THE AMERICAN ORB-WEAVER GENERA COLPHEPEIRA, AND GASTERACANTHA NORTH OF MEXICO (ARANEAE, ARANEIDAE)

HERBERT W. LEVP

Abstract. Colphepeira has only one species from arranged in loops toward the median side United States. There are four the southeastern in the posterior median eyes. In some of Micrathena north of Mexico, three com- species (Pachygnatha, and also Linijphia) only the mon ones in eastern North America from New- lateral have the canoe-shaped tapetum England to the tropics, one uncommon from Ari- eyes lost all zona to Guatemala. All Micratheria known, per- left. Tetragnatha have tapetum. ;| haps 50 species, are tropical American, the three Some arachnologists consider the absence north each to a extending their range belong of epigynum in Pachijgnatha and Tetra- different species group. Even though tropical and gnatha a primitive feature and the two widespread in the eastern states, M. mitrata and ancestral, related to the haplogyne M. gracilis appear absent from southern Florida. genera sliver of the all relatives have All Micrathena species have only a . Out-comparison ( canoe-shaped tapetum left, the latter a character- a canoe-shaped tapetum, and an epigynum) Araneoidea. The istic of most of the superfamily would indicate that the loss of both struc- cosmotropical Gasteracantha has only one or tures be secondary. To learn more two species in the Americas, G. cancriforinis in the may these the has warmer parts of North America. about relationships, tapetum to be examined (Figs. 4, 5, 19, 20, 75, 76). INTRODUCTION Already Homann's eye studies indicated that Colphepeira belongs to the Araneinae, Homann 1971 ( 1950, ) reported diversity not close to Meta or Theridiosoma as previ- in structure within the secondary eye ously thought. My study of Colphepeiras family Araneidae. The tapetum of the o-enitalia confirms Homann's conclusions. A secondary eyes is usually canoe-shaped: new observation, not previously reported, with a crease through the middle dividing is that Micrathena species have only rem- median the tapetum into two parts facing each nants of a tapetum in the posterior to other (Figs. 4, 5, 75, 76). This is char- eyes. Perhaps this is only an adaptation acteristic for members of the superfamily Micrathenas diurnal habits or perhaps it Araneoidea and some related families in- will be of use for figuring out phylogenies. eluding Agelenidae, Gnaphosidae, Clubion- Also of interest are the relatively large tarsal claws of idae, and Amaurobiidae. Meta and Zij- accessory setae below the giella have a large canoe-shaped tapetum, Micrathena. Micrathena may be a good ex- han- like Theridiidae, however the tapetum ap- perimental for studying the pears reduced in Araneus (and also Col- dUng of silk (a subject about which we of phepeira. Figs. 4, 5) with rows rhabdomes know little), because its setae against which silk strands are held by the median claws are than those of other ^Museum of Comparative Zoology. larger genera. 417 Bull. Mus. Comp. Zool, 148(9): 417-442, September, 1978 418 Bulletin Museum of Comparative Zoology, Vol. 148, No. 9 COLPHEPEIRA, MiCRATHENA AND GaSTERACANTHA NoRTH OF MeXICO • Lcvi 419

Plate 2. Gasteracantha cancriformis (Linnaeus) web, spider removed, dusted with cornstarch. Viscid area 17 cm in diameter. Web was built at 45° angle in porch corner of abandoned Florida house: lower left, floor; above and far right, walls. Notice tufts of silk on frame lines and on some radii.

com- Colphepeira contains only one small- attack-wrap prey ( Robinson, personal sized species and is related to Singa and munication). The lack of attack-wrapping perhaps Mangora. MicratJiena and Gaster- is probably not primitive but a secondary acantha are tropical spiders; Micrathena, loss. Micrathena and Gasteracantha orb- with more than 50 species, are all tropical webs have open hubs (Plates 1, 2). Micra- American. Three of these have successfully thena rests in the center of the web in an their the to unusual Plate 1 and controls extended range from tropics position ( ) temperate eastern North America (Map 2). web tension. Unlike other araneid genera Each of the three belongs to a different Micrathena have strong fourth legs, used species group. I believe Gasteracantha and to hold its position in the web (Plate 1). Micrathena to be specialized Araneidae Both Micrathena and Gasteracantha are which have lost some of their palpal scler- diurnal spiders. for ites secondarily, and Micrathena do not I would like to thank colleagues

Plate 1. Micrathena gracilis (Walckenaer) in web, Virginia. The lower photograph is about life-size. Web dusted with cornstarch. Notice the unusual position of the spider in the open hub and the use of the fourth leg (in upper photograph). No. 9 420 Bulletin Museum of Comparative Zoology, Vol 148,

dorsal were making collections available. P. H. Arnaud examination. The posterior eyes and R. X. Schick of the California Academy illustrated with the left eye flat and the at an anterior is on of Sciences, D. Bixler, J. A. Beatty, J. Carico, right angle, top ( Figs. R. Crabill of the National Musenm of Nat- 4, 19, 75). The left lateral eyes were illus- first the anterior ural History, C. Dondale of the Canadian trated diagrammatically, of the left flat under the then the National Collections, S. I. Frommer ( ) microscope, University of California Riverside collec- spider was shifted for the posterior eye H. flat under the The tions, W. R. Icenogle, B. J. Kaston, (right) microscope. of the Field Mu- illustrations are thus Dybas and J. B. Kethley produced composites The rows of rhabdomes seum of Natural History, T. Kronestedt of (Figs. 5, 20, 76). seen in mounts of the the Natural History Museum, Stockholm, can be microscope with a R. E. Leech, G. Uetz and J. D. Unzicker of eyes compound microscope. and British diction- the Illinois Natural History Survey, S. C. Following American not Johnson, W. Peck of the Exline-Peck col- ary definitions but arachnological lection, N. Platnick of the American Mu- vocabulary, spines are immovable, rigid, or as found on the seum of Natural History, W. T. Sedgwick, pointed humps thorns, Gasteracantha. W. Shear, W. Star^ga of the Polish Academy abdomen of Micrathena and of Science, Warsaw, H. K. Wallace, H. V. The movable heavy setae covering the in- Weems of the Florida State Collection of tegument are called macrosetae. , F. R. Wanless of the British Museum, Natural History. The mapping Colphepeira Archer and typing were done by Susan Hunt. The Colphepeira Archer, 1941, Geol. Surv. Alabama, research and its were made publication Mas. Paper, 18: 12. Type species Epeira ca- possible by National Science Foundation tawba Banks by original designation. The name grants BMS 75-05719 and DEB 76-15568. is feminine. METHODS Diagnosis. Colphepeira differs from many other araneid genera by the closely spaced The method of examination, study, and eyes (Figs. 1-3). Unlike Mangora and illustration are those of other studies in this Singa, it has a hirsute carapace and ab- series on North American orb-weavers. domen with short setae on granules and However in Micrathena and Gasteracantha, tlK> shape of the abdomen is higher than measurement of total length is the length long with posterior dorsal tubercles (Figs. in midline from the anterior margin of the 1, 6, 7). It differs from Mangora by the carapace to between the posterior abdom- lack of the characteristic long trichobothria inal humps or spines. on the third tibia. The tapetum of the secondary eyes was Description. The carapace, abdomen, examined by near-vertical illumination and legs are covered with scattered short (with a fiber-light) on the eye. The spider setae. The setae are cylindrical, distally is kept submerged in alcohol and posi- tapering to a blunt point, basally with a tioned on washed sand, which permits odd narrow neck and sitting in the center of a positions and very minor changes in angles. disc. The sides of the setae have some of is a blunt teeth are ( Because reflections, sand poor (Fig. 9). The eyes subequal background for most observations; the to each other in the female, and the an- should black. terior are in the male. The background be ) The mag- larger posterior nification of the stereoscopic dissecting eye row is recurved. The median eyes are microscope is about 50 times for the larger slightly more than their diameter apart, and species, 100 times for the smaller. If the the anterior medians are less than two eye lens has become opaque in preservation diameters from the laterals, the posterior the spider can be cleared in clove oil for medians 2.5 diameters from the laterals. COLPHEPEIRA, MiCRATHENA AND GaSTERACANTHA NoRTH OF MeXICO • Levi 421

TT The thorax has a shallow depression in the Colphepeira catawba ^^ female (Fig. 3) and a transverse, pro- curved mark in the male. The posterior 1 swollen. The part of the head is slighUy than height of the clypeus is slightly more the diameter of the anterior median eyes like the (Fig. 2). The sternum, carapace, is lightly sclerotized and slightly granulate. The chelicerae are weak, without a basal boss, and have two teeth on the anterior margin, two on the posterior and denticles and one tooth in between (Fig. 8). The chelicerae have a proximal anterior pro- as in Theridiidae jection under the clypeus articles of the i^Fig. 8). The proximal legs, 1. Distribution of Colphepeira catawba (Banks), the femora, are also slightly Map especially Mexico. the north of granulate. The first legs are longest, fourth second in length, the third shortest. The metatarsus and tarsus together are terminal apophysis extends and covers most shorter than the patella and tibia. The of the bulb laterally (Figs. 12-15). abdomen is higher than long with postero- Relatiomhip. A similar large terminal dorsal and posterior tubercles (Figs. 1, 6, apophysis (A in Fig. 15) is found in some like 7). The lung covers are smooth and, species of Singa (Levi, 1972) and Mangora of the those of Meta, lack the transverse grooves (Levi, 1975). The resemblance hamata found in species close to Aroneus. The palpus (Fig. 14) to that of Singa the leaflets of the book-lung in a microscope (Clerck) is striking in the shape of soft con- to consist of series of M , the mount appear parallel small median apophysis ( )

to other side side. , the tenninal tracheae attached each by ductor ( C ) and large apophy- The males are hke females, slightly sis (A). The embolus of Colphepeira is subter- smaller and have a more distinct, transverse, simpler, and Colphepeira lacks a thoracic depression and a slightly higher minal apophysis. Other similarities to Singa clypeus, 1.5 diameters of the anterior me- hamata and Mangora are the lightly scler- dian eyes. The endite has a tooth facing a otized epigynum, with a soft broadly at- tooth on the end of the proximal palpal tached scape and the closely spaced eyes. femur. The distal of the first coxa of margin The genitalia also resemble those Cer- has a hook that fits into a groove on the cidia (except for Cercidias large median second femur. The legs are not modified apophysis). All these related genera Col- that the anterior tibia is except slightly and Cercidia sinuous. phepeira, Singa, Mangora, have the eyes relatively closely spaced, Genitalm. The soft epigynum is covered unlike those of the Aranetis, with setae (Fig. 9) and has a soft annulate larger-sized Gasteracantha and those of scape (Figs. 9, 10). The openings appear Micrathena, numerous other araneid on the posterior face (Figs. 10, 11). There genera. CoJpJie- like are fertilization ducts. The male palpus peira, unlike most araneid genera but distinct has a soft conductor (C in Figs. 14, 15), Mangora, does not have contrasting bearing a basal tooth, a median apophysis ventral abdominal marks. the (M), which has a hook, and a very large After he examined the tapetum of terminal apophysis (A) which covers con- secondary eyes (Figs. 4, 5) Homann (1950) is more ductor and embolus (E) distally. The first reported that Colphepeira 422 Bulletin Museum of Comparative Zoology, Vol. 148, No. 9

closely related to Araneus than to Meta or patella and tibia, 1.1 mm; third, 0.6 mm; Thcridiosoma. fourth, 0.7 mm. Distribution. Only one species of Col- Variation. Some specimens have little pJiepeira is known, C. cotaaba, found in black pigment, others are almost completely Total of females 2.2 to 3.8 the southeastern United States (Map 1). black. length , mm; carapace 1.0 to 1.3 mm long, 0.9 to Colphepeira catawba (Banks) 1.1 mm wide. Total length of males, 1.6 to 2.2 0.9 to 1.2 mm 0.9 Figures 1-15; Map 1 mm; carapace long, mm to 1.0 mm wide. catawba 1911, Pioc. Acad. Natur. Epcira Banks, Diagnosis. This species can only be con- Sci. Philadelphia, 63: 450, pi. 34, fig. 4, $. fused with Dolichognatha species, which Female holot\pe from Ash\'ille, Buncombe are of similar size and Doli- County, North Carolina in the Museum of Com- appearance. parative Zoology, lost. There is no old E. B. cJwgnatha, a relative of Tetragnutha, has Bryant catalog card in the file for this species four small tubercles on the abdomen, and as there is for other Banks types. four Colplwpeira has tubercles postero- il Aranea catawba-. —Roewer, 1942, Katalog der dorsal on each side and in addition a Araneae, 1: 859. pair Colphepeira catawba: —Archer, 1941, Geol. Surv. posteriorly (Figs. 1, 6, 7). DoUcJ^ognatha Alabama, Mus. Paper, 18: 13, pi. 1, figs. 3, 4, species have their chelicerae elongate; 1-3. 1953 Amer. Mus. no. pi. 2, figs. Novitates, Colphepeira do not (Fig. 2). The Doli- 1622: 22, figs. 32-34. chognatha epigynum has a depression with Araiicus catawba: —Bonnet, 1955, Bibliographia a dark on each has Araneorum, 2: 452. spot side; Colphepeira a fleshy scape (Figs. 9, 10). The DoU- Female from Arkansas: car- Description. chognuthu male palpus appears rather black with white apace paired yellowish simple, Tetragnatha-]ike, but with a compli- Sternum patches lacking pigment. spotted cated paracymbium. That of Colphepeira black and Coxae yellow-white. yellow- has a terminal apophysis, median apophysis, white. with narrow Legs yellow-white and a simple paracymbium (Figs. 12-15). black Dorsum of abdomen with rings. Natural History. According to Archer streaks paired (Figs. 1, 7). Venter with 1941 catawba its hori- ( ) ColpJiepeira makes black and white and no distinct marks. spots zontal orb-web near the ground between The abdomen than slightly higher long, tree roots in thin open second-growth woods with four tubercles posterodorsal closely with grassy undergrowth. The web is 7 to on each side and a of tuber- grouped pair 9 cm diameter with about 60 spirals and cles on the face Total posterior (Fig. 7). a small, poorly defined hub. The egg-sac 3.5 mm; 1.2 mm 1.0 length, carapace, long; hangs with debris, suspended on a hori- mm wide. First femur, 1.2 mm; and patella zontal line just above the web. The spider tibia, 1.4 1.3 mm; metatarsus, mm; tarsus, has its retreat under loose bark and feeds 0.5 mm. Second patella and tibia, 1.2 mm; on small ants. 0.8 1.0 third, mm; fourth, mm. Distribution. Southeastern United States, Male from Arkansas with color pattern Virginia, southern Florida to Sonora (Map less distinct than female. Abdomen shape like that of female. Total length, 1.6 mm; Records. Virginia. Fairfax Co.: Great carapace, 0.9 mm long, 0.9 mm wide. First Falls, 9 (N. Banks). Georgia. Troup Co.: femur, 1.0 mm; patella and tibia, 1.2 mm; West Point, 7 Sept. 1949, "$ (A. Archer). metatarsus, 0.6 mm; tarsus, 0.4 mm. Second Florida. Monroe Co.: 2 mi SE of Mara-

Figures 1-15. Colphepeira catawba (Banks) 1-11. Female. 1. Dorsal view. 2. Eye region and chelicerae. 3. Carapace. 4. Posterior median eyes. 5. Left lateral eyes. 6. Abdomen from side. 7. Abdomen, pos- terior view. 8. Left chelicera from posterior. 9-11. Epigynum (with enlarged seta). 9. Ventral. 10. Pos- COLPHEPEIRA, MiCRATHEXA AND GaSTERACANTHA NoRTH OF MeXICO • Lcvi 423

I terior. 11. Posterior view, cleared. 12-15. Male, left palpus. 12. Mesal. 13. Ventral. 14. Mesa), expanded. 15. Ventral, expanded.

Scale lines. 0.1 mm, except Figures 1-7, 1.0 mm.

Abbreviations. A. terminal apophysis; C, conductor; E, embolus; H, hematodocha; M, median apophysis; P, paracymbium; T, tegulum. No. 9 424 Bulletin Museum of Comparative Zoology, Vol. 148,

Ala- it be hidden the curvature of thon, 15 Dec. 1962, jiiv. (W. Ivie). lens, may by The bama. Cherokee Co.; May's Gulfe, 11 Aug. the eyeball (Figs. 19, 32, 46, 60). contains rhabdomes without 1948, 13 Oct. 1949, 9 9 (A. Archer). Tusca- mesal side of a variable loosa Co.: Tuscaloosa, 2 Oct. 1941, 9 (A. tapetum, arranged in rows of 5 to 6 in Archer). Mississippi. Forrest Co.: Camp number loops, few, perhaps in M. about 8 Shelby, 1945-1946, 9, 6 (A. Ajrcher). Wil- number gracilis (Fig. 60), the other kinson Co.: Centreville, Jan.-July 1944, 9 to 9 loops in species (Figs. 19, is un- 32, 46 . The narrow lateral (A.Archer). Arkamus. Carroll Co.: Berry- ) tapetum like that of most of Araneidae ville, Aug. 1938, sununer 1941, Sept. 1944. species ( Fig. lateral be 9, $ (C.Wilton). Texas. Wilbarger Co.: 75). The eyes may separated 6 K. from each other as much as their di- 4 mi NW of Elliott, 21 Oct. 1964, ( W. by the rhabdomes to the sides of the Haller). Sonora. Guaymas, on beach, 13 ameter; 9 not are not in rows. Sept. 1966, (J., W. Ivie), mapped, tapetum arranged is smooth and received after completion of paper. Description. The carapace shiny in the female and has a unique light Micrathena Sundevall rim on each side (Figs. 18, 31, 45, 59). Posterior median eyes are 1.2 to 1.5 times Micrathena Sunde\all, 1833, Conspectus Arachni- the diameter of anterior medians, laterals dum, London, p. 14. Type species Epeira chjpeata Walckenaer, the only species listed in subequal or slightly smaller than anterior of The name is fem- "section one" the genus. medians. The median eyes are separated by inine. The of names synonymy problems generic their diameter to 1.5 diameters. The lat- are discussed by Bonnet, 1957 ( Bibliographia erals are several diameters from medians, Araneorum, 2: 2858). but may be up to slightly more than their Micrathena females differ from Diagnosis. diameter from each other (Fig. 47). The those of other in a smooth, genera ha\'ing height of the clypeus is equal to or slightly with a rim on each shiny carapace light more than the diameter of the anterior side 18, 31, 45, 59) and in particular, (Figs. median eyes (Fig. 57). The chelicerae are from Gasteracantha, by having the carapace slightly longer than wide, strong with three than wide in the female, at times longer to four teeth on the anterior margin, three of 31 or lateral with pairs dimples ( Fig. ) to four on the posterior. The legs are spines (in tropical species) unlike that of usually not banded. However, sometimes any other genera. The female abdomen the they are slightly lighter in color than is usually longer than wide, trapezoidal, or carapace and sometimes slightly granulated square armed with spines, sclerites and a (especially the long femora) bearing very sclerotized around the ring spinnerets (Figs. short setae. The abdomen is often brightly 17, 30, 44, while that of Gasteracantha 58), colored, always modified with spines, tuber- is usually wider than long. Males lack the cles, or folds. The spinnerets are sur- carapace rim and the abdominal spines rounded by a sclerotized ring. and have a sclerotized abdomen smooth, The males are smaller than the females with a around the The male ring spinnerets. and have the abdomen hghtly sclerotized. abdomen is longer than wide, not like that In the males, it is greater in length than in of Gasteracantha. The median eyes are \\'idth, but lacks the spines and tubercles never projecting as are those of Gastera- of the female. Thus it is quite difficult to cantha. The posterior legs of both sexes, associate with the females of the same especially the femora, are longer than the species (Figs. 16, 28, 41, 55). The palpal anterior legs or subequal in length, unlike femur lacks the tooth, and there those of most other araneid genera. The proximal is no tooth on the endite. The first posterior median eyes have the canoe- facing have a sometimes shaped tapetum reduced to a very narrow, coxae sometimes hook, lateral sliver. When viewed through the not; the hook is absent in the four species COLPHEPEIRA, MiCRATHENA AND GaSTERACANTHA NoRTH OF MEXICO • Levi 425

north of Mexico. The distal articles of the attack-wrap (M. Robinson, personal com- legs may not be modified and only some- munication). There is no retreat. All spe- times have macrosetae; in M. funebris the cies, north of Mexico, mature in fall in the first femur has macrosetae on the distal end northern part of their range. Little is known (Fig. 16). of egg-sacs and life histories. Genitalia. The epigynum is usually a Distribution. All species known are heavily sclerotized knob with openings at American. There may be as many as 50 the base of the face or more posterior ( Figs. 21-23, tropical American species, with 34-36, 48-50, 62-&4). There is no annulate only four extending their range into the scape. Together with the shape of the temperate area north of Mexico: three in abdomen, the epigynum is a diagnostic the eastern United States, one in the South- feature but has been west slighted by previous ( Map 2). authors. Note. In all species, the genitalia are The palpal patella has one macroseta in quite variable, and thus the species are M. funebris; in M. gracilis and M. sagittata difficult to delineate. The four species the macroseta is present, but small. The north of Mexico, however, are not closely parac\mbium (P in Fig. 40) differs in related to each other and are easily sep- different species (Figs. 25, 38, 52, 66), arated. unlike other araneid genera but as in Zy- Key to Female Micrathena North of Mexico giella. The bulb lacks a terminal apophysis 1. with but has a transparent flap which arises from Female abdomen 5 pairs of conical the the tubercles (Figs. 58, 59); eastern United base of embolus ( E ) and surrounds States to South America gracilis it. It be with the missing - may homologous Female abdomen with 3 or 2 pairs of terminal apophysis (Figs. 27, 40, 54, 68). spines or tubercles (Figs. 18, 31, 45) — . 2 The rests in with of embolus tip ( E ) the conductor 2(1) Abdomen only two pairs pos- (C). The conductor is sometimes a com- terior conical tubercles (Figs. 30, 31); carapace with 3 pairs of dimples ( Fig. plex sclerite and at its base another sclerite 31); eastern United States to South the may appear, paramedian apophysis America (Map 2) _.._ mitrata - (PM) (Figs. 40, 68). The median apophy- Abdomen with 3 pairs of tubercles or sis 45 and varies greatly in different species: a spines ( Figs. 17, 44, ) carapace

without . - 3 sclerotized, hook in M. M in paired dimples (Figs. 18, 45) split funebris ( 3(2) Abdomen much wider behind than ante- Fig. 27), the tip sclerotized in M. mitrata riorly and with anterior, lateral and large forked in M. (Fig. 40), sagittata (Fig. 54) posterior, dorsal spines (Figs. 44, 45); and reduced in M. gracilis (Fig. 68). The eastern United States to South America 2 ___ sclerites of the palpus are only lightly ( Map ) sagittata - Abdomen as \vide behind as in front with- sclerotized, unlike the sclerotized epigy- out anterior dorsal spines or tubercles, but num, and carapace spines. with dorsolateral, posterior dorsal and Natural History. Micrathena species are posterior ventral tubercles (Figs. 17, 18); diurnal and the spiders rest in the open Arizona, Baja California to Central America 2) funebris hub of the orb-web Plate 1 . (Map ( ) The spider hangs in a characteristic position, controlling Key to Male Micrathena North of Mexico the tension of the web while in the hub. The 1. Abdomen wider behind than in front spinnerets are up, the dorsal surface of the (Fig. 41); median apophysis of palpus

with two branches ( 53, M in 54 ) ; abdomen parallel to the ground (Plate 1) Figs. paracymbium a recur\'ed hook pointing and at an to the web The angle plane. dorsally (Fig. 52); eastern United States orb has radii and many spirals. No doubt to South America ( Map 2 ) sagittata - Abdomen as wide behind as in front the long fourth legs are an adaptation to (Figs. 16, 28, 55); median apophysis and the unusual position in the web. Unlike paracymbium otherwise (Figs. 26, 39, most Micratherw not - araneids, species do 67 ) 2 426 Bulletin Museum of Comparative Zoology, Vol. 148, No. 9

Map 2. Distribution of Micrathena species north of Mexico.

2(1) Abdomen more than twice as long as Micrathetia funebris (Marx in Banks) 55 end of carapace ( Fig. ) ; posterior Figures 16-27, IVIap 2 abdomen longer on \enter than dorsum

56 ; eastern United States to South ( Fig. ) Acrosoma fiinehre Mar.\ in Banks, 1898, Proc. Cali- America (Map 2) gracilis fornia Acad. Sci., 3rd ser., 1(7): 249. Female - Abdomen less than 1.5 times as long as syntypes from Calmilla Mines and Sierra San carapace, not longer on venter ( Figs. 16, Nicholas in the California Academy of Sciences, 28, 29) . -- 3 destroyed; and two syntypes from Mazatlan in 3(2) First femur with distal macrosetae ( Fig. the Museum of Comparative Zoology, examined. 16); paracymbium with large granulate Acrosoma macidata Banks, 1900, Canadian Entom., sculpturing (Fig. 25); median apophysis 32: 100. Female holotype from "Arizona" in the a semicircular hook on proximal end of Museum of Comparati\e Zoology, examined. bulb ( M in 27 ) ; Figs. 26, Arizona, Baja NEW SYNONYMY. California to Central America ( Map 2 ) Bio- fiinebris Micrathena granulata F. P. -Cambridge, 1904, — First femur without distal macrosetae logia Centrali- Americana, Araneidea, 2: 532, pi.

50, 12, S Male from ( Fig. 28 ) ; paracymbium smooth and fig. holotype Teapa, Mexico in the British Natural small ( Fig. 38 ) ; median apophysis with Museum, History, distal end pointed and bent i)ack (Figs. examined. Reimoser, 1917, Verb. Zool. Bot. Ges.

.39, M in 40); eastern United States to Wien, 67: 117. Roewer, 1942, Katalog der

..-. South America ( Map 2 ) mitrata Araneae, 1: 958. Bonnet, 1957, Bibliographia COLPHEPEIRA, MiCRATHENA AND GaSTERACANTHA NoRTH OF MeXICO • Levi 427

Figures 16-27. Micrathena funebris (Marx in Banks). 16. Male. 17-24. Female. 17. Lateral. 18. Dorsal. 19. Posterior median eyes. 20. Left lateral eyes. 21-24. Epigynum. 21. Ventral. 22. Posterior. 23. Lateral. 24. Posterior, cleared. 25-27. Male left palpus. 25. Lateral. 26. Mesal. 27. Submesal, expanded.

Scale lines. 0.1 mm; except Figures 16-18, 1.0 mm.

Abbreviations. C, conductor; E, embolus; H, hematodochia; M, median apophysis; R, radix; S, subtegulum; T, tegulum; Y, cymbium. 428 Bulletin Museum of Comparative Zoology, Vol. 148, No. 9

Araneoruni, 2: 2870. Chickering, 1961, Bull. southern Mexico and Guatemala differs in Mus. Comp. Zool., 125(13): 423, figs. 78-82, having a shorter sclerotized portion of the $ . NEW SYNONYMY. median apophysis and a more sclerotized Micrathena funehris: —Reimoser, 1917, Verh. Zool. Females in Bot. Gesell. Wien, 67: 104. Roevver, 1942, paramedian apophysis. vary Katalog der Araneae, 1: 958. Bonnet, 1957, total length 5.8 to 7.2 mm, carapace 2.4 to Bibliographia Araneoruni, 2: 2867. Chickering, 2.6 mm long, 1.5 to 1.7 mm wide. Males Bull. Mus. Zool., 125(13): 414, 1961, Comp. vary in total length 4.0 to 4.1 mm, carapace figs. 55-59, 9. 1.7 to 2.0 mm long, 1.1 to 1.3 mm wide. Micrathena maculata: — Reimoser, 1917, Verh. This is similar to M. Zool. Bot. Gesell. Wien, 67: 10. Roevver, 1942, Diag^nosis. species Katalog der Araneae, 1: 967. Bonnet, 1957, mitrata. Females differ in their lack of 2: 2871. SYN- Bibliographia Araneoruni, NEW the dimples on the carapace (Fig. ONYMY. paired 18), and the openings of the epigynum are iSlote. sus- not in a 22, There Chickering ( 1961 ) already depression (Figs. 24). is of anterodorsal on the pected that the male named M.

Sides of abdomen almost 16 . parallel ( Fig. ) ciety of Natural History, destroyed. Total length 4.1 mm, carapace 1.7 mm long, Acrosoma mitrata: —Emerton, 1884, Trans. Con- 1.1 mm wide. First femur, 1.6 mm; patella necticut Acad. Sci., 6: 327, pi. 38, fig. 9, $. Emerton, 1902, Common Spiders, p. 189, fig. and tibia, 1.5 mm; metatarsus, 1.0 mm; 438. 9. 0.5 mm. Second and tarsus, patella tibia, Acrosoma reduvianum: —McCook, 1893, American 1.4 mm; third, 0.8 mm. Fourth femur, 1.6 Spiders, 3: 213, pi. 21, figs. 6, 7, 9, i. Not PIcctana reduviana Walckenaer, 1841 (= M. mm; patella and tibia, 1.4 mm; metatarsus, gracilis ) . 1.1 mm; tarsus, 0.5 mm. Micrathcmi mitrata: —P.P. -Cambridge, 1904, Bio- Variation. The palpus of males from logia Centrali- Americana, Araneidea, 2: 538. CoLPHEPEiRA, MiCRATHENA AND Gasteracantha North OF MEXICO • Levi 429

Figures 28-40. Micrathena mitrata (Hentz). 28-29. Male. 28. Dorsal. 29. Lateral. 30-37. Female. 30. Lateral. 31. Dorsal. 32. Posterior median eyes. 33. Left lateral eyes. 34-37. Epigynum. 34. Ventral. 35. Posterior. 36. Lateral. 37. Posteriodorsal. 38-40. Male left palpus. 38. Lateral. 39. Mesal. 40. Submesal, expanded.

Scale lines. 0.1 mm; except Figures 28-31, 1.0 mm.

Abbreviations. C, conductor; E, embolus; M, median apophysis; P, paracymbium; PM, paramedian apophy- sis; R, radix; S, subtegulum; T, tegulum; Y, cymbium. 9 430 Bulletin Miiscwn of Comparative Zoology, Vol. 148, No.

Reimoser, 1917. Verhandl. Zool. Bot. Ges. Wicn, Variation. The dorsal abdominal black 67: 104. Roewer, 1942, Katalog der Araneae, 1: marks are smaller in specimens from Guate- 966. Kaston, 1948, Connecticut State Geol. mala and Panama. Females vary in total Natur. Hi.st. Surv., 70: 220, figs. 694-695, 9, from 4.7 to 6.0 mm S. Bonnet, 1957, Bibliographia Araneoruni, 2: length long, carapace 2872. 1.7 to 2.2 mm long, 1.4 to 1.9 mm wide. Micrathcna reduviancr. —Com.stock, 1912, Spider Males vary in total length from 3.0 to 3.7 $. Comstock, 1940, Book, p. 517, fig. 563, mm, carapace 1.5 to 1.8 mm long, 1.1 to Spider Book, rev. ed., p. 530, fig. 563, 5 . Not 1.2 mm wide. The largest female came Plcctana redtiviana Walckenaer, 1841 {= M. from Mexico. gracilis ) . Diagnosis. Unlike other species north of Female from Virginia: Description. Mexico M. mitrata has three pairs of brown with white thoracic rim. Carapace the 31 . The dimples on carapace ( Fig. ) Sternum dark brown. brown, Legs shghtly female differs from M. funebris in the ab- of abdomen hghter than carapace. Dorsum sence of the anterodorsal tubercle on the white with black marks 31). Sides (Fig. abdomen (Fig. 30) and the presence of white with black marks Venter (Fig. 30). openings of the epigynum in a depression black. There is a thoracic depression and (Figs. 34, 35). The male palpus has a three of dimples on each side of thorax pairs smaller, differently shaped, smooth para- 31). Abdomen with four short pos- (Fig. cymbium (Fig. 38), and a median apophy- terior spines (Figs. 30, 31). Total length sis folded back on itself, its tip sclerotized 5.0 mm, carapace 1.7 mm long, 1.4 mm (Figs. 39, M in 40). wide. First femur, 1.7 mm; patella and Natural History. Micrathena mitrata is tibia, 1.7 metatarsus, 1.2 mm; tarsus, mm; found in deciduous forest, woodland, under 0.5 mm. Second and tibia, 1.4 mm; patella trees, sometimes in shrubs and usually in third, 0.9 mm. Fourth femur, 1.7 mm; the shade. and 1.6 1.1 patella tibia, mm; third, mm; Distribution. From Maine to Wisconsin fourth 0.4 mm. and Kansas, south to Mexico and Panama, Male: carapace brown, posterior median but absent from the Florida peninsula eyes on black spots. Sternum black. Legs (Map 2). brown. Dorsum black with paired white pigment spots. Venter black. Carapace (Walckenaer) with three pairs of dimples (Fig. 2S). First Figures 41-54, Map 2 coxa with a veiy small hook. Abdomen rectangular in dorsal outline (Fig. 28). Plcctana sagittata Walckenaer, 1841, Histoire Naturelle des 2: 174. The Total length 3.5 mm, carapace 1.4 mm long, Insectes, Apteres, name was applied to Abbot illustration of 1.2 mm wide. First femur, 1.3 mm; patella Georgia Spiders, p. 8, fig. 50. Photocopy of the and tibia, 1.2 mm; metatarsus, 0.9 mm; Abbot manuscript in the Museum of Compara- tarsus, 0.5 mm. Second patella, and tibia, tive Zoology, examined. Hentz, 1850, Boston Soc. Natur. 1.0 mm; third, 0.7 mm. Fourth femur, 1.4 Epeira spinea J. Hist, 6: 21, pi. 3, fig. 9, 9. Syntypes from and tibia, 1.0 mm; patella mm; metatarsus, Atlantic states in the Boston Society of Natural 0.8 mm; tarsus, 0.5 mm. History, destroyed.

Figures 41-54. Micrathena sagittata (Walckenaer). 41-42, Male. 41. Dorsal. 42. Lateral. 43. Subadult male. 44-51. Female. 44. Lateral. 45. Dorsal. 46. Posterior median eyes. 47. Left lateral eyes. 48-51. Epigy- num. 48. Ventral. 49. Posterior. 50. Lateral. 51. Posterior, cleared. 52-54. Male left palpus. 52. Lateral. 53. Mesal. 54. Mesal, expanded.

Scale lines. 0.1 mm; except Figures 41-45, 1.0 mm.

Abbreviations. C, conductor; E, embolus; M, median apophysis; R, radix; T, tegulum. COLPHEPEIBA, MlCRATHEXA AND GaSTERACANTHA NoRTH OF MeXICO * Lcvi 431 432 Bulletin Museum of Comparative Zoology, Vol. 148, No. 9

AcTOSoma bovimim Thorell, 1859, Oefv. Svensk Posterior median eyes on black spots. Ster- Vet. Ak. 16: 9. Female Forh., 301, holotvpe num, legs brown. Dorsum of abdomen in from Alabama, lost ( not Natural History lilack, white on lateral margin and posterior Museum, Stockholm). white marks. Sides black, venter black and Acrosoma spinea: —Emerton, 1884, Trans. Con- 1.2 diameters necticut Acad. Sci., 6: 326, pi. 38, figs. 5-8, $, brown. Posterior median eyes to $ ; 1902, Common Spiders, p. 190, figs. 437, of anterior medians. Laterals subequal 440-442, 9, $, web. anterior median eyes. Abdomen trapezoi- Acrosoma sagittatum: —McCook, 1893, American dal, dorsoventrally flattened (Fig. 41). Spiders, 3: 214, pi. 21, figs. 8, 9, $, $. Total 4.7 1.9 mm Micrathena sagittata: —F. P. -Cambridge, 1904, length mm, carapace long, Biologia Centrali-Americana, Araneidea, 2: 536, 1.2 mm wide. First femur, 1.6 mm; patella pi. 51, figs. 20, 21, $, i. Comstock, 1912, and tibia, 1.5 mm; metatarsus, 1.0 mm; web. Spider Book, p. 514. figs. 189, 558-561, 9 , tarsus, 0.6 mm. Second patella and tibia, Reimoser, 1917, Verhandl. Zool. Bot. Gesell. 1.2 mm; third, 0.8 mm. Fourth femur, 1.8 Wien, 67: 140, pi. 9, fig. 29, 9. Fetrunkevitch, 1.4 1930, Trans. Connecticut Acad. Sci., 30: 259, mm; patella and tibia, mm; metatarsus, figs. 111-114, 9, i. Comstock, 1940, Spider 1.0 mm; tarsus, 0.6 mm. Book, rev. ed., p. 527, figs. 189, 558-561, 9, Variation. The abdomen of the female web. Roewer, 1942, Katalog der Araneae, 1: 967. be white to golden orange in color. Kaston, 1948, Connecticut Geol. Natur. Hist. may The abdominal of Surv'. 70: 219, figs. 690-693, 2028, 9,5, web. posterior spines speci- Bonnet, 1957, Bibliogruphia Araneonmi, 2: mens from southern Florida are longer than 2876. those from more northern areas. Interest- Micrathena comstocki Archer, 1951, Amer. Mus. ingly some Mexican specimens have minute Novitates, no. 1487: 10, figs. 15-17, 9. Female like related holotype from Royal Palm State Park [Royal posteroventral spines tropical Palm Area, Everglades National Park], Dade species. Females vary in total length 5.4 to Count\', Florida in the American Museum of 8.6 mm, carapace 2.9 to 3.5 mm long, 2.2 to Natural examined. NEW SYNONYMY. History, 2.7 mm wide. Males vary in total length 4.2 Micrathena sagittata emertoni Archer, 1951, Amer. to 5.9 mm, carapace 2.0 to 2.5 mm long, 1.3 Mus. Novitates, 1487: 10, figs. 18, 22, 9. to 1.6 mm wide. Female holotype from Norvvell, Plymouth Co., Massachusetts, in the American Museum of Diagnosis. Micrathena sagittata females Natural History. NEW SYNONYMY. are recognized by having three pairs of spines with the posterodorsals the largest Description. Female from Virginia: 44, and there are no carapace brown, darker on sides of thorax. (Figs. 45), postero- ventral spines in specimens north of Mexico Sternum, legs brown. Dorsum of abdomen as there are in some related tropical species. white to yellow with black sclcrotized The openings of the epigynum are in de- disks; black anteriorly above carapace and pressions on the posterior face of a bulge posterior spines black (Fig. 45). Sides (Figs. 4.8-51). The male, unlike other black with white patches. Venter black species of the area, has a trapezoidal ab- around spinnerets, with paired white domen, widest posteriorly (Fig. 41) and patches. Abdomen with three pairs of a distinct, biforked median apophysis ( Figs. the ones 44, spines, pcjsterior largest (Figs. 53, M in 54). The paracymbium, unlike Total from between the 45). length pos- that of other North American species, is terior 8.0 mm, 3.1 mm 52 with a .spines carapace recurved, pointing back ( Fig. ) wide. First long, 2.5 mm femur, 3.3 mm; spur on its side. Juveniles also have a patella and tibia, 3.0 mm; metatarsus, 1.9 triangular abdomen (Fig. 43). mm; tarsus, 0.9 mm. Second patella and Natural History. This species is found on tibia, 2.8 mm; third, 1.6 mm. Fourth shrubs in deciduous forest and woods. femur, 3.7 mm; patella and tibia, 3.0 mm; Distribution. From southern New Hamp- metatarsus, 2.1 mm; tarsus, 0.9 mm. shire to Minnesota, Nebraska, south to Male from Virginia: carapace brown. Costa Rica. COLPHEPEIRA, MiCRATHENA AxXD GaSTERACANTHA NoRTH OF MEXICO • Lcvi 433

Micrathena gracilis (Walckenaer) depression small, round (Fig. 59). Dorsvmi of with three of and Plate 1; Figures 55-68; Map 2 abdomen pairs spines two pairs of posteriorly directed spines gracilis Walckenaer, 1805, Tableau des Epcira (Figs. 58, 59). Total length 8.5 mm, cara- Aianeides, p. 65. "An unpublished species from 3.0 mm long. 2.2 mm wide. First Carolina, communicated by M. Bosc." pace 2.3 and 2.2 Plectana 'gracilis Walckenaer, 1841, Histoire Natur- femur, mm; patella tibia, mm; elle des Insectes, Apteres, 2: 193. The name metatarsus, 1.4 mm; tarsus, 0.9 mm. Sec- is to the Abbot illustration of the applied Spiders ond patella and tibia, 2.0 mm; third, 1.4 of Georgia nos. 47, 48. Photocopy of the Mu- mm. Fourth femur, 2.7 mm; patella and seum of Comparative Zoology, examined. 1 2.2 1.5 tarsus, consider this the date of the name. tibia, mm; metatarsus, mm; Pleciana rcduviana Walckenaer, 1841, Histoire 0.8 mm. Naturelle des Insectes, Apteres, 2: 201. Name Male: carapace brown, thoracic region for Abbot illustration of no. 49. Spiders Georgia darker. Legs brown. Dorsum of abdomen Photocopy in Museum of Comparative Zoology, venter blackish. There is a round, examined. whitish, Acrosoma matronale C. L. Koch, 1845, Die Arach- circular thoracic depression (Fig. 55). 1.4 niden, 11: 68, fig. 887. Female from Mexico, Total length 4.8 mm, carapace mm long, lost (not in Berlin Museum). 0.9 mm wide. First femur, 0.8 mm; patella Epeira rugosa Hentz, 1850, J. Boston Natur. Hist. and tibia, 0.9 mm; metatarsus, 0.6 mm; Soc, 6: 21, pi. 3, fig. 10. Type from southern tarsus, 0.4 mm. Second and tibia, states in the Boston Natural History Society, patella 0.6 Fourth 1.2 destroyed. 0.9 mm; third, mm. femur, Acrosoma rugosa: —Emerton, 1884, Trans. Con- mm; patella and tibia, 0.8 mm; metatarsus, Acad. 6: 9; necticut Sci., 326, pi. 38, fig. 10, 0.5 mm; tarsus, 0.4 mm. 1902, Common Spiders, p. 189, fig. 439, ? . Variation. The species is quite variable Acrosoma gracile: —McCook, 1893, American in color. Females in total 7.0 Spiders, 3: 212, pi. 21, figs. 1-4, 9,6- vary length Micrathena 77iatronalis: —Simon, 1895, Histoire to 10.8 mm, carapace 2.6 to 3.7 mm long, Naturelle des Araignees 1: 852, fig. 902, 9. 1.7 to 2.5 mm wide. Males vary in total Micrathena gracilis: —F. P. -Cambridge, 1904, Bi- length 4.2 to 5.1 mm, carapace 1.3 to 1.6 ologia Centrali- Americana, Araneidea, 2: 528, mm long, 0.9 to 1.0 mm wide. pi. 50, fig. 3, pi. 51, fig. 16, 9, 6. Comstock, The female can be 1912, Spider Book, p. 516, fig. 562, 9. Rei- Diagnosis. readily moser, 1917, Verhandl. Zool. Bot. Ges. Wien, 67: recognized by the often gray abdomen with 87, pi. 1, fig. 1, 9 . Comstock, 1940, Spider Book, ten and the spines ( Figs. 58, 59) by laterally rev. ed., p. 529, fig. 562, 9 . Roewer, 1942, Kat- flattened tip of the cone of the epigynum alog der Araneae, 1: 966. Kaston, 1948, Bull. The male as well as Connecticut Geol. Natur. Hist. Surv., 70: 219, pi. (Figs. 62-65). juvenile than 33, figs. 688, 689, 9,6- Bonnet, 1957, Biblio- males have an elongate abdomen more graphia Araneorum, 2: 2868. Chickering, 1961, three times as long as wide (Figs. 55, 56), Bull. Mus. Comp. Zool., 125: 421, figs. 72-77, the palpus has a round hookshaped para- 9, S. (Fig. 66), a complex conductor Micrathena nigrior Chamberlin and Ivie, 1936, cymbium is C difficult to make out ( the basal lobe Bull. Univ. Utah, biol. ser. 3(5): 58, figs. 134- ( ) a . from Barro Colo- and 135, 9 Four female syntypes probably the paramedian apophysis ) in the Amer- rado Island, Panama Canal Zone, minute median apophysis with a filamen- ican Museum of Natural History, examined. tous attachment (Figs. 67, M in 68). Micrathena is Description. Female from Virginia: Natural History. gracilis the web in shaded carapace brown, darker on sides and middle found in dense woods, of thorax. Sternum maculated white and areas, often on bushes. A study of the web brown. Legs brown. Dorsum of abdomen was pubhshed by B. E. Dugdale (1969); had 44 radii and about as whitish with dark spots and dark brown the orb observed orb had a radius of 17 sclerotized spots and dark brown spines many spirals. The (Fig. 59). Sides brown with white spots cm. is found from and dark brown sclerotized spots. Thoracic Distribution. The species 434 Bulletin Museum of Comparative Zoology, Vol 148, No. 9

eastern Massachusetts, Michigan, Wiscon- is only one transverse row of black discs sin, Nebraska, Texas, Sonora, to Panama, on the anterior of the abdomen dorsum; absent from southern Florida (Map 2). the abdomen has one or two pairs of spines laterally and one pair posteriorly (Figs. Gasteracantha Sundevall 71, 72). There is a sclerotized, central bulge on the venter of the abdomen of the Gcistcracantha 1833, Arach- Sundevall, Conspectus female between genital groove and spin- nidum, p. 14. Gasteracantha cancrifonnis is nerets, not present in Micrathena (Figs. 71, the type species since the only other species Unlike Micrathena, Gasteracantha has name originalK' included is G. hexacantha, a 73). synon\ni of G. cancrifonnis. The name is a large canoe-shaped tapetum in the pos- Mitt. Zool. Mus. feminine. Dahl, F. 1914, terior median eyes (Fig. 75). Together Ann. Mus. Berlin, 7: 235-301. Benoit, 1962, with Micrathena, Gasteracantha differs Royal de I'Afrique Centrale, 8 ser., sci. zool., from other in the spinnerets 112: 1-70. Emerit, 1974, Faune de Madagas- genera having a car, 38: 1-216. on a cone or their base surrounded by Vibradctlus Chamberlin, 1925, Bull. Mus. Comp. sclerotized annulus (Fig. 73). 67: 214. des- Zool., Type species by original Description. Gasteracantha species are ignation and monot>py V. carolinus Chamber- brightly colored. The sclerotized, square lin (= Ga.steraca)itJia cancriformis). carapace is high in the head region and has Note. Sundevall cited Latreille, 1831 as a deep thoracic groove (Figs. 71, 72). The author of the name Gasteracantha. How- eyes are subequal, small, the anterior Ga.steracanthe Latreille 1831 Cours their diameter the ever, ( median eyes apart, pos- d'Entomologie, p. 530) is a nomen nudum terior medians more than their diameter. since no species are included; it is thus an All secondary eyes have a canoe-shaped invalid name since it lacks an indication tapetum (Figs. 75, 76). The laterals on Art. also (ICZN, 16, V). Bonnet (1957) each side are far from the medians ( Fig.

cites Latreille 1831 as author. 74 . of the the erroneously ( ) ) The height clypeus equals The list of of Gasteracantha is of anterior synonymy diameter the median eyes ( Fig. incomplete; a complete list is found in 74). The heavy chelicerae of G. cancri- Emerit, 1974. There are two excellent re- formis have five teeth on the anterior visions for the genus available, one of margin, four on the posterior margin. The African species by Benoit and one of Mada- legs are short and thick, and the tarsi are gascan species by Emerit. Both point out very short. The female abdomen is a sclero- that Gasteracantha .species are generally tized shield with sclerotized lateral spines variable and difficult to separate. But this and dorsal sclerotized discs. had been noted Dahl 1914 are notice previously by ( ) Males minute (Figs. 69, 70, in his world-wide study. different scale) and less often collected. Diagnosis. The female carapace is al- The median eye area is slightly projecting most square in outline (Figs. 71, 72) not (Figs. 69, 70). The male lacks the tooth longer than wide nor rebordered on the on the proximal end of the palpal femur sides as that of Micrathena. It is high in and a tooth on the endite present in many front. Unlike related African genera, there Araneidae. None of the coxae and none of

Figures 55-68. Micrathena gracilis (Walckenaer). 55-56. Male. 55. Dorsal. 56. Lateral. 57-65. Female. 57. Eye region and chelicerae. 58. Lateral. 59. Dorsal. 60. Posterior median eyes. 61. Left lateral eyes. 62-65. Epigynum. 62. Ventral. 63. Posterior. 64. Lateral. 65. Posterior, cleared. 66-68. Male left palpus. 66. Lateral. 67. Mesal. 68. Mesal, expanded.

Scale lines. 0.1 mm; except Figures 55-59, 1.0 mm.

Abbreviations. C, conductor; E, embolus; M, median apophysis; R, radix; T, tegulum. COLPHEPEIBA, MiCRATHEXA AND GaSTERACANTHA NoRTH OF MeXICO • Levi 435 436 Bulletin Museum of Comparative Zoology, Vol. 148, No. 9

the distal articles of the legs are modified. close to the conductor in Wagneriana and Since the abdomen lacks the prominent Wixia (Levi, 1976, figs. 69-71, 98). The spines of the females and also tlie ventral hematodocha-like material (C) behind the pnjtuberance, males are difficult to asso- embolus (E) is believed to be the con- ciate with matching females in other parts ductor, because of similar structures in sim- of the world where there are several species. ilar positions in Acanthepeira, Wagneriana, Genitalia. The epigynum is a heavily Wixia, and Scoloderus. Since Gasteracantha sclerotized projection with a median lobe lacks a sclerotized conductor and also seen lobes the ( Figs. 77-80). The openings can be on additional on embolus (including each side of a septum on the posterior face a terminal apophysis), I believe the palpus (Fig. 79). The internal genitalia (Fig. 81) to be simplified secondarily. are difficult to make out because of heavy Relationship. Gasteracantha is close to sclerotization. Micrathena and also to Mastophora. The The palpus is relatively simple. In mesal structure of the palpus, particularly the view of the contracted palpus three sclerites lateral (rather than proximal) position of are visible: distally the filiform embolus the tegulum (Figs. 83, 84), the mesal posi- (Figs. 83, F in 84); in the center a round tion of all sclerites, and the presence of the sclerite with its distal folded conductor edge and paramedian apophysis ( PM ) and sculptured, the paramedian apophysis suggest close relationship with Acanthe- (PM); and proximally the median apophy- peira, Wagneriana, Wixia, and Scoloderus sis in In 1976 . Further indication of a (Figs. 83, M 84). the expanded ( Levi, ) highly palpus (Fig. 84) the radix (R) becomes specialized araneid are the widely separate completely free and transparent hematocha- eyes, the square carapace (Figs. 71, 72, 74) like material, probably the conductor (C), and the modified structure of the abdomen. appears behind the embolus (E). The Natural History. Gasteracantha biology is embolus lacks the parallel lobe (perhaps better known than that of many other the terminal apophysis) of Micrathena. In araneids (Araneus diadeniatus excepted) the expanded palpus, tlie paramedian thanks to the beautiful researches of M. PM Emerit. His studies apophysis ( ) slips down and behind the many on Madagascan median as result of Gasteracantha versicolor are listed in Emerit apopliysis ( M ) pressure from the soft conductor (C) (Fig. 84). (1974). The Mastophora palpus is similar but lacks Species. Gasteracantha is a cosmotropical a paramedian apophysis (Levi, in press). genus. As far as we know, there are only The palpal patella lacks strong setae. The one or perhaps two species in America, paramedian apophysis (PM) was called both known to Linnaeus 200 years ago; G. terminal Emerit tetracantha in apophysis by ( 1968a, ( Linnaeus ) the West Indies 1974). This sclerite is in the same position and G. cancriformis (Linnaeus) found and of similar appearance as the para- from the southern United States to Argen- median apophysis of Acanthepeira and tina. other genera (Levi, 1976; in press). The The literature indicates two species in Acanthepeira paramedian apophysis is the Americas, both originally described by doubtless the same structure as that of the Linnaeus from Jamaica: Gasteracantha complex palpus of Eriophora (Levi, 1970) cancriformis and G. tetracantha. Accord- which was studied by Comstock (1910). ing to the literature, G. tetracantha occurs The origin of this structure may perhaps from California and Arizona to the Greater be seen in the Verrucosa palpus (Levi, Antilles. The California and Arizona rec- 1976, figs. 8, 9) in which it appears to be ords come from specimens of the George the basal end of the conductor. It is also Marx collection, well-known for erroneous COLPHEPEIRA, MiCRATHENA AND GaSTERACANTHA NoRTH OF MeXICO • Levi 437

Gasteracantha cancriformis

Map 3. North American distribution of Gasteracantha cancriformis (Linnaeus). records. None were found in recent collec- Gasteracantha cancriformis (Linnaeus) tions and the species probably does not Plate 2; Figures 69-84; Map 3 occur in the Southwest. (There is a speci- Araiiea cancriformis Linnaeus, 1767, Systema men in the N. Banks collection from "Cal." Naturae, 12 ed., p. 1037. Specimens described which also with Marx. probably originated ) from Jamaica, probably lost. Although large collections were available A. hexacantha Fabricius, 1787, Mantissa Insec- 1: 344. Name with one line of from Jamaica, only one species, G. cancri- torum, given description, but no locality. formis, is found and the Linnaeus record Gasteracantha velitaris C. L. Koch, 1838, Die also be a locality error. Gasteracantha may Arachniden, 4: 33, pi. 269, $. Female from tetracantha occurs in Puerto Rico, the Brazil. Virgin Islands, and the Bahamas to the Plcctana elipsoides Walckenaer, 1841, Histoire Natiirelle des 2: 155. Name north, exactly those areas where G. cancri- Insectes, Apteres, given to fig. 118, p. 13 of Abbot, Drawings of formis is absent. specimens appear Many the Insects of Georgia in America, photocopy to be intergrades having six spines and examined. only a few have completely lost the anterior Plcctana quinqueserrata Walckenaer. 1841, Histoire des 2: 157. Female pair. The intergrades come from the north Naturelle Insectes, Apteres, from in Walckenaer's lost. and the Bahamas, not from the west. I Guyana collection, Plcctana sexserrata Walckenaer, 1841, Histoire to obtain more from the hope specimens Naturelle des Insectes, Apteres, 2: 157. Female region to determine whether there are one from Cayenne. or two species in the West Indies. Plcctana atlantica Walckenaer, 1841, Histoire des 2: 167. Female Numerous names have been given to Naturelle Insectes, Apteres, from St. Domingo. populations of G. cancriformis but as far as Gasteracantha ruhiginosa C. L. Koch, 1845, Die I know there are never two different pop- Arachniden, 11: 55, pi. 878. Female from St. ulations overlapping except perhaps in the Domingo, West Indies. West Indies. The niche of the numerous Epeira cancer Hentz, 1850, J. Boston Natur. Hist. 6: $. Females from African Gasteracantha seems occu- Soc, 23, pi. 3, fig. 13, species South Garolina and southern Alabama in the in the pied by species of Micrathena Amer- Boston Natural History Society, destroyed. icas. Gasteracantlw, insulana Thorell, 1859, Oefv. Svensk 438 Bulletin Museum of Comparative Zoology, Vol. 148, No. 9

Vet. Akad. Forh., 16: 302. Female from Gala- Gasteracantha elliptica Getaz, 1893, An. Inst. Fis.- pagos Islands in the Natural Histor>- Museum, geogr. nac. Costa Rica, 4: 105, 9. Female Stockholm, examined. specimens from around San Jose, Costa Rica, Gasteracantha columhiae Giebel, 1863, Z. Ge- depository imknown. sammt. Xatiuw., 21: 312. A black individual Gasteracantha hiolleyi Banks, 1905, Proc. Entomol.

Soc. 7 : . Female holo- from Colombia, lost ( not in Halle ( Saale ) with Washington, 20, fig. 3, 9 the Giebel collection). type from Cocos Island in the Museum of Com- Gasteracantha kochii Butler, 1873, Trans. Entomol. parative Zoology, examined. Bull. Soc. London, p. 169. New name for G. Iicxa- \'ihradellus carolinus Chamberlin, 1925, Mus. cantha: —C. L. Koch, 1838, Arachniden, 4, pi. Comp. Zool., 67: 214, S- Male holotype from in the of 117, fig. 268. Female from Para [Belem, Bra- South Carolina Museum Comparative zil]. Zoology, examined. Gasteracantha oldendorffi Holmberg, 1876, An. Female from Florida: cara- Agric. Rep. Argentina, 4: 143. Female from Description. Noter del Rio Guayguiraro, [Entre Rios], Ar- pace, sternum, legs brownish black. Dor- gentina, destroyed. sum of abdomen whitish, spines orange- Gasteracantha caUida O. P. -Cambridge, 1879, Proc. yellow, muscle scars black. Venter black Zool. Soc. London, p. 284, pi. 26, fig. 7, 9. with white spots; spines and ventral scler- Female holotype from Trinidad, West Indies, in otized Total 7.2 the Hope Museum, O.vford University, Oxford, projection, orange. length not examined. mm, carapace 3.2 mm long, 3.0 mm wide. Gasteracantha raimondii Horae Taczanowski, 1879, First femur, 2.5 mm; patella and tibia, 2.6 Soc. Entomol. Rossicae, 15: 106, pi. 1, figs. 25, mm; metatarsus, 1.4 mm; tarsus, 0.8 mm. 26, 5 . Five female syntypes from Lima, Choril- Second and 2.4 third, 1.6 los and Montana de Nancha, Peru, in the Polish patella tibia, mm; .'KcademN- of Sciences, examined. mm; fourth, 2.4 mm. Gasteracantha rainwndii unicolor Taczanowski, Male from Florida: carapace brownish 1879, Horae Soc. Entomol. Rossicae, 15: 107. black; sternum black. Legs light blackish Two females from Lima, Peru. brown. Dorsum of abdomen dark with Gasteracantlia proboscidea Taczanowski, 1879, gray white venter black with ventral Horae Soc. Entomol. Rossicae, 15: 108, pi. 1, spots;

fig. 27, S Two male syntypes from Lima, Peru paired white spots. Total length 2.2 mm, in the Polish of examined. Academy Sciences, carapace 1.1 mm long, 0.9 mm wide. First Gasteracantha rufospinosa Marx, 1883, Entomol. femvu-, O.S mm; patella and tibia, 0.8 mm; Amer., 2: 25, figs, a-f, 9 , S . Female and male metatarsus, 0.5 mm; tarsus, 0.3 mm. Second from Crescent City, Florida, lost ( not in Na- tional Museum of Natural History). patella and tibia, 0.7 mm; third, 0.4 mm; GasteracantJia maura McCook, 1893, American fourth, 0.7 mm. Spiders, 3: 210, pi. 13, fig. 12, $. Numerous Variation. There are vast differences in syntypes, "young and old from California, par- color and 85 . and shape ( Fig. ) Hispaniola ticularly the southern part . . . and from the islands off lost in have an the coast," ( not Academy of Jamaican specimens may orange

Natural . all Florida Sciences, Philadelphia ) carapace and legs. Almost Gasteracantha — cancrifonnis: McCook, 1893, specimens have orange spines. While black American Spiders, 3: 211, pi. 14, fig. 9, $. specimens occur occasionally throughout F. P. -Cambridge, 1904, Biologia Centrali-Ameri- the all from Mona Island cana, Araneidea, 2: 525, pi. 51, fig. 14, 9. range, specimens Petrunkevitch 1930. Trans. Connecticut Acad. (west of Puerto Rico) were black with light Sci., 30: 249, figs. 103, 9, S. Comstock, 1940, dorsal patches. Texas specimens were Spider Book, rev. ed., p. 526, fig. 556, 557, 9, bright yellow in color (washed out in al- web. Roewer, 1942, Katalog der Araneae, 1: The narrowest bodies were found 949. Bonnet, 1957, Bibliographia Araneorum, 2: cohol). 1945. on Mona Island and Jamaica, the widest

Figures 69-84. Gasteracantha cancriformis (Linnaeus). 69, 70. Male. 69. Dorsal. 70. Lateral. 71-81. Fe- male. 71. Lateral. 72. Dorsal. 73. Abdomen, ventral. 74. Eye region and chelicerae. 77-81. Epigynum. 77. Anteroventral. 78. Ventral. 79. Posterior. 80. Lateral. 81. Posterior, cleared. 82-84. Male left palpus. 82. Lateral. 83. Mesal. 84. Mesal, expanded. COLPHEPEIRA, MiCRATHENA AND GaSTERACANTHA NORTIf OF MeXICO • Levi 439

Scale lines. 0.1 mm; Figures 69-74, 1.0 mm.

Abbreviations. C, conductor; E, embolus; M, median apophysis; PM, paramedian apophysis; R, radix; T, tegulum. 440 Bulletin Museum of Comparative Zoology, Vol. 148, No. 9

"\g^*«4^l^

Figure 85. Geographic variation of Gasteracantha. Locality data clockwise: Torrey Pines State Park, San Diego Co., California; Laguna Beach, California; Austin, Texas; College Station, Texas; Hattiesburg, Mississippi; Houston Co., Alabama; Charleston, South Carolina; Gainesville, Florida; New Providence, Bahamas; Naples, Florida; Mona Island; La Romana, Dominican Republic; Momance, Haiti; Los Llanos, Cuba; Pinar del Rio, Cuba; Port Antonio, Jamaica; Mandeville, Jamaica; Barro Colorado Island, Panama Canal Zone; Carmelina, Honduras; Chichen Itza, Yucatan; Atoyac, Jalisco; Most Southern Palm Grove, Cameron Co., Texas; Brownsville, Texas; La Paz, Baja California.

Scale line: 5.0 mm.

from Texas to Central America. The short- Laguna Beach, California illustrated (Fig. est stubbiest are found in Mona 85 has an extra the spines ) assymmetrical plate on Island and California; the longest ones in left anterior. Many specimens throughout Florida and the southeastern states. The the North American range have the median most acute are found in Cuban into two spines speci- posterior plate split plates ( Texas, mens. A characteristic of Hispaniola and South Carolina, and Dominican Republic, southern Florida specimens is that the sec- Fig. 85). ond pair of spines is larger than the first Total length of females is 5.8 to 8.6 mm, pair. Since there are clines of these char- carapace 2.3 to 3.1 mm wide. Total length acters in various directions, it is not easy to of males 1.9 to 2.7 mm, carapace 0.8 to 1.0 segregate subspecies except for island pop- mm wide. Size variation is about the same ulations. throughout the southern states. Unusual variations ai^e the additional Species problems. Archer judging by round plates found at times on the dorsum museum labels, considered specimens from of the abdomen. The specimen from the western states to be G. serviUei (Guerin) CoLPHEPEiRA, MicRATHENA AND Gasteracantha North OF Mexico • Levi 441

and those from the eastern states to be G. completes the Hfe cycle in a year. Females mature in late fall or winter and are cancriformis; with several subspecies. How- early ever collections from Austin, Texas had found from October to January. Adult males in October and November both specimens which Archer considered to first appear females are one-third to one-half belong to the western species and to the when adult size. Adult males from a eastern species. Since there is no overlap hang single of the two forms, but instead intermediates strand of silk adjacent to a female's web, are found, I consider all to belong to one one to three per web. The tiny males ac- were con- to species. The eastern specimens complish sperm induction just prior sidered by Archer (unpublished) to belong courtship. Egg-sacs are flattened ovate almost to several different subspecies, but masses of tangled white, yellow, and green labeled as intermediates as many were (e.g. silk, marked with a longitudinal stripe of from northern as to Florida), belonging dark green silk. They are found on the the subspecies. underside of leaves adjacent to the webs. Females can be distinguished Diagnosis. Distribution. Gasteracantha cancriformis from West Indian G. tetracantha (Linnaeus) is found from North Carolina to Southern by the presence of six spines on the ab- California south to northern Argentina domen (Figs. 71, 72); G. tetracantha has

. trans- 3 ) occasionally get only four. Males of G. tetracantha have the ( Map Specimens one female was found fruit sclerites of the palpus, especially the para- ported; among median apophysis, relatively smaller than in Seattle. those of G. cancriformis (Figs. 83, 84), al- though all parts are of the same shape. LITERATURE CITED Natural History. The striking appear- Bexoit, p. L. G. 1962. Monographic des ance, webs and diurnal habits — Africains Ara- conspicuous Araneidae Gasteracanthinae ( make this one of the easily collected tropical neae). Ann. Mus. Roy. de TAfrique Centrale, ser. no. 112: 1-70. spiders. The web is found between branches Sci. ZooL, 8, CoMSTOCK, J. 1910. The palpi of male spiders. on shrubs and even on buildings (Plate 2). Ann. Entomol. Soc. America, 3: 161-185. It is made in the and is usually morning Dahl, F. 1914. Die Gasteracanthen des Berliner inclined at an sometimes near vertical angle, Zoologischen Museums und deren geograph-

. threads are decorated Mitt. Zool. Mus. 7: ( Plate 2 ) The outer ische Verbreitung. Berlin, with flocculent tufts of silk (Marples, 1969) 235-301. Dugdale, B. E. 1969. The weaving of an en- and the spider rests in the open hub. Young masterpiece, a spider's orb-web. Gasteracantha versicolor gineering Madagascan may Natur. Hist. 78: 36-41. a have stabilimentum (Emerit, 1968b). Emerit, M. 1968a. Le comportement sexuel Adult males have been collected in d'une araignee Argiopidae malgache a grand Florida from November to February; in dimorphisme sexuel: Gasteracantha versi- color Walck. (Araneidae). Bull. Mus. Nat. Alabama in August; in Texas in April, June, d'Hist. Natur., 2 ser., 39: 898-907. and October and in California July, during de la bio- . 1968b. Contribution a I'etude Adult females can be collected July. logic et du developpement de I'araignee throughout the year in Florida and Texas. tropicale Gasteracantha versicolor (Argiopi- can- dae). Bull. Soc. Zool. France, 93: 49-68. Muma ( 1971 ) found Gasteracantha 1974. Araneidae Gasteracanthinae. criformis webs in central Florida orchards Faune de Madagascar, no. 38: 209. in between trees and also in mixed trees, Homann, H. 1950. Die Nebenaugen der Ara- less than mesophytic woods at a height of neen. Zool. Jahrb. Abt. Anat. und Ontog., 1 to more than 6 m. The female's web has 72: 289-364. Araneae. Ana- . 1971. Die Augen der 10 to 30 spiral lines, the viscid area span- tomic, Ontogenie und Bedeutung fiir die 30 to 60 cm diameter. The prey caught Z. ning Systematik ( , Arachnida). 201-272. are flies, moths, and beetles. The spider Morphol. Tiere, 69: 442 Bulletin Museum of Comparative Zoology, Vol. 148, No. 9

Levi, H. W. 1970. The Ravilla Group of the cosa, Acanthcpcira, Wagneriana, Acacesia, Oib\vea\er Ceniis Eriophora in Xorth .\merica Wixia, Scolodcnis and Alpaida north of Mex- (Araneae: Araneidae). Psyche, 77: 280-302. ico (Araneae: Araneidae). Bull. Mus. Comp.

. 1972. The orb-weaver genera Singa ZooL, 147(8): 351-391.

and Hypsosinga in America (Araneae: Aranei- . in press. Orb-webs and ph\logeny of dae). Pfitjche, 78: 229-256. orb-weavers. Symp. Zool. Soc. London.

. 1975. The American orjj-weaxer genera MuMA, M. 1971. Biological and behavioral notes Larinia, Cercidia and Mangora north of Mex- on Gastcracantha cancrifonnis (Arachnida, ico (Araneae, Araneidae). Bull. Mus. Comp. Araneidae). Florida Entomol., 54: 345-351. Zooh, 147: 99-1.35. M.\RPLEs, B. J- 1969. Observations on decorated

webs. Bull. Brit. Arachnol. 1 : 13-18. . 1976. The orb-\\ea\er genera Vcrru- Soc,

INDEX

\'alid names are printed in italics. Page numbers insulana, Gasteracantha, 437 refer to main references, starred page numbers to kochii, Gasteracantha, 438 illustrations. maculata, Acrosoma, 426 maculata, Micrathena, 428 matronale, Acrosoma, 433 atlantica, Plectana, 437 matronalis, Micrathena, 433 biolleyi, Gasteracantha, 438 maura, Gasteracantha, 438 bovinum, Acrosoma, 432 Micrathena. 424 callida, Gasteracantha, 438 mitrata, Acrosoma, 428 cancer, Epeira, 437 mitrata, Epeira, 428 cancriformis, Aranea, 437 mitrata, Micrathena, 428, 429* cancrifonnis, Gasteracantha, 437, 439*, 440* nigrior, Micrathena, 433 carolinus, Vibradellus, 438 oldendorffi, Gasteracantha, 438 catawba, Aranea, 422 proboscidea, Gasteracantha, 438 catawba, Araneus, 422 quinqueserrata, Plectana, 437 catawba, Colphepeira, 422, 423* raimondii, Gasteracantha, 438 catawba, Epeira, 422 reduviana, Micrathena, 430 Colphepeira, 420 redu\'iana, Plectana, 433 columbiae, Gasteracantha, 438 redu\ianum, Acrosoma, 428 comstocki, Micrathena, 4.32 rubiginosa, Gasteracantha, 437 elipsoides, Plectana, 437 rufospinosa, Gasteracantha, 438 elliptica, Gasteracantlia, 438 rugosa, Acrosoma, 433 emertoni, Micrathena sagittata, 432 rugosa, Epeira, 433 funebre, Acrosoma, 426 sagittata, Micrathena, 430, 431* funehris, Micrathena, 426, 427* sagittata, Plectana, 430 Gasteracantha, 434 sagitattum, Acrosoma, 432 gracile, Acrosoma, 433 sexserrata, Plectana, 437 gracilis, Epeira, 433 spinea, Acrosoma, 432 435* gracilis, Micrathena, 433, spinea, Epeira, 430 433 gracilis, Plectana, unicolor, Gasteracantha raimondii, 438 426 granvilata, Micrathena, \elitaris, Gasteracantha, 437 hexacantha, Aranea, 437 Vibradellus, 434