Correction
18 Mar 2015: The PLOS ONE Staff (2015) Correction: First Record of the Myrmicine Ant Genus Meranoplus Smith, 1853 (Hymenoptera: Formicidae) from the Arabian Peninsula with Description of a New Species and Notes on the Zoogeography of Southwestern Kingdom Saudi Arabia. PLOS ONE 10(3): e0118389. https://doi.org/10.1371/journal.pone.0118389 View correction
Figures
Abstract
The ant genus Meranoplus is reported for the first time from the Arabian Peninsula (Kingdom of Saudi Arabia) by the new species M. pulcher sp. n., based on the worker caste. Specimens were collected from Al Sarawat and Asir Mountains of southwestern Kingdom of Saudi Arabia using pitfall traps. Meranoplus pulcher sp. n. is included in the Afrotropical M. magretii-group, with greatest similarity to M. magrettii André from Sudan. A key to the Afrotropical species of the M. magretii-group is presented. A brief review of the ant taxa with Afrotropical affinities in southwestern region of the Kingdom of Saudi Arabia is given.
Citation: Sharaf MR, Al Dhafer HM, Aldawood AS (2014) First Record of the Myrmicine Ant Genus Meranoplus Smith, 1853 (Hymenoptera: Formicidae) from the Arabian Peninsula with Description of a New Species and Notes on the Zoogeography of Southwestern Kingdom Saudi Arabia. PLoS ONE 9(11): e111298. https://doi.org/10.1371/journal.pone.0111298
Editor: Ulrike Gertrud Munderloh, University of Minnesota, United States of America
Received: August 11, 2014; Accepted: September 18, 2014; Published: November 6, 2014
Copyright: © 2014 Sharaf et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Data Availability: The authors confirm that all data underlying the findings are fully available without restriction. All relevant data are within the paper.
Funding: This project was supported by the NSTIP Strategic technologies program, grant number (12-ENV2484-02), in the Kingdom of Saudi Arabia. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
Competing interests: The authors have declared that no competing interests exist.
Introduction
Since Smith [1] established the genus Meranoplus, 87 species and subspecies are currently included [2]. This genus is broadly distributed in the Old World tropics, ranging through the Afrotropical, Malagasy, Oriental, and Indo-Australian regions [3]–[5]. The genus is absent from the Palaearctic and Oceania regions, except for a single species, M. levellei Emery, 1883 from New Caledonia [6], [7].
Meranoplus has been historically placed in the tribe Meranoplini, which also includes the genera Dicroaspis Emery, 1908 and Calyptomyrmex Emery, 1887 [8]. Recently, the tribe Meranoplini was restricted to two genera, Meranoplus [9] and a fossil genus Parameranoplus Wheeler, 1915 [10]. The Meranoplus fauna has been revised for most zoogeographical realms including the Afrotropical [3], Oriental [11], [12], Australian [13]–[17], and Malagasy [5]. Among the previous regions, the Australian region is the most taxa rich in regard to the number of species [3], [13].
Available information on the habitats and biology of Meranoplus species is limited. Most species nest directly in the soil, under stones, in rotten wood or in leaf litter [3], [4], [13]. African species have been reported to nest either in the ground with a crater type of entrance, or at the base of plants. Nests of one or two species are constructed among roots with workers ascending trees or low shrubs [3]. The majority of species are generalized omnivores [13], whereas others are considered seed harvesters, e. g. M. diversus-group [4], [13], [18].
The specialized morphology of the genus Meranoplus is related to a specialized behaviors, thanatosis, “playing dead” and and becoming cryptic. Several species when disturbed, fold their legs beneath the promesonotal shield and quickly accumulate dirt on body hairs and remain motionless in a fetal-like position [19], [20].
A thorough diagnosis of the genus Meranoplus has been provided by Bolton [3], [21]. The genus can be recognized by the combination of the following traits, masticatory margin of mandibles armed with four to five teeth; palp formula 5, 3; clypeus large, with median portion usually carinate at each side; antennal scrobes well-developed, usually long; antennae nine-segmented with a three segmented club; eyes large, located behind midlength of head, sometimes close to the posterior corners of head; pronotum and mesonotum fused into a plate or shield that extends posteriorly and laterally so the mesosomal sides, and usually also the propodeum, are concealed in dorsal view; lateral and/or posterior margins of promesonotal shield usually armed with spines, lobes, foliacious processes; petiole sessile usually cuneate in lateral view.
The M. magrettii-group can be distinguished by the following characters [3]: masticatory margin of mandibles armed with four or five teeth; mesosoma when seen from above with the propodeum concealed by the broad promesonotal shield; propodeal spines present; petiole cuneate in lateral view with unarmed dorsal surface; postpetiole broad and nodiform.
The M. magrettii-group has only two species known from the Afrotropical region, M. magrettii a species that is broadly distributed in the region and inhabiting savannah, grassland and dry woodland, and M. peringueyi apparently confined to South Africa [3].
The southwestern mountainous region of the Kingdom of Saudi Arabia (KSA) is one of the most diverse in terms of species diversity and relative abundances of insects. Taxonomic and biogeographical studies have indicated that the insects of this region have strong affinities with the Afrotropical Region [21]–[33]. A recent and comprehensive study of the Afrotropical relationships of the region is by El-Hawagryi et al. [32]. These authors recorded 17 orders of insects representing 129 families and at least 582 species and subspecies from Al-Baha Province. Their biogeographic analysis of the species composition clearly revealed that this region has a close affinity with the insect fauna of the Afrotropical Region.
In the present study, the myrmicine ant genus Meranoplus is recorded for the first time from KSA by the new species M. pulcher sp. n. and represents a new generic record for the Arabian Peninsula. In addition, a synopsis of the similarity of the Afrotropical fauna with the southwestern region of KSA is presented based on available records of ants.
Materials and Methods
Study area
Shada Al Ala Mountain is a parallel extension of Hijaz Mountains to the west. This region is managed as a natural protectorate in southwestern of Al Baha Province, 20 km northwest of Al Mukhwah Governorate. The region has a substantial high diversity of wild plants including Albizia lebbeck (L.) Benth. (Fabaceae), Solenostemon Thonn.(Lamiaceae), Juniperus procera Hochst. ex Endlicher (Cupressaceae), Santalum L. (Santalaceae), Pimpinella anisum L. (Apiaceae), Rhamnus frangula L. (Rhamnaceae), Opuntia ficus-indica (L.) Mill. (Cactaceae), Ricinus communis L. (Euphorbiaceae), Olea europaea ssp. africana (Mill.) P. Green. (Oleaceae), Prunus dulcisn (Mill.) D.A.Webb (Rosaceae), Maerua crassifolia Forssk. (Capparceae), Pandanus tectorius Parkinson (Pandanaceae), Panicum Turgidum Forssk. (Poaceae), Coffea arabica L. (Rubiaceae), Opuntia ficus-indica (L.) Mill. (Cactaceae), Breonadia salicina (Vahl) Hepper & J.R.I.Wood (Rubiaceae), Haloxylon salicornicum (Moq.) Bunge ex Boiss. (Chenopdiaceae), Lycium shawii Roem. & Schult (Solanaceae), Cactus (Cactaceae), and Acacia spp. (Memosaceae).
Sampling procedures
Materials listed in this work, the holotype and 29 paratype specimens were collected during an insect inventory of the southwestern region of KSA by using more than 900 pitfall traps. A single specimen of the new species was collected by an aspirator from a Cactus sp. All the materials is deposited in King Saud University Museum of Arthropods, College of Food and Agriculture Sciences (KSMA), King Saud University, Riyadh, Kingdom of Saudi Arabia, except a single paratype specimen in California Academy of Sciences (CASC), San Francisco, USA.
All measurements and indices are expressed in millimeters and follow the standards of Boudinot & Fisher [5].
Measurements
ATL: Abdominal Tergum IV Length. Maximum length of fourth abdominal tergum measured with anterior and posterior margins in same plane of focus.
ATW: Abdominal Tergum IV Width. Maximum width of fourth abdominal tergum with anterior, posterior, and lateral borders in same plane of focus.
CDD: Clypeal Denticle Distance. Distance between clypeal denticle apices, measured in full-face view.
CW: Clypeus Width. Distance between the apices of the frontal lobes across the clypeus.
EL: Eye Length. Maximum eye length in profile view.
EW: Eye Width. Maximum eye width in profile view.
HL: Head Length. Maximum length of head capsule, excluding mandibles, measured from anterior margin of clypeus to nuchal carina, with both in same plane of focus.
HLA: Head Length, Anterior. Distance between the anterior edges of the eyes to the mandible bases in full-face view.
HW: Head Width. Maximum width of head capsule behind the eyes, in full-face view.
PML: Promesonotum Length. Maximum length of promesonotum from posterior spine/denticle apices to anterolateral denticle apices; all four apices in same plane of focus. ( = PMD, [15])
PPH: Postpetiole Height. Measured from sternal process base to postpetiole apex in lateral view.
PPL: Postpetiole Length. Measured from anterior to posterior inflections of postpetiole node in lateral view.
PWA: Promesonotal Width, Anterior. Maximum width of promesonotal shield between anterolateral denticle apices in dorsal view. ( = PW, [15])
PWP: Promesonotal Width, Posterior. Distance between posterior-most promesonotal spine or denticle apices.
PTH: Petiole Height. Measured from petiole sternum to apex in lateral view.
PTL: Petiole Length. Measured from anterior to posterior inflections of petiole node.
SL: Scape Length. Maximum length of the scape excluding basal constriction.
SPL: Propodeal Spine Length. Workers: distance from inner posterior margin of propodeal spiracle to propodeal spine apex. Gynes: maximum propodeal spine length from basal inflection of spine, to spine apex.
WL: Weber's Length. Maximum diagonal length of mesosoma from anterior inflection of pronotum to posterolateral corner of the metapleuron or the metapleural lobes, whichever is most distant.
Indices
CDI: Clypeal Denticle Index. CDD×100/CML
CI: Cephalic Index. HW×100/HL
CS: Cephalic Size. (HW+HL)/2
EYE: Eye Index. 100× (EL+EW)/CS
OMI: Ocular-Mandibular Index. EL×100/HLA
PMI: Promesonotum Index 1. PWA×100/PML ( = PMI2, [15])
PPI: Postpetiole Index. PPL×100/PPH
PTI: Petiole Index. PTL×100/PTH
PWI: Promesonotum Index 2. PWP×100/PML
SEI: Scape-Eye Index. EL×100/SL
SI: Scape Index. SL×100/HW
Illustrations
Specimens were photographed by Michele Esposito (CASC) using a JVC KYF70B3CCD digital camera attached to a Leica M420 stereomicroscope. All digital images were processed using Auto-Montage (Syncroscopy, Division of Synoptics Ltd, USA) software. Images of the specimens are available in full color on www.antweb.org. The map was created by the ArcGIS 9.2 program, with the help of Prof. Mahmoud S. Abdel-Dayem (King Saud University).Specimens were examined and imaged using scanning electron microscope (SEM) (JSM-6380 LA) visualization to record morphological details of the new species. The JSM-6380 LA is a high-performance scanning electron microscope with a high resolution of 3.0 nm.
No specific permits were required for the described field studies or for the surveyed locations which are not privately-owned or protected in any way or do not have endangered or protected species.
Nomenclatural acts
The electronic edition of this article conforms to the requirements of the amended International Code of Zoological Nomenclature, and hence the new names contained herein are available under that Code from the electronic edition of this article. This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the ICZN. The ZooBank LSIDs (Life Science Identifiers) can be resolved and the associated information viewed through any standard web browser by appending the LSID to the prefix "http://zoobank.org/". The LSID for this publication is: urn:lsid:zoobank.org:pub:80A78374-53CF-4A19-90CB-4A392E626999. The electronic edition of this work was published in a journal with an ISSN, and has been archived and is available from the following digital repositories: PubMed Central, LOCKSS.
Results
Meranoplus pulcher SHARAF sp. n. (Figs. 1–11)
urn:lsid:zoobank.org:act:A5035370-971E-451B-B15E-23F8BBCE5847.
Holotype worker
SAUDI ARABIA, Al-Baha Province, Shada Al Ala, 19°51.066'N, 41°18.037'E, 1325 m, 23.IV.2014, P. T. (Al Dhafer et al. Leg.), deposited in KSMA, King Saud Museum of Arthropods, College of Food and Agriculture Sciences, King Saud University, Riyadh, Kingdom of Saudi Arabia.
Paratypes workers
All the following paratype specimens are deposited in KSMA, 3 workers, same locality and data as the holotype; 1 worker with same data as the holotype except the collecting data 15.II.2014; 1 worker, SAUDI ARABIA, Asir Province, Raydah, 18°11.749'N, 42°23.345'E, 1614 m, 28.IV.2014, P.T. (Al Dhafer et al. Leg.), 3 workers, Shada Al Ala, 19°50.575'N, 41°18.691'E, 1666 m, 23.VIII.2014, P. T. (Al Dhafer et al. Leg.); 9 workers, Shada Al Ala, 19°50.411'N, 41°18.686'E, 1611 m, 23.VIII.2014, P. T. (Al Dhafer et al. Leg.); 3 workers, Shada Al Ala, 19°50.329'N, 41°18.604'E, 1563 m, 23.VIII.2014, P. T. (Al Dhafer et al. Leg.); 5 workers, Shada Al Ala, 19°50.710'N, 41°18.267'E, 1474 m, 23.VIII.2014, P. T. (Al Dhafer et al. Leg.); 5 workers, Shada Al Ala, 19°51.066'N, 41°18.037'E, 1325 m, 23.VIII.2014, P. T. (Al Dhafer et al. Leg.); 4 workers, Asir Province, Raydah, 18°11.618'N, 42°23.420'E, 1772 m, 26.VIII.2014, P. T. (Al Dhafer et al. Leg.); 1 workers, Asir Province, Raydah, 18°11.749'N, 42°23.345'E, 1614 m, 26.VIII.2014, P. T. (Al Dhafer et al. Leg.); unique specimen identifier CASENT 0914336, in (CASC) California Academy of Science Collection, San Francisco, California, USA.
Worker measurements
Maximum and minimum based on all specimens, n = 5, (holotype): TL 3.20-3.70 (3.27), HL 0.77–0.87 (0.80), HW 0.67–0.82 (0.72), HLA 0.25–0.30 (0.25), CW 0.22–0.27 (0.30), CDD 0.12–0.15 (0.12), SL 0.47–0.62 (0.60), EL 0.17–0.22 (0.17), EW 0.12–0.15 (0.15), PML 0.40–0.52 (0.47), PWA 0.62–0.75 (0.70), PWP 0.37–0.47 (0.45), SPL 0.17–0.22 (0.22), WL 0.75–0.87 (0.77), PTL 0.12–0.17 (0.20), PTH 0.30–0.42 (0.37), PPL 0.15–0.22 (0.22), PPH 0.25–0.35 (0.32), ATW 1.02–1.22, (1.12) ATL 0.97–1.15 (1.05), CI 87–94 (90), SI 67–82 (83), OMI 63–80 (68), CDI 0.44–−0.68 (40), SEI 31–43 (28), PMI 144–155 (149), PPI 60–80 (69), PTI 40–46 (54), PWI 82–93 (96), CS 0.72–0.84 (0.76), EYE 38–47 (42) (n = 5).
Diagnosis
Although M. pulcher sp. n. is superficially similar to M. magrettii (Figs. 12, 13), it can be readily distinguished by the following contrasting characters: Colour: M. pulcher is yellow, M. magrettii is light to dark brown; anterior clypeal margin: distinctly concave in M. pulcher, more or less flat to shallowly concave in M. magrettii; subpetiolar process: in M. pulcher short and triangular, in M. magrettii the process is more developed forming a short finger or a less developed process; petiolar sculpture: posterior face of petiolar node areolate-rugose in M. pulcher and smooth in M. magrettii; sculpture of first gastral tergite: superficially and finely shagreenate in M. pulcher, in M. magrettii the sculpture of first gastral tergite varies from dense shagreenate or reticulate punctate.
Description
Head.
Head slightly longer than broad with convex sides and straight posterior margin; anterior clypeal margin distinctly concave with well-developed clypeal carinae; mandibles armed with four teeth; eyes relatively large (EL 0.25–0.26 x HW; EYE 38–47) with 12 ommatidia in the longest row; scapes when laid back from their insertions just reach posterior margin of eyes; scrobal carinae well-developed.
Mesosoma.
Anterior pronotal corners armed with a pair of short triangular teeth; promesonotal shield distinctly broader than long (PMI 144–155) widening behind pronotum; promesonotal suture absent; posterior corners of mesonotum armed with a pair of sharp spines; posterior mesonotal margin between spines strongly concave and without secondary armament; propodeal spines long and sharp originating at level of propodael spiracles and curved upwards; propodeal lobes well-developed.
Waist.
Petiole cuneate in profile, sessile, with a broad anterior margin and a narrow acute dorsum; petiolar and postpetiolar anteroventral processes present; postpetiole nodiform, subrectangular in profile, taller than broad (PPI 60–80).
Sculpture.
Mandibles longitudinally striated; cephalic dorsum densely and finely longitudinally rugulose, posterior margin areolate-rugose; promesonotal shield, posterior face of petiolar node and postpetiole dorsum reticulate rugulose, anterior petiolar face smooth and sides transversally rugulose; first gastral tergite finely and densely shagreenate.
Etymology
The species name is derived from the Greek word “pulcher” that means “beautiful” referring to the attractive appearance of this ant species.
Habitat
Twenty five workers of the new species were collected from Al-Baha Province, Shada Al Ala Protectorate (Fig. 14) and six workers from Raydah Protectorate. Both collections were from pitfall traps placed next to Acacia trees. The soil was extremely dry with abundant dry seeds of shrubs. Despite several hours of observing the nest no additional specimens were found. Pitfall trapping is apparently an efficient method for collecting this group of ants.
Key to the Afrotropical species of Meranoplus magrettii-group based on workers
In the key to Afrotropical species (Bolton [3]:48), Meranoplus pulcher sp. n. will key to couplet 7 along with peringueyi and magrettii. Couplet 7 is modified here to separate the three species of the M. magrettii-group.
7. Mandibles armed with 5 teeth peringueyi Emery
- Mandible armed with 4 teeth 8
8 Colour moderately dark brown, Subpetiolar process more developed, in the form of a short finger (Fig. 12), Posterior face of petiolar node smooth (Fig. 13). (Ghana, Sudan, Uganda, Kenya, Tanzania, Zimbabwe, Botswana, South Africa, Fig. 15) magrettii André
- Colour yellowish, Subpetiolar process present but short (Fig. 7), Posterior face of petiolar node areolate-rugose. (Fig. 8) (Saudi Arabia, Fig. 15) pulcher sp. n.
Zoogeography of the southwestern region of Kingdom of Saudi Arabia
The KSA is located at the interchange of three major biogeographical realms, the Palaearctic, Afrotropical and Oriental. Much mixing of these faunal elements has occurred. Geologically, the Ethiopian and Arabian Peninsula highlands and mountains separated approximately 13 mybp producing the Great Rift Valley through a rifting process as the African continental crust separated [34], [35]. Biogeographically, the southwestern region of the KSA belongs to the Afrotropical region [21]–[23], [25]–[33].
Studies of other taxonomic groups of insects have revealed similar results, for example, the Scythrididae (flower moths) (Lepidoptera: Gelechioidea) were treated for the Palaearctic region [36] and the Afrotropical species Scythris albocanella Bengtsson 2002 was recorded from various localities in southwestern region of KSA. In addition, Marnert et al. [37] studied the pseudoscorpion arachnids (Pseudoscorpiones) of the region and concluded that the southwestern region of KSA has a clear faunal similarity with the Afrotropical region.
At least ten pantropical ant genera have been recorded from this region of KSA including Anochetus Mayr, 1861, Cryptopone Emery, 1893, Cerapachys Smith, 1857 Dorylus Fabricius, 1793, Hypoponera Santschi, 1938, Leptogenys Roger, 1861, Melissotarsus Emery, 1877, Platythyrea Roger, 1863, Polyrhachis F. Smith, 1857 and Tetraponera F. Smith, 1852. Strong biogeographical affinities are with the Afrotropical Region [38]. The sole faunistic work carried out for the knowledge of the ants of KSA [38] recorded 89 species from southwestern region, 18 of which having an Afrotropical distribution (table 1). Despite limited amount of specimens collected from this region mentioned in the above work, preliminary conclusions supported an Afrotropical faunal relationship of the region. Recently, six new species were described from the region [28]–[33], taxa more closely related to Afrotropical congeners (Table 2). An approximate estimate of the relative percentage of the Afrotropical faunal similarity of the region is 31%. Taking into account the unidentified materials accumulated over the last ten years from the region by the authors and also the vast area of the region, some of which has not been surveyed, it is expected this number will increase.
Discussion
Meranoplus pulcher sp. n. is the first member of the genus recorded from KSA and from the vast Arabian Peninsula. Following Bolton [3], it belongs to M. magrettii-group and cannot be identified using the available keys to species of Afrotropical [3], Malagasy [5], Oriental [11] or Australian [13], [17] regions. Meranoplus pulcher sp. n. is similar to M. magrettii André from Sudan to which it will key to in Bolton [3], sharing the following characters: mandibles striate and armed with four teeth, anterior pronotal corners armed with a pair of short triangular teeth, promesonotal shield narrowing behind pronotum, posterior corners of mesonotum armed with a pair of short spines, posterior mesonotal margin concave and unarmed, petiole cuneate in profile and postpetiole nodiform.
The habitats of M. pulcher sp. n. and M. magrettii apparently are not similar. The latter species is restricted to sub-Saharan Africa and has been collected from savannah, open-woodland and grassland habitats [3]. Meranoplus pulcher sp. n.is apparently restricted to juniper woodlands of southwestern mountains of KSA. The author (MRS) has made extensive collections of ants from the southwestern KSA. Typical Afrotropical ant genera mentioned above (e.g. Strumigenys, Anochetus, Pachycondyla, Cerapachys, Dorylus etc.) were commonly encountered. For example, the Afrotropical species, S. arnoldi Forel was reported from Al-Baha Province [39] providing evidence of faunal similarities with the Afrotropical Region. Additional future collections from this area of KSA will no doubt provide further evidence of this biographical connection.
The record of M. pulcher sp. n. of the Afrotropical margettii-group is an additional evidence of the Afrotropical faunal similarities of the southwestern mountains of KSA which is consistent with other faunal influences for the Region [21]–[23], [25]–[33].
Acknowledgments
We are grateful to Prince Bandar Bin Saud Al Saud, Head of the Saudi National Commission for Wildlife Conservation and Development for the appreciated support during the study. The authors are grateful to Barry Bolton and Brendon Boudinot for valuable suggestions that improved the manuscript. Special thanks to Brian Fisher and Michele Esposito (California Academy of Sciences, San Francisco, USA) for assistance in photographing the new species, to Loutfy El-Juhany for identifying plants of the studies area, and to Mahmoud Abdel-Dayem for making the map. The authors are indebted to Boris Kondratieff (Colorado State University) for his valuable comments and critical editing, actually, without his help this work could not have been completed.
Author Contributions
Conceived and designed the experiments: MRS HMAD ASA. Performed the experiments: MRS HMAD ASA. Analyzed the data: MRS HMAD ASA. Contributed reagents/materials/analysis tools: MRS HMAD ASA. Wrote the paper: MRS HMAD ASA.
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