North American camel spiders (Arachnida, Solifugae, Eremobatidae): Systematic revision and biogeography of an understudied taxon


Project Summary



Solifuges (Arachnida: Solifugae), commonly called camel spiders or wind scorpions, are an important group of arachnids found in fragile, often threatened desert ecosystems. They are notoriously difficult to study since they are hard to find and collect (there is a reason they are called wind scorpions), nearly impossible to keep alive in the lab, challenging to identify to species due to limited diagnostic characteristics, and essentially impossible to raise from hatching through maturity in the lab. They are the sixth most diverse order of arachnids; yet there are few scientists worldwide studying any aspect of their biology and only the PI’s lab is focused on this order. One of the experts in the field, Jack Brookhart (a collaborator on this grant) is long retired. Thus, there is a desperate need to train the next generation of scientists in the biology and taxonomy of this large taxon. This project will use a recently published phylogeny of one of the three largest families of solifuges, the Eremobatidae (188 described species), to completely revise the taxonomy of all clades and reassess the phylogeny using Next Generation Ultra-Conserved Elements (UCEs) along with morphological characters, including novel and unexplored setal characters. This grant will allow us to leverage the minimal expertise that still exists to excite a new generation of scientists who will move this field of study forward. Grant support will also allow us to create an innovative and dynamic online identification guide, the Camel Spiders of North America, through the NSF-funded Symbiota Collections of Arthropods Network (SCAN) data portal that will provide a resource for a much broader audience and will support the development and expansion of a website focused on this order (


Intellectual Merit


This study will address systematic and taxonomic challenges in the solifuge family Eremobatidae by leveraging the expertise in the PI’s and co-PI’s labs to train students in cutting edge phylogenetic, taxonomic, and biogeographic methodology. We will utilize information from a recently published phylogenetic analysis of this large family to revise the taxonomy of well supported clades. We will test this phylogeny by using an expanded approach combining molecular and morphological data. We will utilize a newly designed genomic toolkit for arachnids incorporating hundreds of conserved DNA regions called ultraconserved elements, or UCEs. This novel approach allows targeted sequencing at both deep and shallow levels, allowing us to assess the results of the previous multi-locus phylogenetic analysis. We will also explore the utility of morphological characters, including the chaetotaxy of the chelicerae and pedipalps. The existing literature suggests that these setal characters may be diagnostic for solifuge taxa but their phylogenetic utility has not been previously assessed. In addition, we will use one species group to examine the roles of Neogene tectonics and Pleistocene/Holocene climate change on the biogeography and diversification of camel spiders in southwestern North America.


Broader Impacts


We will use newly developed molecular probes (UCEs) as well as a new set of underutilized morphological characters to reassess the phylogeny of this large family, revise the taxonomy of all taxa included in the Eremobatidae, evaluate the impact of past climate change on the biogeography of one group, and provide innovative online resources on the biology, taxonomy, and natural history of eremobatids for a large audience. To achieve these goals, we will train 6 – 12 high school students from demographic groups underrepresented in STEM fields (women, African Americans, Hispanics, and Native Americans), 12 - 24 undergraduate students, two graduate students, and one early career postdoctoral scholar. This project will train these students in modern systematic, taxonomic, and biogeographic methodologies. We will also develop online resources that will help us leverage the expertise that still exists to excite a new generation of scientists who will move this field of study on this taxonomic group forward. Specifically, we will increase digitization of eremobatid records in the NSF-funded SCAN database portal, create an online guide to camel spiders of North America, and redesign an online website ( to make a new, more inclusive website ( that provides information about this group of arachnids. We will also develop a small “Science News” exhibit at the DMNS to highlight project results for the museum’s visitors.


Key Personnel


Lead PI: Paula Cushing, Ph.D., Curator of Invertebrate Zoology, Denver Museum of Nature & Science.

Role: coordinate all aspects of the research and broader impacts, student mentoring, collecting permits, fieldwork, labwork, data analysis, specimen curation, data management, and taxonomic descriptions.



Co-PI: Matthew Graham, Ph.D., Assistant Professor, Eastern Connecticut State University.

Role: student mentoring, fieldwork, lab work, data analysis, biogeographic analysis, and taxonomic descriptions





Other Personnel


Jack Brookhart, Research Associate, DMNS.

Role: student training and mentoring, specimen identification, and taxonomic descriptions.






Warren Savary, Field Associate, California Academy of Sciences.

Role: Expansion and maintenance of solifuge website, solifuge identification, student mentoring, and taxonomic descriptions.