NEARCTIC SPECIES OF THE NEW WOLF SPIDER GENUS GLADICOSA ( ARANEAE : LYCOSIDAE ) * BY

This is the second paper in a projected series of systematic studies of the Nearctic Lycosidae described primarily in the genus Lycosa. Over 50 species of medium to large size wolf spiders from the Nearctic Region have been placed in this genus. However, recent studies indicate that several distinct genera are included under Lycosa. Matters have been complicated at the generic level by C. F. Roewer (1954) who listed 44 new genera of Lycosinae in the Katalog der Araneae. They are nomina nuda, lacking descriptions. Later Roewer (1959, 1960) defined these 44 genera, thus validating the names, and added seven more new ones to the Lycosinae as well. These genera were established primarily on the basis of differences in the number of posterior cheliceral teeth and eye arrangement (particularly eyes of the anterior row). Investigations of North American Lycosidae (Brady 1962, 1972, 1979) indicate that the number of posterior cheliceral teeth is an unreliable character in delimiting genera. Recent studies indicate that color patterns on the dorsal surface of the carapace, length of legs relative to body size, and particularly the structure of the male and female genitalia are most reliable in determining generic relationships. Certain features of the eye arrangement, as well as information about habitat, behavior, and life history are also useful. In the final analysis, it is the unique combination of all these features that should be employed to distinguish genera.

Figures 1-5 and described below.Description.Total length 7.8 to 18.8 mm.Carapace length 4.2 to 8.3 mm; width 3.1 to 6.4 mm.Carapace viewed dorsally, narrow- ing at level of PLE row, smoothly convex along lateral margins, with posterior margin concave; viewed laterally essentially the same height from eye region to posterior declivity (highest point is poste- rior cephalic region in front of dorsal groove with the carapace sloping very slightly anteriorly).Dorsal groove long and distinct.Dorsal color pattern with light uneven submarginal stripes and wide median light colored stripe, narrow between ALE, widening until just anterior to dorsal groove (where it is usually constricted), becoming wider again parallel to groove, and then narrowing as it     1986]   Brady--Nearctic Gladicosa 287 follows thoracic declivity to posterior edge of carapace.Black mark- ings framing median stripe at posterior declivity.Dark areas of carapace brown to dark brown and black.Light stripes pale yellow to yellow-orange (Figs.1-5).
Anterior median eyes (AME) slightly larger than anterior lateral eyes (ALE).Anterior eye row much narrower than posterior median eye row (PME), with dorsal tangent slightly procurved.Posterior lateral eye row (PLE) much the widest (see Tables 1-6).
Chelicerae dark reddish brown to black; anterior and posterior margin each with three teeth, the anterior triad crowded more closely together.
Male palpus with stridulatory file situated retrolaterally at tip of tibia.Cymbium with cluster of macrosetae at tip, and with stridula- tory scraper retrolaterally at base.Male palpal sclerites as seen in ventral view" Palea (pa) concave, largely hidden by embolus, visible along retrolateral margin.Embolus (em) blade-like, tapering to a point, with clockwise orientation (from left to right) in left palpus, which is opposite to that of most Lycosinae.Conductor (co) con- cave, with cuplike portion containing tips of the terminal apophysis (ta) and the embolus.Terminal apophysis large, flattened and paral- leling embolus, with its tip serving partly as a conductor.Median apophysis (ma) with a flattened ridge extending retrolaterally and Psyche [Vol. 93  coming to a point near margin of cymbium (cy); heavily sclerotized spur directed medially (Figs. 30,33,34).

METHODS
The techniques and methods employed in the study of Gladicosa were essentially the same as for Trochosa (Brady 1979) and are described there.Color descriptions are based upon appearance of specimens in alcohol illuminated by microscope lamp.Measurements are listed in millimeters, but for Gladicosa the mean and standard error (SEM) are listed instead of the mean and range as in the previous paper.Methods and techniques of measurement are described in the paper on Trochosa (Brady 1979).Under Records specific localities are given for uncommon species and the peripheral range for common species, otherwise localities of specimens exam- ined are indicated by counties.

ACKNOWLEDGMENTS
This study was made possible by the loan of large numbers of specimens from the Museum of Comparative Zoology, Cambridge, Massachusetts, the American Museum of Natural History, New York City, and the Canadian National Collection, Ottawa, Canada.I wish to thank sincerely the curators of those collections, Dr. H. W. Levi, Dr. N. J. Platnick, and Dr. C. D. Dondale respectively for the use of these materials.The loan of type specimens from the Museum of Comparative Zoology, the American Museum and the Phila- delphia Academy of Natural History is gratefully acknowledged.
Thanks are offered to Mr. Donald Azum for loan of the latter.
I am indebted to the following individuals and institutions for making available regional collections that provided a much better picture of geographical distribution and clarified the relationships Transverse piece (tp) of scape of epigynum irregular in shape (Figs.15-17) or, if rectangular, much wider than long (Figs.18-26)   3 Transverse piece entirely pearlescent in appearance.Longitudi- nal piece (lp) lacking indentations where it joins transverse piece (Figs.6-9) gulosa Transverse piece only partly pearlescent white.Longitudinal piece (lp) with indentations at posterior end where it joins transverse piece (Figs.10-14)  pulchra Transverse piece irregular in shape and broadly joined by longi- tudinal piece (Figs.15-17)   euepigynata Transverse piece somewhat rectangular, much wider than long and narrowly joined by longitudinal piece 4 Width of transverse piece greater than length of longitudinal piece.Longitudinal piece about the same width throughout its length (Figs.18-20)   huberti Width of transverse piece equal to or less than length of trans- verse piece.
diagnostic for this species.The locality given is North America, and that doesn't help.To complicate matters, Emerton (1885) misidenti- fled this species as Tarentula kochi Keyserling and transferred it to the genus Lycosa.Gertsch and Wallace (1935) discussed the syste- matic and nomenclatural problems associated with G. gulosa and suggested using the name Lycosa kochi Emerton for this species since Emerton (1885) had placed the species in a different genus.However, according to Article 49 of the International Code of Zoo- logical Nomenclature (1985): "A previously established species- group name wrongly used to denote a species-group taxon because of misidentification cannot be used for that taxon even if it and the taxon to which the name correctly applies are in, or are later assigned to, different genera, except when a previous misidentifca- tion is deliberately used in fixing the type species of a new nominal genus."Bonnet (1955) points out that the name nigraurata or pure- celli of Montgomery should have been used for the species.Mont- gomery (1904) himself synonymized nigraurata with purcelli and the name purcelli has been used only by Montgomery (1902Montgomery ( , 1904)).
The name gulosa, on the other hand, has been employed numerous times since Gertsch and Wallace's (1935) invocation of kochi, and even by Gertsch (1949) in his book American Spiders.It therefore seems best to retain the name gulosa for this species to promote stability of nomenclature by preserving a long accepted name in its accustomed meaning.
Color.Females.Face yellow or yellow-orange, to pale golden brown.Eye region darker with nacelles black.Chelicerae yellowish brown to dark reddish brown, almost black at distal ends.Condyles yellow or orange, to golden brown.
Legs yellow or pale yellow-orange to yellowish brown, darker distally.Femora with dusky bands on dorsal and lateral surfaces.
Ventral surface lighter yellow.
Labium and endites brownish orange to brown with distal ends yellow to cream.Sternum yellow to light golden brown.
Color.Males.Face yellow to yellow-orange, darker brownish in eye region.Chelicerae with basal areas yellow to orange-yellow, darker brown to reddish brown distally.Condyles orange-yellow to orange.Cymbia of palpi dark brown.
Carapace brown with a broad median yellow stripe and irregular yellowish submarginal stripes obscured by thicker clothing of white hair.

Dorsum of abdomen beige to light brown with black markings
along sides beginning anteriorly and continuing posteriorly.Black markings often more prominent than in female.Posterior of dorsum without distinct chevrons as in other species.Venter of abdomen pale yellow to beige, clothed with white hair which is more abun- dant laterally.
Legs yellow to brownish yellow.Darker dorsally without dusky markings on femora as in female.
Labium and endites orange-yellow to orange-brown with distal ends lighter yellow to beige.Sternum orange to orange-brown.
Measurements.Ten females and ten males from Allegan Co., Michigan.See Table 1.
Diagnosis.Gladicosa gulosa is closest to G. pulchra in size and coloration.The markings of pulchra offer greater contrast, and chevrons are usually visible on the dorsum of the female abdomen (compare Fig. 5 with Fig. 4).The epigyna of the females and the palpi of the males also resemble one another in appearance, but are distinctly different when compared in detail.The epigynum of gulosa has the transverse piece entirely pearlescent white, whereas pulchra has some white, but nearly always shows darker brown sclerotized areas on the transverse piece (compare Figs. 6, 8, 9 with  Figs. 10, 11, 13, 14).In gulosa the embolus is pointed at the end, whereas that ofpulchra is somewhat spatulate in shape (compare Figs. 35, 36 with Figs.37, 38).Kaston (1948) reports gulosa running over dead leaves on forest floors in Connecticut.I have found it in leaf litter of deciduous woods in Michigan.Here it is found in more open Oak woodlands as opposed to the shaded floor of Beech- Maple forests.In Michigan and New England gulosa usually matures late in the fall, overwinters as an adult, and mates in early spring.Kaston (1936) made the following observations of courtship behavior in the species: Immediately upon coming in contact with the female, or within 3 minutes thereof, the male begins to drum his palps rapidly against the floor of the cage.These drumming move- ments are made so rapidly that a distinct purring or humming sound can be heard.The palps are used alternately and are raised only a very short distance during the process.The body is held at an angle so that the posterior end of the abdomen almost touches the floor.As a consequence when the male begins to twitch his abdomen in a vertical plane the tip strikes the floor.However, I could not detect any sounds made by this part of the body.It is highly probable that the vibrations set up in the substratum by the tapping movements of the palps and abdomen are perceived by the female.This may exert an excit- ing influence on her in a manner analogous to that which occurs in web-building species, where the male tweaks the threads of the female's snare.
The male now moves slowly toward the female without courting.When near her he reaches over to touch her.At first she may jump at him and chase him away.Later, if she is receptive she allows him to stroke her legs or abdomen.After this contact with the female the male resumes his courtship movements.Later on, if the male gets more excited he begins to raise his forelegs off the floor about or 2 mm, and lower them quickly.During this process the legs quiver violently.
After 13 minutes of this courting one male began to mount the female, but before he could get into the final copulatory position, she ran away from him.Another male had courted only seven minutes when the female allowed him to mount.The position is the usual one for Lycosids, the male using his palps Psyche [Vol. 93  alternately during the 10 minutes the act lasted.This duration time may not be the usual one for the species, however, for one pair were observed in the field, when collected, which were already in copula and remained so for about another half hour.The sound produced during courtship was also reported by Allard (1936).Observations were made on a collecting trip in the Bull Run Mountains of Virginia during late April.He described the sound as a distinct purring produced by drumming rapidly upon dry leaf surfaces.He reports: The creatures were very wary, but with care I was able to examine their movements critically from a distance of only a few inches.When the spider moved and made its sounds, the fore part of the body quivered perceptibly and the palpi, too, executed gentle up and down movements.The quivering movements brought the chelicerae directly in contact with the dry leaf surface, and the latter alone appeared to be responsible for the rather loud sounds I had heard.According to Allard these tapping sounds could be heard a distance of 10 feet or more.Rovner (1975) investigated sound production in three species of Schizocosa and six species of Lycosa, including gulosa.Previous investigators, as with gulosa above, had regarded such sounds as being solely percussive, generally produced by a tapping or scraping of the palps or the chelicerae against the substratum.High-speed Figs.6-9.Gladicosa gulosa (Walckenaer) 6-7.Female from 4 mi.S of New Richmond, Allegan Co., Michigan, 16 Sept. 1974. 6. Epigynum.7. Internal geni- talia.8. Epigynum of female from Pepperell, Middlesex Co., Massachusetts, Apr. 1973. 9. Epigynum of female from Cove Creek Valley, 15 mi.S of Prairie Grove, Washington Co., Arkansas.Figs.10-14.Gladicosa pulchra (Keyserling). 10.Epigynum of female from Stone Co., Mississippi, 21 Dec. 1964. 11.Epigynum of syntype from North Ameri- ca.12-13.Female from Gainesville, Alachua Co., Florida, 14 June 1935. 12. Internal genitalia.13.Epigynum.14. Epigynum of holotype of Lycosa inso- pita Montgomery [--Gladicosa pulchra (Keyserling)] from Austin, Travis Co.,   Texas.Figs.15-17.Gladicosaeuepigynata (Montgomery).15-16.Female from Camp Verde, Kerr Co., Texas, Dec. 1939.15  Rovner (1975) revealed the prescence of a stridulatory organ at the tibio-tarsal joint.This apparatus consists of a file on the distal end of the tibia and a scraper at the base of the palpal cymbium.Further examination revealed a group of stout spines or macrosetae at the tip of the palpal tarsus.These spines apparently aid in coupling the tarsus to the substratum.Thus, the sound produced by gulosa (C)and other lycosids is not generated simply by drumming, but involves a rapid oscillation at the tibio-tarsal joint facilitated by macrosetae that anchor the palpus to the substratum.Kaston (1948) reports seeing mature females of gulosa from Sep- tember, through winter, to June suggesting that some may live for two years.Egg sacs appear in early April and are produced until late May.Egg sacs vary from 6-10 mm in diameter and egg counts range Figures 4, 10-14, 37-42.Map 2.
Color.The range of color in G. pulchra is greater than that of G. gulosa.I have noted light forms and dark forms of pulchra.
These do not represent a genetic polymorphism but are the extremes in a color continuum.There is no discernible correlation between geographic locality and color pattern among the specimens exam- ined.The darker forms are much more numerous than the light colored ones.The range of color is indicated in the following descriptions.
Color.Female.Face orange-brown to dark reddish brown.Chelicerae dark reddish brown to black with condyles lighter orange-brown.
Carapace dark brown to a dark reddish brown with a broad median yellow stripe suffused with white hair.Irregular lighter submarginal yellow stripes similarly clothed with white hair.Pattern as in Figure 4. Dorsum of abdomen brown to brown mottled with black.
Anterio-lateral areas black, blending with similar black areas on cephalothorax.Five pairs of white spots (in well-marked specimens) beginning in cardiac area and continuing posteriad.White spots connected by dark brown chevrons as in Figure 4. Cardiac area darker brown, outlined by lighter brown or yellowish.
Venter of abdomen dark brown to almost black posterior to epi- gastric furrow.Yellowish anterior to furrow.
Legs light brown with darker black annulations on femora to dark reddish brown without distinct annulations.
Labium and endites light brown to black with pale yellowish distal ends.Sternum yellow brown (golden), dark reddish brown to black.
Color.Male.Face yellow-orange to orange-brown.Dark in ocular area.Chelicerae brownish orange to dark reddish brown.Cymbia of palpi yellow-orange to dark reddish brown.
Carapace orange-brown to dark orange-brown with broad yellow to pale orange median stripe overlaid with white hair.Irregular submarginal stripes of same color, sometimes indistinct.Dorsum of abdomen with median area light to medium brown, bordered by black.Five pairs of white spots beginning in cardiac area and continuing posteriad.Spots joined by black chevrons.
Cardiac area brown, enclosed by lighter pale brown to yellowbrown.Pattern similar to female.Venter of abdomen brown to black posterior to epigastric furrow.Light brown to pale yellow or cream anterior to furrow.Labium and endites yellow-orange to orange with distal ends cream.Sternum yellow-orange to orange.
Measurements.Ten females and ten males from Florida.See  Brady--Nearctic Gladicosa  303   than gulosa (compare  2) and is usually darker in color with a more distinct pattern (compare Fig. 4 with Fig. 3).In most specimens ofpulchra the venter of the abdomen is dark brown to black behind the epigastric furrow, while that of gulosa is yellowish to light brown.Differences between female and male genitalia of these two species are noted under gulosa and in the keys.Natural History.Little is known of the habitat or behavior of pulchra.I've collected this species in Florida from the trunks of deciduous trees where their color blends well with the bark substrate.G. B. Edwards (personal communication) has collected spec- imens from similar microhabitats in Florida.Pat Miller (personal communication) reported collecting both male and female pulchra from the trunks of pine trees at night in Perry, Florida, on December 5, 1982.Montgomery (1904) reported finding pulchra near Austin, Texas, in drier habitats than gulosa and less abundantly.He noted that the females live under stones where they make a shallow horizontal burrow lined with silk.Whether this behavior is consistent throughout the life cycle or represents a temporary adjustment to molting or egg laying is a question to be answered.Gladicosa pulchra is not the abundant inhabitant of deciduous leaf litter, as are gulosa and huberti.Of the species investigated pulchra is the most variable in coloration of the body and structure of the epigynum.It is possible that more than one species is represented in this complex.
Roble (1986) reported rearing Mantispa viridis from a Gladicosa pulchra egg sac.It is the first record of a lycosid spider serving as a host of M. viridis.When the spider died, its egg sac was opened and a mantispid cocoon and 95 surviving spiderlings were found.This corroborates an earlier observation of high spiderling survival within a mantispid-infested egg sac of Lycosa rabida.
Distribution.From Long Island, New York, along the East Coast to Texas in the southwest.Limited in its northern range inland to the southern parts of Kansas and Missouri and northern Kentucky.More abundant in the southeastern United States (Map 2).
Color.Females.Face orange-brown to reddish brown with eye nacelles black.Chelicerae dark reddish brown (mahogany) to black.Condyles orange-brown.
Carapace orange-brown to reddish brown with broad median pale orange stripe from PME to posterior edge.Lighter irregular submarginal stripes less distinct than median.Pattern as in Figure 1.
Dorsum of abdomen brown to dark brown with cardiac area outlined in black.Chevrons faintly indicated along posterior half with white spots marking their lateral edges.Anterior lateral edges of dorsum darker as in Figure 1.Venter pale yellow-orange to darker brown.Lateral areas darker in pale-colored individuals, con- colorous brown in others.
Legs yellow-orange to orange-brown, without darker annulations.
Labium and endites orange-brown to dark reddish brown, with distal ends yellowish to cream.Sternum yellow-orange to light orange-brown.
Color.Males.Face dark orange-brown to very dark reddish brown, eye region black.Chelicerae dark reddish brown to black.Condyles lighter.Cymbia of palpi dark red-brown.
Carapace orange-brown to darker reddish brown with light orange broad median stripe from eye region to posterior edge.Light- er, irregular submarginal stripes, not so distinct as median one.Dorsum of abdomen medium to dark brown with cardiac area lighter, outlined by black line which is enclosed in turn with lighter color extending laterally.Anterior lateral areas marked by black color, which extends more posteriad than in female.Venter of BradymNearctic Gladicosa 307 abdomen orange-brown to dark brown.Central area somewhat lighter.
Legs yellow-orange to orange-brown, somewhat lighter ventrally, without darker bands.
Labium and endites yellow-orange to dark reddish brown, with distal ends pale yellow to cream.Sternum yellow to reddish orange-brown.
Measurements.Ten females and ten males from Georgia and Florida.
Natural History.Nothing concerning the natural history of this species is reported in the literature.I have collected it in leaf litter near the edge of woods in Georgia and in a marshy area near the edge of a pond beneath a pine tree canopy in Florida.The great majority of the adult specimens were collected from February through April (see Records below).
Discussion.Gladicosa bellamyi was placed in the new genus Avicosa by Roewer (1954) with Avicosa avida (Walckenaer) [--Schizocosa] as the type species.Two other North American species now placed in Schizocosa (minnesotensis and wasatchensis mccooki) as well as Lycosa ceratiola and Tarentula pictilis (now Alopecosa pictilis) were also included in this new genus.Avicosa is certainly an artificial conglomeration without systematic foundation.Color.Females.Face orange-brown to dark reddish brown.
Chelicerae dark reddish brown to black.Condyles lighter yellowish.
Carapace dark brown to dark reddish brown with broad median yellow-orange to pale brownish orange stripe from PME to poste- rior declivity as in Figure 2. Indistinct submarginal stripes of same color.

Psyche
[Vol. 93  Dorsum of abdomen pale yellow-brown to medium brown, often with darker brown cardiac mark and darker chevrons posteriorly as in Figure 2. Slight indication of black counter-shading anterio- laterally.Venter of abdomen dark brown posterior to epigastric furrow; median area sometimes mottled with light orange-brown.Lighter yellowish anterior to furrow.
Legs brown to dark brown dorsally.Pale yellowish brown to golden brown ventrally.Legs without distinct bands.
Labium and endites dark reddish brown to orange-brown with distal ends lighter golden to yellow.
Color.Males.Face dark red-brown.Eye region black.Che- licerae dark brown to black with inner distal margins lighter orange- brown.Condyles lighter orange to yellow.Cymbia of palpi brown to dark brown.
Carapace dark reddish brown overlaid with fine black hair.Broad median pale yellow-orange to orange-brown stripe from PME to posterior edge.Dorsum of abdomen beige to light brown.Black countershading in anterio-lateral areas, extending posteriorly farther than in female.
Indistinct chevrons posteriorly.In some specimens the median longi- tudinal area of the dorsum is pale yellow to cream with darker brown at edges and along sides.Venter of abdomen dark brown to black posterior to epigastric furrow, lighter yellowish brown ante- riorly.Lateral areas often somewhat lighter in color.
Legs orange-brown to dark brown dorsally, paler golden to yel- lowish brown ventrally.Without darker bands.Tibia and metatar- sus black, tarsus yellow.Labium and endites orange-brown to dark brown with distal ends lighter yellow to golden.Sternum light orange-brown to darker reddish brown.
of certain populations: Dr.Richard Brown and Ms. Pat Miller of the Entomological Museum, Mississippi State University; Mr. Tim helped with general sorting, compilation of locality data, and prep- aration of distribution maps.National Science Foundation grant DEB-7803561 assisted in defraying expenses of the investigation.A summer grant from the faculty development program at Hope College (1980) helped to initiate this project.

Table 1 .
Measurements of ten females and ten males of Gladicosa gulosa from Allegan Co., Michigan.

Table 2 .
Diagnosis.Gladicosapulchra is closest to G. gulosa in size, col- oration, and genitalic structure.Gladicosa pulchra is a larger species

Table 3 .
Measurements of ten females and ten males of Gladicosa huberti from

Table 4 .
Measurements of ten females and ten males of Gladicosa bellamyi from

Table 5 .
Measurements of ten females of Gladicosa bellamyi from Mississippi.

Table 6 .
Measurements of ten females and ten males of Gladicosa euepigynata from Texas.