Herbert W. Levi.
The Spider Genera Steatoda and Enoplognatha in America (Araneae, Theridiidae).
Psyche 69:11-36, 1962.

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THE SPIDER GENERA STEATODA AND
ENOPLOGNATHA IN AMERICA*
(ARANEAE, THERIDIIDAE)
BY HERBERT W. LEVI
Museum of Comparative Zoology, Harvard University The previous revisions (Levi, 1957a, 19570) of the two genera Enoplognatha and Steatoda considered only the North American species. Since the revisions were published, large South American collections have become available and the types of South American species could be consulted.
The majority of species of both genera are found in the north temperate zone and are fairly well known. The additional species described here from the ne~t~opical area are sometimes intermediate between the two genera. Enoplognatha jvruviana may lack the tooth on the posterior margin of the chelicerae as in Steatoda species. The males of several Steatoda (e.g. 8. andina) have the chelicerae enlarged as is characteristic of Enuplognatha. South American Enoplognatha species are found only in southern Peru and northern Chile (Map I). The genus has no representatives in Central America or the West Indies. Steatoda species are found in all parts of South America, with several endemic species and several that are widespread (8. ancorata, S. grossa, S. muesta).
Unlike Anelosinzus species (Levi, in press) Steatoda species cross the desert or mountain barrier into Chile (Map 2).
The types of species could be examined through the hospitality and cooperation of Dr. G. Owen Evans and Mr, D. Clark of the British Museum (Natural History) ; Prof. G. C. Varley of the Hope Depart- ment of Entonlology, Oxford; Dr. L. Brundin of the Natural History Museum, Stockholm; Prof. M. Birabkn of the Museo de la Plata; Prof. M. Vachon of the Mu&um National d1Histoi1-e Naturelle, Paris; Mr. J. Pr6szyriski of the Polish Academy of Sciences, Warsaw; Dr. L. Forcart of the Naturhistorisches Museum, Basel and Dr. M. Beier of the Naturhistorisches Museum, Vienna. I would like to thank the following for the loan of specimens for study: Prof. M. Vachon, Mus&m~ National dlHistoire Naturelle (MNHN) ; Mr. J. Kekenbosch of the Institut Royal des Sciences Naturelles, Brussels (ISNB) ; Dr. W. J. Gertsch of the American Museum of Natural History (AMNH) ; Dr. 0. Kraus of the Senckenberg Museum, *Manuscript received by the editor November 20, 1961. Pu&e W:l1-36 (1962). hup ttpsychu einclub orgt6WW-Ol Lhtml



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12 Psyche [March
Frankfurt (S^VIF) ; Dr. E. S. R.oss of the California Academy of Sciences (CAS) ; Dr. H. Exline (311-s. D. Frizzell) for specimens from northern Peru and Ecuador collected by herself and others; Dr. R. X. Schick of California; Miss H. Zapfe (Mrs. G. Mann) of Santiago de Chile; Dr. J. Abalos of Santiago del Estero and Prof. H. Stahnke of Arizona. A trip to European museums to exam,ine types was made possible by a National Science Foundation Grant (G-4317) and the research was supported by a grant from the
National Institutes of Health (E-1944).
E no plognatha Pavesi
Enoplognatha Pavesi, 1880, Ann. Mus. Civ. Stor. Nat. Geneva, vol. 15, p. 325. Type species by original designation and monotypy heri id ion mandibulart Lucas.
Diagnosis.
Medium sized to large theridiid spiders. Enoplognatha, like Steatoda, has a large colulus and a subspherical abdomen. Females have a tooth on the posterior margin of the chelicerae. Males usually have enlarged chelicerae, with enlarged teeth and have the paracym- bium on the margin of the cymbium (Levi and Levi, 1962). Species misplaced. Enoplognatha Mia Chamberlin, I 9 I 6 = Ane- lositnus studiosus ( Hentz, I 850) . E noplopatha triangulifera Simon, I 902 = hlosimus recurvatus ( Tullgren, 1 90 I ) . juninensis
peruviana
pun0
zapfei
Map 1.
Distribution of South American Enoplognatha species. Key to American Species of Enoplognatha
I a. Species found in South America . . . . . .. . . . . . .. . . . . . . . .... .. . . . . . . . . . . ... . . 2 i b. Species found in north of Mexico .
. .. . . . . . . .. . . .. . , ,... .. .... . .... . .. . . 5



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19 621
2a.
2b.
3a.
3 b.
4a.
4b.
5 a.
5b.
6a.
6b.
7a.
7b.
8a.
8b.
9a.
9b.
I oa.
lob.
I la.
nb.
I 2a.
Levi - Steatoda and Enoplognatha
13
E~igynum with a large central depression (Figs. 12, 13, 15) 3 Epigynum otherwise .................................................................. 4 Epigynum with central depression surrounded by a lip (Figs. .................................... 12, I 3) ; ducts coiled (Fig. I I ) ; Peru .......................................................... E. jmninensis (Keyserling) Epigynum with central depression lacking a lip posterior (Fig. ............
15) ; ducts with a sclerotized loop (Fig. 14) ; Chile E. mpfei n. sp.
.............................................................................. Fused ducts extending anterior and surrounding seminal i-ecep- tacles (Fig. 9) ..................................................... E. pun0 n. sp. Fused ducts not extending anterior of seminal receptacles (Fig. 7) .................................................. E. peruviana Chamberlin Yellow-white coloration with black or red lines on carapace and abdomen (1957a, fig. 4) ; introduced in northeastern and Pacific coast states ......................................... E. ovata (Clerck) Gray, brown or black coloration .............................................. 6 ........................................................................................ Males 7
Females .................................................................................... 14
Lacking paracy~nbiumt on cyrnbial margin (Fig. 4) ; Arizona, southeastern California .................................. E. maricopa n. sp. With paracymbium on cymbial margin .................................... 8 Conductor with a stalked distal apophysis ( 1957a) fig. 13-16) 9 Conductor otherwise ......................................................... 10 Conductor apophysis widened distally and truncate ( 1g57a, figs. I 5, I 6) ; California ................... .å£ se/?na Chambei-lin and Ivie Conductor apophysis pointed distally ( Ig57a, figs. 13, 14) ; introduced in Oregon ................................ E. thoracica (Hahn) Chelicesae with two subequal teeth on posterior margin ( I 957a, figs. 24, 25) ............................................................................ I1 Chelicerae with one large mesa1 tooth (lg57a, figs. 46-48) 12 Abdomen with a spotted dorsal pattern (1957a) fig. 33) ; median apophysis in ventral view only in distal half of alveolus (1957a, fig. 26) ; widespread throughout United States and southern Canada, Mexico ............................... E. ~~zurmorata ( Hentz) Abdomen with a foliate dorsal pattern (1957a, fig. 37) ; median apophysis in ventral view extending into proximial half of alveolus ( 195721, fig. 28) ; east of Rocky Mountains to New- foundland ................................................. E. tecta (Keyserling) Ectal podon of conductor with long axis almost parallel to axis of cymbium (1957a, figs. 42-45) ; western states, rare in east ....................................... E. joshua Chamberlin and Ivie



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14 Psyche
[March
Long axis of ectal portion of conductor at almost right angles ..............................
to axis of cymbium ( I 957a, figs. 38, 40) I 3
Embolus more proximal, conductor with a longer ectal portion (figs. 38, 39) ; South Dakota to Utah at lower elevations ........ ............................................ E. wyuta Chamberlin and Ivie Embolus more distal, conductor with a shorter ectal portion (1957a, figs. 40, 41) ; Alaska to Greenland, in Rocky Moun- tains above timberline and in spruce-fir forest, Minnesota, Pennsylvania. New England . E. intrepida ( Smensen) Epigynum with a light posterior transverse bridge (1957a, fig. 55) ; western states, rare in east E. Joshua Chamberlin and Ivie Epigynu~ni otherwise ........................................................... I 5 Epigynum with an oval depression having dark marks of openings on the anterior of the depression (1957a, fig. 20) ; California E. sdma Chamberlin and Ivie
Epigynum otherwise 16
Epigynum with a transverse swelling, with a seam on the anterior and openings on each end of seam ( 19j7a. fig. 18) ; introduced in Oregon
E. thoracica ( Hahn)
Epigynum otherwise 17
Epigynu~n with distinct dark marks indicating openings I9 Epigynum with diffuse dark marks I 8
Swelling of epigynum divided by a transverse groove ( I 95 7a, figs. 31, 32), spotted dorsal abdominal pattern ( I957a. fig. 33) ; widespread throughout United States and southern Canada, Mexico E. marinorata (Hentz)
Epigynum swelling undivided with only a slight depression posterior ( I 957a, figs. 35, 36), foliate dorsal pattern of abdomen (1957a, fig. 37) ; east of Rocky Mountains to Newfoundland E. tecta (Keyserling)
Dark mark of epigynum anterior to a light swelling (Fig. 2) ; Arizona, southeastern California
E. maricopa 11. sp.
Dark mark anterior to a transverse ridge 20
Posterior edge of dark mark of epigynum straight ( I 95 7a, fig. 50) ; South Dakota to Utah at lower elevations E. wyuta Chamberlin and Ivie
Dark mark dumb-bell shaped ( 1957a, fig. 52) ; Alaska to Greenland, in Rocky Mountains above timberline and in spruce- fir forest, Minnesota, Pennsylvania, New England E. in trepida ( Serensen)




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19621 Levi - Steatoda and Enoplognatha 15 Enoplognatha selnza Charnberlin and Ivie Additional record. California. Los Angeles Co.: Big Tujunga Canyon, San Gabriel Mts., 3 June 1955 (R. X. Schick). Enoplog'natha marmorata ( Hentz) and
Enoplognatha tecta (Keysel-ling)
Although these are common species in the eastern United States, we still have few observations on their habits. Mature E. tecta have been collected on a garage door in Cambridge, Massachusetts and numerous specimens were collected rolled up in leaves in herbaceous vegetation along forest edge about 40 cm above ground in southern Minnesota, in an area where E. marmorata was collected under boards in gardens.
Additional records of E. marmorata. Canada. British Columbia: Muskeg near Little Prairie (D. Rounds). Mexico. Hidalgo: Guer- rero Mill (W. M. Mann).
Enoplognatha maricopa new species
Figures 1-5
Type. Male from Phoenix, Maricopa County, Arizona, I I March 1960 (H. L. Stahnke) in the Museum of Comparative Zoology. Description. Male carapace, sternum, legs red-brown. Abdomen mottled purplish. Sclero'tized portion of epigastric area and plate above pedicel on abdomen red-brown. Carapace with a circular thoracic depression. Anterior median eyes smallest, anterior laterals largest. Anterior median eyes a little less than a diameter apart, their radius from laterals. Posterior median eyes a little less than a diameter apart, one diameter from laterals. Chelicerae heavy but not elongated, with one large anterior tooth (Fig. 5). Total length 3.4 mm. Cara- pace I .8 mm long, 1.3 mm wide. First patella and tibia 1.4 mm ; second 1.3 mm; third 1.0 mm. Fourth femur I .3 mm; patella and tibia I .6 mm; metatarsus I .2 mn1 ; tarsus 0.5 mm. Female carapace, sternum and legs mottled brown or yellowish. Abdomen with a black pattern on dorsum as in Enoplognatha wynta Chamberlin and Ivie (1957a, fig. 53). Anterior median eyes smaller than others. Anterior medians one and one-half diameters apart, one diameter from laterals. Posterior median eyes one diameter apart, a little more than one diameter from laterals. Chelicera with one anterior tooth and three denticles between tooth and base of fang.



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16 Psyche [March
Posterior margin of chelicera with one small tooth (missing on one side of one specimen). Total length one female 2.9 mm, another 3.8 mm.
Carapace of the latter 1.5 mm long, I .2 mm wide. First femur 1.6 mm; patella and tibia I .8 mm; metatarsus I .o mm~; tarsus 0.8 mm. Second patella and tibia I .5 mm ; third I .I mm ; fourth I .8 mm. Comments. It is not certain that the male and female described here belong together.
However, since large collections of spiders are available from southern and southeastern Arizona and no speci- mens of Enoplognatha have been found, I assume that the distribution of Enoplognatha maricopa is north from the type locality, where the females were found, a poorly collected area. Diagnosis. The palpus differs from other North American species in that there is no paracymbium on the margin of the cy~nbiumi (Fig. 4). However, the transparent edge of the cymbium is difficult to see. On the mesa1 side of the bulb is a sclerite, probably part of the conduc- tor, which has two projections, one toward the venter and one toward the distal end of the palpus
(Fig. 3). The female differs from
Enoplognatha intrepida (Smensen) in that it has two dark openings connected by a black groove; posterior to the groove is a swelling having a lighter center (Fig. 2).
Records. California. Placer Co., near Emigrant Gap, 8 July 1956, 2 9 (V. Roth, W. J. Gertsch, AMNH) ; Tahoe City, 8 July 1956, 9 (W. J. Gertsch, V. Roth, AMNH).
Enoplognatha peruviana Chamberlin
Figures 6-8
Enoplognatha teruviana Chamberlin, 1916, Bull. Mus. Comp. Zool., 60: 232, pi. 16, figs. 8-11, 52. Female type from Urubamba, 3100 m alt., [Cuzco], Peru, in the Museum of Comparative Zoology, examined. Description. Carapace, sternum, legs red-brown. Abdomen black with a dorsal pattern as in Enoplogmtha tecta (Keyserling), ( 1g57a, fig. 37). Anterior median eyes two-thirds diameter apart, one diameter from, laterals.
Posterior median eyes two-thirds diameter apart, one and one-half diameters from laterals. Anterior median eyes slightly smaller than others. Cheliccra with three large teeth on anterior margin, the first and third larger than the middle one; none on posterior margin of the type. However, another specimen seems to have the posterior tooth. Colulus relatively small with two setae.
Total length of female type 6.5 mm. Carapace 3.6 mm long, 2.4 m~m wide. First femur, 2.7 nlm; patella and tibia, 3.4 mm;



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19621 Levi - Steatoda and Enoflognatha 17 metatarsus, 2.0 mm ; tarsus, I. I mm. Second patella and tibia, 3.0 mm ; third, 2.3 mm; fourth, 3.2 mm.
The male illustrated (Fig. 6) may belong to this species; this is uncertain, as is the locality where it has been found. Records. Peru. "Pampa Machei, 4400 m," Feb. 1947, d' (Bi-echt, AMNH). Cuzco: Cuzco, 3500 m, July 1947, Q (W. Weyrauch, AMNH).
Enoplognatha puno new species
Figures 9, 10
Type. Female from 32 km north of Desaguadero, Puno, Peru, 27 Feb. 1951 (E. I. Schlinger, E. S. Rcss), in the California Acad- emy of Sciences. The specific name is a noun in apposition, after the type locality.
Description. Carapace, sternum, legs dark brown. Abdomen brownish black with an indistinct dorsal pattern as in E. peruviana. Venter with some white pigment on each side. Anterior median eyes much smaller (two-thirds their diameter) than oval posterior eyes. Anterior median eyes two-thirds diameter apart, more than their diameter from laterals. Posterior median eyes less than their shorter diameter apart, two and one-half times their shorter diameter from laterals. Chelicerae with three large anterior teeth and a minute denticle on posterior margin. Total length 6.1 mm. Carapace 2.6 mm long, 2.0 mm wide. First femur 2.3 mm; patella and tibia 2.9 mm ; metatarsus I .7 mm ; tarsus I .O mm. Second patella and tibia 2.5 mm; third 2.1 mm; fourth 3.0 mm.
Diagnosis. Only the epigynum (Fig. 10) and ducts of the internal genitalia (Fig. 9) separate this species from E. peruviaw; the pattern and the structure are very similar. It is possible that this species belongs to a geographical race of E. peruviana, Enoplognatha juninensis (Keyserli'ng)
Figures I I - I 3
Lithyphantes juninensis Keyserling, 1884, Die Spinnen Amerikas, Theridiidae 2(1) : 143, pi, 6, fig. 90, 9. Female lectotype here designated from Maraynioc, Junin, Peru in the Polish Academy of Sciences, Warsaw, examined.
Enoplognatha juninensis, Simon, 1894, Histoire Naturelle des Araignies, 1 : 578.
The specimen examined from, Pasco, Peru has the abdomen dark gray with a light area on each side on dorsum ; the venter is also dark



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I 8 Psyche [March
gray with a light area on each side between cpigynurn and spinnerets. The chelicerae have three large teeth on the promargin, a minute Figs, 1-5,
Enoplognutha maricopa new species. I. Female genitalia, dorsal view. 2. Epigvnuni. 3, 4. Left palus. 3. Ventral view. 4. Ectal view. 5. Left male chelicera, posterior view,
Figs. 6-8. E. ficruwiana Charnberlin. 6. Palpus (doubtful determination). 7. Female genitalia, dorsal view. 8, Epigynum. Figs. 9-10. E. puno new species. 9. Female genitalia, dorsal view. 10. Eoievnum.
~'iis. 11-13. E. juÌömens'- (Keysding). It. Female genitalia, dorsal view. 12, 13. Epigynum.
Figs. 14-15. E. sapfci new spec&. 14. Female genitalia, dorsal view. IS. Epigynum.
tooth on the posterior margin. The colulus is large, almost trans- parent white, with two setae from the base. Records. Peru. Pasto: near Huayllay, 4400 rn, 10 Aug. 1953,



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19621 Levi - Steatoda and Enoplopatha I9 0 (H. W. Koepcke, SMF) . Junin. Maraynioc, 0 paratype with 9 paratype (BMNH) .
Enoplognatha zapfei new species
Figures 14, I 5
Tyw Female from Putre, Province Tarapac;, Chile, Feb. 1948 (Avilis), in the Museum of Comparative Zoology. The species is named after Miss H. Zapfe of Santiago de Chile. Description. Carapace dark yellow, darker aroitnd anterior median eyes. Chelicerae red-brown. Sternum, legs dark yellow. Abdomen gray with darker gray mottled pattern on doi-sum and a pair of longitudinal lighter bands between genitalic area and spinnerets. Anterior median eyes slightly smaller than others. Anterior lateral eyes on slight tubercles. Anterior median eyes slightly less than one diameter apart, a little more than one diameter from laterals. Posterior median eyes three-quarters diameter apart, two diameters from laterals. Chelicerae with three teeth on anterior margin, a blunt indistinct tooth posterior. Total length 9.2 mm. Carapace 4.5 mm long, 3.4 mm wide. First femur, 4.1 mm; patella and tibia, 5.0 mm; metatarsus, 3.4 mm ; tarsus, I .6 mm. Second patella and tibia, 4.3 mm ; third, 3.7 mm; fourth, 5.1 mm-
Diagnosis. This large species of Enoplognatha can be separated from E. juninensis by the lack of posterior lip in the opening of the depression of the epigynum (Fig. 15) and by lack of the internal coiled duct. The ducts of E. zapfei are heavily sclerotized adjacent to the seminal receptacles (Fig. 14).
Steatoda Sundevall
Steatoda Sundevall, 1833, Conspectus Arachnidurn, p. 16. Type species desig- nated by Thorell, 1869, On European Spiders, p. 93. 8. castanea (Clerck, 1757).
Diagnosis. Medium-sized to large theridiid spiders, usually dark in coloration. Colulus very large. Chelicerae armed with a tooth or several teeth on anterior margin.
But unlike Enoplognatha, it
has no teeth on the posterior margin of female chelicerae. Abdomen subspherical.
Males often with enlarged chelicerae and a sclerotized ring around pedicel on anterior end of abdomen. The abdomen of
most species has a white line around anterior of dorsum, in addition to other lines or spots (Levi and Levi, 1962). Note. The common species of North America were described in



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20 Psyche [March
a previous paper (Levi 1957b). In this paper the ventral view of the genitalia (cleared epigynum) was illustrated, not the dorsal as in most of my papers on theridiid spiders. Gertsch (1960) has taken issue with my treatment of species of the "fulva" group in the previous paper and has named several new species. However, insufficient ecological and life history data are available for the possible species involved to pel-m.it making a final decision now (Levi, 1960). Thus the only species recognized are those of my previous revision. Steatoda brasiliana Keyserling, I 884, Die Spinnen Amerikas Thei-idiidae, 2 ( I ) : I I 5, pi. 5, fig. 75, d. Male type from Rio de
Janeii-o, Brazil in the Naturhistorisches Museum, Vienna, examined is a male Steatoda bipunctata (Linn.) from Europe (NEW SYN- ONYMY). The female described by Keyserling in 1886 as 8. brasil'iana is probably a specimen of S. moesta (Keyserling). Species misplaced: Lithyphantes juninensis Keyserling = Enoplog- natha jununensts (Keyserling) .
Steatoda rubra Keysel-ling, 1886, Die Spirinen Amerikas, Theri- diidae, 2(2) : 239, pi. 20, fig. 294, 9. Female type from Blumenau, Brazil = Theridion rubra (Keyserling) .
Key to American Species of Steatoda
I a. Epigynurn with a prominent, wide, transverse bridge ; areas anterior and posterior to bridge shallow depressions ( 1g57b, figs. 43, 44, 46, 47, 49, 50,) ; palpus with thread-like embolus on venter, evenly curved, with a membrane inside embolus loop from base supporting a portion of embolus ( 1957b, figs. 32-41 ) ; North America .................................................. 5. fulva group ib. Epigynum without transverse bridge or if with bridge, then areas anterior and posterior to' it not depressed; embolus other- wise or if similar South American ......................................... 2 2a. Seminal receptacles thickened, sclerotized, coiled ends of ducts (1957b, figs. 88, 92, 98, 116) ; epigynum having a U-shaped depression ( 1957b,, figs. 105, 107- 109) 0,s a shallow circular depression containing anterior a deeper depression on each sid,e of a septum (rarely absent) ( 1957b, figs. 93, 97, 99-103) or epigynum as in 1957b, fig. 89; palpus with a prominent U- shaped radix on mesa1 side (1957b, figs. I 19, 124, 129, 134, 145, I 55) ; North America ....................... 8. bipunctata group Map 2.
Distribution of South American Steatoda species.



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22
2b.
3 a*
3b*
4a*
4b.
9.
5b.
6a.
6b.
7a-