Research Article |
Corresponding author: Edmundo González-Santillán ( edmundo.gonzalez@ib.unam.mx ) Academic editor: José Antonio Ochoa
© 2015 Edmundo González-Santillán, Fernando Alvarez-Padilla.
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:
González-Santillán E, Alvarez-Padilla F (2015) The male of Megacormus granosus (Gervais, 1844) with comments on its hemispermatophore (Scorpiones, Euscorpiidae). ZooKeys 504: 75-91. https://doi.org/10.3897/zookeys.504.9027
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The male of Megacormus granosus is described for the first time and the female redescribed. A homology scheme proposed recently is applied to hemispermatophore structures. The specimens were collected in an oak forest from Pico de Orizaba Volcano at an average altitude of 2340 m. All adult males were collected by pitfall traps, whereas all adult females and both sex immatures were collected using Berlese funnels, suggesting that males are comparatively more mobile within the leaf litter layer, probably due to mating season.
Euscorpiidae, Orizaba, Veracruz, Mexico
The Mexican genus Megacormus (Karsh, 1881) is comprised of four species, M. granosus (Gervais, 1844); M. segmentatus Pocock, 1900; M. gertschi Díaz Nájera, 1966 and M. grubbsi Sissom, 1994. All species of Megacormus are restricted to the slopes of the Sierra Madre Oriental and the costal lowlands of the Gulf of Mexico, ranging in altitude from 300 m to over 2300 m. They prefer habitats with high relative humidity such as oak, oak-pine and evergreen tropical forest, and are found in these communities within the leaf litter, outcrop crevices, decaying logs or under rocks. Their distribution includes the states of Hidalgo, Oaxaca, Puebla, Querétaro, San Luis Potosí, Tamaulipas, and Veracruz (
Megacormus granosus was originally described as a species of Scorpio Linnaeus, 1758 from a single female (
The hemispermatophore constitutes one of the most informative character systems in suprageneric phylogenetic studies, at least in Bothriuridae Simon, 1880 and Vaejovidae Thorell, 1876 (
The unsatisfactory working terminology and concomitant absence of homologies in the hemispermatophore of the Order Scorpiones have promoted a plethora of nomenclatures, hindering the correct interpretation of homologies (
All scorpions were collected by sifting leaf litter processed with Berlese funnels and pitfall traps. Specimens are deposited in the Laboratorio de Aracnología de la Facultad de Ciencias, Universidad Nacional Autónoma de México (UNAM), Mexico. Measurements in mm were taken with an ocular micrometer. Illustrations were produced with a Nikon SMZ 1000 dissecting stereomicroscope and a Nikon Eclipse E200 with a camera lucida. Photomicrographs were taken with a digital camera Nikon DS-U3 under LED illumination and Ultraviolet (UV) light using a 20W Techno Lite® bulb mounted in a desktop stand lamp. Hemispermatophores were digested with pancreatine (
Nomenclature follows
Specimens were collected in a 15–20-year old oak forest near the boundary of the Pico de Orizaba National Park ca. two kilometers southwest from Atotonilco de Calcahualco, Veracruz. Two plots, of one hectare each, were established with the following central coordinates. Plot I 19°8'17.4"N, 97°12'16.2"W, altitude 2,300 m. Plot II 19°8'30.2"N, 97°12'21.5"W, altitude 2388 m. Three expeditions were conducted, two in May 21–30 th and October 4–14th 2012 and the third in February 15–24th 2013.
Scorpio granosus
Chactas granosus: Karsch 1879: 111.
Megacormus granosus:
MEXICO: Holotype female, depository unknown (
M. granosus is most similar to M. segmentatus by sharing pedipalp chela fingers margin straight lacking proximal notch and median lobe; nineteen trichobothria on patella retrolateral and seven on ventral surface. It can be separated from M. segmentatus by having entirely densely granular surfaces on the femur, patella, tibia dorsal surfaces, prolateral leg surfaces, carapace, and tergites (Figures
The following redescription supplements
Color and infuscation: Base color yellowish to orange. Carapace: tergites, prolateral surface of legs, sternum, genital operculum, pectinal basal piece, fused lamella, metasoma, and telson, with dense, marbled infuscation (Figures
Chelicerae: Manus dorsal surface smooth, lustrous, with three macrosetae distally, decreasing in size from median to lateral surface. Movable finger, retrolateral margin with subdistal and medial denticles triangular, subequal; distal and basal denticle slightly larger; prolateral margin with three smaller, triangular, subequal denticles, situated in distal half of the finger; retrolateral distal finger size half of prolateral distal finger. Fixed finger margin with three denticles, proximal two adjacent and distal separate; distal denticle elongate and sharp (Figure
Carapace: Length equal to 0.9 times the posterior width. Surface shagreened, with enlarge scattered granules covering entire surfaces (Figure
Coxosternal region: Sternum pentagonal, subequilateral, length equal to 0.8 times the width, with five to ten pairs of microsetae. Median sulcus of sternum with anterior and posterior margins broadened, moderately deep (♂), or very deep (♀). Coxa IV two and half times longer than coxa II. Coxae I–IV surfaces with scattered granules and margins densely granular; coxa II, prolateral subproximal margin with three oblique slit-like structures, adjacent to a moderate (♀) or low (♂) granular protuberance; coxae II–IV, prolateral carinae strongly granular (Figures
Pedipalps, Femur prolateral, dorsal and retrolateral intercarinal surfaces shagreened (Figure
Trichobothrial pattern Type C, neobothriotaxic. Femur trichobothria d, e, and i positioned proximally, equidistant; d on dorsal surface, e on ventral prolateral carina, i ventral to dorsal prolateral carina (Figure
Legs: Basitarsi, prolateral ventral and retrolateral ventral spinule rows partial, distal half with two or three sparse spinules on legs I–III, absent on leg IV; retrolateral and retrolateral dorsal rows absent on I–IV; macrosetal counts on legs I–IV, respectively: dorsal 2:2:2:2, retrolateral dorsal, 2:2:3:3; retrolateral ventral, 5:5:5:5; prolateral ventral, 4:4:4:4, all macrosetae not pigmented, translucent and shaped as tines; dorsal and retrolateral dorsal macrosetae arranged in two separate parallel rows on legs I–IV. Telotarsi I–IV, each with single irregular ventromedian row of scattered spinules and one ventrodistal spinule, flanked by prolateral and retrolateral rows of six macrosetae. Ungues short and curved.
Genital operculum: Wider than long, with four (♂) or six (♀) pairs of short and translucent macrosetae; sclerites free longitudinally, anterior margin fused on distal two thirds (♂) or fused longitudinally by a loose pleura folding into a valve covering the genital opening (♀). Genital papillae present, protruding posteriorly (♂) or absent (♀) (Figures
Megacormus granosus (Gervais, 1844). 20 ♂ dorsal coxosternal region, genital operculum, pectines, and sternites III–VI 21 ♀ pectines and genital operculum 22 telson lateral view 23–25 metasomal segments I–V dorsal, lateral, and ventral view. Scale bars: 0.5 mm, except Figure 21: 0.25 mm.
Hemispermatophore: Distal lamina 1.1 times the length of trunk; tapering distally, basal constriction well-developed (Figures
Pectines: Basal piece with three or five pairs of macrosetae, proximal surface granular, V-shaped (♂) or isosceles trapezoidal (♀). Marginal and median lamellae nearly fused into one piece with a fine, shallow furrow (♂), or completely fused, indistinguishable furrow (♀). Fulcra absent. Pectinal teeth: three to four (♂) or one to four (♀). Pectines relatively short, fused lamella aligned with midpoint of coxa IV (Figures
Tergites: I–VI, intercarinal surfaces shagreened, densely covered with minute and coarse granules, posterior margin with rows of irregular granules (Figures
Sternites: Sternite III, surface around pectines shagreened; sternites IV–VI, surfaces smooth to weakly granular medially, shagreened laterally; spiracles minute, ovoid, 2.0 times longer than wide; sternite V, ventral surface distinct hyaline glandular area posteromedially, densely cover with micropores (♂) or absent (♀) (Figure
Metasoma: Length 0.9 times greater than mesosomal length (Table
Measurements (mm) of six adult males and three females of Megacormus granosus (Gervais, 1844).
♂/♂/♂/♂/♂/♂/♀/♀/♀ | ||
---|---|---|
Carapace | length | 3.40/3.25/3.20/3.00/3.25/3.15/4.20/4.30/3.90 |
Anterior | width | 2.00/1.80/1.80/1.60/1.80/1.75/2.40/2.50/2.35 |
Posterior | width | 3.60/3.35/3.35/3.20/3.35/3.25/4.40/4.20/4.25 |
Femul | length | 2.70/2.65/2.70/2.45/2.60/2.55/3.30/3.60/3.05 |
width | 1.10/1.00/0.95/0.90/1.00/1.00/1.30/1.30/1.30 | |
height | 2.80/2.80/2.60/2.60/2.85/2.65/3.50/3.80/3.35 | |
Patella | width | 1.40/1.25/1.40/1.30/1.40/1.30/1.90/1.80/1.80 |
Chela | length | 5.30/5.20/5.05/4.70/5.15/5.00/6.50/7.10/6.4 |
Manus | width | 1.70/1.60/1.55/1.50/1.55/1.50/2.20/2.30/1.95 |
height | 1.40/1.20/1.35/1.05/1.45/1.15/1.70/1.70/1.75 | |
Fixed finger | length | 2.30/2.30/2.30/2.15/2.20/2.15/3.00/3.00/2.80 |
Movable Finger | length | 3.00/2.95/3.00/2.70/2.85/2.85/3.90/4.10/3.75 |
Coxa II | length | 1.30/1.35/1.25/1.20/1.40/1.25/1.50/1.70/1.75 |
Coxa IV | length | 2.70/2.55/2.60/2.45/2.55/2.40/3.40/3.60/3.40 |
Sternum | length | 0.80/0.70/0.70/0.65/0.75/0.85/1.00/0.90/1.05 |
Sternum | width | 0.90/0.90/0.90/0.90/0.85/0.70/1.00/0.70/1.15 |
Mesosoma | length | 5.20/4.70/5.35/4.35/5.30/4.90/7.00/7.70/6.80 |
Metasoma | length | 8.50/4.48/4.60/4.23/4.30/4.10/8.60/9.10/4.25 |
Segment I | length | 0.90/1.00/1.10/1.10/1.05/1.00/1.00/1.20/1.05 |
width | 2.00/1.95/1.95/1.65/1.95/1.80/2.10/2.20/2.10 | |
height | 1.70/1.50/1.60/1.50/1.55/1.60/1.70/1.70/1.85 | |
Segment II | length | 1.20/1.20/1.25/1.25/1.15/1.10/1.20/1.20/1.30 |
width | 1.90/1.85/1.85/1.60/1.75/1.70/1.90/2.00/1.90 | |
height | 1.70/1.35/1.55/1.40/1.35/1.45/1.50/1.70/1.65 | |
Segment III | length | 1.40/1.35/1.40/1.45/1.30/1.20/1.30/1.30/1.30 |
width | 1.80/1.75/1.75/1.60/1.70/1.65/1.80/1.90/1.80 | |
height | 1.60/1.30/1.55/1.40/1.40/1.35/1.50/1.60/1.65 | |
Segment IV | length | 1.80/1.90/1.95/1.65/1.80/1.80/1.90/1.90/1.75 |
width | 1.70/1.60/1.65/1.50/1.55/1.50/1.60/1.70/1.60 | |
height | 1.70/1.40/1.55/1.40/1.50/1.35/1.50/1.60/1.60 | |
Segment V | length | 3.20/3.50/3.50/3.00/3.30/3.10/3.20/3.50/3.10 |
width | 1.50/1.60/1.65/1.45/1.55/1.45/1.50/1.50/1.55 | |
height | 1.60/1.25/1.50/1.35/1.45/1.25/1.40/1.50/1.55 | |
Telson | length | 4.20/3.90/4.25/3.80/4.00/3.80/4.50/4.50/4.30 |
Vesicle | length | 2.90/2.25/2.45/2.30/2.35/2.20/2.90/2.60/2.30 |
width | 1.90/1.65/1.65/1.55/1.70/1.50/1.50/1.60/1.50 | |
height | 1.30/1.30/1.45/1.35/1.35/1.30/1.30/1.30/1.25 | |
Aculeus | length | 1.30/1.65/1.80/1.50/1.65/1.60/1.60/1.90/2.00 |
Total | length | 21.30/16.33/17.40/15.38/16.85/15.95/24.30/25.60/19.25 |
Telson: Vesicle globose, length 1.4 times greater than width (Table
Megacormus granosus has been reported in the vicinities of the National Park Pico de Orizaba, on the slopes of the Trans-Mexican Volcanic Belt facing the Gulf of Mexico, and between Orizaba and Huatusco, Veracruz.
All adult males were collected by pitfall traps, suggesting high motility within the leaf litter. They were particularly abundant in the May 2012 expedition. This behavior in males and the period of the year may be related to the mating season of the species. All adult females and immatures of both sexes were collected exclusively using Berlese funnels, suggesting these are comparatively less mobile. A total of 72 Berleses and 180 pitfalls were used to sample two hectares, of which 18 (25%) and 11 (5%), caught 18 and 9 specimens respectively. These yields are consistent with low population density of this species; adult males are particularly rare. The habitat of, and behavior exhibited by, this species as well as its cryptic morphology (color resembling substrate; relative small size) are congruent with a humiculous ecomorphotype (
The catalog of the scorpions of the world (
The illustrations of the M. granosus hemispermatophore presented in Figures
We would like to thank the community of Atotonilco de Calcahualco for allowing collection on their land and providing logistics in the field: Mr. Isidoro Contreras, Ms. Mercedes Contreras, Mr. Nicolas, and Ms. Noemi. We thank Mr. Armando Fuertes from the National Park Pico de Orizaba for introducing us to the Atotonilco community. The following students assisted during field expeditions: Uriel Garcilazo Cruz, Miguel Hernández Patricio, Dulce Flor Piedra Jiménez, Francisco Andrés Rivera Quiroz, Francisco Javier Salgueiro Sepulveda, and Mariana Servín Pastor. Funding for this study was provided by UNAM-DGAPA-PAPIIT, project IN213612. We thank P. Sharma for editing English, and the editor and two anonymous reviewers for improving earlier drafts of the manuscript.