Journal of Arachnology

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Journal of Arachnology 33(2):347-376. 2005
doi: 10.1636/04-80.1

EVOLUTION OF ORNAMENTATION AND COURTSHIP BEHAVIOR IN SCHIZOCOSA: INSIGHTS FROM A PHYLOGENY BASED ON MORPHOLOGY (ARANEAE, LYCOSIDAE)

Gail E. Stratton

Department of Biology, University of Mississippi, University, Mississippi 38677, U.S.A.

Abstract

A phylogenetic analysis for the North American Schizocosa species was undertaken by scoring 49 morphological characters for 31 taxa representing all of the Nearctic species of Schizocosa plus individuals that are hybrids between S. ocreata and S. rovneri. Rabidosa rabida, Allocosa georgicola and Gladicosa pulchra were used as outgroups. Three clades are recognized: a large clade from eastern North America (Clade A) within which is nested the S. ocreata clade; Clade B, which includes the widespread S. avida and the western S. mccooki, and a smaller, third clade, Clade C. Sexual ornamentation occurs on the first legs of mature males of several species within the Schizocosa and takes the form of pigmentation and or bristles primarily on the tibia of leg I; there is at least one species with bristles in each of the three main clades. Mapping the occurrence of male ornamentation on the preferred phylogeny suggests that ornamentation evolved 5 or 6 separate times and was subsequently lost 2 or 3 times. The ornamentation is concentrated in the S. ocreata clade, a clade defined by a finger like projection on the paleal process of the male pedipalp. Courtship behavior is known for 20 of the 31 taxa. All species studied utilize chemical communication and seismic signals for communication; some species also have distinct visual signals. Seismic signals are produced by palpal drumming (as is seen in several species within Clade B), or by stridulation (seen in Clade A). Visual signals consisting of movements of the first pair of legs are common in species that are distinctly ornamented. This study provides the first phylogenetic study of a North American genus of wolf spider and provides morphometric comparisons of the North American species in Schizocosa.

Received: 22 September 2004; Revised: 1 April 2005

Keywords: Cladistics, sexual selection, secondary sexual characteristics, evolution of behavior, spiders, multimodal signal, seismic signal



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Appendix 2


Specimens examined for this study and literature references for figures. Deposition of specimens are as indicated: GES (collection of G. Stratton), PRM (collection of P. Miller), AMNH (American Museum of Natural History), FSCA (Florida State Collection of Arthropods), and MCZ (Museum of Comparative Zoology). Coordinates are provided for as many collecting localities as possible, either by transcribing data included on collecting labels, or searching for localities on the USGS National Mapping Information website (http://geonames.usgs.gov/) search under “U.S. and territories query.” Coordinates not included on original labels are in brackets.



Schizocosa aulonia Dondale 1969

U.S.A.: Illinois: Madison County, Poag, SE 1/4 NW 1/4SE SE 1/4 Sec. 13 T4N, R9W, [38°48′N, 90°02′W], pitfall, 12–19 July 1993, David Landes, 2 ♂ (GES).



Schizocosa avida (Walckenaer 1837)

U.S.A.: Mississippi: Pontotoc County, 3 miles NE of Pontotoc off Highway 9, field near Mubby Creek, R3E T95 Sect 15 [34°18′N, 88°56′W], 13 June 1992, G. Stratton, P. Miller, W. Miller, 1 ♂(GES); Missisippi: Marshall County, 3 miles S. of Waterford [34°36′N, 89°29′W], 9–12 June 1994, G. Stratton, 1 ♀ (GES). Published figures: Dondale & Redner (1978, figs. 10–12, 51–54).



Schizocosa bilineata (Emerton 1885)

U.S.A.: Michigan: Eaton County, T2N R6W Sect 6 [42°36′N, 85°02′W], on porch of farm, 29 June 1989, G. Stratton, 1 ♂ (GES); Mississippi, Panola County, 3 miles NE. of Como, west side of Nelson Creek, on Compress Road, T6S R7W Sec 23 & 24 [34°32′N, 89°54′W], 15 May 1998, G. Stratton, P. Miller, 1 ♀ (GES). Published figures: Dondale & Redner (1978, figs. 8, 47, 48).



Schizocosa cespitum Dondale & Redner 1978

CANADA: Saskatchewan: Matador, 40 miles N. of Swift Current, short grass prairie, P.W. Riegert, 1 ♂, 3 ♀ paratypes (AMNH). Published figures: Dondale & Redner (1978, figs. 20, 69, 70).



Schizocosa chiricahua Dondale & Redner 1978

U.S.A.: Arizona: Chochise County, Chiricahua Mts, Southwest Research Station [31°53′N, 109°12′W], 5400′, in swimming pool, 15 July 1967 & June 7 1968, V. Roth, 2 ♂ paratypes (AMNH); Arizona: Graham County, near Safford [32°50′N, 109°42′W], 14 July 1956, W. Gertsch & V. Roth, 2 ♂ paratypes, 1 ♀ paratype (AMNH). Published figures: Dondale & Redner (1978, figs. 18, 19, 66– 68).



Schizocosa communis (Emerton 1885)

CANADA: Ontario: 9 miles NW. of Bloomfield, 44°04′N, 77°18′W, 17 July 1965, Jean & Wilton Ivie, 1 ♂ (AMNH); U.S.A.: Pennsylvania: Berks County, Lenhartsville [40°34′N, 75°53′W], July 1964, Vaurie, 1 ♀ (AMNH). Published figures: Dondale & Redner (1978, figs. 14, 55–58).



Schizocosa crassipalpata Roewer 1951

CANADA: Ontario: Rednersville [44°07′N, 77°27′W], field pitfall, 3–19 June 1964, C.D. Dondale, 2 ♂, 2 ♀ (FSCA). Published figures: Dondale & Redner (1978, figs. 9, 49, 50).



Schizocosa crassipes (Walckenaer 1837)

U.S.A.: Florida: Alachua County, 0.25 miles E. of River Styx on Highway 346 [29°31′17″N, 82°15′47″W], 10 May 1994, G.B. Edwards & P. Cushing, 3 ♂, 4 ♀ (GES). Published figures: Dondale & Redner (1978, figs. 2, 27–30); Stratton (1991, fig. 4).



Schizocosa crassipes (Walckenaer 1837)

U.S.A.: Mississippi: Grenada County, T21N R2E sect. 12, 13N &R3E Sec 7S, 18N [33°44′N, 90°00′W], deciduous woods on hillside, 16 May 1993, G. Stratton, P. Miller, W. Miller, B. Grantham, 1 ♂, 1 ♀ (GES).



Schizocosa duplex Chamberlin 1925

U.S.A.: Mississippi: Lafayette County, 8 miles SE. of Oxford, T10S R3W Sec 35, 34°36′N, 89°29′W, in pine litter,18 May 1993, G. Stratton, E. Hebets, 1 ♂, 2 ♀ (GES). Published figures: Dondale & Redner (1978, figs. 6, 42–44).



Schizocosa floridana Bryant 1934

U.S.A.: Florida: Marion County, Hopkins Prairie, Ocala National Forest [29°10′N, 81°47′W], 25 March 1983. G. Stratton, litter, 4 ♂, 2 ♀ (GES). Published figures: Dondale & Redner (1978, figs. 3, 31–34, 35).



Schizocosa humilis (Banks 1892)

U.S.A.: Pennsylvania: Bucks County, east of Jamieson, Horseshoe Bend, Neshaminy Creek, May 1954, W. Ivie (GES). Published figures: Dondale & Redner (1978, figs. 7, 45, 46).



Schizocosa maxima Dondale & Redner 1978

U.S.A.: California: Solano County, Fairfield [38°15′N, 122°02′W], April–August 1955, K.W. Haller, 1 ♂paratype (AMNH); California: Tuolumne County, S. of Highway 108, 5 miles E of Sonora, elev. 2000′, drowned in swimming pool, 14 October 1973, W. Icenogle, 1 ♀paratype (AMNH). Published figures: Dondale & Redner (1978, figs. 16, 17, 63–65).



Schizocosa mccooki (Montgomery 1904)

U.S.A.: New Mexico: Santa Fe County, Santa Fe [35°31′N, 105°56′W], in pinyon scrub, 17 June 1979, D. Lowrie, 3 ♂, 1 ♀ (GES). Published figures: Dondale & Redner (1978, figs. 13, 15, 59– 62).



Schizocosa mimula (Gertsch 1934)

U.S.A.: Colorado: Otero County, Highway 109, 5 June 1967, 1 ♂ (AMNH).



Schizocosa minnesotensis (Gertsch 1934)

U.S.A.: Wyoming: Lincoln County, Kemmerer [41°48′N, 110°32′W], 24 August 1983, R. Parmenter, 2 ♂, 2 ♀ (AMNH).



Schizocosa ocreata (Hentz 1844)

U.S.A.: Ohio: Clermont County, Cincinnati Nature Center [39°07′N, 84°15′W], 6 May 1980, G. Stratton, 1 ♂, 1 ♀ (GES). Published figures: Stratton (1991, figs. 3, 9).



Schizocosa ocreata (Hentz 1844)

U.S.A.: Mississippi: Washington County, Leroy Percy State Park, N. of Highway 12 near entrance of park, T15N R7W, 90°50′W, 33°10′N, 9 April 1993, on knoll by flooded bottomlands, G. Stratton, P. Miller, 1 ♂, 1 ♀ (GES).



Schizocosa ocreata (Hentz 1844)

U.S.A.: Florida: Alachua County, 0.25 miles E. of River Styx on Highway 346 [29°31′17″N, 82°15′47″W], 1 March 1993, G.B. Edwards, P. Cushing, 9 ♂, 3 ♀ (GES).



Schizocosa retrorsa (Banks 1911)

U.S.A.: Mississippi: Lafayette County, 8 miles SE. of Oxford, T10S R3W Sec 35, 34°36′N, 89°29′W, pitfall in pine woods, 28 June–5 July 1993, 1 ♂,1 ♀, G. Stratton (GES). Published figures: Dondale & Redner (1978, figs. 21, 75–78).



Schizocosa rovneri Uetz & Dondale 1979

U.S.A.: Kentucky: Boone County, 5 miles W. of Taylorsport, floodplain of Ohio River, “Sandy Run” [39°05′N, 84°41′W], 3 May 1996, K. Delaney, 1 ♂, 1 ♀ (GES). Published figures: Stratton (1991, figs. 2, 7).



Schizocosa salsa Barnes 1953

U.S.A.: Mississippi: Hancock County, mouth of Jordan River [30°16′N, 89°37′W], on Juncus marsh island, 25 June 1993, 1 ♂ (PRM). Published figures: Dondale & Redner (1978, figs. 24, 79, 80).



Schizocosa saltatrix (Hentz 1844)

U.S.A.: Mississippi: Lafayette County, 8 miles SE. of Oxford, T10S R3W Sec 35, 34°36′N, 89°29′W, “Lonesome 80,” pine deciduous woods, 16 March 1996, G. Stratton & P. Miller, 1 ♂, 1 ♀ (GES). Published figures: Dondale & Redner (1978, figs. 4, 39–41).



Schizocosa sp. nr saltatrix

U.S.A.: Mississippi: Wilkinson County, 5 miles E of Doloroso on Smith Rd, S. of Homochito River, 31°20′N, 92°45′W, uplands deciduous forest, 10 April 1993, G. E. Stratton, P.R. Miller, 1 ♂, 1 ♀ (GES).



Schizocosa segregata Gertsch & Wallace 1937

U.S.A.: Florida: Levy County, 28 April 1934, # 298, H.K. Wallace, 1 ♂, 1 ♀, paratypes (poor condition); Texas: Edinburg [26°18′N, 98°10′W], 1934, S. Mulaik, 1 ♂ (poor condition) (AMNH). Published figures: Dondale & Redner (1978, figs. 23, 81, 82).



Schizocosa stridulans Stratton 1984

U.S.A.: Illinois: Mason County, Sand Ridge State Forest [40°24′N, 89°52′W], 7 June 1985, G. Stratton, L. Hartz, 1 ♂, 1 ♀ (GES). Published figures: Stratton (1991, figs. 1, 5, 6, 13).



Schizocosa uetzi Stratton 1997

U.S.A.: Mississippi: Lafayette County, 8 miles SE of Oxford, T10S R3W Sec 35, 34°36′N, 89°29′W, “Lonesome 80,” mixed pine and hardwoods, 4 July 1992, G. Stratton, 1 ♂, 1 ♀ (GES). Published figures: Stratton (1997, figs. 1–5).



Schizocosa ocreata × Schizocosa rovneri hybrids

Cross between S. ocreata ♀ from Ohio and S. rovneri ♂ from Kentucky, 1980–1982, 1 ♂ (GES). Cross between S. rovneri ♂ from Kentucky and S. ocreata ♀from Ohio, 1980–1982, 1 ♀ (GES).



Allocosa georgicola (Walckenaer 1837)

U.S.A.: Mississippi: Lafayette County, 8 miles SE. of Oxford, T10S R3W Sec 35, 34°36′N, 89°29′W,“Lonesome 80,” pitfall in mixed pine and hardwoods, 16–24 September 1992, G. Stratton, P. Miller, 1 ♂,1 ♀ (GES). Published figures: Chamberlin & Ivie (1944, fig. 57). Pedipalps and epigyna of the closely related H. helluo (Walckenaer 1837) are figured in Kaston (1948, figs. 1065, 1066, 1090, 1105) and Dondale & Redner (1990, figs. 43–47).



Gladicosa pulchra (Keyserling 1877)

U.S.A.: Tennessee: Hardeman County, Chickasaw State Park [35°22′N, 88°50′W], pine deciduous woods, 1 November 1992, G. Stratton, 1 ♂, 1 ♀ (GES). Published figures: Brady (1986, figs. 3, 10–14, 39–42).



Rabidosa rabida (Walckenaer 1837)

U.S.A.: Mississippi: Lafayette County, 8 miles SE. of Oxford, T10S R3W Sec 35, 34°36′N, 89°29′W, pine deciduous mixed, 5 August1991, G. Stratton, P. Miller, G. Miller, 1 ♂, 1 ♀ (GES). Published figures: Brady & McKinley (1994, figs. 1, 6, 11–14) and Kaston (1948, figs. 1077, 1079, 2006).

Figures 1–6.—Dorsal view of males of several of the species used in the phylogenetic study: 1. Allocosa georgicola; 2. Gladicosa pulchra; 3. Schizocosa avida; 4. Schizocosa retrorsa; 5. Schizocosa rovneri; 6. Schizocosa uetzi

Figures 7–10.—Venters of selected species: 7. Male of Schizocosa crassipes; 8. Male of Schizocosa retrorsa; 9. Female of Schizocosa bilineata (epigynum has been dissected); 10. Female of Schizocosa avida

Figures 11–14.—Left pedipalps of males from selected species of Schizocosa. Localities are indicated in Appendix 2: 11. S. avida; 12. S. retrorsa; 13. S. saltatrix; 14. S. duplex. ELL = ear-like lobe, IPE = intromittent portion of embolis, MA = median apophysis, TA = terminal apophysis. Scale lines = 500 μm

Figures 15–18.—Pedipalps of males from selected populations of Schizocosa ocreata and S. crassipes. Localities as indicated in Appendix 2: 15. S. ocreata (FL); 16. S. ocreata (MS); 17. S. crassipes (FL); 18. S. crassipes (MS). Scale lines = 500 μm

Figures 19–22.—Epigyna of selected species. Localities as indicated in Appendix 2: 19. S. avida, 20. S. retrorsa, 21. S. crassipes (MS); 22. S. duplex. MS = median septum, TP = transverse piece. Scale lines = 250 μm

Figures 23–31.—Shape and size of tibial bristles in all taxa with the tibial bristles in the Nearctic Schizocosa: 23. S. aulonia; 24. S. salsa; 25. S. bilineata; 26. S. segregata; 27. S. crassipes (MS); 28. S. stridulans; 29. S. ocreata (FL); 30. S. ocreata (OH) ; 31. S. ocreata (MS)

Figure 32. Single most parsimonious tree from heuristic searches including all taxa, excluding secondary sexual characters and applying preferred weighting (somatic characters = “2,” genitalic characters = “3.” The tree shows clades A, B, & C as well as the S. ocreata clade. Bootstrap values above 50% are shown

Figure 33. Mapping of the ornamentation characters “pigment on femur” and “pigment on tibia,” seen in mature males on the preferred phylogeny

Figure 34. Mapping of representative tibial bristles, seen in mature males, on the preferred phylogeny.

Figure 35. Cladogram of S. ocreata clade + basal species showing presence and extent of bristles on leg I

Figure 36. Distribution of major courtship elements on the preferred phylogeny. The symbol “D” refers to species in which the males are known to show palpal drumming during courtship, “S” refers to species in which males are known to show palpal stridulation. The “+” symbol shows there is arching of leg 1, “++” shows species which have extensive visual signals

table

Table 1. Body size (carapace length, carapace width and ratio of length/width) of male and female Schizocosa and outgroups used in this study. All measures are in mm. Measures of female S. aulonia, and S. maxima are from Dondale & Redner (1978), measures from S. salsa are from Barnes (1952)

table

Table 2. Tibial length and relative tibial length (tibial L/carapace L) of males of Schizocosa and outgroups. All measures are in mm

table

Table 3. Length, width and length/width of tibia and tarsus of pedipalp for Schizocosa and outgroups. All measures in mm

table

Table 4. Epigynal measures including total length of epigynum, and width of epigynum, the ratio of epigynal length/width, and depth of the epi gynal hood. All measures in mm. Values for three species (S. salsa, S. segregata, S. aulonia) were not available and for these species, ratio of length to width was calculated from published figures in Dondale & Redner (1978)

table

Table 5. Comparison of elements of courtship behavior (both seismic and visual) in species of Schi zocosa, Gladicosa and Allocosa

table

Table 5. Continued

table

Appendix 1.—Data matrix of character states for the phylogenetic analysis of Schizocosa wolf spiders and outgroups

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Online publication date: 1-Jul-2009.
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E. A. Hebets. (2008) Seismic signal dominance in the multimodal courtship display of the wolf spider Schizocosa stridulans Stratton 1991. Behavioral Ecology 19:6, 1250-1257
Online publication date: 8-Sep-2008.
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Kevin J. Delaney, J. Andrew Roberts, George W. Uetz. (2007) Male signaling behavior and sexual selection in a wolf spider (Araneae: Lycosidae): a test for dual functions. Behavioral Ecology and Sociobiology 62:1, 67-75
Online publication date: 11-Nov-2007.
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Eileen A. Hebets. (2007) SUBADULT FEMALE EXPERIENCE DOES NOT INFLUENCE SPECIES RECOGNITION IN THE WOLF SPIDER SCHIZOCOSA UETZI STRATTON 1997. Journal of Arachnology 35:1, 1-10
Online publication date: 1-Apr-2007.

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Volker W. Framenau, Eileen A. Hebets. (2007) A REVIEW OF LEG ORNAMENTATION IN MALE WOLF SPIDERS, WITH THE DESCRIPTION OF A NEW SPECIES FROM AUSTRALIA, ARTORIA SCHIZOCOIDES (ARANEAE, LYCOSIDAE). Journal of Arachnology 35:1, 89-101
Online publication date: 1-Apr-2007.

Abstract & References : Full Text : PDF (545 KB) : Rights & Permissions 

Phillip W. Taylor, J. Andrew Roberts, George W. Uetz. (2006) MATING IN THE ABSENCE OF VISUAL CUES BY SCHIZOCOSA OCREATA (HENTZ 1844) WOLF SPIDERS (ARANEAE, LYCOSIDAE). Journal of Arachnology 34:3, 501-505
Online publication date: 1-Dec-2006.

Abstract & References : Full Text : PDF (55 KB) : Rights & Permissions 

 
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