Local mate competition in the solitary parasitoid wasp <Emphasis Type="Italic">Ooencyrtus kuvanae</Emphasis>

Local mate competition (LMC) occurs when brothers compete with each other for mating opportunities, resulting in selection for a female-biased sex ratio within local groups. If multiple females oviposit in the same patch, their sons compete for mating opportunities with non-brothers. Females, in the presence of other females, should thus produce relatively more sons. Sex ratio theory also predicts a more female-biased sex ratio when ovipositing females are genetically related, and sex-ratio responses to foundress size if it differentially affects fitness gains from sons versus daughters. The mating system of the parasitoid wasp Ooencyrtus kuvanae meets assumptions of LMC. Females insert a single egg into each accessible egg of gypsy moth, Lymantria dispar, host egg masses. Wasps complete development inside host eggs and emerge en masse, as sexually mature adults, resulting in intense competition among brothers. We tested the hypothesis that O. kuvanae exhibits LMC by manipulating the number of wasp foundresses on egg masses with identical numbers of eggs. As predicted by LMC theory, with increasing numbers of wasp foundresses on an egg mass, the proportions of emerging sons increased. In contrast, the presence of a sibling compared to a non-sibling female during oviposition, or the size of a female, did not affect the number or sex ratio of offspring produced. The O. kuvanae system differs from others in that larvae do not compete for local resources and thus do not distort the sex ratio in favor of sons. With no resource competition among O. kuvanae larvae, the sex ratio of emergent son and daughter wasps is due entirely to the sex allocation by ovipositing wasp foundresses on host egg masses.     

Synergistic effects of ultraviolet radiation and conditions at low tide on egg masses of limpets (<Emphasis Type="Italic">Benhamina obliquata</Emphasis> and <Emphasis Type="Italic">Siphonaria australis</Emphasis>) in New Zealand

The effects of ultraviolet radiation (UVR), desiccation and conditions in tidal pools on embryonic survival were examined for two common pulmonate limpets that lay intertidal benthic egg masses on rocky shores in New Zealand: Benhamina obliquata and Siphonaria australis. Field surveys and manipulative experiments were conducted between December 2006 and September 2007 in the Wellington region of New Zealand (41°17′S, 174°47′E). Egg mass deposition sites in the field were species-specific: B. obliquata deposited eggs primarily in shaded crevices, whereas S. australis predominantly deposited egg masses in the sun and in tidal pools. For both species, however, embryonic mortality was greater in egg masses that had been in full sun compared to shade. For S. australis, there was also high mortality in egg masses in tidal pools or desiccated compared to those that remained submerged in flowing seawater at low tide. In outdoor experiments, embryonic mortality was also always greatest for egg masses exposed to full sun, and lowest for those in shaded treatments. Mortality was also higher if egg masses were in simulated tidal pools, and for S. australis, if desiccated, compared to those submerged in flowing seawater. Periods of particularly sunny conditions with high temperatures also resulted in higher overall mortality. Finally, egg masses of both species that were initially deposited in the shade had greater mortality in response to subsequent UV exposure compared to egg masses initially deposited in full sun. Results from this study suggest that the egg masses of these two species are highly vulnerable to UVR, as well as other intertidal stressors. Embryos of both of these species may be at risk of high mortality particularly during summer when extreme conditions of UV intensity and high temperature coincide with low tide cycles. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Two modes of development in the morphospecies Crepidula dilatata (Gastropoda: Calyptraeidae) from Southern Chile

In Chinquihue Bay, a sheltered locality in Southern Chile, female Crepidula dilatata Lamarck brood egg masses which differ in intracapsular development. In some, all eggs develop and are hatched as free veliger larvae (indirect development). In others, only some of the eggs develop and hatch as young adults (direct development), the rest being consumed as nurse eggs. Two possible interpretations are considered: intrapopulation variation of developmental pattern in a species, and the coexistence of two sibling species. Stages of the intracapsular development are described and illustrated. Fecundity varies according to size of the female. For females brooding eggs with indirect development it is estimated at between 3840 and 85575 embryos per spawn; for females with eggs undergoing direct development, embryo production per egg mass is estimated at between 70 and 812. A comparison is made between females brooding the two kinds of egg masses. Minor differences exist in adult coloration, shape of egg capsules, egg diameters, spawning season and their abundance and distribution in the intertidal. Brooding females with eggs displaying indirect development are significantly larger than those with eggs of direct development. The latter females form chains subtidally but not intertidally as do females bearing eggs with indirect development. On these females with pelagic larvae, chains frequently include sedentary males which reach large sizes. On basal females with direct development, chains contain mostly other females, suggesting that matings are temporary and by errant males. These results are compared with existing information in the literature for other species both in this genus and in other marine invertebrates. It is concluded that two sympatric sibling species with different modes of development are present in the morphospecies C. dilatata Lamarck in Southern Chile.Devoted to Dr. E.F. Kilian, Zoologisches Institut der Universität Gießen, FRG.     

Reproductive biology of the spiny lobster, <Emphasis Type="Italic">Panulirus Penicillatus</Emphasis>, in the southeastern coastal waters off Taiwan

The reproductive biology of spiny lobster, Panulirus penicillatus, was studied based on 2,068 lobsters, ranging from 34.28 to 131.60 mm carapace length (CL), sampled in Taitung coastal waters from September 2003 to December 2004. The overall sex ratio approximated 1:1 (χ² = 0.02, P > 0.05), but the monthly sex ratios in 2004 showed significant differences and males were predominant in sizes larger than 80 mm CL. Reproductive activity, assessed using histology, a gonadosomatic index and percentage of ovigerous females, indicated that the mature females could be found in every month and that the major spawning occurred from May to September. The presence of re-developing/re-ripe ovaries by month and size-specific spawning time suggest that larger mature females (>60 mm CL) spawn at least three times a year while smaller new mature females spawn at least once a year. For females, the estimated sizes at 50% physiological and functional maturity were (mean ± SE) 56.46 ± 0.56 mm CL and 66.63 ± 1.07 mm CL. The estimated sizes at functional maturity were between 72 and 74 mm CL for males. The number of eggs per spawning event (brood size, BS) was related to CL by the equation Y _BS = 2.4 × 10^-3CL^4.18 (r ² = 0.902, n = 12). Female lobsters with CL ranging from 60 to 80 mm made the greatest contributions to egg production because of their high brood size and active reproductive activity. A minimum legal size should be established for the fishery to protect egg production potential of lobster population in the southeastern coastal waters off Taiwan.     

Reproduction,development and population biology of the Caribbean ophiuroidOphionereis olivacea,a protandric hermaphrodite that broods its young

Reproduction and development of the ophiuroidOphionereis olivacea H. L. Clark, a species that broods its young, were investigated in Florida from January through December 1985 and in Belize in June 1985 and April 1986.O. olivacea has a maximum disc diameter of 5.2 mm and is a protandric hermaphrodite. Sex reversal occurs at disc diameters ranging between 2.2 and 4.0 mm; it has 400 µm diameter oocytes and is ovoviviparous. As expected from an obligate out-crosser, this ophiuroid has a similar allocation to male and female reproduction, in terms of gonad volume. Reproduction of the Florida population was monitored for one year.O. olivacea is a sequential brooder and incubates its young during the coldest months of the year. The females produce a single clutch of juveniles each year. Spawning starts in November and peaks in January and February. The ova are spawned synchronously from all the ovaries of each female into the bursae and are retained there. Fertilization is accomplished by intake of sperm into the female bursae.O. olivacea develops through a ciliated embryonic stage that appears to be a modified vitellaria larva lacking ciliary bands. These modified larvae swim in seawater in the bursae. Direct development in this species has resulted from heterochrony in the loss of ophiopluteus features. The number of brooding females in the population increases in November and reaches a maximum in March. JuvenileO. olivacea emerge from the bursae at a disc diameter of 480 µm and each arm has three segments. Recruitment was first detected in April, and juveniles were particularly abundant in June. Small size inO. olivacea is associated with paedomorphic heterochrony with precocious maturation of the males at a disc diameter of 1.5 mm, ~2 mo after taking up their free existence. Brooding as a life history mode inO. olivacea is examined and the adaptations of this species to its warm-water habitats are discussed.     

Paternal care and egg survival both increase with clutch size in the fathead minnow,Pimephales promelas

Summary In some species of fishes with paternal care, females prefer to spawn with males who are already defeding eggs; moreover, in many species, paternal care increases with the number of eggs that a male is defending. If egg survival depends on the level of paternal care, and is largely independent of egg number, then egg survival should increase with clutch size. This result would provide a potential adaptive mechanism for female preference for males with eggs. I examined the effects of clutch size on paternal care and egg survival in the fathead minnow, Pimephales promelas, and found that both increased with male clutch size.     

Fecundity of a euphasiid crustacean,Nematoscelis difficilis,in the North Pacific Ocean

The number, size, and carbon and nitrogen contents of eggs in the pouches of a euphausiid crustacean, Nematoscelis difficilis Hansen, were examined. A clear linear relationship exists between the number of eggs in the pouch and the body weight of the maternal euphausiid. The eggs are not spherical in shape, and the size of eggs is greater in larger egg masses. The carbon content of the egg masses (50.2%) is higher than that of the bodies (40.7%). The carbon: nitrogen ratio (C/N) is also higher in the egg masses than in the bodies of euphausiids. The carbon content of the eggs is equivalent to 28.4% of the body carbon, the nitrogen content to 19.2% of the body nitrogen.     

Developmental types of shallow-water asteroids of McMurdo Sound,Antarctica

The mode of development was ascertained for 14 of the 16 species of sea stars known to occur in shallow waters of McMurdo Sound, Antarctica (77°51S; 166°40E). The species were collected between September 1984 and December 1985. Females of three species,Odontaster validus, O. meridionalis andPorania antarctica, spawn small to moderate eggs (0.17 to 0.55 mm), have a high fecundity, and produce feeding larvae. Females of an undescribedPorania species spawn a few eggs (150 to 310) that are 0.55 mm in diameter and develop into demersal non-feeding larvae. Females ofDiplasterias brucei andNotasterias armata produce a few (<300) large eggs (2.8 to 3.5 mm) and brood their young. Females of the remaining eight species have moderate fecundity and produce pelagic non-feeding larvae, as determined from egg type (buoyant, 0.54 to 1.28 mm diam) and direct observations of spawning and development. The high incidence (11 out of 14 species; 79%) of non-feeding development is consistent with predictions that environmental conditions in high-latitude regions are unfavorable for planktotrophic development. Nonetheless, most of the species surveyed (11 out of 14) had pelagic larvae, which contradicts inferences of unusual selection for benthic development in the Antarctic.     

Food access, brood size and filial cannibalism in the fantail darter, Etheostoma flabellare

We compared the occurrence of filial cannibalism in fed and starved male fantail darters (Etheostoma flabellare). All males in the experiment consumed eggs, and 56% ate all of their eggs. A male's initial body condition did not explain the number of eggs that he ate. Neither did non-fed males eat more eggs than fed males. Fed males were able to maintain better body condition during the experiment, but the change in body condition also depended on the number of eggs eaten. Thus, males who ate more eggs were able to maintain better body condition.The most important determinant of whether or not a male ate all of his eggs was his initial egg number. Males with small egg masses ate all of their eggs whereas males with large egg masses were only partial cannibals. There was, however, no difference in the total number of eggs eaten by total and partial cannibals. We conclude that eggs are only partially eaten for energetic reasons. We also suggest that small egg masses are completely consumed because the costs of caring for a small egg mass may exceed the expected reproductive benefits of a small egg mass. Received: 26 January 1996 / Accepted after revision: 2 November 1996     

Does spawning behavior minimize exposure to environmental stressors for encapsulated gastropod embryos on rocky shores?

Adults of motile intertidal invertebrates are able to seek shelter to avoid environmental stress associated with low tides, but embryos within egg masses are effectively sessile for the duration of their encapsulation. Gastropod egg masses from 34 taxa on two rocky shores in SE Australia (34°37′08″S, 150°92′03″E and 34°35′45″S, 150°53′20″E) were surveyed over 2 years (June 2002–May 2004) to test the hypothesis that eggs are deposited in patterns that minimize exposure to environmental stress. Egg masses were expected to be predominantly deposited in shaded habitats not prone to environmental extremes. It was also anticipated that the deposition of egg masses in habitats exposed to UVR, desiccation, and/or extremes in temperature would occur when exposure to these abiotic factors was minimized. Among the taxa investigated, only four species spawned in full sun (Bembicium nanum, Nerita morio, Siphonaria zelandica and S. denticulata). Summer had the highest UVR index, water temperature, and air temperature as well as the lowest daytime tides. Univariate and multivariate analyses confirmed that egg mass abundance was highest during summer, with no change in egg mass size. This study shows that those species depositing egg masses on the surfaces of rock platforms do not adjust the seasonal timing or macrohabitat location of their spawning to avoid physiologically stressful conditions, particularly UVR. Alternate reasons for the evolution of egg mass deposition behavior in apparently sub-optimal habitats are discussed, and it is almost certainly the complex interplay of a variety of highly species-specific factors that is responsible for the patterns observed.     

Evolution of egg size and fertilisation efficiency in sea stars: large eggs are not fertilised more readily than small eggs in the genus<Emphasis Type="Italic"> Patiriella</Emphasis> (Echinodermata: Asteroidea)

Fertilisation kinetics theory suggests that, when sperm are limiting, the larger eggs of broadcast-spawning marine organisms ought to be fertilised more frequently than smaller eggs, because they provide a bigger target for searching sperm. Whilst this effect has been demonstrated within species, it is not known if this pattern holds among species. We tested whether a large difference in egg size between congeneric seastars with contrasting planktotrophic and lecithotrophic modes of development results in differences in the likelihood of eggs being fertilised in sperm-limiting situations. Measurement of egg sizes and sperm swimming speeds led to the prediction that the sperm–egg collision rate constant for Patiriella calcar (420-µm-diameter egg) should be nine times greater than for P. regularis (140-µm-diameter egg). Although the eggs of P. calcar should be fertilised at greater rates in low sperm concentrations, they were not. When gametes were allowed to mix for 10 s, the hypothesis that P. calcar eggs required less sperm than P. regularis to ensure 50% of eggs were fertilised was rejected. When gametes were mixed for 5 min, P. regularis eggs were more frequently fertilised, but the difference was not statistically significant. We conclude there must be a difference between these species in the likelihood that when a sperm finds a conspecific egg it can successfully fertilise. This apparent uncoupling of egg size and likelihood of fertilisation suggests that fertilisation is not a major constraint on the evolution of egg size in these seastars.Communicated by G.F. Humphrey, Sydney     

Reproductive cycle and maternal effects on offspring size and number in the neogastropod Buccinum undatum (L.)

This paper examines relationships between maternal phenotype and offspring phenotype in the commercially exploited marine neogastropod Buccinum undatum (L.). The study also aimed to estimate the effects of female size on subsequent recruitment, and to determine the reproductive cycle of B. undatum in Swedish waters. The evaluation of gonad development in field-sampled animals from August 1998 to February 2000 revealed a definite annual reproductive cycle with a single major egg-laying period in the autumn. In addition, adult whelks of varying sizes from two locations and in two consecutive years were kept in aquaria to study the influence of female size on fecundity. Twenty-seven females deposited eggs in the laboratory. Offspring quality, measured as egg and hatchling size, was independent of female size, although larger females produced more eggs and hatchlings than smaller females, and thus probably account for a disproportionately large share of potential recruitment; the relative fecundity (number of eggs or hatchlings per gram of body weight) was constant with female size. I found no evidence for a longer time interval between successive egg-layings in larger females, i.e. mature females of all sizes probably spawn equally often. The present results therefore indicate that spawning-stock biomass is probably a better measure of spawning-stock size than the number of mature individuals in the population. Effects of an individual female's characteristics on stock productivity have largely been overlooked by fishery managers but could help to explain the weakness often found in stock-recruitment relationships.     

Flexible reproductive strategies in tropical and temperate Sepioteuthis squids

A major difficulty confronting the determination of cephalopod reproductive life history is assessing over what portion of the life span an individual is reproductively mature and actively depositing eggs. This paper assesses the potential of the tropical Sepioteuthis lessoniana and two genetic types of the temperate Sepioteuthis australis, to spawn multiple batches of eggs at discrete times throughout the adult life span. This is achieved by histological examination of the ovarian gametogenic cycle and detailed morphological assessments of the reproductive system, in conjunction with other biological information. The genetic type of S. australis found at the northern limits of its Australian distribution showed evidence of a high correlation between body size and quantity of mature eggs, suggesting that eggs may be accumulating to be laid in a single batch. Although maturation was also a size-related process in S. lessoniana and Tasmanian S. australis, oviduct size was not correlated with body weight in mature females, which is indicative of multiple spawning. Further supporting evidence includes relatively low gonadosomatic indices, the heavier weight of the ovary relative to the oviduct, and the feeding activity of mature animals. Mature S. lessoniana and S. australis individuals were present at each location over very wide age and size ranges. In Tasmanian waters, there were distinct seasonal differences in the reproductive biology of S. australis. Summer-caught individuals had much higher gonadosomatic indices and may have been laying larger batches of eggs compared with winter-caught individuals. Summer-caught females also showed a negative correlation between egg size and egg number within the oviduct, suggesting that some individuals were producing fewer, larger eggs and others many smaller eggs. Evidence suggests that considerable flexibility is inherent in the reproductive strategy of both S. lessoniana and S. australis. Received: 11 May 2000 / Accepted: 12 September 2000     

Possible mutualism between females of the subsocial membracid Polyglypta dispar (Homoptera)

Summary Female Polyglypta dispar membracids facultatively guard egg masses or oviposit into masses guarded by other females. Defending females repel at least some enemies of both eggs and nymphs. Eggs of guarding females may be partially protected from parasitism by the presence of additional eggs laid by other females. Females sometimes desert egg masses or groups of nymphs, and undefended masses are sometimes adopted by other females. Some individuals remain on or near the plant where they grew up, and littermates sometimes oviposit into the same egg mass. Some copulating pairs are littermates, while others are not. Females with larger numbers of mature and nearly mature eggs in their ovaries are less likely to defend egg masses, and females desert smaller groups of nymphs more often.     

Reproductive biology of deep-water calanoid copepods from the Arctic Ocean

Reproductive modes and egg production were studied in 15 species of meso- and bathypelagic copepods from nine Calanoida families in the Arctic Ocean. During shipboard incubation, females of seven species released eggs freely into the water and females of three species produced membrane-bound egg sacs. One species, Aetideopsis rostrata, produced a mass of eggs looking like an egg sac, but the “sac” lacked a membrane and disintegrated within 2 h. Females of four additional species were encountered with membrane-bound egg sacs in the preserved samples. In most families, only one reproductive mode, either egg-carrying (the Euchaetidae, Augaptilidae), or broadcast spawning (the Heterorhabdidae, Spinocalanidae, Scolecitrichidae, Tharybidae, Bathypontiidae) was observed. In contrast, different genera of the Aetideidae family demonstrated different reproductive modes, with broadcast spawning predominant in the benthopelagic species, and both broadcast spawning and egg-brooding in the planktonic species. Clutch size and egg diameter varied widely between species in both broadcast spawners and egg-brooders. In broadcast spawners, the clutch size varied from 1 to 95 eggs female⁻¹, while the average egg diameter ranged from 152 to 440 μm. The clutch size for egg brooders varied between 3 and 82, while average egg diameter varied from 258 to 732 μm. Deep-water broadcast spawners produced much larger eggs compared to surface-dwelling broadcast spawning species. This larger egg size may result in a reduction, or elimination, of feeding during naupliar stages, thereby improving the survival potential of deep-water species.     

Size at maturity,fecundity and reproductive potential of a protected population of the spiny lobster<Emphasis Type="Italic"> Palinurus elephas</Emphasis> (Fabricius, 1787) from the western Mediterranean

Protected lobster populations are expected to contribute to the replenishment of fished populations through increased egg production. We studied the reproductive biology and egg production potential of a population of the spiny lobster Palinurus elephas protected from fishing since 1990 in the Columbretes Islands Marine Reserve (western Mediterranean). An index of spawning potential was derived to compare egg production potential in the Reserve and in western Mediterranean exploited populations. Females' physiological maturity (ability to reproduce) and functional maturity (ability to mate and bear eggs) occurred at a carapace length (CL) of 76–77 mm. Males' physiological maturity occurred at a slightly larger size, 82.5 mm CL. In the Reserve, P. elephas' individual fecundity increases linearly with body size up to the females' maximum size, although maximum reproductive yield (eggs per body gram) was reached at intermediate sizes. Size-specific fecundity in the protected population was similar to that of lightly fished populations off Ireland and greater than that of western Mediterranean exploited populations. The female size class of 105–110 mm CL contributed most to egg production in the protected population and is well above the minimum landing size (MLS) for western Mediterranean fisheries. Newly mature females (below MLS) generate a very small fraction (1%) of the egg production from the Reserve. Given the pattern of exploitation in western Mediterranean fisheries, egg production potential depends more on the quantity than on the mean size and fecundity of the available females. The role of the greater availability of large males for mating in unfished populations is discussed in terms of the females' individual fecundity and mating success.     

Reproductive biology in two species of deep-sea squids

Deep-sea squids, Moroteuthis ingens and Gonatus antarcticus, were collected in the slope waters off the Falkland Islands and their reproductive systems preserved and investigated onshore. Changes in oocyte length-frequencies at maturation and spawning, and their fecundity were studied. These squids, as well as many other species, are characterised by a synchronous oocyte growth and ovulation. Oviducts are not used for ripe egg accumulation and consequently the universal scale of Lipinski (1979) cannot be applied to assign female maturity. M. ingens spawns near the bottom; its fecundity is 168–297 thousand eggs. Maximum egg size is 1.8–2.7 mm. G. antarcticus spawns midwater; its fecundity is 10–25 thousand eggs. Egg size is 3.2–3.3 mm. In M. ingens spawning takes place in the austral autumn and winter, in G. antarcticus—in austral winter. Our data and the literature data show that the so-called “synchronous ovulation” probably occurs in all deepwater squids. This pattern is very rare among fish, but is quite common among benthic octopods that brood their egg masses.     

Laboratory observations on spawning,brood size and egg hatchability of the Antarctic krill Euphausia superba from Prydz Bay,Antarctica

Forty-three female Euphausia superba were collected from three sites in Prydz Bay, Antarctica in January 1985. Most gravid females took 3 to 14 d to spawn after capture and some less mature females released eggs after 21 d. Most females spawned only once; overall mean brood size was 2 809 eggs and mean hatchability for eggs from the premoult females was 69.7%. For seven females spawning a second time, overall mean brood size was 597 eggs. Mean hatchability for second-spawn eggs from five premoult females was 67.1%. Possible origins of the second-spawned eggs were small numbers of matured eggs left in the ovaries after the first spawning or eggs developed from those of peripheral nucleolus and vitellogenic stages. Throughout the 8 wk of observations, there was no apparent development of eggs in females which had released all their eggs at one spawning.     

Nest-size variation reflecting anti-predator strategies in social spider mites of<Emphasis Type="Italic"> Stigmaeopsis</Emphasis> (Acari: Tetranychidae)

The social spider mites (Acari: Tetranychidae) of Stigmaeopsis weave dense nests on the underside of host leaves. Four species occur on the leaves of bamboo in Japan: Stigmaeopsis longus, S. celarius, S. takahashii and S. saharai. We initially reconfirmed the occurrence of distinct variation in nest size among the species. Based on the hypothesis that this variation plays a role in protecting the spider mites from predators, we looked at the behavior of the natural enemies that occur on the host plants along with members of Stigmaeopsis. We found considerable variation in the ability of nests to protect the spider-mite eggs. The smallest nests protected the eggs against three predators, whereas the largest nests protected the eggs against only one predator species. So, decreases in nest size increased egg defense. Thus we concluded that nest-size variation reflects a strategy for reducing predation.Communicated by D. Gwynne     

“Egg dumping” in lace bugs (Gargaphia solani,Hemiptera: Tingidae)

Summary Data are presented which document that females of the subsocial lace bug, Gargaphia solani lay eggs in the nests of conspecifics whenever the opportunity arises. Because of an inverse relationship between time invested in maternal care and fecundity, maternal behavior in G. solani is an ecologically expensive trait that is adaptive only in the face of heavy predation on eggs and nymphs. By facultatively utilizing the maternal defensive behavior of conspecifics, it is possible for egg donors to protect their progeny from predators without limiting fecundity. Whenever possible, females oviposit in recently established egg masses of conspecifics. While guarding their own eggs, egg recipients inadvertently protect the eggs of egg donors. Egg donors need not establish and guard their own masses as long as there are females with egg masses in the vicinity. Instead, egg donors are free to lay as many eggs as physiologically possible by avoiding long periods of maternal care.Published with the approval of the Director of the Delaware Agricultural Experiment Station as Miscellaneous Paper No. 1048, Contribution No. 544 of the Department of Entomology and Applied Ecology, University of Delaware, Newark, Delaware, USA