Herbert W. Levi.
The Ravilla Group of the Orbweaver Genus Eriophora in North America (Araneae: Araneidae).
Psyche 77:280-302, 1970.

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THE RAVILLA GROUP OF THE ORBWEAVER GENUS ERIOPHORA IN NORTH AMERICA
( ARANEAE : ARANEIDAE) *
BY HERBERT W. LEVI
Aluseu~n of Comparative Zoology, Cambridge, Mass. 02138 The contradicting synonymies presented in the two recent catalogs of spiders (Roewer, 1942; Bonnet, 1955) reflect the confusion in names and species of Eriophora in the literature. Co~nstock (1940) indicates that specimens of Eriophora have been found as far north as Oregon. Presumably this error can be traced to the use of the George Marx collection by McCook and Keyserling. The Marx collection, housed in the U.S. National Museum, has unreliable locality labels. No recent collections of Eriophora from central and northern California have been seen.
The object of this revision was to find the differences anlong species of these very large, common orbweavers of the southern United States. The types of the old names were examined in hope of pro- viding the stability and unversality required of animal names. Only adult specimens that I have myself examined were used for mapping ranges. Unfortunately, South American collections of Araneidae in North American museums are unsorted and only a few specimens provided records for the southern limits of the distributions. Dr. M. H. Robinson of the Smithsonian Tropical Research Institute, Panama Canal Zone, called my attention to his immediate need of an Eriophora revision to make it possible for him to publish his behavior observations. I take this opportunity to thank Dr. J. G. Sheals and Mr. D. Clark for the hospitality extended when I worked at the British Museum (Natural History), and to thank Mr. Clark for numerous favors. Collections were made available by Dr. W. J. Gertsch and Dr. J. A. L. Cooke of the American Museum of Natural History, Dr. M. Grasshoff of the Sen~kenber~ischen Natur- forschenden Gesellschaft, Frankfurt, Dr. J. Pr6szyn'ski, Polish Academy of Sciences, Warsaw, Dr. D. C. Robinson and Mr. C. E. Valerio of the biology department of the University of Costa Rica, San Jose, Dr. H. V. Weems, Florida State Museum, Gainesville, Prof. H. I<. Wallace, University of Florida. Additional specimens were made available by Dr. L. van der Hammen, Natural History *Manuscript received by the editor December 1, 1970.



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19701 Levi - Genus Eriophora 28 I
Fig. la.
Eriophora fuliginea feeding on a bat. On the left are two web parasites belonging to the genus Argyrodes (Theridiidae) . Photograph taken in the Panama Canal Zone by Dr. R. Akre. Museum, Leiden, Dr. M. Moritz, Zoologisches Museum der Humboldt Universitiit, Berlin, Prof. I<. D. Schiapelli and Mrs. B. G, dePikelin, of the Museo Argentine de 'Ciencias Naturales, Buenos Aires, Mr. D. Bixler and Dr. R. E. Leech. This investiga- tion was supported in part by Public Health Service Research Grant AI-01944 from the National Institute of Allergy and Infectious Diseases, and published with a grant from the Museum of Com- parative Zoology.
At present I consider Parawixia a synoym of Eriophora. How- ever, the species customarily included in Parawixia are not included in this paper. They are much less common and most are South American, the most northern ones coming from Mexico. Collections from these areas have not been sufficiently sorted and few specimens are available. The publication of this paper is necessitated by the immediate need for names and means of identification of these common species.




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282 Psyche [September
Eriophora Simon
Er'iophora Simon, 1863, Histoire Naturelle des Araignies, ed. 1, p. 261. New subgenus of Epeira (=Araneus). The type of Eriophora is Epeira ravilla C. L. Koch as designated by F. P.-Cambridge, 1903, Biologia Centrali-Americana, Araneidea, vol. 2, p. 461. The gender is feminine. All Eriophora species differ from Araneus by having a paramedian apophysis in the palpus (PM in Figs. 2, 4). The female has the second leg almost the same length as the first (Fig. I b) ; the third is always shortest. The median eyes are separated by about their diameter or less. The clypeus width is less than the diameter of the anterior median eyes. The abdomen shape varies from bluntly pointed anteriorly, acutely pointed posteriorly, and widest in the anterior third, to almost spherical (Fig. 32). The posterior of the abdomen is generally high above the spinnerets (Figs. 22-24). There are often posterior or dorsal tubercles on the abdomen: In E. ravilla there is one posterior to another; in E. edax (Figs. 42-44) there is always one distinct tubercle and often more. The venter has a median black patch (Figs. I 6, 3 I ) . The lateral eyes of males are on a short stalk, the medians on a projection. The small chelicerae are long, attenuated and concave anteriorly, providing space for the large palps. The first coxae of males have a hook on the distal margin (absent in E. nephiloides). The second tibiae are modified, most so in E. ravilla (Fig. 8), only swollen in E. nephiloides. The fourth coxae have a row of transverse macrosetae in E. edax
(Fig. 48) and thin setae in E.
nephiloides (Fig. 58). An unusual character is the presence of two strong macrosetae on the fourth trochanter of males (Figs. 7, 48) (except in E. nephiloides, which has only some weak setae). Juvenile specimens of Eustala and Eriophora are often confused. Genitalia. The epigynum is very small in E. nephiloides; in most other species it has a relatively long scape. The scape is attached to the anterior of the base and bends backward; in Verrucosa and in species previously considered to belong to Parawixia, it is attached ventrally. In E. raviZla and E. fuliginea the scape breaks off, prob- ably in mating, and about half the females of these two species in collections lack the scape.
It does not break off in the other two
species, though E. edax may very rarely have the tip of the scape broken off. The base of the epigynum contains the seminal receptacles (SR in Figs. 5, 6). The shape of the seminal receptacles probably differs in different species, as reflected by the proportions of the base; this possibility was not pursued because the base is so



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19701 Levi - Genus Eriophora
283
Fig. lb. Eriophora nephiloides. Photograph taken at Barro Colorado Island, Panama Canal Zone by Dr. M. H. Robinson.



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284 Psyche [September
heavily sclerotized that the character would be almost useless for separating species. The openings are ventral, under the scape (Fig. 5). The openings are bordered posteriorly by a triangular piece (Figs. 5, 14). There are no sclerotized basal lamellae as in some species of Araneus and Verrucosa.
The patella of the palpus has two macrosetae of which one may be much longer than the other.
The palpus of Eriophora was first described by Comstock ( 1910, 1912, 1940). In all species of Eriophora the palpus is similar in structure. The radix (R of Figs. 2-4), stipes (I) and median apophysis (M) are elongated compared to Araneus, but the tegulum (T) is compact (Figs. 2-4). As a result, the median apophysis has shifted laterally from the ventral position it occupies in Araneus, and if viewed from the mesal side seems to be behind the embolus (E in Fig. 4). A separate piece, called the paramedian apophysis (PM) by Comstock, is present. In Verrucosa, which has a similar palpus, the piece is not separate, but is an extension of the proximal end of the conductor. Perhaps because parts of the bulb are elongated, the cymbium (Y) is very narrow and canoe-shaped. Unlike Araneus, Eriophora has little distal hematodocha (DH) and the terminal apophysis (A) and embolus (E) are on a complex V-shaped structure that has only limited articulation with the stipes (I). A spur of the terminal apophysis is probably homologous to the sub- terminal apophysis of Araneus. If there is a tip that breaks off the embolus in mating, it must be small. None was seen, but emboli were not examined under a compound microscope. Data from the collections indicate that throughout the range of the genus (except possibly the most northern parts), adult males can be found throughout the year and there is no definite season of maturity as in the species related to Araneus diadematus. Web. The web of E. fuliginea (according to M. H. Robinson, personal communication) has an open hub and usually is very asym- Figs. 2-6. Structure of Eriophora genitalia, diagrammatic. 2-4. Left palpus. 2. Expanded, mesal. 3. Expanded, lateral. 4. Contracted, mesal. 5-6. Base of epigynum.
Figs. 7-9. Eriophora milla (C. L. Koch). 7. Male, fourth coxae and trochanters, ventral view. 8. Male, left second tibia and tarsus, ventral view. 9. Female, abdomen with epigynum, ventral view. Abbreviations: A, terminal apophysis; C, conductor; D, sperm duct; DH, distal hematodocha; E, embolus; H, basal hematodocha; I, stipes; M, median apophysis ; PM, paramedian apophysis ; P, paracymbium ; R, radix ; S, subtegulum ; SR, seminal receptacle ; T, tegulum ; Y, cymbium. Size Indicators: 1 mm.




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19701 Lcvi - Genus Eriophora 285




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286 Psyche [September
metrical.
The hub is in the upper third of the web. The web may be as large as 1.6 to 2 m in diameter. The spider occupies a retreat to the side.
Distribution. There are specimens of Eriophora species in col- lections from New Guinea, New Caledonia, New Zealand and Australia. One species belonging to the genus has recently been described from South Africa.
Key to North American species of the ravilla group of Eriophora Males .......................................................................................... 2 ........................................................................................ Females 5 Fourth coxa with a transverse row of 3 macrosetae (rarely, only 2 in northern specimens) (Fig. 48) ....................... edax Fourth coxa without a transverse row of macro,setae (Figs. 7, 34) .............................................................................................. 3 Median apophysis of palpus short, framed by other structures (Fig. 49) .................................................................... nephiloides Median apophysis of palpus long extending beyond other struc- tures (Figs. 10, 25, 35) ......................................................... 4 Median apophysis with a spur (Figs. 2, 10) ; tip of embolus drawn out, recurved (Fig. 11) ...................................... ravilla Median apoph~sis without a spur (Fig. 25) ; tip of embolus with a flap on its side (Fig. 26) ................................ fuliginea Carapace, coxae yellow, sternum black; epigynum of minute size (Figs. 51, 54) .......................................................... nephiloides Coloration otherwise; epigynum of normal proportions . . . . 6 Posterior of dorsum of a,bdomen with a distinct hump (Fig. 42) ; scape of epigynum usually with a keel, sometimes circular in cross section, rarely broken off (Fig. 37) .................... edax Posterior of dorsum with only a slight hump or none at all (Figs. 22-24) ; epigynal scope without a keel or often broken off (Figs. 12, 27) ........................................................ 7 "Bse of epigynum wrinkled (Fig. I 3) ......................... ravilla Base of epigynum not wrinkled, but flat shield-shaped (Fig. 28) f z~liginea
............................................................................... Eriofihora ravilla ('C. L. Koch)
Map I ; Figures 7-24
? Epeira Circulata Wakkenaer, 1841, Histoire Naturelle des Insectes Apteres, vol. 3, p. 79. Female syntypes from America lost and also Abbot illus- trations no. 170, 363 from Georgia. The Abbot manuscript is at the British Museum, Natural History. A copy of it in the Museum of Comparative Zoology was examined. Nomen dubium.



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19701 Levi - Genus Eriophora 287
Epeira rawtila C. L. Koch, 1845, Die Arachniden, vol. 11, p. 73, fig. 890, 2. Female holotype from Mexico in Royal Collection in Berlin, lost, likely destroyed. However, a specimen from Veracruz, Mexico, in the L. Knch collection in the British Museum, Natural History, with the label probably by C. L. Koch saying E. rawifla, examined. McCook, 1M3, American Spiders, vol. 3, p. 161, pi. 5, figs. 7, 8, 9, 8. ? Epeira septha Hentz, 1847, J. Boaton Soc. Natur. Hist., p. 470, pi. 30, fig. 9, 9. The types from Alabama and North Carolina have been tientroyed, Nowen dv-b'ium.
Eriophwa rav'ilh,-Simon, 1864, Histoire Natureile des Araign&es, 1st ed., p. 261. R Pi-Cambridge, 1904, Biologia Centrali-Americana, Araneidea, vol. 2, p. 465. Roewer, 1942, Katalog der Arancae, vol. 1, p. 866. Bonnet, 1955, Bibliographia Araneorum, vol. 2, p. 1785. Efeira nicaraguensis Keyserling, 1885, Verhandl. Zoo}. BOL Gea. Wien, vol. 34, p. 523, pl. 13, fig. 31, Q . Female holotype from Nicaragua in the Museum of Comparative Zoology, examined. Keyserling, 1B93, Spinnen Arnerikas, vol. 4, p. 214, pi. 10, fig. 159, 9. NEW SYNEWYMY. Epeira balausfina McCook, 188& Proc. Acad. Natur. Sci. Philadelphia, p. 198. Female syntypes from Florida, Swan Island, Caribbean, and San Dottiingn, lost; the type locality is here restricted to [tropical] Florida. McCook, 1893, American Spiders, vol. 3, p. 155, pi. 4, fig. 2, Q. NEW
SYNONYMY.
tipeira I&nrioluta 0. Pi-Cambridge, 1889, Biologia Central]-Americana, Araneidea, vol. 1, p, 27, pi. 6, fig. IS, juv. Types from Guatemala in the British Museum, Natural History, lost. Nomen dubiu-ia. Keyserling, 1893, Spinnen Amerikas, vol. 4, p. 100, pi. 5, fig. 74, 8. McCook, 1893, American Spiders, vol. 3, p. '160 (in part), pi. 5, fig. 5, 9, 8. Epr'iru wddtita 0. PXarnbridge, 1889, Biologia Centrali-Americana, Ara- ~~eidra, vol. !, p. 46, pi. 6, fig. 14, juv. Juvenile syntypes from Guate- mala in the British Museum, Natural History, lost. Eriofhorti baIaustina,- Banks,
1909, Rep. Centr. Exp. Sta. Cuba, p. 161, Rpeirn winax 0. Pi-Cambridge, 1893, Biologia Centrali-Americana, Aranei- den, vol. 1, p. 112, pi, IS, fig. 1, 9. Three female syntypes from Acagiiizotla, Guerrero, Mexico, in the British Museum, Nautrat History, uxnmined. HEW SYNONYMY.
l?rinfo/iiira mhax, - F. Pi-Cambridge, 1903, Biologia Centrali-Americana, Araneidea, vof. 2, p. 464, pi. 44, fig. 3, $. Roewer, 1942, Katalog der Araneac, vol. 1, p. 867. Bonnet, 1955, Bibliographia Araneorum, vol. 2, p. $784,
/transit nicarag~usis, - F. P.-Cambridge, 1904, Biologia Centrali-Ameri- cam, Araneidea, vol. 2, p. 519. Roewer, 1942, Katalog der Araneae, vn!. 2, p. 848.
KĖ io$/torft drcufata,- Comstocfc, 1912, Spider Book, figs. 108-111, 538 ; 1940, Spider Book, rev. ed., figs.
108-111, 538. Probably not Ep~ira
cireiilata Waickenaer.
? Aruncus perfoliatw Franganillo, 1936, Los Aracnidps de Cuba, p. 70. Specimen in Cuban Academy of Sciences, Havana, with numbered code lacking key. Nomen dabium.
? Aruneus aftunciuaius Franganillo. 1963, Los Aracnidos de Cuba, p. 71. Specimen disposition, see above. Nomen dubh. Araiif~i nicaragu ensis, - Bonnet, 195 5, Bibliographia Araneorum, vol. 2, p. 549.




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288 Psyche
[September
Map 1.
Distribution of Eriophora ravilla (C. L. Koch) Note. Walckenaer's ( I 841 )
description of Epeira circzilata sug-
gests the pattern illustrated by Fig. 20, but the specimen is lost and Abbot's illustration, fig. 170 made from a Georgia spider, also re- seinbles this pattern. However, this species is not usually found as far north as Georgia. 'Chamberlin and Ivie (1944, Bull. Univ. Utah, l3iol. Ser., vol. 7, no. 5, p. 105) interpreted the description to be iw Emidla. Also I have not seen any other specimens with this pattern from the southeastern United States. It seems best not to use this doubtful name. The illustration of Epeira seflima Hentz fits tin's species, the description may be a composite. Hentz reported the species from Alabama, where it occurs, and North Carolina, where it does not occur. Archer ( 1940) considered E. septima a synonym of Araneus trif olium.
In
the L. Koch collection in London there are two specimens labeled E. ravilla, one from Cuba, and one from Veracruz, Mexico. Since the type locality is Mexico, I assume the Veracruz specimen



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19701 Levi- Genus Eriophora 289
might be the lost type, and that L. Koch received the specimens from the elder Koch. The one from Cuba is Eriophora edax. McCook
(1893) first suggested that balaustina might be milla of C. L. Koch.
Description. Female from Florida. Carapace red-brown with white hairs. Chelicerae dark brown. Sternum and coxae red-brown. Legs red-brown with femora darker and with white hairs and white setae.
Dorsum of abdomen dark gray to black. Venter with a tri- angular black mark on gray with some white pigment surrounding it between epigynum and spinnerets. The triangular mark is pointed posteriorly. Anterior median eyes largest, posterior medians 0.6 diameters of anterior medians, laterals radius of anterior medians. Anterior median eyes one diameter apart, posterior medians two- thirds diameters apart. Abdomen is suboval, slightly pointed an- teriorly and on sides.
Total length 17 mm. Carapace 6.8 mm long, 5.8 mm wide. First femur, 7.6 mm; patella and tibia, 9.6 mm; metatarsus, 6.0 mm; tarsus, 1.7 mm. Second patella and tibia, 8.6 mm; third, 4.7 mm; fourth, 7.8 mm.
Male from Cuba. Coloration almost like that of female. Legs banded. Abdomen dark gray. Carapace with a deep longitudinal thoracic groove. Eye proportions like those of female. The second tibia is illustrated in Fig. 8. Abdomen oval, widest anteriorly. Total length 10 mm. Carapace 5.3 mm long, 3.7 mm wide. First femur, 6.8 mm; patella and tibia, 9.0 mm; metatarsus, 6.3 mm; tarsus, 1.4 mm. Second patella and tibia, 6.8 mm; third, 4.0 mm ; fourth, 5.8 mm.
Variation. Females measure total length, 12-24 mm. Some females are almost white, others nearly black. Some have an irregular asymmetrical longitudinal white pigment patch which may be a narrow line or widest anteriorly just over the heart area. Males are 9-13 mm in total length. In some the legs lack banding. The abdomen is white to dark gray. Puzzling is the coloration (Fig. 20) having two dark-bordered, slightly sclerotized humps, one after another. All but one with this coloration were juveniles, the only adult being from Veracruz, Mexico.
Diagnosis. Females are readily told from Eriophora f uliginea by the wrinkled base of the epigynum (Figs. 13-15), males by the spur on the median apophysis (Fig. 10).
Natural History. The species has been collected in live oak trees of scrub at a salt marsh in Levy 'County, Florida and on a hammock above dense undergrowth in Dade County, Florida. In Alachua



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290 Psyche [September
County, Florida, H. I<. Wallace (personal communication) observed a pompilid wasp [Poecilopompilus interruptus (Say) ] attacking a spider while in the web and on the ground. Distribution.
Gulf coast, Florida, Central Mexico, and West Indies to South America.
Border records. Maryland. Bethesda, 6 August 1943, $ (J. M. Davis. Grenada: near St. George's. Brazil. Rio de Janeiro, [ I 865-1 8661, $ (Thayer Exped.) .
Eriophora fuliginea (C. L. Koch), new combination Map 2; Figures la, 25-34
Epeira fiiliginea C. L. Koch, 1893, Die Arachniden, vol. 5, pi. 58, fig. 375, Q. Three female syntype specimens from Brazil were in the Bavarian collection in Munich, presumably destroyed. - Taczanowski, 1873, Horae SOC. Entomol. Ross., vol. 9, p. 129.
Epeira hispida C. L. Koch, 1845, Die Arachniden, vol. 11, p. 71, fig. 889, 8. Male type from Brazil in the Zoologisches Museum, Humboldt Uni- versitat, Berlin ; examined. NEW SYNONMY. Epeira ursina Keyserling, 1865, Verhandl. 2001. Bot. Ges. Wien, vol. 15, p. 822, pi. 19, figs. 3-5,
$. Female holotype from [the Spanish Colony] New Granada, in the British Museum, Natural History, examined. Keyserling, 1893, Spinnen Amerikas, vol. 4, p. 229, pi. 11, fig. 170, 9.
NEW SYNONYMY.
Epeira trapezoidrs Karsch, 1879, Stettiner Entomol. 2. vol. 40, p. 107. Female holotype from Santa Marta, Colombia, in the Zoologisches Museum, Berlin, examined. NEW SYNONYMY
Epeira messallna Hasselt, 1888, Tijdschr. Entomol. 31: 181, pi. 6, figs. 1, 2, Q. Female holotype from Surinam, in the Natural History Museum, Leiden, examined. NEW SYNONYMY.
Epeira purpurascens 0. P.-Cambridge, 1889, Biologia Centrali-Americana, vol. 1, p. 33, pi. 7, figs. 4, 5, 9, 8. F.emaIe syntypes from Bugaba, Panama, in the British Museum (Natural History), examined. Key- serling, 1893, Spinnen Amerikas, vol. 4, p. 226, pi. 11, fig. 168, 9, $ [purpurescens]. NEW SYNONYMY.
Araneus fuligineus,- Simon, 1895, Histoire Naturelle des Araignks, vol. 1, p. 803. Bonnet, 1955, Bibliographia Araneorum, vol. 2, p. 5'05. Araneus purpurascens,- Simon, 1897, Proc. Zool. Soc. London, p. 873. Figs. 10-24. Eriophora ravilla (C. L. Koch). 10. Male, left palpus, mesa1 view. 11. Embolus. 12. Epigynum. 13-15. Base of epigynum. 13. Ventral. 14. Posterior. 15. Lateral. 16. Female, ventral view of abdomen. 17-21. Female, dorsal view of abdomen. 17. (Florida). 18. (Texas). 19. (Nayarit). 20. (Veracruz). 21. (Nicaragua). 22-24. Female, lateral view of abdomen. 22. (Jamaica). 23. (Nayarit). 24. (Veracruz). Size Indicators: Figs. 10-15, 0.5 mm. Figs. 17-24, 1 mm.



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19701 Levi - Genus Eriophora




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292 Psyche [September
Map 2.
Distribution of Eriophora fuliginea (C. L. Koch) Eriophora purpurascens, - F. P.-Cambridge, 1903, Biologia Centrali-Ameri- cana, vol. 2, p. 463, pi. 43, figs. 14, 15, 9, 8. Roewer, 1942, Katalog der Araneae, vol. 2, p. 867. Bonnet, 1955, Bibliogra~hia Araneorum, vol. 2, p. 1784.
? Araneus fuligineus var. rhomboidalis Franganillo, 1936, Los Aracnidos de Cuba, p. 67. The Franganillo collection is in the Cuban Academy of Sciences, Havana, but with a numbered code and without a key to the numbers. Nomen duhium.
? Araneus fuligineus var. sanguineus Franganillo, 1936, Los Aracnidos de Cuba, p. 68. Specimen deposition, see above. Nomen dubium. Aranea fuliginea, - Roewer, 1942, Katalog der Araneae, vol. 2, p. 842. Aranea ursina,-Roewer, 1942, Katalog der Araneae, vol. 2, p. 855. Araneus ursinus,-Bonnet, 1955, Bibliographia Araneorum, vol. 2, p. 626. Note. There are no specimens o'f Epeira fuliginea in the Staats- ammlungen des Bayerischen Staates, Munich, according to Dr. Egon Popp. Presumably these were specimens collected by the Spix and Martius expedition to equatorial Brazil. The oldest specimens



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19701 Levi - Genus Eriophora 293
Figs. 25-34. Eriophora fuliginea (C. L. Koch). 25. Male, left palpus, m e view. 26. Embolus. 27. Epigynum. 28-30. Base of epigynum. 28. Ventral. 29. Posterior. 30. Lateral. 31-33. F,emale, abdomen 31 Ventral. 32. Dorsal. 33. Lateral. 34. Male, fourth coxa and trochanter. Size Indicators: Figs. 25-30, 0.5 mm. Figs. 31-34, 1 mm.



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294 Psyche [September
of this species examined bearing this name are those of Taczanowski in the Polish Academy of Sciences, Warsaw. The Franganillo names may refer to this species or E. raviZZa. Description. Female from Panama Canal Zone. 'Carapace, ster- num, and legs brown. Carapace with some long white hairs from thoracic groove toward sides of head and shorter ones on the anterior margin. Dorsum of abdomen black with a well defined narrow longitudinal white line. Area on venter between epigynum and spinnerets with a triangular black mark pointing posteriorly, and a white pigment spot on each side anterior of the spinnerets (Fig. 3 I ) .