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THE ORB-WEAVER GENERA ARANIELLA and NUCTENEA (Araneae: Araneidae)

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THE ORB-WEAVER GENERA ARANIELLA and NUCTENEA (Araneae: Araneidae) THE ORB-WEAVER GENERA ARANIELLA AND NUCTENEA (Araneae: Araneidae) HERBERT W. LEVI Abstract. The included species in Araniella and berg Museum; C. Dondale and R. Leech, Nuctenea have traditionally been included in Ara- Canadian National collections, Ottawa; W. neiis, but males differ in lacking the embolus cap. G. H. M. Martelli of The lack of embolus cap can be related to differ- Hackman; Locket; ences in mating behavior. Those orb-weavers witii the Zoological Museum of the University cap (Amnetis) can mate only once with each of Florence; W. W. Moss of the Academy Nuctenea males a can mate palpus; lacking cap of Natural Sciences, Philadelphia; W. Peck, several times. Exline-Peck collection; R. X. Schick of Four species of Araniella are known, one of the California of diem Holarctic, the others Palearctic. Some Euro- Academy Sciences; J. pean populations are of interest as there are indi- Proszynski and W. Star^ga of the Polish cations that the species hybridize. Academy of Sciences; B. Vogel; H. K. Wal- Of the si.x known to to species belong Nuctenea, lace and H. V. Weems, Florida State Col- three are Holarctic, and three Palearctic. One of lection of Information was the Holarctic species may be a recent introduc- Arthropods. T. of tion to North America; another may be cosmopoli- provided by Kronestedt the Natural tan. History Museum, Stockholm; M. Moritz of Other to these two species belonging genera the Zoological Museum of the Humboldt may be hidden among the two thousand si^ecies University, Berlin; and F. Wanless and D. placed in Araneus and mostly poorly described. Newman of the British Musevmi, Natural Lorna R. Levi and Lm INTRODUCTION History. Mackay edited the paper. This investigation and Araniella and Nuctenea species have its publication were supported in part by traditionally been placed in Araneus. They National Science Foundation grant number include our commonest orb-weavers. Nev- GB-3616L ertheless the species are not well known, One of the striking and puzzling features and in looking through the collections of these common orb-weavers is the enor- available, I found that many specimens mous individual variation in genitalic were misidentified. structiu-e (Figs. 8-15, 17, 18, 67-76'), while The species of both genera are mainly there is little variation in the size and shape Palearctic with some Holarctic species. of the whole animal. This variation is I would like to thank the following for found in Holarctic Araniella (Usplicata and providing specimens for this study: D. A. cucurhitina of Europe as well as in Bixler; W. J. Gertsch and J. A. L. Cooke of Holarctic Nuctenea patagiata and Holarctic the American Museum of Natural History, N. cornuta. Not infrequently specimens of Cornell and Utah University collections; these common species are sent to the E. J. Carico; M. Grasshoff of the Sencken- museum by collectors who believe them to Bull. Mas. Comp. Zool., 146(6) : 291-316, November, 1974 291 292 Bulletin Museum of Comparative Zoology, Vol. 146, No. 6 be a new species. In their unusual varia- and the United States. To judge by the tion, Araniella and Nuctenea contrast with labels, N. cornuta is less dependent on the small Aroneus species (Levi, 1973). houses in Alaska and probably competes Even though the species are fairly wide- with the native Araneus species. In north- spread, the differences between small eastern America all three Nuctenea species Araneus species are far less than is found are usually found on buildings, but this among individuals of a single collection of is not true throughout their ranges. Nu- N. cornuto. The larger-sized Araneus ctenea patagiata may be found under bark species (A. nordmanni, A. saevus) are in- in woods. termediate in this 1971 . is at times respect ( Levi, ) It most unfortunate that Large variation was not found in any of names have to be changed as a result of the widespread, common thcridiids such revisionary studies. as Achaearanea tepidarioriim, although it does occur in the Tidarren species and Araniella Chamberlin and Ivie ovata. Variation in the Enoplognatha AmnicUa Chamberlin and Ivie, 1942, Bull. Univ. genitalic structure of individuals was Utah, biol. ser., 7(1): 76. Type species Epeira The found in the zone of overlap between displicata Hentz, by original designation. name is of feminine Araneus gemma and Araneus gemmoides; gender. all evidence indicates hybridization (Levi, Note. Chamberhn and Ivie (1942) do 1971 . Araniella also not reasons for E. ) species may hybridize give separating displicatus in Europe. Perhaps the individual varia- from Araneus other than that the species tion of Araniella cucurhitina of Europe is is close to those of Neoscona. I agree with due to separation of northern and southern this opinion. populations during the Pleistocene and the Diagnosis. There are no good superficial later hybridization occurred after the reced- characters that separate female Araniella ing of the ice. In none of the species of from the small species of Araneus. Females Araniella or Nuctenea is the variation have a glabrous carapace and an oval geographic. But I have not studied various abdomen, widest in the middle, lacking populations in detail. Shortly after Pet- setae and lacking a folium pattern, but iimkevitch 1925 wrote on the remarkable ( ) usually with paired black spots (Plate 1, variation of genitalia of Agelena naevia, Figs. 1, 16). The epigynum has a short, Seyler (1940), following up a hint from wide, wrinkled scape (Figs. 8, 25, 34, 40). Gertsch (1934), correctly found that what The scape is not always clearly set off from Petrunkevitch called one species was in the base of the epigynum. Unlike Araneus fact several. But I doubt that populations species, however, Araniella has, besides a of Araniella displicata or Nuctenea patagiata single pair of seminal receptacles, a pair and N. cornuta consist of sibling species. of sclerotized sacs (Figs. 7, 27, 33, 39) Of considerable interest is the relation- between the external entrance from outside ship of Araniella species in Europe. Mr. to the connecting duct and the seminal Locket made me aware of this. While A. receptacles. The entrance of the duct to inconspicua and A. alpica appear distinct the sclerotized sacs on each side is a slit. on the Continent, intermediates are found The ducts are hard to see in very sclerotized in Great Britain. epigyna (A. cucurhitina). While in many American araneid and Unlike other araneids I have examined, theridiid species specimens from the Gulf Araniella has three macrosetae (Figs. 31, Coast and Florida are noticeably smaller 42) on the patella of the male palpus. than those from other parts of North Amer- Species of most genera have only two or ica, Alaskan specimens of N. cornuta are one. However, the palpal femur has a smaller than those from southern Canada basal ventral tooth facing a tooth on the Orb-weaver Auaniella and Nuctenea • LcxA 293 endite as in Araneus. The palpus resembles that of Neoscona (Berman and Levi, 1971, fig. 31) in having the sclerites nearly fused, a small flap-like terminal apophysis (A in Figs. 20-22), and in laekiug a distal hematodoeha (Fig. 21). This eontrasts with the huge terminal apophysis and distal hematodoeha in Araneus and Nu- ctenea (Figs. 58, 61). AranicUa speeies also differ from Neoscona and Araneus in the hook-shaped, selerotized, median apoph- the M , directed toward ysis ( ) dorsally eymbium (Figs. 20, 22), and in the un- usually complex, large conductor (C). The embolus (E) lacks a cap. An embolus cap is always present in virgin males of Araneus species; the cap breaks off in mating. Araniella males have a hook on the distal margin of the first coxa, and the second femur has a matching depression. The legs of males are longer than those of females, and bear many macrosetae. Most Araniella are green or yellow to reddish in color when alive, similar to small species of Araneus. The color, except the black spots, washes out in alcohol. All species are about the same size and pro- portions (Figs. 1, 16). Natural history. All species build a small web between leaves; the web may be liorizontal. The spiders are active at day- time. The egg sac has a loose woollen ap- pearance (Plate 1).^ Distribution. The known species are all Palearctic, except A. displicata which is Plate 1. Araniella displicata (Hentz). Above, female Holarctic in distribution. with egg sac. Below, male. Both laboratory photo- graphs, from Massachusetts. Misplaced species. Araniella geayi Ca- ' which I liad anticipated that Araniella males, poriacco, 1954, Comm. Pontifica Acad. Sci., resemble males of Nuctenea species in lacking a 16: 104 from Guyana has as a holot^'pe a cap on the embolus, would likewise be able to juvenile specimen belonging to the genus mate repeatedly. Therefore, it was with consid- Araneus in the Mu- erable interest that, after submitting my own deposited Zoological I R. Blanke Neue of ex- manuscript, received ( 1973, seum of the University Florence, Ergebnisse zum Sexualverhalten von Araneus cu- amined. ctirbitinus, Forma et Functio, 6: 279-290). iilanke did indeed observe differences between the Key to Species of Araniella behavior of A. cucurbitinus and that of Araneus species: The female approaches the male, the 1 Male, conductor of palpus with a distal male taps the tarsi of females, and the male mates lobe directed at and close to palpal pa- several times. tella (Figs. 37, 42); female with epigy- 294 Bulletin Museum of Comparative Zoology, Vol. 146, No. 6 Map 1. North American distribution of Araniella displicata (Hentz). niim scape having proximal part widest, than part of base showing on each side much wider than tip (Figs. 34, 40) — _ 2 of it (Figs.
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