Journal of the Lepidopterists' Society 39(2), 1985, 65-84
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JOURNAL OF THE LEPIDOPTERISTS' SOCIETY Volume 39 1985 Number 2 Journal of the Lepidopterists' Society 39(2), 1985, 65-84 MAINTAINING SPECIES INTEGRITY BETWEEN SYMPATRIC POPULATIONS OF HYALOPHORA CECROPIA AND HYALOPHORA COLUMBIA (SATURNIIDAE) IN CENTRAL MICHIGAN JAMES P. TUTTLE 728 Coachman #4, Troy, Michigan 48083 ABSTRACT. The available literature suggests that Hyalophora cecropia and Hya lophora columbia have identical ethological requirements with respect to reproductive behavior and F j hybrids have often been produced in the laboratory. As a result, investi gators have long been puzzled by the rarity of natural hybrids since the two species occur sympatrically over much of the northern Great Lakes region. Field observations in Montcalm County, Michigan, identified a series of progressive barriers which operate to prevent hybridization. The prezygotic isolating mechanisms, seasonal isolation, daily allochronic flight behavior, and mechanical incompatibility re strict most of the interaction between the two species. However, when individuals oc casionally overcome those barriers and a successful hybrid mating occurs, several post zygotic isolating mechanisms are then tested. As a result of the overall effectiveness of these systems, the gene pool of each species is protected, thereby allowing the two species to co-exist. The large moths of the genus Hyalophora (Saturniidae) are repre sented by two species within the state of Michigan. The cecropia moth, Hyalophora cecropia (L.), is widely distributed, while Hyalophora co lumbia (S. I. Smith) is a much more localized species. When Moore (1955) published his checklist of the Michigan moths, he listed 18 county records for H. cecropia from such varied locales as Dickinson County in the western portion of the Upper Peninsula to metropolitan Wayne County in the southeastern portion of the Lower Peninsula. He also cited statewide adult capture dates ranging from 27 May to 5 August. l The author's recent review of additional material in public and private collections has increased to 48 the number of counties from which H. cecropia has been reported (Fig. 1). The pa- 1 The 5 August date is based upon a reared specimen probably maintained under artificial conditions. Such late dates are highly unlikely in nature. 66 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY COLUMBIA 0 CECROPIA FIG. 1. Distributional map of H. cecropia and H. columbia in Michigan. Montcalm County (darkened) is the site of the present study. rameters of the adult flight period were also expanded when wild capture dates as early as 22 May were discovered. The distribution of H. cecropia within the state is greatly enhanced by the acceptance of a wide array of larval foodplants. Tietz (1972) lists 75 specific larval food plant records, which include many of the native Michigan broadleaved trees and shrubs. This adaptability to varied plant communities is extremely important to a cosmopolitan species such as H. cecropia. Although there are still a number of counties from which H. cecropia has not been reported, this would appear to be due to a lack of col lecting. Based upon the adaptability of the species, the lack of rest ric- VOLUME 39, NUMBER 2 67 tive topographical barriers within the state and the distribution of ex isting county records, it is reasonable to assume that H. cecropia occurs in all of Michigan's 83 counties. In contrast to the large amount of data available on H. cecropia, the existing Michigan records of the distribution and flight period of H. columbia are extremely limited. Based exclusively on wild collected cocoons, Moore (1955) cited records for H. columbia from Lapeer, Montcalm, Oakland, Washtenaw, and Wayne counties. Since the pub lication of Moore's checklist there have been records from Ingham (Collins & Weast, 1961), Dickinson, Jackson, Livingston, Mecosta, Ne waygo (Ferguson, 1972), Clare, Menominee, Schoolcraft (M. C. Niel sen, in litt.), and Isabella counties (Ted Herig, pers. comm.) (Fig. 1). Most of these additional records are also based upon wild collected cocoons. In fact, prior to the present study, there were only six reports of H. columbia adults having been collected in Michigan. Three adults have been collected at light: a male collected in Menominee County on 10 June 1971; a female collected in Schoolcraft County on 21 June 1971; and a male (specimen lost) collected in Dickinson County (M. C. Niel sen, in litt.). In addition, in June 1978, single H. columbia males were attracted on each of three successive nights to a H. cecropia female in Montcalm County (Frank Hedges, pers. comm.). Unlike the widespread distribution of H. cecropia, the range of H. columbia is limited by its dependency on larch (Larix laricina) as the larval foodplant. Although there are records of H. columbia being reared in captivity on choke cherry (Prunus virginiana) (Collins & Weast, 1961) and weeping willow (Salix babylonica) (Norman Trem blay, pers. comm.), it is now generally accepted that the larvae feed exclusively on larch in nature. Many old food plant records were erroneously based on cocoons found attached to plants growing in association with larch. This is clearly reflected in the type description of H. columbia, when Smith (1863) cites "Nemopanthes canadensis and Rhodora canadensis" as larval food plants and only incidentally mentions larch as a possible alternate host. This can now be explained by the tendency of full-fed larvae to wander in search of an appropriate cocoon spinning site. In Michigan's Upper Peninsula larch abounds as a dominant tree species in conifer bog and swamp community associations (Barnes & Wagner, 1981). As a result of the widespread distribution of larch in this area, H. columbia should be expected to occur regularly. The relatively few records from this region are almost certainly due to limited collecting. The existing county records for H. columbia in Michigan's Lower 68 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY Peninsula occur as two separate pockets which include seven counties in the southeast and five counties in the central portion of the penin sula, In these areas, larch occurs only in poorly drained "kettle hole" type situations. Such bogs may be isolated by several miles of unsuitable habitat and are historically very susceptible to human influence. An examination of specimens at the University of Michigan reveals several records for H. columbia from Wayne County prior to 1900. However, drainage and the resulting development have eliminated all of the suitable habitat. As a result, there are no records of H. columbia from Wayne County in this century. H. columbia populations in Michigan's Lower Peninsula are, there fore, restricted to those undisturbed areas where larch occurs in suffi cient numbers to effectively support the insect. One such locale is the Stanton State Game Area in Montcalm County (Fig. 1). In this area, which serves as the site of the present study, H. columbia occurs sym pat ric ally with H. cecropia. Within zones of contact between closely related species such as H. cecropia and H. columbia, effective mate recognition behavior must be established if both species are to co-exist over an extended period of time (Mayr, 1970). Yet, the available literature suggests that H. cecropia and H. columbia lack such discriminatory behavior. The existing data indicate that H. cecropia and H. columbia adults emerge at the same time of the year (Moore, 1955; Collins & Weast, 1961; Ferguson, 1972). Both species also mate during the hours im mediately preceding dawn (Collins & Weast, 1961; Ferguson, 1972). In addition, even the earliest breeders were able to obtain interspe cific Hyalophora hybrids in the laboratory (Morton, 1895; Soule, 1907). Since that time, almost all possible crosses, backcrosses, and reciprocal crosses have been attempted (Sweadner, 1937; Weast, 1959; Collins & Weast, 1961; Wright, 1971; Kohalmi & Moens, 1975). Investigators thus learned that congeneric males would readily re spond to calling2 non-specific females. Sweadner (1937) effectively ex ploited this ability by using H. cecropia females to attract wild H. columbia and H. gloveri nokomis (Brodie) males in western Ontario and Manitoba. More recently, H. cecropia females have been success fully used to attract wild H. columbia males in Manitoba (Collins, 1973), Ontario (Kohalmi & Moens, 1975), Quebec (Gilles Deslisle, pers. comm.), Wisconsin (Ferge, 1983), and Michigan (Frank Hedges, pers. comm.). Yet, in spite of these apparent behavioral similarities and the relative ease with which hybridization occurs in captivity, very few valid nat- 2 Extension of the ovipositor resulting in the release of a sexual pheromone. VOLUME 39, NUMBER 2 69 ural hybrids between H. cecropia and H. columbia have been reported. The author agrees with Ferguson (1972) who states, "Careful scrutiny of the literature reveals that at least some of the records of natural hybrids may be false, especially those based on cocoons only. Free man's experience of having a normal cecropia emerge from what looked like a hybrid cocoon on larch, already noted, casts suspicion on all reports based on cocoons of intermediate appearance." Ferguson (1972) does cite two records of hybrids emerging from cocoons collected in Mecosta County, Michigan, and Fraserburg, On tario. He also mentions a wild collected hybrid male collected on 13 June in Oakland County, Michigan. The author has examined a wild male collected on 17 June 1973, in Oakland County, Michigan, which appears to reflect a hybrid background. During the present study in Montcalm County, only one of 259 Hyalophora adults examined showed any hybrid influence. In the spring of 1981, the author observed the emergence of Hya lophora adults in outdoor cages in Ann Arbor, Michigan. An extended emergence pattern was observed in the caged H. cecropia adults. A comparison with the concentrated emergence of the H. columbia adults in the adjacent cage suggested partial temporal isolation. These obser vations supplied the impetus for the present study which offers data and discusses the reproductive isolating mechanisms which allow such closely related species to co-exist.