Herbert W. Levi.
American Neoscona and corrections to previous revisions of Neotropical orbweavers (Araneae: Araneidae).
Psyche 99:221-240, 1992.

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AMERICAN NEOSCONA AND CORRECTIONS TO
PREVIOUS REVISIONS OF NEOTROPICAL ORB-WEAVERS (ARANEAE: ARANEIDAE)
Museum of Comparative Zoology
Harvard University
Cambridge MA
One of the first revisions of American araneids was that of Neoscona (Berman and Levi, 1971). At the time I was undecided whether to revise the Nearctic and Neotropical species together or separately. Soon it became evident that, although undesirable, a geographic division had to be made. Most of the Nearctic species were well represented in collections but there were only a few specimens of the Neotropical species. Collections of the American Museum of Natural History and the Museum of Comparative Zool- ogy were the main ones used. At present large collections of Neotropical spiders are available on loan which we did not have in 1970. Grasshoff's revision (1986) of African Neoscona found that Africa and America share some species but that Africa has species belonging to the subgenus Afraranea which are not found in the Americas. But since most of the Neoscona species are found in the area of eastern Asia and western Pacific, our knowledge of Neoscona species is still limited. Neoscona species are easily transported and are part of the unfortunate homogenization of the world fauna. I suspect that the new Neoscona marcanoi from His- paniola, and N. oaxacensis (Keyserling) may be the only native species found in the American tropics. This paper is an addition to the previous revision (Berman and Levi, 1971). The records of specimens included in previous maps (Berman and Levi, 1971) have not been individually listed; most came either from the AMNH or MCZ collections. In the present paper, the previous localities are plotted again, together with the new records, which are listed together with their collections. p~
Manuscript received I5 May 1992
22 1




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222 Psyche [vo~. 99
This paper also includes corrections and additions to previous monographs. After completion of a generic revision, I invariably get specimens sent to me that cannot be determined with my keys and illustrations. Most turn out to be new species. Also I find omissions, errors and misprints. However, the omission of a previ- ously described species, and perhaps one whose genitalia have not been illustrated before, may lead to a redescription as a new species and a possible later synonymy. Also some species turn out to be misplaced, a discovery made only after a specimen of the other sex becomes known. This also has to be corrected. Credits. I would like to thank the curators of the following museums and the individuals who loaned specimens. AD, A. Dean, Texas A and M University, College Station, Texas AMNH, American Museum of Natural History, New York; N. Platnick, L. Sorkin
BMNH, The Natural History Museum, London, England; P. Hillyard, F. Wanless
CAS, California Academy of Sciences, San Francisco; W. J. Pulawski, D. Ubick
CD, C. Deeleman, Ossendrecht, Netherlands CV, C. Valderrama A., Bogota, Colombia
FSCA, Florida State Collection of Arthropods, Gainesville, Florida; G. B. Edwards
IBNP, Inventario Biologico Nacional, Paraguay; J. A. Kochalka INPA, Institute Nacional de Pesquisas da Amazonia, Manaus, Est. Amazonas Brazil; J. A. Raphael
JAK, J. A. Kochalka, Ciudad Universitaria, Paraguay JMM, J. M. Maes, Leon, Nicaragua
MACN, Museo Argentine de Ciencias Naturales, Buenos Aires, Argentina; E. A. Maury
MECN, Museo Ecuatoriano de Ciencias Naturales, Quito, Ecuador; L. Aviles
MCN, Museu de-Ciencias Naturais, Porto Alegre, Rio Grande do Sul, Brazil; E. H. Buckup
MCZ, Museum of Comparative Zoology, Cambridge, Massachu- setts
MEG, M. E. Galiano; Buenos Aires, Argentina MLP, Museo de Universidad Nacional, La Plata, Argentina; R. F. Arrozpide




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19921 Levi 223
MNHNC, Museo Nacional de Historia Natural, Habana, Cuba; G. Alay On
MNSD, Museo Nacional de Historia Natural, Santo Domingo, RepUblica Dominicans
MNRJ, Museu Nacional, Rio de Janeiro, Brazil; A. Timotheo da Costa
MUCC, Museo de Biologia, Universidad Central, Caracas, Venezuela, J. Racenis
MUSM, Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru; D. Silva D.
MZSP, Museu de Zoologia da Universidade de Siio Paulo, S5o Paulo, Brazil; P. Vanzolini, L. Neme, J. L. M. Leme PAN, Polska Akademia Nauk, Warszawa, Poland; A. Riedel, W. Starega, J. Proszynski, A. Slojewska, E. Kierych SMF, Forschungsinstitut Senckenberg, Frankfurt am Main, Germany; M. Grasshoff
USNM, National Museum of Natural History, Smithsonian Institu- tion, Washington, D.C.; J. Coddington
ZMK, Zoologisk Museum, Kobenhavn, Denmark; H. Enghoff I would also like to thank J. Martinez for a California record and E. Buckup for determining Neoscona in the MCN and sending me the records. E. Buckup, A. Lise, and L. Leibensperger made me aware of some errors, misprints and omissions in previous mono- graphs. A. Dean, C. Deeleman, H. Hofer, and A. A. Lise con- tributed specimens. E. Buckup, M. Grasshoff, and L. Leibensperger read the paper and made suggestions for improvement.
Neoscona Simon
Neoscona Simon, 1864: 261. Type species designated by F. P.- Cambridge, 1904: Epeira arabesca F. P.-Cambridge. Diagnosis. Male palpal patella with two macrosetae and with oval median apophysis of palpus bearing a proximal "upper" recurved tooth (similar in all species, M in Fig. 6). Neoscona dif- fers from other araneid genera with two palpal macrosetae (Ara- neus, Aculepeira, Larinia, Metepeira) by the structure of the palpus (Figs. 6-8, 16; 1971, figs. 4-6): a large area of the bulb is a folded basal hematodocha (H), and only a small area has sclerites.



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224 Psyche [VOI. 99
Unlike in other genera, the sclerites are partly hidden by a wide cymbium (Y, Fig. 6) which touches a small median apophysis (M); the radix is fused to the stipes or has a joint; the stipes is distally three-partite with a conical embolus (E), an embolus lamella (L), and a terminal apophysis (A), the last two only lightly sclerotized. There is almost no distal hematodocha between embolus and termi- nal apophysis.
Female Neoscona differ from other genera by a curved, spatula- shaped epigynum whose sclerites appear fused (Figs. 1-3). The epigynum of N. nautica (Fig. 10) can be confused with that of Ocrepeira species.
Unrecognizable. Neoscona nigrovariata Mello-Leitiio, 1941:258.
The species north of the Isthmus of Tehuantepec are: Neoscona arabesca, N. domiciliorum, N. crucifers (=N. hentzii), N. nautica, N. oaxacensis, N. orizabensis, N. pratensis, N. utahana. Misplaced by Berman and Levi (1971) was N. redempta (Gertsch and Mulaik) which is Ocrepeira redempta (Levi, in press).
The Central American species are N. arabesca, N. nautica, and N. oaxacensis.
The West Indian species are N. moreli, N. nautica, and N. mar- canoi n. sp., with only N. marcanoi native, the other two introduced.
The South American species are N. oaxacensis and the intro- duced N. moreli and N. nautica. The distinguishing characters of these species are the dorsal pattern of the abdomen, the shape of the epigynum and the conductor (C in Figs. 6-8, 16), the terminal apophysis (A), and the embolus lamella (L) of the palpus. Key to the species of South American Neoscona 1 . Females ................................................................................. .2 - Males ...................................................................................... 4 2(l)Abdomen rounded, Araneus-like; only slightly longer than wide (Fig. 11). Epigynum about as wide as long (Fig. 10) .nautica
..................................................................................... -
Abdomen oval and elongate (Figs. 15, 19, 23); epigynum dis- .............................
tinctly longer than wide (Figs. 14, 18, 22). .3




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Levi




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226 Psyche [vo~. 99
3(2)Abdomen with dorsal median band having wavy border, wider anteriorly than posteriorly (Fig. 23); epigynum in posterior view 2.5 times as long as wide (Fig. 22; 1971, figs. 78-80) oaxacensis
................................................................................ -
Abdomen with dorsal median band having a straight border (Fig. 19); epigynum length in posterior view less than two times its width (Fig. 18; 1971, figs. 91-95) ..................... moreli ........................ 4(l)Fourth coxa with a soft tubercle (1971, fig. 9) 5 -
Fourth coxa without tubercle ......................................... nautica 5(4)Abdomen with median light band with straight border (Fig. 19, 1971, fig. 100); palpus in lateral view with conductor (C) almost straight, embolus lamella (L) pressed flat against tegu- lum (T) and only side of terminal apophysis (A) visible (Fig. 20) ................................................................................ ..moreli -
Abdomen with median light band with wavy border (Fig. 23, 1971, figs. 88-90); palpus in lateral view with conductor S- shaped and having the proximal end with a curved shelf; only side of lamella (L) and top of terminal apophysis (A) visible in lateral view of palpus (Fig. 24) ................................ oaxacensis Neoscona rnarcanoi new species
Figures 1-9; Map 1
Holotype. Female holotype and eight female and one male paratypes from Terreno Salado between Neiba and Duverge, Dominican Republic, Hispaniola, 25 Oct. 1980 (E. Marcano F.), in MNSD, 29 paratypes in MCZ. The species is named after the collector.
Description. Female holotype: Carapace light yellowish, darker in median band and on sides. Sternum orange, underlain by median white pigment band. Coxae orange; legs yellow. Dorsum of abdomen with median band of large white patches, smaller one on the sides (Fig. 4); venter black with four white patches and a pair of white spots on each side of spinnerets (Fig. 5). Posterior median eyes 0.9 diameter of anterior medians, laterals 0.8 diameter. Ante- rior median eyes slightly less than their diameter apart, two diame- ters from laterals. Posterior median eyes 0.6 diameter apart, 3 diameters from laterals. Height of clypeus 0.6 diameters of anterior median eye. Abdomen an elongate oval (Fig. 4). Total length 10.3



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Levi 227
Map 1, Distribution of Neotropical Neoscona species. mm. Carapace 5.1 mm long, 4.0 wide, behind lateral eyes 1.8 wide. First femur 5.7 mm, patella and tibia 6.9, metatarsus 5.7, tarsus 1.4. Second patella and tibia 5.9 mm, third 3.5, fourth 5.8. Male paratype: Color as in female, but venter of abdomen gray with white spots touching. Posterior median eyes 0.7 diameter of anterior medians, laterals 0.7 diameter. Anterior median eyes their diameter apart, 1.3 diameters from laterals. Posterior median eyes their diameter apart, 2.3 diameters from laterals. Height of clypeus 0.6 diameter of anterior median eye. Endite with tooth, palpal femur with tooth. First coxa with hook, fourth with a tubercle on posterior face. Second tibia thicker than first, swollen, with prolat- era1 macrosetae (Fig. 9). Abdomen oval. Total length 7.0 mm. Carapace 3.6 mm long, 2.7 wide, behind lateral eyes 1.1 wide. First femur 4.8 mm, patella and tibia 6.3, metatarsus 5.7, tarsus 1.3. Second patella and tibia 4.3 mm, third 2.6, fourth 4.6.



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228 Psyche [Vol. 99
Note: This is probably a native species, not introduced. Males and females were collected together. Illustrations were made from holotype and paratype.
Diagnosis. The female differs from that of N. moreli by a shorter abdomen, marked dorsally with light spots (Fig. 4), and by a shorter, wider epigynum (Fig. 1) which is wider at the lateral lobes than at its base (Fig. 2). The male differs by having the con- ductor with a dark, sclerotized lobe (Fig. 7) and by the shape of the terminal apophysis (A) and embolus lamella (L) (Figs. 7, 8). The second tibia has a dense field of setae (Fig. 9). Paratype. DOMINICAN REPUBLIC: 1 km S Las Matas de Far- tan, 25 Aug. 1970, Q (B. Patterson, MCZ). Neoscona nautica (L. Koch)
Figures. 10-13; Map 1
Epeira tristis Taczanowski, 1873: 13 1. Female holotype from Iles de Salut, French Guiana in PAN, examined. Name preoccupied by Epeira tristis Blackwall, 1862. Synonymized by Berman and Levi, 1971.
Epeira nautica L. Koch, 1875: 17. pi. 2, fig. 2., Q. Female holotype from Suakin [Sudan], in BMNH, examined.
Epeira volucripes Keyserling, 1885: 528, pi. 13, fig. 27, Q. Female syntypes from Panama and Haiti in MCZ, examined. Neoscona volucripes: - F. P.-Cambridge, 1904: 473, pi. 44, fig. 18, 0.
Aranea tristimonae Petrunkevitch, 19 1 1: 320. New name for Epeira tristis Taczanowski, preoccupied. Synonymized by Berman and Levi, 197 1.
Araneus marcuzzii Caporiacco, 1955: 355, fig. 35, 9. Female holo- type from Cartipano, Sucre, Venezuela, in MUCC, examined. Synonymized by Berman and Levi, 197 1.
Neoscona nautica: - Berman and Levi, 1971: 498, figs 11 1-120, Q, 0. Grasshoff, 1986: 46, figs. 60-63, Q, 0, map 6. Diagnosis. The female abdomen is almost as wide as long (Fig. 11) and can be confused with that of Araneus species; unlike in other American Neoscona the terminal apophysis of the palpus is very small (Fig. 12); the second tibia (Fig. 13) lacks the dense field of short setae present in N. moreli.



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Figures 10-13. Neoscona nautica. 10, epigynum, posterior. I I, abdomen of female, 12, male left palpus, lateral. 13, male left second patella and tibia, anterior. Figures 14-17, N. arabesca. 14, epigynum, posterior. 15, abdomen of female. 16, male palpus, lateral 17, male second patella and tibia, anterior. Figures 18-21, N. nwreii. 18, epigynum, posterior. 19, abdomen of female. 20, male palpus, lateral. 21, male second patella and tibia, anterior. Figures 22-25, N, oaxmensis. 22, epigy- num, posterior. 23, abdomen of female. 24, male palpus, lateral. 25. male second patella and tibia, anterior.
Scale lines 1 mm; genitalia 0.1.
Abbreviations. A, terminal apophysis. C, median apophysis. L, embolus lamella. M, median apophysis. T, tegulum.




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230 Psyche [vo~. 99
Note. The illustrations were made from a female from Depto. Antioquia, the male from Depto. Valle, Colombia, the male tibia and patella from Puerto Rico.
Distribution. Cosmotropical: widespread in tropical areas of Africa; in America from Virginia, Texas, and the West Indies to Argentina. It is probably introduced to America. New records. CUBA Holquin: Farallones de Moa (MNHNC). Habana: Ojo de Agua de Nico, S. A. Bafios (MNHNC). HAITI Port au Prince (MCZ). MONA ISLAND (MCZ). GUATEMALA Suchitepequez: San Julian (AMNH). Peten: Tikal ruins (MCZ). Chimaltenango: Yepokapa (AMNH, USNM). NICARAGUA Bonanza (AMNH); El Castillo (JMM, MCZ). COSTA RICA Here- dia: La Selva (AMNH). San Jose: San Jose (CAS). PANAMA Boca del Toro: Coroente Grande, Changuinola (MCZ); Rio Chan- quinola nr. Quebrada El Guabo (AMNH). Chiriqui: David (AMNH), Balboa (AMNH). VENEZUELA Carabobo: San Esteban (AMNH). SURINAM Morowijne: Langaman (AMNH). COLOM- BIA Cksar: Valledupar (JAK). Cundinamarca: Tibirita, in house (MCZ). Guajira: Maicao (AMNH). Antioquia: Mutata Cancheras (MCZ); Medellin (MCZ). Chocd; Mutis, Bahia Solano (CAS); between Cucurrupi and Noanama, Rio San Juan (AMNH); Cafio nr. Cucurrupi, Rio San Juan (AMNH); 20 km N Palestina, Rio San Juan (AMNH). Valle: Centr. Hid. Anchicayii (MCZ). Amazonas: Araracuara, 270 m (CV). Nariiio: Reserva Nat. La Planada, Ricau- rte, 1350 m (CV). ECUADOR Guayas: Guayaquil (CAS, MACN); Colonche (CAS). Manabi: Crucita, 0å¡52'S 80å¡32' (MCZ); El Carmen (MECN). BRAZIL Amazonas: Mauks (INPA). Bahia: Ita- maraju (MNRJ). Rio de Janeiro: Rio de Janeiro (ZMK); Petr6polis (AMNH). Mato Grosso: Chapada dos Guimariies (AMNH). Santa Catarina: Corupa (AMNH). SLio Paulo: Juquia (MZSP); Piquete (MZSP); Santos (MZSP); Raiz de Serra (MZSP). Rio Grande do Sul: Santa Rosa (MCN). PARAGUAY Central: Asuncion (MNRJ). Alto Paranh: km 12 de Stroessner, Centre Forestal de Alto Parang (IBNP) . ARGENTINA Chdo ba: General Paz (MLP) . Neoscona arabesca (Walckenaer)
Figures 14-17; Map 1
Epeira arabesca Walckenaer, 1841: 74. The name was given to the specimen illustrated by L. A. G. Bosc, unpublished. Araignkes



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de Caroline, p. 13, pl. 5, fig. 2, in the library of The Natural History Museum, London.
Neoscona arabesca: - Berman and Levi, 1971: 474, Figs. 14 to 35, Q, 0.
Diagnosis. This is the smallest and commonest North American species. The female has an egg-shaped abdomen, with characteris- tic pairs of diagonal marks (Fig. 15 and 1971, figs. 34, 35). The male has the edge of the embolus lamella (L) and terminal apoph- ysis (A) almost parallel and the conductor (C) relatively straight (Fig. 16). The male left second tibia (Fig. 17) lacks the field of short setae present in some other Neoscona species. Note. The illustrations of the female were made from an indi- vidual from Costa Rica, that of the male palpus from a specimen from Talapa, Mexico, the patella and tibia from a male from Cuba. Natural History. Specimens came from a sorghum field in Hon- duras; others were found in meadows and open areas. Distribution. Common from southern Canada and the United States south to Costa Rica, Bahamas, Cuba, Hispaniola, and Jamaica.
New records. GUATEMALA Antigua: Finca Salinas, 29 June 1980, 3~ (J. Schuster, FSCA). HONDURAS 37 km E Tegucigalpa, sorghum fields, 9, 2 Dec. 1982 (R. W. Jones, MCZ). COSTA RICA San Jos6: San Jos6, Q, o (E. Schmidt, AMNH). Neoscona crucifera (Lucas)
Epeira crucifera Lucas, 1839: 42, pl. 6, fig. 3. Specimens from the Canary Islands.
Epeira hentzii Keyserling, 1864: 97, pl. 5, fig. 10, 1 1, Q. Female lectotype designated by Berman and Levi, 197 1 in BMNH. Syn- onymized by Grasshoff, 1986.
Neoscona hentzii: - Berman and Levi, 1971 : 478, figs. 5 1-58. Neoscona cruci$era: - Grasshoff, 1986: 62, figs. 79-84, Q, 0, map 9; Martinez (in press).
Diagnosis. The abdomen is almost as wide as long. Natural history. Martinez reports (personal communication Feb. 1992) that the species is getting more abundant in southern Cali- fornia and builds webs higher up than the webs of N. oaxacensis. Distribution. New England to California and central Mexico; Canary Islands Madeira and Porto Santo, Africa. Presumably this



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232 Psyche [vo~. 99
species has been carried from America to Africa. Heretofore unknown from California.
New record. CALIFORNIA Long Beach, Rancho Los Cerritos, 1 Nov. 1991, 5~ (M. J. Martinez? MCZ).
Neoscona rnoreli (Vinson)
Figures 18-21 ; Map 1
Epeira rnorelii Vinson, 1863: 166, 309, pl. 4, fig. 4. Specimens from Reunion.
Araneus neotheis Petrunkevitch, 19 1 1 : 305. New name for Epeira theisi: - Keyserling, which was thought misidentified. First syn- onymized by Grasshoff, 1986.
Neoscona neotheis: - Berman and Levi? 1971: 490, figs. 91-100, 970.
Neoscona rnoreli: - Grasshoff, 1986: 561, figs. 71-78, Q? 0. Diagnosis. The abdomen is slender and usually has a median light band with almost straight margins (Fig. 19). The conductor of the male palpus is relatively straight and the terminal apophysis has a concave edge (Fig. 20). The second tibia of the male has a dense field of short setae (Fig. 21).
Note. Figure 18 was made from a female from Depto. Valle, Colombia, Fig. 19 from Puerto Rico and Fig. 20 from a male from Depto. Meta, Colombia, Fig. 21 from a male from Cuba. Natural History. Specimens were collected in Puerto Rican mountains, in an open abandoned parking lot, in high grasses and herbacious vegetation, and in a soybean field in Palmira, Valle, Colombia.
Distribution. Widespread in Subsaharan Africa? Madagascar, Reunion, Mauritius and Seychelles, West Indies, and with scat- tered records in South America (Map). I assume the species is an introduction to the Americas, judging by the limited American, and widespread African distribution.
New records. CUBA Santiago de Cuba: La Gran Piedra (MCZ). DOMINICAN REPUBLIC Monte Cristi: ca. desemboca Jura del Rio Yaque de1 Norte (F. Alay6n7 MNHNC). BAHAMA ISLANDS South Caicos (AMNH). VIRGIN ISLANDS St. Croix (AMNH, ZMK), St. Johns (AMNH), Tortola (AMNH), Little Camonoe (AMNH). NEVIS Charlestown (AMNH). MARTINIQUE Fort de



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19921 Levi 233
France (MCZ). GRENADA (AMNH). VENEZUELA Bolivar: Maripa, Rio Cauca (MCZ). GUYANA Rupununi: Mountain Points, 30 km from Dadanawa (G. F. Mees, CD). COLOMBIA Meta: Carimagua, (Eberhard 607> MCZ); 15 km SW Puerto Lopez (Eber- hard 1573, MCZ); Hacienda Mozambique, 15 km SW Puerto Lopez (W. Eberhard, MCZ); 20 km N Rio Muco, 20 km S El Por- venir (W. Eberhard, MCZ). Valle: Lago Calima (W. Eberhard, MCZ); Jamundi, Finca Ceibalito (H. Bastidas, MCZ); Palmira (MCZ). PERU Loreto: Barranca (CAS). Tacna: S Camiara (AMNH). BRAZIL Terr. Amapa: Macapii (J. Becker, MEG) in nest of Eumenes canaliculatus. Bahia: Itamaraju (MNRJ). Minas Gerais: Vi~osa (MNRJ). Mato Grosso: Chapada dos Guimarges (M. Alvarenga, AMNH). Rio de Janeiro: Cidade (AMNH, MCZ). SZo Paulo: Guarujii (AMNH). Rio Grande do Sul: Cap30 da Canoa (MCN); 1tapuii7 Porto Alegre (MCN); Pelotas (AMNH); Santa Maria (MCN); Torres (MCN); Viamiio, (MCN); Xangrilii (MCN). PARAGUAY Concepcih: Fonciere, imm. (E. Reimoser, MCZ). ARGENTINA Chaco: Basail (M. Biraben, MACN). Corrientes: Solari (M. Biraben, MLP).
Neoscona oaxacensis (Keyserling)
Figures. 22-25; Map 1
Epeira oaxacensis Keyserling, 1863: 121, pl. 5, figs. 15, 16, Q. Female holotype from Oaxaca, Mexico in BMNH, examined. Epeira cooksonii Butler, 1877: 76, pl. 13, fig. 2, Q. Six female syn- types from Albemarle Island, Galapagos, in BMNH, examined. Synonymized by Berman and Levi, 197 1.
Epeira adiantoides Taczanowski, 1878: 148, pl. 1, fig. 4, 9. Female syntypes from Chorillos [now suburb of Lima], Peru in PAN, examined. Synonymized by Berman and Levi 197 1. Neoscona oaxacensis: - Berman and Levi, 1971: 486, figs. 78-90, Q, 0.
Diagnosis. the median light band of the abdomen has lobed margins (Fig. 231, and the palpus an S-shaped conductor (Fig. 24). The second tibia of the male has a field of short setae (Fig. 25). Variation. Specimens from the Galapagos Islands are among the largest ones in collections and females have a slightly shorter epigynum. Males have the edge of the terminal apophysis concave,