A. R. Brady.
The Lynx Spider Genus Oxyopes in Mexico and Central America (Araneae: Oxyopidae).
Psyche 82:189-243, 1975.

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THE LYNX SPIDER GENUS OXYOPES IN MEXICO
AND CENTRAL AMERICA (ARANEAE:OXYOPIDAE)l BY ALLEN R. BRADY
Dept. of Biology, Hope College
Holland, Michigan 49423
INTRODUCTION
This is the second in a projected series of papers on the Neo- tropical Oxyopidae. In this study 16 species of Oxyopes are recorded from Mexico and Central America. Of these, 11 are predominately Mexican and Central American, four are more abundant in America, north of Mexico, and one is widely distributed throughout Mexico, North and 'Central America. Numerous specimens from Mexico and Central America were examined during this study. Collections available from the Museum of Comparative Zoology :and the American Museum of Natural History provided an abundance of specimens from certain areas, such as the Panama Canal Zone. However, some regions of Mexico 'and Central America are poorly collected. Additional specimens came from field studies during several summers in Mexico (1969-71), Costa Rica ( 19721, and Panama ( 1972)
Named species of Oxyopes from Mexico and Central America were often inadequately described and figured and there was some confusion as to the correct names of several species, males and fe- males having been separately described. In addition, little was known of the geographic ranges nor was there any clear concept of the evolutionary relationships of Neotropical Oxyopes to each other or to their North American relatives.
Certain species of Oxyopes lare extremely abundant in areas of grassy or herbaceous vegetation. For example, 0. salticus and 0 . flavus constituted as much as 20% of the total number of spiders taken in sweeping vegetation (0.5-1 m in height) in some coastal areas of Guerrero and Veracruz. Judging from their abundance, these spiders are evidently important predators of insects within these habitats. The species of Oxyopes observed were all active dur- ing the day. They ran swiftly over the plants where they occurred and when disturbed exhibited characteristic erratic leaps which tallow them to be immediately recognized in the field. Although Oxyopes do not actively pursute their prey by sight, they are much ^Manuscript received by the editor March 25, 1975 189




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more active than thomisids, and exhibit a general "searching" be- havior when observed. Long spines on the legs, another field char- acteristic of lynx spiders, art apparently used to subdue prey. Much remains to be learned about prey capture and other forms of be- bavior .
The systematic findings in this work are based largely upon morphological studies and distributional data. Ecological and be- havioral information has been incorporated where available. Mem- bers of the 0. acleistus species group ;and the 0. tridens group are distinguished as much by their ecology as their morphology. Species
in the former group are found in areas of moderately luxuriant herbaceous vegetation, while species of the latter group have been found chiefly in rather dry, rocky areas with relatively scanty veg- etation. Color patterns of the two species groups generally reflect their surroundings. Species of the 0. tridens group are generally black and white, fitting well the rather drab, gray backgrounds where they occur. Members of the 0. acleistus group blend more appropriately with their background coloration of green, and are often deceptively iridescent in sunlight. Members of the tridens group have a more robust fourth pair of legs, presumably fitted for saltatory locomotion in a more open habitat. Systematic studies, such as this one, often provide a foundation for more detailed investigations into the biology of organisms. I hople that this paper will not only fill the need for a basic morpho- logical revision, but stimulate further investigations of the ecology and behavior of Oxyopes.
ACKNOWLEDGMENTS
To fully acknowledge help with this paper, I must trace its inception back toa study begun in 1963 while I was a student at the Museum of Comparative Zoology, Harvard University. The major thrust of this early work resulted in the publication of a monograph on the Oxyopidae of North America, north of Mexico (Bnady, 1964). A grant from the Evolutionary Biology Committee, Biological Lab- oratories, Harvard University, made possible a three week visit to the British Museum (Natural History) in the summer of 1963. Here several of the type specimens of Oxyopes of F.0.P.-Cambridge were examined and drawn for the first time (Figs. 80-83, 84-87, 96797) *
The initial writing and preparation of this paper for publication began in 1973. The types of F.0.P.-Cambridge were examined



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19751 Brady - Oxyopes 19 1
again. I appreciate the courtesy of Mr. F. R. Wanless, Department of Zoology (Arachnida) , British Museum (Natural History) for making this possible
I am especially grateful to Dr. H. W. Levi of the Museum of Comparative Zoology, H~arvard University, who has loaned speci- mens, offered helpful criticism, and aided in other ways the prepara- tion of this publication. Special acknowledgement is due to the late Dr. A. M. Chickering whose collections, now housed in the Museum of Comparative Zoology, provided a wealth of material. I am also indebted to Dr. W. J, Gertsch and Dr. N. I. Platnick for the loan of specimens from the American Museum of Natural History. During the 1972-73 academic year I was appointed Visiting Pro- fessor in the Department of Zoology, University of Florida at Gainesville. For faciliti'es and various courtesies there, I wish to thank Dr. Brian K. McNab, who was then Acting Chiairman. For their helpfulness and encouragement of research, I would like to especially thank Dr. Jonathan Reiskind, Dr. H. K. Wallace, and Dr. John Anderson also of the University of Florida. Splecial thanks are due to my wife Sara who has aided in proof reading and accepted my long absences during the preparation of this paper.
A 1974 Summer Faculty Grant from Hope College aided in prep- aration of this paper. National Science Foundation Grant GB-13925 helped to defray expenses connected with this study and permitted its publication.
METHODS
Measurements. All measurements listed are in millimeters. When available ten specimens of each sex were measured. The purpose of this was to give some idea of the variation to be expected within a species population. Measurements are presented in tabular form to facilitate comparisons and aid in their use. Collection Records. Collection records for Mexico and Central America only are listed. Records north of Mexico are in earlier publications. Localities are listed geographically from north to south and from east to west. The number of specimens collected at each locality is indicated; the lower case "0" representing immature specimens.
Drawings. Epigyna and palps were drawn after they were scraped free olf hair and large setae removed. Internal genitalia of the female were drawn after clearing in clove oil.



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Psyche
SPECIES GROUPS OF OXYOPES
The 16 species of Oxyopes reported from Mexico and Central America in this paper can be combined into five distinct groups. These species groups are based primarily upon the shape or form of the body (morphology), color pattern, relative lengths of legs, and structure of the mlale and female reproductive organs. For the sake of completeness, five species of Oxyopes recorded only from north of Mexico are included in the discussion below. Acleistus Group. This species group may be arbitrarily subdivided into two subgroups: one containing chiapas, summus, occidens, and aglossus; the other containing tibwlis, acleistus, bifidus, and bifissus. In the first sub-group only aglossus is not recorded from Mexico or Central America. All of the second group are found in Mexico. In light of comparative morphology, genitalic structure, color pat- terns, and geographical distribution, a dendrogram representing the relationships of the eight species in this species group is shown in Diagram I.
Important considerations in grouping species in Diagram I were as follows: (I) chiapas and summus have similar internal female genitalia (compare Fig. 33 to Fig. 31) and the male palpi are alike (compare Figs. 92, 93 to Figs. go, 91 ) ; (2) occidens and aglossus are similar in coloration (compare figs. 7, 8 to figs. 9-12 of Brady, 1964), female reproductive organs (compare figs. 23, 24 to figs. 21, 22 of Brady, 1964), and male palpi (compare figs. 33, 34 to figs. chiapas summus occidens aglossus tibialis acleistus bifidus bifissus ACLEISTUS
SPECIES GROUP
Diagram 1.




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19751 Brady - Oxy opes 1 93
39, 40 of Brady,
1964) ;
(3) tibialis and acleistus are much alike in color pattern (compare Figs. 5, 6 with figs. 1-6 of Brady, 1964); female reproductive organs (compare Figs. 35, 36 with figs. 13-20 of Brady, 1964), and male palpi (compare Figs. 94-97 with figs. 35-38 of Brady, 1964) ; (4) bifidus and bifissus have similar color patterns (compare Figs. I, 2 with Figs. 3, 4) internal female geni- talia (compare Figs. 25, 27 with Fig. 37) and male palpi (compare Figs. 78-81 with Figs. 84-89).
It should be stressed
that these are observations that lend credi- bility to Diagram r as it is conceived.
The relationships of the spe-
cies is actually much more complex.
For example, it appears that
tibialis may be near thfe ancestral stem of both acleistus and aglossus- occidens, although it shows more similarities to the former species. The di'agram is a simplification of relationships. This is certainly a
case where additional specimens subjected to numerical analyses of multiple characters might yield important information about rela- tionships.
Tridens Group. This species group may also be arbitrarily sub- divided into two subgroups: one containing apollo and artemis; the other cougar, lynx, tridens, pardus, panther, ocelot, cornutus, and felinus. In the first subgroup only apollo has been found in Mexico. In the second subgroup pardus and lynx are represented by unique specimens (Brady, 1964), and cougar is recorded only from the southeastern United States ( Brady, I 969). Utilizing comparative morphology, genitalic structure, color pat- terns, and distribution patterns, a dendrogram representing the hy- pothetical relationships of the eight species in the second subgroup is shown in Diagram 2. The well developed tibial apophyses and palpal sclerites of 0. apollo and 0. artemis males, as well as strongly developed fourth legs in both sexes, place them near the eight species in Diagram 2. There is considerable resemblance in color among the species of the tridens group that I have seen in the field. This is not so apparent in preserved specimens. Members of this group are not often taken in sweep-nets, but seem to frequent rather dry areas with scanty, low-growing vegetation. Some are found at high ele- vations. When more information is available, this group may be as well defined ecologically as morphologically. Important considerations in grouping species in Diagram 2 were as follows: (I ) cougarJ lynx, tridens, pardus, and panther have well- developed tibial apophyses on the male palpi; (2) cougar nand lynx have very similar palpi (compare figs. 9, 10 of Brady, 1969, with figs. 66, 67 of Brady, 1964) ; (3) tridens and pardus have similar



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I94 Psyche [June
palpi (compare figs. 70, 71 and figs. 68, 69 of Brady, 1964) ; (4) tri- dens and panther have color patterns (compare figs. 62, 63 of Brady, 1964 with figs. 5, 6 of Brady, 1969 and Figs. 15, 16) and epigyna alike (compare figs. 53-55 of Brady, 1964 with figs. 16, I 7 of Brady, 1969 and Figs. 45, 46). It is possible that panther is the female of pardus.
Also ( I ) ocelot, cornutus, and felinus have predominant black and white color patterns, but they lack prominent tibial apophyses. Oxyopes felinus has a small knob on the tibial segment (Figs. 102, 103), (2) the male palpi (Figs. 100-IO~), as well as the epigyna and internal genitalia (Figs. 39-44) of these three species iare much alike. Oxyopes ocelot is the most distinctive of the three. Color patterns in Oxyopes seem to be a conservative feature and are useful in determining relationships. It should be pointed out, however, that color is often altered by scraping off hair during handling and to 'a lesser extent by preservation in alcohol. Also, one needs an adequate sample of specimens to determine the color pat- tern norm, because of individual variation within a population and geographic variation within the species. The structure and larrange- ment of male palpal sclerites and .apophyses as well as external and internal structure of the female genitalia are the most reliable cri- teria for determining species rel~ationships. Because' of a need for more specimens from certain critical areas in Mexico and 'Central America the proposed dendrograrns must be considered strictly tenta- tive as based on the available material. cougar lynx tridens pardus panther ocelot cornutus fel TRIDENS
SPECIES GROUP
Diagram 2.




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19751 Brady - Oxyopes I95
The remaining three species of Oxyopes found in Mexico and Central America are 0. scalaris, 0. salticus, and 0. flavus. They are different from all other species found in this region.. In context with the other species they would constitute distinct monotypic spe- cies groups.
Oxyopes scalaris is essentially a temperate species whose southern limit of range extends into the northern Mexican plateau (Map 4). Oxyopes salticus is found in the United States and occurs throughout Mexico and Central America (Map 3). Oxyopes flavus does not occur north of Mexico (Map 4).
UNIDENTIFIABLE SPECIES
Nathan Banks (1898) described 0. acutus based upon several young males from La Chaparos, Sierra Laguna, and Hermosillo, Mexico and an adult female from Baja California. The female which is figured by Banks, is 0. scalaris Hentz. One of the imma- ture males is labelled as a type in thle collection of the Museum of Comparative Zoology. The color pattern of this specimen matches that of several species in the 0. tridens complex. Oxyopes acutus has appeared only as Ian entry in faunal lists in the zoological litera- ture. The name has not been associated with any specimens since the original description by Banks. For that reason and because of the difficulty in accurately determining the species that the immature type represents, I think it best to consider 0. acutus .as a nonzen dubwm.
The type female of 0. actophilus described by Chamberlin ( 1924) has apparently been lost. The description, based upon an immature female type, is not adequate enough to determine the species with any accuracy. Therefore, I think that 0. actophilus is also best considered a nomen dubium.
KEY TO SPECIES OF OXYOPES
..................................................... ia. Leg I longer than leg IV. 2
...................................................... I b. Leg IV longer than leg I. 9
2a. Body length 7.2 mm or less. .................................................... 3 2b. Body length greater than 7.2 mm.
Color yellow with black
markings.
Pattern as in Figures 13, 14. Epigynum as in Fig- ures 47-54. ........................................................................ flavu~



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Psyche [June
Distinct mid-ventral stripes on fmemora 1-11. ........................ 6 No distinct mid-ventral stripes on femora 1-11. ................... 4 Thin vertical black stripes beginning under AME and con- tinuing to subdistal region of chelicerae as in Figure 3. ........ 5 No thin black stripes as in Figure 3. Russet brown, dark brown, or gray in general appearance (figs. 89, go, 97-99 of Brady, I 964) ................................................................. scalaris Found in Arizona, Baja California.
Probably limited to So-
noran Desert region (figs. 7, 8, 23, 24 of Brady, 1964). ........ ........................................................................................ occidens Found south of Sonoran Desert region (Map I). Figures 3, 4, 37, 38. ........................................................................ <^7wus Epigynum with an anteriorly directed scape (Fig. 24). Color pattern as in Figures I I, I 2. ........................................ salticus Epigynum without an anteriorly directed scape. Color pattern ............................................................................ not as abov~e. 7
Found in northeastern Mexico (Map I ) . Color pattern as in figure 1-6, epigynum as in figures I 3-20 of Brady, 1964. ........ acleistus
........................................................................................ Not found in northeastern Mexico. Occurring westward or southward o,f acleistus. ............................................................ 8 Color pattern as in Figures
I, 2. Epigynum as in Figures
25-30. ................................................................................ bifidus Color pattern as in Figures 5, 6. E~igynum as in Figures 35, 36. .............................................................................. tibialis Color pattern as in Figures 7, 8. Epigynum as in Figures 33, 34. ............................................................................ chiapas Color pattern as in Figures 9, 10. Epigynum as in Figures 31, 32. ............................................................................ swnmus Color pattern as in Figures 15, 16. Epigynum as in Figures 45, 46. ............................................................................ panther Color pattern as in Figures 17, 18.
Epigynum as in Figures
41, 42. ................................................................................ OC~?/O? Color pattern as in Figures 19, 20. Epigynum as in Figures 43, 44. cornutus
............................................................................ Color pattern as in Figures 21, 22. Epigynum as in Figures 39, 40. .............................................................................. felinus Color plattern as in figures 45, 46 of Brady, 1964. Epigynum as in figures 53, 55 of Brady, 1964. ........................ acleistus



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Brady - Oxyopes
I oa.
KEY TO SPECIES OF OXYOPES
MALES
...................................................... Leg I longer than leg IV.
2
.................................................... Leg IV longer than leg I.
10
...................................................... Body length 5 mm or less.
3
Body length greater than 5 mm.
Color yellow with black
markings. Pattern as in Figures 76, 77. Palpus illustrated in ............................................................ Figures 106, 107. flavus
Distinct mid-ventral black stripes on femora 1-11. ................ 6 No distinct mid-ventral striples on femora 1-11. .................... 4 Thin vertical black stripes beginning und,er AME and con- tinuing to subdistal region of chelicerae as in Figures 57, 59. 5 No thin black stripes as in Figures 57, 59. Russet brown, dark brown, or gray in general applearance (figs. 87, 88, 1016, 107 ............................................................. of Brady, I 964) scalaris
Found in Arizona, Bajia California. Probably limited to So- noran Desert region (figs. 25, 26, 33, 34 of Brady, 1964). .... ....................................................................................... occidens Found south of Sonoran Desert region (Map I). Figures .................................................................... 57-60, 84-89. bifissus
With .a well-developed i-etrohteral tibial apophysis as in Fig- ures 94-97. ................................................................................ 9 Without a well-developed retrolateral tibial apophysis as in Figures 94-97. ........................................................................ 7 Tarsus of palpus deeply concave, and with a brush of stiff hairs on the ventral surface of the patella. Color pattern as in ................
Figures 73-75. Palpus as in Figures 98, 99. salticus
Tarsus of palpus not deeply concave, and without a brush of ....................
stiff hairs on the ventral surface of th'e patella. 8
................................
Palpus illustrated in Figures 78-81.
bifidus
..........................
P,alpus illustrated in Figures go, 91.
sumnzus
............................
Palpus illustrated in Figures 92, 93.
chiapas
Found in northern Mexico .and northward (Map I ) . Palpus illustrated in figures 35-38 of Brady, 1964. ................ acleistus Found in southern Mexico and southward (Map I). Palpus .......................................... illustrated in Figures 94-97. tibialis
With a conspicuous apophysis on the patella of the palpus ....................................
(figs. 70, 7 I of Bna,dy, 1964).
tridens




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lob. With a small apophysis on the patella of the palpus (figs. 74, 75 of Brady, 1964) or without a patellar apophysis. ............ I I I 1,a. With a small .apophysis on the patella of the palpus as in fig- ures 74, 75 of Brady, 1964. Color pattern as in figures 72, 73 of Brady, 1964. ............................................................ apollo I I b. Without an apophysis on the patella of the palpus. ............ I 2 ........................
12a. Palpus illustrated in Figures 100, 101. ocelot ........................
12b. Palpus illustrated in Figures 102, 103. felinus- 12c. Palpus illustrated in Figures 104, 105. .................... cornutus SPECIES DESCRIPTIONS
Oxyopes acleistus Chamberlin
Map I.
Oxyopes acleistus Chamberlin, 1929, Ent. News, 40:19, fig. 3, S. Female holotype from Sanford, Seminole Co., Florida, in the American Museum of Natural History, examined. Roewer, 1954, Katalog der Araneae 2 (a) :330. Bonnet, 1958, Bibliographia Araneorum, 2 (4) :3223. Oxyopes aureus Brady, 1964, Bull. Mus. Comp. Zool., 131(13) :457, figs. 1-4, 13-17, 27, 28, 35, 36,
$ 9. Female holotype from Bentsen State Park, 6 mi. SW of Mission, Hidalgo Co., Texas, in the Museum of Compara- tive Zoology. NEW SYNONYMY.
Discussion. After seeing both individual and geographic variation of the epigynia of 0. bifidus (Figs. 25-30), I am led to believe that the differences between 0. aureus and 0. acleistus are, at best, subspecific. The fact that the color patterns and the male palpi of these two species are much alike supports this position. After seeing the variance of certain taxonomic ch'aracters within some Mexican and Cmentral American species, I am impressed with the similarities between 0. aureus and 0. acleistus. Therefore, I consider them conspecific here.
See Brady, 1964, pages 454-460 for figures, measurements, color descriptions, diagnoses, notes on natural history, and collection rec- ords north of Mexico.
Distribution. Florida, Louisiana, Texas and northern Mexico. (Map 1).
Records. MEXICO, Tamaulipas. Rio Gualoliejo near Forblon, 16 April 1939, doo (LID, B. Brown). San Luis Potosl. Huichi-
huayan, I 9 May 1952, d (MAC, WJG, RS) ; 10 mi. N of Valles, 23 July 1945, (A. M. Dame). Nuevo Lion. Horsetail Falls, 7 July 1972, 9 9 (ARB).




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Brady - Oxyopes
Oxyopes occidens Brady
Map I.
Oxyopes occidens Brady, 1964, Bull. Mus. Comp Zool., 131 (13) :464, figs. 7, 8, 23-26, 33,
34, 8 9. Male holotype from Sabino Pond, Sabino Canyon, Santa Catalina Mtns., Pima Co., Arizona, in the Museum of Compara- tive Zoology.
Discussion. For figures, measurements, color descriptions, diag-