The Lynx Spider Genus Oxyopes in Mexico and Central America (Araneae: Oxyopidae)

This is the second in a projected series of papers on the Neotropical Oxyopidae. In this study I6 species of Oxyopes are recorded from Mexico and Central America. Of these., II 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 (I969-7I), Costa Rica (I972), and Panama (1972). Named species of Oxyopes from M’exico and Central America were often inadequately described and figured and there was some confusion as to the correct names of several species, males and females 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 are extremely abundant in areas of grassy or herbaceous vegetation. For example, O. salticus and O. flavus constituted as much as 20% of the total number of spiders taken in sweeping vegetation (o.5-I m in height) in some coastal are:as 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 during the day. They ran swiftly over the plants where they occurred and when disturbed exhibited characteristic erratic leaps which allow them to be immediately recognized in the field. Although Oxyopes do not actively pursue their prey by sight, they are much


INTRODUCTION
This is the second in a projected series of papers on the Neotropical Oxyopidae. In this study I6 species of Oxyopes are recorded from Mexico and Central America. Of these., II 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 (I969-7I), Costa Rica (I972), and Panama (1972). Named species of Oxyopes from M'exico and Central America were often inadequately described and figured and there was some confusion as to the correct names of several species, males and females 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 are extremely abundant in areas of grassy or herbaceous vegetation. For example, O. salticus and O. flavus constituted as much as 20% of the total number of spiders taken in sweeping vegetation (o.5-I m in height) in some coastal are:as 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 during the day. They ran swiftly over the plants where they occurred and when disturbed exhibited characteristic erratic leaps which allow them to be immediately recognized in the field. Although Oxyopes do not actively pursue their prey by sight, they are much IManuscript received by the editor March 25,1975 I89 Psyche [June more active than thomisids, and exhibit a general "searching" behavior when observed. Long spines on the legs, another eld characteristic o.f lynx spiders, are apparently used to subdue prey. Much remains to be learned about prey capture and other forms o beh, avior.
The systematic findings in this work a.re based largely upon morphological studies and distributional data. Ecological and behavioral information has been incorporated where, available. Members of the O. acleistus species group and the O. tridens group are distinguished as much by their ecology as their morphology. Species in the former group are ound in areas o modergtely luxuriant herbaceous vegetation, while species o the latter' group have. been ound chiefly in rather dry, rocky areas with relatively scanty vegetation. Color patterns of the two. species groups generally reflect their surroundings. Species of the O. tridens group are generally black and white, fitting well the rath'er drab, gray backgrounds where they occur. Members of the O. acleistus group blend more appro,priately with their background coloration o.f green, and are oten deceptively iridescent in sunlight. Members o.f 'the tridens group have a more robust ourth 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 hope that this paper will not only ill the need for a basic morphological revision, but stimulate further investigations of the ecology and behavior of Oxyopes.

ACKNOWLEDGMENTS
To ully acknowledge help with this paper, I must trace its inception back to a study begun in I963 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 (Brady,I964 A I974 Summer Faculty Grant rom Hope College aided in preparation o this paper. National Science Foundation Grant GB-I3925 helped to deray expenses connected with this study and permitted its publication.

METHODS
Measurements. All measurements listed are in millimeters. When available ten specimens o each sex were measured. The purpose o this was to give some idea o the vari,ation t.o be expected within a species population. Measurements are presented in tabular orm to acilitate comparisons and aid in their use.

Collection Records. Collection records xCor Mexico and Central
America only are listed. Records north o Mexico are in earlier publications. Localities are listed geographically rom north to south and i:rom east to west. The number ox specimens collected at each locality is indicated; the lower case "o" representing immature specimens.
Drawings. Epigyna and palps were drawn .ater they were scraped ree of hair and large setae removed. Internal genitalia o the emale were drawn ater clearing in clove oil.

SPECIES GROUPS OF OXYOPES
The 16 species o1: Oxyopes reported rom Mexico and Central America in this paper can be combined into ve distinct groups.
These species groups are based primarily upon the shape or form o( the body (morphology), color pattern, relative lengths o1: legs, and structure o1: the m.ale and emale reproductive organs. For the sake o.1: completeness, ive species o1: Oxyopes recorded only rom north o Mexico are included in the discussion below.
A cleistus Group. This species group may be arbitrarily subdivided into two subgroups: one containing chiapas, summus, occidens, and aglossus; the other containing tibialis, acleistus, bifidus, and bifissus. In the qrst sub-group only aglossus is not reco.rded 1:rom Mexico. or Central America. All o1: the second group are found in Mexico.
In light ot: comparative morphology, genitalic structure, color patterns, and geographical distribution, a dendrogram representing the relationships o1: the eight species in this species group is shown in Diagram I. Important considerations in grouping species in Diagram were as ollows: (I) chia#as and summus have similar internal female genitalia (compare Fig. 33  It should be stressed that these are observations that lend credibility to Diagram as it is conceived. The relationships of the species is actually much more complex. For example., it appears that tibialis may be near the ancestral stem of both acleistus and aglossusoccidens, although it shows more similarities to the orme.r species. The diagram is a simplification o relationships. This is certainly a case where additional specimens subjected to numerical analyses of multiple characters might yield important information about relationships. Tridens Groul. This species group may also be arbitrarily subdivided into two subgroups: one containing a#ollo and artemis; the other cougar, lynx, tridens, pardus, lanther, ocelot, cornutus, ,and [elinus. In the first subgroup only a#ollo has been found in Mexico. In the second subgroup #ardus and lynx are represented by unique specimens (Brady,I964), and cougar is recorded only from the southeastern United States (Brady,I969).
Utilizing comparative morphology, genitalic structure, color patterns, and distribution patterns, a dendrogram representing the hypothetical relationships of the eight species in the second subgroup is shown in Diagram 2. The well dev'eloped tibial apophyses and palpal sclerites of O. a#ollo and O. 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 o the tridens group that I have seen in the field. This is not so apparent in preserved specimens. Members of this group are not otten taken in sweep-nets, but seem to requent rather dry areas with scanty, low-growing vegetation. Some are tound at high elevations. 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) cougar, lynx, tridens, pard'us, and panther have welldeveloped tibial apophyses on the male palpi; (2) cougar and lynx have very similar palpi (compare figs. 9, IO Brady, 1964 with figs. 5, 6 of Brady, 1969 andFigs. 15, 16) and epigyna alike (compare figs. 53-55 of Brady, 1964 with figs. 16, 17 of Brady, 1969 andFigs. 45, 46). It is possible that panther is the female of pardus. Also (I) ocelot, cornutus, and [elinus have predominant black and white color patterns, but they lack prominent tibial apophyses. Oxyopes [elinus has a small knob on the tibial segment (Figs. lO2, IO3), (2) the male palpi (Figs. IOO-IO5), as well as the epigyna and int.ernal genitalia (Figs. 39-44) of these three species are much Mike. 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 pattern norm, because of individual variation within a population and geographic variation within the species. The structure and arrangement of male palpal sclerites and apophyses as well as external and internal structure of th'e female genitalia are the most reliable criteria for determining species relationships. Because of a need for more specimens from certain critical areas in Mexico and Central America the proposed dendrograms must be considered strictly tentative as based on the available material. 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 Carapace cream to pale orange with darker brown markings as illustrated in Figure 2.
Dorsum o abdomen white to yellow medially, the cardiac area translucent; bounded on each side by brown as in Figure 2. Venter with broad, dark brown median stripe rom epigastric furrow to base of spinnerets; bounded by lighter cream to yellow.

1975]
Brady---Oxyo#es 2Ol Legs cream to yellow. Dark median stripe on ventral surface ot emora I and II.
Labium and endites yellow to pale orange. Sternum cream to pale yellow.
Color. Male. Face and chelicerae yellow to pale orange with darker brown stripes as illustrated in Figure 55. Eye region dark brown to black. Cymbia ot palpi dark brown to black.
Carapace yellow to pale orange with aint indications ot more dusky markings as shown in Figure 56.
Carapace yellow to pale orange with dark brown hairs forming pattern as illustrated in Figure 4.
Dorsum ot: abdomen white to pale yellow, bordered by dark brown as in Figure 4. Vent'er white to pale yellow, speckled with brown.
Legs yellow to pale orange with irregul,ar dusky spots, most conspicuous ventrally.
Labium cream to pale orange. Endites cream to pale orange, outlined with dark brown, Sternum white to pale yellow, with dark brown margin.
Color. Males of O. bifissus occur in two color phases. The darker of these forms is described first, then the lighter phase.
Dark male. Face and chelicerae dark brown with vertical black stripes as in Figure 57. Eye region dark brown with iridescent scales. Cymbia of palpi dark brown to black. Distal ends of chelicerae lighter.
Carapace with median area light brown, bordered by dark brown to black as shown in Figure 58.
Dorsum of abdomen light brown in center to dark brown along sides as in Figure 58; overlaid with iridescent scales. Spinnerets yellow to light orange-brown. Venter dark brown to black with pale lines originating at corners of epiga.stric furrow and continuing to base of spinnerets. Median area heavily clothed with iridescent scales. Scales impart a green or pink iridescence in preserved specimens. Color. Female. Face and chelicerae pale yellow to light yello.worange with dark stripes rom AME to subdistal region of chelicerae as in Figure 5. Eyes with black nacelles, overlaid with white spatulate hair.
Carapace cream to light yellow-orange, with broad dark submarginal stripes as in Figure 6.
Dorsum o abdomen with cardiac area translucent, accented by white. Median area white to pale yellow, outlined by mottled brownish yellow anteriorly and dark brown posteriorly as in Figure   6. Spinnerets pale yellow. Ven:er with broad, mottled brown or solid brown median stripe from epigastric urrow to base o. spinnerets.
Legs yellow. Femora. I-III with mid-ventral dark lines.
Labium and endites yellow to light yellow-orange. Sternum cream to pale yellow.
Color. Male. Face and chelicerae with black reticulate pattern and indistinct vertical stripes as in Figure 6. Some specimens dark brown to black without stripes apparent. Cymbia o palpi black.
Eyes with black nacelles.
Carapace yellow to yellow-orange in some; dark brown with lighter yellow surrounding thoracic groove in others.
Dorsum o abdomen brown mottled with cream to yellow, giving overall gray-brown appearance. Spinnerets yellow. Venter with same mottled appearance as dorsum, with a wide median brown stripe' rom epigastric urrow to base o spinnerets.
Legs yellow with median ventral stripe on femora I-III. Labium  Color. Female. Face and chelicerte pale cream to yellow with vertical black stripes from AME to subdistal ends of chelicerae as in Figure 7. Eye nacelles black, ov'erlaid with white hair.
Carapace pale yellow. Dorsum of abdomen white to cream with brown along sides as in Figure 8. Spinnerets pale yellow. Venter with broad cream colored stripe outlined with brown, extending rom epigastric furrow to base of spinnerets. Lateral areas white.
Legs cream to pale yellow. Brown stripe on ventral surface of femora I and II in one specimen.
Labium brown. Endites pale yellow. Sternum cream. Color. Male. Face and chelicerae pale yellow to yellow-orange. Black vertical stripes from AME to subdistal region of chelicerae as in Figure 63. Eye region black. Iridescent spatulate hairs between PME. Cymbia of palpi dark brown to black.
Carapace yellow to yellow-orange. Some purplish iridescence in one male.
Dorsum of abdomen with cardiac area translucent gray, surrounded by white pigment with lateral areas dark brown to black as in Figure 64. Other  Color. Female. Face and chelicerae pale orange-yellow, lighter cream along lower edge .of clypeus. A pair of black lines from AME to subdistal region of chelicerae as in Figure 9. Eye nacelles black.
Carapace pale orange-yellow with darker brown submarginal stripes as in Figure O.
Dorsum of abdomen with a broad median white stripe from base to spinnerets, outlined with dark brown. Sides mottled with brown as in Figure O.
Venter with a broad median black stripe from epigastric furrow to base of spinnerets, mottled with cream in center. Median stripe flanked by pale cream.
Legs yellow with mid-ventral black line on femora I and II. Line faintly visible on femora III. Labium and endites cream to pale yellow with lighter distal ends. Sternum ivory to cream.
Color. Male. Face and chelicerae pale orange-yellow; lighter along lower edge of clypeus. Two black lines from AME to subdistal end o.f chelicerae as in Figure 65. Eye tubercles black.
Carapace pale orange-yellow with few scatt'ered dusky hairs. Faint iridescent sheen of green and purple.
Dorsum o.f abdomen with a broad white stripe rom base to tips of spinnerets, bordered laterally by mottled light brown extending along sides as in Figure 66.
Venter with wide black median stripe from genital area. to base of spinnerets, bounded by pale yellow.
Legs yellow with mid-ventral black stripe on emora I-III.  (Brady, 1969 Carapace cream to golden yellow with broad dark brown submarginal stripes as in Figure 22.
Dorsum of abdomen with cardiac area translucent cream, outlined with brown. Median area white to cream with lateral areas darker, mottled with dark brown or black against white as in Figure   22. Venter with dark brown median stripe, divided longitudinally into three narrower stripes, bounded on its edges by white or cream. Sides mottled brown and cream.
Legs yellow. Distal ends of femora and tibia with irregul'ar dark markings. A pair of dark ventro-lateral stripes on femora I-III. Carapace yellow-orange with wide dark brown submarginal stripes as in Figure 7o. Color. Female. Face and chelicerae white with a. pair of wide dark brown stripes from _ALE to subdistal ends of chelicerae as illustrated in Figure I9.
Carapace with median area white, bounded by wide dark brown submarginal stripes, with white at margins as in Figure 2o.
Dorsum of abdomen with median white stripe outlined in black, becoming dark brown laterally. Two pairs of white marks posteriorly as in Figure 2o. Venter with median area dusky, bounded by pale cream, becoming darker (light brown) laterally.
Legs yellow to light brown, overlaid with dark hair. Femor,a with dark stripe along anterio-ventral surface.
Labium and endites yellow. Sternum cream to pale yellow.
Color. Male. Face with dark brown or black median area bounded by white and chelicerae with vertical black stripes as in Venter with dark brown median stripe outlin'ed in pale cream to yellow and becoming darker laterally.
Legs yellow to brown, overlaid with dusky hair, without distinct darker markings.
Labium and endites pale yellow. Sternum cream to pale yellow. Diagnosis. Oxyopes cornutus, particularly the male, has a black and white color pattern simil.ar to O. tridens and O. pardus. The male of O. cornutus lacks a patellar apophysis such as that found in 0. tridens and 0. pardus. Oxyopes cornutus appears to be most closely related to 0. felinus in male palpal structure and female epigynal structure. It can, however, be readily separated rom 0. Holotype. Female from 12 mi. W of Portal, Cochise Co., Arizona, 13 July 1968, (L. D. Mikelson) in the Museum of Comparative Zoology. The specific name is a noun in apposition after the Panther.
Discussion. In an earlier paper on the apollo species group I illustrated and described an unknown female Oxyopes from Portal, Arizona (Brady, 1969). Since the females of O. pardus, O. lynx, and O. felinus were unknown at that time, I thought it best not to assign a distinctive name to a single, female, specimen that belonged to the same species group. During the present study four additional females of this same species were found in collections from Mexico.
Although the male is still not known, I have decided to name the females in order that they can be identified. The actual relationship of these five females to O. pardus will be determined only when males and females are found in .association.
Measurements. For measurements of the holotype see Brady, 1969 yellow, overlaid with white appressed hair. Brown stripes rom AME to subdistal region o chelicere as in Figure 15. Eye nacelles black, covered with white hair. Carapace yellow, overlaid with white spatulate-shaped hair. Wide dark brown submarginal stripes as in Figure 6.
Dorsum o( abdomen with broad median white stripe outlined with brown; brown mottled with white along sides as in Figure I6.
V'enter with a wide mottled brown and yellow median stripe rom epigastric urrow to base o spinnerets. This stripe surrounded by pale cream laterally.
Legs yellow, shaded with brown. Color. Female. Face and chelicera.e yellow to yellow-orange with faint darker vertical stripes from AME to subdistal region .of chelicerae as in Figure 7. Eye nacelles black. Lower edge o,f clypeus lighter, pale yellow.
Carapace pale yellow to yellow with a pair of wide brown submarginal stripes as in Figure I8. Dorsum of abdomen dark brown with a wide median white stripe as in Figure 8. Sides of abdomen pale yellow with irregular brown spots. Venter with a wide brown stripe from epigastric furrow to base of spinnerets, outlined by white or pale yellow.
Legs yellow to yellow-orange with mid-ventral black stripe on femora I and II.
St'ernurn cream to pale yellow.
Color. Male. Face and chelicerae pale yellow to light orange without darker markings, with lower edge of clypeus cream as in Figure 7. Palpi light to dark brown. Carapace yellow to pale yellow-orange with darker brown submarginal stripes as in Figure 72.
Dorsum of abdomen with broad white median stripe bordered by dark brown to black as in Figure 72. Venter with wide median brownish stripe from epigastric furrow to base of spinnerets, bordered by lighter cream.
Legs pale yellow with mid-ventral black lines on femora I-II. Labium and endites pale yellow. Sternum cream to pale yellow. Darl} lale. A single darker colored male has the face and chelicerae orange brown without distinct darker markings. Cymbia of palpi black.
Carapace yellow-orange with dark brown submarginal stripes. Dorsum with bro'ad white median stripe overlaid with darker scale-like hairs. Venter with median stripe dark brown to black.
Diagnosis. Oxyol)es ocelot has the well-developed fourth pair of legs and the bro.ad median white stripe of the abdomen flanked by black that is characteristic of the tridens species group. The face is not strongly marked as in other members of the group, however. The p'alpal organ of the male and to a lesser extent the epigynum of this species place it .closest to O. cornutus Carapace cream to pale yellow with our gray stripes as indicated in Figure I2. Gray stripes ormed of appressed hairs, which are rubbed away in some specimens. Dorsum with median area white with darker brown outlining cardiac area. White, pale yellow-orange, and brown spatulate hairs orming pattern illustrated in Figure I2. Venter with wide, dark brown median stripe beginning at epigastric furrow and continuing to base of spinnerets. Brown stripe is sometimes mottled.
Legs cream to pale yellow with black stripe on ventral surfaces of femora I-III. Labium  Carapace pale yellow to light yellow-orange. A few faint dusky markings contributed by spatulate-shaped hairs as in Figure 74.
Dorsum of abdomen cream to pale yellow, dusky laterally .and along sides of abdomen. Scale like hairs providing a metallic green or pink sheen. Cardiac area outlined as in Figure 74. Venter with broad median brown stripe from epigastric furrow to base of spinnerets. The stripe outlined in white, with dusky color laterally.
Legs cream to pale yellow without stripes or dusky ma.rkings in most specimens. Mid-ventral black stripe on femora I and II of some.
Labium cream to pale yellow. Endites cream to pale yellow, outer margins darker brownish. Sternum yellow. Dark Male. Face black between AME to clypeus and chelicerae with black markings as shown in Figure 75. Cymbia of palpi black.
Carapace pale yellow to light yellow-orange, with four longitudinal dusky stripes. Dorsum of abdomen iridescent, mostly dark gray-brown to black. Venter dark gray-brown to black with white lines outlining median stripe conspicuously seen in lighter forms.
Legs cream to pale yellow. Patellae-tibiae and femora of palpi with black markings dominating lighter color.
Labium and endites black with distal ends pale cream. Sternum yellow with black periphery.
Diagnosis. Oxyoles salticus resembles members of the acleistus group with mid-ventral black stripes on the femora and vertical black lines on the ace. It can be immediately distinguished by the peculiar shape of the epigynum (Figs. 23, 24) and the characteristic palpus of the male (Figs. 98, 99). The male. palpus has a stout conical projection on the cymbium and a stout brush of short, stiffened bristles (not figured) on the ventral surface of the palpal tibia. Oxyopes salticus should be placed in a separate species group to reflect its relationship to other Mexican and Central American species. Natural History. Oxyopes salticus is one of the most common spider inhabitants of tall grass and herbaceous vegetation and is found throughout Mexico and Central America wherever this type of vegetation occurs. The local abundance of these spiders gives evidence of their role, as one of the important predators of insects  Color. Male. Face and chelicerae cream to yellow with black stripes from AME to distal third of chelicerae as in Figure 76. Eye region black overlaid with white spatula.te-shaped hairs.
Carapace cream to yellow with dusky markings as in Figure 77-Dorsum .of abdomen white to cream with darker brown marks along sides as in Figure 77. Venter with wide dark brown or black stripe rom genital area to base o.f spinnerets. Median stripe bounded by white with yellow along sides.
Legs pale yellow to yellow without darker markings or a few scattered black spots, especially on ventral surfaces.