Research Article |
Corresponding author: Matthias Seidel ( seidelma@natur.cuni.cz ) Academic editor: Andrey Frolov
© 2017 Matthias Seidel, Mary L. Jameson, Rachel L. Stone.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Seidel M, Jameson ML, Stone RL (2017) A new cryptic species and review of the east-Andean leaf chafer genus Mesomerodon Ohaus, 1905 (Coleoptera, Scarabaeidae, Rutelinae). ZooKeys 671: 61-85. https://doi.org/10.3897/zookeys.671.11815
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The Neotropical scarab beetle genus Mesomerodon Ohaus (Scarabaeidae: Rutelinae: Rutelini) is distributed in the western (lowland) Amazonian region from Colombia to Bolivia. Based on our research, the genus includes three species including a new cryptic species from Ecuador. We use niche modeling to predict potential suitable habitat and identify environmental factors associated with the distribution of Mesomerodon species. We characterize the genus, provide a key to species, diagnose each species, describe a new species, provide spatial and temporal distributions, and discuss distributions of the species in relation to Amazonian landscape biodiversity.
El género neotropical de escarabajos Mesomerodon Ohaus (Scarabaeidae: Rutelinae: Rutelini) se distribuye en las zonas bajas del oeste de la Amazonía, de Colombia a Bolivia. Con base en nuestro estudio, este género se compone por tres especies, incluyendo una nueva especie críptica de Ecuador. Utilizamos el modelado de nicho para predecir el hábitat adecuado potencial e identificar los factores ambientales asociados con la distribución de las especies de Mesomerodon. Aquí caracterizamos al género, proporcionamos una clave dicotómica para las especies, diagnosticamos cada especie, describimos una nueva especie, proveemos información sobre la distribución espacial y temporal de las especies y discutimos la distribución de las especies en relación a la biodiversidad de paisajes de la Amazonía.
Amazonian lowland, pelidnotine chafer, sexual dimorphism, niche modeling
Selva baja amazónica, escarabajos pelidnotinos, dimorfismo sexual, modelado de nicho
The South American genus Mesomerodon Ohaus (Figs
Species in the genus Mesomerodon possess many external similarities in form, but the male genitalia are sufficiently different as to warrant species status for three, distinct operational taxonomic units that we treat as species.
We provide a synthesis of the biodiversity of the genus, including descriptions, key to species, diagnoses, and images. As a result of our research, the genus Mesomerodon includes three species, all of which are distributed in the western (lowand) Amazonia, including a new unexpected and cryptic species.
Characters. Morphological characters formed the basis of this work. The broadest range of potentially phylogenetically informative morphological characters was used for morphological analyses and comparisons. Morphological terminology is based primarily on
Characters and specimens were observed with 6–48× magnification and fiber-optic illumination. Digital images of specimens and structures were captured using the Leica Application Suite V3.8. Images were edited in Adobe Photoshop CS2 (background removed, contrast manipulated).
Species concept. Species are characterized by combinations of characters including the form the male protarsomeres and form of the male parameres in caudal and lateral views, and form of the ventral sclerite of the phallobase. Identification of female specimens required associated males from the same collecting event (place and date). We use the phylogenetic species concept (
Locality data. Locality data for all specimens examined as part of this study were translated into decimal latitude and longitude using GoogleEarth (http://www.google.com/earth/index.html) and provided (Suppl. material
Distribution modelling. To model the potential distribution and to identify environmental factors associated with Mesomerodon species, we used the maximum entropy algorithm (MaxEnt;
Type specimens. Friedrich Ohaus provided a legacy for understanding the biodiversity of Rutelinae with over 170 published papers and research collections (for biography see
Collections (Suppl. material
CCECL Musée des Confluences, Lyon, France (Cédric Audibert)
DBPC Denis Bouchard Personal Collection, Autouillet, France
DCCC David Carlson Personal Collection, Fair Oaks, California, USA
DJCC Daniel Curoe Personal Collection, Palo Alto, California, USA
JWPC Jim Wappes Personal Collection, San Antonio, Texas, USA
MLJC Mary Liz Jameson Personal Collection, Wichita, Kansas, USA
MSPC Matthias Seidel Personal Collection, Prague, Czech Republic
NMPC Department of Entomology, National Museum (Natural History), Prague, Czech Republic (Jiří Hájek)
SLCC Stephane LeTirant Collection, Montreal, Canada
UNSM University of Nebraska State Museum, Lincoln, Nebraska, USA (Brett Ratcliffe, M. J. Paulsen)
USNM U.S. National Museum, Washington, D.C. (currently housed at the University of Nebraska State Museum for off-site enhancement) (Floyd Shockley and Dave Furth)
Mesomerodon Ohaus, 1905: 319. Type species M. spinipenne Ohaus, 1905: 320–321 (by monotypy).
Length from apex of clypeus to apex of pygidium 17.0–20.0 mm (♂) and 19.0–24.0 mm (♀); width at mid-elytra 10.0–12.0 mm (♂) and 11.0–14.0 mm (♀). Color: Dorsal surface tan to ochre (cream or whitish when alive) with or without weak green reflections, ventral surface castaneous with weak metallic green or red reflections; specimens tend to darken with age. Form (Figs
Generic characters for Mesomerodon. 9 Left mandible of M. gilletti, dorsal view (broadly rounded externally with 2 interior, acute teeth; molar region broad) 10 Maxilla of M. gilletti, ventral view (with 6 teeth; galea not fused) 11 Labrum, dorsal view, of M. gilletti (apex emarginate medially) 12 Mentum, ventral view, of M. gilletti (subrectangular in shape, broadest at middle, apex emarginated) 13 Spiculum gastrale of M. gilletti 14 Wing of M. spinipenne showing venation and inset showing dense, thick setae associated with ScA and setose region anterior to RA3+415 Protarsomere 5 of M. barclayi sp. n., male, showing well defined, ventromedial emargination 16 Mesofemur of M. gilletti male, ventral view, showing acute process projecting posteriorly on posterior margin 17 Elytral apex of M. gilletti, lateral view, showing spiniform callus
Biology for species in the genus is not known. Adults likely feed on plant foliage, but no host has been recorded. Because adults are attracted to lights at night, it is likely that feeding occurs at night. Larvae are not described, but likely feed on compost and/or roots.
From the Greek, “mesos” meaning middle or in the middle, “mero” meaning femur, and “odon” meaning tooth. The name refers to the spinose process on the posterior margin of the mesofemur in males, a synapomorphy for species in the genus. The gender is neuter.
(Fig.
Within the Andean corridor, the genus level distribution model is congruent with the specimen-based distribution (Fig.
Species in the genus Mesomerodon are distinguished from other pelidnotine leaf chafers based on the acute, spiniform processes on the apical callus of the elytra in males (Fig.
Key to Mesomerodon species
1 | Mesofemur without process projecting posteriorly on posterior margin (♀) | 2 |
– | Mesofemur with acute process projecting posteriorly on posterior margin (Fig. |
3 |
2 | Distributed in Bolivia and Peru (Fig. |
M. spinipenne (♀) Ohaus |
– | Distributed in Ecuador and Colombia (Fig. |
M. gilletti (♀) Soula or M. barclayi (♀) Seidel, Jameson, & Stone, sp. n. |
3 | Protarsomere 2 ventrally with striate region at apex (Fig. |
M. spinipenne (♂) Ohaus |
– | Protarsomere 2 ventrally lacking striate region at apex (Figs |
4 |
4 | In lateral view, parameres curved posteriorly (Fig. |
M. gilletti (♂) Soula |
– | In lateral view, parameres sinuate (Fig. |
M. barclayi (♂) Seidel, Jameson, & Stone, sp. n. |
Clave para las especies de Mesomerodon
1 | Mesofémur sin proceso proyectándose posteriormente en el margen posterior | 2 |
– | Mesofémur con un proceso agudo proyectado posteriormente en el margen posterior (Fig. |
3 |
2 | Distribuido en Bolivia y Perú (Fig. |
M. spinipenne (♀) Ohaus |
– | Distribuido en Ecuador y Colombia (Fig. |
M. gilletti (♀) Soula o M. barclayi (♀) Seidel, Jameson, & Stone, sp. n. |
3 | Protarsómero 2 ventralmente con una zona estriada en el ápice (Fig. |
M. spinipenne (♂) Ohaus |
– | Protarsómero 2 ventralmente sin una zona estriada en el ápice (Fig. |
4 |
4 | En vista lateral, parámeros curvados posteriormente (Fig. |
M. gilletti (♂) Soula |
– | En vista lateral, parámeros sinuados (Fig. |
M. barclayi (♂) Seidel, Jameson, & Stone, sp. n. |
Holotype male and 19 paratypes (10 males, 9 females). Holotype male at
(based on 11 males and 9 females). The holotype does not differ significantly from the generic description and suffices for the description of this cryptic species (see “Remarks”). Descriptive details specific to the holotype specimen are indicated. Length from apex of clypeus to apex of pygidium 18–22 mm (♂) (holotype: 19 mm) and 22–25 mm (♀); width at mid-elytra 11–12 mm (♂) (holotype: 11 mm) and 12–15 mm (♀). Color: Cream colored (holotype), tan, or ochre; ventral surface castaneous with weak metallic green reflections. Form: Elytral apices with one short spine (holotype) or swelling (females). Legs: Protarsomere 2 of male ventrally lacking well-defined striae at ventral apex (Fig.
Males of Mesomerodon barclayi sp. n. are differentiated from other Mesomerodon species by the following combination of characters: Protarsomere 2 ventrally lacking a striated region at the ventral apex (striated in M. spinipenne; apical region lacking striae in M. gilletti [Figs
It is our honor to dedicate this species to Max Barclay (Curator, Coleoptera, Department of Entomology), who invited the first and second authors to the 1st Scarab Symposium at the
(Fig.
(Suppl. material
ECUADOR: Morona-Santiago, Orellana, Pastaza
Based on label data, this species is known to be active in the months of February, August, and November.
Based on label data, adult M. barclayi sp. n. is usually collected at light, thus suggesting activity and feeding at night. Individuals probably occur throughout the year. Immature stages are unknown. Specimens have been recorded at an elevation of 300 m.
As with other cryptic species, the overall body form of M. barclayi sp. n. is similar to other species in the genus. The form of the male genitalia (Fig.
Mesomerodon gilletti Soula, 2008: 21 (original combination)
Holotype male, allotype female, and eight paratypes (four male, four females. Holotype male at CCECL with label data: a) “Tena (E), 9/91, (750m)” (handwritten), b) male genitalia card mounted c) “Holotype, 2007, Mesomerodon gilletti S., Soula” (typeset and handwritten on red label). Allotype female at CCECL with label data: a) “Tena (E), 9/91, (750m)” (handwritten), b) “Allotype, 2007, Mesomerodon gilletti S., Soula” (typeset and handwritten on red label). Paratype male at CCECL with label data: a) “Tena (E), 9/91, (750m)” (handwritten), b) male genitalia card mounted, c) “Paratype, 2007, Mesomerodon gilletti S., Soula” (typeset and handwritten on red label). Paratypes (2 females) at CCECL with label data: a) “Tena (E), 9/91, (750m)” (handwritten), b) “Paratype, 2007, Mesomerodon gilletti S., Soula” (typeset and handwritten on red label). Paratypes (2 males) at CCECL with label data: a) “Misahuali (E.), 5/91” or “Misahuali (Eq.), 5/91” (handwritten), b) male genitalia card mounted, c) “Paratype, 2007, Mesomerodon gilletti S., Soula” (typeset and handwritten on red label). Paratype male at CCECL with label data: a) “EQUATEUR: Prov. NAPO, MISAHUALLI ile ANACONDA, Alt. 350 m.; 17–22.9.1990, Leg. Joss” (typeset), b) male genitalia card mounted, c) “Paratype, 2007, Mesomerodon gilletti S., Soula” (typeset and handwritten on red label). Paratypes (2 females) at CCECL with label data: a) “Misahuali (E.), 5/91” (handwritten), b) “Paratype, 2007, Mesomerodon gilletti S., Soula” (typeset and handwritten on red label).
(based on 50 males and 19 females). Length from apex of clypeus to apex of pygidium 18.0–20.0 mm (♂) and 21.0–24.0 mm (♀); width at mid-elytra 10.0–11.0 mm (♂) and 12.0–14.0 mm (♀). Legs: Protarsomere 2 of male lacking well-defined striae at ventral apex (Fig.
Mesomerodon gilletti males are differentiated from other Mesomerodon species by the following combination of characters: Protarsomere 2 of male ventrally with striated region poorly defined or lacking at apex (striated in M. spinipenne; lacking in M. barclayi sp. n.; [Figs
(Fig.
(Suppl. material
COLOMBIA: Putumayo
ECUADOR: Napo, Orellana, Sucumbíos
Based on label data, this species is known to be active in the months of February, May, June, August, September, October, and November.
Based on label data, adult M. gilletti are found associated with lights at night. Immature stages are unknown.
Mesomerodon spinipenne Ohaus, 1905: 320–321 (original combination).
Lectotype male (designated by
(based on 52 males and 161 females). Length from apex of clypeus to apex of pygidium 17.0–20.0 mm (♂) and 19.0–24.0 mm (♀); width at mid-elytra 10.0–12.0 mm (♂) and 11.0–14.0 mm (♀). Legs: Protarsomere 2 (♂) with well-defined striae at ventral apex (Fig.
Mesomerodon spinipenne males are differentiated from other Mesomerodon species by the following combination of characters: Protarsomere 2 with well-defined striae at ventral apex (Fig.
(Fig.
(Suppl. material
BOLIVIA: Cochabamba (Chapare), Huánuco (
PERU: Ayacucho (La Mar), Cusco (Quispicanchi), Huánuco (Leoncio Prado, Puerto Inca), Madre De Dios (Manú, Tambopata), Pasco (Oxapampa), Ucayali (Padre Abad), Santa Cruz (
Based on label data, this species is known to be active in all months except January and February.
Based on label data, adult M. spinipenne are active at night and can be collected at lights. Immature stages are unknown.
Based on body measurements provided by
It is possible that the specific epithet, “spinipenne”, refers to the apex of the elytra in the male which possess an apical spine or tubercle. The Latin root “spini” refers to spine or thorn, and the Latin root “penna” refers to wing. This character state is not unique to M. spinipenne; instead, it is a synapomorphy for all species in the genus.
The genus Mesomerodon is composed of three very similar species, two of which that have evaded discovery since the description of the genus by
Co-distributed cryptic species complexes may be a function of the western (lowland) Amazonian region with its aseasonal climate, humid forest, and heterogeneous vegetation. The distribution of two species of Mesomerodon in Ecuador coincides with highest global diversity of passerine birds and anurans (
Future studies associated with these cryptic species are needed to examine divergence, population structure, and sister group relationships of the genus. Molecular data will allow association of males and females for each species. Focused fieldwork could determine distributional limits and yield ecological data which will assist in understanding the origin and cause of sympatry.
We thank the curators, collections managers, and individuals listed in the “Materials and Methods” section for their loans of specimens and for their assistance. Max Barclay (
Distribution data for Mesomerodon species (Coleoptera, Scarabaeidae, Rutelinae)
Data type: occurrence data
Explanation note: Specimen level data used for niche model, phenology, and overall distribution.