ISSN 0973-1555(Print) ISSN 2348-7372(Online) HALTERES, Volume 7, 99-105, 2016  DAVID EMMANUEL M. GENERAL

A review of the Harpegnathos Jerdon, 1851 (: Formicidae) in the Philippines, with the description of two new species

David Emmanuel M. General

*University Researcher, University of the Philippines Los Baños Museum of Natural History, College, Los Baños, Laguna 4031, Philippines, and Research Associate, National Museum of the Philippines, Taft Ave, Ermita, Metro Manila 1000, Philippines.

E-mail: [email protected]

Abstract

Harpegnathos honestoi and H. alperti are described from Mt. Isarog, Luzon Island, Philippines. They are the fourth and fifth worker-based species of the genus from the Philippines. Harpegnathos honestoi is the first species known to be at least sub-arboreal. A key to the Philippine species is provided.

Keywords: Harpegnathos honestoi, Harpegnathos alperti, Philippines, dichotomous key, Formicidae, Mt. Isarog, new species.

Received: 15 February 2016; Revised: 24 May 2016; Online: 26 May 2016.

Introduction Harpegnathos Jerdon, 1851 is a genus Luzon Island (Donisthorpe, 1942). Despite of unmistakably distinctive ponerine . the high species diversity, these ants are rarely These large ants bear long curved forceps-like encountered in the Philippines. All four mandibles endowed with more than 50 sharp species are known only from the holotypes teeth, huge ovate eyes at the anterolateral (Chapmani, 1963; Donisthorpe, 1937; Wheeler corners of the head, a long tubular mesosoma, and Chapman, 1925). With the possible and long spindly legs that they use effectively exception of H. venator chapmani, none have to jump. They range from India and Sri Lanka been encountered again since the original to Southeast Asia. collection for lack of attention (DEMG, Three nominal worker-based species unpublished notes). and subspecies and one male-based species are known from the Philippines (General and Materials and Abbreviations Alpert, 2012). None of these are known to be Specimens were examined and sympatrically associated, each worker-based measured using a Wild M-5A species having been found on different islands stereomicroscope with ocular micrometer. (see Figure 9): H. venator chapmani Images were created using a Canon 7D digital Donisthorpe, 1937 on Luzon Island, H. camera attached to a Leica MZ16 macgregori Wheeler & Chapman, 1925 on stereomicroscope. Montage images were Biliran Island, and H. empesoi Chapmani, rendered using Helicon Focus 6. Images were 1963 on Mindanao Island (Chapman, 1963; edited with Adobe Photoshop CS6 Extended. Donisthorpe, 1937; Wheeler and Chapman, Coordinates for historical collections were 1925). Despite the fact that these species are obtained from the Philippine Gazetteer allopatric, I consider them good species (DIVA-GIS 2014). Coordinates for the because of their clear morphological present specimens refer to the base camp to differences. These morphological differences discourage poachers. are summarized in the identification key (see below). The male-based species, H. The following measurements and indices are medioniger Donisthorpe, 1942 is known from reported.

99

David Emmanuel M. General

Measurements (in millimeters) margin of petiolar denticle to EL Eye length along the maximum posterior face of petiole diameter. PW Maximum width of pronotum in EW Maximum width of compound eye. dorsal view. HFL Maximum length of hind femur in SL Length of scape, excluding the anterior view. basal neck and condyle. HL Maximum head length in full face TL The total outstretched length of the (dorsal) view, measured from the ant from the mandibular apex to anterior-most point of the clypeal the gastral apex; when measured in margin to the posterior-most point profile the sum of mandibular of head capsule. length + head length + mesosomal HW Maximum head width in full face length + lengths of waist segments (dorsal) view, measured behind the + length of gaster eyes. Indices MandL Mandible length, the straight line CI Cephalic Index: HW/HL x 100. length of the mandible at full EI Eye Index: EL/HW x 100. closure, measured in the same MI Mandibulo-cephalic Index: MandL plane for which the HL *100/HL measurement is taken (i.e. full face SI Scape index: SL/HW x 100 view), from the mandibular apex to the anterior edge of the frontal Repositories lobe, or to the transverse line AMNH American Museum of Naturl connecting the anterior-most points History, New York, NY, USA. in those taxa where the margin is BMNH Natural History Museum, London, concave medially. UK. ML Mesosomal length measured from MCZC Museum of Comparative Zoology, the anterior edge of the pronotum Harvard University, Cambridge, (excluding the collar) to the MA, USA. posterior edge of the propodeal PNM National Museum of the Philippines, lobe. Manila, Philippines. MLO Mandibular outside length, USNM United States National Museum of maximum absolute chord length of Natural History, Washington, D.C., left mandible measured from USA. lateral insertion to apex, in oblique lateral view so that the entire List of species of Harpegnathos known from mandible is in focus the Philippines MOW Median ocellus width, maximum H. alperti sp. n. width of median ocellus H. empesoi Chapman 1963 (examined) MtL Maximum length of gaster, from H. honestoi sp. n. base of abdominal tergite IV to H. macgregori Wheeler & Chapman, 1925 apex of abdominal tergite VII, (examined) measured in lateral view. H. medioniger Donisthorpe, 1942 (based on PetH Petiole height, petiolar height in the male; not seen) lateral profile measured as the H. venator chapmani Donisthorpe, 1937 (not perpendicular distance from the seen) ventral margin to the highest point of posterolateral tubercles; if Results ventral margin is concave upward Harpegnathos honestoi General sp. n. then measured from a line tangent to the uppermost portion of the urn:lsid:zoobank.org:act:DC10CBBE- curve and oriented as close as 79B9-4B68-8D14-1EC4952FED20 possible to the long axis of petiole. PetL Petiole length, straight line length, Holotype. PHILIPPINES: Luzon Island, in lateral view, from anterior Camarines Sur, Naga City, Panicuason

100

A review of the ant genus Harpegnathos Jerdon, 1851 in the Philippines

1 2

3 4

Figures3 1-4. Harpegnathos honestoi sp. n., holotype. 1. lateral view, 2. frontal head view, 3. dorsal view, 4. labels.

5 6

8

7

Figures 5-8. Harpegnathos alperti sp. n., holotype. 5. lateral view, 6. frontal head view, 7. dorsal view, 8. labels.

101

David Emmanuel M. General

Village, Mt. Isarog, 500-550 m ± 500 m, vegetation. I collected both the spider and its 13°40’11” N, 123°19’47” E ± 4 km, prey as the spider was struggling to wrap the 30.iv.2002, leg. D.E.M. General, (PNM 9021, ant in silk (DEMG, unpublished notes). deposited in PNM). Etymology: The name of the new species is a patronym lovingly dedicated to my father, Type locality: Philippines: Camarines Sur, Honesto C. General, who recently celebrated Naga City, Panicuason Village, Mt. Isarog. his 90th birthday.

Description of worker (Figs. 1-4) Harpegnathos alperti General sp. n. Holotype measurements: TL 20.44, HL 2.81, HW 2.39, CI 85, SL 2.96, SI 124, MandL urn:lsid:zoobank.org:act:6B58C862-57D7- 4.11, MI 146, MLO 4.16, PW 1.98, ML 5.15, 4EFA-BF6A-B5D216BA99A3 PetL 1.40, PetH 0.88, HFL 3.64, MtL 4.58, EL 1.35, EW 0.94, EI 57, MOW 0.11. Holotype. PHILIPPINES: Luzon Island, In full face view, posterior margin of Camarines Sur, Naga City, Panicuason head straight; scape exceeding posterior Village, 500-550 m ± 500 m, 13°40’11” N, margin of head by at least width of scape; 123°19’47” E ± 4 km, 12.iii.2003, coll. ocelli present; frontal lobes broad, covering D.E.M. General, G.D. Alpert, et al. (PNM antennal sockets; frontal carinae short, as long 13015, deposited in PNM). as about twice width of scape, diverging; clypeus narrowly inserted between frontal Description of worker (Figs. 5-8) lobes; triangular labral lobe present; eyes Measurements: TL 19.45, HL 2.81, HW 2.55, extremely large, ovate, occupying anterior CI 91, SL 2.60, SI 102, MandL 3.54, MI 126, lateral margin of head; mandibles converging MLO 3.64, PW 2.03, ML 5.04, PetL 1.35, gradually from attachments; head coarsely PetH 0.88, HFL 3.22, MtL 4.16, EL 1.40, EW striate; antennal scape with sparse, short erect 0.94, EI 55, MOW 0.11 (n=1). and suberect hairs. In full face view, posterior margin of In lateral view, mesosoma long and head straight; scape exceeds posterior margin cylindrical; front coxa well separated from of head by at least width of scape; ocelli mid- and hindcoxae; front coxa long but distal present; frontal lobes broad, covering antennal end not reaching midcoxa; promesonotal sockets; frontal carinae short, as long as about suture deeply impressed; metanotal groove twice width of scape, diverging; clypeus obsolete; dorsal face of propodeum very long; narrowly inserted between frontal lobes; propodeal declivity not bounded by lateral triangular labral lobe present; eyes extremely carinae; metapleural gland orifice opening large, ovate, occupying the anterior lateral laterally, not protected by guard hairs; petiole margin of head; mandibles converging rather longer than tall; anterior subpetiolar process abruptly from attachments; head irregularly triangular; gaster long; sting present and reticulo-punctate; antennal scape with sparse, functional; tarsal claws with median tooth. short erect and suberect hairs. In dorsal view, irregular striae In lateral view, mesosoma long and subparallel on pronotum and mesonotum; cylindrical; front coxa well separated from propodeum transversely striate; petiole longer mid- and hindcoxae; front coxa long but distal than broad; petiole reticulate dorsally and end not reaching midcoxa; promesonotal laterally; first and second gastral tergite suture deeply impressed; metanotal groove coarsely punctate over underlying obsolete; dorsal face of propodeum very long; punctulation. propodeal declivity not bounded by lateral Sparse short hairs on body. Head, carinae; metapleural gland orifice opening mandibles, antennae and pronotum orange; laterally, not protected by guard hairs; petiole rest of body brownish black; legs yellow. longer than tall; anterior subpetiolar process Known only from the holotype. triangular; gaster long; sting present and Queen and male unknown. functional; tarsal claws with median tooth. In dorsal view, irregular striae Bionomics: Collected live from web of a subparallel, but diverging posterior fourth of theridiid spider, about 1 m from ground in low pronotum; irregular striae subparallel on mesonotum; propodeum coarsely punctate;

102

A review of the ant genus Harpegnathos Jerdon, 1851 in the Philippines

petiole longer than broad; petiole coarsely differences with a non-type specimen of H. v. punctate dorsally and laterally; first and rugosus. Unfortunately, the holotype of H. v. second gastral tergite coarsely punctate over chapmani cannot be located at the MCZC, underlying punctulation. AMNH, USNM, or BMNH, precluding a Sparse short hairs on body. Body direct comparison and a confident black; mandibles and antennae chocolate- determination (DEMG, personal observation; brown; legs yellowish chocolate-brown. Natural History Museum Data Portal. 2016).

Comparative Note: This specimen is Etymology: This species is named after my superficially similar to H. venator chapmani mentor and colleague, Dr. Gary D. Alpert, a Donisthorpe, 1937 which Donisthorpe (1937) true friend who hosted all my visits to considered a black variety of H. venator F. Cambridge, MA, USA. Gary’s fortuitous visit Smith, 1858, albeit with sculpturation similar to Naga City and Mt. Isarog firmly redirected to H. v. rugosus Mayr, 1862. The key (see my research interest from spiders to ants. below) summarizes the morphological

Figure 9. Distribution map of Harpegnathos species in the Philippines. Each valid species is known only from its type locality. Darker regions denote areas of higher elevation.

103

David Emmanuel M. General

Figure 10. Inset map showing distribution of two Harpegnathos species on Mt. Isarog. The coordinates for Harpegnathos alperti are representative coordinates for Mt. Isarog Natural Park from the Philippine Gazetteer (DIVA-GIS 2014) and do not indicate the actual collection site for the species.

Key to the Philippine species of - First and second gastral tergite with Harpegnathos, based on the worker punctulation only; in full-face view, posterior margin of head shallowly 1. Dorsum of head reticulo-punctate; emarginated; mandibles concolorous mandibles converging rather abruptly from yellow……………………………………4 attachments……………………………..2 4. Antennal scape barely exceeds posterior - Dorsum of head striate; mandibles margin of head by about the width of converging gradually from scape; propodeal declivity bounded by attachments……………………………..3 lateral carinae; pilosity on body 2. Sculpture on dorsa of pronotum, sparse...... H. empesoi Chapman mesonotum, propodeum and petiole - Antennal scape exceeds posterior margin reticulo-punctate; pilosity on body of head by almost 1/3 length of scape; moderately abundant………..H. venator propodeal declivity not bounded by lateral rugosus Mayr carinae; pilosity on body moderately - Sculpture on dorsum of pronotum with abundant…...... H. macgregori irregular striae diverging posteriorly; Wheeler & Chapman dorsum of mesonotum with irregular but subparallel striae; dorsum of propodeum Discussion coarsely punctuate; petiole with dorsal and Harpegnathos honestoi is lateral coarse punctation; pilosity on body sympatrically distributed with H. alperti, both sparse …………………....H. alperti sp. n. having been collected in sites less than 5 km 3. First and second gastral tergite with coarse apart on Mt. Isarog, Luzon Island, Philippines punctation over underlying punctulation; (Figure 10). This may be the only instance of in full-face view, posterior margin of two Harpegnathos species from a single head straight; mandibles orange with location. Bharti et al. (2016) reported the infuscated base ……….H. honestoi sp. n. presence of H. saltator Jerdon, 1851 and H.

104

A review of the ant genus Harpegnathos Jerdon, 1851 in the Philippines

saltator cruentatus (F. Smith, 1858) in the 551: 1-83. Indian states of Karnataka, Kerala, DIVA-GIS. 2014. Philippine Gazetteer. Maharashtra, and of H. saltator and H. venator Accessed online at http://www.diva- (F. Smith, 1858) in the Indian states of Punjab, gis.org/datadown. Tamil Nadu, and West Bengal. It is not clear Chapman, J.W. 1963. Some new and how closely sympatric their distributions are interesting Philippine ants (Hymenoptera: within the confines of each State. Formicidae). Philippine Journal of Science The presence of at least two species of 92: 247-263. Harpegnathos in close proximity on Mt. Donisthorpe, H. 1937. A new species of Isarog suggests the conservation importance of Harpegnathos Jerd., with some remarks the forest of that single mountain. There is on the genus, and the other known species much to discover on Mt. Isarog. Because the (Hym. Formicidae). Entomologist’s natural history and interaction of these two Monthly Magazine 73: 196-201. species is unknown, it is imperative that the Donisthorpe, H. 1942. Descriptions of a few remaining forest of Mt. Isarog be conserved ants from the Philippine Islands, and a for future field research. male of Polyrhachis bihamata Drury from More field work is needed at both Mt. India. Annals and Magazine of Natural Makiling (the type locality of Harpegnathos History (11) 9: 64-72. venator chapmani) and Mt. Isarog to elucidate General, D.M. and Alpert, G.D. 2012. A the taxonomic status of H. v. chapmani. The synoptic review of the ant genera natural history of the Philippine species of (Hymenoptera: Formicidae) of the Harpegnathos also needs to be studied. Philippines. ZooKeys 200: 1-111. Natural History Museum Data Portal. 2016. Acknowledgments Accessed online at I am extremely grateful to Gary Alpert http://data.nhm.ac.uk/dataset/56e711e6- and Mary Corrigan for graciously hosting my c847-4f99-915a visit to Cambridge, MA for this study. I also 6894bb5c5dea/resource/05ff2255-c38a- thank Gary for teaching me how to use his 40c9-b657- new imaging system and to edit the images for 4ccb55ab2feb?q=Harpegnathos+venator+ publication. I greatly appreciate the chapmani&view_id=6b611d29-1dcf-4c60- constructive comments and suggestions of b6b5- Francisco Hita Garcia and Phil Ward which 4cbb69fdf4fe&filters=collectionCode%3A much improved the paper. Thanks to Stefan BMNH%28E%29%2F. Cover and Jignasha Rana for her assistance in Wheeler, W.M. and Chapman, J.W. 1925. The the MCZ Ant Room and to JJ Dida of the ants of the Philippine Islands. Part I, Institute of Renewable Natural Resources, Dorylinae and . Philippine UPLB for the use of his computer to generate Journal of Science 28: 47-73. the map. Finally, I am very grateful to the Harvard University Grant Committee for providing an Ernst Mayr Travel Grant for my visit to MCZ and to Linda Ford for allowing me access to the MCZ Ant Room.

References Alpert, G.D., Cover, S.P., General, D.M. and Samarita, V. 2006. Ants of the Philippines. Polistes Foundation, Discover Life. Accessed online at http://pick4.pick.uga.edu/mp/20q?act=x_c hecklistandguide=Ants_ Philippines. Bharti, H., Guenard, B., Bharti, M. and Economo, E.P. 2016. An updated checklist of the ants of India with their specific distributions in Indian states (Hymenoptera, Formicidae). ZooKeys

105