1998. The Journal of Arachnology 26 :244-246

RESEARCH NOTE THE COURTSHIP OF A KANSAS POPULATION OF BOREALIS (ARANEAE, SALTICIDAE)

The coecatus group of the jumping than the other four species, or H. borealis genus Habronattus consists of 23 described from Michigan. species, all found in the Western Hemisphere Live specimens of H. borealis were col- (Griswold 1987) . The structure of the palpi in lected during May 1990, 1991 and late April the male is generally diagnostic (see Griswold and May in 1992 and 1993, at the University 1987, figs. 187-188). The epigynum of the fe- of Kansas, Lawrence, Douglas County, Kan- male includes a central, elongated bell-like sas. These were maintained in vials and small structure (see Griswold 1987, fig . 113-115) . petri dishes at room temperature in the labo- Courtship in the coecatus group is poorly ratory, both at the University of Kansas and known, although perhaps better known than in at New Mexico State University . Specimens some other groups . Most of the species in the were fed leafhoppers and Drosophila. At New group have modifications on the third leg of Mexico State University males were placed in the male at the patella-tibia junction, and these plastic petri dishes first, followed by the fe- are displayed to the female during courtship male. This procedure was established as stan- (Griswold 1976 ; Richman 1982 ; Cutler 1988) . dard because some salticid females (although Habronattus Of the five species previously observed, H. not usually females) may attack males as prey if they are placed in the dish coecatus (Hentz 1846), H. borealis (Banks first. Observations were then made directly, 1895), H. brunneus (Peckham & Peckham and courtship details were recorded by notes 1900), H. captiosus (Gertsch 1934) and H. made during the observations . A total of 25 pyrrithrix (Chamberlin 1924), several usually males was observed in courtship with 19 vir- have relatively long courtships, with periods gin, 5 gravid and 5 penultimate females (16 H. approaching 30 min not uncommon with males, 13 virgin, 5 gravid and 4 penultimate brunneus H. pyrrithrix. and During this time females observed at Lawrence and 9 males, 6 the male crouches low with his front legs virgin females and 1 penultimate female at raised and the palpi lowered . The third legs Las Cruces). The were all preserved are raised and lowered, usually alternately ; the and voucher specimens deposited at the patellae are rotated in and out and are nearly American Museum of Natural History (New touching each other at the beginning and end York), the Florida State Collection of Arthro- of the sequence. Because H. borealis lacks pods (Gainesville) and at the Mu- any modification of the third leg we thought seum, New Mexico State University (Las Cru- that the courtship might be different from the ces) . A video film of Habronattus borealis other species . Maddison & Stratton (1988) in- courtship made by Wayne Maddison and Gail dicated that specimens of H. borealis from Stratton using specimens collected in Michi- Michigan did have a courtship more typical of gan was analyzed and compared with our ob- the species group, but also would twitch the servations of this species from Kansas . abdomen down and up during courtship, pro- Specimens of H. borealis from Franconia, ducing a buzzing or purring sound below 500 Grafton County, New Hampshire (type series Hz. They also noted an alternate shuffling of in Museum of Comparative Zoology) ; Bergen the left and right third legs . However, our ob- County, New Jersey ; Suffolk County, New servations on specimens of H. borealis from York; Berrien County and Emmet County, Kansas seem to confirm the hypothesis that Michigan; and Niagara County, Ontario, Can- members of this population have a much fast- ada, were compared morphologically with er courtship with much less embellishment specimens from Douglas County, Kansas 244

RICHMAN & CUTLER-COURTSHIP OF HABRONAITUS 245

Epigyna of representative females from the Table 1 .-Summary of courtship duration times Kansas population were removed and exam- (in seconds) for 33 trials of Habronattus borealis ined from both ventral and dorsal from Kansas. See text for details of type I and type aspects. All appeared to belong to the same II courtship . species based on morphology . H. bo- Type I No Other midwestern USA records of Type I Type II and H display realis (Bruce Cutler Collection) include IL- LINOIS : 1?, Cook County ; KANSAS : 2? Percent of total 42% 18% 18% 21% (penultimate) & 2d (males matured in labo- Time 5-30 10-30 15-45 - ratory), Chautauqua County ; 26 & 2Y, Cof- fey County ; 16, Wabaunsee County ; and 1d, Anderson County. Also 1 d & 3 ? were ex- There seemed to be no opisthosomal bobbing amined from Morris County. NEBRASKA : as reported by Maddison & Stratton (1988) for 3? & 2d (penultimate), Saunders County . Michigan H. borealis. Later attempts to get Observed courtship displays were very this same male to repeat his courtship with short, usually over in 30-45 sec, often in two other females resulted in the same se- shorter time. Only two courtships resulted in quence as seen in earlier courtships ; i.e., he mating (one observed in Lawrence and one in jumped at the apparently very resistant, but Las Cruces), as the females, even when virgin, virgin, females . A complete mating was also appeared to be highly resistant to male ad- observed at Lawrence in 1993, but in this case vances . Males often started tracking females no courtship was observed at all . The male before they saw them, sometimes drumming over a period of about 1 min slowly ap- palpi on areas where females had been . Upon proached the female from the rear, climbed on seeing the female, a male would raise its front top with no interference from the female. Af- legs, spread them 45°, elevate palpi about 45° ter 30 sec the male turned around to face the and do a few (up to 3-4) brief zigzags, ad- rear of the female, tilted the female opistho- vancing toward the female, while moving the soma, and inserted his right embolus into the palpi up and down alternately. The female female epigynum. After 40 min, the male would typically raise her front legs as well . In switched sides using his left palpus . After an- all but two of the encounters, the male then other 50 min, the female became active and jumped at the female ; or, if the female ad- the male shifted to full dorsal rear-facing po- vanced toward the male, he retreated . The first sition and released the female opisthosoma . observations made us think that this might be The female moved or ran actively for 30 min, accidental, the male mistaking the female for after which they broke apart . At least one fe- prey and becoming confused. However, in one male laid eggs after mating . One egg sac with trial the male went further. In this case the 15 eggs was produced on 22 July 1993 . Thir- male turned upside down after jumping on the teen spiderlings hatched from this clutch on female and attempted to go under her from the 24 August 1993, and a second egg sac with 9 front, with one palpus extended toward her eggs was produced 26 August 1996 . It is not epigynum. The female was able to push him known whether the second egg sac hatched. off and retreat at this point. One male (col- A summary of courtships observed is pre- lected 30 April 1993 and molted to maturity sented in Table 1 . As in the observations de- 10 May 1993) mated successfully at NMSU scribed above, courtship was usually minimal. on 29 May 1993 with a female collected 4 Type I courtship is initiated when the male is May 1993 and matured 15 May 1993 . This about 3 cm from the female, male zigzags, courtship was also very short, less than 30 sec first leg raised about 45° and may be waved, in duration. In this case the female lowered palpi splayed to side and waving. Type II dis- her cephalothorax and allowed the male to play initiated when male is about 1 cm from climb over her and insert his palpus . The male female, first legs raised at right angle to body, alternately inserted his left palpus into the fe- held somewhat forward, tarsi flicking down• male's epigynum (2 min), shifted to the right periodically, palpi elevated slightly and third (33 min) and then to the left (32 min). He then legs may be shuffled . held on for another 4 min, as the female slow- The series of courtships filmed by Maddi- ly turned. Finally they separated after 71 min . son & Stratton had a few early movements in 246 THE JOURNAL OF ARACHNOLOGY common with the Kansas population ; the front to possible predators or parasites while court- legs spread and the palpi raised. However, the ing an apparently very resistant female? The courtship continued and in at least one in- question is a difficult one to answer and re- stance, prior to a male mounting a chilled or quires much more research . dead female, the male raised and lowered his ACKNOWLEDGMENTS third legs alternately, much as in other mem- bers of the species group. Maddison (pers . We thank Wayne Maddison of the Univer- comm.) also observed a few courtships in the sity of Arizona and Gail Stratton of the Uni- population of H. borealis from the Boston versity of Mississippi for the loan of a video Mountains of Arkansas, as well as at least one tape of the Northeastern form of H. borealis. courtship using a male from Kansas. He noted We also thank Herbert W. Levi of the Museum that type I courtship or type II courtship might of Comparative Zoology, Harvard University, be used, but he never saw both used by the for the loan of the type specimens and repre- sentatives of northeastern U . populations same individual in the same courtship display . .S As in our observations, type I was used more and Mary Ellen Dix, U .S .D .A . Forest Service, than type II . Lincoln, Nebraska for the Nebraska specimen . The Michigan males seem to have a court- Wayne Maddison read the manuscript and of- ship that is intermediate between the Kansas fered several very helpful suggestions and corrections . Two anonymous reviewers also population and other members of the species improved the manuscript considerably. group . The male bobs his palpi in unison and does use his third pair of legs in the display, Lll'hRATURE CITED despite the fact that there is no special orna- Cutler, B . 1988 . Courtship behavior in Habronat- mentation on the tibiae or patellae . As far as tus captiosus (Araneae : Salticidae) . Great Lakes we can ascertain, the Kansas population has Entomol ., 21 :129-131 . dispensed with this movement entirely . Even Griswold, C.E . 1976 . Biosystematics of Habron- so, the Michigan courtship may be generally attus in California. M .S . Thesis, Univ. Califor- faster than reported in published records of the nia, Berkeley, 187 pp . courtships of other members of the species Griswold, C .E . 1987 . A revision of the genus Habronattus FO .P. Cambridge group. It was difficult to be sure of this as the (Araneae: Salticidae), with phenetic and cladistic female filmed by Maddison & Stratton was analyses . Univ . California Pub . Entomol ., 107:1- apparently chilled, or dead. 344 . Females of other species of Habronattus, Maddison, WP & G.E . Stratton . 1988 . Sound pro- including the members of the coecatus group duction and associated morphology in male in which courtship is known, have also been jumping spiders of the Habronattus agilis group observed to be highly "resistant" to mating, (Araneae: Salticidae) . J. Arachnol., 16 :199-211 . even when virgin (Griswold 1976 ; Richman Richman, D .B . 1982 . Epigamic display in jumping 1982) . In these cases, however, the males con- spiders (Araneae, Salticidae) and its use in sys- tematics . J . Arachnol ., 10:47-67 . tinued courting for as much as a half-hour. The passivity of one female during one suc- David B. Richman : Dept. of Entomology, cessful mating may point to a narrow window Plant Pathology and Weed Science, New of physiological "readiness" in the Kansas Mexico State University, Las Cruces, New populations. Even so, there was almost no Mexico 88003 USA courtship on the part of the male . The question now arises as to why the courtship of the Kan- Bruce Cutler : Electron Microscopy Labo- sas populations has deviated so much from ratory and Department of Entomology, Uni- those of other members of the species group . versity of Kansas, Lawrence, Kansas 66045 On the other hand, why have at least some USA other members of the species group evolved such time-consuming and complicated court- Manuscript received 11 April 1997, revised 30 June ships? Why spend up to a half-hour exposed 1997.