Research Article |
Corresponding author: Niko Kasalo ( niko.kasalo5@gmail.com ) Academic editor: Oliver Hawlitschek
© 2024 Niko Kasalo, Martin Husemann, Thomas van de Kamp, Josip Skejo.
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:
Kasalo N, Husemann M, van de Kamp T, Skejo J (2024) Description of Phelene reinschmidti from Ecuador with notes on the subfamily Lophotettiginae (Orthoptera, Tetrigidae). Evolutionary Systematics 8(1): 119-125. https://doi.org/10.3897/evolsyst.8.124285
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Lophotettiginae Hancock, 1909, endemic to the Neotropics, is one of the most ill-known subfamilies of Tetrigidae. Until now, there have not been any hypothesis on its relationship with other subfamilies. Lophotettiginae comprise only two genera, Lophotettix Hancock, 1909 and Phelene Bolívar, 1906, both with an arduous taxonomic history. Here, we describe a new species, Phelene reinschmidti from Ecuador, and compare it with its congeners: P. turgida from Peru and P. maroon from French Guiana. We found the subfamily Lophotettiginae to most closely resemble the Neotropical subfamily Metrodorinae, meaning that the two are likely related. We also provide a 3D scan of the holotype, a first for Tetrigidae. The 3D model contains all the important taxonomic characters and much more data could be extracted in the future using artificial intelligence-assisted approaches.
3D scan, Neotropics, taxonomy, pygmy hoppers, grouse hopper
Tetrigidae
are a relatively specious orthopteran family with around 2000 described species (
Among the many problematic taxa, the subfamily Lophotettiginae Hancock, 1909 is perhaps the most elusive one. It comprises only two genera, Lophotettix Hancock, 1909 and Phelene Bolívar, 1906, the specimens of which are extremely rarely observed (
The genus Phelene was briefly synonymized under Chiriquia Morse, 1900 (
One important step to facilitate future taxonomic revisions and make them more independent of the geographic position of the researchers is enhancing digitalization efforts. The digitalization of type specimens is of special importance since it allows quick and easy access to the material without risking damaging specimens and without consuming the curators’ time. However, the vast majority of insect type specimens has not been photographed, and many of those that have been digitalized in this way only have dorsal views available, which severely limits their usefulness (
Here, we describe Phelene reinschmidti sp. nov., a new species of this understudied genus and provide a 3D scan of the holotype, a first among Tetrigidae. Furthermore, we review the status of the genus and provide the first hypothesis about the phylogenetic placement of Lophotettiginae.
The specimen was examined and photographed by Mathias Vielsäcker at SMNK using a Canon EOS R5 with either a Canon EF 100MM F/2.8L MACRO IS USM for the dorsal, ventral and lateral images, or a Laowa 25 mm f/2,8 Ultra Macro 2,5–5X for the images of smaller details. Stacked imaging was performed using a WeMacro rail. Images were then stacked using Helicon Focus.
Measurements were made using ImageJ 1.53t software (
The specimen was scanned in six height steps using a filtered polychromatic beam at the high-throughput tomography station at the Imaging Cluster of KIT Light Source. We employed an air-bearing rotary stage (RT150S, LAB Motion Systems) for sample rotation and a fast indirect detector system consisting of a 200 µm LuAG scintillator, a double objective white beam microscope (Optique Peter;
The individual tomographic volumes were registered and merged in Amira 2022.2, which was also employed for pre-segmentation. Pre-segmented labels served as input for semi-automatic segmentation with Biomedisa (
Frontal costa bifurcation in upper third of eye height. Antennal groove at bottom margin of eyes. Antennomeres flattened, apical one white-tipped. Vertex wider than an eye. First segment of anterior and middle tarsi a little elongated. Hind tarsus very long, first and third segment approximately equally long. Elongated slim ovipositor.
Lophotettix Hancock, 1909, type species Lophotettix brevicristatus Hancock, 1909.
Two genera, Lophotettix, Phelene, endemic to Southern and Central America.
Nephele Bolívar, 1887: 252. (junior homonym of Nephele Hübner, 1819);
Phelene
Bolívar, 1906: 392. (replacement name);
Gladiotettix
Hancock, 1907: 40. (diagnosis in English, obsolete replacement name);
Antennal segments flattened, but oval (apical segments triangular in Lophotettix). Low pronotal crest (medium to tall in Lophotettix). Anterior and middle femora smooth (moderately tuberculated in Lophotettix).
Phelene turgida (Bolívar, 1887)
P. maroon Cadena-Castañeda & Tumbrinck, 2021, P. turgida (Bolívar, 1887), P. reinschmidti sp. nov. The genus seems to be restricted to the northern half of South America.
We identified two female specimens of P. turgida from Peru at the Zoologische Staatssammlung München (ZSM), which we present in Fig.
Easily separated from the congeners by the following set of characters: (i) median carina forming a low elongated crest which extends from between the prozonal carinae to the base of hind legs; (ii) vertex a little less than two eyes wide; (iii) brown body without distinct patterns; (iv) rounded pulvilli of hind tarsi; (v) female subgenital plate oval with moderately protruding triangular apex; (vi) alae dark brown with venation of the same color. Differs from P. maroon by characters iii, iv, v, and vi.
Macropronotal. In anterior view, top of vertex approximately at level of upper margin of eyes. Vertex a little less than two eyes wide. Frontal costa bifurcation in upper third of eye height. Facial carinae forming long oval shape. Paired ocelli approximately at half of eye height. Midline of antennal groove at level of bottom margin of eyes. Antennae composed of 13 visible segments; apical six flattened, last segment white. In dorsal view, vertex of isosceles trapezoidal shape. Medial carina visible in anterior half of vertex. Lateral carinae in form of small tubercles. Prozonal carinae as long as eye as seen in dorsal view, strongly convergent caudad. Median carina visible throughout length of pronotum, forming low but long crest in lateral view (extending from between prozonal carinae to base of hind legs). Pronotal surface covered in small tubercles and carinulae. Humeral angles wide and blunt. Lateral lobes rectangular, projected laterally, with thin white border. Tegmina large and oval. Alae reaching pronotal apex; dark brown with dark brown venation. Anterior and middle femora long and smooth. Anterior and middle femora long and smooth. Anterior and middle tarsi with long first segment and extremely long second segment. Hind femur robust, with two tubercles on dorsal margin; moderately sized sharp antegenicular tooth, genicular tooth barely visible. Hind tibia thin, serrated along caudal margin. Hind tarsus long; first segment extremely long, with three elongated pulvilli; third segment missing in the holotype. Ovipositor extremely elongated, serrated. Base of subgenital plate with triangular notch; shallow stitch line extending from it throughout most of the length of the plate. Subgenital plate elongated with moderately protruding triangular apex.
Measurements (in mm). BL (9.21); PL (12.33); PWS (2.19); PWM (2.85); PHM (2.38); VW (0.84); CEW (0.52); AGW (0.23); SW (0.3); FFL (2.16); FFW (0.45); MFL (2.28); MFW (0.49); HFL (4.75); HFW (1.58); TL (1.47); TW (0.58)
Adult female collected in early 1993 by Duffner in Ecuador, Sucumbíos Province on the upper Aguarico. The holotype is deposited at the SMNK (Karlsruhe, Germany). See Fig.
“ECUADOR (Sucumbios) / Oberer Aguarico, Anfang / 1993 leg. Duffner // 101, // Staatl. Museum / für Naturkunde / Karlsruhe; / ex coll. Riede, / Inv. Nr. E-Orthop-2“
Ecuador, Sucumbíos Province. No precise locality known.
Known only from a single specimen.
The new species is named after Prof. Dr. Matthias Reinschmidt, the current director of the Karlsruhe Zoo for his dedication and investments in nature protection in Ecuador and around the world.
Phelene reinschmidti sp. nov. A. Regions where Phelene species have been found, red shapes represent type localities, star–P. reinschmidti, circle–P. turgida, triangle–P. maroon; B. Frontal view; C. Dorsal view; D. Ventral view; E. Abdominal apex in ventral view; F. Lateral view; G. Image of the 3D model we generated.
1 | Tall narrow pronotal crest above tegmina | Phelene turgida |
– | Low elongated pronotal crest | 2 |
2 | Lower half of body distinctly black, upper half black or brown; pulvilli of hind tarsus triangular and sharp; alae dark with bright venation; female subgenital plate oval with slight tubercle at its apex | Phelene maroon |
– | Body uniformly brown; pulvilli of hind tarsus rounded and blunt; alae and their venation dark brown; female subgenital plate oval with moderately protruding triangular apex | Phelene reinschmidti sp. nov. |
The new species is at first glance quite similar to Phelene maroon. The latter was until recently known only from a single male specimen (
Until now, the question of the position of Lophotettiginae in the Tetrigidae phylogeny remained completely unaddressed. The brief synonymization of Phelene with Chiriquia (
Further, we have to consider the genus Metrodora Bolívar, 1887. Some of its species, e.g., M. lutosa Bolívar, 1887, M. gibbinota (Bruner, 1910), and M. panamae (Hebard, 1924) also exhibit the earlier listed similarities with Phelene, while M. gibbinota and M. panamae also have white-tipped antennae. This genus and by extension its tribe, Metrodorini Bolívar, 1887, is composed of likely related but ill-defined taxa, which is hindering further research (
Lastly, all these genera share the mentioned characters with many members of the predominantly African tribe Xerophyllini Günther, 1979, currently attributed to the subfamily Cladonotinae. Therefore, the possibility that Lophotettiginae, Metrodorinae, and Xerophyllini form a monophyletic group should be seriously considered and examined from both morphological and molecular angles. The morphology akin to Morphopus (Bolívar, 1887) (Xerophyllini) could be basal in this group, which could explain all the different derived morphologies within it, as well as its distribution across the southern half of Gondwana. It should be noted that we do not consider Asian Metrodorinae as true members of that subfamily and that all of the mentioned higher taxa require extensive revisions.
We here provide the first 3D data for any Tetrigidae, though more are in preparation. Such 3D data may become an important resource in future taxonomic works, especially in concert with artificial intelligence and machine learning approaches. One of the large advantages of including 3D data in taxonomic descriptions is that virtual specimens will be readily available around the world, without the danger of sending valuable and irreplaceable holotypes and risking them getting lost or damaged. In addition, such virtual specimens are at least a minor backup in case a type gets lost. The 3D models are scaled to size and allow easy and accurate measurement of distances and also have the benefit of allowing for 3D geometric morphometric measurements. Modern high-throughput X-ray imaging methods already offer the possibility to digitize large numbers of insects in a short time (
NK: first and last draft, taxonomy, measurements, figures; MH: last draft, specimen handling; TvdK: last draft, 3D digitalization; JS: last draft, taxonomy.
We thank Marcus Zuber and Elias Hamann for their assistance with the tomographic scans and Tomás Faragó for tomographic reconstruction. We thank Mathias Vielsäcker for imaging the specimens. We acknowledge the KIT Light Source for provision of instruments at their beamlines and thank the Institute for Beam Physics and Technology (IBPT) for the operation of the Karlsruhe Research Accelerator (KARA). We are grateful to Tomislav Domazet-Lošo for reading an early version of this manuscript and giving valuable suggestions. We acknowledge the help of Leibniz Institut zur Analyse des Biodiversitätswandels in publishing this article.
3D reconstruction of Phelene reinschmidti sp. nov.
Data type: pdf
Explanation note: 3D reconstruction of Phelene reinschmidti sp. nov. Click on the image to start the interactive 3D view; use the menu to switch between the standard views.