Susan Glueck.
A taxonomic revision of the orb weaver genus Acacesia (Araneae: Araneidae).
Psyche 101:59-84, 1994.

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A TAXONOMIC REVISION OF THE ORB WEAVER GENUS ACACESIA (ARANEAE: ARANEIDAE)
There are five species of Acacesia which range collectively from southern North America to Argentina. Two are previously known members of the genus, A. cornigera Petrunkevitch and A. hamata (Hentz). Three of these are new species: A. villalobosi and A. yacuiensis, from southern Brazil, and A. benigna from Bolivia and Peru.
Acacesia is a genus of orb-weaving spiders common and endemic to the Americas, proposed by Simon in 1892. Hentz named the type species Epeira foliata in 1847. The genus con- tained the single species A. hamata until Petrunkevitch (1925) described A. cornigera. Subsequently published names, illustra- tions, and descriptions of new species of Acacesia have turned out to be synonyms of the two extant species (Chamberlin and Ivie 1936, Badcock 1932). Levi (1976) redescribed A. hamata in a revi- sion of Nearctic orb weaver genera, mentioning the Neotropical A. cornigera in passing. He hypothesized at the time that "there are three or four additional species of Acacesia, all Neotropical and all similar in appearance." The specimens I have examined for the current taxonomic revision corroborate this statement, and I expand the genus to include three new South American species. This study represents an addition to Levi's ongoing project of revising Neotropical orb weavers.
I here report the results of my taxonomic project for which I examined and illustrated over 350 museum specimens. I describe factors I considered in delimiting new species such as their Harvard College, Harvard University, Cambridge, MA 02138 current address: Department of Entomology, Comstock Hall, Cornell University, Ithaca, NY 14853
Manuscript received 7 July 1993.




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60 Psyche [vol. 101
morphological characters, especially the significant differences in their genitalia, and their geographic distribution. MATERIALS AND METHODS
Most available specimens had been identified only to genus, excepting those members of the two species known when this research commenced in 1989: A. hamata and A. cornigera. The spiders were preserved in 80% ethanol; some were over fifty years old, so that original color patterns were faded and body parts had darkened with age. They were examined using an American Opti- cal dissecting microscope, on low power (8.4X magnification) and medium power (48X magnification). Descriptions and measure- ments were made following H. W. Levi's suggested guidelines: the color and patterning of the abdomen were noted, along with the shape of the cephalic region, relative distance between eyes, pres- ence of leg macrosetae, etc. The entire thickness of the animal was measured to determine carapace height. When possible the overall length of ten males and ten females of each species was measured, to give some idea of size variation among the species. In addition, in a male and female of each species, the articles of the first leg were measured, and the patella-tibia length of the other legs. Due to the great natural variation within populations, which renders leg length an uninformative character, other specimens were not simi- larly measured.
Collecting localities were recorded in the computer program Filemaker 11, then organized by country, latitude, and longitude. In many cases gazetteers were consulted to provide latitude and lon- gitude when only place names had been indicated. These locations were used to create dot maps on outlines of the North and South American continents to indicate the range of the species. For each species the genitalia and dorsal pattern were drawn. In males the left palpus was removed and drawn in mesa1 view, then turned so that the paracymbium could be illustrated. In females the epigynum was drawn in ventral and posterior (up-turned) view. In some cases drawings of the dorsal pattern were composites of sev- eral specimens to take individual variations into account. New species were identified on the basis of genitalic and dorsal- pattern differences. Further specimens were compared with draw-



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ings of the holotypes, and double-checked by H. W. Levi before locality information was recorded.
I added three new species, Acacesia villalobosi, A. yacuiensis, and A. benigna, to the two previously known members of the genus, A. cornigera Petrunkevitch and A. hamata (Hentz). The new species were collected solely on the South American continent, whereas A. hamata and A. cornigera range further north along the isthmus of Panama and into North America. In 1892 Simon split up the huge genus Epeira (the name previ- ously used by Hentz) into a number of genera including Acacesia. To single out individual species I relied upon differences in geni- talia. Levi (1976) had noted in his diagnosis "In Acacesia alone among related genera the paracymbium is modified, and not simply hook-shaped". The paracymbium ornaments the posterior edge of the base of the cymbium which surrounds the palpus (P in Figs. 8, 16). I used the evidence of variations of the paracymbium to sepa- rate males of three new species: A. yacuiensis, A. villalobosi, and A. benigna. Otherwise, the males of the genus showed marked uni- formity in the design of the palpus itself. The main obvious differ- ences were in the shape and proportions of the median apophysis, the large structure (two-pronged in this genus; M in Fig. 1) that may grasp the scape of the epigynum in maneuvering the male's sperm-transferring embolus into contact with the females' sperm duct (Foelix 1982).
An examination of the mapped location of specimens demon- strates the overlap of the various species of the genus, for the most part sympatric. It is possible that strong genitalic (and likely unknown behavioral) differences have maintained reproductive isolation allowing for sympatric speciation, despite the notable homogeneity of dorsal pattern and body form. None of the species were abundant within the lower Amazon basin area; topographic maps indicate a bare region in the watershed. It is possible either that the habitat is different from that usually frequented by the genus, perhaps due to its lower elevation or higher annual rainfall,



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62 Psyche [vo~. 101
or that the topography and vegetation of the region effectively bars collectors from entering. More information about the ecology of collection sites would help to clarify this situation. Preference for certain collection sites certainly has affected the geographic data for other members of the genus. A. villalobosi is a case in point: most of the specimens were collected within a short distance of two major Brazilian cities, $20 Paulo and Porto Alegre. This sug- gests that collectors restricted their efforts to the most accessible regions. The relative paucity of specimens also makes it difficult to make any generalizations about the range of this species. The situ- ation is similar for A. yacuiensis and A. benigna. A more complete biogeographic picture of Acacesia awaits further fieldwork, espe- cially in those regions which appear devoid of specimens. Acacesia Simon
Acacesia Simon, 1892: 795. Type species by original designation and monotypy Epeira foliata Hentz (= E. hamata Hentz). The name is feminine (Levi 1976: 374).
Description (see also Levi, 1976). Dorsum marked with a dag- ger shape, which is outlined in black and surrounded by a triangu- lar folium. Parallel rows of orange-brown dots extend posteriorly on either side of the dagger (Fig. 6). Depending on state of preser- vation and regional or intraspecific variation, branching pattern may appear extending from the tip of the dagger, often terminating in a straight line. Other variations: presence of dark gray shoulder patches and coloration within and outside folium. These variations do not appear marked enough to allow for distinguishing species, although familiarity with specimens may reveal a gestalt that is clear at least to the researcher.
Width of cephalothorax directly behind the posterior lateral eyes less than or equal to one-half the width of the carapace at its widest part (Figs. 6, 11, 14, 19, 22, 25, 30). Males with macroseta on the fourth coxa and trochanter, and one prominent spur, sur- rounded by shorter and thinner macrosetae, on the swollen second tibia. One macroseta on the palpal patella. First coxa laterally elongated forming a hook on the outside corner. Male palpus with bifurcate median apophysis (M in Fig. 1); length of the tines of this apophysis vary according to species, as does the shape of the hinge where the apophysis connects to the palpus. Embolus resem-



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bling a bird's head with a curved and pointed beak, covered by a transparent lamella (E in Fig. 1). Paracymbium varies interspecifi- cally in size and shape (Figs. 8, 16, 21, 27, 33). As with the para- cymbia, the sclerotized edges of the lateral plates of the underside of the female epigynum show great interspecific variation and range of complexity. Scape of the epigynum always extends poste- riorly from the base (Fig. 4).
Only the natural history of A. hamata is known in any detail, as the other species were identified solely from preserved museum specimens. Thus natural history is unavailable for the individual species diagnoses that follow. Due to the overall similarity of dor- sal pattern within the genus, only striking interspecific variation is noted in the species diagnoses.
Diagnosis. Hentz's description of Epeira foliata (1847: 475) serves to diagnose the genus: "abdomen ovate terminating in a joint, with waved black lines, two external almost meeting at the apex, two internal meeting before or near the middle." Much later Kaston (1978: 144) also noted hamata's triangular folium with "undulating margins." The genus is distinguished from its close relatives Ocrepeira and Cyclosa by the foliate dorsal pattern (Fig. 6) which is largely invariant from species to species (Petrunke- vitch, 1925, noted its "generic value"), as well as by the oval abdomen, which forms a shield shape in Ocrepeira and is often marked by tubercles in Cyclosa. In Acacesia the abdomen is an ovoid, rounded-off diamond shape often with "shoulder" humps and a small hump at the extreme anterior end (Figs. 6, 11, 14, 19, 25, 30). The spinnerets are at the posterior end of the abdomen (Fig. 31), as in Ocrepeira, but differing from Cyclosa where they lie in the center, beyond which extends a "tail." Males of related genera have rows of macrosetae on the second tibia, as opposed to Acacesia's main enlarged spur (Fig. 3). The embolus varies little within the genus, contrasted, for example, with Cyclosa, some species of which have extremely long emboli curving around the palpus. The paracymbium has evolved different ornamentation (P in Fig. 2; Figs. 8, 16, 21, 27, 33), whereas in pre- sumed sister genera it appears constant as a simple club-shaped attachment. (Within Acacesia at least, it proves useful as a deter- minant of species). The extended epigynal scape occurs also in Ocrepeira, but overlaps the base in Cyclosa. Whereas in Ocrepeira



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64 Psyche [VOI. 101
the scape may be twice the length of the base, reaching almost to the spinnerets, in Acacesia the scape is rarely much longer than the base (Figs. 4, 9, 12, 17, 23, 28).
White spots on the venter appear constant only in A. villalobosi (Fig. 31), where they are large enough to be visible without the aid of a microscope, and in A. yacuiensis. The other Acacesia species appear to have white stripes extending the length of the venter and surrounding a gray central patch.
Misplaced Acacesia.
Two single specimens do not fit into any of the species: one from La Paz, Bolivia (AMNH) may be a penultimate ready to molt, and one female from Alto Salim6es (MCN 8821) which remains puzzling. It may be a cornigera.
Epeira vegeta Keyserling, 1865: 819, pi. 19, figs. 31-34, 0, Q, was placed in Acacesia by Simon, 1895: 789, figs. 858-859, 0, but was placed by F. P.-Cambridge in Eustala, 1904: 509, pi. 48, figs. 16, 17, Q, 0. Chickering (1955: 511) redescribed and illustrated the Central American Eustala species.
1.
Base of epigynum a trapezoid with posterior edge wider than anterior and wide spatulate scape (Fig. 28); sclerites in poste- rior view of epigynum like reversed quotation marks (Fig. 29); southeastern Brazil (Map 1) ...................................... villalobosi -
Base of epigynum and scape otherwise (Figs. 4, 9, 17, 23); ................... posterior sclerites otherwise (Figs. 5, 10, 18, 24). 2 2. (1) Scape with lip on posterior end (Fig. 9) and anteriorly scle- rotized median plate in posterior view (Figs. 10, 13); central Mexico to southern Brazil (Map 1) ........................... cornigera -
Scape without lip (Figs. 4, 17, 23); median plate in posterior ............................................. view otherwise (Figs. 5, 18, 24) 3 3. (2) Scape shorter than base (Fig. 17); in posterior view sclero- tized median edges of lateral plates kidney-shaped with ridge or wrinkle at anterior end (Fig. 18); southeastern Brazil, north- . .
eastern Argentina (Map 1) ........................................ yacuiensis -
Scape approximately as long as base; base with two visible dark ...........
spots (Figs. 4, 23); lateral plates otherwise (Figs. 5, 24) 4




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Map 1. Distribution of Acacesia species. 4. (3) Visible part of median plate hourglass-shaped (Fig. 5); North and South America (Map 1) ................................ hamata - Median plate otherwise; sclerotized lateral plates resemble Mickey Mouse hat with big ears (Fig. 24); Peru, Brazil, .......................................................... Bolivia (Map 1) .benigna
.....................
1. Paracymbium wider than high (Figs. 8, 21, 33) .2 -
Paracymbium higher than wide (Figs. 16, 27) ........................ 4 2. (1) Paracymbium with distal lobe, otherwise unadorned; margin of proximal lobe convex (Fig. 8); North and South America ....................................................................... (Map 1) .hamata




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66 Psyche [vo~. 101
-
Paracymbium with additional appendage or ornamentation, like thorn or lip (Figs. 21, 33) ................................................ 3 3. (2) Thorn-shaped projection of paracymbium hanging over dis- tal lobe. Margin of proximal lobe concave (Fig. 21); southeast- . .
ern Brazil (Map 1) ................................................... yacuzenszs - No such projection; instead, distal lobe of paracymbium with folded lip at edge, overlapped by partly transparent proximal lobe. Transparent margin of proximal lobe straight, opaque mar- gin concave (Fig. 33); southeastern Brazil (Map l) ... villalobosi 4. (1) Distal lobe of paracymbium longer than proximal lobe, both anteriorly directed (Fig. 16); central Mexico to southern Brazil (Map 1) ................................................................... ..cornigera - Distal lobe of paracymbium and proximal lobe approximately the same length, both anteriorly directed (Fig. 27); Peru, Brazil, Bolivia (Map 1) ................................................ benigna Abbreviations: AME, anterior median eyes; PME, posterior median eyes; LE, lateral eyes. For museum collection abbreviations refer to the Acknowledgements section.
Figure 1. Expanded left palpus of male A. hamata. Cali, Colombia (MCZ). M, median apophysis; C, conductor; E, embolus; R, radix; P, paracymbium; CY, cym- bium; L, terminal apophysis or embolus lamella. Figure 2. Palpus of male A. cornigera. P=paracymbium. Panama (PMY). Figure 3. Left second tibia of male A. cornigera, posterior. Panama (PMY). Figs. 1-3 by Levi.
Figures 4-8. Acacesia hamata (Hentz). 4-6, female. 4. Epigynum, ventral. Chia- pas, Mexico (MCZ). 5. Epigynum, posterior. Same. B, base of epigynum; S, scape of epigynum; MP, median plate; LP, lateral plate. 6. Dorsal. Quezaltepeque, El Sal- vador (CAS). 7, 8, male, left palpus. 7. Mesal. Cali, Colombia (MCZ). 8. Paracym- bium. Naciemente del Rio Frio, Tamaulipas, Mexico (MCZ). Figures 9-16. A. cornigera Petrunkevitch. 9-11, female. 9. Epigynum, ventral. Barro Colorado Island, Canal Zone (MCZ). 10. Epigynum, posterior. Near Gamboa, Panama (MCZ). 11. Dorsal. Sirena, Osa Peninsula, Costa Rica (MCZ). 12. Epigy- num, ventral. Aparo, S3o Paulo, Brazil (MZSP). 13, Epigynum, posterior. Aparo, Siio Paulo, Brazil (MZSP). 14. Dorsal. Aparo, Sao Paulo, Brazil (MZSP). 15, 16, male, left palpus. 15, Mesal. Barro Colorado Island, Canal Zone (MCZ). 16. Para- cymbium. Same.
Scale lines. 1.0 mm, genitalia 0.1 mm.




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Glueck
Acacesia hamata (Hentz)
Figures 1, 4-8
Epeira hamata Hentz, 1847: 474, pl. 3 1, fig. 10, 0. Specimen from Alabama in Bos. SOC. Natur. Hist., destroyed (Hentz 1875 ed. Burgess, vii).
Epeira foliata Hentz. 1847: 475, pl. 31, fig. 14, Q. Female speci- men from Alabama in Bos. SOC. Nat. Hist., destroyed (Hentz 1875 ed. Burgess, vii). (Not E. foliata C. L. Koch, 1845). Erner- ton, 1884: 318, pl. 34, fig. 10, Q, pl. 37, figs. 6-10, 0. McCook7 1894: 154, pl. 4, figs. 7, 8, Q, 0. F. P.-Cambridge, 1904: 502, pl. 48, figs. 1, 0, and 2, Q.
Epeira folqera Marx, 1890: 545, 593. New name for foliata Hentz, preoccupied.
Araneus hallucinor Petrunkevich, 191 1 : 296. New name for Epeira hamata Hentz, thought preoccupied by Araneus hamatus Clerck, (= Singa hamata).
Acacesia foliata: - Cornstock, 1912: 509, figs. 546? 548, Q, 547, 0. Cornstock, 1940: 522-3, figs. 546, 548, Q, 547, 0. Bonnet, 1955: 122.
Acacesia lanceolata Badcock, 1932: 20, fig. 14, Q. Type specimen from Nanahua, Paraguay; in BMNH, examined. (Synonymized in Roewer, 1942, Bonnet, 1955).
Figures 17-22. A. yacuiensis. new species. 17-19, female. 17. Epigynum, ventral. Municipio de Itu, Siio Paulo, Brazil (AMNH). 18. Epigynum, posterior. Same. 19. Dorsal. Composite drawing of specimens from Arroyo Yacui, Parque Nacional Iguazu, Misiones Province, Argentina (M.E. Galiano). 20-22, male. 20. Left palpus, mesal. Baixo Jordiio, Sider6polis, Santa Catarina, Brazil (MCN). 21. Left palpus, paracymbium. Same. 22. Dorsal. Same.
Figures 23-27. A. benigna. new species. 23-25, female. 23. Epigynum, ventral. Zona Reservada de Manu, Puesto Vigil, Pakitza, Madre de Dios, Peru (USNM). 24. Epigynum, posterior. Same. 25. Dorsal. Puesto de Vigilancia 22 "Falso Paquisha" (sic), Alto Rio Comaina, Amazonas, Peru (MHNSM). 26, 27, male, left palpus. 26. Mesal. Est. Biol. Beni, Beni, Bolivia (USNM). 27. Paracymbium. Same. Figures 28-33. A. villalobosi. new species. 28-31, female. 28. Epigynum, ventral. Serra dos Orgiios, Rio de Janeiro, Brazil (MCZ). 29. Epigynum, posterior. Same. 30. Dorsal. Fazenda Sta. Maria Amparo, Siio Paulo, Brazil (MZSP). 31. Abdomen, ventral. Canela, Rio Grande do Sul, Brazil (MCN). 32, 33, male, left palpus. 32. Mesal. Campo do Jordiio, Siio Paulo, Brazil (MZSP).
33. Paracymbium. Same.
Scale lines. 1 .O mm, genitalia 0.1 mm.




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70 Psyche [VOI. 101
Acacesia folqera: -Roewer, 1942: 763.
Acacesia hamata: - Bryant, 1945: 364. Kaston, 1948: 235, fig. 705, 726,0,725, Q. Levi, 1976: 376, figs. 74-87, Q, 0. Araneus nigrolineatus di Caporiacco, 1955. Female holotype from Rancho Grande, Aragua, Venezuela in UCV. NEW SYN- ONYMY.
Description. Female, from Nantahala Gorge, North Carolina, MCZ. Carapace yellow-brown. Clypeus white. Chelicerae light brownish yellow. Labium light yellow-white, endites light brown- ish yellow. Sternum light yellow-white. Coxae light yellow-brown; legs yellow-brown. Dorsum of abdomen light gray-brown, sides off-white, venter tan. PME 1.1 diameters of AME, LE 0.5 diame- ters. AME 2.2 diameters apart, 2.5 from LE. PME 1.0 diameters apart, 3.5 from LE. LE less than 1 diameter apart. Total length 5.4 mm. Carapace length 2.2 mm, width 1.9, height 1.4. First femur 3.9 mm, patella-tibia 4.4, metatarsus 3.2, tarsus 1.0. Length of first leg 12.5. Second patella-tibia 4.3 mm, third 1.8, fourth 3.0. Male from Lago Calima, Depto. Valle, Colombia, 1,400 m elev., MCZ. Color as in female. PME 0.6 diameters of AME, LE 0.5 diameters. AME 2.0 diameters apart, 1.0 from LE. PME 2.0 diame- ters apart, 3.0 from LE. LE less than 1 diameter apart. Total length 4.3 mm. Carapace length 1.9 mm, width 1.8, height 1 .I. First femur 2.3 mm, patella-tibia 3.5, metatarsus 2.3, tarsus 0.5. Length of first leg 8.6. Second patella-tibia 2.2 mm, third 1.2, fourth 1.8. Variation. Total length of ten females 5.3 to 8.6 mm, of ten males, 3.6 to 4.4.
Diagnosis. Specimens of A. hamata lack distinct white spots on the venter of the abdomen. The straight-line folium pattern (Fig. 6) is characteristic and differs from the more sinuous forms on the backs of A. cornigera, A. yacuiensis, and A. villalobosi. A. hamata's abdomen is smoothly oval, not with protruding "shoul- ders" above a constriction, as is common in A. cornigera and A. yacuiensis. Almost total absence of intraspecific variation in the paracymbium (Fig. 8) makes it a good character by which to iden- tify males. It is the simplest form in the genus, with proximal and distal lobe of equal height, and with the proximal lobe forming a convex curve leading into the distal lobe. In the females the edges of the lateral plate of the epigynum overlap the median plate to form an hourglass shape (Fig. 5).




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Natural History. See Levi 1976.
Distribution. The species ranges from eastern Pennsylvania to western Arkansas, south into Florida and the Bahamas, through Mexico into northern Argentina.
Specimens examined. ARGENTINA Chaco: Presidencia Roque Saenz Pefia (MACN); Selvas del Rio de Or0 (Galiano). Misiones: Eldorado (AMNH). BAHAMAS South Bimini (AMNH) ; Freeport (AMNH). BOLIVIA La Paz: Yungas, Puente Villa (IRSNB). BRAZIL Amapd: Oiapoque (AMNH). Espi'rito Santo: Guarapari (MZSP). Goids: Siio Francisco, Jaraguii (MZSP); Fazenda Mon- jolinho, Corumbii (MZSP); Jatai (AMNH). Mato Grosso: Rosiirio Oeste (AMNH); Fazenda Beija-Flor (MZSP). Minus Gerais: Bar- bacena (AMNH); Pedra Azul (AMNH); Vi~osa (AMNH); Lavras (MCZ); Varginha (AMNH). Pard: Belbm, Val-de-Ciies (AMNH). Parai'ba: Indephcia (MCZ). Parand: Rolhdia (AMNH). Pernam- buco: Recife (MNRJ). Rio de Janeiro: Silvestre (AMNH); Floresta dos Macacos (AMNH); Santa Maria Madalena (AMNH); Santo Ant6nio do Imb6 (AMNH); Silva Jardim (AMNH); Nova Iguap, Miguel Couto (AMNH); Repress Rio Grande (AMNH); Mangarat- iba (AMNH); Angra dos Reis (MZSP); Siio Conrad0 (MCZ); Niter6i (MNRJ). Rio Grande do Norte: Fazenda Canaan (MZSP); Fazenda Camiio, Macau (MZSP). Rio Grande do Sul: S. Heupolblo (MZSP); Morro Siio Pedro (MCN); Tapera (MNRJ); Mupm (MCN); Encantado (MCN); Roca Sales (MCN); Montenegro (MCN); General C2mara (MCN); Triunfo (MCN); Mono Alto, Gravatai (MCN); Porto Alegre (MCN); Viamiio (MCN); Santa Maria (MCN); Taquara (MCN); Butiii (MCN). SGo Paulo: Fazenda Joiio, Inda Verde (MZSP); Itaguers, Nova Europa (MZSP); Esta~, Exper., Pirassununga (MZSP); Ilha dos Buzios (MZSP); Barueri (MZSP); Ubatuba (MZSP); Municipio de Itu, Fazenda Pau d9A1ho (AMNH); Siio Roque (AMNH). BRITISH VIRGIN ISLANDS Great Thatch Island: Tortola (AMNH). COLOMBIA Antioquia: La Estrella (MCZ). Cundinamurca: Monte Redondo (MCZ). Mag- dalenu: Pueblo Bello, Sierra Nevada de Santa Marta (AMNH). Valle: Lago Calima (MCZ); Rio Jamundi (MCZ); Cali (MCZ). COSTA RICA Cartago: Turrialba (AMNH). Sun Josk: San Jose (AMNH); Carillo (MCZ). Guanucaste: Cafias: La Pacifica (MCZ). CUBA Pinar del Ri'o: Peninsula de Guanahacabibes (AMNH). Cienfuegos: Soledad (AMNH), ECUADOR Guayas: 8 mi. S



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72 Psyche [vo~. 101
Manglaralto (CAS). GUATEMALA Nebaj (AMNH); Tikal (MCZ). Pet&: Santa Elena (AMNH). EL SALVADOR Quezaltepeque, Q (CAS). GUYANA Yupukari, Rupununi River (AMNH). HON-