Spiders: Biodiversity, Evolution, and Cybertaxonomy
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My research investigates patterns of biodiversity in some of the most species rich communities on Earth: tropical forests. Landscape-scale patterns of community structure - how many species live in a place and how does that community change over space and time - are essential for understanding, managing, and protecting biodiversity. Although large-scale community dynamics are well understood for some charismatic groups, most of the world's biodiversity resides in its arthropod species. Diverse tropical arthropod communities are generally not well known. Such communities are characterized by a few common species and a very large number of rare species. Adequate sampling from such communities is problematic because the dominance of rare species makes it nearly impossible to obtain a complete census of the species present. It is only within the past several years that statistical methods have become available to compare communities between sites based on incomplete samples. I am applying these methods to learn how factors like distance, habitat, disturbance, and structural complexity affect diverse communities. I use a variety of approaches to obtain data on biodiversity patterns including morphology-based sorting, DNA barcoding, and next-generation sequencing of bulk homogenized samples.
Inventories of tropical arthropods generate specimens that can form the basis of taxonomic and phylogenetic research. My work in spider taxonomy follows a cybertaxonomic model. In addition to traditional publications, elements of my taxonomic work are distributed online in a variety of ways. This is important because scientists and students living and working in proximity to the greatest concentrations of biodiversity often have limited access to large academic libraries, but access to the internet is increasingly widespread. With primary data aggregated according to community standards, it becomes possible to reuse and recombine data without restrictions. Biodiversity is one of the most information-rich fields of human knowledge and online resources provide unprecedented power to make this aggregated information more accessible and relevant.
Spider diversity of Vietnam. In 2009, spiders were sampled from three localities: Vu Quang, Cuc Phong, and Cat Ba. My collaborators and I established a one-hectare plot at each location and conducted a structured inventory. Preliminary sorting of specimens to morphospecies has been completed (see Miller & Pham 2011) and specimens are now being DNA barcoded. Images of all species will be posted online. We will estimate spider species richness in each of the three forests and assess species overlap between the study sites. Studies of spider species richness from single locations have been conducted at several locations around the world, but the rate at which diverse arthropod communities change across the landscape is not so well understood. Digital images of morphospecies posted online will help us to disseminate our findings quickly, facilitate collaboration, and promote rapid progress toward a detailed understanding of spider community structure in Vietnam and beyond. All of this will aid land use planners and other decision makers concerned with biodiversity, and serve as a resource for Vietnamese students and researchers.
DNA barcoding. I am collaborating with the NCB Naturalis Barcoding Facility on two projects: spiders of the Netherlands and spiders of Vietnam. The goals of the Dutch spider project are 1) to investigate how time since collection affects the probability of obtaining a barcode sequence from a specimen and 2) to produce barcode sequences for as many Dutch spider species as possible based on a combination of museum and fresh collections. This study will help us to more efficiently use legacy collections for spider DNA barcoding projects and predict when fresh specimens will be required. Unlike the Dutch spider project, where the fauna is well known, the spiders of Vietnam project is dominated by unidentified and undescribed species that have been assigned a morphospecies designation. DNA bracode sequences will test these morphospecies concepts, providing an independent check on whether males and females have been correctly matched and whether similar-looking specimens from different locations are actually conspecific.
Biodiversity of the Gaoligongshan. Since 1998, the California Academy of Sciences has lead an international interdisciplinary biotic inventory in the Gaoligongshan (or Gaoligong mountains) of western Yunnan, China. The Gaoligongshan is located in the heart of one of the world's biodiversity "hotspots". "Hotspots" are areas of extreme biodiversity and endemism under severe threat of habitat destruction. But the hotspot designation itself, while an extremely successful concept for the conservation community, has been criticized for over-reliance on data from vascular plants and tetrapod vertebrates. The hotspot designation criteria were originally selected in part because these taxa are relatively well known and well sampled worldwide, but it is not clear that they are representative of the biota as a whole because sampling and knowledge about the most diverse groups on Earth tends to be incomplete. Our inventory emphasized vascular plants and tetrapods (hotspot taxa) as well as fish, non-vascular plants (bryophytes), and two arthropod groups. From these data, we will be able to determine whether hotspot taxa are representative of a broader spectrum of the biota for conservation assessment and provide a model for assessing biodiversity of both well known and poorly known groups simultaneously.
From Taxonomic Literature to Cybertaxonomic Content. Biodiversity is one of the most information-rich fields of human knowledge, but advances in basic cybertaxonomic infrastructure have only recently provided the tools to organize biodiversity information in ways that respond to a wide range of user groups, including ecologists, land managers, and interested citizens, not to mention the benefits of readily accessible information to the global taxonomic community. The key to unlocking this potential is semantic tagging of both prospective and retrospective taxonomic literature (Miller et al. 2012-b).
Systematics of velvet spiders. The spider family Eresidae is a small (ca. 100 species) and interesting lineage. It includes the genus Stegodyphus, which has evolved sociality multiple times, and the ladybird spider (males of some Eresus species), which are arguably the most beautiful spiders in temperate Europe and Asia. With the remarkable exception of one species known from Brazil, velvet spiders can be found only in Europe, Asia, and Africa. Eresidae was historically divided into two subfamilies: Eresinae and Penestominae. Penestomines have a controversial history. Based on the results of a molecular phylogenetic analysis, my collaborators and I (Miller et al. 2010-a) removed Penestominae from Eresidae. I worked with an international team of researchers to taxonomically review and image representatives of all eresid genera (Miller et al. 2012-a).
Phylogeny of Spiders. I have used molecular sequence data to investigate questions about the phylogenetic affinities of several groups of entelegyne spiders (Miller et al. 2010-b).Penestomus was first described from females only and placed in the family Eresidae. Discovery of the male decades later brought surprises, especially in the anatomy of the male palp which features (among other things) a retrolateral tibial apophysis (RTA). The presence of an RTA is synapomorphic for a large clade of spiders exclusive of Eresidae. A molecular data matrix based on four loci was constructed to test two alternative hypotheses: 1)Penestomus are eresids and the RTA is convergent, or 2) Penestomus belong within the RTA clade. Our analysis supported Penestomus as an RTA clade lineage not close to Eresidae. As a result, Penestomus was removed from Eresidae and promoted to family rank (Penestomidae). In addition, we transferred the subfamily Coelotinae from the Amaurobiidae to the Agelenidae. A taxonomic revision of Penestomus (Miller et al. 2010-a) with cybertaxonomic enhancements (including registration of new species in ZooBank,Encyclopedia of Life species pages, and specimen records explorable in Google Earth) complements this phylogenetic work.
Spiders of the Gaoligongshan and online taxonomy. I recently published a monograph describing the Gaoligong species of four closely related spider families: Theridiosomatidae, Mysmenidae, Anapidae, and Symphytognathidae (the micro-orbweaving spiders; Miller et al. 2009). Over 1000 adult symphytognathoid specimens were collected during the 10 year inventory. These belong to 36 species, all new to science. In addition to the traditional journal publication, data are available through a variety of online sources. New species have been registered on ZooBank, images have been posted on Morphbank, species pages have been created for the Encyclopedia of Life, and a Google Earth layer facilitates interactive exploration of specimen collection data.
Repeated evolution of male sacrifice behavior. While it is true that cannibalism during mating occurs with more or less frequency in many spider species, the notion that black widow spiders always eat their mates is a myth. However, males of the Australian widow spiderLatrodectus hasselti exhibit remarkable mating behavior: during copulation, males execute a somersault that places their abdomen near the mouth of the female, frequently resulting in the death of the male. This is an example of male sacrifice behavior. Male sacrifice behavior, defined as either complicity in cannibalism or spontaneous death associated with copulation, is not limited to widow spiders. An analysis using phylogeny-based statistics suggests that a series of evolutionary prerequisites are associated with the evolution of male sacrifice behavior in spiders, including extreme sexual size dimorphism, accumulation of multiple males in the female web (an indicator of a male-biased effective sex ratio), and mutilation of the male genitalia during mating (Miller 2007-a).
Social Spitting Spiders. On a recent expedition to Madagascar, I discovered social behavior in spitting spiders (family Scytodidae). These spiders were previously known to have some maternal care, but not to cooperate as adults. The evolution of maternal care precedes more advanced forms of sociality in several spider lineages, so this discovery is consistent with a larger evolutionary motif.
Teaching & mentoring
I participate in courses offered through Leiden University including Biodiversity II, Biodiversity III, and Richness of the World, and also the Flora and Fauna course at Groningen University. Stage opportunities are available for students interested in quantitative biodiversity, community ecology, and cybertaxonomy.
I participate as an instructor for the Advanced Field Ecology and Conservation Course at Xishuangbanna Tropical Botanical Garden, Yunnan, China.
Postdoctoral Fellow, California Academy of Sciences, 2007. Advisor: Charles Griswold. Funded by "Biotic Survey of the Gaoligongshan, a Biodiversity Hotspot in Western Yunnan, China," National Science Foundation award BSI 0103795 (P.I.s N. Jablonski and P. Fritsch).
California Academy of Sciences, Schlinger Foundation Postdoctoral Fellowship in Systematic Entomology, California Academy of Sciences, 2005-2007. Advisor: Charles Griswold. Systematics of Widow Spiders (Araneae, Theridiidae, Latrodectus): A Total Evidence Approach.
National Museum of Natural History, Smithsonian Institution, 2004-2005. Advisors: Dr. Jonathan Coddington and Dr. Ted Schultz. Molecular Phylogeny and Taxonomy of Widow Spiders (Araneae, Theridiidae, Latrodectus).
National Museum of Natural History, Smithsonian Institution, 2003-2004. Advisor: Dr. Jonathan Coddington. Funded by "Assembling the Tree of Life: Phylogeny of Spiders," National Science Foundation award DEB 0228699 (P.I.s W. Wheeler, L. Prendini, J. Coddington, G. Hormiga and P. Sierwald).
The George Washington University (GWU), Department of Biological Sciences, Washington, D.C., 1996-2003, Ph.D. Co-advisors: Dr. Gustavo Hormiga, Dr. Jonathan Coddington. Dissertation: Progress in the systematics of erigonine spiders (Araneae, Linyphiidae, Erigoninae).
Western Carolina University (WCU), Department of Biology in Cullowhee, North Carolina, 1994-1996, M.S. Advisor: Dr. Frederick A. Coyle. Thesis: Systematics of the erigonine spider genus Sisicottus (Araneae, Linyphiidae).
The Evergreen State College (TESC), Olympia, Washington, 1989-1994, B.A., B.S.
Zootaxa. Subject editor for amaurobioid spiders (Araneae: Amaurobioidea).
ZooKeys. Subject editor for spiders (Araneae).
Biodiversity Data Journal. Deputy Chief Editor.
Arachnida collection at Naturalis
Loans and collection visits can be arranged through our collection department firstname.lastname@example.org or myself at email@example.com. Prospective visitors are advised to visit /collections/plan-your-visit. Our loan policy is available at /collections/loan-policy.
Miller, J., T. Dikow, D. Agosti, G. Sautter, T. Catapano, L. Penev, Z.-Q. Zhang, D. Pentcheff, R. Pyle, S. Blum, C. Parr, C. Freeland, T. Garnett, L.S. Ford, B. Muller, L. Smith, G. Strader, T. Georgiev & L. Bénichou. 2012-b. From taxonomic literature to cybertaxonomic content. BMC Biology, 10: 87. Download
Miller, J.A., C.E. Griswold, N. Scharff, M. Rezac, T. Szuts & M. Marhabaie. 2012-a. The velvet spiders: an atlas of the Eresidae (Arachnida, Araneae). ZooKeys 195: 1–144, doi: 10.3897/zookeys.195.2342. doi.org/10.3897/zookeys.195.2342. Download
Supplementary data archived on Dryad: Miller J., C. Griswold, N. Scharff, M. Rezac, T. Szuts, M. Marhabaie. 2012. Data from: The velvet spiders: an atlas of the Eresidae (Arachnida, Araneae). Dryad Digital Repository. doi:10.5061/dryad.qj8t7r0q. Link
Phylogenetic data matrix (Nexus format) based on aligned molecular sequence data.Download
KML (Keyhole Markup Language) file for viewing specimen occurrence records interactively in Google Earth. Download
Electronic supplementary documents. Figures S1-S3. Download
Miller, J. & C. Rahmadi. 2012. A troglomorphic spider from Java (Araneae, Ctenidae,Amauropelma). ZooKeys, 163: 1-11. doi: 10.3897/zookeys.163.2265. Download
Dumbacher, J.P., J. Miller, M.E. Flannery & Yang X. 2011. Avifauna of the Gaoligong Shan Mountains of Western China: A hotspot of avian species diversity. Ornithological Monographs, 70: 30-63. Abstract Request pdf
IJland, S., J. Miller, Y.J. van Sluijs & P.J. van Helsdingen. 2011. Enkele spinnen uit Zuid-Limburg, met Monocephalus castaneipes (Araneae, Linyphiidae) nieuw voor Nederland. Nieuwsbrief Spined 31: 19-22.
Dick, C.W. & J.A. Miller. 2010. Streblidae (Bat Flies). Pages 1249-1260 in: Manual of Central American Diptera. Volume 2. B.V. Brown, A. Borkent, J.M. Cummings, D.M. Wood, N.E. Woodley & M.A. Zumbado.
Wang, X.-P., C.E. Griswold & J.A. Miller. 2010. Revision of the genus DraconariusOvtchinnikov 1999 (Agelenidae: Coelotinae) in Yunnan, China, with an analysis of the Coelotinae diversity in the Gaoligongshan Mountains. Zootaxa, 2593:1-127. Download
Miller, J.A., C.E. Griswold & C.R. Haddad. 2010-a. Taxonomic revision of the spider family Penestomidae (Araneae, Entelegynae). Zootaxa, 2534:1-36. Download
Supplementary file for viewing distribution records and online species pages interactively inGoogle Earth. Download
Penev, L., D. Agosti, T. Georgiev, T. Catapano, J. Miller, V. Blagoderov, D. Roberts, V.S. Smith, I. Brake, S. Ryrcroft, B. Scott, N.F. Johnson, R.A. Morris, G. Sautter, V. Chavan, T. Robertson, D. Remsen, P. Stoev, C. Parr, S. Knapp, W.J. Kress, F.C. Thompson & T. Erwin. 2010. Semantic tagging of and semantic enhancements to systematics papers: ZooKeys working examples. ZooKeys, 50:1-16. doi: 10.3897/zookeys.50.538. Download
Miller, J.A., A. Carmichael, M.J. Ramirez, J.C. Spagna, C.R. Haddad, M. Rezac, J. Johannesen, J. Kral, X.-P. Wang & C.E. Griswold. 2010-b. Phylogeny of entelegyne spiders: affinities of the family Penestomidae (NEW RANK), generic phylogeny of Eresidae, and asymmetric rates of change in spinning organ evolution (Araneae, Araneoidea, Entelegynae). Molecular Phylogenetics and Evolution, 55:786-804. doi:10.1016/j.ympev.2010.02.021.Download
Appendix C. Supplementary data: Figs. S1-S7, four alignments in Nexus format.
Miller, J.A., C.E. Griswold & C.M. Yin. 2009. The symphytognathoid spiders of the Gaoligongshan, Yunnan, China (Araneae, Araneoidea): Systematics and diversity of micro-orbweavers. ZooKeys 11: 9-195. doi: 10.3897/zookeys.11.160. Download
Electronic appendices: Primary biodiversity data
File format: Microsoft Excel (1997-2003). doi: 10.3897/zookeys.11.160-app.B.dt. Download
File format: KML (Keyhole Markup Language) version 2.1 for Google Earth. doi: 10.3897/zookeys.11.160-app.C.dt. Download
Dataset on GBIF (Global Biodiversity Information Facility). Link
Penev, L., T. Erwin, J. Miller, V. Chavan, T. Moritz & C. Griswold. 2009. Publication and dissemination of datasets in taxonomy: ZooKeys working example. ZooKeys 11: 1-8. doi: 10.3897/zookeys.11.210. Download
Coddington, J.A., I. Agnarsson, J.A. Miller, M. Kuntner & G. Hormiga. 2009. Undersampling bias: the null hypothesis for singleton species in tropical arthropod surveys. Journal of Animal Ecology, 78:573-584. DOI: 10.1111/j.1365-2656.2009.01525.x. Download
Helsdingen, P.J. van, J. Miller & S. IJland. 2009. Snelle inventarisatie van het Noordoostpolder (Kuinderbos) in Flevoland (Araneae). Nieuwsbrief Spined 27: 16-17.Download
Agnarsson, I. & J.A. Miller. 2008. Is ACCTRAN better than DELTRAN? Cladistics, 24:1-7.Download
Penev, L., T. Erwin, F.C. Thompson, H.-D. Sues, M.S. Engel, D. Agosti, R. Pyle, M. Ivie, T. Assmann, T. Henry, J. Miller, N.B. Ananjeva, A. Casale, W. Lourenço, S. Golovatch, H.-P. Fagerholm, S. Taiti, M. Alonso-Zarazaga & E. van Nieukerken. 2008. ZooKeys, unlocking earth’s incredible biodiversity and building a sustainable bridge into the public domain: From “print-based” to “web-based” taxonomy, systematic, and natural history. ZooKeys Editorial Opening Paper. ZooKeys, 1:1-7. doi: 10.3897/zookeys.1.11. Download
Miller, J.A. 2007-a. Repeated evolution of male sacrifice behavior in spiders correlated with genital mutilation. Evolution, 61:1301-1315. Abstract Request pdf
Appendix S1. Data matrix (nexus format). Download
Miller, J.A. 2007-b. Review of erigonine spider genera in the Neotropics (Araneae: Linyphiidae, Erigoninae). Zoological Journal of the Linnean Society, 149 (Suppl. 1):1-263.Download
Miller, J.A. 2007-c. Synaphridae of Madagascar (Araneae: Araneoidea): A new family record for the Afrotropical region. Proceedings of the California Academy of Sciences, 58:21-48. Download
Ramírez, M.J., J.A. Coddington, W.P. Maddison, P.E. Midford, L. Prendini, J. Miller, C.E. Griswold, G. Hormiga, P. Sierwald, N. Scharff, S.P. Benjamin, & W.C. Wheeler. 2007. Linking of digital images to phylogenetic data matrices using a morphological ontology. Systematic Biology, 56:283-294. Download
Miller, J.A. 2006. Web-sharing sociality and cooperative prey capture in a Malagasy spitting spider (Araneae: Scytodidae). Proceedings of the California Academy of Sciences, 57:739-750. Download
Simmons, M.P., L. Zhang, C.T. Webb, A. Reeves & J.A. Miller. 2006. The relative performance of Bayesian and parsimony approaches when sampling characters evolving under homogeneous and heterogeneous sets of parameters. Cladistics, 22:171-185.Download
Miller, J.A. 2005-a. Cave adaptation in the spider genus Anthrobia (Araneae, Linyphiidae, Erigoninae). Zoologica Scripta, 34:565-592. Download
Miller, J.A. 2005-b. A redescription of Porrhomma cavernicola Keyserling (Araneae, Linyphiidae) with notes on Appalachian troglobites. Journal of Arachnology, 33:426-438.Download
Miller, J.A. & I. Agnarsson. 2005. A redescription of Chrysso nigriceps (Araneae, Theridiidae) with evidence for maternal care. Journal of Arachnology, 33:711-714.Download
Kostanjsek, R. & J.A. Miller. 2004. New records of sheet web spiders from Slovenia (Arachnida: Araneae: Linyphiidae). Natura Sloveniae, 6:19–24. Download
Miller, J.A. 2004-a. The genus Brattia beyond South America (Araneae, Linyphiidae). Journal of Arachnology, 32:526–538. Download
Miller, J.A. & G. Hormiga. 2004. Clade stability and the addition of data: A case study from erigonine spiders (Araneae: Linyphiidae, Erigoninae). Cladistics, 20:385-442. Download
Miller, J.A. & D. Ubick. 2004. Spiders. Pages 105-128 in: Borror and DeLong's Introduction to the Study of Insects. Seventh Edition. C.A. Triplehorn & N.F. Johnson.
Toti, D.S., F.A. Coyle & J.A. Miller. 2000. A structured inventory of Appalachian grass bald and heath bald spider assemblages and a test of species richness estimator performance. Journal of Arachnology, 28:329-345. Download
Miller, J.A. 1999. Revision and cladistic analysis of the erigonine spider genus Sisicottus(Araneae, Linyphiidae, Erigoninae). Journal of Arachnology, 27:553-603. Download
Zujko-Miller, J. 1999. On the phylogenetic relationships of Sisicottus hibernus (Araneae, Linyphiidae, Erigoninae). Journal of Arachnology, 27:44-52. Download
Miller, J.A. & F.A. Coyle. 1996. Cladistic analysis of the Atypoides plus Antrodiaetus lineage of mygalomorph spiders (Araneae, Antrodiaetidae). Journal of Arachnology, 24:201-213.Download
Online resources and software
Spiders, LifeDesks Administrator, 2008-present. http://araneae.lifedesks.org/.
Encyclopedia of Life (http://www.eol.org/), 48 species pages contributed.
Morphbank (http://www.morphbank.net/), 570 images contributed.
ZooBank (http://www.zoobank.org/), 50 nomenclatural acts registered.
Barcode of Life (http://www.boldsystems.org/views/login.php), 100 sequences contributed.
Miller, J.A. 2004-b. Key to the Genera of Erigonine Spiders in the Neotropics (Araneae, Linyphiidae, Erigoninae). Available online athttp://www.gwu.edu/~spiders/interactiveKeys.htm. National Museum of Natural History, Smithsonian Institution.
Hormiga, G, J.A. Miller & F. Alverez-Padilla. 2003. LinyGen: Linyphioid Genera of the World (Pimoidae and Linyphiidae), An Illustrated Catalog. Preview Version. Available online athttp://www.gwu.edu/~linygen/.
Zujko-Miller, C. & J.A. Miller. 2002. PEST: Precision Estimated by Sampling Traits. Available online at http://www.gwu.edu/~spiders/pestDocs.htm.
Miller, J.A. & M. Tschapka. 2001. The Bat Flies of La Selva (Diptera: Nycteribiidae, Streblidae). Beta Version 0.96. Available online at: http://www.biologie.uni-ulm.de/bio3/Batfly/index.html.
Selected Popular Features and Presentations
Mooiste spinnen van Europa komen bovengronds in online bibliotheek. 12 June 2012. Link
Rahman, R. Meet the new official Lou Reed spider breed: Just as cuddly as the man himself! EW.com. 24 May 2012. Link
Prostak, S. New Genus of Velvet Spider Named after Lou Reed. Sci-news.com. 23 May 2012. Link
Velvet spiders emerge from underground in new cybertaxonomic monograph. Pensoft news. 23 May 2012. Link
Accessible Biodiversity Knowledge and the World Wide Web. Naturalis Late Night Show. 28 February 2012. Explore the Prezi
Interview. In: Huang, C. Redback spider fears stalk the Emirates. The National. 18 December 2010. Link
Interview. In: Dolgin, E. 2009. Lazy male spiders avoid dinner date. Nature. 21 October 2009. doi:10.1038/news.2009.1020. Link
Interview. Black Widows - Part 2. Science in Action, California Academy of Sciences podcast. 3 September 2007.
Interview. Black Widows - Part 1. Science in Action, California Academy of Sciences podcast. 27 August 2007.
Interview. Silk Triplines. Science in Action, California Academy of Sciences podcast. 20 August 2007.
Interview. In: Suicide mating. Pp. 16-17. California Academy of Sciences: Member Publication, Summer 2007.
Interview, photography. In: Perlman, D. Academy of Sciences. Spitting spider's surprise. Unlike its brethren, this new discovery has a nurturing side. San Francisco Chronicle. 22 July 2006. Link
Interview, photography. In: Spiders that hunt together stay together. Pp. 19-21. California Academy of Sciences: Member Publication, Fall 2006.
Interview, cover photo and additional photography. In: Tiny Assassins. Pp. 3-4. California Academy of Sciences: Member Publication, Summer 2006.
Tiny Assassins. California Academy of Sciences, Science Now. 2006. Link
California Academy of Sciences Annual Halloween Party and Benefit, San Francisco, California, 2006
Tagliaferro, L. 2004. Spiders and Their Webs. Animal Homes series. J. Miller, consultant. Capstone Press, Mankato, Minnesota.
Insect Expo, National Museum of Natural History, Washington D.C., 2003.
Bugfest, National Mall, Washington D.C., 1997, 2000.
"Spellbinding Spiders," a public presentation at the Mountain Heritage Center, Cullowhee, North Carolina, 1995.
"Spiders!" an arachnological experience for preschool-age naturalists at the Jackson County Public Library, Sylva, North Carolina, 1995.
Please contact the author when you are interested in one of the articles above and it is not available online or in the repository [Repository].
Barcode of Life. http://www.boldsystems.org/views/login.php
Biodiversity Data Journal. http://biodiversitydatajournal.com/
Darwin Core Reference Guide. http://rs.tdwg.org/dwc/terms/index.htm
DORA (The San Francisco Declaration on Research Assessment).http://am.ascb.org/dora/
EMBL-EBI (European Molecular Biology Laboratory-European Bioinformatics Institute) Sequence Analysis. http://www.ebi.ac.uk/Tools/sequence.html
Encyclopedia of Life. http://www.eol.org/
Entrez, NCBI. http://www.ncbi.nlm.nih.gov/sites/gquery
Global Biodiversity Information Facility (GBIF). http://data.gbif.org/welcome.htm
LifeDesks Spiders. http://araneae.lifedesks.org/
Notes from Nature. http://www.notesfromnature.org/
Selling Wine Without Bottles: The Economy of Mind on the Global Net. John Perry Barlow.http://www.virtualschool.edu/mon/ElectronicFrontier/WineWithoutBottles.html
Universiteit Leiden. Voorzieningen. http://www.voorzieningen.leidenuniv.nl/ulcn/
The World Spider Catalog. Norman I. Platnick.http://research.amnh.org/iz/spiders/catalog/INTRO1.html