Abstract

The checkered beetle genus Ababa is revised for the first time. Morphological evidence suggests that Thaneroclerinae and Isoclerinae comprise a monophyletic group. Based on the positional relationships between the pronotal projection and the prothoracic intercoxal process the isoclerine genus Ababa Casey is taken out of synonymy with Isoclerus Lewis. Cleridopsis Champion is considered to be a junior synonym of Ababa Casey. On the basis of the tarsal formula, the genus Allothaneroclerus Corporaal, with a tarsal formula of 5-5-5, is removed from Isoclerinae, whose genera show a 5-4-4 tarsal formula, and placed into the subfamily Thaneroclerinae. Ababa specimens were collected in fungi niches and it is postulated that the elongated stomodaeum found in adults serve as a crop to process soft food such as the eggs, larvae, and pupae of fungivorous prey. Ababa comprises A. adona Opitz, new species, A. epiiska Opitz, new species, A. granaria Opitz, new species, A. latimana (Champion), and A. tantilla (LeConte). A lectotype was designated for Ababa latimana (Champion). It is hypothesized that the ancestral stock of Ababa and Parateneroclerus evolved in the mid-Tertiary forests of South America. Included in this work are: A brief discussion about Ababa natural history, a discussion of Ababa phylogeny based on a phylogenetic tree generated by WINCLADA in conjunction with NONA, a Bootstrap consensus tree, a key to Ababa species, 10 scanning electron micrographs, one distribution map, four photographs of the aedeagus, and six color habitus photographs.

Resumen

El género de Cleridae Ababa es revisado por primera vez. La evidencia morfológica sugiere que los Thaneroclerinae y los Isoclerinae forman un grupo monofiletico. Basado sobre la relación de la posición de la proyección pronotal y del proceso intercoxal protoracico, el género isoclerine Ababa Casey es removido de la sinonimia con Isoclerus Lewis. Cleridopsis Champion es considerado como un sinónimo junior de Ababa Casey. En base a la formula tarsal, 5-5-5, el género Allothaneroclerus Corporaal es removido de los Isoclerinae, que presentan formula tarsal 5-4-4, e incluido en la subfamilia Thaneroclerinae. Los especímenes de Ababa fueron colectados en hongos y se postula que el stomodaeum alargado encontrado en los adultos sirve de estómago para digerir los huevos, larvas y pupas de sus presas micetófagas. El género Ababa incluye A. adona Opitz, especie nueva, A. epiiska Opitz, especie nueva, A. granaria Opitz, especie nueva, A. latimana (Champion), y A. tantilla (LeConte). Se designó un lectotipo para Ababa latimana (Champion). Se presenta la hipótesis de que los ancestros de Ababa y Parathaneroclerus evolucionaron en los bosques de Suramérica del Terciario medio. En el trabajo se incluye: una breve discusión de la biología de Ababa, una filogenia basada en arboles filogenéticos generados con el programa WINCLADA en conjunto con NONA, un árbol de consenso obtenido por análisis Bootstrap, una clave de las especies de Ababa, cuatro dibujos, 10 fotografías con microscopio electrónico, un mapa de distribución, cuatro fotografías de aedeago, y cinco fotografías a color de especímenes.

Palabras clave: Biología, Clasificación, Clave de especies, Cleridae, Cleroidea, Filogenia, Taxonomía.

Introduction

In 2010, Opitz provided evidence for the reestablishment of Thaneroclerinae as a subfamily of the Cleridae and for the monophyly of subfamily Isoclerinae (Opitz, 2010). The purpose of this work here is to make known better the species composition of Isoclerinae and Thaneroclerinae. Now, new observations and findings support the elevation of Ababa Casey and Parathaneroclerus Pic to generic status, under Isoclerinae Kolibáč. The genus Ababa was described by Thomas L. Casey in 1897 based on A. crinita, a junior synonym of A. tantilla (LeConte, 1865). The genus remained monotypic until Maurice Pic, in 1947, introduced A. longipennis, which was synonymized by Jiří Kolibáč (Kolibáč, 1992) under Isoclerus paralleus (Lewis, 1892). Then, William F. Barr discovered that in 1913 Charles Champion described an Ababa species, Cleridopsis latimanus under Cryptophagidae. Heretofore, Ababa remained bitypic until the present work in which three new species are added to the genus. For a different perspective on the higher classification of Isoclerinae and Thaneroclerinae see Kolibáč (1992).

Material and Methods

Although morphological criteria are used to determine species-level discontinuities, I adhere to the biological species concepts as discussed by Standfuss (1896), Dobzhansky (1937), and Mayr (1963). The operational criteria for the delimitation of species involve morphological structure and any other available criteria that suggest reproductive isolation among members of meta-population lineages (de Queiroz, 2007). Experience with morphological structure is generally a reasonable criterion with which to hypothesize reproductive isolation. In this study, consideration for species status involves: integumental color, structure of the pronotum, arrangement of setiferous punctures on the elytral disc, and characteristics of the aedeagus.

Methods involving dissection, measurements involving eye width/frons width, pronotal width, and elytral length, and morphological terminology follow those described in Opitz (2010). Brown (1956) was used to coin scientific names. Abbreviations used in this manuscript are defined as follows: EW/FW = eye width/frons width, measured at 500 x from the front of the head; PW/PL= pronotal width, across the widest portion of the pronotal disc/pronotal length, from midline anterior margin to midline posterior margin; EL/EW= elytral length, from humeral angle to apex)/elytral width, with greatest dorsal width of one elytron. Elytral setae are designated as 1°, those setae adjacent to asetiferous punctures, and 2°, those set in interstitial spaces, i.e., set between punctures (Opitz, 2018).

Habitus photographs were taken with a Leica Z 16 APO microscope equipped with JVC KYF75U-CCD camera and controlled by Syncroscopy Auto Montage software (Cambridge, United Kingdom). The SEM micrographs were produced with a Scanning Electron MicroscopeS-3500N (Hitachi Science Systems, Ltd., Tokyo, Japan). Image stacks, involving the aedeagus, were taken with a Leica® DM2500 compound scope with a 10X objective lens and a Leica® DFC425 camera (Meyer Instruments, Houston, Texas, United States of America), and combined using Zerene Stacker®.

Assessments of Evolutionary States of Characters

Nine character states were organized into a matrix (Table 1), which was then analyzed with NONA (Goloboff, 2003) in combination with WINCLADA version 100.80 (Nixon, 2002) to find the most parsimonious phylogenetic tree. The WINCLADA program produced one tree via heuristic analysis: Maximum trees (hold) = 100, number of replications 1 (mult) = 100, and multiple TBR = TBR (mult max) were used with indices as follows: L-9, Ci-88, Ri-80 (Fig. 11). The nexus data file was prepared with Mesquite v.3.5 (Build 888); PAUP v.4.Oa (Build 161) was used to perform 100 parsimony bootstrap replicates using heuristic searches, each based on 10 random addition sequences using TBR as the branch-swapping algorithm, with all characters equally weighted. This program produced a bootstrap consensus tree (Fig. 6B) with indices noted on the diagram. The methodology of Hennig (1966) was followed for estimations of supraspecific relationships, although I agree with Tuomikoski (1967) who advocates the use of “apotypic” and “plesiotypic” instead of “apomorphic” and “plesiomorphic” because phylogenetic work may not be restricted to morphological criteria.

Table 1.

Character matrix of 9 adult morphological characters of Thaneroclerinae genera, species of Ababa and the outgroup genera Cymatodera and Lecontella.

Character matrix of 9 adult morphological characters of Thaneroclerinae genera, species of Ababa and the outgroup genera Cymatodera and Lecontella.
Character matrix of 9 adult morphological characters of Thaneroclerinae genera, species of Ababa and the outgroup genera Cymatodera and Lecontella.

Character states given the value of “0” are assessed plesiotypic whereas those judged a value of “1” are assessed apotypic. The genera Cymatodera Gray and Lecontella Wolcott and Chapin were used as outgroups to assist in character state assessments and to predict the evolutionary states of characteristics. I relied on the methods of character-state analysis employed by Ekis (now Opitz) (1977), Watrous and Wheeler (1981), and Nixon and Carpenter (1993).

  • Character 0-Tibial spur formula: (0) not 1-2-2; (1) 1-2-2

  • Character 1-Geographic distribution: (0) Old World; (1) New World

  • Character 2-Tarsal formula: (0) 5-5-5; (1) 5-4-4

  • Character 3-Body size: (0) more than 2.0 mm; (1) 2.0 mm

  • Character 4-Pronotal side margins: (0) not roughened; (1) roughened

  • Character 5-Pronotal shape: (0) not narrow; (1) narrow

  • Character 6-Color of elytral disc: (0) unicolored; (1) bicolored

  • Character 7-Geographic distribution: (0) not South American; (1) South American

  • Character 8-Pronotal interstitial space: (0) not arenose; (1) arenose

Repositories of Specimens

I used codens as noted in Arnett, Jr. et al. (1993) to indicate repositories of specimens, with some modifications to accommodate institutional name changes.

  • AMNH-American Museum of Natural History, Department of Entomology, Central Park West at 79th Street, New York, New York 10024-5192, United States of America (Lee Herman).

  • BMNH-British Museum of Natural History, Department of Entomology, SW 5BD, London, United Kingdom (Beulah Garner).

  • BYUC-Brigham Young University, Monte L. Bean Life Science Museum, Insect Collections, 645 East 1430 North, Provo, Utah 84602, United States of America (Shawn M. Clark).

  • CMNC-Canadian Museum of Nature, Insect Collection, Post Office Box 3443, Station D, Ottawa, Ontario, Canada K1P 6P4, Canada (Robert S. Anderson).

  • CMNH-Carnegie Museum of Natural History, Invertebrate Zoology, 4400 Forbes Avenue, Pittsburgh, Pennsylvania 15213, United States of America (Robert Androw).

  • EMEC-Essig Museum of Entomology, University of California, College of Agriculture, Division of Entomology and Parasitology, California Insect Survey, Berkeley, California 94720, United States of America (Pete Oboyski).

  • FMNH-Field Museum of Natural History, Department of Entomology, Roosevelt Road at Lake Shore Drive, Chicago, Illinois 60605, United States of America (Crystal Maier).

  • FSCA-Florida State Collection of Arthropods, Division of Plant Industry/Entomology, Doyle Connor Building, 1911 SW 34th Street, Florida Department of Agriculture, Gainesville, Florida 32614-7100, United States of America (Paul E. Skelley).

  • MCZC-Museum of Comparative Zoology, Harvard University, Entomology, 26 Oxford Street, Cambridge, Massachusetts 02138, United States of America (Philip D. Perkins).

  • RFMC-Roy F. Morris II Collection, 2635 Ewell Road, Lakeland, Florida 33811, United States of America.

  • UGCA-University of Georgia, Collection of Arthropods, Natural History Building, Athens, Georgia, United States of America, 30602-7882 (E. Richard Hoebeke).

  • USNM-National Museum of Natural History MRC 168, Washington, D.C. 20560-0165, United States of America (Floyd Shockley).

  • WOPC-Weston Opitz Collection, Research Associate, Florida State Collection of Arthropods, Division of Plant Industry/Entomology, Florida Department of Agriculture and Consumer services, 1911 SW 34th Street, Gainesville, Florida 32614-7100, United States of America.

  • ZMHB-Museum für Naturkunde, Institute für Systematische Zoologie, Invalidenstrasse 43, D – 10115, Berlin, Germany (Bernd Jaeger).

KEY TO ADULTS OF SPECIES OF ABABA

    KEY TO ADULTS OF SPECIES OF ABABA
  • 1.

    Dorsum bicolorous . . . 2

  • -

    Dorsum unicolorous . . . 3

  • 2.

    Elytron with brown fascia (Costa Rica, Panama) . . . Ababa adona, sp. nov. (Fig. 14)

  • -

    Elytron without brown fascia (Peru, Argentina, Paraguay) . . . Ababa epiiska, sp. nov. (Fig. 15)

  • 3.

    Pronotal interstitial spaces (spaces between punctures) arenose . . . Ababa granaria, sp. nov. (Figs. 16, 17)

  • -

    Pronotal interstitial spaces not arenose . . . 4

  • 4.

    Pronotum narrow (0.5 mm) (Guatemala) . . . Ababa latimana (Champion) (Fig. 18)

  • -

    Pronotum broad (0.8 mm) . . . Ababa tantilla (LeConte) (Fig. 19)

  • ABABA CASEY, 1897 

  • Ababa Casey, 1897: 653. Type Species: Ababa crinita Casey, 1897: 653, by original designation.

  • Borchmann, 1910: 2 (Othniidae). Schaeffer, 1917: 133. Bradley, 1930: 107. Corporaal, 1939: 348. Wolcott, 1947: 76. Barr, 1975: 1. Kolibáč, 1992: 319. Peck & Thomas, 1998: 85. Opitz, 2002: 273; 2010: 86; 2014: 2. Junior synonyms: Prionostichaeus, Wolcott, 1911: 125, synonymy by Corporaal, 1939: 358 (junior subjective synonym). Cleridopsis, Champion, 1913: 94, new synonymy (junior subjective synonym). Wolcottella Lucas, 1920: 669, synonymy by Corporaal, 1949: 358 (junior objective synonym).

  • Diagnosis: Ababa specimens may be distinguished from the members of its sister genus, Parathaneroclerus, by showing an evenly convex pronotal disc. The pronotal disc shows four indentations in specimens of Parathaneroclerus.

  • Redescription: Form: Habitus oblong-short, deep bodied; pronotum campaniform; elytra short rectangulate; elytral posterior slope acute (downward angle of the elytra). Size: Length 1.9–3.2 mm; width 0.6–1.0 mm. Integumental Color: Mostly light castaneous, rarely with dark brown elytral markings. Head: Quadrate (Fig. 1A); postgena expanded; epistomal sutures abbreviated, externally briefly visible at sides; clypeus not discernible externally; frontal preantennal angle acute; eyes small, coarsely faceted, ocular notch minute; gula narrow, gular sutures briefly converging, then parallel; postgular plate (Fig. 1B) well defined, narrow transverse; post-gular process incipient; antenna comprised of 11 antennomeres, capitate; mandible stout, anterior and medial dens well developed, posterior dens shallow, penicillus well developed, basal notch very large; labral anterior margin broadly concave, transverse tormal processes linear, contiguous; maxilla with well-developed laterolacinia, terminal palpomere broad digitiform; labium well developed, terminal palpomere broad digitiform. Thorax: Pronotum campaniform, dorsolateral carina well developed, anteriorly reaches pronotal anterior angle, posteriorly connects to pronotal hem; pronotal projection (Fig. 2B) long, approximates but not connected to distally widened prointercoxal process; prosternum usually expanded towards anterior; coxal cavities spheroid, procryptosternum reduced; mesosternum cylindrically projected towards anterior; elytral disc profusely vested with 2° setae (Opitz, 2018), 1° setae and asetiferous punctures absent; epipleural fold well developed, positioned laterally, anterior ridge absent; legs, tarsal formula 5-4-4 (Fig. 5B), tarsal pulvilli absent, first 4 prothoracic tarsomeres greatly expanded, venter highly sclerotized and densely setose, 5th tarsomere extraordinarily lengthened, unguis without denticle; mesothoracic and metathoracic tarsomeres not expanded ventrally; prothoracic femur (Fig. 4A) very broad, mesothoracic (Fig. 4B), and metathoracic (Fig. 5A) femora slightly broadened; tibiae about as long as length of femora, without carina; tibial spur formula 1-2-2; metathoracic wing venation greatly reduced; metendosternite with slightly-developed furcal lamina. Abdomen: Comprised of 6 visible sternites, 6th sternite positioned under 5th; pygidium scutiform; 6th visible sternite not incised distally; spicular fork well developed, broad, plates slender, spicular apodeme fused entirely, intraspicular plate transverse narrow, slightly setose; aedeagus poorly sclerotized, phallobasic struts continuous, phallobasic apodeme absent, phallobase lobe acuminate, lobes fimbriate, phallobasic dorsum lightly sclerotized, phallobasic venter semimembraneous. Alimentary Canal: Stomodaeum extraordinarily long, about as long as ventriculus; stomodaeal valve not distinguishable externally; ventriculus not papillose; proventriculus comprised of 4 primary and 4 secondary lobes. Mesodermal Male Reproductive Organs: No information available. Mesodermal Female Reproductive Organs: Spermathecal capsule faintly sclerotized; saccular bursa copulatrix absent.

  • Distribution (Fig. 7): This New World genus is distributed from The United States of America to Argentina.

Fig. 1.

Head of Ababa tantilla. A. Frontal view. B. Ventral view.

Fig. 1.

Head of Ababa tantilla. A. Frontal view. B. Ventral view.

Fig. 2.

Prothorax, ventral view. A. Zenodosus sanguineus. B. Ababa tantilla.

Fig. 2.

Prothorax, ventral view. A. Zenodosus sanguineus. B. Ababa tantilla.

Fig. 3.

Prothorax, ventral view. A. Parathaneroclerus impressus. B. Isoclerus pictus.

Fig. 3.

Prothorax, ventral view. A. Parathaneroclerus impressus. B. Isoclerus pictus.

Fig. 4.

Legs of Ababa tantilla. A. Prothoracic. B. Mesothoracic.

Fig. 4.

Legs of Ababa tantilla. A. Prothoracic. B. Mesothoracic.

Fig. 5.

Legs of Ababa tantilla. A. Metathoracic. B. Metathoracic tarsus.

Fig. 5.

Legs of Ababa tantilla. A. Metathoracic. B. Metathoracic tarsus.

Fig. 6.

Phylogenetic trees. A. WINCLADA/NONA. B. BOOTSTRAP CONSENSUS.

Fig. 6.

Phylogenetic trees. A. WINCLADA/NONA. B. BOOTSTRAP CONSENSUS.

Fig. 7.

Known geographic distribution of Ababa species as noted.

Fig. 7.

Known geographic distribution of Ababa species as noted.

  • Ababa adona, new species

    Figs. 6A, 6B, 7, 8A, 9C.

  • Diagnosis: The brown mid-elytral fascia and the infuscation on the elytral apex will distinguish the members of this species from congeners.

  • Description: Size: Length 2.5 mm; width 0.6 mm. Form: As in Fig. 8A. Color: Head and prothorax dark castaneous; antenna brown; pterothorax, legs, and abdomen castaneous; elytra mostly light castaneous, with mid-elytral black fascia, elytral apex infuscated. Head: Cranium imprinted with oblong punctures; eyes small, coarsely facetted, eye much narrower than frons (EW/FW 5/26); antenna capitate, funicular antennomeres subfiliform, progressively shorter towards capitulum, latter lax, capitular antennomeres 9 and 10 cupuliform, antennomere 11 spheroid. Thorax: Pronotum oblong campaniform; disc impressed with oblong punctures; pronotal side margin slightly roughened (PW/ PL 38/44); elytral impressed with large setiferous punctures, punctures not arranged into linear striae and diminish near elytral 1/2 (EL/EW 125/ 30). Abdomen: Pygidium scutiform; aedeagus as in Fig. 9C.

  • Natural History: Seven specimens were captured in Costa Rica at 915 m. One specimen was collected in Panamá in flower fall of a species of the genus Pseudobombax Dugand (Malvaceae), another in the fruiting body of the woody fungus Fomes sclerodermeus Lév. (Polyporaceae).

  • Distribution: This species is known from Costa Rica and Panamá (Fig. 7).

  • Etymology: The trivial name, adona, is a Latin name derived from adonis (= favorite); with reference to my interest in these beetles.

  • Specimens Examined: Holotype Male: Type locality: COSTA RICA, Provincia de San José: San Isidro del General, 5-XII-1963, 3000 feet, Steven L. Wood (FSCA). PARATYPES: Seven specimens. Locality information as in holotype (BYUC, 1). COSTA RICA, Provincia de Alajuela: near Bijagua, 1-VII-1991, J. T. Doyen & D. Richie (EMEC, 1). PANAMÁ, Canal Zone: Barro Colorado Island, 3-6-II-1976, Pseudobombax flower fall, Snyder-Molino, A. Newton (FMNH, 1); idem, 30-I-1980; idem, 10-II-1968, from Fomes sclerodermeus, J. F. Lawrence (MCZC, 1); idem, collection date not noted, K. W. Cooper (WOPC, 1); idem, 22-28-VIII-1984, H. Wolda (MCZC, 1): Canal Zone: Paraiso, 2-IV-1911, E. A. Schwarz (USNM, 1).

Fig. 8.

Habitus of Ababa species. A. Ababa adona. B. A. epiiska. C. A. granaria. D. A granaria.

Fig. 8.

Habitus of Ababa species. A. Ababa adona. B. A. epiiska. C. A. granaria. D. A granaria.

Fig. 9.

Habitus and aedeagi of Ababa species. A. Ababa latimana. B. A. tantilla. C-F. Aedeagi. C. A. adona. D. A. epiiska. E. A. latimana. F. A. tantilla.

Fig. 9.

Habitus and aedeagi of Ababa species. A. Ababa latimana. B. A. tantilla. C-F. Aedeagi. C. A. adona. D. A. epiiska. E. A. latimana. F. A. tantilla.

  • Ababa epiiska, new species

    Figs. 6A, 6B, 7, 8B, 9D.

  • Diagnosis: The dark castaneous forebody (head and prothorax) distinguishes the members of this species from superficially similar specimens of Ababa latimana (Champion). Also, in specimens of A. epiiska the pronotal disc is conspicuously indented paralaterally, which is not the case in A. latimana specimens.

  • Description: Size: Length 2.8 mm; width 0.8 mm. Form: As in Fig. 8B. Color: Light castaneous, except forebody dark castaneous Head: Cranium imprinted with oblong punctures; eyes small, coarsely facetted, eye much narrower than frons (EW/FW 5/26); antenna capitate, funicular antennomeres subfiliform, progressively shorter towards capitulum, latter lax, capitular antennomeres 9 and 10 cupuliform, antennomere 11 spheroid. Thorax: Pronotum oblong campaniform; disc depressed at middle near outer margin and impressed with oblong punctures; interstitial spaces smooth; pronotal side margin slightly serrate (PW/PL 35/45); each elytral disc with shallow umbo near base, disc impressed with large setiferous punctures, punctures not arranged into linear striae, punctures extend to elytral apex (EL/EW 105/22). Abdomen: Pygidium scutiform; aedeagus as in Fig. 9D.

  • Natural History: Seven specimens from Bolivia were captured on fungi set on forest logs, at 366 m. One specimen was collected from Peru at 290 m.

  • Distribution: This species is known from Bolivia, Peru, and Argentina (Fig. 7).

  • Etymology: The trivial name, epiiska, is a Greek compound name that stems from epi (= upon) and iska (= a fungus on a tree); with reference to the niche from which some of these beetles were collected.

  • Specimens Examined: Holotype Male: Type locality: ARGENTINA: Provincia Salta: El Rey National Park, Pozo Verde Trail, km 10, 10-XII-1987, 1200 m, forest fungi on logs, S. and J. Peck (CMNC). Paratype: Eleven specimens. Locality information as in Holotype (AMNH, 1; CMNC, 3: WOPC, 1). BOLIVIA: Provincia de Santa Cruz: 3.7 km SSE Buena Vista, Hotel Flora and Fauna, 17°29.949′S 63°33.152′W, 5-15-XI-2001, tropical transition forest, M. C. Thomas and B. K. Dozier (CMNH, 1; FSCA, 2); idem, 17-20-X-2000, R. Morris (RFMC, 1). PERU: Provincia de Tambopata: Madre de Dios, Rio Tambota Reserve, 30 km SW Puerto Maldonado, 16-20-XI-1979, 290 m, subtropical moist forest, J. B. Heppner (FSCA, 1). PARAGUAY: Departamento Central: Cordillera San Bernardino, shore of Lago Ypacarai, 7-8-XI-1987, in black light trap, J. Kochalka (WOPC, 1).

  • Ababa granaria, new species

    Figs. 6A, 6B, 7, 8C.

  • Diagnosis: The interstitial spaces (spaces between punctures) of the pronotal disc are minutely arenose (Fig. 17). This characteristic will distinguish the members of this species from congeners. Also, from superficially similar specimens of Ababa tantilla LeConte, A. granaria specimens differ by showing a proportionally broader pronotum.

  • Description: Size: Length 2.8 mm; width 1.0 mm. Form: As in Fig. 16. Color: Light castaneous. Head: Cranium imprinted with oblong punctures; eyes small, coarsely facetted, eye much narrower than frons (EW/FW 5/30); antenna capitate, funicular antennomeres subfiliform, progressively shorter towards capitulum, latter lax, capitular antennomeres 9 and 10 cupuliform, antennomere 11 spheroid. Thorax: Pronotum slightly oblong campaniform; disc impressed with oblong punctures; interstitial spaces minutely arenose; pronotal side margin slightly roughened (PW/PL 44/47); elytral impressed with large setiferous punctures, punctures not arranged into linear striae, extend to elytral apex (EL/EW 105/25). Abdomen: Pygidium scutiform.

  • Distribution: This species is known from México (Fig. 13).

  • Etymology: The trivial name, granaria, is a Latin name with a meaning of “grain”; with reference to arenose character of the interstitial spaces of the pronotal disc.

  • Specimens Examined: Holotype Female: Type locality: MÉXICO: Flohr, no other label information (ZMHB). PARATYPE: One specimen. Intercepted in Philadelphia, USA, on banana trash from México, no other label information (WOPC, 1).

  • Ababa latimana (Champion, 1913: 95)

    Figs. 6A, 6B, 7, 9A, 9E.

  • Diagnosis: From superficially similar specimens of Ababa tantilla LeConte (pronotal width 0.8 mm), A. latimana specimens (pronotal width 0.5 mm) differ by showing a proportionally narrower pronotum. Also, the phallobasic lobes are acuminate in A. latimana specimens, which is not the case in specimens of A. tantilla.

  • Redescription: Size: Length 2.5 mm; width 0.8 mm. Form: As in Fig. 9A. Color: Light castaneous. Head: Cranium imprinted with oblong punctures; eyes small, coarsely facetted, eye much narrower than frons (EW/FW 5/25); antenna capitate, funicular antennomeres subfiliform, progressively shorter towards capitulum, latter lax, capitular antennomeres 9 and 10 cupuliform, antennomere 11 spheroid. Thorax: Pronotum slightly oblong campaniform, disc impressed with oblong punctures, side margin slightly roughened (PW/PL 42/ 46); elytral disc depressed behind basal umbo, impressed with large setiferous punctures, punctures not arranged into linear striae, punctures diminish at elytral 1/2 (EL/EW 95/25). Abdomen: Pygidium scutiform; aedeagus as in Fig. 9E.

  • Distribution: This species is known from Guatemala and Nicaragua (Fig. 7).

  • Biology: Specimens of this species have been collected in the fruiting bodies of fungi such as Tremetes cubensis Klotzsch (Polyporaceae), T. corrugata (Mont.) Sacc. (Polyporaceae), and Polyporus lignosus Klotzsch (Polyporaceae).

  • Specimens Examined: Lectotype, Sex Not Discernible: Here designated. Type locality: Guatemala, Rio Maria Linda, 4-III-1881, 500 ft., Champion (BMNH). PARALECTOTYPE: One specimen from Panamá, Volcan de Chiriquí (specimen not found). Gorham had two specimens available when he made his description, but he did not specify in his description which specimen should be the name barer of this species. Therefore, I invoke Recommendation 73F of the ICZN (1999) and designate a lectotype for this nominal species. Other Material Examined: Five specimens.

  • Guatemala: ex Polyporus lignosus, C. P. Secord, no other information (USNM, 1; WFBM, 1); ex Trametes cubensis, no other information (WOPC, 1); Departamento de Baja Verapaz: no other information (USNM. 1). NICARAGUA: Departamento de Estelí: 5 miles N. Estelí, 22-VII-1965, ex Tremetes corrugate, A. Raske (MCZC, 1).

  • Ababa tantilla LeConte, 1865: 96

    Figs. 6A, 6B, 7, 9B, 9F.

  • Diagnosis: From superficially similar specimens of A. latimana specimens (pronotal width 0.5 mm) Ababa tantilla LeConte (pronotal width 0.8 mm), differ by showing a proportionally broader pronotum. Also, the phallobasic lobes are obtuse in specimens of A. tantilla, they are acuminate in A. latimana specimens.

  • Redescription: Size: Length 2.2 mm; width 0.6 mm. Form: As in Fig. 19. Color: Light castaneous. Head: Cranium imprinted with oblong punctures; eyes small, coarsely facetted, eye much narrower than frons (EW/FW 5/26); antenna capitate, funicular antennomeres subfiliform, progressively shorter towards capitulum, latter lax, capitular antennomeres 9 and 10 cupuliform, antennomere 11 spheroid. Thorax: Pronotum slightly oblong campaniform, disc impressed with oblong punctures, side margin slightly roughened (PW/PL 37/ 40); elytral disc slightly depressed behind basal umbo, impressed with asetiferous punctures, punctures not arranged into linear striae, punctures extend to elytral apex (EL/EW 90/25). Abdomen: Pygidium scutiform; aedeagus as in Fig. 9F.

  • Distribution: This species is known from The United States of America southward to Panamá (Fig. 7).

  • Biology: Specimens of this species have been collected in the fruiting bodies of fungi such as Coriolopsis caperata (Berk.) Murrill (Polyporaceae), a species of Phellinus Quél. (Hymenochaetaceae), and bracket fungi.

  • Specimens Examined: Holotype, Sex Not Discernible: Type locality: United States Of America: Washington, D. C (MCZC). OTHER MATERIAL EXAMINED: Eighteen specimens. UNITED STATES OF AMERICA: Alabama: Dallas County: Selma (MCZC, 2); Illinois: No other information available (BMNH, 1): Texas: Liberty County: Near Liberty, ?-?-1919, H. S. Barber (USNM, 2); 22 miles N Liberty, 20-XI-1919, reared from Pyropolyperus ignarius, collector not noted (USNM, 1); Hidalgo County: 19-IX-4-X-1947, George B. Vogt (USNM, 1); Victoria County: 1.9 miles SW Raisin, 1-XII- 1973, G. Gaumer (WOPC, 1); Nacogdoches County: FSA, Expt1 Forest, 19-III-1972, in fungi, Jim Neal (AMNH, 1): Mississippi: Hancock County: 29-VIII-?, collector not noted (FMNH, 1; CMNH, 1): Louisiana: East Baton Rouge Parish: 23-I-1983, from Polyphora, D. A. Rider (WFBM, 1): Florida: Leon County: Tallahassee, 11-XI-1976, burlese hardwood litter, O'Brien and Marshall (CASC, 1): Franklin County: Apalachicola, 4-IV-8-VI-2008, D. R. Miller (UGCA, 1). MÉXICO: Estado de Veracruz: 4 km SW Fortin, 17-VII-1990, 900 m, J. Doyen (EMEC, 1); San Andres Tuxtla, 4-27-1073, 2400′, G. Ekis (WOPC, 1): Estado de San Luis Potosi: El Salto Falls, 12 km NW El Naranjo, 27-VII-1990, 400 m, J. S. Ashe, K. J. Ahn, R. Leschen (SEMC, 1). PANAMÁ: Canal Zone: Barro Colorado Island, 6-II-1968, on Polyporus caperatus, J. F. Lawrence (MCZC, 1)

DISCUSSION

Herein, I speculate about the function of an unusually long stomodaeum, characteristic of the checkered beetle subfamilies Thaneroclerinae and Isoclerinae (Ekis, 1971; Crowson, 1972; Opitz, 2010, 2014). I will link this unusual stomodeal formation with the known information about thaneroclerine and isoclerine natural history. The stomodaeum of the genera Thaneroclerus, Zenodosus, Ababa, and Parataneroclerus were studied. The specimens of these genera show a stomodaeum that is as long as the lengthy ventriculus. Moreover, the posterior limit of their stomodaeum is not bulbous or invested by a conspicuous, externally visible, layer of circular muscles typically found in those checkered beetle species that have a well-developed proventriculus. Three species of Chaetosomatidae, Chaetosoma scaritides, Somatochaetus quadraticollis and Metaxina ornata Broun also show expanded stomodaeums, but in these predatory species there is a significant swelling at the distal end of the stomodaeum, which suggests a well developed proventriculus, which in turn suggest a carnivorous life style that involves integumental components. A short stomodaeum and well-developed, highly muscular, proventriculus is characteristic of many Cleridae that feed on adult lignicolous prey (Opitz, 2010). The members of Chaetosomatidae are the closest living relatives of the Cleridae (Crowson, 1972).

Most of the Ababa specimens available for this study were collected in fungi niches such as those of: Tremetes cubensis Klotzsch (Polyporaceae), T. corrugata (Mont.) Sacc. (Polyporaceae), Polyporus lignosus Klotzsch (Polyporaceae), Coriolopsis caperata (Berk.) Murrill (Polyporaceae), and Phellinus igniarius (Linnaeus) Quél. (Hymenochaetaceae), and bracket fungi. It is reasonable to assume that the elongated stomodaeum, perhaps with the function of an extended crop (Snodgrass, 1935), and poorly developed proventriculus found in Ababa beetles is an indication that Ababa predation involves consumption of soft food, such as the eggs, larvae, and pupae of fungivorous prey. The same is probably true of those thaneroclerines that are known to feed on insects living in groceries, drugs, tobacco, and termite nests (Corporaal, 1938). It is interesting to note that Trametes fungi are food for caterpillars of certain Lepidoptera fungus moths such as Triaxomera parasitella; another possible soft-bodied food for thaneroclerines. All known Cleridae species that are predatory on adult insects, and whose internal anatomy was studied (Ekis, 1971; Opitz, 2014), show a shortened stomodaeum and highly muscular proventriculus.

Isoclerinae is characterized by the apotypic tarsal formula 5-4-4 (plesiotypic is 5-5-5). Within this subfamily, two groups can be distinguished based on the positional relationship between the pronotal projection (Fig. 2A) and prointercoxal process (Fig. 2B). There is no connection between these two prothoracic structures (Figs. 2B, 3A) in the New World isoclerine genera, Ababa and Parathaneroclerus, whereas in the Old-World genera, Compactoclerus, Cyrtinoclerus, Isoclerus, and Lyctosoma, the prointercoxal process and pronotal projection are firmly connected (as in Fig. 3B). The genus Allothaneroclerus Corporaal must be removed from Isoclerinae and placed into the subfamily Thaneroclerinae since the tarsal formula of A. tuberculatus (Schenkling) is 5-5-5 not 5-4-4. The disconnect between the prointercoxal process and pronotal projection is considered an apotypic character state and their connection is considered plesiotypic. However, this derived state must be considered a homoplastic apotypic state (Opitz, 2011) because in the Thaneroclerinae genus Zenodosus there is a wide gap between the two pronotal structures in question (Fig. 2A); these structures are tightly connected in Chaetosoma scaritides, a close relative of the Cleridae and a primitive element in the Cleroidea.

The exclusive South American presence of Parathaneroclerus Pic, the sister genus of Ababa Casey) (Kolibáč, 1998), is strong evidence that the ancestral stock of these two genera evolved in the forests of South America, perhaps in one of the montane Clerofaunas, as depicted by Opitz (2005). Such hardwood tree forests would have provided ample niches for fungal growths essential for the Ababa species. The conjecture of a South American origin for the Ababa species assumes four independent northern dispersals. This presumably occurred after the closure of the Panamanian Portal some 2.75 to 3 million years ago (Schmittner et al., 2004; Schneider and Schmittner, 2006) and before most of the uplift of the Colombian Andes some 6 to 12 million years ago (Gregory-Wodzicki, 2000). Three of the Ababa species, A. latimana, A. granaria, and A. tantilla, managed to cross the Nicaragua depression, which was a formidable barrier for more northern migrations (Whitehead and Ball, 1977). For an excellent discourse about isoclerine biogeography see Kolibáč (1998).

Figure 6B indicates weak bootstrap support, which undoubtedly reflects the paucity of characters used in this treatise. However, the phylogenetic interpretations are provided represent tentative relationships statements to be improved upon when addition differences in characters and character states are discovered. On the basis of geographic (Fig. 7) and morphologic analyses (Figs. 6A, 6B), we may predict that the ancestral stock of Ababa was characterized as follows: The ancestral terrain involved forests of South American highlands, body color was entirely light castaneous, eyes were very small and coarsely facetted, the antennal capitulum was lax, the profemur was swollen, asetiferous punctures were present on the elytral disc, and the phallobasic apodemes were fused. Ancestral Ababa (ancestor A, Fig. 6A) diversified to produce progenitor B and Ababa tantilla. Ancestor B evolved ancestor C and Ababa granaria characterized by having the interstitial spaces of the pronotum arenose. Further evolution involved the diversification of progenitor C into Ababa latimanus and progenitor D. The latter ancestor diverged to produce Ababa adona and Ababa epiiska.

Acknowledgments

I am very grateful to the curators, listed in the section of “Repositories of Specimens”, for entrusting me with collections in their charge. A special thanks to Douglass R. Miller and Jiří Kolibáč for their review of the manuscript. I thank Jean-Michel Maes for the translation of the abstract into Spanish, my wife Galena for technical assistance, Paul E. Skelley for departmental courtesies, and the Florida Department of Agriculture, DPI, for institutional affiliation and support.

Literature Cited

Literature Cited
Arnett
, Jr.,
R. H.
,
,
Samuelson
G. A.
, and
Nishida
,
G. M.
(
1993
).
The Insect and Spider collections of the World.
-
Second edition, Flora and Fauna Handbook no. 11 Sandhill Crane Press
,
Gainesville, FL, United States of America
,
310
pp.
Barr
,
W. F.
1975
.
Family 73. Cleridae
,
18
pp.
In
,
Arnett
,
Jr., R. H.
(ed.)
Checklist of the beetles of North and Central America and the West Indies, Volume 4.
The click beetles, fire-flies, checkered beetles, and related groups. North American Beetle Fauna Series. Flora and Fauna Publications
.
Gainesville, Florida, United States of America
.
Borchmann
,
F.
1910
.
Nilionidae, Othnidae, Aegialitidae, Petriidae, Lagraiidae
,
In
,
S.
Schenkling
(ed.).
Coleopterorum Catalogus.
Pars 2. W. Junk
,
Berlin, Germany
,
32
pp.
Bradley
,
J. C.
1930
.
A manual of the genera of beetles of America north of Mexico. Keys for the determination of the families, subfamilies, tribes, and genera of Coleoptera with a systematic list of the genera and higher groups.
Daw, Illston and Company
,
Ithaca, New York, United States ofAmerica
,
360
pp.
Brown
,
R. W.
1956
.
Composition of scientific words. 2nd ed.
-
Reese Press, Baltimore
,
Maryland, United States of America
,
882
pp.
Casey
,
T. L.
1897
.
Coleopterological Notices, VII.
Annals of the New York Academy of Sciences.
9
:
285
794
.
Champion
,
C.
1913
.
Notes on various Central American Coleoptera, with descriptions of new generaand species.
Transactions of the Entomological Society of London-Part I (June)
1913
:
58
169
.
Corporaal
,
J. B.
1939
.
Revision of the Thaneroclerinae (Cleridae, Col.).
Bijdragen Tot de Dierkunde
27
:
347
359
.
Crowson
,
R. A.
1972
.
On the systematic value of the alimentary canal of the Cleridae.
Systematic Zoology
21
:
339
340
.
de Queiroz
K
.
2007
.
Species concepts and species delimitation.
Systematic Biology
56
(
6
):
879
886
.
Dobzhansky
T
.
1937
.
Genetics and the origin of species.
Columbia University Press
,
New York, New York, United States of America
,
364
pp.
Ekis
,
G.
(now
Opitz
,
W.
) and
Gupta
,
A. P.
1971
.
Digestive system of Cleridae (Coleoptera).
International Journal of Insect Morphology and Embryology
1
(
1
):
51
86
.
Ekis
,
G.
(now
Opitz
,
W.
).
1977
.
Classification, Phylogeny, and Zoogeography of the Genus Perilypus (Coleoptera: Cleridae).
Smithsonian Contributions to Zoology
227
:
1
138
.
Goloboff
,
P. A.
2003
.
NONA version 2.0. A tree searching program.
Distributed by the author
.
Gregory-Wodzicki
,
K. M.
2000
.
Uplift history of the Central and Northern Andes: A review.
Geologic Society of America Bulletin
112
(
7
):
1091
1105
.
Hennig
,
W.
1966
.
Phylogenetic Systematics.
University of Illinois Press
,
Urbana, IL, United States of America
.
263
pp.
(ICZN) International Code of Zoological Nomenclature
.
1999
. 4th Edition.
Adopted by the International Union of Biological Sciences.
The International Trust for Zoological Nomenclature
,
London, United Kingdom
,
306
pp.
Kolibáč
,
J.
1992
.
Revision of Thanerocleridae, n.stat. (Coleoptera, Cleroidea).
Mitteilungen der Schweizerischen Entomologischen Gesellschaft
65
:
303
340
.
Kolibáč
,
J.
1998
.
New Australian Thanerocleridae, with notes on the biogeography of the subtribe Isoclerinae Kolibáč (Coleoptera: Cleroidea).
Invertebrate Taxonomy
12
:
951
975
.
LeConte
,
J. L.
1865
.
New species of North American Coleoptera.
Smithsonian Miscellaneous Collections
6
(
167
):
87
177
.
Lewis
,
F. L. S.
1892
.
On the Japanese Cleridae. Annals and Magazine of Natural History.
Series 6.
10
:
183
192
Lucas
,
R.
1920
.
Catalogus alphabeticus generum et subgenerum Coleopterorum orbis terrarium totius.
Archiv für Naturgeschichte. Pars I. 84(1918): 669.
Verlags-Buchhandlung R. Stricker
,
Berlin, Germany
,
696
pp.
Mayr
,
E.
1963
.
Animal species and evolution.
Harvard University Press
,
Cambridge, Massachusetts, United States of America
,
797
pp.
Nixon
,
K. C.
2002
.
Winclada ver. 1.00.08.
Published by the author
.
Nixon
,
K. C.
and
J. A.
Carpenter
.
1993
.
On outgroups.
Cladistics
9
:
413
426
.
Opitz
,
W.
2002
.
Family 73. Cleridae Latreille 1804.
Pp.
267
280
.
In
,
Arnett
,
R. H., Jr.
,
Thomas
,
M. C.
,
Skelley
,
P. E.
and
Frank
,
J. H.
(eds.)
2002
.
American Beetles. Volume 2
.
CRC Press. Boca Raton
,
Florida, United States of America
,
861
pp.
Opitz
,
W.
2005
.
Classification, natural history, and evolution of the genus Aphelocerus Kirsch(Coleoptera: Cleridae: Clerinae).
Bulletin of the American Museum of Natural History
293
:
1
128
.
Opitz
W
.
2010
.
Classification, evolution, and subfamily composition of the Cleridae, and generic content and key of the subfamilies (Coleoptera: Cleroidea).
Entomologica Basiliensia et Collectionis Frey
32
:
31
128
.
Opitz
,
W.
2011
.
Classification, natural history, and evolution of the Epiphloeinae (Coleoptera: Cleridae). Part X. The genus Madoniella Pic.
Entomologica Basiliensia et Collectionis Frey
33
:
133
248
.
Opitz
,
W.
2014
.
Morphologic studies of the alimentary canal and internal reproductive organs of the Chaetosomatidae and the Cleridae (Coleoptera: Cleroidea) with comparative morphology and taxonomic analyses.
Insecta Mundi
0342
:
1
40
.
Opitz
,
W.
2018
.
Classification, natural history, and evolution of the Korynetinae (Coleoptera: Cleridae). Part IV. The new genus Nolafigura Opitz, and fourteen new species of Korynetes.
Faunitaxys
6
(
12
):
1
17
.
Peck
,
S. B.
and
M. C.
Thomas
.
1998
.
A distributional checklist of the beetles (Coleoptera) of Florida. Arthropods of Florida and neighboring land areas. Volume 16.
Florida Department of Agriculture and Consumer Services
.
Gainesville, Florida, United States of America
,
180
pp.
Pic
,
1947
.
Coléoptères du globe.
L'Échange, Revue Linnéenne
63
(
508
):
5
8
.
Schaeffer
,
C.
1917
.
On some North American Cleridae (Col.).
Journal of the New York Entomological Society
25
(
2
):
129
134
.
Schmittner
,
A.
,
Sarnthein
,
M.
,
Kinkel
,
H
,
Bartoli
,
G.
,
Bickert
,
T.
,
Crucifix
,
M.
,
Crudeli
,
D.
,
Groeneveld
,
J.
,
Kosters
,
F.
,
Mikolajewicz
,
U.
,
Millo
,
C.
,
Reijmer
,
J.
,
Schafer
,
P.
,
Schmidt
,
D.
,
Schneider
,
B.
,
Schultz
,
M.
,
Steph
,
S.
,
Tieddemann
,
R.
,
Weinelt
,
M.
, and
Zuvela
,
M.
2004
.
Global impact of the Panamanian seaway closure.
EOS
85
:
526
.
Schneider
,
B.
and
Schmittner
,
A.
2006
.
Simulating the impact of the Panamanian seaway closure on ocean circulation, marine productivity and nutrient cycling.
Earth and Planetary Science Letters
264
:
367
380
.
Snodgrass
,
R. E.
1935
.
Principles of insect morphology.
McGraw-Hill Book Company
,
New York, United States of America
,
667
pp.
Standfuss
M
.
1896
.
Handbuch der paläarktischen GrossSchmetterlinge für Forscher und Sammler.
Gustav Fischer
,
Jena, Germany
,
392
pp.
Tuomikoski
,
R.
1967
.
Notes on some principles of phylogenetic systematics.
Annales Entomologici Fennici
33
(
3
):
137
147
.
Watrous
,
L. E.
and
Q. D.
Wheeler
.
1981
.
The outgroup comparison method of character analysis.
Systematic Zoology
30
:
1
11
.
Whitehead
,
D. R.
and
G. E.
Ball
.
1977
.
The Middle American genus Onypterygia Dejean (Insecta: Coleoptera: Carabidae: Platynini): A taxonomic revision of the species, with notes about their way of life and geographical distribution.
Annals of Carnegie Museum
66
(
3
):
289
409
.
Wolcott
,
A. B.
1911
.
New American Cleridae, with notes on others (Col.).
Entomological News
22
:
115
125
.
Wolcott
,
A. B.
1947
.
Catalogue of North American beetles of the family Cleridae.
Fieldiana: Zoology
52
(
2
):
63
105
.