, only about 240 species are known, which, however, occur throughout the world. They are small to medium-sized and have six eyes, which are arranged in three groups (Fig. 4.3). Scytodids stand out due to their long, thin legs and the noticeably large forebody. This is...
Cellar spiders are a family of web-building spiders found throughout the world. Currently, around 2000 species in almost 100 genera are known. Cellar spiders are easily recognized by their unusually long legs, which is why they are often confused with harvestmen. Harvestmen, however, are not spiders but represent their own order within the arachnid...
Woodlouse hunters are a spider family that today comprises a little more than 600 species in 25 genera. They have only six eyes, which are grouped at the front of the body (Fig. 4.12). The front and hind body are elongate. They are small to medium-sized species that occur in many habitats, where they are mostly nocturnal hunters on the ground, in l...
False wolf spiders are a small spider family, with only 181 known species. They resemble wolf spiders (hence the name) and hunt their prey freely without a web. However, unlike wolf spiders, they are cribellate spiders, meaning they produce a very fine, crimped spider silk known as cribellum wool (Chap. 1). However, this is no longer used to build...
The species-poor family Desidae currently comprises just under 330, mostly medium-sized species and is unusual for three reasons. First, the habitat of many Desidae is in the tidal zone of the world’s oceans. Spiders, as emphasized in every spider book, occur (with the exception of one species) only on land. This assessment seems to have overlooked...
Disk web spiders are a small family of 6 genera with only 125 species, which have a distribution focus in the tropics but are also regularly found in the subtropics. Three-quarters of these species belong to the genus Oecobius. They prefer to live under stones but also on vertical structures such as rocks and tree bark, and they have many adaptatio...
Mimetids are an exciting spider family of only 160 species, but they present a highly unusual prey specialization. These are rather small species that occur throughout the world but have a clear distribution focus in the tropics. A hint to their lifestyle is the conspicuous spikiness of the forelegs, which can be used to capture particularly danger...
The orb-weaver spider family is the third most species-rich in the world and includes over 3100 species. This includes exceptionally colorful, pretty spiders and, in addition to small to medium-sized animals, also some quite large species. Orb-weavers are found throughout the world in almost all habitats. A number of species have also become adapte...
Crevice weavers, Filistatidae, are a small family of nearly 200 species worldwide. They are very well adapted to dry habitats; thus, they are mainly found in deserts and semi-desert regions of the world. Since buildings in human settlements have a certain climatic similarity to the original habitat of Filistatidae, we now find them throughout the w...
A survey of 100 spider experts from various countries and an analysis of the specialist literature provided a wealth of information on the occurrence of spiders in buildings. Based on this, the most common spider species in buildings throughout the world were selected, each of which is presented here with a global distribution map.
Morphology has long been used to classify and identify living organisms. However, taxonomic descriptions are often limited to qualitative descriptions of size and shape, making identification difficult due to the subjective language used to describe complex shapes. Additionally, for some taxa, there are few reliable qualitative characters available...
Advanced sequencing technologies have expedited resolving higher-level arthropod relationships. Yet, dark branches persist, principally among groups occurring in cryptic habitats. Among chelicerates, Solifugae (“camel spiders”) is the last order lacking a higher-level phylogeny and thus, historically characterized as “neglected [arachnid] cousins”....
Considerable progress has been achieved in resolving higher-level relationships of Arthropoda in the past two decades, largely precipitated by advances in sequencing technology. Yet, dark branches persist in the arthropod tree of life, principally among groups that are difficult to collect, occur in cryptic habitats, or are characterized by minute...
The spider Myrmecicultor chihuahuensis Ramrez, Grismado & Ubick (Myrmecicultoridae) was described in 2019 and hypothesized to be a myrmecophile, living inside the nests of Novomessor (Myrmicinae) and perhaps also Pogonomyrmex (Myrmicinae) ants. To test the hypothesis that M. chihuahuensis are chemical mimics of their host ants, we carried out behav...
Species of camel spiders in the family Eremobatidae are an important component of arthropod communities in arid ecosystems throughout North America. Recently, research demonstrated that the evolutionary history and biogeography of the family are poorly understood. Herein we explore the biogeographic history of this group of arachnids using genome-w...
Camel spiders, members of the order Solifugae, are a diverse but poorly understood order of arachnids. The conserved morphology within many groups in the order makes species delimitation and identification challenging. In the North American family Eremobatidae, swollen setae called ctenidia are a common character employed in species delimitation, d...
The Southern Appalachians and adjacent provinces of the southeastern USA are geologically and biologically diverse, with high levels of endemism. Phylogeographic analyses indicate that animals with small distributions in these regions often contain cryptic diversity and that Pleistocene climate fluctuations had significant impacts on their distribu...
Phylogenetic relationships and evolutionary patterns within the arachnid order Solifugae are poorly understood and largely unresolved due to conserved morphology and scarce genomic resources. In this study, we evaluated the role of restriction endonuclease (RE) selection in double-digest restriction-site-associated DNA sequencing (ddRADseq) as a me...
Gravity is very important for many organisms, including web-building spiders. Probably the best approach to study the relevance of gravity on organisms is to bring them to the International Space Station. Here, we describe the results of such an experiment where two juvenile Trichonephila clavipes (L.) (Araneae, Nephilidae) spiders were observed ov...
Little is known about the reproductive behaviour of solifuges, or camel spiders (Arachnida: Solifugae). Behavioural research is limited, due in part to challenges of maintaining specimens in the laboratory where they can be best observed. The present study documents the courtship and mating behaviours of a North American solifuge, Eremobates pallip...
Entry for recent Encyclopedia of Social Insects about myrmecomorphs.
Research progress on the order Solifugae, commonly known as camel spiders, has been hindered by challenges inherent in collecting these fast-moving, nocturnal predators. Recently, pitfall trapping combined with artificial light lures showed promise for improving capture rates, but the hypothesis that camel spiders are attracted to light traps (posi...
The new genus and species Myrmecicultor chihuahuensis Ramírez, Grismado, and Ubick is described and proposed as the type of the new family, Myrmecicultoridae Ramírez, Grismado, and Ubick. The species is ecribellate, with entelegyne genitalia, two tarsal claws, without claw tufts, and the males have a retrolateral palpal tibial apophysis. Some morph...
The Solfugae fauna of Canada includes three known species: Eremobatesdocolora Brookhart and Muma, E.scaber (Kraepelin), and Hemerotrechadenticulata Muma. It is expected that as many as four additional species may be found in Canada. Only one Barcode Index Number is currently known from Canadian specimens.
Camel spiders (Arachnida: Solifugae) are a notoriously difficult group of arachnids to study. They are almost all strictly nocturnal, fast moving predators that are difficult to find, collect, and rear. In this paper, we present methods for both attracting solifuges in desert field sites and collecting them efficiently using a combination of light...
A recent phylogenetic analysis demonstrated that the solifuge genus Eremocosta (Eremobatidae), although not monophyletic, formed a strongly supported group, rendered paraphyletic by the exclusion of E. acuitlapanensis, which we herein determine is misplaced in this genus. We revise the taxonomy of the genus Eremocosta. Nine species of the 13 curren...
Recent syntheses of phylogeographical data from terrestrial animals in the Mojave and Sonoran deserts have revealed a complex history of geologic and climatic vicariance events. We studied the phylogeography of Smeringurus vachoni to see how vicariance events may have impacted a large, endemic rock scorpion. Additionally, we used the phylogeographi...
Nine new species of the Eremobates scaber species group of the solifuge genus Eremobates Banks 1900 are described, eight of them from Mexico. These new species are: E. axacoa, E. bonito, E. cyranoi, E. fisheri, E. hidalgoana, E. jalis-coana, E. minamoritaana, E. zacatecana, and E. zapal and together increase the size of this species group to 23. A...
The Denver Museum of Nature & Science Reports (ISSN 2374-7730 [print], ISSN 2374-7749 [online]) is an open-access, non peer-reviewed scientific journal publishing papers about DMNS research, collections, or other Museum related topics, generally authored or co-authored by Museum staff or associates. Peer review will only be arranged on request of t...
The Nearctic agelenid spider genus Agelenopsis Giebel 1896 is revised, with redescriptions of the 13 known species including: A. actuosa (Gertsch & Ivie 1936), A. aleenae Chamberlin & Ivie 1935, A. aperta (Gertsch 1934), A. emertoni Chamberlin & Ivie 1935, A. kastoni Chamberlin & Ivie 1941, A. longistyla (Banks 1901), A. naevia (Walckenaer 1841), A...
Some male camel spiders (Arachnida: Solifugae) in the families Eremobatidae, Karschiidae, and Solpugidae have clusters of specialized conical or acuminate setae called papillae, on the ventral surface of the metatarsus of the pedipalps. We compared the overall structure of the papillae found on representatives of the three families using scanning e...
The Denver Museum of Nature & Science Annals (ISSN 1948-9293 [print], ISSN 1948-9307 [online]) is an open-access, peer-reviewed scientific journal publishing original papers in the fields of anthropology, geology, paleontology, botany, zoology, space and planetary sciences, and health sciences. Papers are either authored by Museum staff, associates...
We describe the mimetic relationship between the ant-like spider Myrmecotypus iguazu Rubio & Arbino 2009 (Araneae: Corinnidae) and the carpenter ant Camponotus sericeiventris Guérin (Hymenoptera: Formicidae), studied in a subtropical rainforest in Iguazú National Park, Argentina. The morphological adaptations, aspects of coloration, and behavior re...
Spiders are critical predators in agroecosystems. Crop management practices can influence predator density and diversity, which, in turn, can influence pest management strategies. Crop intensification is a sustainable agricultural technique that can enhance crop production although optimizing soil moisture. To date, there is no information on how c...
This paper provides a summary of the extensive theoretical and empirical work that has been carried out in recent years testing the adaptational significance of various spider-ant associations. Hundreds of species of spiders have evolved close relationships with ants and can be classified as myrmecomorphs, myrmecophiles, or myrmecophages. Myrmecomo...
Solifuges, or camel spiders (order Solifugae), keep their pedipalps extended when moving through the environment, utilizing them much the way insects use their antennae. The male also uses his pedipalps during copulation, staying in contact with the female throughout the process. The pedipalps are covered with setae that are assumed to function as...
Accurate characterization of predator-prey linkages in agroecosystems is important prior to the implementation of conservation biological programs. The Russian wheat aphid, Diuraphis noxia (Hemiptera: Aphididae), is a significant pest of wheat and barley in the United States. This research utilized molecular gut-content analysis as a minimally disr...
Accurate characterization of predator-prey linkages in agroecosystems is important prior to the implementation of conservation biological programs. The Russian wheat aphid, Diuraphis noxia (Hemiptera, Aphididae), is a significant pest of wheat and barley in the United States. This research utilized molecular gut-content analysis as a minimally disr...
Juvenile solifuges have rarely been observed hunting in natural conditions. We recorded the hunting behavior of juvenile third or fourth instar solifuges of the genus Chanbria (Eremobatidae) near lanterns set up in the Imperial Sand Dunes, Imperial County, California. At least 10 juveniles were observed between 22:50 and 01:40 h on 18—19 June 2010....
When disturbed, Uloborus glomosus either remain in position at the hub of their orb webs, jump from the web, move to the edge of the web, or shake the web. Juveniles more frequently exhibited moving and jumping responses, whereas the majority of adults jumped from the web or remained in position. Adults with linear stabilimenta tended to shake thei...
Two species of Chinchippus (Ammotrechidae) were studied in central Peru. Both species are endemic to the hyper-arid coastal desert and appear to derive most of their energy and nutrients from maritime prey, such as intertidal amphipods feeding on beach-cast algae or as arthropod scavengers feeding upon seabird and pinniped carcasses. Data on the sp...
The ant hunting behavior of Zodarion rubidum (Araneae, Zodariidae) is described from specimens collected in Colorado, USA. Like other members of this genus, Z. rubidum constructs igloo-shaped stone retreats under rocks and feeds on ants. Details of the prey capture behavior are provided including initial and subsequent reactions of ants to the bite...