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Photoperiodic Responses of Phenologically Aberrant Populations of Pierid Butterflies (Lepidoptera)

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Photoperiodic Responses of Phenologically Aberrant Populations of Pierid Butterflies (Lepidoptera) Great Basin Naturalist Volume 35 Number 3 Article 7 9-30-1975 Photoperiodic responses of phenologically aberrant populations of pierid butterflies (Lepidoptera) Arthur M. Shapiro University of California, Davis, California Follow this and additional works at: https://scholarsarchive.byu.edu/gbn Recommended Citation Shapiro, Arthur M. (1975) "Photoperiodic responses of phenologically aberrant populations of pierid butterflies (Lepidoptera)," Great Basin Naturalist: Vol. 35 : No. 3 , Article 7. Available at: https://scholarsarchive.byu.edu/gbn/vol35/iss3/7 This Article is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Great Basin Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. PHOTOPERIODIC RESPONSES OF PHENOLOGICALLY ABERRANT POPULATIONS OF PIERID BUTTERFLIES (LEPIDOPTERA) Arthur M. Shapiro' Abstr.'^CT.— T^vo local pierid populations in western Norlh Ainerit.i showing regi(jnalh' aix'rrant l)henologies were investigated in the lahoratory. Neither a parlialK hivoltine Piciis iiapi from the Sierra Nevada foothills in El Dorado County. California i sui rnuiidi'd hy luiivoltine populations), nor a vcrnal-univoltine P. occidenlalis from a foothill outlier of llie (Colorado Front Range (below hi- voltine populations) showed unusual resjwnses to controlled developmental regimes in the labora- tory. Their unusual phenologies are hypothesized to be the produi t of microclimate. Failure to under- go genetic adaptation to unusual microclimates is discussed witii ])articulai- reference to the i)res- ence or absence of gene flow from neaib\' normal populations. The timing of life-history phenomena naeus complex) or not (P. protodice Bois- in an insect population is (leterminetl by duval & LeConte, P. occidciitdUs Reakirt). physiological responses to en\ironniental Recent st tidies have shown that univol- stimnli. These proximate controls reflect tinism in both grouj)s is derivative from a genetic basis Ijelievecl to be the jiroduct multivoltinism, accompanying invasion of natural selection for seasonal cycles of a short-summer climate (P. occidcn- appropriate to the environment of the ta/is, Sha})iro, 1975a) or persistence in a population. In the western LTnited States progressively drier one (P. napi, Shapiro, topography has a dramatic impact on cli- 1975b). Such patterns are defined over mate, and great differences may occur broad geographic areas. California over short ground distances. How closely P. napi, for example, is differentiated into can insect populations ada])t to their im- a commonly biv'oltine, heavily pigmented mediate climates on a microgeograj)hic subspecies in the coastal summer-fog belt scale? Phenological adaptation is merely and a univoltine, more lightly marked one case of the more general jiroblem of subspecies in the interior, where summers population differentiation (cf. Ehrlich are clear and hot. The transition between and Raven, 1969; Ehrlich et al., 1975). the subspecies apjjears to be in the form In most organisms, at least prior to the of a stee]) cline through the central advent of electrophoretic genetics, popu- Coast Ranges (Shapiro, in preparation). lation differentiation was assessed on the Recently Lees and Archer (1974) have basis of visible phenotypic characters. reported the existence of phenological dif- Such characters, like the enzyme systems ferences among napi populations on a studied by electrophoresis, are often not mvich finer scale. They have found ap- translatable into specific selection pres- parently relict univoltine populations in sures. In markedly seasonal climates the suitable (bog-heath) habitats completely nature of selective pressures acting on surrounded by multivoltine ones in the phenology may be very apparent. Where British Isles. Their preliminary interpre- local deviations from the broad geograph- tation of this situation is that it provides ic pattern of voltinism are observed in a (n ideiice for midtiple invasions of Britain species, the potential exists for the demon- h\ napi stocks having different pheno- (or ecotypic) stration of microgeograj)hic logical characteristics and source regions. differentiation. This is the fourth paper In th(> course of recent work on pierid in a series exploring the evolution of sea- |)hen()logy and e\olution. the existence sonality in the butterfly genus Pieris in of regionally aberrant populations has western North America. b(>en i)r()ught to my attention in both the In various multi\oltine Pieridae both and protodicc-occidcntalis groups. In phenotype and diapause are under photo- napi ( ases the populations appear to be periodic control. The two sets of develop- both repeating mental options (diapause /direct develop- uniciiie, rather than forming a pattern as in British P. napi. They would ment; vernal /esti\ a 1 j)henotype) may be physiologically coupled {Pieris napi Lin- therefore seem to be good candidates for 'Departmcnt of Zoology. Univcisily n[ Cnlifoniin. D.ivi';. Ca 310 . Sept. 1975 311 local genetic tliflcreiitiatioii under al\[j lurliuni offn iiialc W. Br. Rorippa na.s- ical microclimates turliu!ih(i(jii(ih(nni Schii :. & Thell.) gr'ows in [xTincUient stn ims (Fig. 1). Most .,r ih,> uiid ,1,1, hutterflies are ricns iiapi ii he Sierra Foothills idonlicil to lal)orat( bred Sierran Pieris napi from interior (-alitornia are. "cdstnrid' I h'ig. 1 i as noted above, luiivoltine and moiio- On J<) Man h l<)7) niii(> in.de and three phenic in nature. Under laboratory (au- lemale lirst-hrood, vernal phenotyjie napi ditions they can lie reared without dia- were colIectcMl in one of these ravines. pause; then they produce the eslixai These included two coj)ulating j)airs in phenoty|)(» ''castoria'' Airtually unknown which the females were soft-winged, in- in the wild in the interior (Sha])iro. dicating that they had developed in the 1975b). In June 1974 Mr. William Pat- ravine^ itself. The eggs from these females terson of Sacramento, California, took were used in photoperiod experiments se\-eral 'Vy/.s/o/vV/"" of ( ( wild hoth sexes in Table 1 ) . Rearing methods are de- the canyon of the American Ri\ er below scribed m Shaj)iro. 'l975a and 1975b.) Auburn in the Sierra Nevada foothills Hie results are entirely typical for Sier- (El Dorado County, 650 feet). The oc- ran stock and do not suggest that x-Xmer- currence of a second brood there was con i(an Ri^er material has a greater pro- firmed in 1975. P. napi is connnon in the pel isit\' to flevelop directly than do stocks canyon, producing its usual \ernal pheiid tVoju purely uniAoltiiie localities, at least type in March, llie second brood, whi( li under our laboratory regimes. However, is much scarcer, unlike the first is ex- ihis is not particularly surprising. The tremely localized ^vithin llu^ camon -al second brooci of napi in the American present being known from onh' two Ri^er gorge is nukh rarer than the first, densely shaded ravines where the intro- indicating that is is onl}' partial; its num- duced cruciferous weed watercress (Nas- bers also fluctuate from year to year. B B (^5SS£S^IJ4;>rSjgx%,c-;.v|f^Vr^ Fig. 1. Locations of ravines ("B") where !)ivoItiiie Pieris tuipi Ill the American River gorge; univoltine napi are generally distriluited at low (Icnsity. l'S(;S 7. life "Aulnun"' quadrangle. 312 GREAT BASIN NATURALIST Vol. 35, No. 3 Table 1. Incidence of diapause ( D) and non- the host plant at the optimum time failed pupae in bivoltine (American diapause (ND) to turn up ;ni\ napi immatures, although River, 650') and univoltine (Placerville, 1800') ten Pieris rapac lar^'ae Pieris nopi from El Dorado Co., Calironia, reared were found.) on watercress at 27 C under two photoperiods. These circumstances suggest that the production of a second brood here is ac- Photophase: Continuous 15 hr cidental, resulting from the peculiarly Stock: Pupae: D ND D ND cool and moist conditions within the ra- American River 15 29 16 \'ines. There is no evidence that the bi- Placerville 10 23 18 Aoltine sites are in any sense isolated from adjacent univoltine ones, nor is it clear that there is successful re])roduction (In 197:5 three trips by Patterson and by the second brood in all years nor even Shapiro in season turned up only tNvo that there is genetic continuity from year males and one female. A later search of to year in the ravines; perhaps a few pu- t ^P'- Fig. 2. Wild second-brood Pieris napi from the American River gorge, collected by W. Patterson in June 1974 fmales at top; dorsal (left) and ventral (right) surfaces). The heavily marked female is atypical for an inland population. Fig. 3. Phenotypes of representative lab-reared nondiapause Pirris napi from the Ameiican Riv( stock; 27 C. continuous light; dorsal (left) and ventral (right). Sept. 1975 SHAPIRO: BUTTERFLIES 313 pae will develop directly there whenever any female napi happens to colonize them. Experienced California collectors (R. L. Langston, B. Walsh) agree that even near the coast some localities produce second-brood napi every year and others only rarely or sporadically. Exj)eriments have shown both developmental and phenotypic differences between coastal and inland stocks but not among the coastal stocks themselves. Watercress is known to be host of P. napi in various Sierran sites up to about •5,000 feet (Shapiro, 1975c). The only other record of a Sierran ^'castoria" known to me is a fresh male taken flying Table 2. Incidence of diapause (D) and non- diapause (ND) pupae in two split broods of a Barbarea verna-ieeAin^ univoltine Pieris napi (Gates Canyon, Inner Coast Ranges, Solano Co., 750') reared at 27 C on continuous light. None of the differences was significant. Brood Host Pupae: ND Brassica kabet-^ 7 Nasturtium officinale^' 5 Brassica kaber''^ 6 Lepidiuni latifolium^'-'' 5 314 GREAT BASIN NATURALIST Vol.
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