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PSYCHE

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H. W. Levi.
Additional Notes on the Orb-Weaver Genera Araneus, Hypsosinga and Singa North of Mexico (Araneae, Araneidae).
Psyche 82(2):265-274, 1975.

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ADDITIONAL NOTES ON THE ORB-WEAVER
GENERA ARANEUS, HYPSOSINGA, AND SINGA
NORTH OF MEXICO (ARANEAE, ARANEIDAE)*
BY HERBERT W. LEVI
Museum of Comparative Zoology, H,arvard University After my first Araneus study was published in 1971 I was informed by W. J. Gertsch and V.
Roth that the males of A.
illaudatus are found with females of my newly described A. pima. However, owing to the great size difference and abdominal pattern difference, I remained stubbornly unconvinced thlat they could belong to the same species until spiderlings from an egg-sac raised by P. Witt grew up into female A. pima and male A. lllaudatus. This persuaded me but raised some new questions, as the hand-reared males are larger, the females smaller, than in wild populations (see below).
While establishing this synonymy (in Levi, 1971), I take this opportunity to report on an additional new species of Araneus from the Southwest (Levi, 1973), as well as to report range extensions from the American Museum Sings and additional Araneus collec- tions and from the R. V. Chlamberlin (University of Utah) col- lection, now again available. I will publish later on a number of tiny new species of Araneus, most collected as provisions by wasps. I would like to thank W. J. Gertsch, V. Roth, N, Platnick, of the American Museum of Natural History, S. Frommer, F. Enders and P. Witt for specimens, E. Mayr and Lorna R. Levi for reading the manuscript. National Science Foundation Grant BMS 75-057 I g supported the researches and publication. For unknown reasons, size differences between sexes and among individuals of the same sex within populations are much greater in orb-weavers in the superf'amily Araneoidea than in other spiders, although adult size variations are common in arachnids. These dif- ferences, due to variation in the number of instars before maturity, have been commented on by others (e.g. Gerhardt, 1929, 1930; Probst, 1972 in Isometrus scorpions).
*Manuscript received by the editor September 10, 1975 265
Pu&e 2:26S-27 t 1975). http:llpsyche cnlclub orBig11-26 html



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Psyche [June
Since large differences in size may be accompanied by allometric differences it often becomes very difficult to decide whether a large and a small female belong to the same species. Also, the large and
small specimens have differences in numbers of macrosetae. In addition, in the Araneidae there are often large differences in size between the sexes as the dwarfed males mature already after only a few molts, sometimes after fewer than half the number in females. Larger males, having undergone more molts, may resemble females more than they resemble smaller males (Gerhardt, 1929). Oddly, the smallest female and one of the larger males of A. illaudatus came out of the same egg-sac, hand-raised in P. Witt's laboratory in North Carolina. This poses the question, to what extent the variable number of instars is environmentally determined? The egg-sac came from a building of the Southwestern Research Station near Portal, Arizona, where males are usuially small and females large.
According to older literature the minute size of many male orb- weavers prevents suitors from being mistaken for prey by th,e large female. This idea, which originated with Charles Darwin and Fried- rich Dahl (Gerhardt, 1929)) is not convincing to me. In Gerhardt's experiments neither large nor small males were attacked. It is much more likely that the differences in size, especially in warm climate spiders, is a secondary by-product of selection for different rates of ontogeny. A quick maturation of males, etc. is an adaptation 'against inbreeding. The tiny males mature after a few molts, perhaps months before sibling females undergo their 8 to 11 instars, and are forced to look for mature females from earlier broods or even from a different generation. [Witt (personal communication) reports that in his cultures of Araneus diadematus males mature long before -
females.] But then why do small males come in various sizes (as in A. iZZaudatus), and why do hand-raised females vary in size? Is there genetic polymorphism for rate of development? -
-- ---
Figs. 1-4. Left palpus, apical view. 1. Araneus gemmoides Chamberfin and Ivie. 2. Araneus gemma (McCook). 3, 4. Araneus illaudatus (Gertsch and Mulaik). 3. (Portal, Arizona). 4. (hand-raised specimen, Portal, Arizona).
Figs. 5-11. Araneus abigeatus new species. 5-8. Epigynum. 5. Ventral view, cleared. 6. Ventral view. 7. Posterior view, cleared. 8. Posterior view. 9-11. Left palpus. 9. Mesal. 10. Ventral. 11, Embolus in mesa1 view. Size indicators, 0.1 mm.




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Levi - Araneus, Hypsosinga, Singa
267




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Psyche
Araneus illaudatus (Gertsch and Mulaik)
Figures 3, 4
Aranea illaudata Gertsch and Mulaik, 1936, Amer. Mus. Novi'tates, no. 863, p. 19, figs. 36, 37, 8. Male holotype from Edinburg, Texas in the American Museum of Natural History, examined. Araneus pima Levi, 1971, Bull. Muse Comp. Zool., vol. 141, p. 176, plates 1, 4, figs. 218-232,
?, 8, map 8. Female holotype from Madera Canyon, Santa Rita Mountains, Pima Co., Arizona in the American Museum of Natural History. NEW SYNONYMY.
Araneus iZlaudatus,~Levi, 1971, Bull. Mus. Comp. Zool., vol. 141, p. 176, figs. 238-240, $ (not figs. 233-237, 9 which is A. ablgeatus, n. sp.). Note. Some A. illaudatus males had shorter embolus caps than others. The distribution maps (Levi, 1971) of A. illaudatus and A. pima should be combined.
Diagnosis. Males of the three similar species can be separated by the length of the terminal ~apophysis in apical view. Araneus gem- moides Chamberlin and Ivie has the shortest, widest terminal apophy- sis (Fig. I ) and largest palpal tibia ( 1971, figs. 199-200). Araneus gemma (McCook) has a pointed, curved terminal apophysis, widest at the base (Fig. 2). Araneus illaudatus has a long curved terminal apophysis, the sides more or less parallel (Figs. 3, 4). Size ranges.
Total length of females varied I 1.5 to 27 mm. The smallest female was hand-raised from Chihuahua Mts., Arizona and another collected 'at Gateway, Oregon, both 11.5 mm. Total length of California females in mm: 14, 17, 22; Utah: 13, 23; Arizona, Fort Grant: 22; Huachuca Mts.: 18, 18, 18, 20, 23; Chihuahua Mts. : 13, 19, 21, 22, 23 ; Santa Rita Mts. : 18, 20, 24; Grand Canyon : I 5 ; Texas : 14, 18 ; Mexico, Chihuahua : 18. Not al females were measured, only one from each collection. There appears to be no geographic pattern.
Total length of males 2.9 to 7.9 mm. Measurements of males in mm: Utah, Watson : total length 4.2, carapace 1.9 long, 1.5 wide; Arizona, Santa Rita Mtns.: total length 7.9, carapace 4.1 long, 3.0 wide; Chihuahua Mts.: total length 2.9 to 3.9 mm, carapace I .6 to I .g long, I .2 to I .6 wide, three specimens; Texas : total length 3.6 to 4.0 mm, carapace 1.9 to 2.0 long, 1.3 to 1.5 wide, two speci- mens; specimen from Chihuahua Mts. hand-raised in North Caro- lina: total length 6.6 mm, carapace 3.6 long, 2.8 wide (Fig. 4). All males were measured; the 6.6 mm and 3.8 mm males are illus- trated in Figures 3, 4.




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Lev; - Araneus, Hypsosinga, Sings
Araneus abigeatus new species
Figures 5-1 I
Araneus illaudatus, - Levi, 1971, Bull. Mus. Comp. Zool., vol. 141, p. 176, figs. 233-237, 9, epigynum and abdomen; not figs. 238-240, 8. Holotype. Female from Rustler Camp, Chiricahua Mts., Cochise Co., Arizona, 9 Sept. 1950 (W. J. Gertsch) in the American Museum of Natural History. The name abigeatus is a Latin verb for cattle stealing, rustling.
Description. Female. Carapace dark brown, marbled on lighter brown with some light setae. Sternum dark brown, coxae light. Legs banded brown on yellow-white with some white-tipped setae. Dor- sum of abdomen with a median longitudinal line of white spots and four pairs of dark brown chevrons, the posterior pair fused in mid- dle on dark brown speckled background ( 1971, fig. 236). Venter with a black band from epigynum to spinnerets, and a pair of white spots side by side. Spinnerets dark brown. Eyes subequal in size. Anterior median eyes slightly more than their diameter apart. The abdomen has a pair of anterior dorsal humps. Total length I I .O mm. Carapace 3.7 mm long, 3.5 wide. First femur, 4.5 mm; patella and tibia, 5.5 ; metatarsus, 3.7; tarsus, 1.4. Second patellca and tibia, 5.0 mm ; third, 2.9 ; fourth, 4.9.
Male. Coloration similar to that of female, except carapace more uniform light brown. Thorax with a longitudinal groove. Eye size and spacing as in female. First coxa with a hook, second femur with groove. Second coxa with a small posterior, proximal tubercle, no spur. Second tibia swollen with macrosetae on venter. Total length 7.0 mm.
Car,apace 3.8 mm long, 3.2 wide First femur, 5.0 mm; patella and tibia, 6.5 ; metatarsus, 3.6; tarsus, 1.3. Second patella and tibia, 5.0 mm ; third, 2.9; fourth, 4.5. Diagnosis. The constricted neck of the scape of the epigynum covering a large hollow space (Figs. 6, 8) separates this species from Araneus illaudatus. The internal genitalia (Figs. 5, 7) 'are heavily sclerotized and were cleared but not macerated as only few speci- mens were available.
The male will key out to A. washington! in Levi ( 197 I ) but the palpus is similar to that of A. nordmanni. The paalpus has a terminal apophysis prong similar to that of A. nordmanni but differs by the shape of the shorter embolus (Fig. I I) and shape of conductor (Figs. 9, 10).
Records. Known from type locality only. Paratypes collected on 25 Aug. 1952, I 9 (B. Malkin), 23 Aug. 1968, I Q (V. Roth), 4 Aug. 1973, $2 , c? (S. Frommer) .




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270 Psyche
A ranens washingtoni Levi
Additional Record. Ontario : Batchawana, 6. Araneus santarita (Archer)
Additional Record. Arimna: Cochise Co.: Upper Cave Creek, Chiricahua Mts. (V. Roth).
Araneus pratensis ( Emerton)
Additional Records. Ontario : Thessalon. Mississippi: Wilkin- son Co. : Centreville.
Araneus mammatus (Archer)
Additional Record. New Mexico: Santa Fe Co.: Hyde Memorial State Park, 8 miles northeast of Santa Fe. Araneus mariposa Levi
Additional Record.
Oregon: Jackson Co: Sams Valley to Dead
Indian Spring, 2 Q (J. Schuh) .
Araneus apache n. sp.
Figures I 2-20
Type. Male holotype with a female paratype from Rustler Camp, Chiricahua Mts., g Sept. 1950 (W. J. Gertsch) in the American Museum of Natural History.
Description of female paratype. Carapace yellow-white with a wide median longitudinal brown band pointed posteriorly toward thoracic depression. Sternum brownish black. Legs with dark bands. Coxae yellow-white. Dorsum of abdomen with transverse marks as in Figure 17. Venter brownish black with transverse white mark behind epigynum (Fig. 16). Posterior median eyes subequal to anterior medians. Anterior laterals 0.8, posterior laterals 0.9 di- ameters of anterior medians. Anterior median eyes 1.4 di'ameter apart, 1.6 from laterals. Posterior median eyes slightly less than their diameter apart. The height of the clypeus is less than diameter of anterior median eyes. Abdomen is almost round, dorso-ventrally slightly flattened. Toea1 length 7.5 mm. Carapace 3.1 mm long, 2.4 wide. First femur, 3.2 mm; patella and tibia, 4.2; metatarsus, 3.0; tarsus, 1.0. Second patella and tibia, 3.4 mm; third, 2.0; fourth, 3.0




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Figs. 12-20.
Araneus apache new species. 12-15. Epigynum. 12. Ventral view, cleared.
13. Ventral view. 14. Posterior view, cleared. 15. Posterior view. 16. Female abdomen, ventral. 17. Female carapace and abdomen, dorsal. 18-20. Left male palpus. 18. Subventral view. 19. Ventral view. 20. Embolus, subventral view.
Fig. 21.
Hypsosinga pygmaea (Sundevall), dorsal view of genitalia. Size indicators, 0.1 mm, except Figs. 16, 17, 1.0 mm.



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272 Psyche [June
Males. Males are less distinctly marked than females. The eye size ratios and distances are about the same as those of female. The first coxa has a hook, the second tibia has macrosetae but is not swollen. Total length 4.0 mm. Carapace 2.3 mm long, 1.9 wide. Second patella and tibia, 3.0 mm; third, 1.7 ; fourth, 2.5. Variation. There is little variation in size of the few specimens available. The dorsum is much lighter in some individuals than in others; and in the pale specimens, the median carapace band and the dark spots on the abdomen barely show.
All, however, have la dark
sternum.
Diagnosis, This species is close to Araneus pegnia (Walckenaer), but males differ by having three prongs on the median apophysis (Fig. 18) rather than two and in details of the shape of other distal palpd structures (Figs. 18-20). Females differ from those of A. pegnia by having a wider scape (Fig. 13) and 'also a brownish black sternum and brownish black venter of the abdomen surrounding a transverse white mark (Fig. 16). Males differ from those of A. thaddeus, which have a similar three-pronged median apophysis, by the very differently shaped embolus
(Fig. 20), bearing a worm-
shaped lamella.
Such an embolus lamella is present in A. pegnia also, but is not illustrated in Levi (1973, figs. 436, 437). The species keys out to 3, A. pegnia and A. thaddeus, in my key to females of smlaller species of Araneus (Levi, 1973, p. 488) and to 3 A. thaddeus in my key to males of smaller species of Araneus (Levi, 1973, P. 489)
Note. This is not A. montanus Archer. The holotype of A. montanus was re-examined and is A. thaddeus as synonymized before (Levi, 1973). Females of A. apache
I had previously placed in
A. pegnia. Araneus pegnia does not seem to occur in southeastern Arizona; it is found in California and Sonora (Levi, 1973, map 6). Records. Paratypes from Arizona. Cochise Co. : Chiricahua Mts., 29 Aug. 1938, (R. H. Crandall). Graham Co.: Pinecrest, Graham Mt., 13 Sept. 1950, 2 $
(W. J. Gertsch), Shannon Camp.
Graham Mt., 13 Sept. 1950, 5 $2 (W. J. Gertsch). Pima Co.: Santa Catalina Mts., 5 Sept. 1938, 2
Q (R. H. Crandall).
Singa keyserlingi McCook
Additional Records. New Jersey. Ocean Co. : Lakehurst. Bur- lington Co. : Lebanon State Forest. Minnesota. Goodhue Co. : Lake Pepin. Carlton Co. : Carlton.




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19751 Levi - Araneus, Hypsosinga, Sings 273 Singa eugeni Levi
Additional Records. New York. Kings Co.: Bergen Beach (Brooklyn). New Jersey. Atlantic Co. : Oceanville. North Garo- Una. Carteret Co. : Beaufort.
Hypsosinga pygnzaea ( Sundevall )
Figure 21
Theridion pygmaea Sundevall, 1831, Kongl. Svenska Vet. Akad. Handl. p. 121. Type from Sweden, assumed lost.
Singa variabilis Emerton, 1884, Trans. Connecticut Acad. Sci., vol. 6, p. 322, pi. 34, fig. 16, pi. 37, figs. 19-21, 9, 5. Five syntypes from New
Haven, Connecticut in the Museum of Comparative Zoology. NEW SYNONYMY.
Singa pygmaea, - Wiehle, 1931 in Dahl, Tierwelt Deutschland, vol. 23, p. 47, figs. 64, 65, 9, 5. Roewer, 1942, Katalog der Araneae, vol. 1, p. 875. Locket and Millidge, 1953, British Spi,ders, vol. 2, p. 155, figs. 102q 103b, 104b, ?, 5.
Hypsosinga variabilis,-Levi, 1972, Psyche, vol. 78, p. 242, figs. 44-57, 9, 5, map 2.
Note. Mr. G. H. Locket recently lent me British specimens of this species and they turned out to be the same as our common H. variabilis. There are only minor, probably geographic, differences. Distribution. The distribution of this species is holarctic. The synonymy of 13. variabilis with H. pygnzaea was overlooked pre- Additional American Records. Nova Scotia. Berwick. Tennes- see. Lake Co.: Reelfoot Lake. Georgia. Chatham Co.: 3 mi. south- east of Savannah. Nebraska. Loup Co.: Taylor. Wyoming. Yel- lowstone Natl. Park : Yellowstone Lake. Grand Teton Natl. Park: Moran. Utah. Salt Lake Co.: Salt Lake City. Cuba. Pinar del Rio : Vifiales.
Hypsosinga f unebris ( Ke~serling )
Cercidia funebris Keyserling, 1893, Spinnen Amerikas, vol. 4, p. 37, pi. 2, fig. 32, 9. Female holotype from Crescent City, Florida, Marx collection of the United States National Museum kept in the American Museum of Natural History, examined.
Araneus singaeformis Scheffer, 1904, Entomol. News, vol. 15, p. 259, pi. 77, figs. 4-6, 9. Female syntypes from Wallace County, Kansas in the Museum of Comparative Zoology.
Hypsosinga singaeformis, - Levi, 1972, Psyche, vol. 78, p. 246, figs. 58-71, 9, 5. Map 3.




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Additional Reds. Alberta. Medicine Hat. Maine. Acadia Natl. Park: Mt. Desert Island. Connecticut. South Meriden. New Jersey. Ocean Co.: Lakehurst. Tennessee. Bedford Co.: Shelby- ville. Minnesota Carlton Co.: Barnum. Texas. Kerr Co.: Kerr- ville. Hildalgo Co. : Edinburg. Utah. Salt Lake Co.: Alta. Cali- f ornia. Monterey co. : Monterey.
Hypsosinga alberta Levi
Additional Record. Yukon Territory. South shore, Atlin Lake, willow, heath, dwarf shrubs, 20 Aug. 1974, $ (D. E. Bixler).
Hypsosinga groenlandica Simon
Additional Record. Yuklon Territory. Herschel, 24 July I 971 (W. R. M. Mason). Utah. Cache Co.: Franklin Basin, 16 July 1975, I 9 (G. F. Knowlton) .
GERHARDT, U.
1929. uber Grossenvarianten der Mannchen von Nephila madagas- cariensis Vinson. 2001. Anz. vol. 86, p. 80-82. 1930. Biologische Beobachtungen an Nephila madagascariensis Vins. Sitzungsber. Gesellsch. naturforsch. Freunde Berlin, p. 358-360. LEVI, H. W.
1971. The Diadematus Group of the Orb-Weaver Genus Araneus North of Mexico (Araneae: Araneidae). Bull. Mus. Comp. 2001. VO~. 141, p. 131-179.
1972. The Orb-Weaver genera Sings and Hypsosinga in America (Araneae: Araneidae). Psyche, vol. 78, p. 229-256. 1973. Small Orb-Weavers of the Genus Araneus North of Mexico (Araneae: Araneidae). Bull. Mus. Comp. 2001. vol. 145, p. 473- 552.
PROBST, P.
1972. Zur Fortpflanzungsbiologie und zur Entwicklung der Giftdriisen beim Skorpion Isometrus maculatus (DeGeer, 1778). Acta Trop- ica, vol. 29, p. 1-87.




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