Funnel-Web Spiders from Sam Houston National Forest and San Antonio, Texas 3

— BugsInTheNews is a VIEWER-PARTICIPANT WEBSITE. This article by Jerry Cates, Marney M. (SHNF), and Cliff M. (San Antonio, TX), was first published in July 2009, and revised last on 8 July 2012. © Bugsinthenews Vol. 10:07.

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Marney (a gentleman) wrote:

• “I was out, with my family, at Double Lakes Recreation Area in the Sam Houston National Forest on July 3rd, and shot these pictures of a spider in its ‘lair’. I found your website during an Internet search and was wondering if you can identify this critter.”

I thanked Marney for the excellent photos, and asked if he could resend them, not as email photo attachments (which some mail servers automatically reduce to a low resolution for email transmission) but as image file attachments (that the mail server will send without modifying). He did that immediately, and enlargements of selected portions of both photographs are posted here, showing important anatomical features not visible on the email attachments initially received.

I was struck, on first viewing these, by the similarity these photos bear to a set of images I remember seeing, in the late 1940’s as a young child, in a nature encyclopedia. The encyclopedia, which I recall to have been published in the 1920’s, provided a well-written article on funnel-web weavers, and told of the biology of such spiders, how they build their funnel-web snares, and the mechanics of how they kill and consume their prey. Such accounts, supplied with beautiful drawings and well-illustrated photographs, caused my interest in nature to blossom and grow. Now I wonder if the content of this page, meager as it is, might grip other young minds, with anything similar in way of effect.

Willis J. Gertsch, at the time Curator Emeritus, Department of Insects and Spiders, American Museum of Natural History, published in 1979 an excellent book on arachnids, titled “American Spiders”. In this book, Gertsch devoted a total of five pages to the funnel-web weavers. He mentions in those pages that the Agelenidae (which, then as now, was devoted to the funnel-web weavers) comprise a numerous and diverse family of over 400 species found in the United States. Presently, due to a number of important revisions to arachnid taxonomy, the Agelenidae of North America is represented in but 9 genera and 85 species. The Amaurobiidae (10 N.A. genera, 97 species), though once included within the Agelenidae, are now accorded family status on their own, being recognized as mostly builders of cribellate retreats with radiating tangles, and — for an ecribellate subset — sheet webs with one or a plurality of retreats, or small sheet webs that they hang from, inverted.

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Note that the photos displayed on this page, as with images posted on all web pages on bugsinthenews, may be enlarged for more detailed viewing by hovering your cursor over them, then clicking.

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It is not possible to assign a specific genus or species from these photos. The crucial anatomical features needed for such assignments are simply not visible.

Species in the genus Tegenaria have eight eyes of roughly equal size, though the anterior lateral eyes (ALE) are slightly larger; both eye rows are roughly straight or gently procurved. The remaining eight Agelenidae genera represented in North America also have eight eyes, but their ALE are both dramatically larger and relatively close together, with the much smaller anterior median eyes (AME) immediately above them.

In the first photograph posted here, the reflection of Marney’s flash appears to emanate from two large ALE, with another reflection that could be produced by two smaller PLE above and between them. However, it is unwise to infer anything about the respective size of this spider’s eyes from these reflections, as the eyes themselves are not visible in sufficient resolution to allow comparisons to be made between them.

The funnel web, in general, is said to be similarly constructed by all species in the Agelenidae. Gertsch described it as having both an entrance and an exit, the latter hidden from view but supplying a propitious back door exit from a nasty situation should the need arise.

Spread before the entrance is an expansive sheet web that beguiles naive insects and other organisms who mistake it for a nice place to land, meander across, or to rest upon.

The spider, whose tarsi are unencumbered by any structures beyond their three claws, races over the sheet web with facility, capturing the foolish interlopers and producing therefrom a nifty repast. This facility comes with a price, as agelenids — lacking the brush of fine hairs,  at the tips of their feet (scopulae, as described in Foelix, 1996, p. 18, 20-21) — cannot scale smooth surfaces such as glass or porcelain the way many other spiders do. Thus cosmopolitan agelenids that invade our homes and businesses often become ensnared by kitchen bowls, porcelain sinks, and bathroom tubs; once they fall into these smooth-sided articles they are unable to climb out.

The sheet web gracing the threshold of the funnel retreat is, according to Gertsch, a work of a lifetime for the funnel-web weaver. She constantly adds to the throughout her existence, rather than daily tearing it down and rebuilding from scratch the way most orb-weaving spiders do.

Cliff M. wrote:

• “The spider in these photos (Cliff sent a number of images, but the image above is the best of the lot and the others do not materially add to it) has built its web outside my new home in NE San Antonio. The spider’s body is about an inch across, leg-body-leg. I hope you can tell me something about it. I recently turned (Cliff’s age has been redacted for privacy purposes), and have had a camera in my hand since age 10. I’ve taken over 5,600 shots with this camera alone.”

I thanked Cliff for the excellent photos, and asked if he could take some more, especially of the frontal head that will show the arrangement of the eyes. Then I explained I was only a few years his junior, and I, too, had kept a camera nearby since my tenth Christmas, when I found a Kodak Hawkeye under the tree with my name on it. Two years later, using money saved from a paper route, I bought an Argus C-3, and have sought to hone my skills at photography ever since. The Argus C-3 was retired years ago, but it is still a beautiful instrument.

As to the photo at hand, it is of a funnel web weaver in the family Agelenidae, similar in many respects to the specimen Marney M. photographed at Sam Houston National Forest on July 3rd. Notice, for example, that the patella of each leg is darkened, as is the distal portion of each tibia, and the tarsus of each palp, in both spiders.

There are important differences, too. Cliff’s spider’s legs have additional dark bands, for one thing, to the point that every leg segment has a dark mark at its distal end and the tibia is marked at mid-shaft as well. The alternating dark and light stripes on the carapace are reversed, for another; Cliff’s spider has a dark, narrow median stripe bordered on each side by a narrow pale stripe, then by a broad dark stripe. Marney’s spider, however, has a pale median stripe bordered by a broad, dark marking.

Notes on the spider family Agelenidae:

The science of arachnology is in a dizzying state of flux. Nowhere is that more evident than in the assignment of individual spiders to family, genus, and species. When we speak of a spider as being an agelenid, as we are doing here, one might presume the term to have a well-defined, undisputed meaning. In the main, perhaps, it does. But on delving more deeply, one easily gets the impression that the seemingly rigid walls around that meaning are so riddled with cracks as to require constant remodeling (i.e., revisions). The old saw about broken eggs and omelettes is, methinks, apropos, as the following review illustrates.

The family name, Agelenidae, was first described in 1837, some 174 years ago, and derives from the Latin word, agel, for “a small field.” Over the intervening generations, its reach has expanded and shrunk as new spiders were added and old ones were either discarded (on recognizing them as synonyms of existing species), used to create new families or genera, or moved to other, more appropriate taxa. Over that same period, many of the spiders in the family have had their binomial names altered, some gently, others in a more dramatic fashion. Note that one or more investigators misspelled the family name as Agalenidae, and the generic name that derives from it (Agelena) as Agalena.

Following is a terse summary of a number of texts on the Agelenidae (each reference is listed in full at the end of this article, with links to on-line sources if available), wherein we explore some of the taxonomical revisions that have taken place:

Emerton, in his 1883 book on spiders and their habits, is concerned with generalities and devotes less than one page (p. 26) to the Agelenidae, which he spells Agalenidae, describing them thusly:  “AGALENIDAE. Long-legged, brown spiders, with two spinnerets longer than the others, and extending out behind the body. They make flat webs, with a funnel-shaped tube at one side, Fig. 24, in which the spider waits. Fig. 9 is Agalena nævia, the common grass spider.”

Later, on page 54, he describes the web drawn in his Fig. 24 in these words:

“COBWEBS. The simple nests and tubes that have been described are made by spiders, most of which spin no other webs. The larger and better known cobwebs for catching insects are made by comparatively few species. On damp mornings in summer the grass-fields are seen to be half covered with flat webs, from an inch or two to a foot in diameter, which are considered by the weatherwise as signs of a fair day. These webs remain on the grass all the time, but only become visible from a distance when the dew settles on them. Fig. 24 is a diagram of one of these nests, supposed, for convenience, to be spun between pegs instead of grass. The flat part consists of strong threads from peg to peg, crossed by finer ones, which the spider spins with the long hind-spinnerets, Fig. 20, swinging them from side to side, and laying down aband of threads at each stroke.”

“The web is so close and tight, that one can hear the footsteps of the spider as she runs about on it. At one side of the web is a tube leading down among the grass-stems. At the top the spider usually stands, just out of sight, and waits for something to light on the web, when she runs out, and snatches it, and carries it into the tube to eat. If any thing too large walks through the web, she turns around, and retreats out of the lower end of the tube, and can seldom be found afterward. In favorable places these webs remain through the whole season, and are enlarged, as the spider grows, by additions on the outer edges, and are supported by threads running up into the neighboring plants. Similar webs are made by several house-spiders, and are enlarged, if let alone, till they are a foot or two feet wide, and remain till they collect dirt enough to tear them down by its weight.”

Emerton’s 1902 book, published 19 years after his 1883 book on spiders and their habits, is occupied with describing species, and devotes sixteen pages (pp. 91-106) to the individual Agelenidae (he persists in spelling it Agalenidae). In these pages, he describes, by name, seven separate agelenids (misspelling the generic name as in 1883), omitting attribution: (1) Agalena nævia, the common grass spider mentioned in passing in his 1883 book, (2) Tegeneria derhamii, (3) Tegeneria (Cælotes) medicinalis, (4) Tegenaria (Cælotes) longitarsus, (5) Tegenaria (Cicurina) complicata, (6) Hahnia bimaculata, and (7) Hahnia cinerea.

Comstock’s 1912 book on spiders, published 12 years after Emerton, 1902, devotes 17 pages (pp. 582-598) to the Agelenidae (he spells the family and generic names correctly). Though most likely benefitting from Emerton’s previously published works, he makes no direct references to them. In the third paragraph of his exposition he states that the family includes such notables as Argyroneta aquatica¹ of Europe. Then he lists “the nine genera of our fauna,” and discusses each separately, describing one or more species in more detail, usually without attribution: (1) Cybæus giganteus, (2) Agelena nævia, whose male palpal characters are so varied that he wonders if more than one species may be involved, (3) Coras medicinalis, here noting that “Hentz, who first described the species and proposed the name medicinalis for it, states that ‘for some time the use of its web as a narcotic in cases of fever was recommended by many physicians in this country; but now it is probably seldom used.'”²; (4) Tegenaria derhami, here spelled with a single “i” rather than the “ii” used by Emerton, (5) Cœlotes fidelis, (6) Chorizomma californica, (7) Cicurina arcuata, (8) Cryphœca montana Emerton, and Cryphœca peckami, and (9) Hahnia agilis, and Hahnia cinerea.

¹The species Argyroneta aquatica Latrielle, 1804, also known as the diving bell spider, was for many years included in the Agelenidae as a separate genus comprising a single species, but was transferred to the Cybaeinae by Grothendieck & Kraus, in 1994. The Cybaeinae, traditionally considered a subfamily of Agelenidae, was made a subfamily of Dictynidae by Lehtinen in 1967 and elevated to family status by Forster in 1970. Platnick (2011b).

²Hentz first described the species as Tenenaria medicinalis in 1821, the establishment date for the species name. Later, in 1837, Walckenaer placed it in the genus Clubiona, but his position was not accepted by others, and in 1841 he revised the name to Tegenaria nemorensis. Hentz, in 1847, and again in 1867, reasserted his original nomenclature, Tegenaria medicinalis, while Keyserling, in 1887, proposed its placement in the genus Coelotes, with the species name urbanus. In 1898 Simon published a revision placing the species under the genus Coras, giving priority to the specific name established by Hentz in 1821: thus Coras medicinalis. Emerton (1902), as previously noted, described the species as Tegenaria (Cælotes) medicinalis, implying that the generic placement was in dispute, and that he was unaware of Simon’s 1898 revision. Comstock, in 1912, recognized Simon’s 1898 revision as superior, as have other investigators since that date to the present.

Kaston (1978) devotes 9½ pages (pp. 164-173) to the Family Agelenidae. Therein he describes the following species, with attribution: (1) Yorima angelica Roth, (2) Cybaeus reticulatus Simon, (3) Cryphoeca montana Emerton, (4) Cicurina brevis Emerton, (5) Cicurina arcuata Keyserling, (6) Cicurina utahana Chamberlin, (7) Cicurana robusta Simon, (8) Agelenopsis pennsylvanica C. L. Koch, (9) Agelenopsis naevia¹ Walckenaer, (10) Agelenopsis aperta Gertsch, (11) Rualena cockerelli Chamberlin & Ivie, (12) Calilena restricta Chamberlin & Ivie, (13) Novalena intermedia Chamberlin & Gertsch, (14) Hololena curta McCook, (15) Hololena hola Chamberlin & Gertsch, (16) Wadotes hybridus Emerton, (17) Wadotes calcaratus Keyserling, (18) Coras medicinalis Hentz, (19) Coras lamellosus Keyserling, (20) Coelotes juvenalis Keyserling, (21) Tegenaria domestica Clerck, (22) Calymmaria cavicola Banks, (23) Calymmaria emertoni Simon, and (24) Calymmaria californica Banks.

¹This species was first described by Walckenaer in 1805. under the species name nævia, nomen nudem. He published a complete description of the species, assigning it to the genus Agelena, in 1841, the formal establishment date for the name. Thorell proposed reassigning it to the genus Agelenopsis in 1877, which proposal was formalized in 1941 (100 years after Walckenaer’s original establishment date) by Chamberlin & Ivie.

Gertsch (1979) devotes 4½ pages to the funnel-web weavers (pp. 213-218), noting that the family “includes more than 400 species,” then lists or describes, without attribution, (1) Agelenopsis pennsylvanica, (2) Tegenaria domestica, (3) Tegenaria chiricahuae, (4) Tegenaria agrestis, (5) Tegenaria pagana, (6) Tegenaria gigantia, (7) Tegenaria larva, (8) the genus Calymmaria, (9) the genus Cicurina, (10) the genus Blabomma, (11) the genus Yorima, (12) and the genus Cybaeozyga. He then points out that the Hahniidae family, though distinct from the Agelenidae, is very similar, and that Hahnia cinerea is a member of the Hahniidae family.

Taxonomy:

• Phylum Arthropoda (described by Pierre André Latreille [November 20, 1762 – February 6, 1833], a French zoologist, in 1829, using two Greek roots: αρθρον “arthron” = jointed + ποδ “pod” = foot: in reference to animals with jointed feet, but in the more narrow context of invertebrates, which have segmented bodies as well as jointed appendages);
• Subphylum: Chelicerata (described by Richard Heymons [1867 – 1943], a German zoologist, in 1901, using the Greek noun χηλη “chele” = a claw, talon, or hoof + the Latin suffix ata — which by convention is suffixed to the names of animal subdivisions — in reference to animals with specialized appendages before the mouth that are used in feeding, capturing and securing prey, and, in the case of spiders, injecting venom and digestive agents into their prey); 
• Class Arachnida (described by Jean Léopold Nicolas Frédéric Cuvier (August 23, 1769 – May 13, 1832), a.k.a. Georges Cuvier, a French naturalist and zoologist, in 1812, using the Greek noun αραχης “araches” = a spider, in reference to all eight-legged arthropods, including such disparate animals as ticks, mites, scorpions, harvestmen, solpugids, and spiders) 
• Order Araneae (described by Carl Alexander Clerck [1709 – 22 July 1765], a Swedish entomologist and arachnologist, in 1757, using the Latin aranea = a spider or a spider’s web, to refer to eight legged arthropods that spin webs); 
• Family Agelenidae (described by Carl Ludwig Koch [September 21, 1778 – August 23, 1857], a German entomologist and arachnologist, in 1837 (Koch, 1837), presumably following Walkenaer in applying the Latin word agel = a small field + the patronymic suffix -idae, from the Greek ιδες — used by convention in zoological nomenclature to indicate a family name — in reference to spiders that are characteristically found in grassy fields); below is Koch’s original description in German, from page 24 of Part I of his 1837 book, Übersicht des arachnidensystems (Arachnid Systematics):

“Familie IV.

Trichterspinnen. Agelenides.

1. Unterfamilie. Eigentliche Trichter- spinnen,

Die Weibchen verfertigon sich meistentheils grosse horizontale ziemlich dichte Gewebe, welche im Hintergrunde in eine trichterförmige Röhre auslaufen. Man findet sie in den Winkeln der Gebäude, zwischen hohlliegenden Wurzeln, in Mauerlöchern, auf der Erde in Höhlungen ausgehend, seltener auf Gesträuchen. Die Weibchen verlassen ihr Gewebe freiwillig nicht, und verstecken sich, wenn sie Gefahr merken, in ihr Trichtergewebe. Die Männchen sind herumirrend; man sieht sie gewöhnlich den Tag über mit ausgebreiteten Beinen ruhend an Wänden, Mauern u. dgl.”

• A rough translation of the original German into English follows:

“Family IV.

Funnel weaver. Agelenides.

1. Subfamily. Actual funnel- spiders,

The females make large, rather dense, horizontal webs, which in the background have the form of a funnel-tube that narrows distally. One finds them in the corners of buildings, in the hollows between roots, in holes in walls, emanating from concavities in the ground, more rarely on dense shrubbery. The females do not leave their webs voluntarily, but hide, if they sense danger, in their funnel-web retreats. The males are more daring; one usually sees them throughout the day, their legs spread outward as they rest on walls and similar surfaces.”

• Genera: 64 worldwide (Platnick, 2011), incl. 9 in North America (Bennett & Ubick, 2005);

1. Gen. Acutipetala Dankittipakul & Zhang, 2008: 2 species (Thailand)
2. Gen. Agelena Walckenaer, 1805: 69 species
3. Gen. Agelenella Lehtinen, 1967: 1 species (Yemen, Socotra)
4. Gen. Agelenopsis Giebel, 1869: 13 species in North America (Bennett & Ubick, 2005); embolus a conspicuous large open circle, epigynum with coupling cavity on posterior margin (Bennett & Ubick, 2005).
5. Gen. Ageleradix Xu & Li, 2007:
6. Gen. Agelescape Levy, 1996:
7. Gen. Alloclubionoides Paik, 1992:
8. Gen. Aterigena Bolzern, Hänggi & Burckhardt, 2010:
9. Gen. Azerithonica Guseinov, Marusik & Koponen, 2005:
10. Gen. Barronopsis Chamberlin & Ivie, 1941: 4 species in North America (Bennett & Ubick, 2005); embolus in tight spiral coil basally; atrium with deeply notched anterior margin furrow (Bennett & Ubick, 2005).
11. Gen. Benoitia Lehtinen, 1967:
12. Gen. Bifidocoelotes Wang, 2002:
13. Gen. Calilena Chamberlin & Ivie, 1941: 16 species in North America (Bennett & Ubick, 2005); male palp with RTA (retrolateral apophysis) complex, distal and lateral components occupying most of retrolateral face of palpal tibia; embolus short, simply curved; transparent fulcrum present (may be very small) at base of embolus; genital bulb with medium apophysis and conductor, but lacking conspicuous tegular process; epigynum with slender elongate scape projecting posteriorly from anterior median margin; 2 teeth on retromargin of cheliceral fang furrow (Bennett & Ubick, 2005).
14. Gen. Coelotes Blackwall, 1841:
15. Gen. Coras Simon, 1898:
16. Gen. Draconarius Ovtchinnikov, 1999:
17. Gen. Femoracoelotes Wang, 2002:
18. Gen. Hadites Keyserling, 1862:
19. Gen. Himalcoelotes Wang, 2002:
20. Gen. Histopona Thorell, 1869:
21. Gen. Hololena Chamberlin & Gertsch, 1929: 30 species in North America (Bennett & Ubick, 2005); male palp retrolateral tibial apophysis bipartite with basal component very large and distinctly separate from distal component; embolus sinuous with tip enclosed in transparent fulcrum arising anteriorly from surface of tegulum; epigynum large with atrium broad and divided longitudinally by more-or-less complete ridge and with pair of stout postero-lateral spurs; atrium of some Rualena very similar; 3 teeth on retromargin of cheliceral fang furrow (Bennett & Ubick, 2005).
22. Gen. Huangyuania Song & Li, 1990:
23. Gen. Huka Forster & Wilton, 1973:
24. Gen. Hypocoelotes Nishikawa, 2009:
25. Gen. Inermocoelotes Ovtchinnikov, 1999:
26. Gen. Iwogumoa Kishida, 1955:
27. Gen. Kidugua Lehtinen, 1967:
28. Gen. Leptocoelotes Wang, 2002:
29. Gen. Lineacoelotes Xu, Li & Wang, 2008:
30. Gen. Longicoelotes Wang, 2002:
31. Gen. Lycosoides Lucas, 1846:
32. Gen. Mahura Forster & Wilton, 1973:
33. Gen. Maimuna Lehtinen, 1967:
34. Gen. Malthonica Simon, 1898:
35. Gen. Melpomene O. P.-Cambridge, 1898: 1 species in North America (Bennett & Ubick, 2005); male palp with retrolateral tibial apophysis simplex, confined to distal half of retrolateral face of palpal tibia; embolus slender, compoundly corved; transparent fulcrum absent; tegular processes well developed; epigynum with anterior hood but lacking slender, elongate scape; 3-4 teeth on retromargin of cheliceral fang furrow (Bennett & Ubick, 2005).
36. Gen. Mistaria Lehtinen, 1967:
37. Gen. Neorepukia Forster & Wilton, 1973:
38. Gen. Neotegenaria Roth, 1967:
39. Gen. Neowadotes Alayón, 1995:
40. Gen. Notiocoelotes Wang, Xu & Li, 2008:
41. Gen. Novalena Chamberlin & Ivie, 1942: 5 species in North America (Bennett & Ubick, 2005); male palp with retrolateral tibial apophysis not clearly bipartite, basal and distal components not distinctly separated; embolus evenly curved, transparent fulcrum absent; epigynum much smaller and lacking broad, medially ridged atrium and postero-lateral spurs; 2-4 teeth on retromargin of cheliceral fang furrow (Bennett & Ubick, 2005).
42. Gen. Olorunia Lehtinen, 1967:
43. Gen. Oramia Forster, 1964:
44. Gen. Oramiella Forster & Wilton, 1973:
45. Gen. Orepukia Forster & Wilton, 1973:
46. Gen. Orumcekia Koçak & Kemal, 2008:
47. Gen. Paramyro Forster & Wilton, 1973:
48. Gen. Pireneitega Kishida, 1955:
49. Gen. Platocoelotes Wang, 2002:
50. Gen. Porotaka Forster & Wilton, 1973:
51. Gen. Pseudotegenaria Caporiacco, 1934:
52. Gen. Robusticoelotes Wang, 2002:
53. Gen. Rualena Chamberlin & Ivie, 1942: 9 species in North America (Bennett & Ubick, 2005); distinguished by retrolateral tibial apophysis confined to distal half of palpal tibia; epigynum with atrium undivided or only partially divided, and with posterior margin bordered by a thickened ridge; 2-3 teeth on retromargin of cheliceral fang furrow (Bennett & Ubick, 2005).
54. Gen. Spiricoelotes Wang, 2002:
55. Gen. Tamgrinia Lehtinen, 1967:
56. Gen. Tegecoelotes Ovtchinnikov, 1999:
57. Gen. Tegenaria Latreille, 1804: 6-7 species in North America (Bennett & Ubick, 2005); distinguished by eight eyes in two straight or procurved rows in frontal view, & sternum with contrasting, variable pattern of pale median stripe and lateral spots (Bennett & Ubick, 2005), [and] by the presence of plumose hair on the body and legs (Roth, 1968).
58. Gen. Textrix Sundevall, 1833:
59. Gen. Tikaderia Lehtinen, 1967:
60. Gen. Tonsilla Wang & Yin, 1992:
61. Gen. Tortolena Chamberlin & Ivie, 1941: 1 species in North America (Bennett & Ubick, 2005); nearctic male apparently unknown, but embolus probably in form of transverse figure-8; atrium marked by paired ridges spiraling upwards to small copulatory openings (Bennett & Ubick, 2005).
62. Gen. Tuapoka Forster & Wilton, 1973:
63. Gen. Urocoras Ovtchinnikov, 1999:
64. Gen. Wadotes Chamberlin, 1925:

• Species: Unk. number of species (some est. about 500) worldwide (but see Platnick, 2011), incl. 85 in North America (Bennett & Ubick, 2005).

Behavior:

Common Names: in process

Distinguishing Characteristics: in process

Distribution: in process

Physiology: in process

Mythology: in process

Similar Families: in process

References:

• Aitchison, C. W. 1981. Feeding and growth of Coelotes atropos (Araneae, Agelenidae) at low temperatures. J. Arachnol., 9:327-330.
• Beccaloni, Jan. 2009. Arachnids. University of California Press.
• Bennett, R. G. 1987. Systematics and natural history of Wadotes (Araneae, Agelenidae). J. Arachnol., 15:91-128.
• Bennett, Robert G. 1988. The spider genus Cybaeota (Araneae, Agelenidae). J. Arachnol, 16:103-119.
• Bennett, Robert G. 1989. “Emerit’s glands” in Cybaeota (Araneae, Agelenidae). J. Arachnol., 17:225-235.
• Bennett, Robert G., and Darrel Ubick. 2005. Agelenidae, p. 56-59 in D. Ubick, P. Paquin, P. E. Cushing, and V. Roth (eds.) Spiders of North America: an identification manual. American Arachnological Society.
• Borradaile, L. A., and F. A. Potts. 1958. The Invertebrata; a manual for the use of students. Cambridge at the University Press.
• Buchsbaum, Ralph, et al. 1987. Animals Without Backbones, Third Edition. The University of Chicago Press.
• Bulloch, W. S. 1958. Practical Invertebrate Anatomy. McMillan & Co. Ltd.
• Catley, Kefvn M. 1992. Supercooling and its Ecological Implications in Coelotes atrropos (Araneae, Agelenidae). J. Arachnol. 20:58-63.
• Comstock, John Henry. 1912. The Spider Book. The New Nature Library, Vol. Seven, Part Two. Doubleday, Page & Company.
• Emerton, James H. 1883. The Structure and Habits of Spiders. S. E. Cassino & Co, Publishers.
• Emerton, James H. 1902. The Common Spiders of the United States. Dover Publications Inc.
• Foelix, Rainer F. 1996. Biology of Spiders, Second Edition. Oxford University Press.
• Fraser, J. G. 1987. Courtship and copulatory behavior of the funnel-web spider, Hololena adnexa (Araneae, Agelenidae). J. Arachnol., 15:257-262.
• Gaver, Melissa M., & Laurel D. Hansen. 2005. Efficacy of Insecticides to Control Tegenaria agrestis and Latrodectus hesperus (Araneae: Agelenidae and Theridiidae). Proc. 5th Int. Conf. on Urban Pests.
• Gertsch, Willis J. 1979. American Spiders, Second Ed. Van Nostrand Reinhold Company.
• Heiss, John Stabe, and Michael L. Draney. 2004 Revision of the Nearctic Spider Genus Calymmaria (Araneae, Hahniidae). J. Arachnol. 32:457-525.
• Hillyard, Paul. 1994. The Book of the Spider; from Arachnophobia to the Love of Spiders. Random House Inc., N.Y.
• Ibarra N., G. 1985. Egg feeding by Tegenaria spiderlings (Araneae, Agelenidae). J. Arachnol., 13:219- 223.
• Kaston, B. J. 1978. How to know the spiders. McGraw Hill Company.
• Koch, Carl Ludwig. 1837. Übersicht des arachnidensystems. Nürnberg,C.H. Zeh [etc.]1837-50.
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