Journal of lnsect Behavior, Vol. 5, No.

6, 1992

Short Communication

Transfer of a Larval Mantispid During Copulation of Its Spider Host

Sonja J. Scheffert'z
Accepted May 18, 1992; revised May 18, 1992 KEY WORDS: Climaciella brunnea; rnantispids; Schizocosa species behavior; spider parasites.

INTRODUCTION Within the Mantispidae (Neuroptera), larvae in the subfamily Mantispinae feed on the eggs of spiders. Two methods are known by which larval mantispids gain access to spider eggs. In some species, the larva actively searches out a spider egg sac, penetrates the silk covering, and enters the sac. In other species, the larva climbs aboard a living spider and remains there until an egg sac is produced. During the construction of the egg sac the larva leaves the spider and enters the sac. Particular mantispid species may be obligate egg sac penetrators, obligate spider-boarders, or facultative penetrator/boarders, using either strategy (Redborg and MacLeod, 1983). While mantispid larvae aboard spiders are known to feed on the hemolymph of their hosts, development beyond the first instar does not proceed until feeding begins on spider eggs (Redborg and MacLeod, 1984). Because only female spiders construct egg sacs, natural selection in spiderboarding species should favor larvae that preferentially end up on female spiders. Selective boarding of female spiders has not been detected in the mantispid species studied so far (Redborg and MacLeod, 1985; S. Scheffer, personal observation). A possible alternative to selective boarding of female spiders is the transfer by mantispid larvae from male to female spiders during copulation. Transfer behavior during copulation was not found in Mantispa uhleri Banks (Redborg and MacLeod, 1985), and it has not been looked for in any other species.
~Department of Biological Science, University of Cincinnati, Cincinnati, Ohio 45221. 2Current address: Department of Ecology and Evolution, State University of New York, Stony Brook, New York 11794.
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The purpose of this study was to investigate the possibility of transfer behavior in the mantispid Climaciella brunnea (Say), an obligate spider-boarder.
METHODS C. brunnea, a mimic of Polistes wasps (Wheeler, 1889; Opler, 1981), is found from the western and central United States south to Central America (Opler, 1981). C. brunnea larvae on male spiders were obtained for this study in two ways. In the summer of 1987, a number of adult mantispids emerged from the egg sacs of laboratory-mated female Schizocosa ocreata (Hentz) (Lycosidae), which had been collected as subadults at the Cincinnati Nature Center (Hamilton County, Ohio). These mantispids were allowed to mate; subsequently, eggs were laid on the wall of a plexiglass container by a single female. Following hatching, the mantispid larvae were allowed to board S. ocreata spiderlings, which were then reared in the laboratory until the following spring when they reached maturity. Additional male spiders (S. ocreata and S. rovneri Uetz and Dondale) carrying mantispids were collected in May of 1988 (at the Cincinnati Nature Center and in northern Kentucky, respectively). The mantispids, often positioned on the spider's pedicel or carapace (Redborg and MacLeod, 1983; S. Scheffer, personal observation), are readily evident. All mantispidbearing spiders used in this experiment carded only one mantispid. Twenty-three pairs of male and female spiders were placed individually in 15-ml plastic culture dishes and left for 3 to 4 h until copulation or, in two cases, until cannibalism occurred. One member of each pair carded a mantispid, while the other did not. Following copulation all spiders were checked for the presence of the mantispid. In no case was more than one mantispid found per pair.

RESULTS Fourteen copulations occurred in which the male was initially carrying a mantispid. In 12 instances (85.7%) the mantispid was found on the female and was missing from the male following copulation. In 2 of these 12 cases the female ate the mate before I had determined whether or not the larva had transferred during copulation. When these females had finished feeding they were checked for the presence of the larva. In both cases the mantispids had transferred successfully. Because larvae are known to transfer from a cannibalized spider to its predator [two additional cases in this study; Redborg and MacLeod (1985) with M. uhleri], it is unclear whether these two transfers occurred during copulation or cannibalism. After discarding these two cases, 10 successful transfers remain of 12 copulations (83.3 % transfer). Larval transfer was never seen when the initial carder was a female spider (n = 9).

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Larval transfer behavior was directly observed in only one case. During courtship, and at the start of copulation, the mantispid remained fairly stationary on the pedicel and posterior portion of the carapace; this is the usual condition for mantispids on spiders not engaged in mating-related activities. During copulation, the mantispid became very active, repeatedly moving rapidly around and underneath the male carapace until it finally disappeared under the male; it was later found on the female.

DISCUSSION Because larval mantispids seem not to board female spiders preferentially, a great number of larvae must board male spiders, thereby lacking direct access to eggs (which are required for development beyond the first instar). The transfer of larval mantispids from their male spider hosts to female spiders during copulation is an important strategy, allowing the larvae to position themselves to enter egg sacs. This is the first report of larval transfer behavior during host copulation in a mantispid species. The transfer behavior shown by C. brunnea was not detected in a previous study involving M. uhleri (Redborg and MacLeod, 1985). The difference in transfer behavior between C. brunnea and M. uhleri could be due to a number of factors. M. uhleri is a facultative spider-boarder; it is able to enter previously constructed egg sacs. Redborg and MacLeod (1985) suggest that, following the death of its male spider host, a larva may be free to wander in search of an egg sac (or a new host). This alternative route providing access to eggs is not open to C. brunnea, which do not recognize previously formed egg sacs or eggs as food (Redborg and MacLeod, 1983). Selection for larval transfer behavior may be more intense in a species in which successful larval development absolutely requires that a larva is carried by a mature female spider. Another factor that may influence the evolution of larval transfer behavior is the number of spider hosts used by each mantispid species. The host range of M. uhleri is very broad; it has been collected from spiders in at least eight families (Redborg and MacLeod, 1985, Table 27). In contrast, C. brunnea has been collected exclusively from spider species within the family Lycosidae (George and George, 1975; Redborg and MacLeod, 1983; LaSalle, 1986; this study). Restriction of C. brunnea to spider hosts within a single family may provide a consistent cue indicating host copulation that is not available to M. uhleri, which uses a far more diverse group of spiders as hosts. Additionally, lycosid copulation is commonly of a long duration, allowing ample time for transfer. Restriction of C. brunnea to this single spider family may have made the evolution of larval transfer behavior more likely.

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ACKNOWLEDGMENTS T h a n k s are due to Cole Gilbert for initially identifying the mantispids. George Uetz kindly provided laboratory space. D o u g F u t u y m a , Leo Shapiro, and Keith Karoly m a d e valuable c o m m e n t s on the manuscript. A Sigma Xi G r a n t - i n - A i d o f Research supported m y research on reproductive b e h a v i o r o f Schizocosa spiders from which this project sprang. The Cincinnati Nature Center allowed m e to collect spiders from their grounds.

REFERENCES

George, L. D., and George, N. L. (1975). Notes on the three hundred and fifty-eighth meeting: A new record of mantispid reared from spider. Pan-Pacif Entomol. 51: 90. LaSalle, M. W. (1986). Note on the mantispid Climaciella brunnea (Neuroptera: Mantispidae) in a coastal marsh habitat. Entomol. News 97: 7-10. Opler, P. A. (1981). Polymorphic mimicry of polistine wasps by a neotropical neuropteran. Biotropica 13: 165-176. Redborg, K. E., and MacLeod, E. G. (1983). Climaciella brunnea (Neuroptera: Mantispidae): A mantispid that obligately boards spiders. J. Nat. Hist. 17: 63-73. Redborg, K. E., and MacLeod, E. G. (1984). Maintenance feeding of first instar mantispid larvae (Neuroptera, Mantispidae) on spider (Arachnida, Araneae) hemolymph. J. Arachnol. 11: 337341. Redborg, K. E., and MacLeod, E. G. (1985). The developmental ecology of Mantispa uhleri Banks (Neuroptera: Mantispidae). III. Biol. Monogr. 53: 1-130. Wheeler, W. M. (1889). Two cases of insect mimicry. Proc. Wisc. Nat. Hist. Soc., April, pp. 217-221.