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Molecular Phylogeny of the North American Enchenopa Binotata (Homoptera: Membracidae) Species Complex

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Molecular Phylogeny of the North American Enchenopa Binotata (Homoptera: Membracidae) Species Complex SYSTEMATICS Molecular Phylogeny of the North American Enchenopa binotata (Homoptera: Membracidae) Species Complex 1 2 CHUNG PING LIN AND THOMAS K. WOOD Ann. Entomol. Soc. Am. 95(2): 162Ð171 (2002) ABSTRACT The North American Enchenopa binotata (Say) species complex is a model of sympatric speciation in which phytophagous insects are hypothesized to diverge through host-plant special- ization resulting from changes in host plant usage that alter life history timing. A robust phylogeny is needed to evaluate the historical relevance of the prediction that sister taxa differ in critical life-history traits. Phylogenetic analysis using parsimony and likelihood criteria of 2305 nucleotides in sequences from mitochondrial COI, COII, tRNA-Leucine, and 12S genes revealed two pairs of sister taxa. Both pairs of sister taxa differ from each other in the timing of egg hatch in the spring that is mediated by differences in host-plant phenology. Host plant mediated timing of egg hatch results in asynchronous life histories among sister taxa facilitating reproductive isolation. Sister taxa of Enche- nopa from Celastrus and from Viburnum differ in their diurnal and temporal spans during which mating occurs. Mating of Enchenopa from Liriodendron takes place after that of its sister species on Cercis. These results support the hypothesis that speciation could have been initiated through a shift to a host plant that alters life-history timing. KEY WORDS Homoptera, Enchenopa, speciation, sympatric CLADES OF SPECIES are interrelated through historical, mographically divergent and reproductively cohesive genealogical, and geographical connections that in- making cause and effect difÞcult to interpret (Wood ßuence how extant species complexes respond to evo- 1993). Species in this complex not only have well- lutionary processes through time (Hillis 1997). Thus, developed host plant preference/philopatry during the historical phylogenetic context of a species or its mating and oviposition but also have been experimen- relationship within a clade is essential to interpreting tally demonstrated in choice tests to mate by species. the results of comparative studies dealing with evo- Females experimentally transferred during oviposi- lutionary processes (Brooks and McLennan 1991, tion to inappropriate host plants do not successfully Brooks et al. 1995). One of the fundamental underly- produce offspring (Wood 1980; Wood and Guttman ing processes of evolutionary biology is speciation. 1982, 1983). Other differences such as substrate-borne This is a difÞcult area because deÞnitions of species are mating signals, morphological and color pattern dif- varied and often reßect underlying assumptions con- ferences among nymphs have also been demonstrated cerning the mode of speciation (Templeton 1989). For (Wood 1980, Pratt and Wood 1992, Hunt 1994). How- example, the historical debate over whether geo- ever, discrete adult morphological differentiation in graphical isolation is a requisite for speciation or not this complex of species has not developed (Pratt and has been controversial for many years (Mayr 1982). Wood 1993). The challenge for species complexes like Many of the examples of purported sympatric specia- E. binotata is to Þnd sufÞcient phylogenetically infor- tion are difÞcult to evaluate from a phylogenetic per- mative characters to provide a robust analysis of re- spective because organisms such as the host races of lationships within the clade to evaluate mechanisms of Rhagoletis pomonella (Walsh) (Diptera: Tephritidae) speciation. have not achieved species status regardless of deÞni- The E. binotata species complex is amodel of sym- tion (Bush 1969). In others, such as the Enchenopa patric speciation in which phytophagous insects are binotata (Say) species complex, each species is de- hypothesized to diverge through host-plant special- ization resulting from changes in host-plant usage (Wood 1980; Wood and Guttman 1981, 1982, 1983; Nomenclatural acts or proposed reclassiÞcations inferred in this article are not considered published within the meaning of the 1985 Wood et al. 1990; Wood and Keese 1990; Wood 1993; International Code of Zoological Nomenclature (Article 8b). This Tilmon et al. 1998; Wood et al. 1999). The hypothe- work will be published elsewhere in accordance with article eight of sized speciation mechanism is that asynchronous mat- the Code. ing is induced by differences in plant phenology 1 Department of Entomology, Comstock Hall, Cornell University, Ithaca, NY 14853 (e-mail: [email protected]). which, interacting with philopatry during reproduc- 2 Department of Entomology and Applied Ecology, University of tion, allows divergence in host-plant associated per- Delaware, Newark, DE 19717. formance traits (Wood et al. 1990; Wood and Keese 0013-8746/02/0162Ð0171$02.00/0 ᭧ 2002 Entomological Society of America March 2002 LIN AND WOOD:MOLECULAR PHYLOGENY OF Enchenopa COMPLEX 163 Table 1. Host plants of the Enchenopa binotata species com- The genus Enchenopa is placed within the tribe plex (Wood 1993) Membracini in the subfamily Membracinae on the basis of morphology (Metcalf and Wade 1965, Deitz Genus Species Family 1975, McKamey 1998). There is debate over whether Ptelea trifoliata (L.) Rutaceae the tribe Membracini is monophyletic and its rela- Juglans nigra (L.) Juglandaceae Juglans cinerea (L.) Juglandaceae tionship to the other tribes within the subfamily Carya illinoensis (Wang) K. Koch (Dietrich and McKamey 1995). Because pronotal ovalis (Wang) Sarg. shape is a primary character for distinguishing genera, cordiformis (Wang) K. Koch Enchenopa is not well delineated from other genera laciniosa (Michx.) Loud. ovata (Mill.) K. Koch such as Campylenchia and Enchophyllum. Thus, it is Celastrus scandens (L.) Celastraceae possible that many species presently assigned to Liriodendron tulipifera (L.) Magnoliaceae Enchenopa, Campylenchia and Enchophyllum are mis- Robinia pseudoacacia (L.) Leguminosae placed. Although adults of the nine species in the E. Cercis canadensis (L.) Leguminosae Viburnum cassinoides (L.) Caprifoliaceae binotata complex differ in body length and pronotal rufidulum (Raf.) shape, the genitalia (Pratt and Wood 1993) and other lentago (L.) morphological features do not provide diagnostic prunifolium (L.) characters suitable for the development of a rigorous phylogenetic hypothesis. Attempts have been made to understand relationships within the E. binotata species 1990; Wood 1993; Tilmon et al. 1998; Wood et al. 1999). complex using allozymes, female pronotal shape and The North American E. binotata species complex is nymphal characters (Wood 1993). comprised of nine species associated with eight plant Relationships within the E. binotata species com- genera distributed among six plant families (Table 1). plex were Þrst inferred using allozyme data (Guttman Because the species in this complex have not been et al. 1981, Wood 1993) using Campylenchia latipes formally named, we refer to them by their host-plant (Say) as an outgroup to derive a distance matrix. genus or by host-plant species as in the case of the two There is considerable debate on whether allozymes species using Juglans nigra (L.) and J. cinerea (L.) provide appropriate information reßecting evolution- (see footnote for nomenclatural disclaimer). The E. ary history (Swofford et al. 1996). The two major binotata species on Carya and Viburnum are polyph- concerns are whether or not to transform allozyme agous in that they occur on several species within their data to a genetic distance and how evolutionarily im- respective plant genera but the remaining species ap- portant is the presence/absence of alleles versus the pear to be monophagous (Wood 1993). The Enche- frequency of alleles (Mickevich and Johnson 1976, nopa species and their host plants are sympatric Swofford and Berlocher 1987). Unless sample sizes are throughout the eastern United States (Wood 1993) large, taxa that are in reality polymorphic for some (Fig. 1). There is substantial overlap of the distribution alleles may be scored as Þxed if one allele is rare. The of Enchenopa species with that of its host plant (Little same holds true if only a few populations within a 1971). species are sampled since the frequency of an allele In North America, the univoltine E. binotata species can vary dramatically among populations over a geo- begin their life history in the spring when overwin- graphic range (Swofford and Berlocher 1987). In the tered eggs hatch synchronously within a 10 d period allozyme study of the E. binotata complex, sample size on their host-plant. Nymphs mature to the adult stage and number of localities appear to be adequate to Ϸ1 mo after egg hatch and males eclose 1Ð2 d before counter these objections, but a distance matrix cannot females. Mating begins Ϸ1 mo after adult eclosion. be analyzed by modern character-based cladistic anal- Males can mate several times, whereas females mate ysis, and homology assessments remain controversial. once (Wood 1993). Oviposition begins Ϸ1 mo after The only cladistically based analysis is a phylogeny the onset of mating. Females insert eggs into woody using seven Þrst and 24 Þfth-instar nymphal characters stems, cover eggs with froth, and deposit multiple egg (Pratt and Wood 1992). masses throughout the fall until mid-November In the above studies (Pratt and Wood 1992, Wood (Wood and Patton 1971). Eggs of each of the nine 1993), C. latipes was used as an outgroup because it Enchenopa species hatch at different times in the was the only
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