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Classification, natural history, and evolution ofNeorthopleurinae Opitz (Coleoptera, Cleridae). Part III. TheGenera Agaphalera Opitz, Allochotes Westwood, KataspinulaOpitz, Lebasiella Spinola, Loedelia Lucas, Patuleius Fairmaire,Rifkindius Opitz, and Romanaeclerus WinklerAuthor(s): Weston OpitzSource: Pan-Pacific Entomologist, 89(4):244-258. 2013.Published By: Pacific Coast Entomological SocietyDOI: http://dx.doi.org/10.3956/2013-20.1URL: http://www.bioone.org/doi/full/10.3956/2013-20.1

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Classification, natural history, and evolution of Neorthopleurinae Opitz(Coleoptera, Cleridae). Part III. The Genera Agaphalera Opitz,

Allochotes Westwood, Kataspinula Opitz, Lebasiella Spinola, LoedeliaLucas, Patuleius Fairmaire, Rifkindius Opitz, and

Romanaeclerus Winkler

WESTON OPITZ

Kansas Wesleyan University, Department of Biology, 100 East Claflin Avenue,

Salina, Kansas 67401-6196 U.S.A.

e-mail: opitz@kwu.edu

Abstract. A treatise involving eight genera that form a monophyletic group within

Neorthopleurinae is presented. Three new species of the subfamily are described: Agaphalera

corallina, sp. nov., A. cymatilis, sp. nov., and Loedelia westcotti, sp. nov. An intergeneric

phylogeny is proposed and a key to genera is included. Keys to the species of Agaphalera Opitz2009b and Loedelia Lucas 1920 are included.

Key Words. Coleoptera, Cleridae, Neorthopleurinae, checkered beetles, new species,

taxonomy, phylogeny.

INTRODUCTION

This is the third of a series of works that attempts to make better known the

checkered beetle taxa of subfamily Neorthopleurinae Opitz. In essence, the work

represents a spinoff project from a more comprehensive treatise that reassesses the

subfamilial classification of World Cleridae (Opitz 2010). The eight genera treated

herein form a monophyletic group among the 22 genera that comprise the

Neorthopleurinae Opitz. The more speciose genus of this monophyletic group,

Allochotes Westwood 1875, is currently under revision by other authors. For now,

keys are provided of the less speciose genera to which new species are added.

SPECIMENS AND METHODS

This study is based on 169 specimens. The species descriptions are brief while the

generic characteristics are comprehensively described by Opitz (2009b). Generic level

morphological diversity is extensive among many groups of the Cleridae, whereas

species level diversity in often minimal. The species concept follows Standfuss (1896:

115) and Mayr (1963: 19). Methods involving dissection, measurements, descriptive

terminology, and techniques of illustrations follow Opitz (2010: 35).

Specimens are deposited in the British Museum of Natural History (BMNH)

(Maxwell V. L. Barclay), California Academy of Sciences (CASC) (David H.

Kavanaugh), California State Collection of Arthropods (CSCA) (Stephen Gaimari),

Canadian Museum of Nature (CMNC) (Robert S. Anderson), Field Museum of

Natural History (FMNH) (James H. Boone), Florida State Collection of Arthropods

(FSCA) (Michael Thomas, Paul E. Skelley), Jacques Rifkind Collection (JNRC),

Kansas University-Biodiversity Institute (SEMC) (Zachary Falin), Museum d’His-

toire Naturelle (MNHN) (Antoine Mantilleri), Texas A & M University (TAMU)

(Edward G. Riley), University of California, Riverside (UCRC) (Doug Yanega), Utah

THE PAN-PACIFIC ENTOMOLOGIST89(4):244–258, (2013)

State University (EMUS) (Carol D. VanDohlen), Weston Opitz Collection (WOPC),

and William F. Barr Entomology Museum (WFBM) (Frank W. Merickel).

PHYLOGENETICS OF GENERA

The tenets involving phylogenetic analysis follow Hennig (1966) and the character

matrix (Table 1) was analyzed via NONA (Goloboff 2003) in combination with

Winclada version 1.00.08 (Nixon 2002). The analysis resulted in a single tree

(Fig. 25), which involves 20 steps, index of consistency of 100, and an index of

retention of 100. Heuristic analysis [maximum tree (hold)] 5 100, number of

replications 9 (mult) 5 100, and multiple TBR (mult max) was used.

CHARACTER STATES

Twenty morphological and two geographical character states were used to analyze

the evolutionary relationships among the eight genera included in this treatise.

Outgroups involve the remaining 14 genera of Neorthopleurinae (Opitz 2009b).

Character states assessed ‘‘0’’ are considered plesiotypic (Tuomikosky 1967), while

those evaluated as ‘‘1’’ are judged apotypic (Table 1). Methods that involve

determination of the evolutionary states of characters are amply noted in Ekis (now

Opitz) (1977: 166) and Nixon and Carpenter (1993: 413).

Table 1. Character matrix for 22 morphological characters of genera included in this study.

Taxa Characters

0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21Outgroup 1 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0Agaphalera 1 1 1 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0Allochotes 1 1 1 0 1 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0Kataspinula 1 1 1 0 0 1 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0Lebasiella 1 1 1 1 0 0 1 1 0 0 0 0 0 0 0 1 1 1 0 0 0 0Loedelia 1 1 1 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1Patuleius 1 1 1 0 1 0 0 1 0 0 1 1 0 0 0 0 0 0 0 0 0 0Rifkindius 1 1 1 1 0 0 1 1 0 0 0 0 0 0 0 1 0 0 1 1 0 0Romanaeclerus 1 1 1 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0

Character 0 Pronototergostenal suture: (0) complete; (1) not complete

Character 1 Shape of postgular process: (0) not petiolate; (1) petiolate

Character 2 Body form: (0) oblong long; (1) oblong shortCharacter 3 Ovipositor: (0) not very long; (1) very long

Character 4 Epipleuron: (0) not partially inflexed; (1) partially inflexed

Character 5 Distribution: (0) Old World; (1) New World

Character 6 Asetiferous punctations: (0) seriate; (1) aseriate

Character 7 Hind body form: (0) not oval; (1) oval

Character 8 Elytral humerus: (0) not bulging; (1) bulging

Character 9 Phallic plate: (0) not spinous; (1) spinous

Character 10 Land: (0) continental; (1) insularCharacter 11 Epipleuron: (0) not extensively ridged; (1) extensively ridged

Character 12 Apex of tibiae: (0) without bifid setae; (1) with bifid setae

Character 13 Phallic membrane: (0) without spines; (1) with spines

Character 14 Tibial spur formula: (0) 2-2-2; (1) 1-1-1

2013 OPITZ: TAXONOMY OF NEORTHOPLEURINAE (COLEOPTERA: CLERIDAE) 245

KEY TO GENERA

1. Last maxillary palpomere digitiform. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

- Last maxillary palpomere not digitiform . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

2. Elytral asetiferous punctations aseriate . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

- Elytral asetiferous punctations seriate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

3. Seventh antennomere larger than eight (Mexico) . . . . . Rifkindius Opitz 2009b

- Seventh antennomere not larger than eight . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. Extremity of prointercoxal process triangular; pronotal projections nearly

touch prointercoxal process; forebody yellow (Costa Rica, Honduras,

Mexico, Nicaragua, U.S.A.). . . . . . . . . . . . . . . . . . . . Lebasiella Spinola 1844

- Extremity of prointercoxal process not triangular; pronotal projections

distant from prointercoxal process; forebody black (Mexico) . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kataspinula Opitz 2009b

5. Epipleuron inflexed; funicular antennomeres not widened (Madagas-

car) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Patuleius Fairmaire 1902

- Epipleuron not inflexed, in ventral position; funicular antennomeres

widened (Cameroon, Democratic Republic of the Congo, Cote D’Ivoire,

Equatorial Guinea, Kenya, Sao Tome, South Africa, Tanzania). . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Romanaeclerus Winkler 1960

6. Body rotund; antenna serrate (Bhutan, Brunei, Burma, Cambodia, China,

India, Japan, Malaysia, Nepal, North Vietnam, New Guinea, Philippines,Sri Lanka, Sumatra) . . . . . . . . . . . . . . . . . . . . . . Allochotes Westwood 1875

- Body not rotund; antenna not serrate, capitates . . . . . . . . . . . . . . . . . . . . . 7

7. Tarsal unguis bifid (Mexico, U.S.A.) . . . . . . . . . . . . . . . Loedelia Lucas 1920

- Tarsal unguis not bifid (Mexico) . . . . . . . . . . . . . . . Agaphalera Opitz 2009b

TAXONOMY

One intent of this publication is to elucidate the phylogenetic relationships among

the eight genera listed in the title. The phylogeny of the other 14 genera of

Neorthopleurinae is presented in part II of this series of publications (Opitz in

preparation) whose purpose is to bring to light Neorthopleurinae systematics. The

genera included in this work were recently described and copiously illustrated

elsewhere (Opitz 2009b). In this treatise I make known generic level phylogenetics,

increase the species inventory of Neorthopleurinae by three, and present for the firsttime keys of the known species of Agaphalera and Loedelia. A generic description of

Agaphalera and Loedelia is provided to assure a continuance of understanding of

those generic-level characteristics relevant to the new species descriptions. Rifkindius

and Kataspinula remain monotypic. It needs to be clarified that Opitz (2009b: 185,

Character 15 Intraspicular plate: (0) present; (1) absent

Character 16 Dorsal plate of phallobase: (0) not elongated; (1) elongated

Character 17 Funicular antennomere 5: (0) not larger than six; (1) larger than six

Character 18 Tegmen: (0) lobed distally; (1) not lobed distallyCharacter 19 Antennomeres 9 and 10: (0) not acuminate anterodistally; (1) acuminate

anterodistally

Character 20 Basal denticle: (0) small; (1) large

Character 21 Unguis: (0) not bifid; (1) bifid

246 THE PAN-PACIFIC ENTOMOLOGIST Vol. 89(4)

2010: 50) erroneously noted that the tarsal pulvillar formula for Romanaeclerus asbeing 0-0-0, when in fact it is 2-2-2 (Opitz 2009a). Taxonomically, what remains to be

done with this monophyletic branch of Neorthopleurinae is to comprehensively

revise the Asiatic genus Allochotes and the Madagascan genus Patuleius.

Agaphalera Opitz (Opitz 2009b:142)

(Figs. 2, 3, 7–10, 13–15, 19, 21, 22)

Type Species. Lebasiella janthina LeConte 1866:99.

Diagnosis. Specimens of Agaphalera resemble superficially those of Loedelia. But,

in Agaphalera specimens the denticle of the unguis is large and somewhat truncate

(Fig. 19) not acuminate as in specimens of Loedelia (Fig. 20).

Figure 1. Habitus of Loedelia maculicollis (LeConte).

2013 OPITZ: TAXONOMY OF NEORTHOPLEURINAE (COLEOPTERA: CLERIDAE) 247

Description. Size: Length 3.2–5.5 mm; width 1.4–2.4 mm. Form (Figs. 21, 22):

Oblong short, robust, deep bodied, about 2 times longer than broad. Vestiture: Disc

of cranium and pronotum densely vested with stout setae, elytra vested only with 1usetae. Head (Fig. 13): Cranium subspheroid, frons very wide, indented with shallow

setiferous punctations, latter not widely separated; gula (Fig. 15) very small, process

short, narrow, and minutely forked; labrum short, medial incision curvate concave;

mandible, body short, anterior, medial, and posterior dens well developed, penicillus

well developed; maxilla, laterolacinia present, terminal palpomere subsecuriform

(Fig. 14); labium, terminal palpomere narrow triangular; eyes small, finely facetted;

antenna (Figs. 2, 3) capitate, capitulum lax and short, capitular antennomeres 9 and

Figures 2–12. Various organs. 2–4 Antennae. 2) Agaphalara cymatilis. 3) A. corallina. 4)Loedelia westcotti. 5–6) Mesodermal reproductive organs of Loedelia maculicollis (5 male, 6 female).7–12) Aedeagi. 7–8) Agaphalera corallina (7 phallus, 8 tegmen). 9–10) A. cymatilis (9 phallus, 10tegmen). 11–12) Loedelia westcotti (11 phallus, 12 tegmen).

248 THE PAN-PACIFIC ENTOMOLOGIST Vol. 89(4)

10 subacuminate at their anterodistal angle. Thorax: Pronotum transverse, convex,

side margins evenly rounded, sculptured with very small round setiferous

punctations, prointercoxal process not expanded distally; pronotal projections

short, do not contact prointercoxal process; elytron sculptured with small asetiferous

Figures 13–15. Various organs of Agaphalera janthina. 13) Head. 14) Mouthparts. 15)Gular process.

2013 OPITZ: TAXONOMY OF NEORTHOPLEURINAE (COLEOPTERA: CLERIDAE) 249

punctations, latter aseriate, 1u setae always adjacent to asetiferous punctations, 2usetae absent, epipleural fold expanded in proximal half and extended to elytral distaltwo-thirds; metendosternite with furcal lamina; legs, tibial spur formula 2-2-2, tarsal

pulvillar formula 3-3-3, unguis with large truncate denticle. Abdomen: Aedeagus

shorter than length of abdomen, phallobasic lobes minutely fimbriate; lateral plates

Figures 16–18. Various organs of Loedelia mexicana. 16) Head. 17) Mouthparts. 18) Gular process.

250 THE PAN-PACIFIC ENTOMOLOGIST Vol. 89(4)

of spicular fork acuminated laterally, spicular apodemes fused, interspicular plate

minute and rod-shaped; ovipositor, ventral and dorsal laminae multilobed, laminal

rod present; distal margin of pygidium and 6th visible sternite rounded, not incised.

Alimentary Canal: Stomodaeal valve comprised of 4 primary lobes. Mesodermal

Male Internal Reproductive Organs: No information available. Mesodermal Female

Internal Reproductive Organs: No information available.

Agaphalera corallina, sp. nov.

(Figs. 3, 7, 8, 21)

Holotype -. Mexico, Puebla, Hwy. 190, 7 km SSE Acatlan, 1280 m, 5 July 1992,

C. L. Bellamy, colls. (LACM).

Paratypes. Ten specimens. Mexico: Guerrero: 9.6 km NE Tixtla, 16-VII-1984, J.

Woolley (WOPC, 2); 9.6 km E Xochipala, 18-VII-1985, J. B. Wooley (WOPC, 1):

Puebla: Hwy. 190, 7 km SSE Acatlan, 5-VII-1992, 1280 m, C. L. Bellamy (JNRC, 3;

WOPC 2); idem, R. L. Wescott (WOPC, 1): Morelos: Tlaquiltenango, Huaxtla,

Figures 19–20. Ungues. 19) Agaphalera janthina. 20) Loedelia mexicana.

2013 OPITZ: TAXONOMY OF NEORTHOPLEURINAE (COLEOPTERA: CLERIDAE) 251

18 37486u N, 99 05642u W, Selva Baja Caducifolia, Golpeando vegetacion, 22-VII-

2009, V. H. Toledo (JNRC, 1).

Diagnosis. Specimens of this species resemble superficially the members of

Agaphalera cymatilis, sp. nov., but in specimens of A. corallina the epipleural margins

are subconvex, whereas in cymatilis specimens the epipleural margins are subparallel.

Description. Size: Length 5.0 mm; width 2.0 mm. Form (Fig. 21): Integumental

Color: Antennae and terminal palpomeres brown, forebody and legs yellow-brown,tarsi infuscated, elytra, pterothoracic venter, and abdomen dark blue. Integumental

Structure: Cranium and pronotum minutely punctated; elytral surface punctations

shallow, punctations aseriate; antenna capitate, capitulum short and lax, funicular

Figures 21–24. Habiti. 21) Agaphalera corallina. 22) A. cymatilis. 23) Loedelia westcotti. 24)Loedelia westcotti.

252 THE PAN-PACIFIC ENTOMOLOGIST Vol. 89(4)

antennomeres subfiliform; pronotum convex, side margins slightly convex;

epipleural margins somewhat parallel in basal half, then rapidly incurved towards

apex. Male Genitalia (Figs. 7, 8): Aedeagus slightly sclerotized; tegmen bilobed

distally, lobes fimbriate; phallic apex digitiform; phallobasic rod large and cordate;

phallobasic apodeme broadly dilated distally.

Variation. The elytral punctations are slightly more pronounced in one specimen

from Acatlan.

Etymology. The trivial name corallinus (5 coral-red) is a Latin adjective. I refer to

the coloration of the forebody of this beetle.

Agaphalera cymatilis, sp. nov.

(Figs. 2, 9, 10, 22)

Holotype U. Mexico, Colima, Lago La Maria, 41009, 22 July 1995, beating Acacia,

J. Rifkind, A. Reifschneider, colls. (LACM).

Paratypes. Seven specimens. Mexico: Colima: Lago La Maria, 22-VII-1995, beatingAcacia, 1250 m, J. Rifkind, A. Reifschneider (WOPC, 1): Jalisco: 12 km S Autlan, 16-

VII-1990, E. Giesbert (WOPC, 1): Nayarit: 15 km N of Chapalilla, 24-VII-1993, beating

Acacia pennatula, oak/pine forest, Rifkind, Bellamy, Reifschneider (WOPC, 1); idem,

25-VII-1993 (JNRC, 2): Jalisco: 12 km S Autlan, 16-VII-1990, E. Giersbert (FSCA, 2).

Diagnosis. See previous species.

Description. Size: Length 5.0 mm; width 1.8 mm. Form (Fig. 22): IntegumentalColor: Antennae (Fig. 2) and terminal palpomeres brown, forebody yellow-brown,

legs brown, elytra, pterothoracic venter, and abdomen dark blue. Integumental

Structure: Cranium and pronotum minutely punctated; elytral surface deeply

punctated, punctations aseriate; antenna capitate, capitulum short and lax, funicular

antennomeres subfiliform; pronotum convex, side margins strongly convex;

epipleural margins somewhat parallel in basal three-fourths, then gradually incurved

towards apex. Male Genitalia (Figs. 9, 10): Aedeagus slightly sclerotized; tegmen

bilobed distally, lobes fimbriate; phallic apex digitiform; phallobasic rod small andcordate; phallobasic apodeme broadly dilated distally.

Variation. The mesoscutellum and legs of one of the paratypes are yellow-brown.

Etymology. The trivial name cymatilis (5 blue) is a Latin adjective. I refer to the

predominantly blue coloration of this beetle.

KEY TO THE SPECIES OF AGAPHALARA OPITZ

1. Pronotum yellow-brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

- Pronotum blue-black. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

2. Epipleural margins subparallel; hindbody somewhat rectangulate (Fig. 22)

(Mexico) . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Agaphalera cymatilis, sp. nov.

- Epipleural margins subconvex; hindbody somewhat oval (Fig. 21) (Mexico). . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Agaphalera corallina, sp. nov.

3. Elytral humeral angle yellow-brown (Mexico). . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . Agaphalera quadrimaculata (Pic 1939)

- Elytral humeral angle blue-black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. Elytral disc with a violaceous tinge, never with yellow macula (Mexico). . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . Agaphalera janthina (LeConte 1866)

- Elytral disc with blue tinge, often with yellow macula (Mexico). . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . Agaphalera unimaculata (Pic 1940)

2013 OPITZ: TAXONOMY OF NEORTHOPLEURINAE (COLEOPTERA: CLERIDAE) 253

Loedelia Lucas 1920:380

(Figs. 1, 4–6, 11, 12, 16–18, 20, 23, 24)

Type species. Necrobioides mexicana Gahan 1910:76; Wolcott 1947:88; Corporaal

1950:312; Opitz 2002:280, 2009b:161.

Necrobioides Gahan 1910:76.

Diagnosis. Within Neorthopleurinae, only in members of Loedelia is the denticle of

the ungues acuminate (Fig. 20).

Description. Size: Length 5.0–6.0 mm; width 2.0–3.0 mm. Form (Fig. 1): Oblong

short, robust, deep body, about 2 times longer than broad. Vestiture: Disc of

cranium and pronotum densely vested with stout setae, elytra vested only with 1usetae. Head (Fig. 16): Cranium subspheroid, frons very wide, indented with very

small setiferous punctations, latter widely separated; gula (Fig. 18) very small,

sutures oblique, gular process short, narrow, and minutely forked; labrum short,

medial incision curvate concave, medial tormal processes transverse confluent,

epipharyngeal plate not distinguishable; mandible, body short, anterior, medial, and

posterior dens well developed, penicillus very well-developed; maxilla, laterolacinia

present, terminal palpomere subsecuriform (Fig. 17); labium, terminal palpomere

narrow triangular; eyes small, very finely facetted; antenna (Fig. 4) capitate,

capitulum lax and short. Thorax: Pronotum transverse, convex, side margins evenly

rounded, sculptured with very small round setiferous punctations, prointercoxal

process not expanded distally; pronotal projections short, do not contact

prointercoxal process; elytron sculptured with small circular asetiferous punctations,

latter aseriate, elytral 1u setae always adjacent to asetiferous punctations, 2u setae

absent; epipleural fold expanded in proximal half and extended to elytral distal two-

thirds; metendosternite with very small furcal lamina, furcal anterior plate extended;

legs, tibial spur formula 2-2-2, tarsal pulvillar formula 3-3-3, unguis bifid, basal

denticle acuminate. Abdomen: Aedeagus (Figs. 11, 12) shorter than length of

abdomen, phallobasic lobes profusely fimbriate; apices of spicular fork lateral plates

broadened, spicular apodemes fused together, intraspicular plate minute and rod-

shaped; ovipositor, ventral and dorsal laminae multilobed, laminal rod not present;

distal margin of pygidium and 6th visible sternite rounded, not incised. Alimentary

Canal: Six cryptonephridial Malpighian tubules. Mesodermal Male Internal

Reproductive Organs (Fig. 5): Two pairs of accessory glands, medial pair biramous.

Mesodermal Female Internal Reproductive Organs (Fig. 6): Spermathecal capsule

barrel shaped, well sclerotized, spermathecal gland attached to apex of spermathecal

capsule.

Loedelia westcotti, sp. nov.

(Figs. 11, 12, 23, 24)

Holotype -. Mexico, Baja Calif. SUR, SE Shore Bahia Concepcion, 23-IV-1985,

beat Prosopis in bloom, R. L. Westcott collector (CASC).

Paratypes. Nine specimens. Mexico: Baja California Sur: SE Shore Bahia

Concepcion, 23-IV-1985, beat Prosopis in bloom, R. L. Westcott (JNRC, 1; WFBM,

2; WOPC, 3); idem, 1.2 rd. mi. E Rosario, 5-IX-1988, blacklight, E. G. Riley

(TAMU, 2; WOPC, 1).

Diagnosis. In the members of this species the elytral disc is mostly yellow-brown,

which is unique in the genus.

254 THE PAN-PACIFIC ENTOMOLOGIST Vol. 89(4)

Description. Size: Length 5.5 mm; width 2.2 mm. Form (Figs. 23): Integumental

Color: Antennae, tarsi, and terminal palpomeres dark brown, remainder of

integument mostly yellow-brown, and with infuscation on cranial vertex, large area

of pronotal disc, distally on femora, mesepisternum, metepisternum, elytral disc

heavily infuscated at humeral angle, sutural margin near and including mesoscu-

tellum, and before elytral apex. Integumental Structure: Cranium and pronotum

minutely punctated; elytral surface punctations shallow, punctations aseriate,

interstitial spaces arenose; antenna capitate, capitulum short and lax, funicular

antennomeres subfiliform; pronotum convex, side margins slightly convex;

epipleural margins somewhat convex. Male Genitalia (Figs. 11, 12): Aedeagus

slightly sclerotized; tegmen bilobed distally, lobes fimbriate; phallic apex digitiform;

phallobasic rod large and bifid distally.

Variation. The expression of the black infuscations on the elytral and pronotal disc

is very variable. Sometimes this infuscation on the elytra is reduced to small

midbasal macula (Fig. 24).

Etymology. The trivial name is a dedicative epithet to honor Richard L. Westcott

for his many contributions to field Entomology.

KEY TO THE SPECIES OF LOEDELIA LUCAS

1. Pronotum concolorous dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

- Pronotum bicolorous or completely yellow . . . . . . . . . . . . . . . . . . . . . . . . . 3

2. Elytral disc concolorous, blue-black often with a violaceous tinge (Mexico). . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Loedelia mexicana (Gahan 1910)

- Elytral disc bicolorous, mostly black, with a yellow macula (Mexico). . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Loedelia peninsularis Barr 1950

3. Elytral disc mostly yellow-brown (Figs. 23 and 24) (Mexico) . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Loedelia westcotti, sp. nov.

- Elytral disc entirely dark colored . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. Pronotal disc mostly yellow-brown and with a central spheroid macula

(U.S.A.) . . . . . . . . . . . . . . . . . . . . . . . . . . Loedelia discoidea (LeConte 1881)

- Pronotal disc variously infuscated, infuscation usually extended from

anterior margin of pronotum (Mexico, U.S.A.) . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . Loedelia maculicollis (LeConte 1874)

EVOLUTIONARY CONSIDERATIONS

Perhaps the most outstanding information intrinsic in the phylogenetic hypothesis

of the taxa in this treatise (Fig. 25) is their clear separation into two geographical

components; the Old World genera and the New World genera. Such distributional

and evolutionary relationship evidence suggest that the taxa under consideration

have had an extensive evolutionary history that might have extend back to an era

when the continents were amassed into the great ancient Southern Hemisphere

land mass known as Gondwanaland. The evolution and diversification of these

neorthopleurines are probably linked to ancient angiosperm proliferations and to the

rise of ancient forests of gymnosperms. It is widely known that as a group the

Cleridae is mostly xylophilous (Balduf 1935) and that angiosperms flourished and

radiated throughout Gondwanaland (White 1988). Other discussions that link

ancestral Cleridae to a probable Gondwanan origin is found in Opitz (2003).

2013 OPITZ: TAXONOMY OF NEORTHOPLEURINAE (COLEOPTERA: CLERIDAE) 255

There are two synapotypies (Kavanaugh 1978, Opitz 2007) that link the generic

taxa treated herein to their sister outgroup, a petiolate shape of the post-gular

process and incomplete pronotosternal. The first divergence of the Gondwanan

ancestor (ancestral species A) produced a line of neorthopleurines in which the

ovipositor became greatly lengthened [the outgroup, Dermestoides generic group

(Opitz, in press)]. The second line (ancestral species B) evolved an oblong-short body

and became dichotomously distributed. The Old World group generated taxa

towards ancestor C, in which the epipleuron became inflexed, whereas in the New

World genera the elytral punctations became aseriate. Subsequently ancestor C

proliferated the Romanaeclerus group in which the apex of the tibiae generated bifid

setae. This ancestor also generated progenitor D in which the hind body took on a

more oval shape. Then ancestral species D diverged toward the Allochotes group, in

Figure 25. Phylogenetic diagram of genera.

256 THE PAN-PACIFIC ENTOMOLOGIST Vol. 89(4)

which the humerus became bulgy and the phallic plate spinous, and the Madagascan

Patuleius group, in which the epipleuron became extensively ridged.

The ancestral stock of the New World taxa (ancestral species E) proliferated the

Mexican Agaphalera group, characterized by a large basal denticle and the anterodistal

angle of antennomere 9 and 10 became subacuminate. Ancestor E also generated

ancestor F, which eventually evolved the progenitors of the Loedelia group, in which

the unguis became bifid, and the progenitors of ancestor G. The latter evolved theMexican Kataspinula group, in which the phallic membrane evolved two rows of

spines, and progenitor H. Ancestral species H generated the Lebasiella group. Among

these species, the intraspicular plate was lost and the dorsal plate of the phallobase

became elongated. The monotypic Rifkindius group forms the complementary stock, in

which the tegminal lobes were lost and funicular antennomere 5 enlarged.

ACKNOWLEDGMENTS

My gratitude to John Dorshorst for his review of the manuscript and to Gregory

Zolnerowich (Kansas State University) for providing SEM time.

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