Allen R. Brady.
Nearctic species of the new wolf-spider genus, Gladicosa (Araneae: Lycosidae).
Psyche 93:285-320, 1986.

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NEARCTIC SPECIES OF THE NEW WOLF SPIDER
GENUS GLADZCOSA (ARANEAE: LYCOSIDAE)*
Department of Biology, Hope College
Holland, Michigan 49423
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 Nearc- tic 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 che- liceral 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.
Gladicosa gen. nov.
Lycosa (part) Walckenaer 1837: 338. Emerton 1885: 485. Marx 1890: 562; 1892: 160. Stone 1890: 423, 426. Montgomery 1902: 538, 546, 566; 1904: 277-280; 1905: 174; 1909: 514. Banks 1901: 184; 1910: 55, 57; 191 1: 454. Chamberlin 1904: 147; 1908: 225,226,265; 1924: 28. Petrunkevitch 191 1: 560. Comstock 1913: 631,639; *Manuscript received by the editor July 15, 1986 285




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1940: 644, 650. Bishop and Crosby 1926: 207. Wood 1926: 174. Crosby and Bishop 1928: 1067. Elliott 1930: 5; 1932: 423. Worley and Pickwell 193 1: 9 1,93. Chickering 1932: 351. Gertsch 1934: 7, 8; 1949: 82. Gertsch and Wallace 1935: 20-22; 1937: 10. Kaston 1935: 191; 1936: 103, 114; 1938: 184; 1948: 322, 328; 198 1 : 322, 328. Allard 1936: 67. Jones 1936: 69. Chamberlin and Ivie 1944: 142, 144. Bonnet 1957: 2607, 2635, 2645. Fitch 1963: 108-109. Whitcomb, Exline, Hunter 1963: 656. Whitcomb and Bell 1964: 45. Dorris 1965: 408; 1968: 36. Drew 1967: 194. Harrison 1969: 14- 16. Bultman, Uetz, Brady 1982: 26. Leimonia (part) Simon 1864: 352.
Trochosa (part) Montgomery 1904: 301, 305. Chamberlin and Ivie 1942: 35. Avicosa (part) Roewer 1954: 236.
Hogna (part) Roewer 1954: 258.
Scaptocosa (part) Roewer 1954: 293.
Varacosa (part) Roewer 1954: 306.
Alopecosa (part) Bonnet 1955: 248.
Type species. Gladicosa gulosa (Walckenaer) Etymology.
The generic name is a combination of gladius (Latin for sword) referring to the unique sword-shaped embolus of the male palpus, and cosa derived from the generic name Lycosa. It is considered feminine.
Diagnosis. Gladicosa may be distinguished from other lycosid genera by the following combination of characters: (1) the swordlike or bladelike form of the embolus (em) and its clockwise orientation in ventral view of the left palpus of the male (Fig. 33), (2) the modi- fication of the terminal apophysis (ta), which is also broadly flattened and parallels (and partly supports) the embolus (Figs. 33,34), (3) the rectangular or wedge shape of the transverse piece (tp) of the scape of the epigynum, together with its white pearlescent appearance, in whole or part (Fig. 10) and (4) the dorsal color pattern illustrated in 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



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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.
Legs when compared to body dimensions relatively longer than in Trochosa; without distinct annulations; yellow, yellow-orange to golden brown in color. Order of leg length IV-1-11-111. Tibial spina- tion in female: leg I, 2-2-2 ventral, 1-0 or 1-1 prolateral; leg I1 2-2-2 ventral, 1 - 1 prolateral; leg I11 2-2-2 ventral, 1 - 1 prolateral, 1 - 1 retro- lateral, 1-1 dorsal; leg IV 2-2-2 ventral, 1-1 prolateral, 1-1 retrolat- eral, 1-1 dorsal. Tibial spination in the male is the same with the addition on leg I of 1-1 retrolateral and leg I1 1-1 retrolateral. Dorsal abdominal pattern variable according to size and hirsute- ness, but generally with anterio-lateral black markings aligned with those on carapace, cardiac area well marked, and often with pattern of chevrons as indicated in Figures 1-5. Dark colors on dorsum of abdomen brown to black, lighter colors cream to tan or beige. Ven- ter of abdomen cream to light brown in gulosa, huberti, and euepi- gynata; dark brown to black in pulchra and bellamyi. Region anterior to epigastric furrow of contrasting darker or lighter color respectively.
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



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coming to a point near margin of cymbium (cy); heavily sclerotized spur directed medially (Figs. 30, 33, 34). Epigynum of female with scape shovel-shaped with elongate lon- gitudinal piece (lp) (handle) and rectangular or trapezoidal trans- verse piece (tp) (blade). The transverse piece is unusual in being wholly or partly translucent white or pearlescent in appearance (Fig. 10). Spermathecae (s) smooth and round to ovoid (Fig. 7), rarely elongate ovoid (Fig. 15); usually their diameter apart. 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. Measure- ments 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.
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 of certain populations: Dr. Richard Brown and Ms. Pat Miller of the Entomological Museum, Mississippi State University; Mr. Tim



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Lockley, Deta State Research Center, USDA, Stoneville, Missis- sippi; Dr. G. B. Edwards and Dr. H. K. Wallace of the University of Florida, Gainesville; and Dr. Andrew Penniman, Defiance College, Defiance, Ohio.
Special thanks are extended to Dr. C. D. Dondale and Dr. H. W. Levi who consented to review the manuscript and offered construc- tive criticism and friendly advice. I am also grateful to Mr. F. R. Wanless for sending specimens of Lycosa pulchra Keyserling from the L. Koch collection maintained in the British Museum (Natural History). A note of special appreciation to Ms. Amy Youatt, who helped with general sorting, compilation of locality data, and prep- aration of distribution maps.
National Science Foundation grant DEB-780356 1 assisted in defraying expenses of the investigation. A summer grant from the faculty development program at Hope College (1980) helped to initiate this project.
Transverse piece (tp) of scape of epigynum rectangular, about equal in length and width (Figs. 6- 14) ................. .2 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. Lungitudi- 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. Longitudinal piece wider anteriorly, narrowing posteriorly (Figs. 2 1-26) ...................... bellamyi



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Both embolus (em) and terminal apophysis (ta) bladelike, paralleling one another with each separate and drawn out to .........................
a point (Figs. 27,28,35-42) .3
Embolus bladelike, but terminal apophysis not resembling it; the two not as distinctly separated as above (Figs. 29-34, ............................................. 43-46) 2
Relatively small species. Total length 7.8 to 11.0 mm (Figs. 29-34). Not reported from central Texas. ........ beZZamyi Relatively large species. Total length 10.4 to 13.9 mm (Figs. ...........
43-46). Distribution central Texas euepigynata Embolus with relatively short, pointed tip (Figs. 27, 28) ........................................ huberti Embolus with longer drawn out tip that is curved at end. ... .4 Tip of embolus pointed; median apophysis (ma) with large ret- ......................
rolateral spur (Figs. 35-36) gulosa
Tip of embolus flattened; median apophysis (ma) with small ...................
retrolateral spur (Figs. 37-42) pulchra
Gladicosa gulosa (Walckenaer), comb. nov. Figures 5, 6-9, 35, 36. Map 1.
Lycosa gulosa Walckenaer, 1837: 338. Male holotype from North America, de- stroyed. Marx 1890: 562. Chamberlin 1908: 225, 226, 265, pi. 21, figs. 4, 7, Q Q. Montgomery 1909: 5 14. Petrunkevitch 19 1 1 : 560. Comstock 19 13: 63 1,639, figs. 720 g-h, $5; 1940: 644,650, figs. 720 g-h, Q Q. Bishop and Crosby 1926: 207. Wood 1926: 174. Crosby and Bishop 1928: 1067. Elliott 1930: 5; 1932: 423. Worley and Pickwell 193 1 : 9 1, 93. Chickering 1932: 35 1. Gertsch 1934: 7; 1949: 82. Gertsch and Wallace 1935: 20. Kaston 1935: 191; 1936: 103, 1 14; 1938: 184; 1948: 322,328, pi. 57, figs. 1 106-1 109, $8; 1981: 322,328, figs. 1 106-1 109, Q 3. Allard 1936: 67. Fitch 1963: 108-109, fig. 46. Whitcomb, Exline, Hunter 1963: 656. Whitcomb and Bell 1964: 45. Don-is 1965: 408; 1968: 36. Drew 1967: 194. Harrison 1969: 14-16. Bultman, Uetz, Brady 1982: 26. Leimonia gulosa: Simon 1864: 352.
Lycosa kochi: Emerton 1885: 485, pi. 46, figs. 6-6c, Q 3; 1902: 74, figs. 179, 180, Q. Stone 1890: 423,426, pi. 15, fig. 3. Marx 1892: 160. Gertsch and Wallace 1935: 21, figs. 39,42,
Q. Not Lycosa kochi Keyserling.
Lycosa helluo: Banks 190 1 : 184 (part). Lycosa nigraurata Montgomery, 1902: 538, 546, pi. 30, fig. 53, Q. Male holotype from Medford, Burlington Co., New Jersey (N.J. Stone), examined. Synony- mized with Lycosa purcelli Montgomery by Montgomery 1904: 305. Lycosa purcelli Montgomery, 1902: 538, 566, pi. 30, figs. 30, 31, 9 8. Female syntype from Philadelphia, Philadelphia Co., Pennsylvania, May, 1888, and



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19861 Brady- Nearctic Gladicosa 29 1
male syntype from Point Pleasant, Ocean Co., New Jersey, 30 April 1889 (N.J. Stone), examined. Synonymized with Lycosa kochi: Emerton by Gertsch and Wallace 1935: 2 1.
Trochosa purcelli, Montgomery, 1904: 30 1, 305. Lycosapulchra: Chamberlin 1904: 147 (part); Banks 1910: 57 (part). Varacosa gulosa: Roewer 1954: 306.
Alopecosa gulosa: Bonnet 1955: 248.
Discussion. The nomenclatural history of G. gulosa is complex. Walckenaer's (1837) seven-line description without figures is not diagnostic for this species. The locality given is North America, and that doesn't help. To complicate matters, Emerton (1885) misidenti- fied 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 namepurcelli has been used only by Montgomery (1902, 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.
Carapace light brown to brown, with broad yellow to yellow- orange median stripe. Narrow irregular submarginal yellow stripes suffused with brown. Posterior declivity with black patches as in Figure 3.




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Fig. I,
Gladicosa huberti(Chamberlin), female from Bar M Ranch near Boston, Thomas Co., Georgia, 2 Mar. 1973.
Fig. 2. Gladicosa behnyi (Gensch and Wal- lace), female from 2 mi. N of Stoneville, Washington Co., Mississippi, 9-1 1 May 1983. Fig. 3. Gladicosa pulchra (Keyserling), female from Gainesvilie, Alachua Co., Florida, 14 June 1935. Fig. 4.
GWicosa guhu (Walckcnaer), female from 4 mi. S of New Richmond, Allegan Co., Michigan, 16 Sept. 1974. Fig. 5.
GIadicosa
euepigynata (Montgomery), Camp Verde, Ken- Co., Texas, kc, 1939.



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Dorsum of abdomen light brown to brown with pair of black anterior-lateral patches as in Figure 5. Anterior cream to yellow spots mark depressions of internal muscle attachments. Cardiac area faintly indicated. Venter of abdomen cream or light beige to pale yellowish brown. Few scattered darker spots. Overlaid with fine coat of white hair.
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, 1 1, 13, 14). In gulosa the embolus is pointed at the end, whereas that of pulchra is somewhat spatulate in shape (compare Figs. 35, 36 with Figs. 37, 38).




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Table 1.
Measurements of ten females and ten males of Gladicosa gulosa from Allegan Co., Michigan.
Females: Mean SEM Mean SEM
Ant. Eye Row
PME
PLE
POQ
Car. Width
Car. Length
Body Length
Pat.-Tibia I1
Pat.-Tibia 111
Males: Mean SEM
Femur I
Pat.-Tibia I
Meta. I
Tarsus I
Total I
Femur IV
Pat.-Tibia IV
Meta. IV
Tarsus IV
Total IV
Mean SEM
Ant. Eye Row
PME
PLE
POQ
Car. Width
Car. Length
Body Length
Pat,-Tibia I1
Pat.-Tibia 111
Femur I
Pat.-Tibia
Meta. I
Tarsus I
Total I
Femur IV
Pat.-Tibia IV
Meta. IV
Tarsus IV
Total IV i
Natural History.
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



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Map 1.
Distribution of G. gulosa.
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 1 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



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