Non-Theraphosidae Mygalomorphs of North America

© 2015 by Andrew Olson

North America has a rich collection of fauna in all the families of . This paper introduces these families, with a special emphasis on the taxa of the USA. To see what non­Theraphosidae mygalomorphs appear in your state, please consult this large table: http://cacoseraph.x10host.com/ntmygs/ntmygs_usa.php ​

Taxonomical placement of mygalomorph families found in the USA. Family arrangement based on Platnick’s World Catalog and is indicative of the “closeness” of relation between various families. Kingdom: Animalia . Convenience taxonomical level: Invertebrata # . Phylum: Arthropoda . . Class: Arachnida . . . Order: Araneae . . . . Suborder: . . . . . Infraorder: Mygalomorphae ...... Family: Atypidae / Atypidae­ Pursewebs or Atypical ­ “Atypical” ​ ​ ​ ​ ​ ​ ! . . . . . Family: / Antrodiaetidae ­ Folding trapdoors ​ ​ ​ ​ ​ ​ ! . . . . . Family: Mecicobothriidae / Mecicobothriidae ­ Dwarf tarantulas ​ ​ ​ ​ ​ ​ ! . . . . . Family: / Dipluridae ­ Funnel­web tarantulas ­ “Two Tails” ​ ​ ​ ​ ​ ​ ! . . . . . Family: Cyrtaucheniidae / Cyrtaucheniidae ­ Waferlid/Wafer trapdoors ​ ​ ​ ​ ​ ​ ! . . . . . Family: / Ctenizidae ­ Corklid trapdoors/Trapdoors ​ ​ ​ ​ ​ ​ ...... Family: Euctenizidae / Euctenizidae ­ ????????? ​ ​ ​ ​ ​ ​ ...... Family: / Nemesiidae ­ False tarantulas ​ ​ ​ ​ ​ ​ ...... Family: / Barychelidae ­ Brushed trapdoors/Brushfoot trapdoors ​ ​ ​ ​ ​ ​ ! . . . . . Family: Theraphosidae / Theraphosidae ** ­ Tarantulas ​ ​ ​ ​ ​ ​ Key: WSC Link / Wiki Link ­ Common name(s) ­ “Translation of family name” # Invertabrata is not a genuine taxon, but is used as a way to conveniently describe all critters that lack a backbone/notochord/major central never channel/etc ! Taxa so marked occur in Arizona. ** Not the focus of this discussion

Atypidae / Atypidae- Pursewebs or Atypical tarantulas- “Atypical” ​ ​ ​ ​ Atypids are known as purseweb because of the type of web and burrow they create. They will make a silk lined burrow in which the silk lining extends out and up away from the burrow entrance to form an above ground, exposed length of tubeweb. This tubeweb almost always ends up a piece of structure like a rock or tree trunk. Hunting involves waiting to sense the passage of a prey item across the exposed portion of tube web. The spider will then bite their prey through the webbing, then once subdued will drag the victim through the webbing and into their burrow. Because of this hunting method atypids have VERY large chelicerae compared to other spiders of similar size. This gives them what some perceive as a fierce look. Atypids lack most of the setae on the cephalothorax that is present in Theraphosidae.

Marshal Hedin ­ Atypidae Web Patrick Edwin Moran ­ Sphodros rufipes, mature male CCASA2.5G* GNUFDL*

Antrodiaetidae / Antrodiaetidae - Folding trapdoors ​ ​ ​ ​ Antrodiaetids are an interesting mix of shape and behavior of Atypids and other, less closely related trapdoor spiders. They have more massive chelicerae and fangs like Atypids but make actual trapdoors on their burrows. The doors are thought to possibly help retain moisture and explain the increased range of Antrodiaetids compared to Atypids. Some Antrodiaetids will block their burrows with plugs made of either silk or silk and soil to increase the insulating factor. Antrodiaetus tend to make their burrows topped with a turret while Aliatypus do not.

Marshal Hedin CCA2.0G* Marshal Hedin CCA2.0G* Antrodiaetus unicolor Aliatypus torridus

Mecicobothriidae / Mecicobothriidae - Dwarf tarantulas ​ ​ ​ Mecicobothriids appear similar to very small tarantulas, though they are not as hirsute. Their main spinnerets are longer than the longest of any Theraphosid, also. The largest are around 1­1.5” DLS (diagonal leg span). They tend to live under structures or in preexisting holes in a sheetweb construct. Not unusual to find them within 30m of a water feature. Mecicobothriids make lenticular, fixed eggsacs. The smallest species average under ten eggs per sac and the largest species average approximately 80 eggs per eggsac. Mecicobothrids lack the scopulae that tarantulas possess and thus can not walk or run up smooth vertical surfaces.

Andrew Olson ­ Megahexura fulva, size Andrew Olson ­ Megahexura fulva, mating comparison

Andrew Olson ­ Megahexura fulva, in situ Andrew Olson ­ Megahexura fulva, detail shot of tergal plates

Dipluridae / Dipluridae - Funnel-web tarantulas - “Two Tails” ​ ​ ​ ​ Diplruids look very similar to tarantulas, but generally are smaller and have much larger main spinnerets. Diplurids use these large spinnerets to line their burrows with copious amounts of silk. A diplurid burrow will usually have sheet web extending out from the entrance, in comparison to US tarantulas which usually line their burrows but do not have much web extending beyond. Diplurids can be found under structures and do not tend to dig or use preexisting rodent burrows as much as US tarantulas. Diplurids in the US tend to be brown and yellowish colors, however there is a beautiful jet black species in Mexico and Diplurids from the other continents can be beautiful colors.

Public Domain Microhexura montivaga

Cyrtaucheniidae / Cyrtaucheniidae - Waferlid/Wafer trapdoors ​ ​ ​ ​ Cyrtaucheniids are limited to a single species of record in the USA, now that all genera with representatives have been moved to Euctenizidae. Cyrtauchenius talpa is listed in the World Spider Catalog as occurring in the USA, but the author was able to find no definite records to indicate which states.

Ctenizidae / Ctenizidae - Corklid trapdoors/Trapdoors ​ ​ ​ ​ Ctenizids (hint: the “C” is silent) are classic trapdoor spiders. They tend to make thick doors to their burrows out of web and surrounding substrate. This can make their burrows extremely well camouflaged and hard to find. When their burrows occur on slopes, especially in disturbed areas like around streams and rivers, the ground level can wear away leaving a little tower capped by the trapdoor. Ctenizids leg structure is such that they can hold hold their door closed with full force against their predators (or collectors). If care is taken and the substrate cooperates an entire burrow and spider can be removed by a collector. One of the most interesting genera of Ctenizids to appear in the USA is Cyclocosmia. Trapdoors in this genus have a flattened, hardened posterior abdomen which can be used to defensively plug or block their burrow.

N. M. Hentz, public domain LA Dawson CCASA2.5G* Cyclocosmia truncata, original illustration, 1875 Ummidia sp., showing classic Ctenizid shape

Euctenizidae / Euctenizidae - ????????? ​ ​ ​ Formerly a subfamily of Cyrtaucheniidae, Euctenizids tend make either wafer or cork type trapdoors. Euctenizids can be some of the most interestingly colored species of non­Theraphosidae mygalomorphs found in the United States. The genus tends to be more brightly colored and sometimes possesses an interesting herringbone pattern on its abdomen. Eucentizids can be distinguished from Ctenizids by their lack of spines on legs I and II. Eutenizid burrows can have side passages and multiple entrances and exits. The genus Apomastus does not make trapdoors at all, ending up with an open, burrow somewhat similar to Atypidae, except the external portion is typically not anchored to anything and is relatively shorter.

Marshal Hedin CCA2.5G* Andrew Olson Aptostichus sp., female Aptostichus sp., female with eggsac

Andrew Olson Marshal Hedin CCA2.0G* Apomastus kristenae, tube web Aptostichus sp., burrow entrance and door

Nemesiidae / Nemesiidae - False tarantulas ​ ​ ​ ​ Nemesiids are probably the most similar to tarantulas in looks and size of all the non­Theraphosidae mygalomorphs in North America. Species that appear in the United States all seem some form of brown and/or black in most light, but can take on a spectacular metallic coloration under the right light. While no more toxic to humans than North American tarantulas, more care should be taken as Nemesiids are much more inclined to bite and otherwise defend themselves. Nemesiids do not possess scopulae to the same degree as Theraphosids and thus typically can not climb smooth, vertical surfaces. The babies of some species, C. longitarsus in particular, can retain a lovely translucent white color for their first several instars.

Andrew Olson Andrew Olson Calisoga (Brachythele) longitarsus Calisoga (Brachythele) longitarsus, mating, male bottom

Barychelidae / Barychelidae - Brushed trapdoors/Brushfoot trapdoors ​ ​ ​ ​ An interesting family, if for no other reason than their range. This family only is represented only in Hawaii out of all the 50 states and D.C. Up until the 1980’s it was thought that Hawaii lacked any mygalomorphs but now several species are known from there. Barychelids have scopulae (the “sticky pads” on their feet) approximately equal to Theraphosids and thus can climb smooth vertical surfaces like glass.

Sean Salamon CCASA2.5G* Nihoa makina, Artist’s representation of this Hawaiian species

Theraphosidae / Theraphosidae - Tarantulas ​ ​ ​ ​ The most well known family of mygalomorphs, Theraphosids are not the subject of this paper.

A subset of the dichotomous key to Mygalomorph families from Raven 1985. Branches that are not applicable to our North American concentration are weeded out (indicated by smaller font and italics).

1 a ) Claw tufts present ...... 2 b ) Claw tufts absent ...... 9 2 a ) Anterior maxillary lobe distinct and produced (fig. 16) ...... 3 b ) Anterior maxillary lobe short, if present (fig. 34) ...... 4

3 a ) Little or no scopulae on tarsi III and leg cuticle noticeably scalelike ...... , part ​ b ) Distinct scopulae on tarsi III and leg cuticle more or less smooth ...... Theraphosidae ​

4 a ) One row or no teeth on paired claws of males or females ...... 5 b ) Two rows of teeth on paired claws of males or females...... 6

5 a ) Clypeus wide and posterior median spinnerets absent ...... Nemesiidae, part ​ ​ b ) Clypeus absent and/or posterior median spinnerets present ...... Barychelidae, part ​ ​ 6 a ) Third claw present on fourth leg ...... Cyrtaucheniidae, part ​ ​ b) Third claw absent on all legs ...... 7 7 a ) Scopulae well developed on legs I and II ...... Barychelidae, part ​ ​ b ) Scopulae on legs I and thin, if present ...... 8 8 a ) Numerous cuspules on labium or eye group subquadrate...... Barychelidae, part ​ ​ b ) No cuspules on labium and eye group rectanguloid ...... Nemesiidae, part ​ ​

9 a ) Booklung apertures small, round ...... ​ b) Booklung apertures normal, slitlike ...... 10 10 a ) Trichobothria absent on most tarsi ...... 11 b ) Trichobothria present on all tarsi ...... 12 11 a ) Anterior maxillary lobe elongate and posterior median spinnerets apically truncate ...... Atypidae ​ b ) Anterior maxillary lobe short, posterior median spinnerets apically rounded, normal ...... Antrodiaetidae ​

12 a ) Eyes in a compact group on a very steeply elevated tubercle and leg scopulae entirely absent ...... Paratropidae, part ​ b ) Eye tubercle arched, low, or absent, or leg scopulae present ...... 13

13 a ) Labium much longer than wide and eyes occupy at least half of head width ...... Actinopidae ​ b ) Labium wider and/or eyes occupy less than half of head width ...... 14

14 a ) Apical segment of PLS short, domed, or triangular (fig. 259) ...... 15 b ) Apical segment of PLS digitiform or longer (fig. 38) ...... 31

15 a ) Cheliceral fangs with two longitudinal keels and rastellum absent ...... ​ b ) Fangs smooth, rastellum present or absent ...... 16

16 a ) Males ...... 17 b ) Females ...... 23

17 a ) Median haematodocha extends down embolus and sometimes divided basally by narrow sclerotized band (fig. 243) ...... ​ , part b ) Median haematodocha small, normal ...... 18

18 a ) Cheliceral furrow with two rows of teeth ...... 19 b) Cheliceral furrow with one row of teeth ...... 20

19 a ) Caput short and arched high (fig. 258) ...... Ctenizidae, part ​ ​ b ) Caput long and arched but low (fig. 226); Africa, Madagascar, South and Central America ...... Cyrtaucheniidae, part ​

20 a ) Each paired claw with teeth on both lateral faces ...... 21 b ) Each paired claw with one row ofteeth or one tooth ...... 22

21 a ) Caput arched and fovea broad and strongly procurved ...... Cyrtaucheniidae, part ​ ​ b ) Caput low and fovea short, and weakly procurved, straight, recurved...... Nemesiidae, part ​ ​ 22 a ) Metatarsus I with distinct process or rastellum on a distinctly raised mound; North and Central America ...... Cyrtaucheniidae, part ​ ​ b ) Metatarsus I without process and rastellum consists of long spines on a low mound; Africa, Madagascar, India, Oriental region ​ ...... Nemesidae, part

23 a ) A pair of eyes (ALE) set far in advance of others so making the group longer than wide ...... Idiopidae, part ​ ​ ​ b ) Eyes not in such a long group ...... 24 24 a ) Scopula present on tarsi I ...... 25 b ) Scopula absent on tarsi I ...... 29 ​ 25 a ) Each paired claw with teeth on both faces ...... 21 b ) Each paired claw with teeth only on medial keel ...... 26 26 a ) Fovea straight or recurved ...... 27 b ) Fovea strongly procurved ...... 28 27 a ) Preening combs present; North and Central America ...... Cyrtaucheniidae, part ​ ​ b ) Preening combs absent; Madagascar, India, Australia, New Zealand ...... Idiopidae, part ​ 28 a ) Cuspules extend for full length of maxillae; North and Central America ...... Cyrtaucheniidae, part ​ ​ b ) Cuspules extend only to half length of maxillae; North and Central America ...... Idiopidae, part ​ 30 a ) Fovea strongly procurved ...... Ctenizidae, part ​ ​ b ) Fovea recurved or straight ...... Idiopidae, part ​

31 a ) Each paired claw with teeth only on medial keel ...... 32 b ) Each paired claw with teeth on both faces ...... 36

32 a ) Fovea longitudinal ...... 33 b ) Fovea transverse or absent ...... 34

33 a ) Apical segment ofPLS bipartite or pseudosegmented ...... Mecicobothriidae ​ b ) Apical segment of PLS integral ...... Dipluridae, part ​ ​

34 a ) Six spinnerets ...... , part ​ b ) Four or two spinnerets ...... 35

35 a ) Cuspules numerous on labium ...... Hexathelidae, part ​ b ) Cuspules very few or absent on labium ...... Dipluridae, part ​ ​

36 a ) PLS very long, extending higher than abdomen ...... Dipluridae, part ​ ​ b ) PLS shorter ...... 37 ​

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A NOTE ON IMAGES Unless otherwise noted, images sourced from Wikimedia Commons and are used under the Creative Commons ​ ​ ​ ​ ​ Attribution­Share Alike 3.0 Unported. No implication of the original authors’ endorsement is meant. ​

REFERENCES

Platnick’s World Spider Catalog https://research.amnh.org/iz/spiders/catalog_15.0/INTRO3.html http://www.wsc.nmbe.ch/

Wikipedia’s Mygalomorphae page https://en.wikipedia.org/wiki/Mygalomorphae

The spider infraorder Mygalomorphae (Araneae) : cladistics and systematics. Bulletin of the AMNH ; v. 182, article 1 Raven, Robert J. http://digitallibrary.amnh.org/dspace/bitstream/handle/2246/955//v2/dspace/ingest/pdfSource/bul/B182a01.pdf?sequen ce=1

A Reanalysis of Mygalomorph Spider Families (Araneae) PABLO A. GOLOBOFF http://digitallibrary.amnh.org/dspace/bitstream/handle/2246/5017/N3056.pdf?sequence=1

NATURAL HISTORY OF THE CALIFORNIAN TRAPDOOR SPIDER GENUS ALIATYPUS (ARANEAE, ANTRODIAETIDAE ) Frederick Coyle and Wendell R. Icenogle http://americanarachnology.org/JoA_free/JoA_v22_n3/JoA_v22_p225.pdf

A Reconsideration of the Classification of the Spider Infraorder Mygalomorphae (Arachnida: Araneae) Based on Three Nuclear Genes and Morphology Jason E. Bond, Brent E. Hendrixson, Chris A. Hamilton, Marshal Hedin http://www.plosone.org/article/fetchObject.action?uri=info:doi/10.1371/journal.pone.0038753&representation=PDF

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