CLASSIFICATION, NATURAL HISTORY, AND EVOLUTION OF THE GENUS APHELOCERUS KIRSCH (COLEOPTERA: CLERIDAE: CLERINAE) WESTON OPITZ Kansas Wesleyan University, Department of Biology, 100 East Cla¯in Avenue, Salina KS 67401-6196 ([email protected]) BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 293, 128 pp., 287 ®gures, 1 table, 27 maps Issued May 11, 2005 Copyright q American Museum of Natural History 2005 ISSN 0003-0090 2 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 293 Habitus of Aphelocerus coactus, new species. CONTENTS Abstract ....................................................................... 6 Resumen ....................................................................... 7 Introduction .................................................................... 8 Literature Review ............................................................. 8 Materials and Methods ........................................................ 8 Assessment of Species-Level Discontinuities .................................... 10 Phylogenetic Methods ........................................................ 11 Repositories of Specimens .................................................... 11 Natural History ................................................................ 13 Mimetic Associations ........................................................ 13 Floral Associations .......................................................... 15 Field Observations ........................................................... 16 Natural History Information from Specimen Labels .............................. 17 Diagnosis of Aphelocerus ....................................................... 17 Description of Aphelocerus ..................................................... 18 Species Groups of Aphelocerus .................................................. 20 Troublesome Couplets in Key to Species ......................................... 23 Key to Species Groups and Species .............................................. 23 Description of Aphelocerus Species .............................................. 30 coalitus group Aphelocerus leucomelas (Chevrolat) ......................................... 30 Aphelocerus coalitus, new species ........................................... 32 echinatus group Aphelocerus echinatus, new species .......................................... 34 Aphelocerus affaniatis, new species .......................................... 34 ciliaris group Aphelocerus ciliaris, new species ............................................ 35 Aphelocerus citimus, new species ............................................ 37 bispineus group Aphelocerus delicatulus (Barr) .............................................. 39 Aphelocerus aeneus, new species ............................................ 39 Aphelocerus bispineus, new species .......................................... 41 Aphelocerus acuticolis, new species ......................................... 41 acanthus group Aphelocerus primigenius, new species ........................................ 42 Aphelocerus sturnus Kirsh .................................................. 42 Aphelocerus acanthus, new species .......................................... 43 Aphelocerus inconstans (Gorham) ........................................... 45 Aphelocerus cirritus, new species ........................................... 47 Aphelocerus patulus, new species ........................................... 47 Aphelocerus chiriqui, new species ........................................... 48 sculptilus group Aphelocerus sculptillus, new species ......................................... 50 Aphelocerus sabulosus, new species ......................................... 51 Aphelocerus chondrus, new species .......................................... 51 Aphelocerus cohibilis, new species .......................................... 53 irroratus group Aphelocerus irroratus, new species .......................................... 53 immarginatus group Aphelocerus immarginatus (Chevrolat) ....................................... 55 yungas group 3 4 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 293 Aphelocerus acutus, new species ............................................ 56 Aphelocerus torosus, new species ........................................... 56 Aphelocerus olanchoensis new species ....................................... 58 Aphelocerus vietus, new species ............................................. 58 Aphelocerus turnbowi, new species .......................................... 59 Aphelocerus cornuatus, new species ......................................... 61 Aphelocerus humerus, new species .......................................... 61 Aphelocerus domus, new species ............................................ 62 Aphelocerus capillus, new species ........................................... 62 Aphelocerus coactus, new species ........................................... 64 Aphelocerus scalenus, new species .......................................... 67 Aphelocerus yungas, new species ............................................ 67 Aphelocerus extensivus, new species ......................................... 68 batesi group Aphelocerus lividus, new species ............................................ 69 Aphelocerus nitidus (Chevrolat) ............................................. 70 Aphelocerus sagittarius, new species ......................................... 72 Aphelocerus triangulus, new species ......................................... 73 Aphelocerus batesi, new species ............................................. 73 Aphelocerus ebenus, new species ............................................ 75 Aphelocerus catie, new species .............................................. 75 Aphelocerus monteverde, new species ........................................ 76 Aphelocerus arenatus, new species .......................................... 78 Aphelocerus bufustis, new species ........................................... 78 Aphelocerus anticus, new species ............................................ 81 Aphelocerus naevius, new species ........................................... 81 Aphelocerus protenus, new species .......................................... 82 Aphelocerus argus, new species ............................................. 84 panus group Aphelocerus panus, new species ............................................. 84 Aphelocerus hespenheidei, new species ....................................... 85 Aphelocerus collaris, new species ........................................... 85 formicoides group Aphelocerus dispilis, new species ............................................ 87 Aphelocerus formicoides, new species ........................................ 87 cheliferous group Aphelocerus akis, new species .............................................. 88 Aphelocerus chelonus, new species .......................................... 89 Aphelocerus cheliferous, new species ........................................ 89 scutellaris group Aphelocerus scutellaris (Chevrolat) .......................................... 90 Aphelocerus improcerus, new species ........................................ 91 Aphelocerus propinquus, new species ........................................ 93 Aphelocerus calvus, new species ............................................ 93 myrmecoides group Aphelocerus myrmecoides, new species ...................................... 95 Aphelocerus inbatus, new species ........................................... 98 Aphelocerus eriodes, new species ........................................... 98 Aphelocerus discapillus, new species ....................................... 100 Evolutionary Considerations ................................................. 100 Character Selected for Phylogenetic Analysis ................................ 101 Phylogenetic Analysis ..................................................... 103 Phylogenetic Scenario for the Evolution of Aphelocerus and its Species Groups . 103 2005 OPITZ: APHELOCERUS 5 Zoogeographic Considerations ............................................. 104 Geophysical Orismology .................................................. 105 Grupos de Especies de Aphelocerus ........................................... 108 Pasos Problamaticos en la Clave de Especies .................................. 110 Clave de los Grupos de Especie y de las Especies .............................. 111 Acknowledgments ............................................................ 117 References ................................................................... 117 Maps ........................................................................ 120 ABSTRACT The genus Aphelocerus is rede®ned to include 66 species as follows: A. leucomelas (Chev- rolat); A. coalitus, n.sp.; A. echinatus, n.sp.; A. affaniatis, n.sp.; A. delicatulus (Barr); A. aeneus, n.sp.; A. bispineus, n.sp.; A. extensivus, n.sp.; A. primigenious, n.sp.; A. ciliaris, n.sp.; A. acuticolis, n.sp.; A. citimus, n.sp.; A. sturnus Kirsh; A. acanthus, n.sp.; A. inconstans (Gor- ham); A. cirritus, n.sp.; A. patulus, n.sp.; A. chiriqui, n.sp.; A. sculptilus, n.sp.; A. sabulosus, n.sp.; A. chondrus, n.sp.; A. cohibilis, n.sp.; A. irroratus, n.sp.; A. immarginatus (Chevrolat); A. acutus, n.sp.; A. torosus, n.sp.; A. olanchoensis, n.sp.; A. vietus, n.sp.; A. turnbowi, n.sp.; A. cornuatus, n.sp.; A. humerus, n.sp.; A. domus, n.sp.; A. capillus, n.sp.; A. coactus, n.sp.; A. scalenus, n.sp.; A. yungas, n.sp.;