Herbert W. Levi.
The Orb-Weaver Genera Singa and Hypsosinga in America (Araneae: Araneidae).
Psyche 78:229-256, 1971.

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THE ORB-WEAVER GENERA SINGA AND 4HYPSOSINGA IN AMERICA (ARANEAE : ARANEIDAE)*
BY HERBERT W. LEVI
Museum of Comparative Zoology, Harvard University The North American spiders commonly placed in Skga belong to four different genera. It was first thought best to publish on these species together with the small species of Araneus. However, the slowness with which the Araneus studies proceed makes it advisable to publish on Singa and Hypsosinga first. Most of the specimens of these genera belonging to the American Museum appear to be lost and only small collections other than those of the Museum of Comparative Zoology were available. I would like to thank the following for loan of additional specimens: Dr. J. A. Beatty; Mr. D. E. Bixler; Mr. D. Buckle; Dr. J. A. L. Cooke for collections of the American Museum of Natural History and Cornell University; Dr. R. Crabill for specimens from the U.S. National Museum; Dr. B. Cutler; Dr. C. D. Dondale of the Canada Dept. of Agriculture; Dr. B. J. Kaston; Dr. W. W. Moss for collections from the Academy of Natural Sciences, Phila- delphia; Dr. W. Peck for specimens from the Exline collection; Dr. J. Pr6szyhski for collections of the Polish Academy of Sciences; Miss Susan Riechert; Mr. V. Roth; Dr. W, Shear; Dr. C. Ti-iple- horn of Ohio State University; Prof. S. L. Tuxen for specimens from the Zoological Museum of the University of Copenhagen; Dr. J. D. Unzicker for specimens from the Illinois Natural History Survey; Prof. M. Vachon and Dr. M. Hubert for collections of the Museum National d'Histoire Naturelle, Paris; Dr. "R. Vogel; and Dr. H. I<. Wallace. I am also indebted to Mr. J. Denis, Mr. G. Ptihringer, and especially to Miss Susan Riechert for helpful information. The research and publication were supported by Public Health Service Grant AI-01944 from the National Institute of Allergy and Infectious Diseases.
Shga C. L. Koch
Singa C. L. Koch, 1836, Arachniden, vol. 3, p. 42. Type species Singa hamata (Clerck) designated by Thorell, 1869. On European Spiders, p. 58. The name is of feminine gender.
Diagnosis. The anterior median eyes are the largest, the posterior medians the same size or smaller, the laterals about 0.6 diameter of the anterior medians. In Sings the median ocular quadrangle is wider *Manuscript received by the editor February 8, 1972. 229




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230 Psyche [December
in front (Figs. 20, 30) ; in Hypsosinga it is rectangular or wider behind. The carapace of the female is shiny and has no thoracic depression (Figs. 22, 32) ; in the male it has a short longitudinal line. The first coxa of the Singa male, unlike that of the Hypso- singa male, has a hook on the side. The height of the clypeus is equal to or less than the diameter of the anterior median eyes (Figs. 20, 30) ; in >Hypsosinga it is higher. The second tibia of the Sings male may be modified; in Hypsosinga the first tibia may be modified. In Sings the sides of the abdomen are almost parallel and the abdomen overhangs the spinnerets. There are two dorsal longi- tudinal black bands on the abdomen of Singa (Figs. 21, 22, 31, 32). The female genitalia differ from those of Hypsosinga in that Singa has a scape on the epigynum (Figs. 6, 1 I, 25). The palpus of the Singa male has an enormous terminal apophysis (A in Figs. 3, 5, 9), ending in a sclerotized spine and located in the contracted palpus on the outer side of a rather narrow tegulum (Fig. 5). Below the terminal apophysis, a heavily sclerotized hook, perhaps the subterminal apophysis (SA in Figs. I, 3, IO), lies hidden in the contracted bulb. The embolus may have a lamella (Figs. I, 9) ; it does not seem to have a part that breaks off in mating. The stipes (I in Fig. I) is a distinct sclerite in S. hamata, but is apparently fused to the embolus in S. keyserlingi
(Fig. 9). The tegulum is
widest near the base of the conductor (Figs. 2, 3, 10). Natural History. Surprisingly little is known about the habits of Sings. They make a complete orb. Nielsen (1932, Biology of Spiders, 2; fig. 330) has a picture of the retreat of S. hamata. Both American species prefer moist locations, and adults are found through- out the season. G. Puhringer (personal communication) told me that Singa phragnziteti Nemenz is common on reeds along the Neu- siedler See in Austria. It prefers a site above water and has to be collected from a boat. I suspect that American species have similar habits, which may account for the few specimens in collections. Distribution. Species are known only from Eurasia and temperate North America. All others described are probably misplaced. Misplaced American Species. (This list of names follows Roewer, 1942, Katalog der Araneae. The types of the species, unless indicated otherwise, have been examined. Species placed in Hypsosinga are not listed. )
abbreviata Keyserling, I 879 = Theridiosomatidae. k r y i Archer, 1958 = Metepeira bengryi (Archer), new com- bination.




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19711 Le-vi - Orb-Weaver Genera 23 1
calix Walckenaer, 1841 = Alpaida calix, new combination. crewii Banks, 1903; the type is lost, the description is not recog- niza ble.
dotana Banks, 1914 = Theridion dotanum.
d~odecim~uttata Keyserling, 1879 = Alpaida duodecimguttata (Keyserling), new combination.
erythrothorax Taczanowski, 1873 = Alpaida erythrothorax (Tac- zanowski) , new combination.
essequibensis Mello-Leitiio, 1948 = ?Alpaida essequibensis, type specimens unavailable.
ftava 0. P.-Cambridge, 1894 = Araneus ftavus (0. P.-Cam- bridge).
ftoridana Banks, 1896 = Araneus fioridana (Banks). leucogramma White, 1841 = Alpaida leucogramma (White), new combination.
listerii McCook, I 893 = Araneus pratensis Emerton. longicauda Taczanowski, I 878 = generic placement uncertain. marm ota Taczanowski, I 873 = Alpaida marmorata (Taczanow- ski), new combination.
maura Hentz, 1847 = Alpaida calix Walckenaer. nzoesta Banks, 1893. The type has been destroyed, the descrip- tion cannot be recognized.
mollybyrnae McCook, 1893 = Metazygia pallidula (Keyserling). NEW SYNONYMY. Type locality in error; not District of Colum- bia, probably Colombia.
niveosigillaa Mello-LeitiTo, I 941 = ?Alpaida niveosigellata, type specimens unavailable.
pratensis Emerton, I 884 = Araneus pratensis (Emerton) . paticola Simon, 1895 = Araneus pratensis (Emerton). tremens Holmberg, 1876. The type has been destroyed. vanbruysseli Becker, I 879 = Cyclosa turbinata (Walckenaer) , NEW SYNONYMY.
vittata Taczanowski, 1873. The type is lost. The two common species north of Mexico, calix and patensis, are obviously misplaced. Judging by the structure of the genitalia, Singa pratensis is close to Araneus sturmi (Hahn) of Europe. Dif- ferences are the shape of the abdomen and lack of body setae, but the shape, setation, and coloration of the abdomen are quite variable in the many species of these small Araneus. Araneus sturmi is the type species of the genus Atea. At present it does not seem wise or even feasible to fragment the genus Araneus. It would lead to




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232 Psyche [December
proliferation of names without meaning for relationships. They are obviously monophyletic.
Sinpa caZix belongs to a South American genus, one of the largest genera of orb weavers in the Americas. As far as I know at present, the oldest name is Alpaida 0. P.-Cambridge, 1889. But numerous
other generic names have been used for this genus; Lariniacantha Archer (1951, Amer. Mus. Novitates, no,. 1487, p. 15) most re- cently. The genus is much closer to Awnthepeira than to Sings. However, I am still hestitant about the placement until I have more knowledge of the webs and habits of the species in the genus. Key to American Species of Sings
la. Base of epigynum trapezoidal, with sides sclerotized (Figs. II- 14) ; median apophysis of palpus with one hook (Figs. 23, 24) . . . . . . . . . . . .. . . . . . . . . , . . , . . , . . , , ,. . . . . . . ....... , . , . . . . keyserZingi ib. Base of epigynum with a lobe on each side (Fig. 25) ; median apophysis of palpus with two hooks (Figs. 33, 34) ........ eugeni Singa hamata (Clerck)
Figures 1-8
Araneus hamatus Clerck, 1757, Aranei Svedici, p. 51, pi. 3, fig. 4. Female
type specimens from Sweden believed lost. Bonnet, 1955, Bibliographia
Araneorum, vol. 2, p. 513.
Singa hamata,- C. L. Koch, 1836, Die Arachniden, vol. 3, p. 42, figs. 197, 198, 9, 8. Wiehle, 1931, in Tierwelt Deutschlands, vol. 23, p. 42, figs. 54-57, 9, 5. Roewer, 1942, Katalog der Araneae, vol. 1, p. 873. Locket and Millidge, 1953, British Spiders, vol. 2, p. 157, figs. 102b, 103c, 105c, 9, 5.
This species, very similar to the two American ones, is known only from Eurasia.
Sings keyserlingi McCook
Figures 9-24, Map I
Singa keyserlingi McCook, 1893, American Spiders, vol. 3, p. 230, pi. 19, fig. 2, 9. Female holotype from St. Louis, Missouri, in the Academy of Natural Sciences, Philadelphia; examined and labelled as type. Singa campestrls Emerton, 1915, Trans. Connecticut Acad. Sci., vol. 20, p. 153, pi. 3, fig. 3, 8. Male syntype from Rat Portage, Ontario, in the Museum of Comparative Zoology; examined. NEW SYNONYMY. Note. McCook described S. keyserlingi as a new species. How- ever, in the first paragraph of the description he finds it necessary "to propose a new name" for the species Keyserling illustrated and called erroneously Sings rubella (Hentz). The specimens McCook



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19711 Levi - Orb-Weaver Genera
233
Figs. 1-8. Singa hamafa (Clerck). 1-5. Left palpus. 1-3. Expanded. 4. Mesal view. 5. Ventral view. 6-8. Epigyaum. 6. Ventral. 7. Posterior. 8. Dorsal.
Figs. 9-10, Singa keyseriwgi McCook, palpus, expanded. Abbreviations: A, terminal apophysis; C, conductor; DH, distal hema- todocha; E, embolus; H, basal hematodocha; I, stipes; M, median apophysis; P, paracymbium ; R, radix ; S, subtegulum ; SA, subterminal apophysis; T, tegulum; Y, cymbium.
Size Indicators: 0.1 mm.




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234 Psyche
[December
had on hand and figured were specimens of what is here considered to be S. keyserlingi, and not what Keyserling called rubella. In the United States National Museum there is a specimen marked Singa rubella Hentz, St. Louis, Mo., collected by Marx. Another label in the vial reads "Cotype 1688 U.S.N.M." The specimen contained is a female of what is here called S. eugeni, and there is no evidence that it is a syntype of S. keyserlingi Mc'Cook. McCook reports having seen specimens from the District of Columbia in the Marx collection. Keyserling also reports that the Singa rubella he examined
came from the District of Columbia from the Marx collection.
Figure 13 was prepared from the type of S. camfiestris. Description. Female from Wisconsin. Carapace orange ; head region, clypeus black.
Chelicerae, labium, endites dark brown. Ster- num orange. Legs orange, distal articles darker. Dorsum of abdo- men with two wide black bands separated by a narrow white band (Fig. 22). Sides of black bands have lateral white band (Fig. 21). Venter has a black patch which may have a white line on each side. Anterior median eyes slightly more than one diameter apart, one diameter from laterals. Posterior median eyes their radius apart, two and one-half diameters from laterals. Total length 5.5 mm. Carapace 2.4 mm long, I .7 mm wide. First femur, I .4 mm ; patella and tibia, 2.1 mm; metatarsus, 1.3 mm; tarsus, 0.7 mm. Second patella and tibia, I .9 mm; third, 1.3 mm; fourth, 1.7 mm. Male from Wisconsin. The coloration is like that of the female except that black on the head seems limited to the eye region. The
chelicerae are black only distally, orange at the base. The dorsum
of the abdomen is all black. The anterior median eyes one and one- third diameters apart, one diameter from laterals. Posterior median eyes their radius apart, two and one-half diameters from laterals. The palpal patella has two weak macrosetae. The first and second
legs have strong macrosetae on prolateral surface, but are not bent or swollen.
Total length 3.9 mm. Carapace 2.2 mm long, 1.6 mm wide. First femur, 1.5 mm; and tibia, 2.0 mm; metatarsus, 1.4 mm; tarsus, 0.7 mm.
Second patella and tibia, 1.8 mm; third, 1.2 mm; fourth, 1.5 mm.
Variation. In the female the scape of the epigynum varies greatly in length and shape (Figs. 11-15) ; the internal ducts may be either heavily sclerotized or transparent, sac-like and difficult to make out. Females vary from 5.1-6.0 mm in total length, carapace I .5-I .7 mm



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19711 Levi - Orb-Weover Genera 235
Figs. 11-24,
Singa kfysfrhgi McCook. 11-22. Female. 11-19, Epigyniim. 11-14. Ventral 11. (Illinois). 12. (Wisconsin). 13. (Ontario). 14, 15. (South Dakota). IS. Posterior. 16-19. Cleared. 16. Subventral. 17. Ventral. IS. Dorsolateral. 19. Posterior. 20. Face. 21. Abdomen, lateral. 22. Dorsal. 23-24. Left palpus. 23. Maal. 24. Ventral. Size indicator^: 0.1 mm, except for Figs. 21-22, 1 mrn.



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236 Psyche [December
wide.
Males vary from 2.3-4.0 mm in total length, carapace I .3-I .6 mm wide.
Diagnosis.
Females differ from those of 8. eugeni in the shape of the base of the epigynum, trapezoidal with the lateral margins sclerotized (Figs. I I- I 4) . The male differs from that of S. eugeni by having only one hook on the median apophysis (Figs. 23, 24). Natural History.
Sings keyserlingi has been collected from open woods, in low shrubs and by sweeping grass on lakeshores. Mature males have been collected in all months between May and August. Females have been collected through September. Distribution.
From Edmonton, Alberta, Smoky Falls, Ontario to Black Warrior National Forest (Winston Co.), Alabama (Map 1).
Singa eugeni sp. n.
Figures 25-34, Map I
Singa rubella,-Keyserling, 1893, Spinnen Amerikas, vol. 4, p. 284, pi. 14, fig. 209, 9. Not Epeira rubel1,a Hentz. Type, Male holotype and female paratype from T8N, R5E. SgNWg, Iowa County, Wisconsin (Susan Riechert), in the Mu- seum of Comparative Zoology. The species is named after Count Eugen Keyserling.
Description. Female. Carapace orange with a wide black band covering eye region (Fig. 32), narrowing behind. Clypeus black. Chelicerae brown-black. Labium black. Sternum yellowish with dark brown margin. Legs yellow. Dorsum of abdomen with two longi- tudinal dark bands; at each end the bands are darker and approach each other (Fig. 32). The bands are separated by a white pigment line.
The sides are white (Fig. 31) ; the venter is yellowish with an indistinct dark area in the middle. The anterior median eyes are one and one-quarter diameters apart, less than one diameter from laterals.
Posterior median eyes are less than one-quarter diameter apart, one and one-half diameters from laterals. Total length 4.6 mm. Carapace 2.0 mm long, 1.4 mm wide. First femur, 1.3 mm; patella and tibia, 2.2 mm; metatarsus, 1.2 mm; tarsus, 0.6 mm. Second patella and tibia, I .g mm; third, 1.2 mm ; fourth, I .7 mm. Male.
The coloration of the male is like that of female. The carapace is narrower in front than in the female. The anterior median eyes overhang the chelicerae. The anterior median eyes are more than one diameter apart, their radius from laterals. The pos-



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19711 Levi - Orb- Weaver Genera 237
1 I
U! /å
I I /' Singa keyserlingi
Map. 1. Distribution of Sd'nga kcyscrlingi (McCook) and S. cugeni sp. n.



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238 Psyche [December
terior median eyes are one-quarter diameter apart, one and one-half diameters from laterals. The second tibia is thick, slightly curved, with macrosetae on prolateral side, but only very slightly modified. The palpal patella has one strong and one very weak seta. There are eight black, sclerotized dorsal muscle attachments on the abdo- men. Except for the narrower carapace, the male looks like the female. Total length 5 mm. Carapace 2.2 mm long, 1.6 mm wide. First femur, 2.2 mm; patella and tibia, 3.4 mm; metatarsus, 2.3 mm; tarsus, 0.9 mm. Second patella and tibia, 2.9 mm; third, 1.4 mm; fourth, 1.9 mm.
Variation. In some individuals only four black spots remain of the two abdominal bands, two anterior and two posterior. If present the soft projection from the terminal apophysis, seen in ventral view (Fig. 34), may be either a flap or a rod. Females are from 4.3- 6.5 mm in total length, carapace I .I-I .6 mm wide; males are from 3.6-5.4 mm in total length, carapace 1.2-1.8 mm wide. Diagnosis. Females are distinguished from those of S. keyserlingi by the smaller epigynum with a lobe on each side of scape (Fig. 25), as opposed to a sclerotized diagonal margin seen in S. keyserlingi. The median apophysis of the palpus has two hooks, one on each end (Figs. 33, 34). In the related S. nitidula of Europe the median apophysis is of different shape and the embolus narrower. Natural 'History. Wisconsin specimens came from open bottom- land forest, along backwaters of river edges, and edge of marsh; Georgia specimens came from Spartma stems. Barrows ( 191 8, Ohio J. Sci., 18: 310) reported that a Sinpa from Cedar Point, Ohio, almost certainly this species, made "a small orb in tops of dune grass (Andropogon). During the day it stays in the hollow stems of dead grass." The spiders have also been collected in Penn- sylvania by the wasp Efiisyron quinquenotatus (Say). The males are mature in September and October in the north. Adult females
have been collected from May to October. Localities collected. Pennsylvania; Erie Co.: Presque Isle State Park. Ohio. Erie Co.: Cedar Point. D.C. Washington. Georgia. Mclntosh Co. : Sapelo Island. Michigan. Clinton Co. : Rose Lake. Eaton Co. : Calumet. Livington Co. : George Reserve. Midland Co. Wisconsin. Iowa Co.: 9, 8 paratypes. Jefferson Co. (Map I).
Hypsosinga Ausserer
Ifypsosinga Ausserer, 1871, Verhandl. 2001. Bot. Gesell. Wien, vol. 21, p. 823 (subgenus). Type species Singa sunguinea (C. L. Koch)



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Levi - Orb-Weaver Genera 239
\"
Figs. 25-34, Siaqa euqeni sp. n. 25-32. Female Ventral. 26. Posterior. 27-29. Cleared. 27. Dorsal. 28. Ventral. 29. -
Posterior. 30. Face. 31. Abdomen, lateral. 32. Dorsal. 33-34, Left palp~~s. 33. Mesal. 34. Ventral.
Six Indicators: 0.1 mm, except fcr Figs. 31, 32, 1 mm.



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240 Psyche [December
designated by Petrunkevitch, 1911, Bull. Amer. Mus. Natur. Hist., vol. 29, p. 275. The name is of feminine gend,er. Note. Both Wiehle ( 1931 ) and Roewer's Katalog der Araneae spell the generic name Hyposinga. Roewer changed the spelling of many generic names from long accustomed usage to the spelling of that of the original author. But here Roewer changed the original spelling. Ausserer consistently spelled the name with "s" and also indicated that a main character of the subgenus is the high clypeus (hypo, Greek for high).
Diagnosis. I3ypsosinga differs from Sings in having the posterior median eyes the largest ( 1.2-2 diameters of anterior median eyes, Figs. 52, 64, 93, 105). The ocular quadrangle is wider behind than in front, or rectangular. The clypeus height in Hypsosinga is 1.5 to 3 diameters of the anterior median eyes (Figs. 52, 64, 81, 105), but only about one diameter of the anterior median eyes in Singas It is always slightly higher in males than females. As in Singa, but unlike most Araneus, I.ypsosinga has the carapace smooth and rather wide in front, wider than the eye area (Figs. 54, 66) ; there is no thoracic depression, or sometimes a small longitudinal black mark in the male. Unlike many Araneus, the males of the North American species have no hooks on the first coxae. The first tibiae of males of I$. singaeformis and H. groenlandica are swollen (Fig. 71). In many araneids, it is the second tibia that is modified. The abdomen in Hypsosinga unlike that of Singo tends to be oval, widest in the middle, with either two dorsal longitudinal bands or four dark spots (Figs. 54, 66, 83, 95, 106). Like Sings, unlike Armeus, Hypsosinga frequently has the eye region black. The epigynum differs from those of both Sings and Araneus in lacking a scape (Figs. 49, 61, 102). The palpus differs from that of Sings in having a smaller terminal apopthysis (A) and a spur on the ventral
face of the tegulum
(Figs. 35, 36, 39). Hypsosinga
differs from all other genera of Araneidae in having a large tsans- parent scale attached to the base of the embolus (Fig. 69) ; the scale breaks off in mating and lodges in the epigynum (Figs* 99, 100; Levi, 1972). A scar remains on the embolus (Figs. 70, 98). The median apophysis is small in all species (M) Figs. 35, 38) 39). The palpal patella has two setae.
Description. All species are quite simila,r in general appearance and unlike Neoscona species differ more from each other in genitalic differences than in abdominal patterns. The carapace is orange, lacking hair; eye region black, and rarely in individuals, the black



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19711 Levi - Orb-Weaver Genera 241
may extend to a point on thorax) or only surround eyes (H. groen- Zandica, Fig. 93). Sternum ora,nge to dark brown. Legs orange, rarely with longitudinal lines in H. singaeformis, and very rarely banded in some individuals of H. rubens. Juveniles may have white pigment spots on dorsum of carapace and sometimes on sternum. Abdomen with little hair and with two longitudinal black bands indistinctly separated by a lighter area, but fused posteriorly (Figs. 66, 83). Bands very distinctly set off toward lighter sides (Figs. 53, 65, 82). The dark bands are usually darkest anteriorly and pos- teriorly. There may only be four black spots on dorsum, or they may be compIetely missing; sometimes abdomen entirely black. The sides may have an additional black band (Fig. 53). Venter of abdomen black with a white longitudinal line on each side (Figs. 53, 94). Males have abdominal pattern Iess distinct. Smaller individuals of 12. rubens and females of H. dberta may have the dorsal muscle spots sclerotized. Total length of females is 2.4-.5.1 mm, of maIes 2.2-3.5 mm.
Natural History. Hytsosinga make a complete orb probably with a retreat. A11 species are most commonly collected by sweep- ing vegetation. Males are mature in spring) the females throughout the season.
Distribution. Species of Hypsosinga are only known from Eurasia, North Africa and North America.
Keys to American species of Hypsosinga
Female epigynum with a median depression (Fig. 102). MaIe unknown ; Western Canada ............................................ alberta Female epigynum with a median, raised septum (Figs. 49, 61, go) ............................................................................................... 2 Epigynum with septum having sides almost parallel and septum about one-third width ,of epigynum
(Fig. 49) ; male embolus