Herbert W. Levi.
More American Spiders of the Genus Chrysso (Araneae, Theridiidae).
Psyche 69:209-237, 1962.

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MORE AMERICAN SPIDERS OF THE GENUS
CHRYSSO (ARANEAE, THERIDIIDAE)*
BY HERBERT W. LEVI
Museum of Comparative Zoology, Harvard University In a previous paper (Levi, 1955) I revised the genus Chrysso, which I thought at the time to be a compact group of closely related species quite distinct from Theridion, and all coming from America. How- ever, soon after publishing a description of the genus Arctachaea for only two species (Levi, 1g57), I discovered species intermediate between the new genus and Chrysso.
Though species bridging genera have been found in various groups of theridiids, it was a surprise to see the description of Argyria venusta Yaginuma, 1957 (Figs. 3-5) from Japan, a species close to Chrysso and having similar habits, but less specialized in genit a 1' ic structure. Soon several other species were examined that are best placed in Chrysso unless we want numerous monotypic genera. One of these species is Argyrodes nigra 0. P.-Cambridge (Figs. I, 2) from Ceylon (with genitalia superficially resembling those of Theridion punctospar- sum Emerton) though it has the palpus still less specialized than does Chrysso venusta (Yaginuma), Allied to Chrysso nigra (0. P.-Cam- bridge) is the American Chrysio nigro^ternum Keyserling (Fig. 13). Several of the species now placed in Chrysso have the cymbium modi- fied [e.g. Chrysso perplexa (Keyserling), Fig. 181, and bridge the gap to the two species previously placed in Arctachaea. Further, a male of Chrysso cambridgei ( Petrunkevitch) (Figs. 43-47) bridges the gap between the latter gioups and the species tentatively placed in Meotipa. Meotipa is a genus of doubtful validity; the male of the type species remains unknown. The three species placed in Meotipa differed from Chrysso mainly by their longer legs and by a somewhat different shape of abdomen.
There is still some doubt that the groups considered in this paper are monophyletic. Certainly all are related to Theridion, from which they can be separated by the modified shape of the abdomen and by the more specialized palpus. The specialization of the palpus, how- ever, may go in different directions. Most species resemble the 7". frondeum group. A distally truncate cymbium, otherwise rare in theridiids, is common in Chrjls~o and probably indicates relationship. Another unusual character, grooves on the sides of the abdomen, is found in some of the species here considered to belong to Chrysso and *Manuscript received by the editor April 18,1962.



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210 Psyche [December
may be further evidence of common origin. In addition, several
species have teeth on the posterior margin of the chelicerae. I would like to thank Dr. W. J. Gertsch for the loan of collections from the American Museum of Natural History (AMNH) ; Dr. A. M. Chickering for large collections from Panama and Jamaica now in the Museum of Comparative Zoology ; for additional collections Mrs. D. L. Frizzell (Dr. H. Exline) of Rolla, Missouri; Mrs. Friz- zell and Dr. E. S. Ross for specimens from the California Academy of Sciences (CAS) ; Dr. 0. Kraus of the Senckenberg Museum, Frankfurt (SMF) ; and Dr. R. V. Chamberlin for a specimen of the University of Utah collection
(UU) ; Dr. E. Dsesco, Paris, for
making drawings of Meotipa species; Prof. M. Vachon of the Museum National d7Histoire Naturelle, Paris (MNHN) ; Prof. G. C. Varley of the Hope Department of Entomology, Oxford, and Dr. G. 0. Evans, Mr. E. Browning, and Mr. I<. Hyatt of the British Museum, Natural History for their hospitality while visiting their institutions. A. Riedel and J. Pr6szyiiski of the Polish Academy of Sciences, War- saw (PAS) ; Dr. L. Brundin, Riksmuseum, Stockholm, Mr. 7'. Yaginuma, Osaka, and Dr. J. V. Scorza, Caracas loaned valuable specimens. A National Science Foundation grant (G-4317) made possible a trip to Europe to examine types and the National Institute of Health grant (E-1944) facilitated completion of the study and publication of the manuscript.
Chrysso 0. P.-Cambridge
Chrysso 0. P.-Cambridge, 1882, Proc. 2001. Soc. London, p. 429. Type species by original designation Chrysso albomaculata 0. P.-Cambridge. Meotipa Simon, 1892, Histoire Naturelles des Araignkes, 1: 519. Type species by original designation Meotipa picturata Simon. Physcoa Thorell, 1895, Descriptive Catalog of the Spiders of Burma, p. 83. Type species by original designation Physroa sc'intillans Thorell. Argyria Yaginuma, 1957, Acta Arachnologica, 15: 11. Type species by origin- al designation Argyria wenusta Yaginuma. Homonym of Argyr.a Hueb- ner, 1818, and Argyria Robinean-Desvoidy, 1863. Argyroaster Yaginuma, 1958, Acta Arachnologica, 15: 37. New name for Argyria Yaginuma preoccupied.
Arctachaea Levi, 1958, Psyche, 64: 102. Type species by original designation Arctachaea pelyx Levi.
Diagnosis. Chrysso differs from Theridion by a hump or posterior dorsal extension of the abdomen ; from Achasaranea by having a much more complicated palpus containing a radix; from Argyrodes and Spintharus by lacking a colulus (Levi and Levi, 1962). Comments. The types of C/zrysso nigriceps Keyserling and Helvibis sdcata Keyserling have been examined since publishing my paper in



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19621 Levi - Chrysso 21 I
1957. The specimens given those names were correctly identified. The following species have recently been misplaced in Chrysso: Chrysso maronica Caporiacco, 1954 (Comm. Pontificia Acad. Sci. 16: 74) is ldchaearanea hirta (Taczanowski) . NEW SYNONYMY. Cf pentagonu Caporiacco, 1954, ibid., 16: 75 belongs to Achaearanea. While many species of Chrysso are strikingly colored, the coloration usually cannot be used for determination or as a key character. As in most thesidiids, it is variable, e.g. Chrysso compressa (Keyserling) has a striking black sternum and abdomen with a black venter in Brazil, but the venter is yellow in Peruvian specimens. Uyemura (1957) has observed rapid color change in Chrysso venusta (Yaginuma) when disturbed. Probably other species of Chrysso can also change color. Species of Chrysso are absent from the United States and Canada except C. albomaculata and C. clementinae (Map) found in the south- eastern states and the two species previously placed in Arctachaea found from Wyoming to Alaska.
None are known
from Chile or
southern Argentina. Most species are found in the hotter parts of America, where the spider fauna is least known, and our knowledge of the species and their distribution is fragmentary. Key to the American species of Chrysso
Mesal tip of male cymbium with teeth ( 1957, figs. I 1-13), fe- ........................
male chelicerae toothless ; Wyoming to Alaska 2
Cvmbium without teeth; female chelicerae usually with teeth; ..........................
southeastern United States to South America 3
Mesal corner of cymbium projecting (1957, figs. 12, 13) ; female ................................ abdomen higher than long C. pelyx (Levi)
Mesal corner of cymbium rounded ( 1957, fig. I I ) ; female abdomen longer than high .... C. nordica (Chamberlin and Ivie) Males .......................................................................................... 4 Females .................................................................................... 25 Thread-shaped portion of embolus originating in center of bulb, looping towards outside and base before continuing straight .......................................
towards distal end ( I 955, figs. 1-5)
5
Embolus otherwise .................................................................. I I Width of area enclosed by embolus loop almost equals width of basal portion of embolus ( 1955, fig. I 5) ; Hugnuco, Peru ........ .............................................................................. C. mariae Levi Width of area enclosed by embolus loop at least twice width of basal portion of embolus thread (1955, figs. 7, 9, 13) ............ 6 Median apophysis with a large thorn ( 1955, fig. 13) ; Amazon, Peru ........................................ C. sulcata (Keyserling)



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212
6b.
7 a.
7b.
8a.
8 b.
9a.
9b-
I oa.
1 ob.
I 1a.
I ib.
I 2a.
12b.
I 3a.
13b.
I 4a.
14b.
I 5a.
15b.
I 6a.
I 6b.
I 7a.
I 7b.
18a.
18b.
Psyche [December
........................................... Median apophysis without thorn 7
A thorn on ectal side of palpal bulb (1955, figs. 7, g) ........... 8 No thorn on ectal side of bulb .................................................. 9 Radix with evenly rounded swelling; ectal thorn large (1955, ................................................ fig. 7 ) ; Panama
C. vallensis Levi
Radix with round boss ; ectal thorn small ( I 955, fig. 9) ; Pana- ............................
ma, Peru
C. diplosticha Chamberlin and Ivie
Radix with a swelling; conductor almost lanceolate in ventral view ( 1955, fig. I I ) ; coloration as in 1955 fig. 12 ; Panama . . ................................. C. indicifer Chamberlin and Ivie Radix without swelling; conductor hook-shaped (1955, fig. ...................................................... 1-6) ; coloration otherwise 10
Radix with a mesa1 fold, (1955, figs. 1-4) ; abdomen usually ............................................ with spots; North Carolina to Brazil
............................................ C. albomaculata 0. Pi-Cambridge Radix without fold (1955, figs. 5, 6) ; abdomen without dis- .......
Crete spots; Panama to Argentina
C. vexabilis Keyserling
Prominent ventral embolus biforked (Figs. 8, 13) ................ 12 .................................................................. Embolus otherwise I 3
Embolus long, reaching base of bulb (Fig. 13) ; southern Brazil ....................................................... C. nigrosternum Keyserling Embolus short, covering distal two-thirds of bulb (Fig. 8) ; Minas Gerais .................................................. C. gounellei sp. n. Distal end of cymbium truncate ............................................ 14 Distal end of cymbium rounded ............................................ 15 Ventral view as in Fig. 18 ; southern Barzil ................................ -
............................................................... C. perplexa Keyserling Ventral view as in Fig. 28; Venezuela ............ C. simoni sp. n. With long thread-shaped embolus of even diameter (Fig. 23) ; Costa Rica to Trinidad ................................. C. questona sp. n. Embolus otherwise ................................................................. 16 Embolus corkscrew-shaped, on distal end of bulb (Figs. 89, 94) .................................................................................................... I7
.................................................................. Embol~~s otherwise I 8 Embolus larger than remaining portion of bulb (Fig. 89) ; abdomen higher than long; Panama to Ecuador .... C. silva sp. 11. Embolus one-third length of remaining portion of bulb (Fig. 95) ; abdomen longer than high ; Costa Rica, Panama ............. C. volcanensis sp. n.
...................................................................... Embolus with distal portion hook-shaped (Fig. 47) ; Mexico to Venezuela ................................. C. cam bridge! ( Petrunkevitch ) EmboIus otherwise ............................................................... 10



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19621 Levi - chrysso 2 13
~ga. Two transparent sickle-shaped structures 011 venter of bulb ............ (Fig. 42) ; Colombia to Bolivia C. vittatula (Roewer)
........................
igb. Palpal bulb without sickle-shaped structures 20 20a. Conductor prominent and with distal hook (Fig. 74) ; cosmo- ..................................
tropical
C. clementinae ( Petrunkevitch )
20b. Conductor otherwise ............................................................... 2 [ 21a. Embolus a ventral bottle-shaped structure (Fig. 69) ; Peru to ............................................ Brazil C. compressa (I<eyserling)
............................................................. 2 I b. Embolus otherwise 22
22a.
Median apophysis not apparent in ventral view (Fig. 30) ; ............................................... southern Brazil
C. arops sp. n.
..............................
22b. Median apophysis visible in ventral view 23 23a. Median apophysis in distal half of bulb, palpus lightly sclero- ....................................
tized (Fig. 79) ; Panama
C. melba sp. n.
23b. Median apophysis mostly on proximal half of bulb; palpus .................................................................... heavily sclerotized 24
........................
24a. Palpus as in Figure 36, 37 ; Colombia to Bolivia ................................................................... C. ecuadorensis Levi .............................................. 24b. Palpus as in Figure 5 I ; Venezuela ...................................................... C. barromachadoi Caporiacco 25a. Epigynum with openings in coiled borders of fossa (Fig. 78 j ; Costa Rica to Trinidad ................................ C. questona sp. n. 25b. Epigynum otherwise .............................................................. 26 26a. Ducts showing through epigynal wall long and coiled, openings indistinct (Figs. 82, 84) ........................................................ 27 26b. Epigynum otherwise ............................................................... 28 27a. Ducts loosely coiled in large loops (Fig. 83) ; Panama ............... C. alecula sp. n.
................................................................. 27b. Ducts tightly coiled (Fig. 85) ; Trinidad .......... C. arima sp. n. 2%. Epigynum with a central black mark (Fig. 45) ; sternum, venter black ; Mexico to Venezuela ... C. cambridgei ( Petrunkevitch) 28b. E~igynum otherwise; sternum and venter usually not black (except C. conzpressa, C. nigrosternum} ................................ 29 29a. Openings of epigynum widely separated (Fig. 88) ; Panama to Ecuador ............................................................. C. silva sp. :I. 29b. Epigynum otherwise ............................................................. 30 30a. Epigynum with a light, oval swelling, bordered all around (Figs. 34, 35, 40) Colombia to Bolivia ........................................... 31 3ob. E~igynum otherwise ............................................................... 32 3 1a. A dark mark anterior to swelling (Figs. 34, 35) ........................ ..................................................... C. ecdorensis Levi 31b. A dark mark posterior to swelling (Fig. 40) ............................... ................................................................ C. vittatula ( Roewer)



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Psyche
[December
Epigynum with a light swelling bordered anterior and sides ..................
( Fig. 50) ; Venezuela
C. barromachadoi Caporiacco
................................................................ Epigynum otherwise 3 3
Epigynum with openings near ~osterior margin in a depression, ducts loop anterior (Fig. 54) ; southern Brazil ............................ ........................................................................... C. ribeirao sp. n. ................................................................ Epigynum otherwise 34
................................................ Epigynum with a central fossa
3 j
........................................................ Epigynum without fossa 41
Two openings in center of fossa (Fig. 93) ; Costa Rica, Panama ..................................................................... C.volcanensis sp. n. Epigynum otherwise ................................................................ 36 A black circular mark on each lateral end of fossa (Fig. '73) ; ............................
cosmopolitan C. clementinae ( Petrunkevitch) Epigynum otherwise ................................................................ 37 Fossa far anterior, usually bordered on sides only (Fig. 27) ; .......................................................... Venezuela C. simoni sp. n.
Fossa in center or posterior, bordered anterior or posterior as .............................................................................. well as sides 38
Fossa in a dark spot; ducts visible posterior to fossa (Figs. 16, I 7) ; Brazil, Argentina ................ C. rubrovittata (Keyserling) Epigynum otherwise ................................................................ 39 Narrow ducts leading from sides into fossa (Figs. 57, 62) .... 40 Ducts wider and leading from anterior into fossa (Figs. 64, 66, 67), Peru to Brazil ........................ C. coqressa (Keyserling) Fossa bordered on sides and posterior (Fig. 57) ; southern Brazil .................................................. C. pulchra (Keyserling) Fossa bordered on sides and anterior (Fig. 62) ; Minas Gerais .. .......................................................................... C. caraca sp. n. Epigynal openings in a slit anterior to a dark area (Fig. 60) ; ducts simple (Fig. 59) ; San Fernandez Isl. ................................ .......................................................... C. backstromi (Berland) ........
Epigynal openings otherwise ; usually with internal sacs 42
Epigynum a central transverse connected dark mark as in Fig- ure 21 ; Brazil ................................................. C. arztorzio sp. 11. Epigynum otherwise ................................................................ 43 Epigynum with two central dark curved marks (Figs. 10, 12) ; southern Brazil ......................... C. nigrosternum (Keyserling) Epigynum otherwise ................................................................ 44 Epigynum with an anterior dark mark and light posterior as in 1955, fig. 33; head region blackened; Colombia, Ecuador ........ C. nigriceps Keyserling
............................................................... Epigynum otherwise, coloration usually otherwise ................ 45



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Levi - Chrysso 215
Epigynum with two lines, diverging posterior (19 55, fig. 31 ) ; ................
Panama to Peru
C. diplosticha Chamberlin and Ivie
Epigynum otherwise ............................................................... 46 Epigynum with a transverse dark mark ~osterior to two spots ..............................
( I 9 j j, fig. 29) ; Panama
C. vallensis Levi
Epigynum otherwise ................................................................ 4 7 Epigynum with a transverse line anterior to two spots ( 1955. ........................................ fig. 37) ; southern Brazil
C. sicki Levi
Epigynum otherwise .............................................................. 48 Epigynum with a pair of longitudinal lines in light area posterior to transverse dark mark (195 j, fig. 35) ; dorsum of abdomen with two black long lines which fuse at posterior end; Panama ................................................ C. indicifer Chamberlin and Ivie Epigynum otherwise ............................................................... 49 Carapace dark brown; abdomen with a longitudinal dark band ....................................
( 195 j, fig. 20) ; Peru
C. huanuco Levi
............................................................... Coloration otherwise 50
Abdomen without discrete white spots; Panama to Argentina ............................................................... C. vexabilis Keyserling Abdomen usually with white spots; North Carolina to Brazil .......................................... C. albomaculata 0. Pi-Cambridge Chrysso albomaculata 0. P.-Cambridge
Argyrodes elegans Taczanowski, 1872, Horae SOC. Ent. Rossicae, 9: 118, pi. 5, fig. 11, 9. Female syntypes from Uassa, French Guiana [now Rio Uach, Amapa, Brazil] in the Polish Academy of Sciences, Warsaw, examined. NEW SYNONYMY. Not Theridion elegans Blackwall, 1862. Chrysso albomaculata 0. P.-Cambridge, 1882, Proc. Zoo]. Soc. London, p. 429, fig. 6, 9, 8. Male, female syntypes from the Amazon in the Hope Department of Entomology, Oxford, examined. - Levi, 1955, Jour. New York Ent. Soc., 68: 61, figs. 1-4, 18-19, 25-27, 9, 8. Chrysso elegans, -Keyserling, 1884, Die Spinnen Amerikas, Theridiidae 2 (I) : 151.
Theridion elegans, - Simon, 1903, Histoire Naturelle des Araigneks, 2: 990. Theridion emendattun Roewer, 1942, Katalog der Araneae 1 : 429. New name for Argyrodes elegans Taczanowski. NEW SYNONYMY. Distribution. Southeastern United States, Mexico, West Indies, northeastern South America to Bahia, Brazil. Additional records. Lesser Antilles. Grenada: St. George's ( N. I>. H. Krauss, AMNH) . Trinidad: Ilt. St. Benedict (J. G. Myers). Venezuela. Carabobo : San Esteban, I 888 (E. Simon, MNHN) . Grand Elang. Dist. Fed.: hacienda Corosal, N of Mt. Silla, 1888 (E. Simon, MNHN). British Guiana. Georgetown (A. II. Nadlei, AMNH) . Surinam. Paramaribo. Brazil. Ceara : Sierra Corn-



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216 Psyche [December
munaty (E. Gounelle, MNHN ) . Pernambuco : Recife ( SMF ; A. M. Nadler, AMNH) . Bahia: Salvador (A. M. Nadler, AMNH) ; Terra Nova (MNHN).
Chrysso vexabilis Keyserling
Chrysso vexabilis Keyserling, 1884, Die Spinnen Amerikas, Theridiidae, 2 (1) : 155, pi. 7, fig. 96, Q, 3. Syntypes from N. Granada [now Ecuador, Colombia, Panama, Venezuela] in the British Museum, Natural History, examined. - Levi, 1955, Jour. New York Ent. Soc., 63 : 63, figs. 5, 6, 23, 24, Q, 8.
Steatoda rubicunda F. P.-Cambridge, 1902, Biologia Centrali-Americana, Araneidea, 2: 386, pi. 36, fig. 21,
8. Male type from Bugaba, Panama
in the British Museum, Natural History, examined. (Not Theridion rubicunda Nicolet, 1849.) NEW SYNONYMY.
Theridion rubicundulum Roewer, 1942, Katalog der Araneae, 1: 498. New name for Theridion rubicunda (F. P.-Cambridge). NEW SYNONYMY. Examination of Keyserling's type indicated that this species had been correctly determined. The internal female genitalia of Argentine specimens like the ones from Panama, have the posterior portion of the sacs of the internal genitalia ending in narrow extensions (Levi, 1955, fig. 23) quite different from the broadly rounded sacs of C. albomaculata. Unlike C. albomaculata it has no white spots on abdo- men.
Distribution. Panama to Argentina.
Additional records. Venezuela. Dist. Fed. : hacienda Corosal, q, 6 (E. Simon, MNHN). Ecuador. Guayas: Milagso, 9, d (H. E., D. L. Frizzell) ; Rio Chimbo, 6 (Edwards, MNHN). El Oro: Arenillas (E. L. Moore). Brazil. Bahia: Salvador, 9, (E. Goldi, MNHN) ; Rio Salobro, $? (E. Gounelle, MNHN, doubtful det.). Argentina. Cordoba: Cosquin, 9.
Chrysso diplosticha Chamberlin and Ivie
Chrysso diplosticha Chamberlin and Ivie, 1936, Bull. Univ. Utah, biol. ser., 3 (5) : 36, fig. 83,
9. Female type from Barro Colorado Island, Panama Canal Zone in the University of Utah Collection. - Levi, 1955, Jour. New York Ent. Soc., 63: 65, figs. 9, 30, 31, 9, 3. Distribution. Nicaragua, Panama, Trinidad and Dept. Huinuco, Peru.
Additional records. Nicaragua: Musawas, Waspuc River, c? (B. Malkin, AMNH). Lesser Antilles. Trinidad: Simla near Arima, ~7 (A. M. Nadler, AMNH) .
Chrysso indicifer Chamberlin and Ivie
Chrysso indicifer Chamberlin and Ivie, 1936, Bull. Univ. Utah, biol. ser., 3 (5) : 36, figs. 82, 96, 9. Female type from Barro Colorado Island in the



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19621 Levi - Chrysso 217
University of Utah collection. -Levi, 1955, Jour. New York Ent. Soc., 63: 66, figs. 11, 12, 17, 34, 35, 9, 8.
Distribution. Panama to Peru.
Additional records. Venezuela. Dist. Fed. Caracas, (E. Simon, MNHN) . Carabobo: La Cumbre, S. of San Esteban, (E. Simon, MNHN) . Ecuador.
Guayas: Milagro, (H. E., D. L. Frizzell) ; Rio Chimbo, (MNHN). El Oro: Arenillas, (E. L. Moore) ; Pasaje, (R. Walls). Peru. Huinuco: Monzdn Valley, Tingo Marfa, 9, c? (E. I. Schlinger, E. S. Ross, CAS).