A Monograph of the Genus Polyphylla Harris in America North of Mexico (Coleoptera: Scarabaeidae: Melolonthinae) Ronald M
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University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln Bulletin of the University of Nebraska State Museum, University of Nebraska State Museum 1988 A Monograph of the Genus Polyphylla Harris in America North of Mexico (Coleoptera: Scarabaeidae: Melolonthinae) Ronald M. Young Follow this and additional works at: http://digitalcommons.unl.edu/museumbulletin Part of the Entomology Commons, Geology Commons, Geomorphology Commons, Other Ecology and Evolutionary Biology Commons, Paleobiology Commons, Paleontology Commons, and the Sedimentology Commons Young, Ronald M., "A Monograph of the Genus Polyphylla Harris in America North of Mexico (Coleoptera: Scarabaeidae: Melolonthinae)" (1988). Bulletin of the University of Nebraska State Museum. 38. http://digitalcommons.unl.edu/museumbulletin/38 This Article is brought to you for free and open access by the Museum, University of Nebraska State at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Bulletin of the University of Nebraska State Museum by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. BULLETIN OF VOLUME 11, NUMBER 2 The University of Nebraska State Museum FEBRUARY, 1988 Ronald M. Young A Monograph of the Genus Polyphylla Harris in America North of Mexico (Coleoptera: Scarabaeidae: Melolonthinae) Ronald M. Young A Monograph of the Genus Polyphylla Harris in America North of Mexico (Coleoptera: Scarabaeidae: Melolonthinae) BULLETIN OF The University of Nebraska State Museum VOLUME 11, NUMBER 2 FEBRUARY, 1988 Frontispiece.-Polyphylla decemlineata (Say). Male. BULLETIN OF VOLUME 11, NUMBER 2 THE UNIVERSITY OF NEBRASKA STATE MUSEUM February, 1988 ABSTRACT A Monograph of the Genus Polyphylla Harris in America North of Mexico (Coleoptera: Scarabaeidae: Melolonthinae) Ronald M. Young The North American species of the genus Po/yphyl/a Harris, 1841, are defined and revised for the first time since Fall (1928). All name bearing types were examined except those of P. ok/ahomensis Hatch and P. subvittata LeConte (synonyms of P. hammondl), which are lost. A lectotype is designated for P. gracilis Horn, and neotypes for P. decemlineata (Say), P. occidentalis (L.), and P. vari%sa (Hentz). New synonyms of P. decemlineata are P. comstockiana von Bloeker, P. matrona Casey, and P. potsiana Casey. New synonyms of P. crinita LeConte are P. pacifica Casey and P. ruficol/is perversa Casey. P. al/eni Cazier is a new synonym of P. diffracta Casey. P. crinita nigra Casey is raised to specific rank and P. inco/umis relicta Casey, P. inco/umis robustu/a Casey, P. martini von Bloeker, and P. santacruzae von Bloeker are its new synonyms. New synonyms of P. hammondi LeConte are P. /atifrons Casey, P. ru fescenta Tanner, P. speciosa Casey, and P. speciosa acomana Casey. P. marginata is a nomen nudum. Descriptions of first known females are given for P. barbata Cazier, P. devestiva Young, and P. erratica Hardy and Andrews. P. mesca/erensis is a new species from southeastern New Mexico. Of the 71 names established since 1767 which represent Po/yphyl/a, 28 are recognized as valid. Females are unknown for 12 species. These species cluster morphologically and geographically into four species complexes. A key to larvae is provided for the four species having described larvae. A key to adults is provided, and all taxa are described or redescribed and illustrated by photographs. Available biological information is included. Distributional data and maps based on 14,094 specimens are provided. A cladistic interpre tation and hypothetical phylogeny of the genus are constructed. Copyright © 1988 by the Board of Regents of the University of Nebraska Library of Congress Catalog Card Number ISSN 0093-6812 Manufactured in the United States TABLE OF CONTENTS Introduction ............................................................................ 1 Acknowledgements ................................................................... 12 Review of literature ................................................................... 13 Survey of morphological characters ................................................... 14 Nomenclature ........................................................................ 15 Po/yphyl/a Harris ...................................................................... 20 Generic nomenclature ............................................................ 20 Generic diagnosis ................................................................ 21 Generic description .............................................................. 21 Key to larvae ..................................................................... 22 Key to adults ..................................................................... 23 Species exclusively east of the Mississippi River (the occidentalis species complex) ........................................... 28 P comes Casey ...................................................... 28 P gracilis Horn ........................................................ 30 P occidentalis (L.) .................................................... 33 P vari%sa (Hentz) ................................................... 35 Species primarily west of the Mississippi River .................................... 39 The decemlineata species complex .......................................... 39 P arguta Casey ....................................................... 39 P decemlineata (Say) ................................................. 40 P monahansensis Hardy and Andrews ................................ 51 The diffracta species complex ................................................ 52 P avittata Hardy and Andrews ........................................ 52 P barbata Cazier ..................................................... 53 P crinita LeConte ..................................................... 55 P devestiva Young .................................................... 59 P diffracta Casey ..................................................... 60 P hirsuta Van Dyke ................................................... 63 P. mescalerensis new species ......................................... 64 P. modulata Casey .................................................... 65 P. nigra Casey ........................................................ 67 P. nubila Van Dyke .................................................... 69 P. pottsorum Hardy and Andrews ...................................... 70 P. ratcliffei Young ..................................................... 71 P. rugosipennis Casey ................................................ 72 P. sobrina Casey ...................................................... 73 P. stellata Young ...................................................... 74 The hammondi species complex ............................................. 75 P. anteronivea Hardy and Andrews .................................... 75 P. brownae Young ..................................................... 75 P. cavifrons LeConte .................................................. 77 P. erratica Hardy and Andrews ........................................ 78 P. hammondi LeConte ................................................. 79 P. squamiventris Cazier ............................................... 90 Distribution and isolating mechanisms ................................................. 90 Pheromones .......................................................................... 92 Phylogenetic analysis ................................................................. 92 Biogeography ........................................................................ 95 Dispersal and differentiation of species ................................................ 99 References .......................................................................... 106 INTRODUCTION vide clues to understanding the species. The work of The melolonthine genus Polyphylla Harris is found Downes and Andison (1941), Johnson (1954), Young worldwide between ca. 15 and 53 degrees North lat (1966,1972), and Hardy and Andrews (1978), among itude. In the New World the southernmost record is others, now enables us to predict which general hab Honduras and the northernmost is Quesnel, British itat types will support Polyphy/la. I have an impres Columbia (Fig. 1). sion that accumulation of dissimilarity in these ani Distribution is typically discontinuous with many mals is a fairly rapid process in isolated interbreeding species restricted to specific, isolated, sandy envi populations; therefore it may be predicted that ad ronmental refugia. Of the 28 species in America north ditional species will be discovered with intensified in of Mexico, 8 (ca. 30%) are broadly distributed across vestigations of sand/grass associations. It may also three to many states. Five species (18%) are narrow be predicted that our knowledge of immature stages, ly distributed, typically found in only two or three adult feeding habits and periodicity, and the females states. The remaining 15 species (ca. 54% of the which remain unknown in 12 species, will increase. North American fauna), are endemic to one or two Endemism of a taxon to a relatively small area is localities which tend to be unique sandy or sand dune common in insects; perhaps so common that ento areas. mologists began to take it for granted. Yet when one All science is a step process with current scientists stops to consider, it seems quite remarkable that of correcting, refining, and adding to the work of their all the spectrum