a new species of archedinus (coleoptera: scarabaeoidea: trichiinae) from oaxaca, mexico

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A new species of Archedinus (Coleoptera: Scarabaeoidea:Trichiinae) from Oaxaca, MexicoAuthor(s): Miguel Angel Morón and Fernando Z. Vaz-de-MelloSource: Pan-Pacific Entomologist, 83(2):110-119. 2007.Published By: Pacific Coast Entomological SocietyDOI: http://dx.doi.org/10.3956/0031-0603-83.2.110URL: http://www.bioone.org/doi/full/10.3956/0031-0603-83.2.110

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A new species of Archedinus (Coleoptera: Scarabaeoidea: Trichiinae)from Oaxaca, Mexico

MIGUEL ANGEL MORON* AND FERNANDO Z. VAZ-DE-MELLO

Instituto de Ecologıa, A.C. Apdo. Postal 63, Xalapa, Veracruz 91000, Mexico

e-mail: [email protected], [email protected]

Abstract. Archedinus howdeni n.sp. from humid forests in the Chimalapas mountains, Oaxaca,Mexico, is described, illustrated and compared with A. relictus Moron and Krikken from thecloud forests of El Triunfo Biosphere Reserve, Chiapas, Mexico. A key is included to thespecies of Incaini (Trichiinae) found in the Central American Nucleus; as well as an analysis ofthe phylogenetic relationships among incaine genera.

Key Words. Insecta, Incaini, Archedinus, new species, Mexico.

Resumen. Se describe e ilustra Archedinus howdeni n.sp., colectada en los bosques humedos delas montanas Chimalapas, Oaxaca, sureste de Mexico. Se destacan las diferencias con A.relictus Moron y Krikken que habita en los bosques nebulares de las montanas de El Triunfo,Chiapas, Mexico. Se incluye una clave para identificar las especies de Incaini (Trichiinae)presentes en el Nucleo Centroamericano; y se presenta una primera prospeccion de relacionesfilogeneticas entre los generos de Incaini.

INTRODUCTION

Following Krikken (1984), Moron and Krikken (1990) united the genera Inca

LePeletier & Serville, 1828, Pantodinus Burmeister, 1847, Golinca Thomson, 1878

and Archedinus Moron & Krikken, 1990 in the tribe Incaini Schoch, 1896. Ricchiardi

(2002) included the genus Coelocratus Burmeister, 1842 in the same tribe and

presented a key to all incaine genera. According to Krajcik (1999), Inca clathrata

sommeri Westwood, Pantodinus klugi Burmeister and Archedinus relictus Moron &

Krikken are distributed in Mesoamerica (total three species excluding the one

described here). Inca c. clathrata (Olivier), I. c. sommeri, I.c.quesneli Boss and

Ratcliffe, I. bonplandi Gyllenhall, I. besckii Schaum, I. burmeisteri Burmeister, I.

irrorata Chevrolat, I. pulverulenta (Olivier), Golinca bifrons (Olivier), G. davisi

Waterhouse, G. ishiharai Nagai and Coelocratus rufipennis (Gory and Percheron),

are distributed in South America (total twelve species).

Colleagues collected four male specimens of an undescribed species of the genus

Archedinus in the Chimalapas mountains in Oaxaca during the course of vertebratefield studies These specimens came as a nice surprise because heretofore the genus

included only a single relict species, A. relictus, from the El Triunfo mountains in

Chiapas. The aim of this paper is to describe this new species of Archedinus and to

consider its phylogenetic relations with the other Central American Incaini.

The terms used in this description are those of Howden (1968), Krikken (1984)

and Moron & Krikken (1990). Figures are images obtained with Automontage

system and Jeol scanning electron microscope; measurements were made with an

ocular micrometer or electronic caliper.

The type series is deposited in the following collections: Henry and Anne Howden

Collection/Canadian Museum of Nature, Ottawa (HAHC/CMNC); Miguel A.

* Corresponding author.

THE PAN-PACIFIC ENTOMOLOGIST83(2):110–119, (2007)

Moron collection/Instituto de Ecologıa, Xalapa, Mexico (MXAL/IEXA); Fernando

Z. Vaz-de-Mello collection, Lavras, Brazil (FVMC).

The phylogenetic analysis was made using NoNa (Goloboff 1993), with Fitch

parsimony and equally weighted characters; and a total of 23 male morphological

characters (females of Coelocratus and A. howdeni are unknown) (Table 1.). Selected

outgroups are Osmoderma eremicola and Trichius fasciatus (the last one considered

as most external for tree rooting); the ingroup comprises both Archedinus species and

the type species of other genera, except for Inca, which is represented by its type and

one other species. Clade support was inferred by the Bremer index (decay index) up

to ten extra steps.

Archedinus howdeni Moron & Vaz-de-Mello, sp. nov.

(Figs. 1, 3, 5)

Holotype. Male. Total length 25.0 mm, humeral width 10.5 mm. Shiny black,

dorsally glabrous, surface coarsely and irregularly punctate. Clypeal margin

anteriorly upturned with sinuate border, dorsal surface with irregular deep punctures

(Fig. 1), anterior surface wide, sparsely punctate and deeply excavated towards

ventral border. Clypeal horn absent. Maximum width of head capsule 4.5 mm;

Table 1. Characters used for phylogenetic analysis of relationships between genera of Incaini.

Number Description/state 0 1 2

0 Antero-ventral clypeal border Nearly straight Widely sinuate Deeply sinuate1 Form of clypeal projections None Central horn or

proccessPareate horns

2 Frontal surface Concave Convex3 Transverse frontal carina Absent Present4 Supraocular ridges Absent Scarce Strong5 Antennal club Shorter than steam Longer than

steam6 Disk of mentum Concave Flat7 Pronotal disk With depressions Convex With process8 Anterior medial pronotal border Not pointed Pointed9 Pronotal setae Absent Present10 Pronotal color Not metallic Metallic11 Anterior angles of pronotum Rounded Projected Angled but not

projected12 Form of probasisternal process Acute Rounded13 Length of probasisternal

processShorter than

procoxalanterior face

Longer thanprocoxalanterior face

14 Pterosternal setae Scarce Abundant15 Elytra surface Not velvety Velvety16 Elytral striae Well marked None Weakly

indicated17 Elytral surface Shiny Dull18 Apex of profemur With blade With claw19 Base of protibia Straight Sinuate Toothed20 Fore tibia apico-internal tooth Absent Present21 Length of parameres Shorter than basal

pieceAs long as basal

piece22 Apex of parameres Entire Bifurcate

2007 MORON & VAZ-DE-MELLO: NEW ARCHEDINUS FROM OAXACA 111

Figures 1–2. Dorsal view of head and pronotum of Archedinus spp.: 1. A. howdeni n. sp.; 2. A.relictus. Scale lines 5 1 cm.

112 THE PAN-PACIFIC ENTOMOLOGIST Vol. 83(2)

maximum width of clypeus 2.2 mm. Distal segment of maxilary palpus fusiform,

enlarged, 0.9 mm length, and third segment of labial palpus subcylindrical, rounded,

not connate with the second one, enlarged, 0.7 mm length. Antennae 10 segmented,

antennal club enlarged (Fig. 1), 1.4 times longer than preceding six segments

combined; segment 7 wider than long, with narrowed, short anterior projection;

segment 6 longer than wide, with apex obliquely truncated, without anterior

projection; combined segments 4 and 5 nearly as long as the segment 3, segment 3

longer than wide, pedicel nearly globular, as wide as long; scape progressively

enlarged toward distal border.

Figures 3–4. Lateral view of parameres of Archedinus spp.: 3. A. howdeni n.sp.; 4. A. relictus.


Pronotum about 1.3 times wider than long; dorsal surface with wide, deeply and

irregularly punctate groove on midline and two shallow depressions on each side

(Fig. 1). Proepisternum with deep, rounded, large punctures. Probasisternal

projection long, with rounded apex, and many reddish setae. Elytra (viewed from

above) with borders uniformly curved, with 10 wide, deep striae, each with irregular

row of rounded punctures and indistinctly covered with waxy secretion; striae I–V

extend from anterior border to apical callus; striae VI–IX extend from humeral

callus to apical callus; stria X extends along lateral margin, but does not reach apex;

interstriae shiny with sparse microscopic punctures; preapical area shiny, irregularly

punctate; elytral apex rounded; epipleura narrowing posteriorly. Mesosternum

depressed on midline, abruptly upturned near anterior borders of mesocoxae. Wings

well developed. Metasternum convex at sides, narrow and shallowly grooved on

midline; lateral surface with rounded punctures and many slender, erect setae.

Profemur with preapical rounded, blade-like dorsal projection. Protibia with wide

basal notch on inner side. All tarsi robust, with large sickle-shaped claws.

Abdominal sternites convex, glabrous, finely punctate medially and pitted toward

sides; last sternite with posterior border slightly raised and briefly sinuate at middle.

Pygidium glabrous, shiny, with dense fine micropunctures mixed with sparse, large

round punctures, finely rugose toward basal angles; without waxy spots; apical

margin nearly straight.

Figures 5–6. Distal view of parameres of Archedinus spp.: 5. A. howdeni n.sp.; 6. A. relictus.


Aedeagus with large basal piece, slightly convex; tectum shallowly concave;

parameres elongate, sinuose, with apex entire, narrowed (Figs. 3, 5); inner sac

without sclerotized structures.

Female. Unknown

Variation. Paratypes are similar to the holotype except as follows: total body

length: 25–27.0 mm; humeral width: 10–11.7 mm. Pronotal posterior depressions

can be reduced or, in one case, absent; pygidial sculpture is fossulate on proximal

half and fossulate-rugose on basal angles in one paratype.

Specimens examined. 4 males. Holotype, male; MEXICO. OAXACA: Municıpio

de San Miguel Chimalapa, near Benito Juarez, 1600 m, 27-VI-2005, A. Abundis and

R. Dıaz collectors; deposited MXAL/IEXA. Paratypes (3): same data as holotype,

deposited HAHC/CMNC, MXAL/IEXA, FVMC.

Etymology. This species is named in honor of Dr. Henry F. Howden, in

recognition of his important contributions to the study of American Trichiinae.

Remarks. Archedinus howdeni is similar to A. relictus (Figs. 2, 4, 6) in body shape,

pronotal and elytral sculpture, proportions of antennae, and form of legs; but

externally it differs from A. relictus by having the anterior clypeal border upturned;

a shallow fronto-clypeal cavity medially divided by a fine keel; two pronotal

depressions on each side; proepisterna with scattered rounded, deep, large punctures;

probasisternal projection with rounded apex; profemur with rounded preapical

projection; metasternum shallowly furrowed on the midline; elytral striae without

waxy secretion; and rounded elytral apex. In contrast, A. relictus has the clypeus with

a bifurcate horn-like projection; deep undivided fronto-clypeal cavity; one pronotal

depression on each side (Fig. 2); proepisterna fossulate (see note above);

probasisternal projection with acute apex; profemur with claw-shaped preapical

projection; metasternum deeply furrowed on the midline; elytral striae with gray

waxy secretion; and tooth-like elytral apex. In addition, A. relictus has parameres

with bifurcate apices (Figs. 4, 6), while A. howdeni has parameres with acute, entire

apices.

The distributions of known Mesoamerican Incaini show two different patterns:

one is clearly South-American in origin (represented by Inca), while the other seem

to be originally Mesoamerican (Pantodinus and Archedinus).

Archedinus relictus is known only from the cloud forests from the biosphere

reserve ‘‘El Triunfo’’, Municipio de Angel Albino Corzo, Chiapas, Mexico, and

neighboring shaded coffee plantations between 1000–2000 m altitude (Fig. 7).

Archedinus howdeni was collected in tropical subdeciduous forest at 1600 m altitude

in the Chimalapas mountains near the Oaxaca-Chiapas border (Fig. 7). Pantodinus

klugi is a common species in some mountains of the departments of Solola,

Guatemala, Chimaltenango, San Marcos and Quetzaltenango, Guatemala, at 1800–

2300 m altitude; but apparently it is a rare species in the state of Chiapas, Mexico,

where it has only been found in Boqueron and Tacana volcanoes (Fig. 7) between

1800–2400 m altitude. Those two genera appear to form a well-defined group

(Moron and Krikken 1990) in the Incaini and do not include non-Mesoamerican

representatives. The present distributions of Archedinus and Pantodinus suggest an

ancient origin with post- Pleistocenic restriction to the higher mountains of Central

American Nucleus.

In contrast, Inca clathrata sommeri is found sporadically along the eastern or

northern lower slopes of the mountains and some inner tropical canyons of


Tamaulipas, San Luis Potosı, Hidalgo, Puebla, Veracruz, Oaxaca and Chiapas,

between 100–1300 m altitude (Moron 1983, Moron et al. 1997), without records

from the Yucatan Peninsula or Peten area, and some records in Belize (Howden1968), Alta Verapaz and Izabal, Guatemala (E. Cano personal communication) and

Olancho and Yoro, Honduras (R. Cave personal communication), between 1200–

1450 m altitude (Fig. 7). All other species of Inca are exclusively South American.

The present distribution of I. c. sommeri suggests an origin in northern South

America with Pleistocenic extension into the subtropical and tropical montane

humid forests of Mesoamerica.

Larvae of Inca c. sommeri have been found inside rotten logs and stumps of Pinus

oocarpa Schiede. (Pinaceae) and Brosimum sp (?) (Moraceae) (Moron 1983), whilethe larvae of Archedinus relictus were collected under rotten log of Guarea sp.

(Meliaceae) (Moron 1995). Adults of A. howdeni were obtained under bark of rotten

logs of undetermined species.

KEY TO THE SPECIES OF INCAINI OF MESOAMERICA

1 Elytra deeply striate. Dorsal color dark brown, shiny. Male with or withoutsingle clypealhorn . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

19 Elytra not striate. Pronotum velvety black, with 3–5 waxy yellow stripes.

Elytra dull, reddish black or purple with small yellow spots irregularly

Figure 7. Distribution of the species of Incaini in Mesoamerica. Archedinus relictus (a),Pantodinus klugi (c), Inca clathrata sommeri (d), Archedinus howdeni (g).


distributed. Male with two divergent clypeal horns covered with dense

yellow hairs on inner sides. . . . . . . . . . . . . Inca clathrata sommeri Westwood

2 Pronotum with distinct basal margin, with deep and sparse punctures. Male

pronotum with two latero-dorsal rounded prominences anteriorly excavated

and with setae; female pronotum widely convex. Male clypeal horn

elongated, rounded apically, curved upwards, with preapical short process

on dorsal border. . . . . . . . . . . . . . . . . . . . . . . . .Pantodinus klugi Burmeister

29 Pronotum without distinct basal margin, very coarsely punctate foveate,

without projections in male and female. Male clypeus with anterior border

elevated or weakly horned (Figs. 1, 2) . . . . . Archedinus Moron & Krikken, 3

3 Apex of probasisternal process acute. Profemur with claw-like preapical

projection. Striae with gray waxy secretion. Male clypeus with short

bifurcate horn (Fig. 2). Apex of parameres bifurcate (Figs. 4, 6). . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .A. relictus Moron & Krikken

39 Apex of probasisternal process rounded. Profemur with rounded preapical

projection. Striae without waxy secretion. Male clypeus with increased

anterior border briefly sinuate (Fig. 1). Apex of parameres acute (Figs. 3,

5) . . . . . . . . . . . . . . . . . . . . . . .A. howdeni Moron & Vaz-de-Mello, sp. nov.

PHYLOGENY OF INCAINI

A character matrix is presented in Table 2. Two characters (0 and 6) are codified

as unknown for Coelocratus, because they can not be discerned from available

literature and photographs (specimens were unavailable for this study).

Analysis was performed with exhaustive search (mswap+command), and only one

tree was found (Length 5 48, CI 5 68; RI 5 58; Fig. 8.). Characters supporting the

monophyly of the Incaini are antennal club long, base of protibia sinuate or toothed,

and parameres long. Incaini are clearly divided in two clades, one represented by

Pantodinus+Archedinus (Bremer 5 1), supported by deep elytral striae (unambigu-

ous) and antero-ventral clypeal border sinuate (homoplasic, shared with Inca

clathrata); and the other represented by Golica+Coelocratus+Inca (Bremer 5 1),

supported by the apex of profemur internally with a claw (homoplasic, shared with

Archedinus relictus).

The monophyly of Archedinus (Bremer 5 3) is supported by strong supraocular

ridges, long probasisternal process (both unambiguous), and scarce pterosternal

setae (homoplasic, shared with Coelocratus).

Table 2. Data matrix used for phylogenetic analysis of relationships among genera of Incaini.

0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22

Trichius fasciatus 0 0 1 0 0 0 1 1 0 1 0 0 1 0 1 1 2 1 0 0 0 0 0Osmoderma eremicola 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0Coelocratus rufipennis ? 0 1 1 0 1 ? 1 0 0 0 2 0 0 0 0 1 1 1 1 0 1 0Golinca bifrons 0 2 0 0 1 1 0 0 1 0 0 1 1 0 1 0 1 1 1 1 0 1 0Inca clathrata 1 2 1 1 0 1 0 1 0 0 1 1 1 0 1 1 1 1 1 2 1 1 1Inca bonplandi 0 2 1 1 0 1 0 0 0 0 0 1 1 0 1 1 1 1 1 1 1 1 1Pantodinus klugi 1 1 1 0 1 1 1 2 0 0 0 1 1 0 1 0 0 0 0 1 0 1 0Archedinus relictus 2 1 0 0 2 1 0 0 0 0 0 0 0 1 0 0 0 0 1 2 0 1 1Archedinus howdeni 1 1 0 0 2 1 0 0 0 0 0 0 1 1 0 0 0 0 0 1 0 1 0


In the Golica+Coelocratus+Inca clade, a group formed by Coleocratus+Inca

(Bremer 5 1) is supported by the presence of a frontal transverse carina

(unambiguous) and by the absence of supraocular ridges (comparted with Trichius).

Monophyly of Inca (Bremer 5 2) is supported by velvety elytral surface (comparted

with Trichius), presence of an apico-internal tooth in the foretibia (unambiguous),

and bifurcate parameres (homoplasic, comparted with Archedinus relictus).

The clade formed by Archedinus+Pantodinus is exclusively Mesoamerican; while

the Golica+Coelocratus+Inca clade is primarily South American, with only one

species of Inca, and maybe some of Golinca, occurrying in Central America.

ACKNOWLEDGMENTS

We thank Alejandro Abundis Santamarıa and Roman Dıaz (both at Instituto de

Ecologıa, A.C., Xalapa, Mexico), collectors of the new species, with the hope that its

description will encourage future collecting and maybe show them that scarabs are

much more fun taxonomically than vertebrates. Thanks also to Robert E. Woodruff

and Paul Skelley (Florida State Collection of Arthropods, Division of Plant

Industry, Gainesville, Florida) for the automontage and scanning electron

microscope images included in this paper; to Daniel Curoe (Mexico City) for the

donation of the Golinca specimen used for the phylogenetic analysis; to Enio Cano

(Universidad del Valle, Guatemala) and Ronald D. Cave (University of Florida, Ft.

Pierce) for the data of the Inca specimens from their collections; and to Dr. Gonzalo

Halffter (Instituto de Ecologıa) for general logistical support to FZVM. This paper is

a result of the projects ‘‘Coleoptera Lamellicornia de America Latina’’ supported by

Figure 8. Phylogeny of Incaini. Characters shown in black are non-homoplasic, those shown inwhite are homoplasic. Characters numbers are above character symbol. Length 5 48, CI 5 68; RI5 58.


Instituto de Ecologıa (account 902-08-011) and ‘‘Analisis de las relaciones entre

diversidad alfa, beta y gamma a distintos niveles de escala espacial. V. etapa’’supported by CONABIO (EE005/06). FZVM receives a Capes grant (Ministerio da

Educacao, Brazil, BEX 120802-0).

LITERATURE CITED

Goloboff, P. 1993. NoNa ver. 2.0. Software and documentation.Howden, H. F. 1968. A review of the Trichiinae of North and Central America (Coleoptera:

Scarabaeidae). Memoirs of the Entomological Society of Canada 54:1–77.Krajcik, M. 1999. Cetoniidae of the World. Catalogue-Part II (Coleoptera: Cetoniidae). Typos

Studio Most, Most, Czech Republic, 72 pp + XXIII pp.Krikken, J. 1984. A new key to the suprageneric taxa in the beetle family Cetoniidae, with

annotated lists of the known genera. Zoologische Verhandelingen. Leiden, 210:1–75.Moron, M. A. 1983. Los estados inmaduros de Inca clathrata sommeri Westwood (Coleoptera:

Melolonthidae: Trichiinae); con observaciones sobre el crecimiento alometrico del imago.Folia Entomologica Mexicana 56:31–51.

Moron, M. A. 1995. Larva and pupa of Archedinus relictus Moron & Krikken (Coleoptera:Melolonthidae; Trichiinae, Incaini). The Pan-Pacific Entomologist 71(4):237–244.

Moron, M. A. & J. Krikken. 1990. A new Mesoamerican genus of Trichiinae (Coleoptera:Scarabaeoidea). Folia Entomologica Mexicana 78:71–84.

Moron, M. A., B. C. Ratcliffe & C. Deloya. 1997. Atlas de los escarabajos de Mexico. ColeopteraLamellicornia. Vol. I Familia Melolonthidae. CONABIO y Sociedad Mexicana deEntomologıa, A.C. Mexico, 280 pp.

Ricchiardi, E. 2002. Notes on the genus Coelocratus Burmeister, 1842 (Coleoptera: Scarabaeidae).Cetoniimania 2(1):3–7.

Received 26 Sep 2006; Accepted 6 Dec 2006. Publication date 12 June 2007.


a new species of archedinus (coleoptera: scarabaeoidea: trichiinae) from oaxaca, mexico

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