Anochetus (Hymenoptera: Formicidae) in the Arabian Peninsula, with Description of a New Species from Oman

PROC. ENTOMOL. SOC. WASH. 119(1), 2017, pp. 78–89

ANOCHETUS (HYMENOPTERA: FORMICIDAE) IN THE ARABIAN PENINSULA, WITH DESCRIPTION OF A NEW SPECIES FROM OMAN

urn:lsid:zoobank.org:pub:4D887E81-F8AC-4899-83CE-883DBD3755DA

MOSTAFA R. SHARAF,JOE MONKS,ABDULRAHMAN S. ALDAWOOD, AND ANDREW POLASZEK

(MRS, ASA) Plant Protection Department, College of Food and Agriculture Sciences, King Saud University, Riyadh 11451, P. O. Box 2460, Kingdom of Saudi Arabia (corresponding author e-mail: [email protected]); (JM) School of Life Sciences, the University of Nottingham, Nottingham, U.K.; (AP) Department of Life Sciences, Natural History Museum, London SW7 5BD U.K.

(MRS) urn:lsid:zoobank.org:author:E2A42091-0680-4A5F-A28A-2AA4D2111BF3 (JM) urn:lsid:zoobank.org:author:D5FC6B75-1909-4DF8-A953-1D6363270C2D (ASA) urn:lsid:zoobank.org:author:477070A0-365F-4374-A48D-1C62F6BC15D1 (AP) urn:lsid:zoobank.org:author:AFDEB328-39B5-4902-A907-26603BCC99D6

Abstract.—Anochetus Mayr, 1861 is revised for the Arabian Peninsula based on the worker caste. Three species are recognized, Anochetus annetteae n. sp., A. sedilloti Emery, 1884, and A. traegaordhi Mayr, 1904. Anochetus annetteae n. sp. is described from Hibra Village, the Nakhl Region, Sultanate of Oman, based on the worker caste. An illustrated key to the Arabian species of Anochetus is given, including a distribution map for the three Arabian species. New distribution records for A. traegaordhi for the Kingdom of Saudi Arabia are also presented. Key Words: Ponerinae, Palaearctic Region, Middle East, date palm, revision, Saudi Arabia, Yemen DOI: 10.4289/0013-8797.119.1.78

The specialized trap mandibles and the constructed in soil, in rotten wood or shape of the head make the ant genus twigs, in leaf litter, in crevices of bark, Anochetus Mayr one of the more distinctive under logs, in interstices of palm leaf- genera in the subfamily Ponerinae. The bases, or in termite nests (Brown 1978, genus contains 115 described extant species Lattke 1986, Shattuck and Slipinska and eight described fossil species (Bolton 2012). These nests frequently contain low 2014, www.antwiki.org, accessed 17 May numbers of workers, usually fewer than 2016) distributed worldwide in the tropical 100 (Brown 1978). Their feeding habits and subtropical regions (Brown 2000). are also variable (Brown 1978). Most The nesting habits of Anochetus spe- species are predators of other smaller cies are diverse (Brown 1978). Nests are terrestrial arthropods (Shattuck 1999), VOLUME 119, NUMBER 1 79 some are specialized predators on ter- species, A. boltoni and A. goodmani. mites (e.g., A. traegaordhi Mayr, 1904) This revision was the first treatment to (Schatz et al. 1999), and a few species combine both morphological and molec- feed on seeds or plant debris (e.g., A. ular data. Fernandez (2008) presented renatae Shattuck and Slipinska, 2012, a key to the Neotropical species. Shattuck (Heterick 2009)). Often relatively few and Slipinska (2012) revised the Australian specimens are available for study because Anochetus fauna keying 14 species in- of the cryptic nature of these ponerine ants cluding eight new species. Recently, and therefore little information on biology Schmidt and Shattuck (2014) provided a list and ecology is available (Brown 1978). of the world fauna, notes on global distri- The worker caste is very similar to butions, ecology, behavior, and phylogeny. Odontomachus Latreille, 1804 in head De Andrade (1994) revised and keyed the shape and mandibular structure. The Anochetus fossils known from Dominican mandibles of both genera are elongate and amber, describing six new species. linear, situated midlength on either side of The first recorded Anochetus species the head and armed with two or three from the Arabian Peninsula were A. vertical teeth (Fisher and Smith, 2008). sedilloti Emery, 1884 and A. traegaordhi Anochetus can be readily separated from Mayr, 1904 (Collingwood 1985) from Odontomachus by the petiole structure. Fayfa Region (Jazan Province, the Workers of Anochetus have two or three southwestern region of the Kingdom of large teeth arranged vertically, whereas Saudi Arabia (KSA)). The above men- Odontomachus has a single petiolar tooth tioned records were based on only three or spine. For additional discussion see specimens, two A. traegaordhi and one Brown (1978), Bolton (1994), and Fisher A. sedilloti. Later, A. traegaordhi was and Smith (2008). recorded from Yemen by Collingwood In addition to the above petiolar char- and van Harten (2001). No additional acter, Anochetus can be recognized in the material was collected from the region worker caste from other ponerine genera by since the Yemeni record. the combination of the following morpho- In the present study, the Arabian species logical characters (Brown 1978, Bolton of Anochetus are reviewed. The genus is 1994): Nuchal carina forming a broad, recorded for the first time from Oman with uninterrupted U-shaped ridge across the the new species, A. annetteae sp. n. posterodorsal extremity of the head; the V-shaped apophyseal lines absent; petiolar MATERIAL AND METHODS node either rounded or with two or three teeth. Measurements and indices The taxonomic history of the genus Anochetus is extensive. New synony- All measurements and indices are in mies and notes on variation within the millimetres, and follow the standard mea- genus were given by Brown (1964). The surements of Brown (1976, 1978) and sole global revision of Anochetus (Brown, Kugler and Ionescu (2007). Digital colour 1978) treats the species of most zoogeo- images of lateral and dorsal views of the graphical regions, and includes notes on entire body and full-face views of the head species groups, biology, ecology, and of each species were created using a Leica distribution. In their revisionary work DFC450 digital camera with a Leica Z16 of the Madagascar fauna, Fisher and APO microscope and LAS (v3.8) soft- Smith (2008) recognized five species ware. These images are also available from the region, and described two new onlineonAntWeb(www.antweb.org)and 80 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON are accessible using the unique identifying specimen code. The species names follow the online catalogue of ants of the world (Bolton 2014) available on www.AntCat. org. Distribution maps were made using DIVA-GIS (version 7.5.0.0).

Measurements (Figs. 1, 2) EL Eye length, maximum diameter of eye in full-face view. HL Head length, maximum length of head in dorsal view, measured in a straight line from mid-point of anterior clypeal margin to mid- point of level of posterior lobes. HW Head width, maximum width of head behind eyes measured in full-face view. MDL Mandible length, straight line length of mandible from apex to anterior clypeal margin, measured in dorsal view. ML Mesosoma length, diagonal length of mesosoma in proﬁle from point at which pronotum meets cervical shield to posterior base of metapleuron. PNH Petiolar node height, maximum Figs. 1–2. Morphological measurements used in Anochetus. 1, head in full-face view. 2, body in proﬁle. height of petiolar node measured in lateral view from highest point Indices: to outline of ventral lobe. PNL Petiolar node length, measured as CI Cephalic index (HW3100/HL). maximum thickness in lateral view. EI Eye index (EL3100/HW). PNW Petiolar node width, measured in MDI Mandibles index (MDL3100/HL). dorsal view and excluding ante- PNI Petiolar node index (PNH3100/ rior and posterior peduncles. PNL). PW Pronotum width, maximum width SI Scape index (SL3100/HW). of pronotum measured in dorsal view. Abbreviations of Museums following SL Scape length, maximum straight Lattke (2000): line length of antennal scape excluding basal constriction or neck BMNH Natural History Museum (British to condylar bulb. Museum, Natural History), TL Total length, sum of lengths of London, U.K. mandible, head, mesosoma, peti- KSMA King Saud University Museum ole and gaster. of Arthropods, Plant Protection VOLUME 119, NUMBER 1 81

Department, College of Food and (M.R. Sharaf) (MRS0311). Paratypes: 2 Agriculture Sciences, King Saud workers with same data as the holotype. University, Riyadh, Kingdom of Type depository.—The holotype is Saudi Arabia. deposited in KSMA, 1 paratype speci- MSNG Museo Civico di Storia Naturale men in BMNH and 1 in ONHM. Giacomo Doria, Genoa, Italy. Holotype worker.—Measurements: ONHM Oman Natural History Museum, EL 0.12, HL 0.92, HW 0.85, MDL 0.47, Muscat, Sultanate of Oman. ML 1.17, PNH 0.45, PNL 0.17, PNW 0.27, PW 0.50, SL 0.67, TL 4.20; Indices: RESULTS CI 92, EI 14, MDI 51, PNI 265, SI 79. Paratype workers.—Measurements: KEY TO ARABIAN SPECIES OF ANOCHETUS EL 0.10–0.12, HL 0.90–1.15, HW 0.82, BASED ON WORKER CASTE MDL 0.50–0.52, ML 1.10–1.20, PNH 1. Small species (TL 3.35–4.20, HL 0.90– 0.45–0.47, PNL 0.15, PNW 0.25–0.27, 1.15, HW 0.82–0.85); eyes small with six PW 0.50, SL 0.72–0.82, TL 3.35–3.57; to eight ommatidia in the longest row (EI Indices: CI 71–91, EI 12–15, MDI 45– 12–15) (Fig. 3) ...... annetteae n. sp. 56, PNI 300–313, SI 88–100 (n=2). – Larger species (TL 5.00–7.50, HL 1.46– Worker.—Head: Subrectangular in 1.75, HW 1.25–1.50); eyes larger with full-face view; posterior margin strongly more than 20 ommatidia in the longest row concave; lateral margins of head feebly (EI 17–25) (Fig. 4) ...... 2 concave and broad at eye level; scapes 2. Head, mandibles, antennae and legs orange, relatively short (SI 79–100), when laid mesosoma and petiole red, gaster dark brown; back from their insertions failing to eyes large (EI 25); scapes short (SI 79) not reach posterior margin of head; second surpassing the posterior margin of head; in funicular segment as long as broad; re- profile, petiole with broadly rounded dorsum maining funicular segments distinctly (Fig. 5); mesosoma, mesopleuron, meta- longer than broad; eyes small (EI 12–15) pleuron and petiole finely and irregularly with six to eight ommatidia in longest costulate (Fig. 5) ...... sedilloti Emery row; mandibles long, broader apically, – Body uniformly yellow, gastral tergites with with three teeth, the middle tooth half yellow tint; eyes small (EI 17–22); scapes the length of the other two teeth; ventral distinctly longer (SI 100–116) surpassing the mandibular margin when seen in lateral posterior margin of head; in profile, petiole view with six blunt denticles; concealed with a narrowly rounded dorsum (Fig. 6); by dorsal internal mandibular margin in mesosoma faintly and irregularly rugulose; full-face view. Mesosoma:Inprofile mesopleuron, metapleuron and petiole smooth with a flat dorsal outline, and with (Fig. 6) ...... traegaordhi Mayr shallow but distinct promesonotal suture and metanotal groove; promesonotal Anochetus annetteae Sharaf, new suture appears as transverse line in species dorsal view; propodeal dorsum meets declivity in a right angle; propodeal urn:lsid:zoobank.org:act:99F4FB10- spiracle circular, situated above mid- 513A-41CF-80AF-AECF1CA5D9FE line of metapleuron in profile. Peti- (Figs. 7-9) ole: Triangular in profile, with sharp Holotype worker.—Sultanate of Oman, pointed dorsum; subpetiolar process Hibra village, Nakhl Region, 05.iv. well-developed and rounded ventrally; 2016, 23.49327°N, 57.83421°E, 190 m, petiole in profile as high as propodeum. 82 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 3– 6. Head and body structures of Anochetus spp. 3, head of Anochetus annettae n. sp. showing small eyes. 4, head of A. sedilloti showing large eyes. 5, body proﬁle of A. sedilloti showing mesosomal sculpture and petiole. 6, body proﬁle of A. traegaordhi showing mesosomal sculpture and petiole.

Sculpture: Parallel rugae between Colour: Head, mesopleuron, lower frontal carinae running mesad behind half of metapleuron, petiole, and legs midlength of head, remaining cephalic yellow, mesosomal dorsum, propleuron surface and mandibles smooth and and upper half of metapleuron yellow shining; pronotal and mesonotal dorsa, brown; first and second gastral tergites and propleura irregularly rugulose; pro- dark brown, remaining of gaster yellow podeal dorsum reticulate-rugose; pro- brown. podeal declivity in dorsocaudal view Diagnosis.—Anochetus annetteae is with about 16 transverse rugae; meso- a member of the A. graeffei-group as pleuron distinctly smooth and shining; defined by Brown (1978). The new spe- metapleuron transversally striate; petiole cies cannot be identified with available nearly smooth and shining; gaster super- keys to the Anochetus species of Asia ficially shagreened, dull with few smooth (Brown 1978), the Arabian Peninsula and shining small areas. Pilosity: Cephalic (Collingwood and Agosti 1996), the surface, antennae, and mandibles with Mediterranean and the Middle East appressed pubescence; mesosomal dor- (Kugler and Ionescu 2007), the Indian sum and petiole with abundant, short, (Bharti and Wachkoo 2013), and the suberect hairs; gaster with abundant hairs Madagascan (Fisher and Smith 2008) which are longer than mesosoma hairs. regions. Anochetus annetteae appears VOLUME 119, NUMBER 1 83

view; the irregular and weaker pronotal and mesonotal sculpture. Anochetus yerburyi has a red-brown mesosoma, dark brown gaster, and yellow head and petiole; the eyes are distinctly larger (EI 21) with about 16 ommatidia in the longest row; the scapes longer when laid back from their insertions, reaching the posterior margin of the head; the pronotal sculpture is regularly circular. Etymology.—The patronym annetteae honors Dr. Annette Patzelt, the Scientiﬁc Director of the Oman Botanic Garden in Muscat. Habitat and Biology.—The type locality (Fig. 10) of the new species is a large date palm plantation (Phoenix dactylifera L.). A single specimen was found in moist soil under the base of a date palm tree and collected by digging around a palm tree where an unidentiﬁed species of termite was also observed nesting. The two paratype specimens were found foraging on the ground and collected by leaf litter sifting.

Anochetus sedilloti Emery, 1884 (Figs. 4, 5; 11-13) Anochetus sedilloti Emery 1884: 377, ﬁg. (w.) Tunisia. Palearctic. Syntype worker. Tunisia, Gabes Gafsa road, casent0903976 (MSNG) [image examined]. Forel 1907: 202 (m.); Santschi 1907: 325 (m.). Senior synonym of Figs. 7–9. Anochetus annettae n. sp. 7, body Anochetus indicus Brown 1978: 559 (see also in proﬁle. 8, body in dorsal view. 9, Head in full- p. 594). face view (Michele Esposito, photographer, casent0919949: www.AntWeb.org). Measurements.—EL 0.32, HL 1.46, HW 1.27, MDL 0.76, ML 1.66, PNH 0.55, PNL 0.33, PNW 0.40, PW 0.73, SL 1.00, TL most similar to A. yerburyi Forel, 1900, 5.00; Indices: CI 87, EI 25, MDI 52, PNI described from Sri Lanka, but is easily 167, SI 79. (casent0907413). distinguished by the paler color, the Worker.—Head: Posterior margin of smaller eyes (EI 12–15, with six to eight head deeply emarginate; eyes rounded, ommatidia in the longest row); the relatively large (EI 25) with about 25 shorter scapes that fail to reach the ommatidia in the longest row; scapes posterior margin of the head in full-face when laid back from their insertions just 84 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 10. Type locality of Anochetus annettae n. sp. (M. Sharaf, photographer). reaching median part of posterior margin characterized by moderate body size, of head. large eyes, subtruncate or broadly Mesosoma: Promesonotum flat, slop- rounded petiolar nodes, well-developed ing back to feeble metanotal groove; body sculpture, and pubescence which propodeal dorsum about 1.5 3 of de- either is decumbent or appressed. clivity length, both making an obtuse Previous records.—KSA, Jazan angle in profile. Petiole: With a rounded Province, Fayfa, 17.266667°N, 43.1°E, node and higher than propodeum in 28.iii.1983, (C. A. Collingwood). profile. Sculpture: Cephalic surface and Distribution.—Tunisia (Emery 1884), mandibles smooth and shining, with area Circum-Saharan Africa, Kenya, Eritrea, between frontal carinae finely striate; Senegal, India (Brown 1978), KSA, North mesosoma and petiole finely and irreg- Africa (Collingwood and Agosti 1996). ularly costulate; gaster smooth and Comment: This species was not collected shining. Pilosity: Cephalic surface with during the present study and the species abundant appressed pubescence; meso- diagnosis given here is based on an image soma bare except for three to four pairs of the syntype worker (casent0903976). of setae on promesonotum; first gastral tergite bare, remaining gastral tergites with few scattered long setae; entire Anochetus traegaordhi Mayr, 1904 gastral surface with abundant appressed (Figs. 6; 14-16) pubescence. Colour: Head, mandibles, antennae, and legs orange, mesosoma Anochetus traegaordhi Mayr 1904: 2 (w.) and petiole red, gaster dark brown. Sudan. Afrotropical. Type not located. An Diagnosis.—Anochetus sedilloti is a image of a syntype worker of A. gracili- member of the A. sedilloti species cornis, (CASENT0915168), a junior syno- group as defined by Brown (1978). It is nym of A. traegaordhi was examined. This VOLUME 119, NUMBER 1 85

Worker.—Head: Posterior margin of head strongly concave; eyes of moderate size (EI 17–22); scapes short, fail to reach posterior margin of head. Meso- soma: Promesonotum feebly convex in profile; metanotal groove impressed; propodeal dorsum about 2.5 3 longer than declivity; propodeal outline ob- tusely angulate. Petiole: In profile, anterior face broadly concave, posterior face feebly convex; petiolar node narrowly rounded. Sculpture: Cephalic surface smooth and shining except for fine striations between frontal carinae, running posteriorly and outwards to level of posterior margin of eyes; mesosoma faintly and irregularly rugulose; mesopleuron smooth; petiole and gaster smooth and shining. Pilosity: Cephalic surface without hairs or pubescence, anterior clypeal margin with few simple hairs, mandibles and antennae with appressed pubescence, mesosoma with scattered short hairs (hair length 0.08), gaster with scattered hairs. Color: Uni- form yellow, gastral tergites yellow with Figs. 11–13. Anochetus sedilloti, syntype brown tint. worker. 11, Body in profile. 12, body in dorsal view. Diagnosis.—Anochetus traegaordhi 13, head in full-face view (Zach Lieberman, pho- is a member of the A. ghilianii-group as tographer, casent0903976: www.AntWeb.org). mentioned by Brown (1978). The A. ghilianii-group can be recognized by the medium or large body and eye sizes, the specimen has the following data: Renk, short cephalic longitudinal striations that Aeg. Sudan, Dr. F. Santschi. Senior syno- fail to reach nuchal carina, the smooth nym of A. silvaticus: Brown 1964: 215; of and shining cephalic surface, and the A. angusticornis, (and the junior synonym thick, upright, compressed petiolar node A. sudanicus): Brown 1978:559. as seen from profile. Worker.—Measurements: EL 0.25– Material examined.—KSA, Al Bahah 0.30, HL 1.50–1.75, HW 1.25–1.50, Province, Al Mukhwah, Thi Ayn Ar- MDL 0.90–1.05, ML 2.00–2.25, PNH chaeological Village, 19.929583°N, 0.65–0.95, PNL 0.30–0.40, PNW 0.25– 41.442861°E, alt. 744 m, 20. ix. 2011, 0.40, PW 0.60–0.80, SL 1.45–1.60, TL (4w) (M. R. Sharaf); KSA, Allaith 5.35–7.50; Indices: CI 77–93, EI 17–22, gov., Adam, Wadi Elarj, 20.45296°N, MDI 57–63, PNI 157–317, SI 100–116 40.81555°E, alt. 450 m, 9.xi.2012, (n=9). The specimens used for measure- (4w) (M. R. Sharaf); KSA, Jazan ments were recent collections from the Province, Wadi Aljora, near Abadan, KSA. 17.29263°N, 43.07010°E, alt. 465 m, 86 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

12.xi.2012, (1w) (M.R.Sharaf.) (KSMA). Previous records.—KSA, Jazan Prov- ince, Fayfa, 17.28797°N, 43.14434°E, 30. iii.1983, (C.A.Collingwood)(Colling- wood 1985); Yemen, Khamis Bani Sa’ad, 15.184722°N, 43.51°E, 23.iv.1999, 28. iii.2000, (A. van Harten) (Collingwood and van Harten 2001). Distribution.—Sudan, Eritrea, (Brown 1978), Kenya, Mozambique, South Africa, Tanzania, Zimbabwe (www. antweb.org, accessed 11.05.2016), Saudi Arabia (Collingwood and Agosti 1996), Yemen (Collingwood and van Harten 2001). Habitat and Biology.—This species was collected nesting in soil that was moist and rich in organic matter. A nest series was found also under a large Ficus tree (Moraceae) in Thi Ayn Archeological Village (Al Bahah Prov- ince) collected by digging. Some workers were found foraging in leaf litter at Wadi Aljora.

DISCUSSION Anochetus is considered a rare ant genus in the Arabian Peninsula. Rela- tively few specimens have been collected or are available in collections from the region. No doubt this is due to the well-known cryptic nature of the genus (Brown 1978, Deyrup 2002, Torres et al. 2000) and often low number of individuals per nests (Brown 1978, Shattuck and Slipinska 2012). Date palm plantations in the type locality of A. annetteae provide a cool shaded microclimate supplied with Figs. 14–16. Anochetus traegaordhi worker. regular water from the traditional 14, body in proﬁle. 15, body in dorsal view. 16, Aﬂaj irrigation system (Wilkinson head in full-face view (Will Ericson, photographer, 1977, Norman et al., 1998, Gebauer casent0263886, KSA: www.AntWeb.org). et al. 2007). Although crops other than dates may be present (see Gebauer et al. 2007), often large areas of the plantations are left unmanaged allowing VOLUME 119, NUMBER 1 87

Fig. 17. Distribution of Anochetus species in the Arabian Peninsula. a rich ground flora to develop. Although this species may help clarify this re- no systematic surveys are available for lationship. these plantations in Oman, the lush flora and the cooler and wetter conditions ACKNOWLEDGMENTS should support an abundant and diverse invertebrate fauna, warranting addi- The authors extend their sincere ap- tional studies. preciation to the Deanship of Scientific Presently, three species are known Research at King Saud University for its from the Arabian Peninsula (Fig. 17), funding this Research group NO (RGP A. traegaordhi from KSA and Yemen, -1436-029). We are indebted to Boris A. sedilloti from KSA and A. annetteae Kondratieff (Colorado State University) from Oman. Anochetus traegaordhi is for useful suggestions and careful editing. more common of the three species. The authors are grateful to Michele More than 60 workers of this ant were Esposito and Brian Fisher (California observed at the Thi Ayn Archaeologi- Academy of Sciences) for photographing cal Village, Al Bahah Province, KSA. the new species. We thank Nasser Al Anochetus annettae may be a predator Rashedi and Saif Al-Hatmi (Oman Botanic of termites, a behavior previously Garden) for help in the field. Special thanks suggested by Schatz et al. (1999) for to Mahmoud Abdel-Dayem for making the other species of Anochetus. Future distribution map. The authors are grateful studies on the biology and behavior of to the Anglo Omani Society, London UK, 88 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON for a grant that facilitated our fieldwork in Emery, C. 1884. Materiali per lo studio della Oman. fauna Tunisina raccolti da G. e L. Doria. III. Rassegna delle formiche della Tunisia. [part]. Literature Cited Annali del Museo Civico di Storia Naturale 21[=(2)1]: 373–384. Bharti, H. and A. 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