Monomorium (Hymenoptera: Formicidae) of the Arabian Peninsula with description of two new species, M. heggyi sp. n. and M. khalidi sp. n.

We present a revised and updated synoptic list of 44 Arabian Monomorium species, including two new species of the M. salomonis species-group: M. heggyi sp. n., and M. khalidi sp. n. We propose the following new synonyms: M. abeillei André (= M. wahibiense Collingwood & Agosti syn. n.); M. areniphilum Santschi (= M. fezzanense Collingwood & Agosti syn. n., = M. hemame Collingwood & Agosti syn. n. = M. marmule Collingwood & Agosti syn. n.); M. bicolor Emery (= M. phoenicum Santschi syn. n.); M. harithe Collingwood & Agosti (= M. najrane Collingwood & Agosti syn. n.); M. niloticum Emery (= M. matame Collingwood & Agosti syn. n.); and M. nitidiventre Emery (= M. yemene Collingwood & Agosti syn. n.). An illustrated key and distribution maps are presented for the treated species. Ecological and biological notes are given when available. The majority of Arabian Monomorium species (24) are endemic to the peninsula. All except one of the remaining species are more broadly ranging Afrotropical and Palearctic species, supporting the view of Arabia as a biogeographical crossroads between these two regions. Monomorium floricola (Jerdon), the sole species of Indomalayan origin, is recorded for the first time from the Arabian Peninsula.


INTRODUCTION
Mayr is one of the largest genera of ants, with 326 valid species and subspecies, as well as three fossil species (Bolton, 2020). Monomorium species predominantly inhabit tropical, subtropical, and warm temperate regions of the world (Bolton, 1987(Bolton, , 1994Brown, 2000). The majority of Monomorium species are native to the updated taxonomic revision with likely more than 100 undescribed species and many valid species needing reevaluation (Bolton, 1987;Hita Garcia, 2020, personal observation).
The oldest records of Monomorium species from the Arabian Peninsula were of M. carbonarium (Smith) and M. niloticum Emery (Forel, 1907). Collingwood (1985) reported 17 valid Monomorium species from the Kingdom of Saudi Arabia (KSA) and  listed 49 valid Monomorium species from the entire Arabian Peninsula, including 32 new species. Additional articles have included records from the KSA Sharaf & Aldawood, 2013b;Sharaf et al., 2015Sharaf et al., , 2018aSharaf et al., , 2018b, Oman (Sharaf et al., 2018b;Monks et al., 2019), the Socotra Archipelago (Collingwood et al., 2004;Sharaf et al., 2017a), the United Arab Emirates (UAE) (Collingwood et al., 2011), and Yemen (Aldawood, Sharaf & Collingwood, 2010). Collingwood (1985) and  recognized a total of 53 Monomorium species on the Arabian Peninsula, of which 17 were described from countries in the region. However, the taxonomic status of several species has remained uncertain due to brief descriptions with insufficient differential diagnoses, apparent ambiguities in the taxonomic keys, and lack of species-group assignment (Sharaf et al., 2020a(Sharaf et al., , 2020b. The present study aims to clarify the current status of the Arabian Monomorium by providing the following: diagnoses of Arabian Monomorium and Monomorium species-groups, a synoptic checklist of Arabian Monomorium, an illustrated identification key to species based on the worker caste, new taxonomic treatments proposing new synonymies and describing two new species, and biogeographical analyses, including distribution maps.

MATERIALS AND METHODS
The species names follow the online catalogue of ants of the world (Bolton, 2020). We made digital color images of each species using a Leica DFC450 digital camera with a Leica Z16 APO microscope and LAS (v3.8) software. The images are available online on AntWeb (http://www.AntWeb.org) and are accessible through unique specimen identifiers (e.g., CASENT0922288). Distribution maps were made using DIVA-GIS (version 7.5.0.0).

Measurements and indices
All measurements are in millimeters and follow the standard measurements of Bolton (1987) and Sharaf et al. (2018a). CI = Cephalic Index (HW/HL × 100). EI = Eye Index (EL/HW × 100). EL = Eye Length; maximum diameter of eye in lateral view. EM = Eye-Mandible Distance; distance between anterior margin of eye and mandibular insertion in lateral view. HL = Head Length; maximum length of head, excluding mandibles in full-face view. HW = Head Width; maximum width of head directly behind eyes in full-face view. and for Monomorium khalidi sp. n. is urn:lsid:zoobank.org:act:3B5BB529-D842-4146-B8F7-ACCAC9CD5BA7. The online version of this work is archived and available from the following digital repositories: PeerJ, PubMed Central and CLOCKSS.

Diagnosis of Arabian Monomorium species-groups
The M. monomorium species-group can be readily recognized by the following combination of characters in the worker caste (Sharaf et al., 2018a): monomorphic, with size variation; head longer than broad; mandibles smooth and masticatory margin armed with four teeth; antennae with 10-12 segments, terminating in a three-segmented club; median clypeal portion raised anteriorly and longitudinally bicarinate; eyes present with variable size, located in front of the midlength of the sides in full-face view, and with four or more ommatidia in the longest row; head smooth and shining; metanotal groove well-defined, with distinct cross-ribs; propodeal spiracle circular to subcircular; propodeal dorsum meeting declivity in a rounded angle; promesonotum and propodeal dorsum smooth; body pilosity variable but usually distinct; petiole, postpetiole and gastral tergites usually smooth. The M. salomonis species-group can be diagnosed by the following character states in the worker caste (Bolton, 1987): monomorphic, with minor size variation; palp formula 2,2 or 1,2 in some minute species; mandibles usually sculptured; masticatory margins of mandibles armed with four teeth which decrease in size from apex to base; median clypeal portion raised, projecting anteriorly; cephalic dorsum usually sculptured, ranging from dense blanketing reticulate-punctuation to feeble superficial reticular patterning; eyes prominent of medium to large size, usually with six or more ommatidia in the longest row; eyes circular to oval in shape; head longer than broad; scapes usually relatively long (SI > 80); metanotal groove moderately impressed to absent; metanotal cross-ribs inconspicuous to absent; propodeal spiracle circular to subcircular; propodeum rounded to angular between dorsum and declivity; propodeal dorsum usually sculptured but never transversely striate; petiolar spiracle situated at the node or immediately in front of the anterior face of the node; body pilosity variable in distribution and density, but usually reduced on the head and mesosoma; mesosoma, petiole and postpetiole usually sculptured; first gastral tergite usually shagreenate or finely sculptured. 8. Eyes larger, with a ring of seven to eight ommatidia encircling a single row of 2 ommatidia, and in profile closer to mandibular insertions (EM 0.05); meso-and metapleuron smooth; petiole and postpetiole smooth and each with one pair of standing hairs (Fig. 2C) (Fig. 2D) 16. Underside of head without long, standing hairs (Fig. 4A)         Remarks. Monomorium wahibiense is represented by a single worker deposited in WML and accompanied by a red card and handwritten label by C. Collingwood indicating that this specimen represents the syntype. The label's data are consistent with the data for the type in the original description in terms of collecting locality (Oman, Wahiba sand) and collector (M.D. Gallagher), but not the collection date, which we consider a typographical error. Comparing the mentioned type material with the image of the type material of M. abeillei André, we found the two species share the same morphological characters, which can be summarized as follow: scapes relatively short, when laid back from their insertions just reaching posterior head margin; eyes of moderate size with about 10-11 ommatidia in longest row; cephalic surface between frontal lobes faintly striated whereas in some individuals the striations are absent; promesonotum and mesonotum forming continuous flat line in profile; mesosoma with single pair of standing hairs on pronotal humeral angles; metanotal groove feebly impressed; petiole with single pair of backward directed hairs; postpetiole with two pairs. However, the eyes are slightly smaller in M. abeillei, and the central cephalic sculpture is feebly microreticulate-striolate than in the relatively large-eyed M. wahibiense and the superficially sculptured cephalic surface but we consider these two traits as variable characters. Herein, we propose treating M. wahibiense as a junior synonym of M. abeillei André.

Remarks.
A thorough examination of the type material of M. fezzanense, M. hemame, M. marmule, and M. areniphilum yielded no evidence for heterospecificity; they are indistinguishable. All four taxa share the following characters: median portion of anterior clypeal margin shallowly concave; eyes large with 12-15 ommatidia in longest row; promesonotum and anterior portion of mesonotum in profile feebly convex; posterior portion of mesonotum sloping steeply to broadly and deeply impressed metanotal groove; mesosoma without hairs; petiole with single pair of backward directed hairs; postpetiole with two pairs of hairs.
Herein M. fezzanense, M. hemame, and M. marmule are treated as junior synonyms of M. areniphilum. It is worth mentioning that in the original description of M. marmule,  gave a brief differential diagnosis with M. areniphilum based on variable characters such as the presence of mesosomal pubescence, the petiole and postpetiole color and pilosity.
Geographic Distribution. A species originally described from Tunisia and recorded from most countries of the Arabian Peninsula including KSA, Kuwait, Oman, and Yemen (Collingwood, 1985;, and the UAE (Collingwood et al., 2011). It is also reported from North Africa and the Afrotropical Region (Bolton, 1987).  Geographic Distribution. This species was originally described from Kazakhstan and recorded from Oman (Collingwood, 1985;, the UAE (Collingwood et al., 2011), Turkey (Kiran & Karaman, 2020), Israel (Vonshak & Ionescu-Hirsch, 2009). The present material represents a new record to the KSA.
Remarks. Monomorium barbatulum looks similar to some members of the genus Trichomyrmex in terms of the following characters: polymorphic with 12-segmented antennae that lacking the well-defined terminal club; masticatory margin of mandibles armed with 3-4 teeth; propodeum unarmed. However, M. barbatulum lacks a critical diagnostic character for the genus Trichomyrmex which is the absence of the transverse striations on propodeal dorsum. With the availability of more material for a comprehensive taxonomic investigation, together with additional molecular evidence(s), we will be able to resolve properly the taxonomic status of the species. Remarks. The type material of M. bicolor and M. phoenicum are clearly conspecific. They share the same diagnostic characters as follow: scapes relatively long, when laid back from their insertions surpassing posterior head margin by about length of pedicel; head in full-face view with eyes just breaking sides; cephalic surface dull, finely and densely punctate; median anterior clypeal margin distinctly concave; area between frontal carinae finely longitudinally striated; mesosoma without standing hairs; metanotal groove acutely impressed; propodeal dorsum in profile meeting declivity in continuous curve; petiole and postpetiole each with single pair of back directed hairs; first gastral tergite with hairs scattered over tergite surface; biocolored species, with head, mesosoma, petiole and postpetiole yellow-red or yellow-brown, gaster dark brown to black. Herein, we propose M. phoenicum as a junior synonym of M. bicolor.
Geographic Distribution. Monomorium bicolor was originally described from Eritrea and is a widespread species commonly encountered in open, sandy areas through the Afrotropical Region (Bolton, 1987). In the Arabian Peninsula, it is known from the KSA and the UAE (Collingwood, 1985;Collingwood et al., 2011). Monomorium brunneolucidulum Monomorium brunneolucidulum Collingwood & Agosti, 1996 Remarks. In their brief original description of the enigmatic species M. brunneolucidulum from Oman,  neither gave successful diagnostic characters nor illustrations for species recognition. In addition, the type-material is apparently lost. Due to a lack of type material and species diagnostic characters, it is impossible to confirm the identity of the species. Until the type material of this species is available we prefer to treat it as a nomen dubium. Geographic Distribution. This species was originally described from India. It is a successful tramp species of putative Southeast Asian origin that is widely distributed throughout tropical and subtropical regions worldwide (Deyrup, Davis & Cover, 2000;Wetterer, 2010a KSA: Riyadh (24.714, 46.675, 21.i.1980, A.H. Talhouk, 2w); Yemen: Taiz (13.578, 44.018, 20.iii.1993, C.A. Collingwood, 2w).
Remarks. Monomorium harithe was described from the KSA and Yemen, while M. najrane was described from Najran (KSA) near the Saudi-Yemeni borders . The comparison of the type material of the two species reveals a straightforward synonymy. The two species share the following characters: anterior median clypeal margin distinctly concave; scapes distinctly short, when laid back from their insertions failing to reach posterior head margin; mesosoma without standing hairs; metanotal groove feebly impressed but distinct; propodeal dorsum with distinct furrow; mesosoma, petiole and postpetiole finely and densely punctate; petiole and postpetiole each with single pair of back directed hairs; gaster smooth and shining.
Geographic Distribution. This Arabian endemic species is only known from the KSA and Yemen . Diagnosis. Monomorium heggyi is diagnosed by the following character combination: scapes when laid back from their insertions just reaching posterior margin of head; head in full-face view with eyes located nearly at midlength; promesonotal outline feebly convex or flat, sloping posteriorly to narrow and shallowly impressed metanotal groove.

Monomorium heggyi
Worker. Head. Head in full-face view distinctly longer than broad, with concave posterior margin and shallowly convex sides and feebly concave posterior margin; median portion of clypeus with anterior free margin slightly indented; eyes of moderate size, in profile with convex dorsal sides and straight ventral side, maximum diameter 0.21 x-0.30 x HW, with 7 ommatidia in longest row; head in full-face view eyes located nearly at midlength of head and just breaking sides; scapes when laid back from their insertions just reaching posterior head margin. Mesosoma. Promesonotal outline feebly convex or flat, slopping posteriorly to narrow and shallowly impressed metanotal groove; propodeal dorsum in profile convex making a continuous curve with propodeal declivity and with defined lateral margins. Petiole. Petiole with high rounded node in profile; subpetiolar process broad and blunt. Postpetiole. Postpetiolar node lower than petiolar node in profile and nearly as broad as petiole in dorsal view. Sculpture. Mandibles feebly longitudinally sculptured; cephalic surface with faint vestiges of superficial reticular patterning, almost entirely effaced, area between frontal carinae finely longitudinally striate; clypeus smooth; entire mesosoma, petiole and postpetiole sharply and densely reticulate-punctate; gastral tergites smooth and shining. Pilosity. Dorsum of head without standing hairs behind the level of the frontal lobes; several pairs of long hairs on the anterior clypeal margin and on mandibles; antennae with dense appressed pubescence; mesosoma, petiole and postpetiole without standing hairs of any description; first gastral tergite without standing hairs except for sparse appressed pubescence; pilosity of remaining gastral tergites restricted to the posterior margins. Color. Uniformly yellow.
Remarks. Monomorium heggyi belongs to the M. salomonis species-group (Bolton, 1987). It is most similar to M. rabirium Bolton, 1987 from Botswana from which it is readily distinguished by the longer scapes (SI 80-108) that reach the posterior head margin in full-face view, the posteriorly shifted eyes located nearly at the midlength of head in full-face view. Monomorium rabirium has shorter scapes (SI 92-97) that fail reaching the posterior head margin in full-face view, and eyes conspicuously located in front of midlength of head in full-face view. Among the Arabian species of the M. salomonis species-group, M. heggyi is superficially similar to M. elghazalyi from the Socotra Archipelago from which it can be easily separated by the larger eyes (EI 21-31), the shallowly impressed metanotal groove, and the densely sculptured mesosoma, petiole and postpetiole. Monomorium elghazalyi has smaller eyes (EI 19-20), broadly and deeply impressed metanotal groove, and a smooth body surface.
Etymology. The patronymic name honors Dr. Essam Heggy, the Egyptian space scientist at NASA.
Habitat. The type locality, Shada Al A'la, is a Nature Reserve (Fig. 17)  Diagnosis. Monomorium khalidi can be distinguished by the combination of the following characters: short scape failing to reach posterior head margin in full-face view; abundant mesosomal pilosity; straight outline of promesonotum; densely reticulatepunctate surfaces of head, mesosoma, petiole, and postpetiole; promesonotum dorsally with at least five to six pair of hairs, promesonotum and propodeum each with three pairs. Worker. Head. Head nearly as long as broad, or little longer than broad with concave posterior margin and feebly convex sides; median portion of clypeus with anterior free margin distinctly concave; eyes of moderate size, in profile view with convex dorsal sides and straight ventral side, maximum diameter 0.20 × HW, with 9-10 ommatidia in longest row); head in full-face view with eyes failing to break head sides; scapes when laid back from their insertions failing to reach posterior margin. Mesosoma. Promesonotal outline flat, slopping posteriorly to narrow and feebly impressed metanotal groove; propodeal dorsum flat and short, longitudinally concave, with sharply defined lateral margins. Petiole. Petiole with high rounded node in profile. Postpetiole. Postpetiole as broad as petiole in dorsal view. Sculpture. Cephalic surface between and immediately behind frontal lobes finely longitudinally striate; cephalic surface and sides, entire mesosoma, petiole and postpetiole sharply and densely reticulate-punctate; first gastral tergite shagreened and relatively shining. Pilosity. Cephalic surface with several pairs of standing hairs behind level of frontal lobes; posterior margin of head with three pairs of standing hairs; underside of head with about five pairs of short hairs; promesonotum dorsally with at least five to six pair of hairs, promesonotum and propodeum each with three pairs; petiole and postpetiole each with two-three pairs of backward directed hairs; first gastral tergite with numerous standing hairs which are evenly distributed over the sclerite in front of the apical transverse row. Color. Bicolored species, head, mesosoma, petiole, postpetiole and appendage light red brown, gaster black.
Remarks. This new species is a member of the M. salomonis species-group (Bolton, 1987). Monomorium khalidi is closest to M. junodi Forel, 1910 from South Africa in terms of the relatively small eyes, the short scapes that fail to reach posterior margin of head, the acute metanotal groove, and densely punctate body surface. The two species have short scapes that fail to reach posterior margin of head in full-face view, metanotal groove feebly impressed; body surface densely punctate; and propodeal longitudinally concave, with sharply defined lateral margins. However, M. khalidi can be easily distinguished from M. junodi by the following characters: body bicolored with head, mesosoma, petiole, postpetiole and appendages light red-brown contrasting with the black gaster; head in full-face view with eyes failing to break sides; promesonotum dorsally with at least five to six pair of hairs, promesonotum and propodeum each with three pairs; promesonotal outline flat. Monomorium junodi is uniformly brown to dark brown, head in full-face view with eyes breaking sides; promesonotum dorsally with two pair of hairs, propodeum without hairs; promesonotal outline feebly but distinctly convex. Among the Arabian Monomorium species, M. khalidi is superficially similar to M. nitidiventre in terms of body size, surface sculpture, eye shape but the former can be readily recognized by the reduced stiff pilosity.
Etymology. The patronymic name honors Khalid Amr (born at 04/11/2012), the son of the second author.
Habitat. The type locality of M. khalidi is Shada Al A'la ( Fig. 19), the same locality where M. heggyi was collected.
Ecological and Biological notes. The broad geographic distribution of the species can be interpreted in the light of the species diverse habitat preferences, including the deserts, mountainous, and cultivated sites. Several worker series were found nesting in either dry or humid soil beneath rocks in an undisturbed site in the KSA where a broad diverse of plant species exists including Acacia ( Remarks. The synonymy of M. yemene with M. nitidiventre is straightforward since both are morphologically similar and indistinguishable. Both share the following key characters: posterior margin emarginated in full-face view; median portion of anterior clypeal margin distinctly concave; metanotal groove deeply impressed; head, mesosoma, petiole, and postpetiole densely reticulate-punctate and covered with abundant pale standing hairs. Note: However, the locality label of the paratype specimen (CASENT0913865) (Madinat Al shiraq) is not matching the locality data mentioned in the original description but the collecting data and collector are congruous with the description, therefore, this specimen is treated as one of the type material of M. yemene.

DISCUSSION
Monomorium is one of the most diverse ant genera in the world, but it is rarely the most speciose genus on a regional scale. However, the Arabian Monomorium fauna, based on review of previous literature data and on our current work, includes 44 species, making it the most diverse known ant genus of the Arabian Peninsula (Collingwood, 1985;Collingwood et al., 2011;Sharaf & Aldawood, 2013b;Sharaf et al., 2015Sharaf et al., , 2017aSharaf et al., , 2018aSharaf et al., , 2018b. Currently, the genus represents 14% of the total number of species reported from the region (312 spp.) (Collingwood, 1985;Collingwood et al., 2011;Sharaf & Aldawood, 2013cSharaf et al., 2015Sharaf et al., , 2017aSharaf et al., , 2017bSharaf et al., , 2018aSharaf et al., , 2018bSharaf, Aldawood & Hita Garcia, 2019;Sharaf et al., 2020c). This value is lower than the 20% mentioned by . This reduction is the result of numerous taxa in previous studies now treated as junior synonyms of other species (Sharaf et al., 2015(Sharaf et al., , 2017a(Sharaf et al., , 2018a; this study).
Biogeographically, the biota of the Arabian Peninsula does not constitute a cohesive unit (Larsen, 1984;Sharaf et al., 2020aSharaf et al., , 2020b. Instead, the Arabian Peninsula is often considered to be at the nexus of two terrestrial biogeographic realms, the Palearctic and the Afrotropic. In fact, Olson et al. (2001) places the northern and central Arabian Peninsula in the Palearctic bioregion (along with Europe, northern Africa, Asia north of the Himalayas, and neighboring islands) and the southern and eastern coasts of the Arabian Peninsula in the Afrotropic bioregion (along with sub-Saharan Africa, southern Iran, southwestern Pakistan, and neighboring islands). Our Geographic distribution data (see also Table 1) of Arabian Monomorium supports this basic division, with 12 Palearctic species more common in the north and central deserts and the five Afrotropic species more common in the southern region and along the coasts. The majority of the endemic Arabian ant species has faunal similarities with taxa from the Afrotropic bioregion that has been earlier documented by several studies (Larsen, 1984;Collingwood, 1985;Sharaf, Aldawood & Taylor, 2012;Sharaf, Aldawood & El-Hawagry, 2012a, 2012bSharaf et al., 2020aSharaf et al., , 2020bEl-Hawagry et al., 2013Hájek & Reiter, 2014). Therefore, it is not surprising that a large proportion of the endemic Arabian Monomorium species (13/24) have been found in the mountainous ranges of southwestern KSA that extend to Yemen.  Both new Monomorium species reported here, M. heggyi and M. khalidi, were collected in the Shada Al-A'Ala Nature Reserve (SANR), a protected area consisting of an isolated granite mountain massif in southwestern Saudi Arabia. Its location, elevational range (470-2,222 m) and high rainfall resulting in diverse microclimates and a high biodiversity (SWA, 2020). As a unique biodiversity hotspot, the SANR contains~495 plant species (~22% of the total reported Saudi Arabian flora), including 43% of the threatened plant species and 19 endemic plants (Thomas, El-Sheikh & Alatar, 2017). The SANR also protects a diverse fauna, including rare and endemic vertebrates, including the griffon vulture (Gyps fulvus (Hablitz)), the Arabian leopard (Panthera pardus nimr (Hemprich and Ehrenberg)), and the Arabian wolf (Canis lupus arabs Pocock) (SWA, 2020). The SANR invertebrate fauna has attracted relatively little attention, but recent insect biodiversity inventories and monitoring research projects conducted by King Saud University Museum of Arthropods resulted in two important faunistic studies that recorded 119 Diptera species (El-Hawagry et al., 2016) and 62 carabid beetle species (Abdel-Dayem et al., 2019). Further studies are planned to be carried out at SANR to explore additional levels of biodiversity.
In addition to the native Monomorium species, there are two Monomorium known from the Arabian Peninsula that are cosmopolitan tramp species, spread around the world through human commerce: M. pharaonis and M. floricola. The pharaoh ant, M. pharaonis, is a common domestic pest. Although it was first described from Egypt, its original native range is uncertain (Wetterer, 2010b). We report the first known Arabian record of M. floricola, an Indomalayan species, from a single site in Oman. Although widespread around the world, M. floricola is rarely considered a serious pest. However, because this species is very small, slow moving, cryptically colored, and primarily arboreal, its abundance and ecological importance may be underappreciated (Wetterer, 2010a). Human activities, population movements, and global trade around the world greatly contribute to the further spread of tramp species outside their native habitats (Sharaf et al., 2020c(Sharaf et al., , 2020d.
Considering the high degree of endemism encountered, it is likely that the known Arabian Monomorium fauna will increase in the future with further exploration of poorly surveyed areas of the Arabian Peninsula, especially the southwestern mountains of the KSA, Yemen, and the mountainous regions of Oman and the UAE. We hope that the present study will serve as a cornerstone of future taxonomic treatments of Monomorium in the Arabian Peninsula.
Brendon E. Boudinot performed the experiments, analyzed the data, authored or reviewed drafts of the paper, and approved the final draft. James K. Wetterer performed the experiments, analyzed the data, prepared figures and/ or tables, authored or reviewed drafts of the paper, and approved the final draft. Francisco Hita Garcia conceived and designed the experiments, performed the experiments, analyzed the data, prepared figures and/or tables, authored or reviewed drafts of the paper, and approved the final draft. Hathal M. Al Dhafer performed the experiments, authored or reviewed drafts of the paper, and approved the final draft. Abdulrahman S. Aldawood conceived and designed the experiments, performed the experiments, analyzed the data, authored or reviewed drafts of the paper, and approved the final draft.

Data Availability
The following information was supplied regarding data availability: All data supporting the work results are presented in the in the figures and Materials and Methods and Results sections.