A peer-reviewed open-access journal ZooKeys 413: |—170 (2014) doi: 10.3897/zookeys.4 13.7172 #7,00Ke y S WWwW.ZOO keys.o rg Launched to accelerate biodiversity research The hyper-diverse ant genus Tetramorium Mayr (Hymenoptera, Formicidae) in the Malagasy region - taxonomic revision of the T. naganum, T. plesiarum, T. schaufussii, and T. severini species groups Francisco Hita Garcia'?, Brian L. Fisher!* | Entomology, California Academy of Sciences, 55 Music Concourse Drive, San Francisco, CA 94118, U.S.A. T Attp://zoobank.org/B7ADF56F-935D-4BD8-ADB3-50E96F8BB463 £ Attp://zoobank.ore/7 CFCBBE5-5443-4F94-B2AC-04BFD4D520F0 Corresponding author: Francisco Hita Garcia (fhitagarcia@gmail.com) Academic editor: /. Borowiec | Received 1 February 2014 | Accepted 20 April 2014 | Published 4 June 2014 Attp://zoobank. org/5791 CE9C-1 CCO-4720-9583-8A585DA79446 Citation: Hita Garcia FE, Fisher BL (2014) The hyper-diverse ant genus 7étramorium Mayr (Hymenoptera, Formicidae) in the Malagasy region - taxonomic revision of the ZT’ naganum, T. plesiarum, T. schaufussii, and T. severini species groups. ZooKeys 413: 1-170. doi: 10.3897/zookeys.413.7172 Abstract The taxonomy of the Tetramorium naganum, T. plesiarum, T: schaufussii, and T. severini species groups are revised for the Malagasy region. A total of 31 species are treated, of which 22 are newly described and nine redescribed. This increases the richness of the hyper-diverse genus Tetramorium in the Malagasy region to 106 species, which makes it the most species-rich genus in the region. Twenty-nine of the treated species are endemic to Madagascar, one is endemic to the Comoros, and one species is found predominantly in Madagascar but also on the island of Reunion. The 7’ naganum species group contains five species, which are mainly distributed in the rainforests and montane rainforests of eastern and northern Madagascar: T alperti sp. n., Ti dalek sp. n., T’ enkidu sp. n., T. gilgamesh sp. n., and 1 naganum Bolton, 1979. The T. plesiarum species group holds five species: T’ bressleri sp. n., T. hobbit sp. n., T. gollum sp. n., T mars sp.n., and 7. plesiarum Bolton, 1979. All five are arid-adapted species occurring in the southwest and west of Madagascar. The second-most species-rich group in the region is the T’ schaufussii species group with 20 species, most of which inhabit rainforests or montane rainforests of eastern and northern Madagascar. This group includes two species complexes each containing ten species: the 7’ cognatum complex with the species 7 aspis sp. n., T’ camelliae sp. n., T cognatum Bolton, 1979, T. freya sp. n., T. gladius sp. n., T. karthala sp. n., T’ myrmidon sp. n., T. proximum Bolton, 1979, 7. rumo sp. n., and T. tenuinode sp. n.; Copyright F Hita Garcia, B.L Fisher. This is an open access article distributed under the terms of the Creative Commons Attribution International License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 2 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) and the 7’ schaufussii complex with the species T’ merina sp. n., I monticola sp. n., T. nassonowii Forel, 1892 stat. n., 7’ obiwan sp. n., T. pseudogladius sp. n., 1. rala sp. n., T. schaufussii Forel, 1891, T sikorae Forel, 1892 (= /atior (Santschi, 1926)), 7’ scutum sp. n., 7. xanthogaster Santschi, 1911. The last group treated in this study is the 7’ severini species group, which contains only the species 7’ severini (Emery, 1895). This very conspicuous species is widely distributed in the rainforests and montane rainforests of eastern and northern Madagascar. All four groups are fully revised with group diagnoses, illustrated spe- cies-level identification keys, and detailed descriptions for all species that include multifocused montage images and distribution maps. Keywords Comoros, Madagascar, Reunion, taxonomy, Tetramoriini Table of contents WiGkG Gc LOL ins Fee ee arene See ve aati oe Bee tee aan A Nara Slave teen fa le = FADD TEV ATI OMS 01 EPO LOTEES otra e on tie tee sue ate a ete deet em ental Ona a ake scree soc 5 I Tate tell citer an eU TO Sot secs Peas Dene Aaes stro eae a dhas, MA ant tin eel tay sh ealh ce MS sence a aa 5 Synopsis of Malagasy Tetramorium species treated in this study .........ceeeeeeeeeereeees 7 Revision of the Tetramorium naganum SPeCi€S QLOUP ........scceeessesseesecseceeeeseeessenes 8 LELTAINOTIUINNAPANUM SPECIES OLOUP: 2 vosepapsimeda.csteosn tape sbedieeen ssh opbesilbsn toniopee Unt 8 Identification key to species of the 7’ naganum species group (workers) ..... 10 Tetramorium alperti Hita Garcia & Fisher sp. 1. ......seesesscessesesseeeseesseeseceees 11 letramorium dalek ita Gatcia 86 Fisher sp. ti... .s.2..-cese-nonctinatons teenpsbonotenenes LS letramorium enkidu Pitta Gateia- 86 Fishersp. 11. %2::..0-2heressnensesaeesbeeneretes 19 Tetramorium gilgamesh Hita Garcia & Fisher sp. nu ....esceceseeseeeeseeeeeseceees 22 Letrantoriuim nagariwi BOWOM ai: c. ise cocdets ch Bas cack antaescapenckecaededtaaeotasuadennddtuas 26 Revision of the Tetramorium plesiarum species QrOUP .....scsseesceseeseeseeessesseeseeees 30 LELYATROTININ PICSIATUINSPECIES- CLOUP in. swe php said coves utente lghvond WAeresdesmeraoeeveeteswcnd tes 30 Identification key to species of the 7’ plesiarum species group (workers)..... 31 lPiramorium, bressier: Lita-Garcla-GCFIs her Sp. fis. : teva: cen cevaeBsurhtadess cossenadee 33 Tetramorium gollum Nita Garcia & Fisher sp. Micsssersocesetecssupeoscansoosenpeeseares 39 Tetramorium hobbit Hita Garcia & Fisher sp. 0. ..s.eeeeeseesceescceseeeeeeeseeeeeeees 42 letramoriam mars Hita Gatcia GOP sher spit, fius.nnshtoe. us tved pote Fodsnoieedecha see 46 LOCA ILO FINI P1Css Ay TDG MOM van aceon bene tan ens tinett bain se caeeanadedebhaet adhoc se 50 Revision of the Tetramorium schaufussii species QrOUP....1..ssssceseeseeeseeseeseeseeees 55 LELIAINOTIUIN RCD AU USSLE SPECIES CLOUD: pine coup eshagetcuca ts uedubnsnioes sertenianatenblytceba Shpedks 55 Key to species complexes of the Tetramorium schaufussii species group ....... >) Tetra mor iUUCOPNALUM. SPECIES COMPLEX: 5 -ien Beck an sree Teenie gbede th eanainds tie near -® 57 Identification key to species of the 7’ cognatum species complex (workers) ...58 letramorinm aspis Tita Gatcia 86 Fisher Sp. 17. ....edeissosexcadsasevesevesocoseauns 61 Tetramorium camelliae Hita Garcia & Fisher sp. ne...sceeeeeeeseeeseeeseeeeees 65 Letramorium cognatum Ne fol fro}: Baal by 9A9 bid Ae een Nn eed Seer RenMeh cca 68 Tetramorium freya Hita Garcia & Fisher sp. 1. ...sceseeeeeeeseeeseeseeeseeseeees 76 Taxonomy of Malagasy Tetramorium 3 Tetramorium gladius Hita Garcia & Fisher sp. 1. ...eeeeeeeesesseeseeeseeseeees 79 Tetramorium karthala Hita Garcia & Fisher sp. n.....seeseesseeseeeeeeeseeeeees 82 Tetramorium myrmidon Hita Garcia & Fisher sp. 0... eeeeeeseeseeseeseeees 86 Letramorium proximum GUT ee ese CR ALUN, Salah aan te chaohveoknet toe 90 letramorium rue Wlita:Gatcla:Ga@ Fisher Spe Aiie2s. cepteseesbecaeczeeresbvacnees ee Tetramorium tenuinode Hita Garcia & Fisher sp. 1.......eeeeseeeseeseeeeeees 101 Tetramorium schaufussti species COMpLeX........eseescesseseeecseeeseeseeeseenees 107 Identification key to species of the 7. schaufussii species complex (workers)....108 Tetramorium merina Hita Garcia & Fisher sp. 1. ....eeeeeeeeseeeseeseeeeeeees 111 Tetramorium monticola Hita Garcia & Fisher sp. 1.......eeeeseeseeseeeseeees 116 Tetramorium nassonowii Forel, 1892 stat. 1. .....ccccccsececcseecccsececcsccenecss 121 Tetramorium obiwan Hita Garcia & Fisher sp. 0. .....eeeeeeeseeeeeeseeeeeeees 125 Tetramorium pseudogladius Hita Garcia & Fisher sp. n. woe. 129 Tetramorium rala Hita Garcia & Fisher sp. 1......eeseeeesseeceeeeeeeseeeeeeees 132; Terramioriumischaupussi Potel ASI cap. cecediicebesstenss tetsssatorecsstegntevstsnrdh 136 Tetramorium scutum Hita Garcia & Fisher sp. n......eeeeeeeeeeeeseeseeeeeeees 143 letramorium sikorae Forel; 1892 ..ccccccccccesvelaceaosseeccacesvacccocsveccscesesecenees 146 letramorium xantbogaster Santschi gl 91 x. ayuvsiveersavaswentswearsiea ees 150 Revision of the Téetramorium severini SpeCi€S QLOUP .......sscseseessceseeseeeeeeneesseeseees 156 LCRA OF IUTIESCUCTIAL SPECIES, CLOUD... vrbeunessrercnoteaniverrenenssnitidbdeseuasstsned ones 156 LECPAINONLIESEUETING WPANET CUO D 9) rssh vs testa onsite soa eee ree tayedgeensG IHS 157 INGRTIGWIEAEEIVEMES. antes ensure ttteeiea cy sxe dae quhs atign Panels sapiens Tal aetancenes van chstigs Saget adensies 168 PLATE TICES Sa ut hie etc bsehaey tected tas eanett aunts phan tds nela eget MMAR cou Je Ne teeta ace Me chaeBiszehare 168 4 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Introduction The hyper-diverse genus Tetramorium Mayr is widely distributed throughout all bio- geographic regions, and with currently 520 described species (Bolton 2014) one of the most species-rich ant genera. Tetramorium is most diverse in the subtropics and trop- ics of the Old World, especially the Afrotropics and Madagascar, but is species-poor in the New World. Most regional faunas were revised by Bolton (1976, 1977, 1979, 1980), who provided a good taxonomic foundation on which later works could build (e.g. Csdsz et al. 2007; Csdsz and Schulz 2010; Hita Garcia et al. 2010; Hita Garcia and Fisher 2011; Bharti and Kumar 2012; Sharaf et al. 2012). The Malagasy Tetramo- rium fauna was also first revised by Bolton (1979), who treated eight species groups containing 36 species, of which 29 were endemic to Madagascar. The synonymisation of Triglyphothrix Forel (Bolton 1985) under Tetramorium added an additional species group with one tramp species; two additional tramp species were reported much later (Blard et al. 2003; Roberts and McGlynn 2004), for a total of 39 species known prior to 2011. Recently, we started a large-scale taxonomic revision of the genus Tetramo- rium for the Malagasy region based initially on more than 160 morphospecies with more than 40,000 mounted specimens (Hita Garcia and Fisher 2011). We proposed 14 species groups as a foundation and provided a preliminary identifica- tion key to the Malagasy groups. In the same study we also revised the taxonomy of the 7. bicarinatum, T. obesum, T. sericeiventre, and T. tosii species groups. One species was newly described while another was synonymised, leaving the species count for the region at 39. Based on that work, the 7: bessonii, T. bonibony, T. dysalum, T. kelleri, T. marginatum, T. tortuosum, T. tsingy, and T. tosii species groups were revised shortly afterward (Hita Garcia and Fisher 2012a, 2012b). The latter two studies dealt with 58 species, of which 45 were described as new, and increased the species count for the Malagasy region to 84. We also proposed additional species groups for a total of 18 for the region (not 19 as noted in Hita Garcia and Fisher 2012b). In this study we revise the taxonomy of four more Malagasy species groups: 7. naganum, T. plesiarum, T. schaufussii, and T: severini species groups. All are fully re- vised with group diagnoses, illustrated species-level identification keys, multifocused montage images for all species, descriptions of 22 new species, redescriptions of nine previously known species, and distribution maps for all species. The species treated here increase the total species count for the genus Tetramorium in the Malagasy region to 106, which means that at present this genus holds the highest described species rich- ness in the region. Furthermore, this study increases the number of recently revised Malagasy Tetramorium species groups to 16 (Hita Garcia and Fisher 2011, 2012a, 2012b), leaving only two species groups untreated: the species-rich 7: ranarum group and the less diverse but abundant 7! simillimum group. These groups will be revised in an upcoming study. Taxonomy of Malagasy Tetramorium 5 Abbreviations of depositories The collection abbreviations follow Evenhuis (2013). The material upon which this study is based is located and/or was examined at the following institutions: BMNH._ The Natural History Museum (British Museum, Natural History), London, Guke CAS California Academy of Sciences, San Francisco, California, U.S.A. MCSN_ Museo Civico di Storia Naturale Giacomo Doria, Genoa, Italy MCZ Museum of Comparative Zoology, Cambridge, Massachusetts, U.S.A. MHNG Muséum d’Histoire Naturelle de la Ville de Genéve, Geneva, Switzerland NHMB Naturhistorisches Museum, Basel, Switzerland Material and methods The material examined in this study is based on ant inventories carried out in the Mala- gasy region from 1992 to 2012, which included more than 6,000 leaf litter samples, 4,000 pitfall traps, and 9,000 additional hand collecting events (see Fisher 2005 for additional details). All new type material and all imaged specimens can be uniquely identified with specimen-level codes affixed to each pin (e.g. CASENT0078328). In the presented descriptions we list all of the available specimen-level codes for the whole type series. It should be noted, however, that the number of stated paratype workers does not necessarily match the number of listed specimen-level codes because several pins hold more than one specimen. Digital colour montage images were created using a JVC KY-F75 digital camera and Syncroscopy Auto-Montage software (version 5.0), or a Leica DFC 425 camera in combination with the Leica Application Suite software (version 3.8). All images presented are available online and can be seen on AntWeb (http://www.antweb.org). The distribution maps provided at the end of the study (Figs 61-66) were generated with R software (R Core Team 2014). The measurements were taken with a Leica MZ 12.5 stereomicroscope equipped with an orthogonal pair of micrometers at a magnif- cation of 100x, rarely 80x. Measurements and indices are presented as minimum and maximum values with arithmetic means in parentheses. In addition, all measurements are expressed in mm to two decimal places. The measurements and indices used in this study follow Bolton (1975, 1979, 1980), Giisten et al. (2006), and Hita Garcia and Fisher (2011, 2012a, 2012b, 2013): HL Head length: maximum distance from the midpoint of the anterior clypeal margin to the midpoint of the posterior margin of head, measured in full-face view. Impressions on anterior clypeal margin and posterior head margin re- duce head length. PSL PTH PTL PTW PPH PPL PPW OI CI SI DMI LMI PSLI PeNI LPel DPel PpNI LPpI DPpl PPI Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Head width: width of head directly behind the eyes measured in full-face view. Scape length: maximum scape length excluding basal condyle and neck. Eye length: maximum diameter of compound eye measured in oblique lateral view. Pronotal width: maximum width of pronotum measured in dorsal view. Weber’s length: diagonal length of mesosoma in lateral view from the poster- oventral margin of propodeal lobe to the anteriormost point of pronotal slope, excluding the neck. Propodeal spine length: the tip of the measured spine, its base, and the centre of the propodeal concavity between the spines must all be in focus. Using a dual-axis micrometer the spine length is measured from the tip of the spine to a virtual point at its base where the spine axis meets orthogonally with a line leading to the median point of the concavity. Petiolar node height: maximum height of petiolar node measured in lateral view from the highest (median) point of the node to the ventral outline. The measur- ing line is placed at an orthogonal angle to the ventral outline of the node. Petiolar node length: maximum length of the dorsal face of the petiolar node from the anterodorsal to the posterodorsal angle, measured in dorsal view ex- cluding the peduncle. Petiolar node width: maximum width of dorsal face of petiolar node measured in dorsal view. Postpetiole height: maximum height of the postpetiole measured in lateral view from the highest (median) point of the node to the ventral outline. The measuring line is placed at an orthogonal angle to the ventral outline of the node. Postpetiole length: maximum length of postpetiole measured in dorsal view. Postpetiole width: maximum width of postpetiole measured in dorsal view. Ocular index: EL / HW x 100 Cephalic index: HW / HL x 100 Scape index: SL/ HW x 100 Dorsal mesosoma index: PW / WL x 100 Lateral mesosoma index: PH / WL x 100 Propodeal spine index: PSL / HL x 100 Petiolar node index: PTW / PW x 100 Lateral petiole index: PTL / PTH x 100 Dorsal petiole index: PTW / PTL x 100 Postpetiolar node index: PPW / PW x 100 Lateral postpetiole index: PPL / PPH x 100 Dorsal postpetiole index: PPW / PPL x 100 Postpetiole index: PPW / PTW x 100 Note that the petiole and postpetiole were measured differently. For the petiole, only the petiolar node was measured, excluding the peduncle, as the node has proved Taxonomy of Malagasy Tetramorium q to be of high diagnostic value (Hita Garcia et al. 2010). Measurements of the whole petiole, peduncle plus node, would mask important differences between species. In contrast, we measured the whole postpetiole because it was rounded in most species and without a distinct peduncle-like structure. As a consequence, some information might be lost in the few species with a moderately or strongly anteroposteriorly com- pressed postpetiole. Even so, the postpetiole measurements as defined still permit bet- ter comparisons for most species. Pubescence and pilosity are often of high diagnostic value within the genus Tetramo- rium (e.g. Bolton 1976, 1980, 1985; Hita Garcia et al. 2010, Hita Garcia and Fisher 2011, 2012a). The varying degree of inclination of pilosity is particularly important for the diagnosis of species, complexes, or groups. In this context we use the terms “erect”, “suberect”, “subdecumbent”, “decumbent”, and “appressed” following Wilson (1955). Synopsis of Malagasy Tetramorium species treated in this study Tetramorium naganum species group Tetramorium alperti sp. n. Tetramorium dalek sp. n. Tetramorium enkidu sp. n. Tetramorium gilgamesh sp. n. Tetramorium naganum Bolton, 1979 Tetramorium plesiarum species group Tetramorium bressleri sp. n. Tetramorium hobbit sp. n. Tetramorium gollum sp. n. Letramorium mars sp. n. Tetramorium plesiarum Bolton, 1979 Tetramorium schaufussii species group Tetramorium cognatum species complex Letramorium aspis sp. n. Tetramorium camelliae sp. n. Letramorium cognatum Bolton, 1979 Tetramorium freya sp. n. Tetramorium gladius sp. n. Tetramorium karthala sp. n. Tetramorium myrmidon sp. n. Letramorium proximum Bolton, 1979 Letramorium rumo sp. n. Tetramorium tenuinode sp. n. 8 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Tetramorium schaufussii species complex Tetramorium merina sp. n. Tetramorium monticola sp. n. Tetramorium nassonowii Forel, 1892, stat. n. Tetramorium obiwan sp. n. Tetramorium pseudogladius sp. n. Tetramorium rala sp. n. Tetramorium schaufussii Forel, 1891 Tetramorium sikorae Forel, 1892 = Tetramorium latior (Santschi, 1926) Tetramorium scutum sp. n. Tetramorium xanthogaster Santschi, 1911 Tetramorium severini species group Tetramorium severini (Emery, 1895) Revision of the Tetramorium naganum species group Tetramorium naganum species group Diagnosis. Eleven-segmented antennae; anterior clypeal margin medially impressed; frontal carinae always well developed, diverging posteriorly and usually approaching or reaching corners of posterior head margin; antennal scrobe present, weakly to very well developed; mesosoma moderately to strongly marginate from sides to dorsum; pro- podeal spines medium-sized to long, elongate-triangular to spinose; propodeal lobes triangular and short; petiolar node in profile high rounded nodiform to rectangular nodiform with moderately to well-rounded margins, in profile 1.5 to two times higher than long (LPel 50-68), in dorsal view between 1.0 and 1.4 times wider than long (DPel 103-139), anterior and posterior faces parallel, anterodorsal and posterodorsal margins usually at about same height (sometimes anterodorsal margin higher); postpe- tiole in profile always more or less globular; mandibles variably sculptured, but mostly unsculptured; cephalic dorsum and dorsal mesosoma with distinct longitudinally ru- gose sculpture; waist segments and gaster always unsculptured, smooth, and shiny; dorsal surfaces of head, mesosoma, and usually waist segments with few to abundant long, standing hairs; pilosity/pubescence on first gastral tergite variable, but with ten- dencies to more inclined pilosity and dense pubescence; sting appendage spatulate. Comments. This small and compact group is restricted in its distribution to east- ern and northern Madagascar. All five species are found only in rainforests or montane rainforests and seem to live mainly in leaf litter. Within the 13 species groups with 1 1-segmented antennae, the 7: naganum group shares the complete lack of sculpture on both waist segments with the majority of groups, but differs from the 7: kelleri, T. ranarum, T. tortuosum, and parts of the Taxonomy of Malagasy Tetramorium 9 T. dysalum groups. These groups contain only species in which either one or both waist segments are clearly sculptured. In addition, the very well developed sculpture on head and mesosoma distinguishes the 7: naganum group from the groups with reduced sculpture: the 7° bessonii, T. marginatum, and T. tsingy groups. Also, T. severini, the only member of the T° severini group, has a longer and lower mesosoma (LMI 35-38) with less margination from the sides to the dorsum, while the species of the T° naga- num group have a higher (LMI 40-46), stouter, and more angled mesosoma. The 7: plesiarum group is characterised by the presence of relatively deep and well-developed antennal scrobes with margins all around and a strongly developed median scrobal carina, a character always absent in the 7’ naganum group. The latter also cannot be mistaken for the T° bonibony group, in which some species possess a very conspicuous bump or protuberance on the pronotal dorsum while the remainder of the species have triangular or cuneiform petiolar nodes. ‘The differentiation between the 7. naganum group and the 7: dysalum group can be more difficult however. The 7: dysalum group is relatively heterogeneous and there are a few species that are morphologically close to some species in the 7: naganum group. Nevertheless, the best method to discrimi- nate between these two groups is to compare gastral pubescence and/or pilosity. In all species of the 7: dysalum group appressed pubescence on the first gastral is scarce and inconspicuous, and pilosity consists of numerous long suberect to erect hairs. By contrast, pubescence and pilosity are very variable in the 7° naganum group, but never with very reduced pubescence and long, standing pilosity as in the 7: dysalum group. The separation from the 7° schaufussii group is likely the most difficult, and 7! na- ganum was a member of that group until recently (Hita Garcia and Fisher 2012a). The five species of the 7: naganum group have much stronger developed frontal carinae, a generally broader head (CI 92-99), a higher mesosoma (LMI 40-46), and usually longer propodeal spines (PSLI 25-37). In the T° schaufussii group most species (but not all) have weaker frontal carinae, a usually thinner head (CI 85-95), a lower meso- soma (LMI 35-42), and usually much shorter propodeal spines (PSLI 7—28). These values overlap in a few species. In fact, most members of the 7° schaufussii group have relatively short spines with a PSLI below 20, a few species have slightly longer spines (PSLI 20-25), and only a few specimens of 7. gladius and T. rumo have a higher PSLI of 26-28. The five species of the group are morphologically very close to each other, and their delimitations are mainly based on different patterns of pilosity/pubescence on the waist segments and the first gastral tergite. Tetramorium dalek is easily separable from the other four species on the basis of the absence of standing hairs on the waist segments and shorter propodeal spines, but 7. alperti, T. enkidu, T. gilgamesh, and T. naganum are morphologically very similar. Indeed, they are often difficult to dis- criminate and there are only very few reliable diagnostic characters, mainly gastral pilosity/pubescence. It is possible that two, three, or all four are conspecific and the abovementioned differences are just intraspecific variation. Nevertheless, we prefer to treat them as four distinct species since gastral pilosity/pubescence is usually very species-specific within the genus Tetramorium and other myrmicine genera and the 10 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) material listed here can be moderately well distinguished by the diagnostics given in the key and the descriptions. However, we cannot rule out that gastral pilosity/ pubescence is highly variable in this group, and if so, our hypotheses may not reflect real species boundaries. Identification key to species of the T’ naganum species group (workers) 1 Propodeal spines relatively shorter (PSLI 25-27); waist segments without long standing hairs, instead with short, appressed to subdecumbent pubes- cence only, sometimes with one or two short erect to suberect hairs (Fig. 1A) (eG GaSe ai estas et sdb snared nents th tioned Shak: Subdodee mnie msn ivedia: T. dalek — Propodeal spines relatively longer (PSLI 27-37); waist segments always with long standing hairs and short appressed to subdecumbent pubescence (Fig. IAS) Bef oa re One Pee Rn co Peer rte oe One Ce RE a oak Sol Pate a eed Ra Pear ps 2 In profile petiolar node relatively thicker, around 1.5 to 1.6 times higher than longs Pel GO-GS MORI, DAG By Ne. seusech, vabeah trseseibahaeenbedis ong dias -buttpoang Wtheteasttas! 3 = In profile petiolar node relatively thinner, around 1.7 to 2.0 times higher thandoea Ee Reli50— 59 )iCFig aC. ID): a si.ca eet score Bi hae. Gk Sa 4, 3 First gastral tergite with moderately long, relatively scattered, appressed to decumbent pubescence in combination with several much longer, fine, and Srecwniaitsn(bigeS rch IMAC ap AS CAL et gewacectaehewe suslecesoueeyar rteseetereeceont T. alperti — First gastral tergite with moderately short, abundant, subdecumbent to su- berect pilosity, and without short, dense, appressed to decumbent pubes- cence or long, fine erect hairs (Fig. 3B) [Madagascar] ............. T. enkidu 4 Eyes larger (OI 25-27); first gastral tergite covered by a mix of short to mod- erately long, abundant, decumbent to suberect pilosity, pilosity appearing disorganized due to varying degrees of inclination and hair length (Fig. 4A) [Mia ascaia beset fesse. th sucess conc S au dts lsu Soca S38 satay satabada neta eat T. gilgamesh — Eyes smaller (OI 21—23); first gastral tergite with short, dense, appressed to decumbent pubescence only, without any long, standing pilosity (Fig. 4B) [Mad aGaseat |Meat ka ade tone tit coat onic dk othe deenoant: T. naganum Figure |. Propodeal spines and waist segments in profile. A 7’ dalek (CASENT0038402) B 7’ naganum (CASENT0280584). Taxonomy of Malagasy Tetramorium 11 Figure 2. Petiole and postpetiole in profile. A 7 alperti (CASENT0042547) B 7 enkidu (CASENT0045673) € 7) naganum (CASENT0280584) D 7. gilgamesh (CASENT0247312). Figure 4. First gastral tergite in profile. A 7) gileamesh (CASENT0247312) B 7 naganum (CASENT0102346). Tetramorium alperti Hita Garcia & Fisher, sp. n. http://zoobank.org/53D9615C-CF34-4216-B511-41081A7031B7 http://species-id.net/wiki/Tetramorium_alperti Figs*2iy SAs15; 61 Type material. Holotype, pinned worker, MADAGASCAR, Antsiranana, Parc Na- tional de Marojejy, Antranohofa, 26.6 km 31° NNE Andapa, 10.7 km 318° NW Manantenina, 14.44333°S, 49.74333°E, 1325 m, montane rainforest, sifted lit- ue Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) ter (leaf mould, rotten wood), collection code BLF09080, 18.XI.2003 (B.L. Fisher) (CAS: CASENT0042547). Paratypes, nine pinned workers with same data as holo- type (BMNH: CASENT0042817; CAS: CASENT0042703; CASENT0042704; CASENT0042708; CASENT0042813; CASENT0042815; CASENT0042827; CASENT0042835; MCZ: CASENT0042821). Non-type material. MADAGASCAR: Antsiranana, Rés. Anjanaharibe-Sud, 9.2 km WSW Befingotra, 14.75°S, 49.46667°E, 1280 m, montane rainforest, 5.X1.1994 (B.L. Fisher); Fianarantsoa, Foret d’Ambalagoavy Nord, Ikongo, Ambatombe, 21.8275°S, 47.33889°E, 625 m, 1.XII.2000 (R. Harin’Hala & M.E. Irwin). Diagnosis. Tetramorium alperti differs from the other three species of the group by the following character combination: propodeal spines long to very long (PSLI 29-37); waist segments with several long erect hairs; first gastral tergite with moder- ately long, scattered, appressed to decumbent pubescence in combination with several much longer, erect standing hairs. Worker measurements (N=10). HL 0.56—-0.68 (0.64); HW 0.54—0.65 (0.62); SL 0.40-0.47 (0.45); EL 0.13-0.15 (0.14); PH 0.31—0.38 (0.35); PW 0.40-0.50 (0.47); WL 0.70-0.84 (0.79); PSL 0.18-0.23 (0.21); PTL 0.15-0.18 (0.16); PTH 0.23-0.28 (0.26); PTW 0.16—-0.19 (0.18); PPL 0.17—0.22 (0.20); PPH 0.23-0.27 (0.25); PPW 0.22—0.28 (0.26); CI 95-97 (96); SI 69-75 (73); OI 22-23 (23); DMI 57-62 (59); LMI 42-46 (44); PSLI 29-37 (33); PeNI 36—40 (38); LPel 62-68 (64); DPelI 103-115 (108); PpNI 54-57 (55); LPpI 74-84 (80); DPpI 126-132 (128); PPI 139-156 (145). Worker description. Head longer than wide (CI 95-97); posterior head mar- gin weakly concave. Anterior clypeal margin with distinct median impression. Fron- tal carinae strongly developed, diverging posteriorly and approaching or ending at posterior head margin; antennal scrobe present, but weak, shallow, and without de- fined posterior or ventral margins. Antennal scapes short, not reaching posterior head margin (SI 69-75). Eyes of moderate size (OI 22—23). Mesosomal outline in profile flat to weakly convex, relatively high (LMI 42—46), and moderately to strongly mar- ginate from lateral to dorsal mesosoma; promesonotal suture and metanotal groove absent. Propodeal spines long to very long, spinose, acute, and often thick (PSLI 29-— 37); propodeal lobes short, triangular, and blunt or acute, always much shorter than propodeal spines. Petiolar node in profile high, rounded nodiform with moderately rounded antero- and posterodorsal margins, around 1.5 to 1.6 times higher than long (LPel 62-68), anterior and posterior faces approximately parallel, anterodorsal and posterodorsal margins situated at about the same height and equally marginate, peti- olar dorsum weakly convex; node in dorsal view around 1.0 to 1.1 times wider than long (DPel 103-115), in dorsal view pronotum between 2.4 to 2.8 times wider than petiolar node (PeNI 36-40). Postpetiole in profile globular to subglobular, approxi- mately 1.2 to 1.3 times higher than long (LPpI 74-84); in dorsal view around 1.2 to 1.3 times wider than long (DPpI 126-132), pronotum between 1.7 to 1.8 times wider than postpetiole (PpNI 54-57). Postpetiole in profile appearing less voluminous than petiolar node, postpetiole in dorsal view between 1.3 to 1.6 times wider than petiolar Taxonomy of Malagasy Tetramorium 13 node (PPI 139-156). Mandibles usually mostly unsculptured and smooth with some weakly striate parts, generally very shiny; clypeus longitudinally rugose/rugulose, with three to five rugae/rugulae, median ruga always well developed and distinct, lateral rugae/rugulae sometimes weaker and interrupted; cephalic dorsum between frontal carinae with seven to nine longitudinal rugae, rugae running from posterior clypeal margin to posterior head margin, often interrupted or with cross-meshes, especially posteriorly; scrobal area mostly unsculptured; lateral head reticulate-rugose to longitu- dinally rugose. Ground sculpture on head absent to weakly punctate. Dorsum of mes- osoma mostly irregularly longitudinally rugose; lateral mesosoma variably sculptured, lateral pronotum and anepisternum mostly unsculptured, smooth and shining with very little sculpture, usually only traces of rugulae present, katepisternum, metapleu- ron, and lateral propodeum noticeably irregularly longitudinally rugose. Forecoxae unsculptured, smooth and shining. Ground sculpture on mesosoma weak to absent. Waist segments and gaster completely unsculptured, smooth and shining. Whole body with numerous, long, and fine standing hairs; first gastral tergite with moderately long, relatively scattered, appressed to decumbent pubescence in combination with several much longer, fine, and erect hairs. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae usually with decumbent to suberect hairs. Head, mesosoma, waist segments, and gaster orange-brown to chestnut brown, mandibles, antennae, and legs always lighter, usually yellowish brown. Etymology. The name of the new species is a patronym dedicated to Gary D. Alp- ert from Cambridge, Massachusetts, U.S.A., to honour his numerous ant collecting activities in Madagascar. Distribution and biology. This new species is only known from three localities (Fig. 61). Two are located in the northeast of Madagascar (Anjanaharibe-Sud and Ma- rorejy) while the third is found much further south (Ambalagoavy). Anjanaharibe-Sud and Marorejy are montane forests ranging from 1280 to 1325 m elevation, whereas Ambalagoavy is located at 525 m. ‘This distributional and elevational pattern suggests that 7° alperti might have been distributed throughout most of the eastern Madagascar humid forests, but is now only found in a few montane forests and one lower rainforest site. Based on the available collection data, 7: alperti seems to be a leaf litter inhabitant. Discussion. Tetramorium alperti cannot be confused with T. dalek since the latter has no long standing hairs on the waist segments and the first gastral tergite, while these are present in T° alperti. Differentiation from the other three species of the group, however, is more challenging. The primary diagnostic characters distin- guishing 7° alperti are the pilosity/pubescence pattern on the first gastral tergite and the shape of the petiolar node in profile. Tetramorium alperti possesses moderately long, relatively scattered, appressed to decumbent pubescence in combination with several much longer and erect hairs. This pattern on the first gastral tergite is not found in 7. enkidu, T. gilgamesh or T. naganum since they either lack long and erect hairs or appressed to decumbent pubescence entirely. In addition, T° alperti also has a thicker petiolar node which is around 1.5 to 1.6 times higher than long (LPel 62-68), compared to T° gilgamesh and T. naganum, in which the petiolar nodes are 14 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Figure 5. 7° alperti holotype worker (CASENT0042547). A Body in profile B Body in dorsal view C Head in full-face view. Taxonomy of Malagasy Tetramorium LS around 1.7 to 2.0 times higher than long (LPel 50-59). The species probably most easily confused with 7! alperti is T. enkidu since both share the same general habitus and the same morphometric range, and the only difference is the pattern of pilosity/ pubescence on the first gastral tergite outlined above. Possibly they are conspecific and the differences in gastral pilosity/pubescence represent intraspecific variation, however, there are some good arguments to separate them as two distinct species. First, both species are found in sympatry in Marojejy, and they are easily identified by the differences in pilosity/pubescence mentioned above without any intermedi- ate forms. Second, patterns of pilosity/pubescence are usually very species-specific within the genus Tetramorium (e.g. Bolton 1976, 1980; Hita Garcia et al. 2010; Hita Garcia and Fisher 2012a, 2012b). We were able to reveal several consistent pat- terns of pilosity/pubescence in the 7. naganum species group, and the pattern of T- alperti is unique to this group, although it resembles the one seen in T. ryanphelanae Hita Garcia & Fisher from the 7° bessonii species group, and a few species from the T. schaufussii species complex. To our knowledge, there is no significant intraspecific variation in the material treated as T° alperti. Tetramorium dalek Hita Garcia & Fisher, sp. n http://zoobank.org/F10C4841-A175-4D98- B3DD- ISPBECIPSEO3 http://species-id.net/wiki/Tetramorium_dalek Figs 1A, 6, 61 Type material. Holotype, MADAGASCAR, Toamasina, Montagne d’Anjanaharibe, 19.5 km 27° NNE Ambinanitelo, 15.1783°S, 49.635°E, 1100 m, montane rainfor- est, sifted litter (leaf mold, rotten wood), collection code BLF08150, 12.—16.II1.2003 (B.L. Fisher et al.) (CAS: CASENT0038402). Paratypes, 17 pinned workers with same data as holotype (BMNH: CASENT0038394; CAS: CASENT0038369; CASENT0038372; CASENT0038376; CASENT0038390; CASENT0038408; CASENT0038413; CASENT0038416; CASENT0038425; CASENT0038429; CASENT0038443; CASENT0038445; CASENT0038450; CASENT0038456; CASENT0038465; MCZ: CASENT0038397; CASENT0038424); and seven work- ers from MADAGASCAR, Toamasina, Montagne d’Anjanaharibe, 18.0 km 21° NNE Ambinanitelo, 15.1883°S, 49.615°E, 470 m, rainforest, sifted litter (leaf mold, rotten wood), collection code BLF08802, 8.—12.III.2003 (B.L. Fisher et al.) (CAS: CASENT0037833; CASENT0037869; CASENT0037903; CASENT0037909; CASENT0037916; CASENT0037933; CASENT0037936). Non-type material. MADAGASCAR: Antsiranana, 1 km W Andampibe, Cap Masoala, 15.69361°S, 50.18139°E, 125 m, lowland rainforest, 29.X1.1993 (G.D. Alpert); Antsiranana, Res. Anjanaharibe-Sud, 9.2 km WSW Befingotra, 14.75°S, 49.4667°E, 1280 m, 4.—-9.XI.1994 (BL. Fisher); Toamasina, Ambohitsitondroina, 6.9 km NE Ambanizana, 15.5851°S, 50.0095°E, 825 m, rainforest, 2.XII.1993 (B.L. 16 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Fisher); Toamasina, Andranobe, 6.3 km S Ambanizana, 15.6813°S, 49.958°E, 25 m, rainforest, 13.-14.X1.1993 (B.L. Fisher); Toamasina, Andranobe, 5.3 km SSE Ambanizana, 15.6713°S, 49.9739°E, 425 m, rainforest, 21.X1.1993 (BL. Fisher); Toamasina, Montagne d’Anjanaharibe, 18.0 km 21° NNE Ambinanitelo, 15.1883°S, 49.615°E, 470 m, rainforest, 8.—12.I]].2003 (B.L. Fisher et al.); Toamasina, Mon- tagne d’Anjanaharibe, 19.5 km 27° NNE Ambinanitelo, 15.1783°S, 49.635°E, 1100 m, montane rainforest, 12.—16.II1.2003 (B.L. Fisher et al.); Toamasina, Reserve Bet- ampona, Camp Vohitsivalana, 37.1 km 338° Toamasina, 17.8867°S, 49.2025°E, 520 m, rainforest, 1.—3.XII.2005 (B.L. Fisher et al.); Toamasina, 19 km ESE Maroantsetra, 15.4833°S, 49.9°E, 350 m, rainforest, 22.1V.1989 (P.S. Ward); Toamasina, Mon- tagne d’Akirindro 7.6 km 341° NNW Ambinanitelo, 15.2883°S, 49.5483°E, 600 m, rainforest, 17.—21.III.2003 (B.L. Fisher et al.); Toamasina, Nosy Mangabe, 15.5°S, 49.766667°E, 300 m, rainforest, 18.IV.1989 (P.S. Ward); Toamasina, Ile Sainte Ma- rie, Forét Ambohidena, 22.8 km 44° Ambodifotatra, 16.8243°S, 49.9642°E, 20 m, littoral rainforest, 21.X1.2005 (B.L. Fisher et al.); Toamasina, F.C. Sandranantitra, 18.0483°S, 49.0917°E, 450 m, rainforest, 21.—24.1.1999 (HJ. Ratsirarson); Toamasi- na, Parc National de Zahamena, Tetezambatana forest, near junction of Nosivola and Manakambahiny Rivers, 17.743°S, 48.7294°E, 860 m, rainforest, 18.-19.II.2009 (B.L. Fisher et al.); Toamasina, Parc National de Zahamena, Onibe River, 17.7591°S, 48.8547°E, 780 m, rainforest, 21.—23.11.2009 (B.L. Fisher et al.). Diagnosis. Tetramorium dalek is easily distinguishable by the following combi- nation of characters: waist segments without long standing hairs, instead with short, appressed to subdecumbent pubescence only, sometimes with one or two short erect to suberect hairs; propodeal spines moderately long to long (PSLI 25—27); first gastral tergite with short, relatively dense, appressed to subdecumbent pubescence and with- out any standing hairs at all. Worker measurements (N=12). HL 0.47-0.56 (0.51); HW 0.45—0.54 (0.49); SE:0.31-0:35 (0:33); EL 011-0:13-(012); PH 0223-030 (026); PW 0:34-0:39 (0.36); WL 0.54—0.68 (0.61); PSL 0.12—0.15 (0.13); PTL 0.12-0.14 (0.12); PTH 0.19-0.22 (0.20); PTW 0.13—0.17 (0.15); PPL 0.13-0.16 (0.15); PPH 0.19-0.21 (0.19); PPW 0.18—-0.22 (0.20); CI 95-97 (96); SI 63-70 (67); OI 23-24 (24); DMI 54-63 (59); LMI 42—46 (43); PSLI 25-27 (26); PeNI 38-46 (42); LPeI 55-68 (61); DPel 108-139 (123); PpNI 53-59 (56); LPpI 70-80 (76); DPpI 131-147 (139); PPI 126-138 (133). Worker description. Head longer than wide (CI 95-97); posterior head mar- gin weakly concave. Anterior clypeal margin with distinct median impression. Frontal carinae strongly developed, diverging posteriorly, and usually approaching or ending at posterior head margin; antennal scrobe present, but weak, shallow, and without de- fined posterior or ventral margins. Antennal scapes short, not reaching posterior head margin (SI 63-70). Eyes of moderate size (OI 23-24). Mesosomal outline in profile flat to very weakly convex, relatively high (LMI 42-46), and moderately to strongly marginate from lateral to dorsal mesosoma; promesonotal suture and metanotal groove absent. Propodeal spines elongate-triangular to spinose, moderately long to long, and Taxonomy of Malagasy Tetramorium aes acute (PSLI 25-27); propodeal lobes short, triangular, and blunt or acute, always much shorter than propodeal spines. Petiolar node in profile high, rounded nodiform to weakly rectangular nodiform, with moderately rounded antero- and posterodorsal margins, around 1.5 to 1.8 times higher than long (LPel 55-68), anterior and poste- rior faces approximately parallel, anterodorsal and posterodorsal margins situated at about the same height and equally marginate (very rarely posterodorsal margin more rounded and lower than anterodorsal margin), petiolar dorsum flat to very weakly convex; node in dorsal view around 1.1 to 1.4 times wider than long (DPel 108-139), in dorsal view pronotum between 2.2 to 2.6 times wider than petiolar node (PeNI 38-46). Postpetiole in profile globular, approximately 1.2 to 1.4 times higher than long (LPpI 70-80); in dorsal view around 1.3 to 1.5 times wider than long (DPpl 131-147), pronotum between 1.7 to 1.9 times wider than postpetiole (PpNI 53-59). Postpetiole in profile appearing slightly more voluminous than petiolar node, postpe- tiole in dorsal view around 1.3 to 1.4 times wider than petiolar node (PPI 126-138). Mandibles distinctly striate; clypeus longitudinally rugose/rugulose, with three to five rugae/rugulae, median ruga always well developed and distinct, lateral rugae/rugu- lae sometimes weaker and interrupted; cephalic dorsum between frontal carinae with seven to ten longitudinal rugae, rugae running from posterior clypeal margin to poste- rior head margin, often interrupted or with cross-meshes, especially posteriorly; scrob- al area mostly unsculptured; lateral head longitudinally rugose to reticulate-rugose. Ground sculpture on head absent to weakly punctate. Mesosoma laterally and dorsally mostly irregularly longitudinally rugose. Forecoxae unsculptured, smooth and shining. Ground sculpture on mesosoma weak to absent. Waist segments and gaster completely unsculptured, smooth and shining. Head and mesosoma with numerous, moderately long and fine standing hairs; waist segments and first gastral tergite with short, com- paratively dense, appressed to subdecumbent pubescence, sometimes several of these short hairs suberect to erect. Anterior edges of antennal scapes and dorsal (outer) sur- faces of hind tibiae usually with decumbent to suberect hairs. Head, mesosoma, waist segments, and gaster generally orange-brown to chestnut brown, mandibles, antennae, and legs always lighter, usually yellowish brown. Etymology. The name of the new species is taken from the popular British TV show “Dr. Who” and refers to a fictional, extra-terrestrial race of evil mutants. During different stages of the revision we considered placing the material listed here as 7. dalek in at least three to four different groups, which caused a significant amount of nui- sance, especially to the first author. Naming this species after an evil, extra-terrestrial, and often annoying race was a logical consequence. ‘The species epithet is an arbitrary combination of letters, thus invariant. Distribution and biology. The new species is found in the lowland and montane rainforests of eastern Madagascar from the southernmost localities Sandranantitra, Betampona, and Zahamena, north to Anjanaharibe-Sud (Fig. 61). In addition, T- dalek has an elevational range from 20 to 1280 m, and seems to live in leaf litter. Discussion. Tetramorium dalek is easily distinguishable within the 7. naganum species group since it is the only species without long, standing hairs on the waist seg- 18 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) 2 4 on Figure 6. 7) dalek holotype worker (CASENT0038402). A Body in profile B Body in dorsal view C Head in full-face view. Taxonomy of Malagasy Tetramorium 19 ments and the first gastral tergite, and in addition, it also has generally shorter propo- deal spines (PSLI 25—27) than the other four species (PSLI 27-37). But it is possible to mistake T° dalek with species from other groups. Its general habitus and in particular its lack of standing pilosity on the first gastral tergite could lead to misplacement in the T. schaufussii complex of the T. schaufussii species group or the T: ibycterum complex in the 7. ranarum group. Indeed, a misidentification with one species from the latter complex is likely. Zetramorium ibycterum superficially shares many characters with T. dalek, and even most morphometric ranges. However, T° ibycterum has very well developed antennal scrobes with clearly defined margins all around, whereas T: dalek has weaker antennal scrobes without clearly defined margins all around. In addition, T. dalek differs from the species of the 7! schaufussii complex in having a broader head (CI 95-97) and higher and stouter mesosoma (LMI 42-46). Consequently, despite morphological similarities to other species groups, we consider T: dalek best placed in the 7. naganum group. Tetramorium enkidu Hita Garcia & Fisher, sp. n. http://zoobank.org/CEF594E4-5B03-42F9-B250-4295A137AAC6 http://species-id.net/wiki/Tetramorium_enkidu Figss2B,.5'B, 275.61 Type material. Holotype, pinned worker, MADAGASCAR, Antsiranana, Forét Ambanitaza, 26.1 km 347° Antalaha, 14.67933°S, 50.18367°E, 240 m, rainforest, sifted litter (leaf mold, rotten wood), collection code BLF10997, 26.X1.2004 (B.L. Fisher) (CAS: CASENT0056450). Paratypes, ten pinned workers with same data as holotype (BMNH: CASENT0056445; CAS: CASENT0056435; CASENT0056436; CASENT0056437; CASENT0056441; CASENT0056448; CASENT0056449; CASENT0056456; CASENT0056467; MCZ: CASENT0056461). Non-type material. Antsiranana, Forét Ambanitaza, 26.1 km 347° Antalaha, 14.6793°S, 50.1837°E, 240 m, rainforest, 26.X1.2004 (B.L. Fisher); Antsiranana, 1 km W Andampibe, Cap Masoala, 15.6936°S, 50.1814°E, 125 m, lowland rainforest, 1.X11.1993 (GD. Alpert); Antsiranana, Forét de Binara, 9.4km 235° SW Daraina, 13.2633°S, 49.6°E, 1100 m, montane rainforest, 5.XII.2003 (B.L. Fisher); Antsira- nana, Parc National Montagne d’Ambre, 3.6 km 235° SW Joffreville, 12.5344°S, 49.1795°E, 925 m, montane rainforest, 20.—26.1.2001 (B.L. Fisher et al.); Antsiranana, Parc National de Marojejy, Manantenina River, 27.6 km 35° NE Andapa, 9.6 km 327° NNW Manantenina, 14.435°S, 49.76°E, 775 m, 15.-18.XI.2003 (B.L. Fisher et al.); Toamasina, Montagne d’Akirindro, 7.6 km 341° NNW Ambinanitelo, 15.2883°S, 49 .5483°E, 600 m, rainforest, 17.-21.1]].2003 (B.L. Fisher et al.); Toamasina, 19 km ESE Maroantsetra, 15.4833°S, 49.9°E, 350 m, 22.1V.1989 (P.S. Ward). Diagnosis. Tetramorium enkidu is distinguishable from the other species of the group by the following combination of characters: eyes small to moderate in size (Ol 22-24); waist segments with several long erect hairs; propodeal spines long to very long 20 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) (PSLI 29-36); in profile petiolar node relatively thick, between 1.5 and 1.7 times higher than long (LPel 60-65); first gastral tergite with moderately short, abundant, subdecum- bent to suberect pilosity, and without short, dense, appressed to decumbent pubescence. Worker measurements (N=12). HL 0.53-0.62 (0.58); HW 0.50—0.60 (0.56); SL 0.33-0.43 (0.39); EL 0.11-0.14 (0.13); PH 0.25-0.34 (0.31); PW 0.36-0.46 (0.43); WL 0.59-0.77 (0.71); PSL 0.16-0.21 (0.19); PTL 0.13-0.16 (0.15); PTH 0.21-0.26 (0.24); PTW 0.14-0.18 (0.17); PPL 0.15-0.21 (0.19); PPH 0.20—0.26 (0.24); PPW 0.21-0.26 (0.24); CI 93-98 (96); SI 64—72 (69); OI 22-24 (23); DMI 58-62 (60); LMI 41-45 (43); PSLI 29-36 (32); PeNI 35-42 (39); LPel 60-65 (63); DPel 104-113 (109); PpNI 54-60 (57); LPpI 73-85 (79); DPpI 120-140 (131); PPI 142-155 (147). Worker description. Head weakly to distinctly longer than wide (CI 93-98); posterior head margin weakly concave. Anterior clypeal margin with distinct median impression. Frontal carinae strongly developed, diverging posteriorly, and usually ap- proaching or ending at posterior head margin; antennal scrobe present, but weak, shallow, and without defined posterior or ventral margins. Antennal scapes very short, not reaching posterior head margin (SI 64-72). Eyes short to moderate (OI 22-24). Mesosomal outline in profile weakly convex, relatively high (LMI 41-45), and mod- erately to strongly marginate from lateral to dorsal mesosoma; promesonotal suture and metanotal groove absent. Propodeal spines spinose, long to very long, and acute (PSLI 29-36); propodeal lobes short, triangular, and blunt or acute, always much shorter than propodeal spines. Petiolar node in profile high, rounded nodiform, with well-rounded antero- and posterodorsal margins, around 1.5 to 1.7 times higher than long (LPel 60-65), anterior and posterior faces approximately parallel, anterodorsal and posterodorsal margins situated at about the same height and equally marginate, petiolar dorsum always distinctly convex; node in dorsal view slightly wider than long (DPel 104-113), in dorsal view pronotum between 2.4 to 2.8 times wider than peti- olar node (PeNI 36-42). Postpetiole in profile globular, approximately 1.2 to 1.4 times higher than long (LPpI 73-85); in dorsal view between 1.2 to 1.4 times wider than long (DPpI 120-140), pronotum around 1.7 to 1.8 times wider than postpetiole (PpNI 54-60). Postpetiole in profile appearing slightly lower and thicker than petiolar node, postpetiole in dorsal view around 1.4 to 1.5 times wider than petiolar node (PPI 142-155). Mandibles usually unsculptured, smooth, and shining, sometimes weakly partially striate (especially basally), rarely fully covered in fine striations; clypeus longi- tudinally rugose/rugulose, with three to six rugae/rugulae, median ruga always well de- veloped and distinct, lateral rugae/rugulae usually weaker and/or interrupted; cephalic dorsum between frontal carinae with seven to nine longitudinal rugae, rugae running from posterior clypeal margin to posterior head margin, often irregular, interrupted or with cross-meshes, especially posteriorly; scrobal area mostly unsculptured; lateral head longitudinally rugose to reticulate-rugose. Ground sculpture on head weak to absent. Mesosoma laterally and dorsally irregularly longitudinally rugose, rarely lateral mesosoma with few unsculptured areas. Forecoxae unsculptured, smooth and shining. Ground sculpture on mesosoma very weak to absent. Waist segments and gaster com- Taxonomy of Malagasy Tetramorium 21 Figure 7. 7) enkidu holotype worker (CASENT0056450). A Body in profile B Body in dorsal view C Head in full-face view. 22 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) pletely unsculptured, smooth and shining. Head, mesosoma, and waist segments with numerous, long, and fine standing hairs; first gastral tergite with moderately short, abundant, subdecumbent to suberect pilosity, and without short, dense, appressed to decumbent pubescence; pilosity appearing disorganized due to varying degrees of inclination. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae with decumbent to suberect hairs. Head, mesosoma, waist segments, and gaster light to dark brown, mandibles, antennae, and legs always of lighter brown. Etymology. The new species is named after the fictional character “Enkidu” who is a central figure in the ancient Mesopotamian poem “Epic of Gilgamesh”, one of the oldest written stories in human history. The species epithet is an arbitrary combination of letters, thus invariant. Distribution and biology. The new species is restricted to the northern part of Madagascar. Its distribution ranges from Montagne d’Akirindro, the area around Maroantsetra, and Cap Masoala north through Ambanitaza, and Marojejy to Binara and Montagne d’Ambre (Fig. 61). ‘The localities are rainforests or montane rainforests situated at altitudes from 125 to 1100. In addition, 7: enkidu appears to live in leaf litter or the ground. Discussion. Tetramorium enkidu is easily identifiable within the species group. The presence of several long, erect hairs on the waist segments and much longer pro- podeal spines (PSLI 29-36) separate T: enkidu from T. dalek (PSLI 25-27). The latter species and 7. naganum both lack standing pilosity on the first gastral tergite, which is present in 7: enkidu. Tetramorium naganum also has a thinner petiolar node, which is between 1.7 to 1.9 times higher than long (LPel 54-58), contrasting with the thicker node of T° enkidu, which is between 1.5 and 1.7 times higher than long (LPel 60-65). Also distinguishable from T: enkidu by a relatively thin petiolar node is T: gilgamesh (LPel 50-58). The latter also possesses larger eyes (OI 25—27) than 7: enkidu (OI 22- 24), but this can be difficult to see without measuring. The last species of the group, T. alperti, shares the thicker petiolar node shape with 7! enkidu, as well as most other characters except gastral pilosity. Indeed, as outlined in the description of 7: alperti, both could be easily combined into one species since their separation is based only on differences in gastral pilosity/pubescence. However, we prefer to describe them as distinct because their distribution ranges overlap and both maintain a species-specific pattern of gastral pilosity/pubescence in sympatry. Tetramorium gilgamesh Hita Garcia & Fisher, sp. n http://zoobank.org/24769905-BB87-467 D-8785- 93586E3E8E65 http://species-id.net/wiki/Tetramorium_gilgamesh Figs 2D, 4A, 8, 61 Type material. Holotype, pinned worker, MADAGASCAR, Toamasina, F.C. Sandranantitra, 18.0483°S, 49.0917°E, 450 m, rainforest, sifted litter (leaf mold, rotten wood), collection code HJR101, 18.—21.1.1999 (AJ. Ratsirarson) (CAS: Taxonomy of Malagasy Tetramorium a CASENT0247312). Paratypes, four pinned workers with same data as holotype (CAS: CASENT0189097; CASENT0218015); and three pinned workers with same data as holotype except collected from the 21.—24.1.1999 and collection code HJR102 (CAS: CASENT0189096; CASENT0218016). Non-type material. MADAGASCAR: Antsiranana, 1 km W Andampibe, Cap Masoala, 15.6936°S, 50.1814°E, 125 m, lowland rainforest, 1.XII.1993 (G.D. Abp- ert); Toamasina, Montagne d’Akirindro 7.6 km 341° NNW Ambinanitelo, 15.2883°S, 49,5483°E, 600 m, rainforest, 17.—21.I11.2003 (BL. Fisher et al.); Toamasina, An- dranobe, 5.3 km SSE Ambanizana, 15.6713°S, 49.9739°E, 425 m, rainforest, 21.X1.1993 (B.L. Fisher); Toamasina, Réserve Spéciale Ambatovaky, Sandrangato river, 16.7727°S, 49.2655°E, 450 m, rainforest, 20.—22.]].2010 (B.L. Fisher et al.); Toamasi- na, Réserve Spéciale Ambatovaky, Sandrangato river, 16.7633°S, 49.2669°E, 520 m, rainforest, 22.11.2010 (B.L. Fisher et al.); Toamasina, Réserve Spéciale Ambatovaky, Sandrangato river, 16.8175°S, 49.295°E, 360 m, rainforest, 25.—-27.11.2010 (B.L. Fisher et al.); Toamasina, F.C. Andriantantely, 18.695°S, 48.8133°E, 530 m, rainforest, 7.-10. XIL.1998 (HJ. Ratsirarson); Toamasina, Montagne d’Anjanaharibe, 18.0 km 21° NNE Ambinanitelo, 15.1883°S, 49.615°E, 470 m, rainforest, 8.—12.I]].2003 (B.L. Fisher et al.); Toamasina, Reserve Betampona, Camp Vohitsivalana, 37.1 km 338° Toamasina, 17.8867°S, 49.2025°E, 520 m, rainforest, 1.—3.XII.2005 (B.L. Fisher et al.); Toamasi- na, Parc National Mananara-Nord, 7.1 km 261° Antanambe, 16.455°S, 49.7875°E, 225 m, rainforest, 14.X1.2005 (B.L. Fisher et al.); Toamasina, F.C. Sandranantitra, 18.0483°S, 49.0917°E, 450 m, rainforest, 18.—24.1.1999 (HJ. Ratsirarson). Diagnosis. ‘The following character combination distinguishes 7: gilgamesh from the other members of the 7° naganum group: relatively large eyes (OI 25-27); propo- deal spines long (PSLI 27-30); petiolar node relatively high and thin, around 1.7 to 2.0 times higher than long (LPel 50-59); waist segments with several long erect hairs; first gastral tergite with short to moderately long, abundant, decumbent to suberect pilosity, and without short, dense, appressed to subdecumbent pubescence; pilosity appearing disorganized due to varying degrees of inclination and hair length. Worker measurements (N=12). HL 0.52-0.56 (0.54); HW 0.49-0.55 (0.51); SL 0.36-0.38 (0.37); EL 0.13-0.14 (0.13); PH 0.26—0.30 (0.27); PW 0.36-0.41 (0.38); WL 0.61—0.66 (0.64); PSL 0.14—-0.17 (0.15); PTL 0.11-0.14 (0.12); PTH 0.21-0.24 (0.22); PI'W 0.13-0.17 (0.14); PPL 0.15-0.18 (0.16); PPH 0.19-0.22 (0.20); PPW 0.20—0.24 (0.22); CI 92-98 (94); SI 68-76 (73); OI 25-27 (26); DMI 58-62 (59); LMI 40-45 (42); PSLI 27-30 (28); PeNI 33-40 (37); LPel 50-59 (55); DPel 112-127 (118); PpNI 54-59 (56); LPpI 74-85 (80); DPpI 125-142 (132); PPI 143-169 (153). Worker description. Head weakly to distinctly longer than wide (CI 92-98); posterior head margin weakly concave. Anterior clypeal margin with distinct median impression. Frontal carinae strongly developed, diverging posteriorly, and usually ap- proaching or ending at posterior head margin; antennal scrobe distinct but relatively shallow, usually with defined margins all around, but ventral margin sometimes poor- ly defined and merging with surrounding sculpture, median scrobal carina generally 24 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) not well developed, if present relatively weak and reaching eye level only. Antennal scapes short, not reaching posterior head margin (SI 68—76). Eyes relatively large (Ol 25-27). Mesosomal outline in profile weakly convex, relatively high (LMI 40-45), and moderately to strongly marginate from lateral to dorsal mesosoma; promesonotal suture and metanotal groove absent. Propodeal spines spinose, long, and acute (PSLI 27-30); propodeal lobes short, triangular, and blunt or acute, always much shorter than propodeal spines. Petiolar node in profile high, rounded nodiform, with well- rounded antero- and posterodorsal margins, around 1.7 to 2.0 times higher than long (LPel 50-59), anterior and posterior faces not parallel, node slightly narrowing from base to apex, usually anterodorsal and posterodorsal margins situated at about same height (very rarely anterodorsal margin weakly higher than posterodorsal margin), petiolar dorsum weakly to moderately convex; node in dorsal view around 1.1 to 1.3 times wider than long (DPel 112-127), in dorsal view pronotum between 2.5 to 3.0 times wider than petiolar node (PeNI 33-40). Postpetiole in profile globular, ap- proximately 1.2 to 1.3 times higher than long (LPpI 74-85); in dorsal view between 1.2 to 1.4 times wider than long (DPpI 125-142), pronotum between 1.7 to 1.9 times wider than postpetiole (PpNI 54-59). Postpetiole in profile appearing thicker and lower than petiolar node, postpetiole in dorsal view around 1.4 to 1.7 times wider than petiolar node (PPI 143-169). Mandibles usually unsculptured, smooth, and shining, rarely with traces of longitudinal rugulae; clypeus longitudinally rugose/ rugulose, with three to six rugae/rugulae, median ruga always well developed and distinct, lateral rugae/rugulae usually weaker and/or interrupted; cephalic dorsum be- tween frontal carinae with eight to nine longitudinal rugae, rugae running from pos- terior clypeal margin to posterior head margin, some rugae irregular, interrupted or with cross-meshes, especially posteriorly; scrobal area mostly unsculptured, smooth and shining; lateral head mostly longitudinally rugose to reticulate-rugose, but with extensive unsculptured areas. Ground sculpture on head absent to weakly punctate. Mesosoma laterally and dorsally irregularly longitudinally rugose, sometimes lateral mesosoma with a few unsculptured areas medially. Forecoxae unsculptured, smooth and shining. Ground sculpture on mesosoma usually weak to absent. Waist segments and gaster completely unsculptured, smooth and shining. Head, mesosoma, and waist segments with numerous, long, and fine standing hairs; first gastral tergite with short to moderately long, abundant, decumbent to suberect pilosity, without short, dense, appressed to subdecumbent pubescence; pilosity appearing disorganized due to varying degrees of inclination and hair length. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae with decumbent to subdecumbent hairs. Head, mesosoma, waist segments, and gaster usually orange to light brown, rarely of darker brown, mandibles, antennae, and legs usually lighter, yellowish brown. Etymology. ‘The new species is named after the fictional character “Gilgamesh”, the main figure in the ancient Mesopotamian poem “Epic of Gilgamesh”, one of the earliest surviving works of literature. The species epithet is an arbitrary combination of letters, thus invariant. Taxonomy of Malagasy Tetramorium 25 Figure 8. 7. gilgamesh holotype worker (CASENT0247312). A Body in profile B Body in dorsal view C Head in full-face view. 26 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Distribution and biology. Tetramorium gilgamesh is distributed in eastern Madagascar (Fig. 61). Its distribution ranges from the southernmost known locality Andriantantely north to Montagne d’Anjanaharibe and Montagne d’Akirindro, and northeast to Andranobe and Andampibe on the Masoala Peninsula. All localities are lowland rainforests situated at elevations of 125 to 600 m. The preferred microhabitat of T. gilgamesh seems to be leaf litter. Discussion. The identification of 7: gilgamesh within the T: naganum species group is fairly straightforward. The best diagnostic character is eye size since T: gilgamesh has the largest eyes of the group with an OI 25-27 (vs. OJ 21—24 in the other four species). Beyond this, it cannot be mistaken for 7: dalek since the waist segments and a first gas- tral tergite of the latter species are covered with short, comparatively dense, appressed to subdecumbent pubescence without standing pilosity. By contrast, 7! gilgamesh has long, erect hairs on the waist segments and the first gastral tergite is covered with short to moderately long, abundant, decumbent to suberect pilosity, but without short, dense, appressed to subdecumbent pubescence. Additionally, the pilosity on the first gastral tergite of T: gilgamesh appears disorganized due to varying degrees of inclination and hair length. The gastral pilosity separates it also from T° alperti, T: enkidu, and T. naganum, which have very different patterns of pilosity/pubescence. Furthermore, 7! alperti and T. enkidu have thicker petiolar nodes (LPel 60-68) than 7: gilgamesh (LPel 50-59). Tetramorium naganum shares the same shape of the petiolar node with T°. gilgamesh, and both are also found in sympatry throughout most of their distribution ranges. However, differences in eye size and gastral pilosity distinguish between both species fairly well. On the basis of the available material it seems that intraspecific variation is gener- ally low in T. gilgamesh. Tetramorium naganum Bolton http://species-id.net/wiki/Tetramorium_naganum Figs 1B, 2C, 4B, 9, 61 Tetramorium naganum Bolton, 1979:150. Type material. Holotype, pinned worker, MADAGASCAR, Toamasina, La Mandraka, 18.912778°S, 47.892222 E 1280 m, montane forest, collection code AB41, 8.[I.1977 (W.L. & D.E. Brown) (MCZ: MCZ_Holotype_32379) [examined]. Paratypes, 16 pinned workers and one dealate queen with same data as holotype (BMNH: CASENT0102346; CASENT0235212; MCZ: MCZ_Paratype_32379) [examined]. [Note: the GPS data of the type locality was not provided by the locality label or the original description. The data presented above is based on our own georeferencing of the rainforest locality La Mandraka and should be considered an approximation and not the exact location of the type locality of T° naganum.| Non-type material. MADAGASCAR: Antsiranana, Rés. Anjanaharibe-Sud, 6.5 km SSW Befingotra, 14.75°S, 49.5°E, 875 m, rainforest, 19.X.1994 (B.L. Fisher); Taxonomy of Malagasy Tetramorium 2F Antsiranana, Rés. Anjanaharibe-Sud, 9.2 km WSW Befingotra, 14.75°S, 49.4667°E, 1280 m, montane rainforest, 5.XI.1994 (B.L. Fisher); Antananarivo, 3 km 41° NE Andranomay, 11.5 km 147° SSE Anjozorobe, 18.4733°S, 47.96°E, 1300 m, 5.-13. XII.2000 (B.L. Fisher et al.); Toamasina, 6.9 km NE Ambanizana, Ambohitsitondroi- na, 15.5851°S, 50.0095°E, 825 m, rainforest, 2.XII.1993 (B.L. Fisher); Toamasina, Ambanizana, Parc National Masoala, 15.5722°S, 50.0069°E, 1020 m, montane rain- forest, 2.—6.I1.2003 (D. Andriamalala et al.); Toamasina, Ambatoharanana, Corri- dor Forestier Analamay-Mantadia, 18.8042°S, 48.4008°E, 968 m, 12.-19.XII.2012 (B.L. Fisher et al.); Toamasina, Analamay, 18.8062°S, 48.3371°E, 1068 m, montane rainforest, 21.I11.2004 (B.L. Fisher); Toamasina, Ankerana, 18.4017°S, 48.806°E, 1035 m, montane forest, 24.-29.1.2012 (B.L. Fisher et al.); Toamasina, Montagne d’Anjanaharibe, 19.5 km 27° NNE Ambinanitelo, 15.17833°S, 49.635°E, 1100 m, montane rainforest, 12.—16.III.2003 (B.L. Fisher). Diagnosis. Tetramorium naganum can be easily diagnosed within the T. naganum group on the basis of the following character combination: eyes small to moderate in size (OI 21—23); propodeal spines relatively long (PSLI 28-33); petiolar node in profile relatively thin, between 1.7 to 1.9 times higher than long (LPel 54—58); waist segments with long, standing hairs; first gastral tergite with short, comparatively dense, appressed to decumbent pubescence, and without any long standing hairs. Worker measurements (N=10). HL 0.57-0.72 (0.63); HW 0.59-0.73 (0.65); SL 0.39-0.51 (0.46); EL 0.13-0.16 (0.14); PH 0.29-0.40 (0.33); PW 0.41-0.53 (0.45); WL 0.73-0.92 (0.78); PSL 0.18—0.22 (0.20); PTL 0.13-0.16 (0.14); PTH 024-028: (0225)* (PAW 0215-0:192 (0:16) PPE 0:18-0:22: (0219)6PPH 025=0:27 (0.24); PPW 0.23-0.28 (0.25); CI 94-99 (97); SI 68-77 (73); OI 21-23 (22); DMI 56-62 (58); LMI 40-45 (43); PSLI 28-33 (30); PeNI 34-38 (35); LPel 54-58 (56); DPel 107-122 (115); PpNI 52-59 (54); LPpI 77-84 (80); DPpI 125-137 (128); PPI 147-159 (153). Worker description. Head weakly to distinctly longer than wide (CI 94-99); posterior head margin moderately concave. Anterior clypeal margin with distinct me- dian impression. Frontal carinae strongly developed, diverging posteriorly, and usu- ally approaching or ending at posterior head margin; antennal scrobe present, but weak, shallow, and without defined posterior or ventral margins. Antennal scapes short, not reaching posterior head margin (SI 68-77). Eyes short to moderate (Ol 21-23). Mesosomal outline in profile weakly convex, relatively high (LMI 40-45), and moderately to strongly marginate from lateral to dorsal mesosoma; promesonotal suture and metanotal groove absent. Propodeal spines spinose, long, and acute (PSLI 28-33); propodeal lobes short, triangular, and blunt or acute, always much shorter than propodeal spines. Petiolar node in profile high, rounded nodiform, with well- rounded antero- and posterodorsal margins, between 1.7 to 1.9 times higher than long (LPel 54-58), anterior and posterior faces approximately parallel, anterodorsal and posterodorsal margins situated at about the same height (very rarely anterodorsal mar- gin higher than posterodorsal margin) and equally marginate, petiolar dorsum always convex; node in dorsal view around 1.1 to 1.2 times wider than long (DPel 107-122), 28 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) in dorsal view pronotum between 2.7 to 3.0 times wider than petiolar node (PeNI 34-38). Postpetiole in profile globular, approximately 1.2 to 1.3 times higher than long (LPpI 77-84); in dorsal view between 1.2 to 1.4 times wider than long (DPpl 125-137), pronotum around 1.7 to 1.9 times wider than postpetiole (PpNI 52-59). Postpetiole in profile thicker and lower than petiolar node, postpetiole in dorsal view around 1.5 to 1.6 times wider than petiolar node (PPI 147-159). Mandibles variably sculptured, ranging from fully unsculptured, smooth, and shining through partially striate to fully striate; clypeus longitudinally rugose/rugulose, with two to six rugae/ rugulae, median ruga always well developed and distinct, lateral rugae/rugulae usually weaker and/or interrupted; cephalic dorsum between frontal carinae with six to nine longitudinal rugae, rugae running from posterior clypeal margin to posterior head margin but very often irregular, interrupted or with cross-meshes, especially posteri- orly; scrobal area usually mostly unsculptured, rarely longitudinally rugose to reticu- late-rugose; lateral head longitudinally rugose to reticulate-rugose. Ground sculpture on head absent to weakly punctate. Mesosoma laterally and dorsally irregularly lon- gitudinally rugose, rarely lateral mesosoma with a few unsculptured areas medially. Forecoxae unsculptured, smooth and shining. Ground sculpture on mesosoma very weak to absent. Waist segments and gaster completely unsculptured, smooth and shin- ing. Head, mesosoma, and waist segments with numerous, long, and fine standing hairs; first gastral tergite with short, comparatively dense, appressed to subdecumbent pubescence. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae usually with decumbent to suberect hairs. Head, mesosoma, waist segments, and gaster usually orange to light brown, rarely of darker brown, mandibles, antennae, and legs always lighter, usually light yellowish brown. Distribution and biology. Tetramorium naganum is found in rainforests or mon- tane rainforests in eastern and north-eastern Madagascar (Fig. 61). The distribution range is disjunctive, very much as in T° alperti and T. dalek. T. naganum is either found in the region around La Mandraka, Analamay, Andranomay, and Andasibe- Mantadia, and Ankerana, or much further north between Anjanaharibe-Sud and the Masoala Peninsula (Ambanizana), with Montagne d’Anjanaharibe being in-between. The reasons behind this discontinuous distribution are not clear yet, but as for T: alperti, it is likely that T. naganum was previously much more common in eastern Malagasy humid forests, and present-day populations represent only relict popula- tions. The altitudinal range of the species with 825-1300 m supports this. However, since T° naganum is not particularly abundant or common where it occurs, it may just be a relatively rare faunal element in eastern Madagascar. If so, further sampling might yield additional material in the future, especially in the geographic areas between the two main populations mentioned above. Like most species in this group, 7: naganum seems to be a leaf litter inhabitant. Discussion. Tetramorium naganum is the only species of the group that was known prior to our revision, and can be seen as the core species of the group. The lack of pi- losity on the first gastral tergite isolates it fairly well from T°: alperti and T. gilgamesh since these possess pilosity in varying degrees of inclination, length, and abundance. Taxonomy of Malagasy Tetramorium 29 Figure 9. 7) naganum holotype worker (CASENT0280584). A Body in profile B Body in dorsal view C Head in full-face view. 30 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) The relatively thin and high petiolar node (LPel 54—58) separates T’ naganum from T. alperti and T. enkidu, which have much thicker petiolar nodes (LPel 60-68). Ad- ditionally, 7! naganum has smaller eyes (OI 21-23) than T! gilgamesh (OI 25-27). The last species of the group, T: dalek, shares the lack of long, standing pilosity on the first gastral tergite with 7° naganum. Nevertheless, both species are very unlikely to be confused with each other. Tetramorium dalek is generally smaller (HW 0.45-0.54; WL 0.54-68), has shorter propodeal spines (PSLI 25-27), and lacks long, standing hairs on the waist segments, whereas 7. naganum is generally larger (HW 0.55-0.72; WL 0.66—0.92), possesses longer propodeal spines (PSLI 28-33), and always has long, standing hairs on the waist segments. Revision of the Tetramorium plesiarum species group Tetramorium plesiarum species group Diagnosis. Eleven-segmented antennae; anterior clypeal margin medially impressed; frontal carinae strongly developed and forming dorsal margin of very well-developed antennal scrobes, scrobes moderately to very deep and with clearly defined margins all around (except in T° gollum); median scrobal carina very well developed and distinctly surpassing posterior eye level, usually ending between posterior eye margin and pos- terior scrobe margin, often approaching the latter; anterior face of mesosoma weakly developed; mesosomal outline in profile moderately to strongly convex, moderately marginate from lateral to dorsal mesosoma; mesosoma relatively high (LMI 41-56); propodeal spines usually medium-sized to long, elongate-triangular to spinose (PSLI 23-32), rarely short (PSLI 18-20); propodeal lobes usually short, triangular to elon- gate-triangular, and acute; petiolar node in profile nodiform to high nodiform, in profile between 1.3 to 1.8 times higher than long, (LPel 56—76), anterior and posterior faces generally parallel, anterodorsal and posterodorsal margins situated at about same height; node in dorsal view typically distinctly wider than long, between 1.1 to 1.5 times (DPel 108-148); postpetiole much broader than long and transverse, between 1.4 to 2 times broader than long (DPpI 141-200); sculpture on mandibles variably developed; cephalic sculpture distinct, between frontal carinae predominantly longitu- dinally rugose; mesosoma with well-developed longitudinally rugose sculpture; waist segments weakly to moderately sculptured, never completely unsculptured; first gastral tergite usually with weak ground sculpture at base or completely unsculptured, in T° gollum basal half of tergite strongly reticulate-rugose; whole body with abundant, usu- ally dense, long, and standing hairs; sting appendage spatulate. Comments. The 7: p/esiarum group is a compact assemblage of five species that resemble one another very closely. All are morphologically very conspicuous elements within the Malagasy Tetramorium fauna, and represent arid-adapted species found mostly in the drier western and southern parts of Madagascar. Surprisingly, 7. ple- starum, which is much less common than TJ. bressleri, T. hobbit, and T. mars, is the Taxonomy of Malagasy Tetramorium Cul *% j La mu n | Figure 10. Mesosoma and petiole in profile. A TZ hobbit (CASENT0019207) B Z mars (CASENT0474279) C T’ plesiarum (CASENT0036474) D T gollum (CASENT0074974) E TU. bressleri (CASENT0055196). only member of the species group that was known prior to this revision, and only from the holotype. The other four species, 7: bressleri, T. gollum, T. hobbit, and T. mars, are newly described here. All five species share a more or less similar habitus with very well developed antennal scrobes, more or less convex mesosomal profiles, usually medium- sized to long propodeal spines, higher than long and broader than long petiolar nodes, and abundant, long, and often dense pilosity. The species delimitations presented here are mainly based on differences in the shape of the petiolar node and sculpture on dif- ferent parts of the body. One intriguing feature of the group is the morphological cline observable in the shape of the petiolar node (see Fig. 10), which ranges from massively enlarged and blocky (7: /obbit) through smaller, but still relatively blocky (7° mars), to much higher and thinner (7° bressleri, T: gollum, T. plesiarum). Tetramorium plesiarum was initially placed in the 7: ranarum species group by Bolton (1979) mainly on the basis of petiolar node shape and pilosity. However, in Hita Garcia and Fisher (2012a) we proposed a T: plesiarum species group based on the very conspicuous antennal scrobes present in all species of the group, a character absent in most other Malagasy Tetramorium. We still think that the T. plesiarum spe- cies group presents a well distinguishable grouping, although the conspicuous antennal scrobes unfortunately do not separate it from all other Malagasy groups. Several spe- cies within the 7: ranarum group (e.g. T. zenatum, T. ibycterum, and two additional, yet-undescribed species) also have distinct antennal scrobes, although in these species the margins around the scrobes are much less distinctive and the scrobes themselves shallower than in the T° plesiarum group. However, the species of the 7. ranarum group with a scrobe 1) either lack any long, suberect to erect pilosity on the first gastral tergite (7° ibycterum and another undescribed species here listed as T: fhg-bilb), or 2) they have long, subdecumbent pilosity (e.g. one yet-undescribed species listed here as T. fhg-vazi), or 3) if they possess long, standing pilosity, the petiolar node is much longer than broad (7° zenatum). Therefore they cannot be confused with any species of the T° plesiarum group. Identification key to species of the T’ plesiarum species group (workers) 1 Gaster conspicuously enlarged, appearing swollen; basal half of first gastral tergite strongly sculptured (Fig. 11A) [Madagascar]... eee T. gollum = Gaster never enlarged or swollen; first gastral tergite not strongly sculptured; if sculpture present, then restricted to base of the tergite (Fig. 11B, C)....... 2 32 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) a Petiolar node blocky, massive and very large; head, mesosoma, and waist segments with conspicuous reticulate-punctate ground sculpture (Fig. 12A) [Middle caseat ie accs cassette teisectte seve teeta ustetithde Sieies deve den cvaldtnen T. hobbit = Petiolar node always much smaller than above; head, mesosoma, and waist segments without conspicuous reticulate-punctate ground sculpture (Fig. 2B eeiGe a ner. tee Sie enna, Niece, Neale, Somer, Ue, NUR, Nee oe, Nema, NA 3 3 Antennal scapes shorter (SI 58-62); petiolar node in profile lower and thicker and in dorsal view longer (LPel 69-76; DPel 108-123); dorsum of peti- olar node fully covered with distinctly reticulate-rugose sculpture (Fig. 13A) [Madagascan bs. seesscmcarssammaes ren secre er cserarseesseS, GIZs o e scarce T. mars - Antennal scapes longer (SI 62—69); petiolar node in profile higher and thin- ner and in dorsal view broader (LPel 56-63; DPel 130-144); dorsum of petiolar node weakly rugulose/rugose, mostly smooth (Fig. 13B, C)........... 4 4 Generally smaller species (HW 0.67—-0.76; WL 0.81—092); eyes relatively larger (OI 21-23); in profile anterodorsal margin of petiolar node clearly protruding anteriorly; mesopleuron and lateral propodeum longitudinally ru- gose with distinct reticulate-punctate ground sculpture, appearing matte and rough; sides of petiolar node with strong reticulate-punctate ground sculpture, appearing fairly matte and rough (Fig. 14A) [Madagascar.......... T. plesiarum — Generally larger species (HW 0.80—1.00; WL 0.92-1.15); eyes relatively smaller (OI 18-19); in profile anterodorsal margin of petiolar node not pro- truding anteriorly; mesopleuron and lateral propodeum with very little ru- gose/rugulose sculpture, mostly unsculptured, smooth, and shining; sides of petiolar node with weak but distinct reticulate-punctate ground sculpture, appearing only slightly matte and mostly smooth and shining (Fig. 14B) [Mata cases Me rrem caesarean ar =.coet crane ceases aca T. bressleri Figure | 1. Body in profile. A 7’ gollum (CASENT0074849) B T° plesiarum (CASENT0172831) C 7 bressleri (CASENT0055196). a rr ay : 7 His =e Figure 12. Head, mesosoma and petiole in profile. A 7 hobbit (CASENT0019207) B T mars (CASENT0474279) € T. bressleri (CASENT0055196). Taxonomy of Malagasy Tetramorium 33 a Figure 13. Mesosoma and petiole in profile. A 7 mars (CASENT0474279) B T° plesiarum (CASENT0172831) C TZ. bressleri (CASENT0055196). A Tetramorium bressleri Hita Garcia & Fisher, sp. n. http://zoobank.org/288AA2 1 B-2B9E-49F 1-8C6A-B22EGFD4619E http://species-id.net/wiki/Tetramorium_bressleri Figs 10E, AC, AZ esis 1AB 5562 Type material. Holotype, pinned worker, MADAGASCAR, Mahajanga, Parc Nation- al de Namoroka, 9.8 km 300° WNW Vilanandro, 16.46667°S, 45.35°E, 140 m, tropi- cal dry forest, sifted litter (leaf mold, rotten wood), collection code BLF06446, 4.-8. X1.2002 (B.L. Fisher et al.) (CAS: CASENT0035677). Paratypes, 27 pinned workers with same data as holotype (BMNH: CASENT0035669; CAS: CASENT0035636; CASENT0035640; CASENT0035642; CASENT0035647; CASENT0035648; CASENT0035649; CASENT0035652; CASENT0035653; CASENT0035654; CASENT0035656; CASENT0035658; CASENT0035660; CASENT0035662; CASENT0035666; CASENT0035667; CASENT0035670; CASENT0035674; CASENT0035675; CASENT0035678; CASENT0035681; CASENT0035686; CASENT0035687; CASENT0035689; CASENT0035694; CASENT0035703; MCZ: CASENT0035646). Non-type material MADAGASCAR: Antananarivo, Reserve Speciale d’Ambohitantely, Forét Ambohitantely, 20.9 km 72°NE Ankazobe, 18.2253°S, 47 .2868°E, 1410 m, montane rainforest, 17.-22.1V.2001 (B.L. Fisher et al.); Antsira- nana, Forét d’Anabohazo, 21.6 km 247° WSW Maromandia, 14.3089°S, 47.9143°E, 120 m, tropical dry forest, 11.-16.1H.2001 (BL. Fisher et al.); Fianarantsoa, Am- pangabe I Non Protected Area, 21.4 km W Itremo, 20.6111°S, 46.6069°E, 1414 m, savannah woodland, 21.—23.III.2010 (A. Ravelomanana); Fianarantsoa, Ampangabe II Non Protected Area, 21.29 km W Itremo, 20.6114°S, 46.6081°E, 1402 m, savannah woodland, 21.—23.II.2010 (A. Ravelomanana); Fianarantsoa, Ampangabe VI Non 34 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Protected Area, 21.16 km W Itremo, 20.6144°S, 46.6104°E, 1379 m, shrubland, 21.— 23.111.2010 (A. Ravelomanana); Fianarantsoa, Ampangoabe V Non Protected Area, 21.37 km W Itremo, 20.6136°S, 46.608°E, 1449 m, shrubland, 22.—23.III.2010 (A. Ravelomanana); Fianarantsoa, Ampotoampoto I National Parc, 8.02 km NW Ilaka- ka, 22.6283°S, 45.1886°E, 917 m, savannah woodland, 26.—28.II.2010 (A. Ravelo- manana); Fianarantsoa, Forét d’Analalava, 29.6 km 280° W Ranohira, 22.5917°S, 45.1283°E, 700 m, 1.-5.1.2003 (B.L. Fisher et al.); Fianarantsoa, Antohatsahomby I Non Protected Area, 22.77 km NW Ambatofinandrahana, 20.5506°S, 46.5856°E, 1550 m, Uapaca woodland, 15.-17.11.2010 (A. Ravelomanana); Fianarantsoa, Forét d Atsirakambiaty, 7.6 km 285° WNW Itremo, 20.5933°S, 46.5633°E, 1550 m, mon- tane rainforest, 22.—26.1.2003 (B.L. Fisher et al.); Fianarantsoa, Parc National d’Isalo, 9.1 km 354° N Ranohira, 22.4817°S, 45.4617°E, 725 m, gallery forest, 27.-31.1.2003 (B.L. Fisher et al.); Fianarantsoa, Parc National d’Isalo, Ambovo Springs, 29.3 km 4° N Ranohira, 22.2983°S, 45.3517°E, 990 m, Uapaca woodland, 9.-14.11.2003 (B.L. Fisher et al.); Fianarantsoa, Mampiarika I Non Protected Area, 28.08 km SW Ambosi- tra, 20.7344°S, 47.0835°E, 1480 m, Uapaca woodland, 31.1.—2.11.2010 (A. Ravelo- manana); Mahajanga, Forét Ambohimanga, 26.1 km 314° Mampikony, 15.9627°S, 47.4382°E, 250 m, tropical dry forest, 13.-15.XII.2004 (B.L. Fisher); Mahajanga, Boeny Region, District of Soalala, Analamanitra forest, 14 km SW Mitsinjo, 16.7°S, 45.7°E, 19 m, dense dry forest, 26.11.-4.111.2008 (AZ. Rin ha); Mahajanga, Parc Na- tional d’Ankarafantsika, Ampijoroa Station Forestiere, 40 km 306° NW Andrano- fasika, 16.3208°S, 46.8107°E, 130 m, tropical dry forest, 26.HI.-1.[V.2001 (B.L. Fisher et al.); Mahajanga, Parc National d’Ankarafantsika, Ampijoroa Station Fores- tiere, 5.4 km 331° NW Andranofasika, 16.2989°S, 46.813°E, 70 m, tropical dry for- est, 26.H1.-1.1V.2001 (B.L. Fisher et al.); Mahajanga, Parc National d’Ankarafantsika, Forét de Tsimaloto, 18.3 km 46° NE de Tsaramandroso, 16.2281°S, 47.1436°E, 135 m, tropical dry forest, 2.1V.-8.1V.2001 (B.L. Fisher et al.); Mahajanga, Parc National de Baie de Baly, 12.4 km 337° NNW Soalala, 16.01°S, 45.265°E, 10 m, tropical dry forest, 26.-30.X1.2002 (B.L. Fisher et al.); Mahajanga, Reserve Forestiere Beanka, 50.2 km E Maintirano, 18.0265°S, 44.0505°E, 250 m, tropical dry forest on tsingy, 19.-26.X.2009 (B.L. Fisher et al.); Mahajanga, Reserve Forestiere Beanka, 52.7 km E Maintirano, 18.0622°S, 44.5259°E, 300 m, tropical dry forest on tsingy, 24.-27.X.2009 (B.L. Fisher et al.); Mahajanga, Reserve Forestiere Beanka, 50.7 km E Maintirano, 17.8802°S, 44.4688°E, 140 m, tropical dry forest on tsingy, 29.X.-1. X1.2009 (B.L. Fisher et al.); Mahajanga, Reserve Forestiere Beanka, 50.2 km E Mainti- rano, 17.8876°S, 44.4726°E, 153 m, tropical dry forest on tsingy, 31.X.2009 (B.L. Fisher et al.); Mahajanga, Reserve Speciale de Bemarivo, 23.8 km 223° SW Besala- mpy, 16.925°S, 44.3683°E, 30 m, tropical dry forest, 19.-23.X1.2002 (B.L. Fisher et al.); Mahajanga, Mahavavy River, 6.2 km 145° SE Mitsinjo, 16.0517°S, 45.9083°E, 20 m, gallery forest, 1.—5.XII.2002 (B.L. Fisher et al.); Mahajanga, Parc National de Namoroka, 9.8 km 300° WNW Vilanandro, 16.4667°S, 45.35°E, 140 m, tropical dry forest, 4.-8.X1.2002 (B.L. Fisher et al.); Mahajanga, Parc National de Namoroka, 17.8 km 329° WNW Vilanandro, 16.3767°S, 45.3267°E, 100 m, tropical dry for- Taxonomy of Malagasy Tetramorium 35 est, 8.-12.X1.2002 (B.L. Fisher et al.); Mahajanga, Parc National de Namoroka, 16.9 km 317° NW Vilanandro, 16.4067°S, 45.31°E, 100 m, tropical dry forest, 12-16. X1.2002 (B.L. Fisher et al.); Mahajanga, Parc National Tsingy de Bemaraha, 3.4 km 93° E Bekopaka, Tombeau Vazimba, 19.1419°S, 44.828°E, 50 m, tropical dry for- est, 6.-10.X1.2001 (B.L. Fisher et al.); Mahajanga, Parc National Tsingy de Bemara- ha, 2.5 km 62° ENE Bekopaka, Ankidrodroa River, 19.1322°S, 44.8147°E, 100 m, 11.-15.X1.2001 (B.L. Fisher et al.); Mahajanga, Parc National Tsingy de Bemaraha, 10.6 km ESE 123° Antsalova, 18.7094°S, 44.7182°E, 150 m, tropical dry forest on tsingy, 16.—20.X1.2001 (B.L. Fisher et al.); Toliara, Parc National d’Andohahela, Forét d’Ambohibory, 1.7 km 61° ENE Tsimelahy, 36.1 km 308° NW Tolagnaro, 24.93°S, 46.6455°E, 300 m, tropical dry forest, 16.—20.1.2002 (B.L. Fisher et al.); Toliara, An- dohahela National Park, Tsimelahy, 24.9368°S, 46.6267°E, 180 m, transition for- est, 16.—-26.11.2003 (AE. Irwin et al.); Toliara, 18 km NNW Betroka, 23.1633°S, 45.9686°E, 825 m, savannah, 24.X].-4.X11.1994 (A.A. lvie e& D. A. Pollock); Toliara, Atsimo Andrefana Region, District of Betioky, 30 km E Betioky, Beza Mahafaly Spe- cial Reserve, 23.6865°S, 44.591°E, 165 m, gallery dry deciduous forest, 1.-7.1.2002 (M. Rinha); Toliara, Beza-Mahafaly, 27 km E Betioky, 23.65°S, 44.6333°E, 135 m, tropical dry forest, 23.1V.1997 (B.L. Fisher); Toliara, Fiherenana, 23.1769°S, 43.9608°E, 100 m, gallery forest, 21.—24.X.2002 (Frontier Project); Toliara, Fiherena- na, 23.177°S, 43.9614°E, gallery forest, 18.—19.VII.2003 (Frontier Wilderness Project); Toliara, southern Isoky-Vohimena Forest, 59 km NE Sakaraha, 22.4667°S, 44.85°E, 730 m, tropical dry forest, 21.1.1996 (B.L. Fisher); Toliara, Makay Mts., 21.2098°S, 45.3418°E, 525 m, gallery forest on sandy soil, 27.XI.-2.XII.2010 (B.L. Fisher et al.); Toliara, Makay Mts., 21.2199°S, 45.324°E, 500 m, gallery forest on sandy soil, 24.X1.-12.X11.2010 (B.L. Fisher et al.); Toliara, Makay Mts., 21.31°S, 45.1295°E, 590 m, dry forest on sandy soil, 3.-6.XII.2010 (B.L. Fisher et al.); Toliara, Vohibasia For- est, 59 km NE Sakaraha, 22.4667°S, 44.85°E, 780 m, tropical dry forest, 13.1.1996 (B.L. Fisher); Toliara, Parc National de Zombitse, near road, 22.8405°S, 44.7312°E, 825 m, spiny deciduous forest, 15.X.2001-23.H1.2002 (R. Harin’Hala); Toliara, Parc National de Zombitse, near ANGAP office, 22.8865°S, 44.6922°E, 840 m, deciduous spiny forest, 9.XI.2001-26.1.2002 (R. Harin’Hala); Toliara, Parc National de Zom- bitse, 19.8 km 84° E Sakaraha, 22.8433°S, 44.71°E, 770 m, tropical dry forest, 5.—9. 11.2003 (B.L. Fisher et al). Diagnosis. Tetramorium bressleri can be recognised by the following combination of characters: larger species (HW 0.80—1.00; WL 0.92-1.15); eyes relatively small (Ol 18-19); petiolar node high nodiform, not blocky and massively enlarged, anterodorsal and posterodorsal margins at about the same height and equally marginate, anterodor- sal margin not protruding anteriorly nor very sharply angled; petiolar node in profile relatively high and thin, between 1.6 to 1.8 times higher than long (LPel 56-61), in dorsal view between 1.3 to 1.5 times wider than long (DPel 135-145); gaster never extremely enlarged and swollen; head and mesosoma without strongly developed and conspicuous reticulate-punctate ground sculpture; usually sculpture on the mesopleu- ron and lateral propodeum mostly absent; basal half of first gastral tergite not strongly 36 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) reticulate-rugose, only base of tergite weakly sculptured; pilosity on first gastral tergite mostly erect. Worker measurements (N=12). HL 0.81—1.00 (0.94); HW 0.80—1.00 (0.94); SL 0.51—-0.64 (0.60); EL 0.15—0.19 (0.18); PH 0.42—0.53 (0.48); PW 0.57-0.73 (0.68); WL 0.92-1.15 (1.07); PSL 0.24—0.32 (0.27); PTL 0.20—0.26 (0.23); PTH 0.35-0.42 (0.40); PI'W 0.28—0.36 (0.32); PPL 0.23—-0.30 (0.27); PPH 0.32-0.41 (0.38); PPW 0.35-0.45 (0.42); CI 98-102 (100); SI 62-65 (64); OI 18-19 (19); DMI 61-66 (64); LMI 43-47 (45); PSLI 26-32 (28); PeNI 45—49 (47); LPel 56-61 (58); DPel 135-145 (138); PpNI 59-63 (61); LPpI 70-75 (73); DPpI 150-158 (152); PPI 125-134 (130). Worker description. Head more or less as long as broad (CI 98-102); posterior head margin weakly concave. Anterior clypeal margin with distinct median impression. Frontal carinae strongly developed and forming dorsal margin of very well-developed antennal scrobes, scrobes moderately to very deep and with clearly defined margins all around; median scrobal carina very well developed and distinctly surpassing posterior eye level, usually ending halfway between posterior eye margin and posterior scrobe margin. Antennal scapes short, not reaching posterior head margin (SI 62-65). Eyes relatively small (OI 18-19). Mesosomal outline in profile weakly to moderately con- vex, rounded and high (LMI 43-47), moderately to strongly marginate from lateral to dorsal mesosoma; promesonotal suture and metanotal groove absent. Propodeal spines elongate-triangular to spinose, long, and acute (PSLI 26-32), propodeal lobes short, triangular, and acute, always much shorter than propodeal spines. Petiolar node in profile high, rectangular nodiform with well defined antero- and posterodorsal mar- gins, between 1.6 to 1.8 times higher than long (LPel 56-61), anterior and poste- rior faces approximately parallel, anterodorsal and posterodorsal margins situated at about the same height, petiolar dorsum flat to weakly convex, anterodorsal margin not protruding anteriorly; node in dorsal view between 1.3 to 1.5 times wider than long (DPel 135-145), in dorsal view pronotum between 2.0 to 2.2 times wider than petiolar node (PeNI 45-49). Postpetiole in profile subglobular and weakly anteropos- teriorly compressed, approximately 1.3 to 1.4 times higher than long (LPpI 70-75); in dorsal view around 1.5 to 1.6 times wider than long (DPpI 150-158), pronotum between 1.6 to 1.7 times wider than postpetiole (PpNI 59-63). Postpetiole in profile appearing slightly less voluminous than petiolar node, postpetiole in dorsal view be- tween 1.2 to 1.4 times wider than petiolar node (PPI 125-134). Mandibles variably sculptured, either unsculptured, smooth, and shining, or partly or fully finely rugu- lose; clypeus longitudinally rugose/rugulose, with five to eight distinct rugae, median ruga always distinct, lateral rugae often weaker and sometimes interrupted; cephalic dorsum between frontal carinae with ten to thirteen longitudinal rugae, rugae running from posterior clypeal margin to posterior head margin, rarely interrupted or with cross-meshes; scrobal area mostly unsculptured, smooth and shiny; lateral head mainly longitudinally rugose. Ground sculpture on head usually weak to absent. Dorsum of mesosoma mostly longitudinally rugose without any distinct ground sculpture, spaces between rugae smooth and shining; lateral pronotum irregularly longitudinally rugose, often with weak to moderate punctate ground sculpture, mesopleuron and lateral pro- Taxonomy of Malagasy Tetramorium 37 podeum usually with very little sculpture, few irregular rugae/rugulae and no ground sculpture. Forecoxae often with weak ground sculpture, but generally very smooth and shining. Petiolar node laterally with conspicuous but relatively weak reticulate- punctate ground sculpture only, appearing weakly matte but still relatively smooth and shiny; dorsum of node medially almost unsculptured, smooth, and shiny, surrounding areas rugulose, ground sculpture on petiolar dorsum neglectable. Postpetiole laterally and dorsally weakly to moderately rugose/rugulose and with conspicuous, moderate reticulate-punctate ground sculpture, appearing relatively matte; base of first gastral tergite usually weakly punctate and/or costulate and/or shagreened, remainder of ter- gite unsculptured, smooth, and shining. Whole body with abundant, long, and fine standing hairs; first gastral tergite with abundant, long, erect hairs and much scarcer, shorter, decumbent to subdecumbent hairs. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae with decumbent to suberect hairs. Body usually of uniform brown, appendages often lighter. Etymology. The name of the new species is a patronym dedicated to Dr. Barry Lee Bressler, retired physicist, former adjunct professor of physics at Virginia Polytechnic Institute and State University, and amateur naturalist, in recognition of his interest in myrmecology and his support of ant taxonomy. Distribution and biology. As mentioned above, it is surprising that most species in the species group, except the holotype of 7: plesiarum, were previously unknown. This is especially true for 7. bressleri. It is by far the most common and abundant species of the 7° plesiarum group. The material available was sampled in many localities and includes more than 500 mounted specimens with many more in alcohol. The distribu- tion ranges of all group members strongly overlap, but the range of 7. bressleri is by far the largest; this species is known from many more localities than the other four (Fig. 62). The southernmost localities are Andohahela, Beza Mahafaly, and Fiherenana, and from there the distribution ranges north through much of western Madagascar up to the northernmost known locality Anabohazo. ‘The eastern limit of the range goes in an almost straight line north from Andohahela through Mampiarika, Ambohitantely, and Ambohimanga to Anabohazo. The new species prefers arid habitats such as tropical dry forests, tropical dry forests on tsingy, gallery forests, spiny deciduous forests, savannah woodland, Uapaca woodland, and spiny thickets. The elevational range of the species is a relatively broad one, ranging from 10 to 1550 m, but most of the material was collected at low elevations (ca. 420 m on average). Tetramorium bressleri was mainly sampled by pitfall trapping and litter sifting, suggesting a ground-active life style. Discussion. Tetramorium bressleri is not likely to be confused with T: gollum, T. hobbit, or T: mars, whereas differentiating between T! bressleri and T. plesiarum can be challenging at first glance. Tetramorium bressleri lacks the enlarged gaster and strong reticulate-rugose sculpture on the basal half of the first gastral tergite seen in 7: gollum, nor does it have the blocky petiolar node shape of 7: mars or the massively enlarged petiolar node of 7: hobbit. The separation from T° plesiarum requires more atten- tion and caution. The main and obvious difference is body size. Tetramorium bressleri is usually a much larger species (HW 0.80—1.00; WL 0.92-1.15) than T° plesiarum 38 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Figure 15. 7 dressleri holotype worker (CASENT0035677). A Body in profile B Body in dorsal view C Head in full-face view. Taxonomy of Malagasy Tetramorium 39 (HW 0.80-1.00; WL 0.92-1.15). There is some overlap, but this is mainly due to a few very small specimens of T° bressleri and one very large specimen of T. plesiarum. Otherwise, both species fall neatly into their respective size ranges. However, body size alone should not be used as a primary diagnostic character, and in this case, we pro- vide more evidence for their heterospecificity. The eyes of T° plesiarum (OI 21-23) are larger than in T° bressleri (OI 18-19), although this is often difficult to assess without measuring. In addition, both can be separated by the sculpture on the mesopleuron and lateral propodeum; this sculpture is usually mostly absent in 7: bressleri, making the area appear very smooth and shiny, while it is usually longitudinally rugose with reticulate-punctate ground sculpture in 7. plesiarum. The sculpture on the sides of the petiolar node varies too, since it is much more reticulate-punctate and matte in 7: plesiarum than in T. bressleri, in which the weak reticulate-punctate ground sculpture looks faintly matte but still relatively smooth and shiny. The shape of the petiolar node in profile is an additional useful character. In 7! plesiarum the anterodorsal margin of the node protrudes anteriorly and is a bit more marginate than the more rounded posterodorsal margin, while in T° bressleri both margins are usually equally marginate and the anterodorsal margin does not protrude anteriorly at all. Considering how common and abundant 7! bressleri is in western Madagascar, it is interesting that it shows very little intraspecific variation and remains very stable in its morphology. Tetramorium gollum Hita Garcia & Fisher, sp. n. http://zoobank.org/60 DC6DB7-5425-4597-B2E3-3 DOOE8AD4E2B http://species-id.net/wiki/Tetramorium_gollum Figs 10D, 11A, 16, 62 Type material. Holotype, pinned worker, MADAGASCAR, Fianarantsoa, Forét d’Analalava, 29.6 km 280° W, Ranohira, 22.59167°S, 45.12833°E, 700 m, tropical dry forest, pitfall trap, collection code BLF0O7381, 1.—5.II.2003 (CAS: CASENT0074849). Paratypes, two pinned workers with same data as holotype (CAS: CASENT0074839, CASENT0074974). Diagnosis. The strongly developed sculpture on the basal half of the first gastral tergite and the extremely swollen gaster clearly distinguish 7: gollum from the other species in the group. Worker measurements (N=3). HL 0.76-0.90 (0.81); HW 0.74—0.90 (0.80); SL 0.47—-0.59 (0.52); EL 0.15—-0.18 (0.16); PH 0.48-0.52 (0.50); PW 0.54—0.66 (0.59); WL 0.85-1.03 (0.93); PSL 0.23-0.26 (0.25); PTL 0.20-0.25 (0.23); PTH 0.35-0.42 (0.38); PI'W 0.27-0.37 (0.31); PPL 0.24-0.27 (0.25); PPH 0.37-0.43 (0.40); PPW 0.40-0.50 (0.45); CI 97-100 (98); SI 64-66 (65); OI 19-20 (20); DMI 62-64 (63); LMI 50-56 (54); PSLI 29-31 (30); PeNI 50-56 (53); LPel 57-61 (59); DPel 130-148 (138); PpNI 75-79 (76); LPpI 63-65 (64); DPpI 167-185 (176); PPI 135-148 (143). 40 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Worker description. Head weakly longer than wide to as long as wide (CI 97— 100); posterior head margin weakly concave. Anterior clypeal margin with distinct median impression. Frontal carinae strongly developed, curving down shortly before posterior head margin, forming dorsal margin of very well-developed antennal scrobes; scrobes moderately shallow; posterior and ventral margins not fully defined, merging with very strong cephalic sculpture. Antennal scapes short, not reaching posterior head margin (SI 64-66). Eyes relatively small (OI 19-20). Mesosomal outline in profile conspicuously convex, rounded, and very high (LMI 50-56), moderately to strongly marginate from lateral to dorsal mesosoma; promesonotal suture and metanotal groove absent. Propodeal spines long (PSLI 29-31), elongate-triangular to spinose, and acute; propodeal lobes short, triangular to elongate-triangular, and acute. Petiolar node in profile high, rectangular nodiform with well-defined antero- and posterodorsal margins, between 1.6 to 1.8 times higher than long (LPel 57-61), anterior and posterior faces approximately parallel, anterodorsal and posterodorsal margins situated at about the same height, petiolar dorsum flat to weakly convex; node in dorsal view around 1.3 to 1.5 times wider than long (DPel 130-148), in dorsal view pronotum between 1.8 to 2.0 times wider than petiolar node (PeNI 50-56). Postpetiole in profile anteroposte- riorly compressed, approximately 1.5 to 1.6 times higher than long (LPpI 63-65); in dorsal view around 1.7 to 1.9 times wider than long (DPpI 167-185), pronotum only around 1.3 times wider than postpetiole (PpNI 75-79). Postpetiole in profile appearing approximately as voluminous as petiolar node but relatively thinner, postpetiole in dor- sal view approximately 1.3 to 1.5 times wider than petiolar node (PPI 135-148). Gaster extremely enlarged and swollen. Mandibles unsculptured, smooth, and shining; clypeus longitudinally rugose, with five to six distinct rugae, median ruga better developed than remainder, rugae without cross-meshes; sculpture on cephalic dorsum between frontal carinae variable: either mainly longitudinally rugose with higher proportion of reticulate-rugose areas towards posterior head margin, or irregularly reticulate-rugose with a higher proportion of longitudinally rugose sculpture restricted anteriorly close to posterior clypeal margin; scrobal area partly unsculptured; lateral head longitudinally rugose to reticulate-rugose. Mesosoma laterally reticulate-rugose to irregularly longi- tudinally rugose with a few shining areas on mesopleuron and propodeum, dorsally reticulate-rugose to irregularly longitudinally rugose. Forecoxae unsculptured, smooth, and shining. Petiolar node laterally with weak, longitudinally rugose sculpture mostly restricted towards dorsum, dorsum of node mostly unsculptured, smooth, and shiny; sculpture on postpetiole better developed, laterally and dorsally reticulate-rugose. Basal half of first gastral tergite and most of first sternite with very conspicuous and strongly developed reticulate-rugose sculpture superimposed on a reticulate-punctate ground sculpture; remainder of gaster unsculptured, smooth, and shining. Ground sculpture on head, mesosoma, and waist segments weak to moderate, except procoxae with weak but distinct rugulose ground sculpture. Whole body with abundant, long, and fine standing hairs; first gastral tergite with a mix of more abundant, long, erect hairs and much scarcer, shorter, decumbent to subdecumbent hairs. Anterior edges of anten- nal scapes and dorsal (outer) surfaces of hind tibiae with decumbent to suberect hairs. Taxonomy of Malagasy Tetramorium 41 Figure 16. 7° gollum holotype worker (CASENT0074849). A Body in profile B Body in dorsal view C Head in full-face view. 42 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Head, mesosoma, and waist segments light to dark brown, mandibles, antennae, and legs light brown to yellow, and gaster much darker brown than remainder of body. Etymology. The new species is named after the fictional character “Gollum” from J.R.R. Tolkien’s novels “The Hobbit” and “The Lord of the Rings”. The species epithet is an arbitrary combination of letters, thus invariable. Distribution and ecology. The new species is only known from the type locality, the Forét d’Analalava, which is a tropical dry forest located at an elevation of 700 m (Fig. 62). The biology of the species is unknown, but foraging might be undertaken on the ground since the only three known specimens were collected from a pitfall trap. Indicated by this lack of material, 7: gollum probably is one of the more rarely encoun- tered Tetramorium species in Madagascar. Discussion. Tetramorium gollum is a very conspicuous species within the 7° ple- siarum group and the whole Malagasy Tetramorium fauna. The extremely swollen gaster with strongly developed sculpture on the basal half of the tergite is easily rec- ognisable within the species group. The only other Malagasy species with such con- spicuous sculpture clearly extending to half of the tergite or more are T° jedi in the T. tortuosum group and T. sericeiventre in the T. sericeiventre group. ‘The latter has 12 antennal segments (vs. 11 in 7: gollum) and T. jedi has, among other characters, a very differently shaped petiolar node (longer than broad in T° jedi vs. broader than long in T. gollum). Beyond sculpture and the enlarged gaster, 7° gollum is morphologically very similar to T. bressleri, for example the petiolar node shape is identical in both suggesting a close relationship. However, they both also differ in the shape of the post- petiole, which is much higher and broader in T! gollum (LPpI 63-65; DPpI 167-185) than in T° bressleri (LPpI 59-63; DPpI 150-158) Tetramorium hobbit Hita Garcia & Fisher, sp. n. http://zoobank.org/125B2BB7-7E47-4273-B0OFA-B2676FD18D8E http://species-id.net/wiki/Tetramorium_hobbit Figs LOA sa 27s 72 62 Type material. Holotype, pinned worker, MADAGASCAR, Toliara, Parc National de Tsimanampetsotsa, Forét de Bemanateza, 20.7 km 81° E Efoetse, 23.0 km 131° SE Beheloka, 23.99222°S, 43.88067°E, 90 m, spiny forest/thicket, sifted litter (leaf mold, rotten wood), collection code BLF06254, 22.—26.III.2002 (B.L. Fisher et al.) (CAS: CASENT0019207). Paratypes, 2 pinned workers with same data as holotype (CAS: CASENT0019210; CASENT0019227); 2 pinned workers with same data as holotype except collected from pitfall trap and collection code BLF06258 (CAS: CASENT0078055; MHNG: CASENT0078059); 5 pinned workers with same data as holotype except collected from ground nest and collection code BLF06270 (CAS: CASENT0445004; CASENT0445005); and 2 pinned workers with same data as holotype except sampled from ground and collection code BLF06337 (MCZ: CASENT0445502; CASENT0445520). Taxonomy of Malagasy Tetramorium 43 Non-type material MADAGASCAR: Fianarantsoa, Ampandravelo I] Non Protected Area, 10.78 km NE Ranohira, 22.5392°S, 45.5155°E, 873 m, shrubland, 20.—22.11.2010 (A. Ravelomanana); Fianarantsoa, Ampotoampoto I National Parc, 8.02 km NW Ilakaka, 22.6283°S, 45.1886°E, 917 m, savannah woodland, 26.—28. 11.2010 (A. Ravelomanana); Fianarantsoa, Ampotoampoto III National Parc, 7.91 km NW Ilakaka, 22.6294°S, 45.189°E, 919 m, savannah woodland, 27.—28.11.2010 (A. Ravelomanana); Fianarantsoa, Ampotoampoto IV National Parc, 7.83 km NW II akaka, 22.6294°S, 45.1912°E, 923 m, savannah woodland, 27.—28.11.2010 (A. Ravelo- manana); Toliara, Parc National d’Andohahela, Forét de Manantalinjo, 33.6 km 63° ENE Amboasary, 7.6 km 99° E Hazofotsy, 24.8169°S, 46.61°E, 150 m, spiny forest/ thicket, 12.-16.1.2002 (B.L. Fisher et al.); Toliara, Forét de Beroboka, 5.9 km 131° SE Ankidranoka, 22.2331°S, 43.3663°E, 80 m, tropical dry forest, 12.-16.[II.2002 (B.L. Fisher et al.); Toliara, Parc National de Kirindy Mite, 16.3 km 127° SE Belo sur Mer, 20.7953°S, 44.147°E, 80 m, tropical dry forest, 6.-10.III.2002 (B.L. Fisher et al.); Toliara, Makay Mts., 21.3411°S, 45.1805°E, 500 m, barren rock with sparse vegetation, burned grass, 28.X1.2010 (BL. Fisher et al.); Toliara, Parc National de Tsimanampetsotsa, Forét de Bemanateza, 20.7 km 81° E Efoetse, 23.0 km 131° SE Beheloka, 23.99222°S, 43.88067°E, 90 m, spiny forest/thicket, 22.—26.II1.2002 (B.L. Fisher et al.). Diagnosis. Tetramorium hobbit is easily recognisable within the T: plesiarum spe- cies group due to its massively developed petiolar node and very conspicuous reticu- late-punctate ground sculpture on head and mesosoma. Worker measurements (N=12). HL 0.79-0.88 (0.84); HW 0.80—0.88 (0.85); SL 0.52-0.58 (0.55); EL 0.16—0.20 (0.18); PH 0.42—0.53 (0.47); PW 0.63-0.72 (0.69); WL 0.96-1.11 (1.04); PSL 0.16-0.25 (0.20); PTL 0.28—0.31 (0.30); PTH 0.40-0.46 (0.44); PTW 0.38-0.45 (0.41); PPL 0.24-0.30 (0.27); PPH 0.39-0.45 (0.43); PPW 0.45-0.54 (0.51); CI 100-102 (101); SI 64-67 (65); OI 19-22 (21); DMI 63-72 (66); LMI 43-50 (46); PSLI 18-29 (23); PeNI 57-66 (60); LPel 63- 71 (68); DPel 133-148 (140); PpNI 69-76 (74); LPpl 60-67 (63); DPpI 175-200 (188); PPI 116-126 (122). Worker description. Head as long as wide to weakly longer than wide (CI 100-— 102); posterior head margin weakly concave. Anterior clypeal margin with distinct median impression. Frontal carinae strongly developed and forming dorsal margin of very well-developed antennal scrobes, scrobes moderately to very deep and with clearly defined margins all around; median scrobal carina very well developed and distinctly surpassing posterior eye level, usually approaching posterior scrobe margin. Antennal scapes short, not reaching posterior head margin (SI 64-67). Eyes relatively small to moderate (OI 19-22). Mesosomal outline in profile conspicuously convex, rounded and often very high (LMI 43-50), moderately marginate from lateral to dorsal meso- soma; promesonotal suture and metanotal groove absent. Propodeal spines either short and elongate-triangular or long and spinose (PSLI 18-29; material from Andohahela and ‘T’simanampetsotsa with PSLI of 28—29; material from other localities PSLI 18—20), spines always with broad base and acute tip; propodeal lobes well developed, elongate- 44 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) triangular, and acute, always shorter than propodeal spines. Petiolar node massively developed, very large, in profile high, rectangular, blocky nodiform with well defined antero- and posterodorsal margins, between 1.4 to 1.6 times higher than long (LPel 63-71), anterior and posterior faces approximately parallel, anterodorsal and postero- dorsal margins situated at about the same height, petiolar dorsum flat to weakly convex; node in dorsal view around 1.3 to 1.5 times wider than long (DPel 133-148), in dorsal view pronotum around 1.5 to 1.7 times wider than petiolar node (PeNI 57-66). Post- petiole in profile subglobular to anteroposteriorly compressed, approximately 1.5 to 1.7 times higher than long (LPpI 60-67); in dorsal view around 1.7 to 1.9 times wider than long (DPpI 175-200), pronotum around 1.3 to 1.4 times wider than postpetiole (PpNI 69-76). Petiole in profile much more voluminous than postpetiole, in dorsal view postpetiole only about 1.1 to 1.3 times wider than petiolar node (PPI 116-126). Mandibles usually finely longitudinally rugulose, fairly smooth and shiny, sometimes unsculptured; clypeus longitudinally rugose, usually with five to seven very distinct and strong rugae, median ruga better developed than remainder, rugae usually with- out cross-meshes; cephalic dorsum between frontal carinae with 11 to 13 longitudinal rugae, rugae running from posterior clypeal margin to posterior head margin, often interrupted or with cross-meshes, especially posteriorly; scrobal area mostly unsculp- tured, only with ground sculpture; lateral head mainly reticulate-rugose. Mesosoma laterally reticulate-rugose to irregularly longitudinally rugose, dorsally mostly longitu- dinally rugose. Forecoxae unsculptured, smooth, and shining, often with weak traces of ground sculpture. In profile, basal half of petiolar node mostly unsculptured, upper half distinctly reticulate-rugose, dorsum of node distinctly reticulate-rugose; sculpture on postpetiole much weaker, laterally and dorsally only weakly rugulose. Gaster usually unsculptured, smooth and shining, sometimes with weak punctate ground sculpture at base of tergite. Ground sculpture on head, mesosoma, and petiolar node very conspicu- ous, usually strongly reticulate-punctate; ground sculpture on postpetiole and gaster usually much weaker but present, though sometimes absent. Whole body with very abundant, long, and fine standing hairs; first gastral tergite with abundant, long, erect hairs and much scarcer, shorter, decumbent to subdecumbent hairs. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae with decumbent to suberect hairs. Body reddish to dark brown, head and gaster usually darker than rest of body. Etymology. This very hairy new species is named after the fictional people from J.R.R. Tolkien’s novels “The Hobbit” and “The Lord of the Rings”. The species epithet is an arbitrary combination of letters, thus invariant. Distribution and biology. The distribution of 7: hobbit is approximately re- stricted to the southern third of the island of Madagascar (Fig. 62). In the south, the distribution range encompasses the type locality Tsimanampetsotsa in the west and Andohahela in the east, while the Makay Mountains and Isalo mark the northernmost known localities. Interestingly, the population in Andohahela seems to be slightly iso- lated from the other, more western localities. However, we consider this to be more of a sampling artifact. Like the remainder of the species group, 7: hobbit prefers arid habitats, such as tropical dry forests, spiny forests and thickets, savannah woodland, Taxonomy of Malagasy Yetramorium 45 C Head in full-face view. and barren rocks with little vegetation. It is found at elevations of 80 to 923 m. Te- tramorium hobbit is likely a ground-active species since it was mainly collected from leaf litter extractions and pitfall traps. 46 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Discussion. Tetramorium hobbit is very unlikely misidentified with the other four species of the group. The massively enlarged petiolar node, in combination with the distinct reticulate-punctate ground sculpture on head and mesosoma, are not seen in any other group member. However, of all group members, 7° mars seems morphologi- cally closest to T: hobbit, especially on the basis of the blockier petiolar node shape seen in profile that both species share. However, in addition to the smaller petiolar node and the lack of reticulate-punctate ground sculpture on head and mesosoma, T. mars also has shorter antennal scapes (SI 58-62; DPel 108-122) and a narrower petiolar node in dorsal view than 7° hobbit (SI 64-67; DPel 133-148). Tetramorium mars (HW 0.69-0.74; WL 0.83-—0.96) is also smaller than 7: hobbit (HW 0.80-0.88; WL 0.96-1.10) One noticeable intraspecific variation can be observed within 7: hobbit. There are two distinct groups with differently developed propodeal spines. The populations from the type localities Tsimanampetsotsa and Andohahela always have relatively long spines (PSLI of 28-29), which contrast with the much shorter spines seen in the mate- rial from in Beroboka, Kirindy, Isalo, and the Makay Mountains (PSLI 18 -20). This difference is very pronounced and constant in all of the examined material. However, since there is no other obvious morphological difference between the short-spined and the long-spined populations, we treat them as conspecific in this study. Tetramorium mars Hita Garcia & Fisher, sp. n. http://zoobank.org/0682CF5D-60A1-40B2-B552-E2A235 1FB575 http://species-id.net/wiki/Tetramorium_mars Figs 10B, 12B, 13A, 18, 62 Type material. Holotype, pinned worker, MADAGASCAR, Toliara, Forét de Kirindy, 15.5 km 64° ENE Marofandilia, 20.045°S, 44.6622°E, 100 m, tropi- cal dry forest, sifted litter (leaf mold, rotten wood), BLF04605, 28.XI.2001 (B.L. Fisher et al.) (CAS: CASENT0474279). Paratypes, 24 pinned workers with same data as holotype (BMNH: CASENT0474298; CAS: CASENT0474278; CASENT0474281; CASENT0474287; CASENT0474289; CASENT0474292; CASENT0474294; CASENT0474304; CASENT0474310; CASENT0474312; CASENT0474322; CASENT0474328; CASENT0474331; CASENT0474335; CASENT0474338; CASENT0474345; CASENT0474347; CASENT0474349; CASENT0474354; CASENT0474357; CASENT0474366; CASENT0474371; MHNG: CASENT0474312; MCZ: CASENT0474286; CASENT0474307). Non-type material. MADAGASCAR: Mahajanga, Parc National d’Ankarafantsika, Forét de Tsimaloto, 18.3 km 46° NE de Tsaramandroso, 16.2281°S, 47.1436°E, 135 m, tropical dry forest, 2.-8.IV.2001 (BL. Fisher et al); Mahajanga, Parc National d’Ankarafantsika, Ampijoroa Station Forestiere, 5.4 km 331° NW Andranofasika, 16.2989°S, 46.813°E, 70 m, tropical dry forest, 26.[II.-1.1V.2001 (B.L. Fisher et al); Mahajanga, Parc National de Baie de Baly, 12.4 km 337° NNW Soalala, 16.01°S, Taxonomy of Malagasy Tetramorium 47 45.265°E, 10 m, tropical dry forest, 26.—30.X1.2002 (B.L. Fisher et al.); Mahajanga, Parc National Tsingy de Bemaraha, 10.6 km ESE 123° Antsalova, 18.7094°S, 44.7182°E, 150 m, tropical dry forest on tsingy, 16.—20.X1.2001 (B.L. Fisher et al.); Toliara, Tulear, Bereboka, 60 km NE Morondava, 18.—23.V.1983 (/.S. Noyes & M.C. Day); Toliara, Atsimo Andrefana Region, District of Betioky, 30 km E Betioky, Beza Mahafaly Special Reserve, 23.6865°S, 44.591°E, 165 m, gallery dry deciduous forest, 29.IV.-19.V.2002 (M. Rina); Toliara, Res. Beza-Mahafaly, Parcel 1, 23.65833°S, 44.62889°E, 160 m, tropical dry forest, 13.11.1993 (G.D. Alpert); Toliara, Res. Beza-Mahafaly, Parcel 1, 23.65°S, 44.63333°E, 130 m, tropical dry forest, 13.X1.1993 (P.S. Ward); Toliara, Beza-Mahafaly, 27 km E Betioky, 23.65°S, 44.6333°E, 135 m, tropical dry forest, 23.1V.1997 (B.L. Fisher); Toliara, Kirindy Forest, 20.07458°S, 44.67611°E, 100m, tropical dry forest, 20.XII.1993 (G.D. Alpert); Toliara, Forét de Kirindy, 15.5 km 64° ENE Marofandilia, 20.045°S, 44.6622°E, 100 m, tropical dry forest, 28.XI.-3.XII.2001 (B.L. Fisher et al.); Toliara, Parc National de Kirindy Mite, 16.3 km 127° SE Belo sur Mer, 20.7953°S, 44.147°E, 80 m, tropical dry forest, 6-10.X1.2001 (B.L. Fisher et al.); Toliara, Forét de Mite, 20.7 km 29° WNW Tongobory, 23.5242°S, 44.1213°E, 75 m, gallery forest, 27.1] —3.III.2002 (B.L. Fisher et al.); Toliara, 48 km ENE Morondava, 20.06667°S, 44.65°E, 30 m, tropical dry forest, 4.-7.1.1991 (D.M. Okon). Diagnosis. Tetramorium mars is distinguishable from the other group members by the following combination of characters: antennal scapes very short (SI 58-62); petiolar node never enlarged and massive; node in profile high, rectangular, blocky nodiform with well-defined antero- and posterodorsal margins; anterodorsal margin not protruding anteriorly nor very sharply angled; petiolar node in profile relatively low, between 1.3 to 1.5 times higher than long (LPel 69-76), in dorsal view only 1.1 to 1.2 times wider than long (DPel 108-123); gaster never enlarged or swollen; ground sculpture on head and mesosoma weak to absent; first gastral tergite without strong reticulate-rugose sculpture, usually completely unsculptured, but if sculpture present, then relatively weak and restricted to base of tergite; pilosity on first gastral tergite mostly erect. Worker measurements (N=12). HL 0.71-0.77 (0.74); HW 0.69-0.74 (0.72); SL 0.42-0.46 (0.44); EL 0.15—-0.17 (0.16); PH 0.35—0.44 (0.38); PW 0.50—0.58 (0.54); WL 0.83-0.96 (0.88); PSL 0.21—0.24 (0.22); PTL 0.22-0.27 (0.24); PTH 0.32-0.37 (0.34); PTW 0.26-0.33 (0.29); PPL 0.22—0.26 (0.24); PPH 0.28-0.33 (0.31); PPW 0.36-0.42 (0.38); CI 96-99 (98); SI 58-62 (60); OI 21-22 (22); DMI 60-64 (62); LMI 42-46 (44); PSLI 28-32 (30); PeNI 51-56 (53); LPel 69-76 (73); DPel 108-123 (117); PpNI 68-72 (70); LPpI 75-80 (78); DPpI 156-165 (160); PPI 129-137 (134). Worker description. Head longer than wide (CI 96-98); posterior head margin weakly concave. Anterior clypeal margin with distinct median impression. Frontal carinae strongly developed and forming dorsal margin of very well-developed antennal scrobes, scrobes moderately to very deep and with clearly defined margins all around; median scrobal carina very well developed and distinctly surpassing posterior eye level. Antennal scapes very short, not reaching posterior head margin (SI 58-62). Eyes rela- 48 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) tively small to moderate (OI 21—22). Mesosomal outline in profile moderately convex, rounded and high (LMI 42-46), moderately to strongly marginate from lateral to dorsal mesosoma; promesonotal suture and metanotal groove absent. Propodeal spines long, elongate-triangular to spinose (PSLI 28-32), spines always with broad base and acute tip; propodeal lobes well developed, elongate-triangular, and acute, always much shorter than propodeal spines. Petiolar node in profile high, rectangular, blocky nodiform with well-defined antero- and posterodorsal margins, between 1.3 to 1.5 times higher than long (LPel 68-76), anterior and posterior faces approximately parallel, antero- and pos- terodorsal margins situated at about same height, petiolar dorsum weakly convex; node in dorsal view around 1.1 to 1.2 times wider than long (DPel 108-123), in dorsal view pronotum between 1.8 to 2.0 times wider than petiolar node (PeNI 51-56). Postpetiole in profile subglobular, approximately 1.3 times higher than long (LPpI 75-80); in dor- sal view around 1.5 to 1.7 times wider than long (DPpI 156-165), pronotum around 1.4 to 1.5 times wider than postpetiole (PpNI 68-72). Postpetiole in profile appearing much less voluminous than petiolar node, postpetiole in dorsal view around 1.3 to 1.4 times wider than petiolar node (PPI 129-137). Mandibles unsculptured, smooth, and shining; clypeus longitudinally rugose/rugulose, with five to seven distinct rugae/rugulae, median ruga better developed than remainder, all rugae without cross-meshes; cephalic dorsum between frontal carinae with eight to twelve longitudinal rugae, rugae running from posterior clypeal margin to posterior head margin, often interrupted or with cross- meshes, especially posteriorly; scrobal area mostly unsculptured; lateral head mainly longitudinally rugose. Ground sculpture on head generally weak to absent, sometimes a few areas weakly punctate. Mesosoma laterally irregularly longitudinally rugose with a few shining areas on mesopleuron and propodeum, mesosomal dorsum longitudinally rugose. Forecoxae unsculptured, smooth, and shining. Ground sculpture on mesosoma absent. In profile, basal half of petiolar node mostly unsculptured, upper half distinctly reticulate-rugose, whole dorsum of node distinctly reticulate-rugose; sculpture on post- petiole much more weakly developed, laterally and dorsally weakly irregularly rugulose. Ground sculpture on waist segments variable, usually weak or absent, often petiole and/ or postpetiole conspicuously reticulate-punctate. Gaster usually unsculptured, smooth, and shining, sometimes base of first gastral tergite weakly punctate. Whole body with very abundant, long, and fine standing hairs; first gastral tergite with abundant, long, erect hairs and much scarcer, shorter, decumbent to subdecumbent hairs. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae with decumbent to suberect hairs. Usually body uniformly light to reddish brown, sometimes darker. Etymology. This new species is named after the ancient Roman god of war, “Mars”. The species epithet is an arbitrary combination of letters, thus invariable. Distribution and biology. Tetramorium mars has a broad distribution in western Madagascar (Fig. 62). It ranges from the southernmost known localities, Beza-Maha- faly and Forét de Mite, through Kirindy Mite and Tsingy de Bemaraha to the north- ernmost localities, Baie de Baly and Ankarafantsika. All known localities are tropical dry or gallery forests at low elevations from 10 to 165 m. The new species was mainly Taxonomy of Malagasy Tetramorium 49 Figure 18. 7 mars holotype worker (CASENT0474279). A Body in profile B Body in dorsal view C Head in full-face view. 50 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) sampled by litter extraction or pitfall trapping, which suggests a ground active lifestyle. Interestingly, 7! mars was not found in Andohahela in the southeast of Madagascar, even though 7° bressleri, T. hobbit, and T. plesiarum occur there. This is surprising since the four species of the group except 7: gollum share more or less the same distri- bution range. Discussion. Within the species group 7. mars is not likely to be confused with the other five species. The lack of strong and conspicuous sculpture on the basal half of the first gastral tergite, an enlarged gaster, and the lower and less broad petiolar node separate 7: mars (LPel 69-76; DPel 108-123) clearly from 7! gollum (LPel 57-60; DPel 130-148). Despite the fact that 7: mars shares some similarities with T. bressleri and T. plesiarum, the latter also have relatively higher, thinner, and wider petiolar nodes (LPel 56-63; DPel 130-144) which distinguish them from T: mars. In addition, 7: mars has much better developed sculpture on the dorsum of the petiolar node than 7! bressleri or T. plesiarum. The species most similar morphologically to T- mars is T. hobbit. Both share a more blocky nodiform petiolar node, but which is still much larger in 7° hobbit. Indeed, this massively developed node, in combination with very conspicuous punctate ground sculpture on head and mesosoma, separates T° hob- bit from T. mars, which has a much smaller petiolar node and only weakly developed ground sculpture on head and mesosoma. Further characters that distinguish 7. mars are the comparatively short antennal scapes (SI 58-62 vs. SI 64-67) and smaller body size (HW 0.69-0.74 vs. HW 0.80-0.88). Despite its relatively broad distribution range, 7. mars displays very little intraspe- cific or geographic variation. Tetramorium plesiarum Bolton http://species-id.net/wiki/Tetramorium_plesiarum Figs 10C, 11B, 13B, 14A, 19, 62 Tetramorium plesiarum Bolton, 1979:150. Type material. Holotype, pinned worker, MADAGASCAR, Causse de Kelifely, 17.30306°S, 45.73444°E, forest humus and litter, dry forest, 20.—30.X1.1974 (A. Pey- rieras) (MCZ: MCZ_Holotype_32372) [examined]. [Note: the GPS data of the type locality was not provided by the locality label or the original description. The data presented above is based on our own geo-referencing of the Kelifely limestone plateau and should be considered an approximation but not the exact position of the type locality.] Non-type material, MADAGASCAR: Mahajanga, Reserve Forestiere Beanka, 50.7 km E Maintirano, 17.8802°S, 44.4688°E, 140 m, tropical dry forest on tsingy, 29.X.-1.X1.2009 (B.L. Fisher et al.); Mahajanga, Parc National de Namoroka, 9.8 km 300° WNW Vilanandro, 16.4667°S, 45.35°E, 140 m, tropical dry forest, 4.—-8. X1.2002 (B.L. Fisher et al.); Mahajanga, Parc National de Namoroka, 17.8 km 329° Taxonomy of Malagasy Tetramorium 51 WNW Vilanandro, 16.3767°S, 45.3267°E, 100 m, tropical dry forest, 8.—12.X1.2002 (B.L. Fisher et al.); Mahajanga, Parc National de Namoroka, 16.9 km 317° NW Vilanandro, 16.4067°S, 45.31°E, 100 m, tropical dry forest, 12.-16.X1.2002 (B.L. Fisher et al.); Mahajanga, Parc National Tsingy de Bemaraha, 10.6 km ESE 123° Ant- salova, 18.7094°S, 44.7182°E, 150 m, tropical dry forest on tsingy, 16.—-20.X1.2001 (B.L. Fisher et al.); Toliara, Parc National d’Andohahela, Forét de Manantalinjo, 33.6 km 63° ENE Amboasary, 7.6 km 99° E Hazofotsy, 24.8169°S, 46.61°E, 150 m, spiny forest/thicket, 12.-16.1.2002 (B.L. Fisher et al.); Toliara, Parc National d’Andohahela, Forét d’Ambohibory, 1.7 km 61° ENE Tsimelahy, 36.1 km 308° NW Tolagnaro, 24.93°S, 46.6455°E, 300 m, tropical dry forest, 16.—20.1.2002 (B.L. Fisher et al.); To- liara, Andohahela National Park, Tsimelahy—Parcel II, 24.9368°S, 46.6267°E, 180 m, transition forest, 5.-15.II.2003 (ME. Irwin et al.); Toliara, Makay Mts., 21.2184°S, 45.3106°E, 510 m, gallery forest on sandy soil, 24.-27.X1.2010 (B.L. Fisher et al.); Toliara, Makay Mts., 21.2176°S, 45.3392°E, 500 m, gallery forest on sandy soil, 28.X1.2010 (B.L. Fisher et al.); Toliara, Makay Mts., 21.3136°S, 45.1478°E, 525 m, gallery forest on sandy soil, 6.XII.2010 (B.L. Fisher et al.); Toliara, Vohibasia Forest, 59 km NE Sakaraha, 22.4667°S, 44.85°E, 780 m, tropical dry forest, 13.1.1996 (B.L. Fisher); Toliara, Parc National de Zombitse, 19.8 km 84° E Sakaraha, 22.8433°S, 44.71°E, 770 m, tropical dry forest, 5.-9.11.2003 (B.L. Fisher et al.). Diagnosis. The following character set separates T. plesiarum from the remainder of the species group: smaller species (HW 0.80—1.00; WL 0.92-1.15); eyes of moder- ate size (OI 21-23); petiolar node high nodiform, not massively enlarged, anterodorsal margin protruding anteriorly and slightly more marginate than posterodorsal margin; petiolar node in profile relatively high and thin, between 1.6 to 1.8 times higher than long (LPel 56-63), in dorsal view between 1.3 to 1.4 times wider than long (DPel 131-137); gaster never extremely enlarged and swollen; head without strongly de- veloped and conspicuous reticulate-punctate ground sculpture; mesopleuron and lat- eral propodeum with moderate punctate ground sculpture; first gastral tergite without strong reticulate-rugose sculpture, usually completely unsculptured, but if sculpture present, then relatively weak and restricted to base of tergite; pilosity on first gastral tergite mostly erect. Worker measurements (N=12). HL 0.66-0.75 (0.73); HW 0.67-0.76 (0.72); SL 0.45-0.49 (0.47); EL 0.15-0.17 (0.15); PH 0.33-0.44 (0.36); PW 0.50-0.57 (0.53); WL 0.81-0.92 (0.85); PSL 0.16-0.21 (0.18); PTL 0.18—-0.22 (0.19); PTH 0.31-0.35 (0.33); PTW 0.23-0.29 (0.26); PPL 0.22—0.26 (0.24); PPH 0.29-0.33 (0.31); PPW 0.32—0.40 (0.35); CI 97-102 (99); SI 64-69 (66); OI 21—23 (22); DMI 60-65 (63); LMI 41-47 (43); PSLI 23-28 (25); PeNI 45-51 (49); LPeI 56-63 (59); DPel 131-137 (133); PpNI 63-70 (66); LPpI 73-79 (76); DPpI 140-157 (149); PPI 129-143 (136). Worker description. Head more or less as long as broad (CI 97—102); posterior head margin weakly concave. Anterior clypeal margin with distinct median impression. Frontal carinae strongly developed and forming dorsal margin of very well-developed antennal scrobes, scrobes moderately to very deep and with clearly defined margins all 52 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) around; median scrobal carina very well developed and distinctly surpassing posterior eye level, usually ending halfway between posterior eye margin and posterior scrobe margin, often approaching posterior scrobe margin. Antennal scapes short, not reach- ing posterior head margin (SI 64-69). Eyes small to moderate (OI 21-23). Mesosomal outline in profile weakly to moderately convex, rounded and high (LMI 41-47), mod- erately marginate from lateral to dorsal mesosoma; promesonotal suture and metanotal groove absent. Propodeal spines elongate-triangular to spinose, moderately long to long, and acute (PSLI 23-28), spines always with broad base and acute tip; propo- deal lobes well developed, triangular to elongate-triangular, and acute, always much shorter than propodeal spines. Petiolar node in profile high nodiform, between 1.6 to 1.8 times higher than long (LPel 56—63), anterodorsal margin situated higher and more angled than posterodorsal, more rounded margin, petiolar dorsum weakly to moderately tapering backwards posteriorly, anterodorsal margin slightly protruding anteriorly, anterior and posterior faces approximately parallel; node in dorsal view be- tween 1.3 to 1.4 times wider than long (DPel 131-137), pronotum between 1.9 to 2.2 times wider than petiolar node (PeNI 45-51). Postpetiole in profile subglobular, approximately 1.3 to 1.4 times higher than long (LPpI 73-79); in dorsal view around 1.4 to 1.6 times wider than long (DPpI 140-157), in dorsal view pronotum between 1.4 to 1.6 times wider than postpetiole (PpNI 63-70). Postpetiole in profile appear- ing slightly less voluminous than petiolar node, postpetiole in dorsal view about 1.3 to 1.4 times wider than petiolar node (PPI 129-143). Mandibles unsculptured, smooth, and shining; clypeus longitudinally rugose, with three to five distinct rugae, median ruga better developed than remainder, rugae without cross-meshes; cephalic dorsum between frontal carinae with ten to twelve longitudinal rugae, rugae running from posterior clypeal margin to posterior head margin, often interrupted or with cross- meshes, especially posteriorly; scrobal area mostly unsculptured, relatively smooth and shiny; lateral head longitudinally rugose to reticulate-rugose. Ground sculpture on head usually present but weak. Mesosoma laterally irregularly longitudinally rugose, usually with moderate punctate ground sculpture on mesopleuron and propodeum, dorsal mesosoma longitudinally rugose without any distinct ground sculpture, spaces between rugae smooth and shining. Forecoxae unsculptured, smooth, and shining. Sides of petiolar node and postpetiole with conspicuous, moderate reticulate-punctate ground sculpture, both nodes appearing relatively matte; petiolar node and postpetiole dorsally weakly rugulose/rugose, but mostly unsculptured, and appearing smooth and shining. Gaster usually unsculptured, smooth and shining, sometimes base of first gas- tral tergite with traces of punctate ground sculpture. Whole body with very abundant, dense, moderately long, and fine standing hairs; first gastral tergite with abundant, long, erect hairs and much scarcer, shorter, decumbent to subdecumbent hairs. Ante- rior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae with decumbent to suberect hairs. Body of uniform brown, appendages often lighter. Distribution and biology. Despite being known only from seven localities and fewer than 20 specimens, 7. plesiarum has a comparatively large distribution range Taxonomy of Malagasy Tetramorium oi) Figure 19. 7. plesiarum holotype worker (CASENT0172831). A Body in profile B Body in dorsal view C Head in full-face view. 54 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) (Fig. 62). However, the populations appear to be relatively disjunctive. The species is known from Andohahela in the southeast of Madagascar and from six additional, more or less scattered localities in the western part of the island ranging from Zom- bitse and Vohibatsia north through the Makay Mountains, Tsingy de Bemaraha, and Beanka to the Kelifely plateau and Namoroka. The small number of specimens suggests that 7. plesiarum is much less common than the sympatric 7. bressleri, which shares much of the distribution range, but is known from many more locali- ties and hundreds of specimens. The rarity of 7. plesiarum might well explain the disjunctive distribution. Unfortunately, there is no “fresh” material available from the type locality since the holotype remains the only known specimen from the Ke- lifely plateau. All localities are tropical dry forests, tropical dry forests on tsingy, or gallery forests ranging from 100 to 780 m elevation. Like the other group members T. plesiarum was mostly collected by pitfall trapping and litter sifting suggesting a ground-active lifestyle. Discussion. Tetramorium plesiarum \acks strong specialisations, such as the strongly sculptured and enlarged gaster of 7. gollum, or the massively developed petiolar node of 7. Aobbit. Furthermore, 7: mars has a lower and wider petiolar node (LPel 69-76; DPel 108-123) compared to T° plesiarum (LPel 56-63; DPel 131-137). The latter also has much less sculpture on the dorsum of the petiolar node. Within the species group 7. plesiarum is morphologically most similar to T- bressleri, and as mentioned in the description of the latter, we have treated them as conspecific through most of the revision The most noticeable difference is certainly body size since T. bressleri (HW 0.80-1.00; WL 0.92—1.15) is much larger than T- plesiarum (HW 0.80-1.00; WL 0.92—1.15). As noted above, there is some slight overlap caused by one large specimen of T. plesiarum and very few small specimens of 7. bressleri, but in the main they fit their respective size ranges. Not consider- ing body size, they also differ in eye size and sculpture on mesosoma and waist segments. The eyes of 7. plesiarum (OI 21-23) are larger than in T° bressleri (OI 18-19). The mesopleuron and lateral propodeum of T° bressleri is usually mostly unsculptured and relatively smooth and shining, whereas this area is longitudinally rugose with reticulate-punctate ground sculpture in 7. plesiarum. Almost the same pattern is seen in the sculpture on the lateral petiolar node as it is significantly more reticulate-punctate and matte in T. plesiarum than in T. bressleri, in which the weak reticulate-punctate ground sculpture appears faintly matte but still relatively smooth and shiny. The sculpture on the sides of the petiolar node varies, too, since it is much more reticulate-punctate and matte in 7. plesiarum than in T. bressleri, in which the weak reticulate-punctate ground sculpture looks faintly matte but still relatively smooth and shiny. The shape of the petiolar node in profile is an additional useful character. Finally, in 7. plesiarum the anterodorsal margin of the petiolar node protrudes anteriorly and is slightly more marginate than the more rounded posterodorsal margin, contrasting with the shape observed in 7. bressleri, in which both margins are usually equally marginate and the anterodorsal margin does not protrude anteriorly at all. Taxonomy of Malagasy Tetramorium 55 Revision of the Tetramorium schaufussii species group Tetramorium schaufussii species group Diagnosis. Eleven-segmented antennae; anterior clypeal margin medially impressed; frontal carinae variably developed, always present and surpassing eye level, usually weak to moderate and posteriorly merging into the surrounding sculpture, in a few species very well developed, noticeably raised and approaching posterior head mar- gin; antennal scrobes usually weak, shallow, and without clearly defined posterior and ventral margins; anterior face of mesosoma weakly developed; mesosomal outline in profile relatively flat, usually comparatively low and elongated (LMI 35-42), usually moderately marginate from lateral to dorsal mesosoma; propodeal spines usually short to moderately long, triangular to elongate-triangular, long and spinose in a few spe- cies (PSLI 7—28); propodeal lobes usually triangular and short; petiolar node in profile rounded nodiform to high rounded nodiform with well-rounded margins, rarely high cuneiform or squamiform, node in profile between 1.2 to 3.0 times higher than long (LPel 33-81), in dorsal view usually wider than long (DPel 111-238), rarely longer than wide (DPel 87-98), anterior and posterior faces parallel in most species; postpeti- ole in profile globular to subglobular, 1.2 to 1.7 times higher than long (LPel 60-83), in dorsal view between 1.2 to 1.7 times wider than long (DPpI 120-168); mandibles always unsculptured, smooth, and shining; cephalic sculpture distinct, between frontal carinae predominantly longitudinally rugose; mesosoma usually with well-developed longitudinally rugose/rugulose sculpture, in some species sculpture mainly irregularly rugose/rugulose, rarely irregularly rugose/rugulose to reticulate-rugose; waist segments and gaster unsculptured, smooth, and shiny; standing pilosity always present on head, but variable on remainder of body, first gastral tergite often without standing pilosity at all, appressed pubescence on first gastral tergite variably developed, varying from very short and scarce to long and dense; sting appendage spatulate. Comments. With 20 recognised species the T° schaufussii species group is second in terms of species-richness in the Malagasy region, following the 7! tortuosum group with 22 (Hita Garcia and Fisher 2012b). This group is of special importance for Mada- gascar since many of its members are very widespread, common, and/or abundant. Almost all species prefer rainforests or montane rainforests, and with few exceptions, live in the forest leaf litter stratum. The T. schaufussii species group is a very distinctive group among the thirteen groups with eleven-segmented antennae, and most of its members are unlikely to be mistaken for any other group. In general, most T: schaufussii group species are small to medium-sized ants, have a comparatively low and elongated mesosoma, and short to very short propodeal spines. Even though these characters are not unique to this group its members are usually easily recognisable. One interesting feature of the 7! schau- fussii group is that the mandibles of all species are completely unsculptured, smooth, and shining. This character alone already separates the T° schaufussii group from the T. bessonii, T. bonibony, T. kelleri, T. tortuosum, and T. weitzeckeri groups, in which 56 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) all species have noticeably sculptured mandibles. In most other groups this character, however, is variable. Some species of the T: dysalum group have unsculptured mandi- bles, but these have either much longer propodeal spines or a much higher mesosoma. Another important group character is the lack of sculpture on the waist segments. The T. schaufussii group shares this character with a number of other groups, but it does distinguish the group from the T° kelleri, T. ranarum, and T. tortuosum groups, as well as from parts of the 7: bonibony and T. dysalum groups. All these groups have at least one waist segment with distinct sculpture, but most of their species have conspicuous sculpture on both. Furthermore, in the 7° schaufussii group the antennal scrobe is ei- ther weakly developed or absent distinguishing it immediately from the T! plesiarum group and a few species of the T° ibycterum complex of the T. ranarum group with moderately to very deep scrobes with clearly defined margins all around. Moreover, all species of the T° schaufussii species group have relatively well devel- oped cephalic and mesosomal sculpture. The sculpture on the cephalic dorsum be- tween the frontal carinae is always well developed, whereas sculpture on the mesosomal dorsum can be weak in some species (but remains distinctly present). This separates the group from the 7! bessonii, T. marginatum, T. tsingy groups, which usually have strongly reduced sculpture on the head and mesosoma. The members of the 7: mar- ginatum group that possess more sculpture have strongly cuneiform and/or triangular petiolar nodes, and are thus not confusable with the T° schaufussii group. The groups being morphologically most similar to the T: schaufussii group are the T. naganum and T. severini groups, as they share the usually high nodiform petiolar node and complete- ly unsculptured waist segments. The only previously known species of both groups, T. naganum and T. severini, were initially recognised as members of the T! schaufussii group by Bolton (1979), but recently split by Hita Garcia and Fisher (2011, 2012a, 2012b). The separation from the 7! severini group is relatively straightforward. The latter species is very large, very darkly coloured, and has very long to extremely long and curved propodeal spines that easily distinguish its members from the 7! schau- fussii group. The distinction between the T° schaufussii and the T. naganum group is less clear. The members of the 7° naganum group usually have much more strongly developed frontal carinae, a generally broader head (CI 92-99), a higher mesosoma (LMI 40-46), and usually longer propodeal spines (PSLI 25-37). Some of these values overlap with a few species of T° schaufussii group, but they separate the bulk of species comparatively well. Bolton (1979) divided the group into two complexes on the basis of the absence (7. cognatum complex) or presence (7° schaufussii complex) of long, standing pilosity on the first gastral tergite. We follow this division and have placed all the new species into these two complexes even though at present we have no more evidence that these complexes represent monophyletic, mutually exclusive clades or lineages within the species group. Indeed, the only separating character is gastral pilosity, and future studies on the group that ideally include molecular data will likely yield different groupings. However, with 20 species the group is relatively species-rich and the clear-cut division into these two complexes facilitates the taxonomic organisation of the species group. Taxonomy of Malagasy Tetramorium Bus Key to species complexes of the Tetramorium schaufussii species group 1 First gastral tergite with appressed pubescence of varying length and without any standing hairs (Fig. 20A, B) [Madagascar and Comoros]........... cesses See cee Ai AA ERR ae RE T. cognatum species complex - First gastral tergite with appressed pubescence of varying length and few to numerous standing hairs (Fig. 20C, D) [Madagascar and Reunion].............. EL aes teh san eS! T. schaufussii species complex Pew ivipal le 0.5 mm Figure 20. First gastral tergite in profile. A 7’ karthala (CASENT0136774) B ZT proximum (CASENT0906146) C 7; merina (CASENT0437232) D T; obiwan (CASENT0447245). Tetramorium cognatum species complex Comments. This species complex contains a compact assemblage of ten species that are separated from the 7° schaufussii species complex by their lack of any long, standing pilosity on the first gastral tergite, a character present in all members of the T° schau- Jussii complex. The ten species of the 7. cognatum complex might be further divided into three smaller groupings: the first of these contains the five small to moderately large species with weak to very weak frontal carinae (T° aspis, T. camelliae, T. cognatum, T. karthala, and T. rumo); the second includes the four larger species with very well developed frontal carinae (T°. gladius, T. myrmidon, T. proximum, and T. tenuinode); the third sub-grouping contains only T° freya, which is the only species of the complex lacking standing pilosity on the mesosoma, but it also possesses reduced frontal carinae and a larger body size. Four species of the complex have very restricted distribution ranges and are en- demics to smaller areas or localities in Madagascar or the Comoros (Figs 63, 64): T. aspis (Andringitra and Ivohibe), T°. camelliae (Ranomafana), T\ karthala (Grand Co- more), and 7: myrmidon (Ambohijanahary), while 7. freya was found only in a few littoral localities ranging from Nosy Be to the northernmost tip of Madagascar. The remaining species (7: cognatum, T. gladius, T. proximum, T. rumo, and T. tenuinode) have much wider distribution ranges, especially 7. cognatum and T. proximum, which are found in almost all rainforests and montane rainforests from which material is available. Interestingly, almost all species appear be restricted to or prefer rainforests or montane rainforests, even the widely distributed 7: cognatum, T. proximum, and T: tenuinode. However, T. cognatum was occasionally collected in drier habitats and T: freya occurs also in littoral and dry forests. 58 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Identification key to species of the T’ cognatum species complex (workers) 1 Mesosoma without any long, standing hairs (Fig. 21A) [Madagascar].... T. freya — Mesosoma always with few to numerous pairs of standing hairs (Fig. 21B, C)...2 Ds Petiolar node strongly squamiform, in profile 2.8 to 3.0 times higher than long (LPel 33-36) and in dorsal view around 2.3 to 2.4 times wider than long (DPel 228-238) (Fig. 22A) [Madagascar]... cece T. camelliae — Petiolar node never strongly squamiform as above, in profile 1.6 to 2.7 (usually 1.6 to 2.2) times higher than long (LPel 37-62) and in dorsal view between 1.1 to 1.7 times wider than long DPel 111-171) (Fig. 22B, C, D).. ee. 3 6) Eyes relatively small (OI 19-20) (Fig. 23A) [Madagascar] ............ T. gladius = Eyes always much larger than above (OI 24-31) (Fig. 23B, C, D)........ 4 4 Larger species (HW 0.58-0.81; WL 0.85-1.07); frontal carinae moderately well to strongly developed, noticeably raised, and usually approaching or endine-atposterior head margin: (hig. 24M, IB) airbase veel essen oveluseeteaeaas 5 = Smaller species (HW 0.43-0.55; WL 0.56-0.81); frontal carinae usually weakly developed, faintly raised, and mostly ending or fading into surround- ing sculpture halfway between posterior eye margin and posterior head mar- aT AG gy Coe 925 ON D) MR Aen yO REO REDRESS 7 5 Usually significantly larger species (HW 0.65—0.81; WL 0.92-1.07); meso- soma usually with five or six pairs of long, standing hairs on pronotum and mesonotuin( Pies 25A))[ Madagascar) f. .ccctct covseredsteveues maveectes T. proximum = Usually smaller species (HW 0.58-0.70; WL 0.76—0.94); mesosoma always with only two pairs of long, standing hairs, one pair on anterior pronotum and one on‘anterior nesonorum (Pig. 25 BRE) tes. st cic tesnceeede ced pecsiees 6 6 Antennal scapes shorter (SI 66-70); petiolar node higher, around 1.8 to 2.2 times higher than long (LPel 45—54); postpetiole in dorsal view around 1.5 to 1.7 times broader than petiolar node (PPI 148-167) (Fig. 26A, B) [Mad apa scart. tt ea hict seas cust t caret Suvese tant dba diet aueaacctlehs cea vas T. tenuinode = Antennal scapes longer (SI 74—76); petiolar node lower, around 1.6 to 1.7 times higher than long (LPel 58-60) postpetiole in dorsal view only around 1.3 to 1.4 times broader than petiolar node (PPI 133-141) (Fig. 26C) A ASCE tact sSotabaiatcatsetatartunanciasese he wind destin At tetas coeat ade T. myrmidon 7 Smaller species (HW 0.43-0.49; WL 0.56—0.67); propodeal spines moder- ately long to long, elongate-triangular to spinose (PSLI 22-26); petiolar node thinly cuneiform and moderately squamiform, in profile around 2.3 to 2.7 times higher than long (LPel 37-43), in dorsal view around 1.6 to 1.7 times wider than long (DPel 156-171); colouration always uniformly whitish yel- low to light yellowish brown (Fig. 27A) [Madagascar] «0.0... cece T. rumo - Larger species (HW 0.48-0.55; WL 0.65-0.81); propodeal spines short to moderately long, triangular to elongate-triangular (PSLI 12-22); petiolar node high nodiform, in profile around 1.8 to 2.2 times higher than long Taxonomy of Malagasy Tetramorium 59 (LPel 46-55), in dorsal view 1.3 to 1.6 times wider than long (DPel 129- 161); colouration variable, but rarely yellowish as above (Fig. 27B, C) ....... 8 8 Antennal scapes shorter (SI 61-67); propodeal spines reduced and very short (PSLI 12-16), propodeal spines and lobes often of same length or spines even shorter, never strongly directed towards each other; petiolar node in dorsal view around 1.3 to 1.4 times wider than long (DPel 129-142) (Fig. 28A, B) (MAA GS car || dit atest. SUahtms aN Rt Sel ape eet TRE as os Rat Sachi T. cognatum - Antennal scapes longer (SI 68-72); propodeal spines short to moderately long (PSLI 18-22), spines either much longer than lobes or only weakly so and spines and lobes strongly inclined towards each other; petiolar node in dorsal view around 1.5 to 1.6 times wider than long (DPel 146-161) (nO) Dae a ee ety, cde ene JR a 9 9 Eyes smaller (OI 25-27); propodeal lobes always only weakly shorter than propodeal spines, and spines and lobes strongly inclined towards each other; dorsal mesosoma with six or more pairs of long, standing hairs on pronotum and mesonotum, and propodeum usually with one or two pairs anteriorly Cig 2 OAs WIAA AC AS Ga) cats skseadeotsdvasany euSoshs uastbeseSate’s sarees dsathiSiscans te T. aspis = Eyes larger (OI 29-30); propodeal lobes short and triangular, always much shorter than propodeal spines, and spines and lobes never strongly inclined towards each other; dorsal mesosoma with two to four pairs of long, standing hairs on pronotum and mesonotum, standing hairs absent from propodeum (Pig 2 9B): (GommorOs |. ih. Seaton veresn entice teaser neath senanetateteontracetes T. karthala Figure 21. Mesosoma in profile. A 7 freya (CASENT0085551) B 7° cognatum (CASENT0217758) C 7. gladius (CASENT0163234). Figure 22. Waist segments in profile. A 7’ camelliae (CASENT0247496) B 7. rumo (CASENT0073025) C 7. aspis (CASENT0344940) D 7 proximum (CASENT0906146). 60 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) ate i _ 1 he # Yl i i ~ ‘ * re “ , i " rs on " hulle \. , fi eae Py ten 2 4 ’ : %, , a — oz A F i ir ee r i oe 7 i if i . ; a) My i= te oi am a r z. \ . 7. ¥ ‘ FE re ay ; i ff “ae he ail r 3 i! il ae: Pe t r 5 ee i OF a] ¥ a ——— = if | , " ‘ : - = ? p r i " I i 2— L. 7H <: ) we A ; ‘at em) Figure 23. Head in profile. A 7’ gladius (CASENT0406982) B 7’ cognatum (CASENT0067891) C 7. myrmidon (CASENT 0028635) D 7 rumo (CASENT0217759). { ) Wires: —— e- _\ \ 5 rin Ss Figure 25. Mesosoma in profile. A 7’ proximum (CASENT0906146) B 7’ myrmidon (CASENT0028635) C 7. tenuinode (CASENT0040115). : . . _ — . — —— ine Figure 26. Waist segments in profile. A 7 tenuinode (CASENT0162682) B ZT tenuinode (CASENT0040115) © 7 myrmidon (CASENT0028635). Taxonomy of Malagasy Tetramorium 61 Figure 27. Mesosomaand petiole in profile. A 7’ rumo (CASENT0217759) B T aspis (CASENT0344940) C 7. cognatum (CASENT0067891) D T karthala (CASENT0137302). Figure 28. Mesosoma and petiole in profile. A 7 cognatum (CASENT0102343) B ZT cognatum (CASENT0067891) € 7 aspis (CASENT0344940) D 7 karthala (CASENT0136774). Figure 29. Head and mesosoma in profile. A 7’ aspis (CASENT0344940) B T° karthala (CASENT0136774). Tetramorium aspis Hita Garcia & Fisher, sp. n http://zoobank.org/018A7596-2EC0-42 13- B928- D034D77868B4 http://species-id.net/wiki/Tetramorium_aspis Fics 22G32/B 28 C294. 30803 Type material. Holotype, pinned worker, MADAGASCAR, Fianarantsoa, R.S. Ivohibe 8.0 km E Ivohibe, 22.48333°S, 46.96833°E, 1200 m, montane rainforest, sifted litter, (leaf mold, rotten wood), collection code BLF01747, 15.-21.X.1997 (B.L. Fisher) (CAS: CASENT0344940). Paratypes, eleven workers with same data as holotype (CAS: CASENT0189115; CASENT0198990; CASENT0198991; CASENT0198995; CASENT0198998; CASENT0217756; CASENT0247396; CASENT0247397; CASENT0247398; CASENT0247399; CASENT0247400); three workers with same data as holotype except sampled from rotten stick on ground and collection code BLF01746 (CAS: CASENT0189124); fourteen workers from Fianarantsoa, 8.0 km NE Ivohibe, 22.4217°S, 46.8983°E, 1200 m, montane rainfor- est, sifted litter, (leaf mold, rotten wood), collection codes BLFO1751 and BLF01753, 62 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) 3.-9.XI.1997 (B.L. Fisher) (BMNH: CASENT0247403; CAS: CASENT0189123; CASENT0189156; CASENT0198992; CASENT0198993; CASENT0198994; CASENT0198996; CASENT0247402; CASENT0247404; CASENT0247405; MCZ: CASENT0247401). Non-type material, MADAGASCAR: Fianarantsoa, 45 km S Ambalavao, 22.21667°S, 47.01667°E, 785 m, rainforest, 25.[X.1993 (B.L. Fisher); Fianarantsoa, Rés. Andringitra 43 km S Ambalavao, 22.23333°S, 47°E, 825 m, rainforest, 5.X.1993 (B.L. Fisher); Fianarantsoa, Rés. Andringitra 40 km S Ambalavao, 22.21667°S, 46.96667°E, 1275 m, montane rainforest, 15.X.1993 (B.L. Fisher); Fianarantsoa, Rés. Andringitra 38 km S Ambalavao, 22.2°S, 46.96667°E, 1680 m, montane rainforest, 23.X.1993 (B.L. Fisher); Fianarantsoa, R.S. Ivohibe, 7.5 km ENE Ivohibe, 22.47°S, 46.96°E, 900 m, rainforest, 7.-12.X.1997 (B.L. Fisher); Fianarantsoa, R.S. Ivohibe 8.0 km E Ivohibe, 22.48333°S, 46.96833°E, 1200 m, montane rainforest, 15.-21.X.1997 (B.L. Fisher); Fianarantsoa, 8.0 km NE Ivohibe, 22.4217°S, 46.8983°E, 1200 m, montane rainforest, 3.—9.XI.1997 (B.L. Fisher). Diagnosis. The following character combination clearly diagnoses T° aspis within the 7! cognatum species complex: relatively large eyes (OI 25—27); frontal carinae weak- ly developed; propodeal spines short, triangular to elongate-triangular, and acute (PSLI 18-21), propodeal lobes always only weakly shorter than propodeal spines, and spines and lobes strongly inclined towards each other; petiolar node in profile high rounded nodiform, weakly squamiform and relatively thin, in profile around 2.0 to 2.2 times higher than long (LPel 46-51), and in dorsal view around 1.4 to 1.6 times wider than long (DPel 146-161); mesosoma with six or more pairs of long, standing hairs on pro- notum and mesonotum, propodeum usually with one or two pairs located anteriorly. Worker measurements (N=12). HL 0.60-0.63 (0.61); HW 0.51-0.55 (0.54); SL 0.36—0.39 (0.37); EL 0.14-0.15 (0.14); PH 0.26-0.30 (0.28); PW 0.40-0.44 (0.43); WL 0.72-0.81 (0.77); PSL 0.11-0.13 (0.13); PTL 0.11-0.13 (0.12); PTH 0.23-0.25 (0.24); PIW 0.16—-0.19 (0.18); PPL 0.16—0.19 (0.18); PPH 0.23-0.24 (0.24); PPW 0.22—0.24 (0.23); CI 85-88 (87); SI 68-72 (70); OI 25-27 (26); DMI 54-57 (55); LMI 35-37 (36); PSLI 18-21 (21); PeNI 40-44 (42); LPeI 46-51 (49); DPel 146-161 (152); PpNI 51-56 (54); LPpI 70-81 (74); DPpI 126-135 (131); PPI 121-137 (129). Worker description. Head much longer than wide (CI 85-88); in full-face view posterior head margin weakly concave. Anterior clypeal margin with distinct median impression. Frontal carinae weakly developed, only faintly raised, slightly diverging posteriorly, merging with surrounding sculpture halfway between posterior eye mar- gin and posterior head margin. Antennal scrobes weak to absent, shallow and without clear and distinct posterior and ventral margins. Antennal scapes very short, not reach- ing posterior head margin (SI 68-72). Eyes relatively large (OI 25—27). Mesosomal outline in profile flat to weakly convex, comparatively low and long (LMI 35-37), moderately marginate from lateral to dorsal mesosoma; promesonotal suture absent; metanotal groove usually present, but very weakly developed. Propodeal spines short, triangular to elongate-triangular, and acute (PSLI 18-21), propodeal lobes triangular Taxonomy of Malagasy Tetramorium 63 to elongate-triangular, always slightly shorter than propodeal spines, in profile spines and lobes strongly inclined towards each other. Petiolar node in profile high rounded nodiform, weakly squamiform and relatively thin, around 2.0 to 2.2 times higher than long (LPel 46-51), anterior and posterior faces approximately parallel, anterodorsal margin usually situated slightly higher and more angulate than posterodorsal margin, petiolar dorsum relatively flat to weakly convex; node in dorsal view around 1.5 to 1.6 times wider than long (DPel 146-161), in dorsal view pronotum around 2.3 to 2.5 times wider than petiolar node (PeNI 40-44). Postpetiole in profile globular, between 1.2 to 1.4 times higher than long (LPpI 70-81); in dorsal view around 1.3 to 1.4 times wider than long (DPpI 126-135), pronotum around 1.8 to 2.0 times wider than post- petiole (PpNI 51-56). Postpetiole in profile appearing more voluminous than petiolar node, postpetiole in dorsal view around 1.2 to 1.4 times wider than petiolar node (PPI 121-137). Mandibles completely unsculptured, smooth, and shiny; clypeus weakly ir- regularly longitudinally rugulose, median ruga never fully developed, usually reduced to few traces, one or two mostly unbroken lateral rugulae present on each side; cephalic dorsum between frontal carinae longitudinally rugulose with seven to nine rugulae, rugulae usually running from posterior clypeal margin to posterior head margin, often irregularly shaped, interrupted or with cross-meshes; scrobal area mostly unsculptured; lateral head reticulate-rugose to longitudinally rugose, often with larger areas with re- duced sculpture. Ground sculpture on head weakly to moderately reticulate-punctate, especially laterally. Dorsum and sides of mesosoma mostly irregularly longitudinally rugose/rugulose, sides of mesosoma often with some reticulate-rugose areas. Ground sculpture on dorsal mesosoma mostly absent, lateral mesosoma usually weakly to mod- erately reticulate-punctate. Forecoxae usually unsculptured, smooth and shining. Both waist segments and gaster fully unsculptured, smooth, and shining. Dorsum of head with several pairs of long, fine, standing hairs; mesosoma with six or more pairs on pronotum and mesonotum, propodeum usually with one or two pairs anteriorly; waist segments and first gastral tergite without any standing hairs at all; first gastral tergite with short, moderately dense, appressed pubescence. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae with appressed hairs. Head, mesosoma, waist segments and gaster usually of uniform brown to dark brown, appendages yellowish to light brown; sometimes waist segments and gaster lighter than head and mesosoma but still darker than appendages. Etymology. The name of the new species is Old Greek and means “shield,” refer- ring to the weakly squamiform condition of the petiolar node. ‘The species epithet is a nominative noun, and thus invariant. Distribution and biology. So far 7. aspis is known only from a few localities in the southeast of Madagascar around Ivohibe and Andringitra, where it was collected in rainforests or montane rainforests at elevations ranging from 785 to 1680 m (Fig. 63). Tetramorium aspis was mainly sampled from leaf litter or general collecting on the ground, which suggests that it might be a ground-active species. Discussion. The new species is clearly distinguishable within the T. cognatum spe- cies complex. Tetramorium aspis with its large eyes (OI 25-27) and presence of six or 64 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Figure 30. 7° aspis holotype worker (CASENT0344940). A Body in profile B Body in dorsal view C Head in full-face view. more pairs of long, standing hairs on the mesosoma is unlikely to be confused with the small-eyed T° gladius (OI 19-20), or T. freya, which lacks standing pilosity on the mesosomal dorsum. Furthermore, the weakly developed frontal carinae separate it Taxonomy of Malagasy Tetramorium 65 from the species with strong frontal carinae 7. myrmidon, T. proximum, and T. tenu- inode. he latter three are also mostly larger species (HW 0.58-0.81; WL 0.76-1.07) compared to T° aspis (HW 0.51—-0.55; WL 0.72-0.81). The remaining four species are morphologically closer to T° aspis. Of these, T. camelliae differs significantly from T- aspis on the basis of petiolar node development. In the latter the node is only weakly squamiform, in profile around 2.0 to 2.2 times higher than long (LPel 46-51), and in dorsal view between 1.4 to 1.6 times wider than long (DPel 146-161). This shape contrasts with the node of T° camelliae, which is around 2.8 to 3.0 times higher than long (LPel 33-36) and around 2.3 to 2.4 times wider than long (DPel 228-238). The best character for separating 7° aspis from the remaining three species is the develop- ment and arrangement of propodeal spines and lobes. In 7° aspis the spines are short, triangular to elongate-triangular, and acute (PSLI 18-21), the lobes relatively long, only weakly shorter than the spines, and the spines and lobes are strongly inclined towards each other. This arrangement is not seen in 7! cognatum, T. karthala or T: rumo (it is present in 7. camelliae though) since they either have short to moderate spines (T: karthala + T. rumo: PSLI 20-26), much longer than the lobes, or the spines are very short (7. cognatum: PSLI 12-16) and approximately same length as lobes or even shorter. Beyond this conspicuous character, 7. aspis also has longer antennal scapes (SI 68-72), a broader petiolar node (DPel 146-161), and a narrower postpe- tiole (DPpI 126-135) than T. cognatum (SI 61-67; DPel 129-142; DPpI 137-153). Tetramorium karthala, which is only found on the Comoros, has only two to four pairs of long, standing hairs on the pronotum and mesonotum while 7° aspis has least least six pairs. The last species, 7: rumo, has very short antennal scapes (SI 60-66), a thinly cuneiform and moderately squamiform petiolar node, and is usually very bright yellow to light brown. Intriguingly, 7: scutum from the T. sikorae complex, which is endemic to the same geographic area, bears a strong superficial resemblance to 7: aspis. However, they are in different species complexes based on their differences in pilosity on the first gastral tergite (present in 7: scutum but absent in 7: aspis). Apart from this key character they also differ slightly in eye size and propodeal spine length. To our best knowledge, there is no apparent intraspecific variation in T° aspis. Tetramorium camelliae Hita Garcia & Fisher, sp. n. http://zoobank.org/E836FA0A-EA89-46D0-86A1-5518D9075C98 http://species-id.net/wiki/Tetramorium_camelliae Bigs 22A,.3),-63 Type material. Holotype, pinned worker, MADAGASCAR, Fianarantsoa, Ranomafa- na National Park, Vohiparara, 21.24444°S, 47.39694°E, 1173 m, montane forest, 16.1X.1992 (E. Rajeriarison) (CAS: CASENT0247496). Paratypes, four pinned work- ers with same data as holotype (BMNH: CASENT0247495; CAS: CASENT0247499; MCZ: CASENT0247498, CASENT0247500). 66 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Non-type material. MADAGASCAR: Fianarantsoa, Ranomafana National Park, Vohiparara, 21.24444°S, 47.39694°E, 1173 m, montane forest, 16.[X.1992 (E. Raje- riarison); Fianarantsoa, Ranomafana National Park, Vatoharanana River, 4.1 km 231° SW Ranomafana, 21.29°S, 47.43333°E, 1100 m, montane rainforest, 27.31 .III.2003 (B.L. Fisher et al.). Diagnosis. The strongly squamiform and transverse petiolar node, which in pro- file is around 2.7 to 3.0 times higher than long (LPel 33-36), and in dorsal view around 2.3 to 2.4 times wider than long (DPel 228-238), isolates T° camelliae from the other members of the 7. cognatum species complex. Worker measurements (N=7). HL 0.56—0.63 (0.61); HW 0.50—0.57 (0.55); SL 0.33—0.40 (0.38); EL 0.13—-0.15 (0.14); PH 0.25—0.29 (0.27); PW 0.40-0.44 (0.43); WL 0.67-0.78 (0.74); PSL 0.10—0.12 (0.10); PTL 0.07—0.09 (0.08); PTH 0.21-0.25 (0.24); PI'W 0.16-0.21 (0.19); PPL 0.15-0.20 (0.18); PPH 0.21-0.25 (0.24); PPW 0.20—-0.26 (0.24); CI 89-91 (91); SI 66-70 (68); OI 25-26 (26); DMI 56-60 (57); LMI 36-37 (37); PSLI 17-19 (18); PeNI 40-48 (45); LPelI 33-36 (35); DPel 228-238 (234); PpNI 50-59 (57); LPpI 69-80 (74); DPpI 133-141 (137); PPI 120-134 (126). Worker description. Head much longer than wide (CI 89-91); in full-face view posterior head margin weakly concave. Anterior clypeal margin with distinct medi- an impression. Frontal carinae moderately developed, diverging posteriorly, merging with surrounding sculpture halfway between posterior eye margin and posterior head margin. Antennal scrobes weak, shallow and without clear and distinct posterior and ventral margins. Antennal scapes very short, not reaching posterior head margin (SI 66-70). Eyes relatively large (OI 25-26). Mesosomal outline in profile flat to weakly convex, comparatively low and long (LMI 36-37), moderately marginate from lateral to dorsal mesosoma; promesonotal suture absent; metanotal groove present, but weakly developed. Propodeal spines short, triangular to elongate-triangular, and acute (PSLI 17-19), propodeal lobes triangular to elongate-triangular and of approximately same length as propodeal spines, in profile spines and lobes strongly inclined towards each other. Petiolar node in profile strongly squamiform, around 2.7 to 3.0 times higher than long (LPeI 33-36), anterior and posterior faces approximately parallel, anterodorsal and posterodorsal margins situated at about same height, petiolar dorsum relatively flat to weakly convex; node in dorsal view transverse and 2.3 to 2.4 times wider than long (DPel 228-238), in dorsal view pronotum between 2.1 to 2.5 times wider than petiolar node (PeNI 40-48). Postpetiole in profile subglobular, between 1.2 to 1.4 times higher than long (LPpI 69-80); in dorsal view around 1.3 to 1.4 times wider than long (DPpl 133-141), pronotum between 1.7 to 2.0 times wider than postpetiole (PpNI 50-59). Postpetiole in profile much more voluminous than petiolar node, postpetiole in dorsal view around 1.2 to 1.4 times wider than petiolar node (PPI 120-134). Mandibles com- pletely unsculptured, smooth, and shiny; clypeus irregularly longitudinally rugulose, median ruga never fully developed, usually reduced to few traces posteriorly or medi- ally, one or two mostly unbroken lateral rugulae present on each side; cephalic dorsum between frontal carinae longitudinally rugose/rugulose with seven to eight rugae/rugu- lae, rugae usually running unbroken from posterior clypeal margin to posterior head Taxonomy of Malagasy Tetramorium 67 Figure 31. 7) camelliae holotype worker (CASENT0247496). A Body in profile B Body in dorsal view C Head in full-face view. 68 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) margin, sometimes interrupted or with cross-meshes; scrobal area partly unsculptured, but mostly merging with surrounding reticulate-rugose to longitudinally rugose sculp- ture present on lateral head. Dorsum and sides of mesosoma longitudinally rugose/ rugulose. Forecoxae unsculptured, smooth and shining. Ground sculpture on head and mesosoma weak to absent. Both waist segments and gaster fully unsculptured, smooth, and shining. Dorsum of head with several pairs of long, fine, standing hairs; mesosoma with seven to eight pairs restricted to pronotum and mesonotum; propodeum, waist segments and first gastral tergite without any standing hairs; first gastral tergite with short, moderately dense, appressed pubescence. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae with appressed hairs. Head, mesosoma, waist seg- ments, and gaster uniform brown to dark brown, appendages yellowish to light brown. Etymology. ‘The new species is dedicated to the first author’s lovely life partner, Tracy Lynn Camellia Audisio from San Francisco, California, U.S.A. Distribution and biology. Tetramorium camelliae is a rare species only known from Ranomafana National Park (Fig. 63) where it was collected twice in montane rainforest at elevations ranging from 1100 to 1173. The little available data indicates that 7: camelliae lives in leaf litter. Discussion. Tetramorium camelliae is easily distinguishable from the other mem- bers of the 7: cognatum species complex. The main diagnostic character is the petiolar node shape, which is strongly squamiform in T: camelliae (LPel 33-36; DPel 228- 238) but not in the remainder of the species complex (LPel 37-62; DPel 111-171). Generally, its smaller body size, weaker frontal carinae, and gastral pubescence ally T- camelliae morphologically with T: aspis, T: cognatum, T. karthala and T. rumo. Tetramorium cognatum Bolton, 1979 http://species-id.net/wiki/Tetramorium_cognatum Figs:24 By. 23B; 27C, 28A, By 32,63 Tetramorium cognatum Bolton, 1979:135. Type material. Holotype, pinned worker, MADAGASCAR, Toamasina, Perinet & vicinity, 18.82667°S, 48.44778°E, rainforest, rotten wood, 19.III.1969 (WL. Brown) (MCZ: MCZ_Holotype_32261) [examined]. Paratypes, eleven pinned workers with same data as holotype except collected from 17.-19.III.1969 (BMNH: CASENT0102343, CASENT0102344, CASENT0235218; MCZ: MCZ._Para- type_32261; NHMB; MHNG: CASENT0911248) [all examined, except NHMB]. [Note: the GPS data of the type locality was not provided either by the locality label or the original description. The data presented above is based on our own geo- referencing of the Périnet=Andasibe area and should be considered an approximation and not the exact position of the type locality.] Non-type material. MADAGASCAR: no loc (Staudinger); Antananarivo, Réserve Spéciale d’Ambohitantely, Forét d’Ambohitantely, 20.9 km 72° NE Ankazobe, Taxonomy of Malagasy Tetramorium 69 18.22528°S, 47.28683°E, 1410 m, montane rainforest, 17.-21.IV.2001 (B.L. Fisher et al.); Antananarivo, Réserve Speciale d’Ambohitantely, 18.18762°S, 47.28576°E, 1580 m, montane forest, 8.[II.2012 (B.L. Fisher et al.); Antananarivo, Réserve Speciale d’Ambohitantely, 18.22444°S, 47.2774°E, 1490 m, montane forest, 9.II.2012 (B.L. Fisher et al.); Antananarivo, 3 km 41° NE Andranomay, 11.5 km 147° SSE Anjozo- robe, 18.47333°S, 47.96°E, 1300 m, montane rainforest, 5.-13.XII.2000 (B.L. Fisher et al.); Antananarivo, Ankalalahana, 19.00711°S, 47.1337°E, 1350 m, Uapaca wood- land, 26.111.2011 (BL. Fisher et al); Antananarivo, Mandraka Park, 18.9019°S, 47 .90786°E, 1360 m, montane shrubland, 11.11.2012 (BL. Fisher et al.); Antanana- rivo, Réserve Naturelle Sohisika, Sohisika 24.6 km NNE Ankazobe, 18.10322°S, 47.18692°E, 1464 m, gallery montane forest, 1.—2.V1.2008 (B.L. Fisher et al.); Anta- nanarivo, Forét de galerie, Telomirahavavy, 23.4 km NNE Ankazobe, 18.12167°S, 47.20627°E, 1520 m, disturbed gallery montane forest, 3.—4.V1.2008 (B.L. Fisher et al.); Antsiranana, Ambondrobe, 41.1 km 175° Vohemar, 13.71533°S, 50.10167°E, 10 m, littoral rainforest, 30.XI.2004 (B.L. Fisher); Antsiranana, Parc National Mont. d’Ambre, 1050 m, 12.11.1977 (W.L. & D.E. Brown); Antsiranana, Parc National Montagne d’Ambre, 3.6 km 235° SW Joffreville, 12.53444°S, 49.1795°E, 925 m, montane rainforest, 20.—26.1.2001 (BL. Fisher et al.); Antsiranana, Parc National Montagne d’Ambre, 12.2 km 211° SSW Joffreville, 12.59639°S, 49.1595°E, 1300 m, montane rainforest, 2.—7.I].2001 (B.L. Fisher et al.); Antsiranana, Parc National Mon- tagne d’Ambre, 12.2 km 211° SSW Joffreville, 12.59639°S, 49.1595°E, 1300 m, mon- tane rainforest, 7.11.2001 (GD. Alpert); Antsiranana, Parc National Montagne d’Ambre, 12.51778°S, 49.17957°E, 1000 m, montane rainforest, 3.—7.II1.2001 (BL. Fisher et al.); Antsiranana, Parc National Montagne d’Ambre, 12.52861°S, 49.17717°E, 1100 m, montane rainforest, 12.II].2001 (B.L. Fisher et al.); Antsiranana, Parc Na- tional Montagne d’Ambre, Ambre grand lac, 12.59656°S, 49.15932°E, 1350 m, mon- tane rainforest, 13.X1.2007 (B.L. Fisher et al.); Antsiranana, Parc National Montagne d’ Ambre, Lac maudit, 12.58502°S, 49.15147°E, 1250 m, montane rainforest, 13.—14. X1.2007 (B.L. Fisher et al.); Antsiranana, Parc National Montagne d’Ambre, Rous- settes, 12.52574°S, 49.17238°E, 1025 m, montane rainforest, 15.XI.2007 (B.L. Fisher et al.); Antsiranana, Parc National Montagne d’Ambre, Petit lac, 12.53664°S, 49.17412°E, 1130 m, montane rainforest, 17.XI.2007 (B.L. Fisher et al.); Antsiranana, Parc National Montagne d’Ambre, Antomboka, 12.51269°S, 49.17807°E, 970 m, montane rainforest, 17.X1.2007 (B.L. Fisher et al.); Antsiranana, Parc National Mon- tagne d’Ambre, Mahasarika, 12.53176°S, 49.17662°E, 1135 m, montane rainforest, 19.X1.2007 (B.L. Fisher et al.); Antsiranana, Parc National Montagne d’Ambre, Pic Bades, 12.5186°S, 49.18625°E, 900 m, montane rainforest, 20.X1.2007 (B.L. Fisher et al.); Antsiranana, Rés. Anjanaharibe-Sud, 17 km W Andapa, 15°45'27.9"S, 49°30'36.7"E, 875 m, rainforest, 5.X1.1994 (G.D. Alpert); Antsiranana, Rés. Anjana- haribe-Sud, 6.5 km SSW Befingotra, 14.75°S, 49.5°E, 875 m, rainforest, 17.-31.X.1994 (B.L. Fisher); Antsiranana, Rés. Anjanaharibe-Sud, 9.2 km WSW Befingotra, 14.75°S, 49.46667°E, 1180-1200 m, montane rainforest, 7.—9.X1.1994 (B.L. Fisher); Antsira- nana, Anjanaharibe, 1.II.-1.I11.2003 (KA. Jackson & D. Carpenter); Antsiranana, 70 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Betaolana Forest, along Bekona River, 14.52996°S, 49.44039°E, 880 m, rainforest, 4—5.II1.2009 (B.L. Fisher et al.); Antsiranana, Forét de Binara, 9.4 km 235° SW Daraina, 13.26333°S, 49.6°E, 1100 m, montane rainforest, 5.XII.2003 (B.L. Fisher); Antsiranana, R.S. Manongarivo, 10.8 km 229° SW Antanambao, 13.96167°S, 48.43333°E, 400 m, rainforest, 8.—13.X.1998 (B.L. Fisher); Antsiranana, R.S. Manon- garivo, 12.8 km 228° SW Antanambao, 13.97667°S, 48.42333°E, 780 m, rainforest, 11-17.X.1998 (B.L. Fisher); Antsiranana, R.S. Manongarivo, 14.5 km 220° SW Anta- nambao, 13.99833°S, 48.42833°E, 1175 m, montane rainforest, 20.X.1998 (B.L. Fisher); Antsiranana, R.N.I. Marojejy, 14°26'43.2"S, 49°47'8.3"E, 375 m, 23.X1.1993 (G.D. Alpert); Antsiranana, Parc National de Marojejy, Manantenina River, 27.6 km 35° NE Andapa, 9.6 km 327° NNW Manantenina, 14.435°S, 49.76°E, 775 m, rain- forest, 15.XI.-11.XII.2005 (B.L. Fisher et al.); Antsiranana, Parc National de Marojejy, Antranohofa, 26.6 km 31° NNE Andapa, 10.7 km 318° NW Manantenina, 14.44333°S, 49.74333°E, 1325 m, montane rainforest, 19.X1I.2003 (B.L. Fisher); Ant- siranana, Sakaramy, 12.44114°S, 49.23197°E, 260 m, tropical dry forest, 11.V.2011 (B.L. Fisher et al.); Fianarantsoa, Rés. Andringitra 40 km S Ambalavao, 22.21667°S, 46.96667°E, 1225 m, montane rainforest, 19.X.1993 (B.L. Fisher); Fianarantsoa, Rés. Andringitra 44 km S Ambalavao, 22.23333°S, 47°E, 800 m, rainforest, 11.X.1993 (B.L. Fisher); Fianarantsoa, Ampandravelo II Non Protected Area, 10.78 km NE Ranohira, 22.53917°S, 45.51548°E, 873 m, shrubland, 20.—22.I].2010 (A. Raveloma- nana); Fianarantsoa, Parc National Befotaka-Midongy, Papango 27.7 km S Midongy- Sud, Mount Papango, 23.83517°S, 46.96367°E, 940 m, rainforest, 14.-16.X1.2006 (B.L. Fisher et al.); Fianarantsoa, Parc National Befotaka-Midongy, Papango 28.5 km S Midongy-Sud, Mount Papango, 23.84083°S, 46.9575°E, 1250 m, montane rainfor- est, 17.-18.X1.2006 (B.L. Fisher et al.); Fianarantsoa, Isalo National Park, Canyon de Sinze, 22°28’ S, 45°33’ E, 800 m, forest, 17.11.1993 (E. Rajeriarison); Fianarantsoa, Isalo IV National Parc, 12 km SW Ranohira, 22.61472°S, 45.31304°E, 867 m, Bis- marckia woodland, 25.11.2010 (A. Ravelomanana); Fianarantsoa, R.S. Ivohibe, 7.5 km ENE Ivohibe, 22.47°S, 46.96°E, 900 m, rainforest, 7.-12.X.1997 (B.L. Fisher); Fianarantsoa, R.S. Ivohibe 8.0 km E Ivohibe, 22.48333°S, 46.96833°E, 1200 m, montane rainforest, 15.-21.X.1997 (B.L. Fisher); Fianarantsoa, 9.0 km NE Ivohibe, 22.42667°S, 46.93833°E, 900 m, rainforest, 12.-17.X.1997 (B.L. Fisher); Fianarant- soa, Manandriana I Non Protected Area, 27.11 km SW Ambositra, 20.73194°S, 47.09413°E, 1590 m, savannah grassland, 9.-11.11.2010 (A. Ravelomanana); Fianarantsoa, Parc National de Ranomafana, Vatoharanana River, 4.1 km 231° SW Ranomafana, 21.29°S, 47.43333°E, 1100 m, montane rainforest, 27.—31.2003 (B.L. Fisher et al.); Fianarantsoa, Parc National Ranomafana, Talatakely, 21.24833°S, 47.42667°E, in guava forest, 9.-26.IV.1998 (CE. Griswold et al.); Fianarantsoa, Parc National de Ranomafana, Sahamalaotra River, 6.6 km 310° NW Ranomafana, 21.23667°S, 47.39667°E, 1150 m, montane rainforest, 31.[I.2003 (B.L. Fisher et al.); Fianarantsoa, Forét Classée Vatovavy, 7.6 km 122° Kianjavato, 21.4°S, 47.94°E, 175 m, rainforest, 6.-8.VI.2005 (B.L. Fisher et al.); Fianarantsoa, Forét de Vevembe, 66.6 km 293° Farafangana, 22.791°S, 47.18183°E, 600 m, rainforest, transition to mon- Taxonomy of Malagasy Tetramorium Fal tane forest, 23.1V.2006 (B.L. Fisher et al.); Mahajanga, Réserve Spéciale Marotandra- no, Marotandrano 48.3 km S Mandritsara, 16.28322°S, 48.81443°E, 865 m, transi- tion humid forest, 7.X1.2007 (B.L. Fisher et al.); Toamasina, Montagne d’Akirindro 7.6 km 341° NNW Ambinanitelo, 15.28833°S, 49.54833°E, 600 m, rainforest, 17.— 21.111.2003 (B.L. Fisher et al.); Toamasina, Ambanizana, Parc National Masoala, 15.57167°S, 50.00611°E, 848 m, montane rainforest, 26.II.-2.II].2003 (D. Andria- malala et al.); Toamasina, 5.3 km SSE Ambanizana, Andranobe, 15.67133°S, 49.97395°E, 425 m, rainforest, 21.XI.1993 (B.L. Fisher); Toamasina, 6.3 km S Am- banizana, Andranobe, 15.6813°S, 49.958°E, 25 m, rainforest, 23.X1.1993 (B.L. Fish- er); Toamasina, 6.9 km NE Ambanizana, Ambohitsitondroina, 15.58506°S, 50.00952°E, 825 m, rainforest, 2-8.XII.1993 (B.L. Fisher); Toamasina, Réserve Spé- ciale Ambatovaky, Sandrangato river, 16.7633°S, 49.26692°E, 520 m, rainforest, 22.— 24.11.2010 (B.L. Fisher et al.); Toamasina, Forét Ambatovy, 14.3 km 57° Moramanga, 18.85083°S, 48.32°E, 1075 m, montane rainforest, 21.]1.2004-12.1V.2005 (B.L. Fisher et al.); Toamasina, Ambatovy, 12.4 km NE Moramanga, 18.84963°S, 48.2947°E, 1010 m, montane rainforest, 3.—6.III.2007 (B.L. Fisher et al.); Toamasina, Ambatovy, 12.4 km NE Moramanga, 18.83937°S, 48.30842°E, 1080 m, montane rainforest, 4.-7.I]].2007 (BL. Fisher et al.); Toamasina, Station Forestiére Anal- amazaotra, Analamazaotra 1.3 km S Andasibe, 18.38466°S, 48.41271°E, 980 m, montane rainforest, 11.-13.X1.2007 (B.L. Fisher et al.); Toamasina, Analamay, 18.80623°S, 48.33707°E, 1068 m, montane rainforest, 21.[11.2004 (Malagasy ant team); Toamasina, F.C. Andriantantely, 18.695°S, 48.81333°E, 530 m, rainforest, 4—10.X11.1998 (HJ. Ratsirarson); Toamasina, Montagne d’Anjanaharibe, 18.0 km 21° NNE Ambinanitelo, 15.18833°S, 49.615°E, 470 m, rainforest, 8.—12.I]I.2003 (B.L. Fisher et al.); Toamasina, Montagne d’Anjanaharibe, 19.5 km 27° NNE Ambi- nanitelo, 15.17833°S, 49.635°E, 1100 m, montane rainforest, 12.—16.II.2003 (B.L. Fisher et al.); Toamasina, Reserve Betampona, Camp Vohitsivalana, 37.1 km 338° Toamasina, 17.88667°S, 49.2025°E, 520 m, rainforest, 1.—3.XII.2005 (B.L. Fisher et al.); Toamasina, Reserve Betampona, Camp Rendrirendry, 34.1 km 332° Toamasina, 17.924°S, 49.19967°E, 390 m, rainforest, 28.X1.2005 (B.L. Fisher et al.); Toamasina, Bevolota, 17.1 km N Andasibe, 18.77071°S, 48.43164°E, 995 m, montane rainforest, 12.X1I.2007 (B.L. Fisher et al.); Toamasina, F.C. Didy, 18.19833°S, 48.57833°E, 960 m, rainforest, 16.—23.XII.1998 (HJ. Ratsirarson); Toamasina, P.N. Mantadia, 18.79167°S, 48.42667°E, 895 m, rainforest, 25.X].-1.XII.1998 (ALJ. Ratsirarson); Toamasina, Sahafina Forest, 11.4 km W Brickaville, 18.81445°S, 48.96205°E, 140 m, rainforest, 13.-14.X]I.2007 (BL. Fisher et al); Toamasina, F.C. Sandranantitra, 18.04833°S, 49.09167°E, 450 m, rainforest, 18.—21.1.1999 (HJ. Ratsirarson); Toamasina, Torotorofotsy, 18.87082°S, 48.34737°E, 1070 m, montane rainforest, marsh edge, 24.—29.III.2004 (Malagasy ant team); Toamasina, Parc National de Zaha- mena, Onibe River, 17.75908°S, 48.85468°E, 780 m, rainforest, 21.—23.II.2009 (B.L. Fisher et al.); Toamasina, Parc National de Zahamena, Tetezambatana forest, near junction of Nosivola and Manakambahiny Rivers, 17.74298°S, 48.72936°E, 860 m, rainforest, 18—19.11.2009 (BL. Fisher et al); Toliara, Réserve Spéciale iD Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) d’Ambohijanahary, Forét d’Ankazotsihitafototra, 35.2 km 312° NW Ambaravaranala, 18.26667°S, 45.40667°E, 1050 m, montane rainforest, 13.—-17.1.2003 (B.L. Fisher et al.); Toliara, Réserve Spéciale d’Ambohijanahary, Forét d’Ankazotsihitafototra, 34.6 km 314° NW Ambaravaranala, 18.26°S, 45.41833°E, 1100 m, montane rainforest, 16.1.2003 (BL. Fisher et al.); Toliara, Rés. Andohahela, 11 km NW Enakara, 24.56667°S, 46.83333°E, 800 m, rainforest, 17.XI.1992 (B.L. Fisher); Toliara, Rés. Andohahela, 10 km NW Enakara, 24.56667°S, 46.81667°E, 420 m, rainforest, 19. X1.1992 (B.L. Fisher); Toliara, Rés. Andohahela, 10 km NW Enakara, 24.56667°S, 46.81667°E, 430 m, rainforest, 25.X1.1992 (B.L. Fisher); Toliara, Rés. Andohahela, 13 km NW Enakara, 24.55°S, 46.8°E, 1250 m, montane rainforest, 30.X1.1992 (B.L. Fisher); Yoliara, Rés. Andohahela, 13 km NW Enakara, 24.55°S, 46.8°E, 1300 m, montane rainforest, 2.X1I.1992 (B.L. Fisher); Toliara, Rés. Andohahela, 13 km NW Enakara, 24.56667°S, 46.81667°E, 900 m, rainforest, 3.XII.1992 (B.L. Fisher); Toli- ara, Rés. Andohahela, 3 km E Mahamavo, 24°45’ S, 46°45’ E, 1050 m, montane rainforest, 5.X1.1993 (P.S. Ward); Toliara, Parc National d’Andohahela, Col du Se- dro, 3.8 km 113° ESE Mahamavo, 37.6 km 341° NNW Tolagnaro, 24.76389°S, 46.75167°E, 900 m, montane rainforest, 21.—25.].2002 (B.L. Fisher et al.); Toliara, Parc National d’Andohahela, Manampanihy River, 5.4 km 113° ESE Mahamavo, 36.7 km 343° NNW Tolagnaro, 24.76389°S, 46.76683°E, 650 m, rainforest, 24.1.2002 (B.L. Fisher et al.); Toliara, Forét lvohibe 55.6 km N Tolagnaro, 24.56167°S, 47.20017°E, 650 m, rainforest, 4.XII.2006 (B.L. Fisher et al.); Toliara, Réserve Spé- ciale Kalambatritra, Befarara, 23.4178°S, 46.4478°E, 1390 m, montane rainforest, 78.11.2009 (B.L. Fisher et al.); Toliara, Réserve Spéciale Kalambatritra, Betanana, 23.4144°S, 46.459°E, 1360 m, montane rainforest, 8.I].2009 (B.L. Fisher et al.); To- liara, Réserve Spéciale Kalambatritra, Ampanihy, 23.4635°S, 46.4631°E, 1270 m, montane rainforest, 9.-10.1I.2009 (B.L. Fisher et al.); Toliara, Réserve Spéciale Kalam- batritra, Ampanihy, 23.463°S, 46.47057°E, 1269 m, montane rainforest, 10.11.2009 (B.L. Fisher et al.); Toliara, Réserve Spéciale Kalambatritra, Ambinanitelo, 23.4502°S, 46.45658°E, 1325 m, montane rainforest, 11.11.2009 (B.L. Fisher et al.); Toliara, Grand Lavasoa, 25.9 km W Tolagnaro, 25.08767°S, 46.749°E, 450 m, rainforest, 30.XI.-2.XII.2006 (B.L. Fisher etal); Toliara, 4.4 km 148° SSE Lavanono, 25.45056°S, 44.97417°E, 60 m, spiny forest/thicket, 17.11.2002 (B.L. Fisher et al.); Toliara, 2.7 km WNW 302° Ste. Luce, 24.77167°S, 47.17167°E, 20 m, littoral rainforest, 9.-11. XII.1998 (B.L. Fisher); Toliara, Manatantely, 8.9 km NW Tolagnaro, 24.9815°S, 46.92567°E, 100 m, rainforest, 27.XI.2006 (B.L. Fisher et al.). Diagnosis. Tetramorium cognatum can be easily separated from the other mem- bers of the 7: cognatum species group by the following character combination: eyes very large (OI 27-29); antennal scapes very short (SI 61-67); frontal carinae weakly developed; propodeal spines reduced to very short, triangular teeth (PSLI 12-16), spines and lobes usually of approximately same length, often spines weakly shorter than lobes, very rarely spines weakly longer than lobes, never strongly directed towards each other; petiolar node high nodiform, in profile around 1.8 to 2.0 times higher than long (LPel 49-55), in dorsal view around 1.3 to 1.4 times wider than long (DPel Taxonomy of Malagasy Tetramorium 73 129-142); dorsal mesosoma normally with four to six (sometimes reduced to two to three) pairs of long, standing hairs occurring from anterior pronotum to metanotal groove, but absent from propodeum. Worker measurements (N=25). HL 0.53-0.61 (0.57); HW 0.48—0.54 (0.51); SL 0.31-0.36 (0.33); EL 0.13-0.15 (0.14); PH 0.25—-0.29 (0.27); PW 0.37-0.42 (0.39); WL 0.65-0.75 (0.70); PSL 0.07—0.10 (0.08); PTL 0.11-0.13 (0.12); PTH 0.21-0.25 (0.23); PTW 0.15-0.17 (0.16); PPL 0.14-0.16 (0.15); PPH 0.20-0.23 (0.21); PPW 0.20—0.23 (0.22); CI 87-91 (90); SI 61-67 (65); OI 27-29 (28); DMI 54-59 (56); LMI 36—40 (38); PSLI 12-16 (14); PeNI 38-44 (41); LPel 49-55 (52); DPel 129-142 (135); PpNI 53-59 (56); LPpI 66-75 (70); DPpI 137-153 (145); PPI 131-147 (137). Worker description. Head much longer than wide (CI 87-91); in full-face view posterior head margin usually very weakly concave. Anterior clypeal margin with dis- tinct median impression. Frontal carinae weakly developed, only faintly raised, slightly diverging posteriorly, and relatively long, often ending shortly before posterior head margin. Antennal scrobes very weakly developed, shallow and without clear and dis- tinct posterior and ventral margins. Antennal scapes very short, not reaching posterior head margin (SI 61-67). Eyes very large (OI 27-29). Mesosomal outline in profile flat to weakly convex, comparatively low and long (LMI 36-40), moderately margin- ate from lateral to dorsal mesosoma; promesonotal suture absent; metanotal groove usually present, but relatively weak. Propodeal spines reduced to very short, triangular teeth (PSLI 12-16), propodeal lobes short and triangular, spines and lobes usually of approximately same length, often spines weakly shorter than lobes, very rarely spines weakly longer than lobes. Petiolar node in profile rounded high nodiform, around 1.8 to 2.0 times higher than long (LPel 49-55), anterior and posterior faces approxi- mately parallel, anterodorsal margin usually sharper than more rounded posterodorsal margin, both margins often situated at about same height, often anterodorsal margin slightly higher, petiolar dorsum usually weakly convex; node in dorsal view around 1.3 to 2 times wider than long (DPel 129-142), in dorsal view pronotum between 2.3 to 2.7 times wider than petiolar node (PeNI 38-44). Postpetiole in profile globular, rarely subglobular, between 1.3 to 1.5 times higher than long (LPpI 66-75); in dorsal view around 1.3 to 1.5 times wider than long (DPpI 137-153), pronotum between 1.7 to 1.9 times wider than postpetiole (PpNI 53-59). Postpetiole in profile usu- ally appearing shorter and less voluminous than petiolar node, postpetiole in dorsal view between 1.3 to 1.5 times wider than petiolar node (PPI 131-147). Mandibles completely unsculptured, smooth, and shiny; clypeus usually irregularly longitudinally rugulose, median ruga often unbroken and well developed, often partly reduced, and often fully absent; lateral rugulae ranging from one to three on each side, often irregu- larly shaped, broken, or reduced to traces; cephalic dorsum between frontal carinae longitudinally rugulose with six to ten fine rugulae, rugulae usually running from pos- terior clypeal margin to posterior head margin, mostly irregularly shaped, interrupted, meandering or with cross-meshes, and sometimes becoming much weaker posteriorly; scrobal area mostly unsculptured, merging with surrounding sculpture; lateral head 74 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) reticulate-rugose to longitudinally rugose, often with larger areas of reduced sculpture; ground sculpture on head weakly to moderately punctate, sometimes completely ab- sent. Dorsum of mesosoma irregularly longitudinally rugulose, sometimes weakly so; lateral mesosoma usually mostly irregularly longitudinally rugulose, lateral pronotum often much weaker-sculptured, almost smooth, and sometimes reticulate-rugulose; ground sculpture on mesosoma usually weakly to moderately punctate, sometimes completely absent. Forecoxae usually unsculptured, smooth and shining, sometimes with traces of ground sculpture dorsally. Both waist segments and gaster fully unsculp- tured, smooth, and shining. Dorsum of head with several pairs of long, standing hairs, dorsal mesosoma normally with four to six (sometimes reduced to two to three) pairs occurring from anterior pronotum to metanotal groove, but absent from propodeum; waist segments and first gastral tergite without any long, standing hairs at all; first gas- tral tergite with moderately long, dense, appressed to decumbent pubescence. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae with appressed to subdecumbent hairs. Body colouration variable, ranging from uniformly bright yellow to very dark brown, if colour brown to dark brown, appendages usually lighter. Distribution and biology. This species is one of the most common Tetramorium encountered in the rainforests and montane rainforests of Madagascar (Fig. 63). Its distribution range encompasses almost all sampled forests in eastern and northern Madagascar, as well as the isolated humid forests in central and western parts of the island (e.g. Ambohijanahary, Isalo). Surprisingly, 7° cognatum also seems to do fairly well outside humid forests since it is also found in a few disturbed gallery forests, in Uapaca woodland and tropical dry forests, and very rarely in spiny forest/thicket and savannah/grassland. Additional sampling in arid habitats might show that T. cognatum is even more widely distributed than currently understood. ‘The species was mostly col- lected from leaf litter and the ground. Discussion. Tetramorium cognatum is an important species within this species complex. As outlined above, it is very widely distributed and common, and co-occurs in sympatry with almost all other members of the complex. It is easily distinguishable from the usually larger T° freya, which also lacks pilosity on the dorsal mesosoma, and T. gladius, the species with the smallest eyes in the complex (OI 19-20). Furthermore, T. cognatum possesses relatively weakly developed frontal carinae, separating it from I. myrmidon, T. proximum, and T. tenuinode. This character allies 7. cognatum with T. aspis, T. camelliae, T. karthala, and T. rumo, to which it is generally closer in mor- phology. Of these species close to T° cognatum, T. karthala (endemic to Grand Comore) has longer antennal scapes (SI 70-74) and propodeal spines (PSLI 20—22) than 7! cog- natum (SI 61-67; PSLI 12-16). Tetramorium camelliae, only found in Ranomafana, and 7: rumo have squamiform or thinly cuneiform petiolar nodes, which in profile are between 2.3 to 3.0 times higher than long (LPel 33-43) while the node of 7! cognatum is only 1.8 to 2.0 times higher than long (LPel 49-55). Tetramorium aspis, which is only found in the area around Andringitra and Ivohibe, has longer propodeal spines (PSLI 18-21) than T° cognatum. More importantly however, the propodeal spines and lobes of T. cognatum are not strongly inclined towards each other as they are in T° aspis. Taxonomy of Malagasy Tetramorium 75 " vd i Figure 32. 7) cognatum paratype worker (CASENT0102343). A Body in profile B Body in dorsal view C Head in full-face view. Considering its local abundance, relative flexibility in habitat requirements, and widespread distribution, 7° cognatum shows startlingly little intraspecific variation. From the southern spiny forest Lavanono and the montane rainforests in Andohahela 76 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) to the northernmost known localities at Montagne d’Ambre/Sakaramy, 7: cognatum is always easily recognisable. However, some noticeable variation in sculpture and body colouration still exists. Colouration ranges from very light yellow through all shades of yellowish orange or brown to very dark brown, almost black. The southeastern and northern populations are mostly light yellow to light brown while most material from the central-eastern populations is mainly brown to very dark brown. There is also some variation in the degree to which sculpture is developed on the clypeus and the sides of the head and mesosoma as outlined in the description above. Tetramorium freya Hita Garcia & Fisher, sp. n. http://zoobank.org/65E4760A-E053-4CB 1-809A-7 DEG9A408EE3 http://species-id.net/wiki/Tetramorium_freya Figs 21A, 33, 63 Type material. Holotype, pinned worker, MADAGASCAR, Antsiranana, Diego-Su- arez, 7 km N Joffreville [camp 2 of Fisher], 12.33333°S, 49.25°E, 360 m, in dry forest, Malaise trap, collection code MA-01-07-08, 6.—20.[1I.2001 (R. Harin’Hala) (CAS: CASENT0085551). Paratype, one pinned worker with same data as holotype except collected 20.III.—7.IV.2001 and collection code MA-01-07-09 (CASENT0083419). Non-type material. MADAGASCAR: Antsiranana, 4 km SW Ambositra (=Jof- freville), 12.51667°S, 49.18333°E, 1000 m, rainforest,1.XII.1990 (P.S. Ward); Ant- siranana, Nosy Be, Réserve Naturelle Intégrale de Lokobe, 6.3 km 112° ESE Hell- ville, 13.41933°S, 48.33117°E, 30 m, rainforest, 19.-24.I11.2001 (B.L. Fisher et al.); Antsiranana, Sakalava Beach (vegetated beach dunes), 12.26278°S, 49.3975°E, 10 m, across sandy trail in dwarf littoral forest, 20.-28.VII.2001 (R. Harin’Hala). Diagnosis. ‘The following character combination distinguishes 7: freya from the oth- er species of the 7: cognatum species complex: frontal carinae weakly developed, merging with surrounding sculpture halfway between posterior eye margin and posterior head margin; petiolar node high nodiform with well rounded margins, in profile around 1.6 to 1.7 times higher than long (LPel 58-62), in dorsal view 1.2 to 1.3 times wider than long (DPel 124-131); lack of any long, standing pilosity on dorsal mesosoma. Worker measurements (N=5). HL 0.61-0.74 (0.68); HW 0.56—0.68 (0.62); SL 0.40—-0.46 (0.44); EL 0.14—0.17 (0.16); PH 0.30—0.35 (0.33); PW 0.38-0.43 (0.40); WL 0.72-0.87 (0.81); PSL 0.12—0.15 (0.13); PTL 0.13—0.16 (0.15); PTH 0.21-0.27 (0.25); PTW 0.17-0.20 (0.19); PPL 0.16-0.21 (0.19); PPH 0.22-0.28 (0.25); PPW 0.23-0.28 (0.25); CI 90-92 (91); SI 68-71 (70); OI 25; DMI 44-54 (50); LMI 40- 42 (41); PSLI 18—20 (19); PeNI 42-50 (46); LPeIl 58-62 (59); DPeI 124-131 (127); PpNI 59-70 (63); LPpI 72-77 (75); DPpI 127-144 (134); PPI 133-140 (136). Worker description. Head much longer than wide (CI 90-92); in full-face view posterior head margin weakly concave. Anterior clypeal margin with distinct median impression. Frontal carinae weakly to moderately developed, merging with surround- ing sculpture halfway between posterior eye margin and posterior head margin. Anten- Taxonomy of Malagasy Tetramorium Foe nal scrobes weak to absent, shallow and without clear and distinct posterior and ventral margins. Antennal scapes very short, not reaching posterior head margin (SI 68-71). Eyes relatively large (OI 25). Mesosomal outline in profile weakly to moderately con- vex, moderately high and stout (LMI 40-42), weakly to moderately marginate from lateral to dorsal mesosoma; promesonotal suture absent; metanotal groove present, but weakly developed. Propodeal spines short, triangular to elongate-triangular, and acute (PSLI 18-20), propodeal lobes triangular and short, always much shorter than propodeal spines. Petiolar node in profile high nodiform with well-rounded margins, around 1.6 to 1.7 times higher than long (LPel 58-62), anterior and posterior faces approximately parallel and rounding smoothly onto dorsum, anterodorsal and pos- terodorsal margins situated at about same height, petiolar dorsum relatively noticeably convex; node in dorsal view 1.2 to 1.3 times wider than long (DPel 124-131), in dor- sal view pronotum between 2.0 to 2.4 times wider than petiolar node (PeNI 42-50). Postpetiole in profile globular to subglobular, between 1.3 to 1.4 times higher than long (LPpI 72-77); in dorsal view around 1.3 to 1.4 times wider than long (DPpl 127-144), pronotum between 1.4 to 1.8 times wider than postpetiole (PpNI 59-70). Postpetiole in profile appearing approximately as voluminous as petiolar node, postpe- tiole in dorsal view around 1.3 to 1.4 times wider than petiolar node (PPI 133-140). Mandibles completely unsculptured, smooth, and shiny; clypeus in parts irregularly longitudinally rugulose, median ruga never fully developed, usually reduced to a few traces, one or two irregular and broken lateral rugulae present on each side; cephalic dorsum between frontal carinae weakly irregularly longitudinally rugulose, rugulae weak, often interrupted or with cross-meshes and becoming much weaker posteriorly; scrobal area mostly irregularly longitudinally rugulose and merging with surround- ing irregularly longitudinally rugose to reticulate-rugose sculpture present on lateral head. Head with very conspicuous punctate ground sculpture. Mesosoma laterally and dorsally with traces of irregular rugulae or reticulate-rugose sculpture overlaying a dis- tinct punctate ground sculpture. Forecoxae in parts weakly longitudinally rugulose and partly unsculptured, smooth, and shining. Both waist segments and gaster fully unsculptured, smooth, and shining. Dorsum of head with several pairs of hairs. Meso- soma, waist segments, and first gastral tergite without any standing hairs at all; first gastral tergite with very short, moderately dense, and appressed pubescence. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae with appressed hairs. Head, mesosoma, waist segments, and gaster uniform brown, appendages yellowish to light brown. Etymology. The name of the new species was inspired and derived from the name of the Old Norse goddess Freyja (which means “Lady” in Old Norse) who was consid- ered the goddess of love, beauty, fertility, war, and death. As a name taken from Norse mythology, it is treated as an arbitrary combination of letters, thus invariant. Distribution and biology. Tetramorium freya is a rare species only known from a few localities at the northernmost tip of Madagascar and from the island of Nosy Be (Fig. 63). The species was collected in rainforests, dry forests, and coastal dunes at elevations from 10 to 1000 m. Also, 7: freya was mostly sampled from Malaise traps or 78 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Figure 33. 7 freya holotype worker (CASENT0085551). A Body in profile B Body in dorsal view C Head in full-face view. Taxonomy of Malagasy Tetramorium 79 from vegetation and only once from leaf litter; considering there are only five known specimens, it seems likely that 7! freya nests and forages in vegetation. Additional sam- pling in lower vegetation might yield more material of this otherwise very rare species. Discussion. ‘The identification of T: freya is easy and straightforward since it is the only species of the species complex without any long, standing pilosity on the meso- somal dorsum. Beyond mesosomal pilosity, 7: freya also has weakly developed frontal carinae and sculpture on the head and mesosoma, which separate it from T° gladius, T: myrmidon, T. proximum, and T. tenuinode. \n addition, T: freya possesses a stouter and higher mesosoma (LMI 40-42) and a lower and thicker petiolar node (LPel 58-62, around 1.6 to 1.7 times higher than long) compared to T° aspis, T: camelliae, T. cog- natum, T. karthala, and T. rumo (LMI 35—40; LPel 33-55, around 1.8 to 3.0 times higher than long). Based on the limited number of specimens available for this study, there seems to be no significant intraspecific variation in T° freya. Tetramorium gladius Hita Garcia & Fisher, sp. n. http://zoobank.org/04D35 BE9-26E7-450E-B5FC-3EE67E7B7934 http://species-id.net/wiki/Tetramorium_gladius Figs 21C, 23A, 34, 63 Type material. Holotype, pinned worker, MADAGASCAR, Antananarivo, 3 km 41° NE Andranomay, 11.5 km 147° SSE Anjozorobe, 18.47333°S, 47.96°E, 1300 m, montane rainforest, sifted litter (leaf mold, rotten wood), collection code BLF02464, 5.-13.XII (BL. Fisher et al.) (CAS: CASENT0406982). Paratypes, one pinned worker with same data as holotype (CAS: CASENT0406981); four pinned worker with same data as holotype but collected ex rotten stick on ground and collection codes BLF02407 and BLF0O2408 (CAS: CASENT0406964; CASENT040697 1); fifteen pinned workers with same data as holotype but collected ex rotten log and collection codes BLF02429, BLF02498, and BLF02500 (CAS: CASENT0406966; CASENT0406967; CASENT0406968; CASENT0406969; CASENT0406974; CASENT0406975; MCZ: CASENT0406976); one pinned worker with same data as holotype but collected ex rotten tree stump and collection code BLF02486 (BMNH: CASENT0406965). Non-type material MADAGASCAR: Antananarivo, Réserve Spéciale d’Ambohitantely, 18.22444°S, 47.2774°E, 1490 m, montane forest, 9.II].2012 (B.L. Fisher et al.); Antananarivo, 3 km 41° NE Andranomay, 11.5 km 147° SSE Anjozo- robe, 18.47333°S, 47.96°E, 1300 m, montane rainforest, 5.-13.XII (B.L. Fisher et al.); Antsiranana, Rés. Anjanaharibe-Sud, 9.2 km WSW Befingotra, 14.75°S, 49.46667°E, 1200 m, montane rainforest, 9.XI.1994 (B.L. Fisher); Antsiranana, Forét de Binara, 9.4 km 235° SW Daraina, 13.26333°S, 49.6°E, 1100 m, montane rainforest, 5.—6. XII.2003 (B.L. Fisher); Toamasina, 6.9 km NE Ambanizana, Ambohitsitondroina, 15.58506°S, 50.00952°E, 825 m, rainforest, 2.XII.1993 (BL. Fisher); Toamasina, 80 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) 5.3 km SSE Ambanizana, Andranobe, 15.67133°S, 49.97395°E, 425 m, rainforest, 21.X1.1993 (B.L. Fisher); Toamasina, Réserve Spéciale Ambatovaky, Sandrangato river, 16.77274°S, 49.26551°E, 450 m, rainforest, 20.—22.11.2010 (B.L. Fisher et al.). Diagnosis. The relatively small eyes (OI 19-20) of T. gladius separate it from the other members of the 7. cognatum species complex. Worker measurements (N=12). HL 0.71-0.87 (0.81); HW 0.65—0.82 (0.75); SL 0.48-0.58 (0.54); EL 0.13-0.16 (0.15); PH 0.32-0.41 (0.37); PW 0.47-0.61 (0.55); WL 0.87—1.08 (1.00); PSL 0.17—0.22 (0.19); PTL 0.14-0.18 (0.15); PTH 0.24-0.33 (0.29); PTW 0.17—0.22 (0.19); PPL 0.20-0.27 (0.23); PPH 0.25-0.34 (0.30); PPW 0.26—0.34 (0.29); CI 92-95 (93); SI 71-74 (72); OI 19-20 (20); DMI 54-56 (55); LMI 36-38 (37); PSLI 21-28 (23); PeNI 31-36 (35); LPel 48-58 (53); DPel 113-133 (124); PpNI 49-56 (53); LPpI 70-82 (78); DPpI 120-133 (125); PPI 148-159 (153). Worker description. Head longer than wide (CI 92—95); in full-face view poste- rior head margin weakly to moderately concave. Anterior clypeal margin with distinct median impression. Frontal carinae moderately to well developed, diverging posteri- orly, either merging with surrounding sculpture halfway between posterior eye mar- gin and posterior head margin or only becoming weaker after posterior eye level but still approaching posterior head margin. Antennal scrobes very weak, shallow, and without clear and distinct posterior and ventral margins. Antennal scapes short, not reaching posterior head margin (SI 71-74). Eyes relatively small (OI 19-20). Meso- somal outline in profile flat to weakly convex, moderately low and long (LMI 36-38), moderately marginate from lateral to dorsal mesosoma; promesonotal suture absent; metanotal groove mostly reduced and absent, if present weakly developed. Propodeal spines moderately long to long, elongate-triangular to spinose, and usually acute (PSLI 21-28), propodeal lobes short and triangular, always much shorter than propodeal spines. Petiolar node in profile nodiform to high rounded nodiform, around 1.7 to 2.1 times higher than long (LPel 48-58), anterior and posterior faces approximately paral- lel, anterodorsal and posterodorsal margins usually situated at about same height and weakly rounded, petiolar dorsum weakly to moderately convex; node in dorsal view between 1.1 to 1.3 times wider than long (DPel 113-133), in dorsal view pronotum between 2.8 to 3.2 times wider than petiolar node (PeNI 31-36). Postpetiole in pro- file globular, around 1.2 to 1.4 times higher than long (LPpI 70-82); in dorsal view around 1.2 to 1.3 times wider than long (DPpI 120-133), pronotum around 1.8 to 2.0 times wider than postpetiole (PpNI 49-56). Postpetiole in profile usually appear- ing more voluminous than petiolar node, postpetiole in dorsal view around 1.5 to 1.6 times wider than petiolar node (PPI 148-159). Mandibles completely unsculptured, smooth, and shiny; clypeus longitudinally rugose/rugulose with three to seven median ruga always fully developed and distinct, and one to three mostly unbroken lateral rugae/rugulae present on each side; cephalic dorsum between frontal carinae longitudi- nally rugose with seven to nine rugae, rugae running mostly unbroken from posterior clypeal margin to posterior head margin, sometimes interrupted or with cross-meshes; Taxonomy of Malagasy Tetramorium 81 Figure 34. 7° gladius holotype worker (CASENT0406982). A Body in profile B Body in dorsal view C Head in full-face view. 82 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) scrobal area partly unsculptured, but mostly merging with surrounding reticulate-ru- gose to longitudinally rugose sculpture present on lateral head. Dorsum and sides of mesosoma mostly irregularly longitudinally rugose. Forecoxae unsculptured, smooth, and shining. Ground sculpture on head and mesosoma usually weak to absent. Both waist segments and gaster fully unsculptured, smooth, and shining. Dorsum of head with several pairs of long, fine, standing hairs; dorsum of mesosoma with six to eight pairs on pronotum and mesonotum, anterior propodeum with one pair; waist seg- ments and first gastral tergite without any standing hairs; first gastral tergite with very short, scarce, appressed pubescence. Anterior edges of antennal scapes and dorsal (out- er) surfaces of hind tibiae with appressed, rarely decumbent hairs. Head, mesosoma, waist segments, and gaster uniform reddish, orange-brown, appendages always lighter, yellowish to light brown. Etymology. The name of the new species is the ancient Latin general word for sword, although it mostly refers to the short swords used by foot soldiers of the Ro- man legion. The name was chosen based on the shape of the propodeal spines in most specimens of T: gladius, which resemble this type of sword. The species epithet is a nominative noun, and thus invariant. Distribution and biology. Tetramorium gladius has a relatively wide but patchy distribution in central, central-eastern, and northern Madagascar (Fig. 63). The south- ernmost localities are Ambohitantely and Andranomay and the northernmost is Binara with Ambatovaky, Ambanizana, and Anjanaharibe-Sud in-between. All localities are rainforests and montane rainforests at elevations ranging from 425 to 1490 m. Te- tramorium gladius seems to be only moderately common and living in the leaf litter. Discussion. The relatively small eyes of 7. gladius (OI 19-20) differentiate it im- mediately from the remainder of the 7. cognatum species complex, in which all species have much larger eyes (OI 24-31). Apart from this, 7: gladius appears to be morpho- logically close to T. myrmidon, T: proximum, and T! tenuinode since they all share a larger body size and very well developed frontal carinae and sculpture. Little morphological variation is observed in T° gladius, with the exception, that the propodeal spines are longer in specimens from Ambatovaky (PSLI 26-28) com- pared to the rest of the material (PSLI 21-24). Tetramorium karthala Hita Garcia & Fisher, sp. n. http://zoobank.org/9B534784-COF8-4B8C-AC93-77B3CBDB6DCC http://species-id.net/wiki/Tetramorium_karthala Figs 20A, 24C, 27D, 28D, 29B, 35, 63 Type material. Holotype, pinned worker, COMOROS, Grande Comore, Karthala, 11.82699°S, 43.4295°E, 1000 m, montane rainforest, sifted litter (leaf mold, rot- ten wood), collection code BLF19734, 14.-15.II.2008 (B.L. Fisher et al.) (CAS: CASENT0137302). Paratypes, three pinned workers with same data as holotype (CAS: CASENT0137426; CASENT0137440; CASENT0137510); two workers with Taxonomy of Malagasy Tetramorium 83 same data as holotype except collected from rotten log and collection code BLF19750 (MCZ: CASENT0135232); six pinned workers from Grande Comore, Karthala, 11.81336°S, 43.41945°E, 1125 m, montane rainforest, sifted litter (leaf mold, rot- ten wood), collection code BLF19700, 13.—14.IJI.2008 (B.L. Fisher et al.) (BMNH: CASENT0136774; CAS: CASENT0136766; CASENT0137214; CASENT0137221; CASENT0137244; CASENT0137263;). Non-type material COMOROS: Grande Comore, Karthala, 11.81336°S, 43.41945°E, 1125 m, montane rainforest, 13.III.2008 (B.L. Fisher et al.). Diagnosis. Tetramorium karthala is diagnosable within the 7. cognatum complex on the basis of the following character combination: very large eyes (OI 29-30); short antennal scapes (SI 70-74); frontal carinae weakly to moderately developed; propo- deal spines short to moderate, elongate-triangular to spinose, and usually acute (PSLI 20-22), propodeal lobes always much shorter than spines and lobes never strongly inclined towards each other; petiolar node rounded high nodiform, in profile around 1.8 to 2.0 times higher than long (LPel 50-54), in dorsal view around 1.5 to 1.6 times wider than long (DPel 148-158); mesosoma with two to four pairs of long, standing hairs on pronotum and mesonotum. Worker measurements (N=10). L 0.59-0.63 (0.61); HW 0.51-0.55 (0.53); SL 0.36—0.39 (0.38); EL 0.15—-0.16 (0.15); PH 0.27—-0.31 (0.29); PW 0.38-0.42 (0.40); WL 0.72-0.79 (0.75); PSL 0.12—0.14 (0.13); PTL 0.12—0.14 (0.13); PTH 0.23-0.26 (0.24); PTW 0.18-0.21 (0.19); PPL 0.15-0.17 (0.15); PPH 0.22-0.25 (0.23); PPW 0.22—0.25 (0.24); Cl 86-87 (87); SI 70-74 (72); OI 29-30 (29); DMI 51-55 (54); LMI 37-39 (39); PSLI 20-22 (21); PeNI 45—49 (47); LPel 50-54 (51); DPel 148-158 (151); PpNI 55-61 (58); LPpI 65-68 (66); DPpI 138-158 (151); PPI 112-131 (124). Worker description. Head much longer than wide (CI 86-87); posterior head margin weakly concave. Anterior clypeal margin with distinct median impression. Frontal carinae weakly to moderately developed, only weakly raised, and ending between posterior head margin and posterior head margin. Antennal scrobes very weakly developed to almost absent, shallow and without clear and distinct posterior and ventral margins. Antennal scapes short, not reaching posterior head margin (SI 70-74). Eyes very large (OI 29-30). Mesosomal outline in profile flat to weakly convex, moderately low and long (LMI 37-39), moderately marginate from lateral to dorsal mesosoma; promesonotal suture absent; metanotal groove present and dis- tinctly developed. Propodeal spines short to moderately long, elongate-triangular to spinose, and usually acute (PSLI 20—22), propodeal lobes short and triangular, always much shorter than propodeal spines, spines and lobes never strongly inclined to- wards each other. Petiolar node in profile rounded high nodiform, around 1.8 to 2.0 times higher than long (LPel 50-54), usually anterior and posterior faces more or less parallel, anterodorsal and posterodorsal margins both well-rounded and usually situ- ated at about same height, petiolar dorsum moderately convex; node in dorsal view around 1.5 to 1.6 times wider than long (DPel 148-158), in dorsal view pronotum between 2.0 to 2.2 times wider than petiolar node (PeNI 45-49). Postpetiole in pro- file subglobular, between 1.4 to 1.6 times higher than long (LPpI 65-68); in dorsal 84 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) view around 1.4 to 1.6 times wider than long (DPpI 138-158), pronotum around 1.6 to 1.8 times wider than postpetiole (PpNI 55-61). Postpetiole in profile usually appearing of more or less same volume as petiolar node, postpetiole in dorsal view around 1.1 to 1.3 times wider than petiolar node (PPI 112-131). Mandibles com- pletely unsculptured, smooth, and shiny; clypeus usually irregularly longitudinally rugulose with three to five, often broken, rugulae, sometimes rugulae reduced to one or two, median ruga/rugula often developed, but usually broken, rarely completely absent; cephalic dorsum between frontal carinae longitudinally rugose/rugulose with seven to nine rugae/rugulae, rugae usually running from posterior clypeal margin to posterior head margin, often interrupted, rarely with cross-meshes; scrobal area mostly unsculptured with ground sculpture only, lateral head reticulate-rugose to longitudinally rugose. Dorsum and sides of mesosoma mostly irregularly longitudi- nally rugose/rugulose. Forecoxae dorsally with few traces of rugulae, otherwise mostly unsculptured, smooth, and shining. Ground sculpture on head and mesosoma very distinct, usually reticulate-punctate with few areas very finely reticulate-rugulose. Both waist segments and gaster fully unsculptured, smooth, and shining. Dorsum of head with numerous pairs of standing, long, fine hairs; mesosoma usually with two to four pairs: one long pair on anterior pronotum and one long pair on anterior me- sonotum, sometimes two shorter pairs present, one on posterior pronotum and one on posterior mesonotum; propodeum, waist segments and first gastral tergite without any standing hairs; first gastral tergite with short, moderately dense, appressed pu- bescence. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae with appressed to decumbent hairs. Head, mesosoma, waist segments and gaster uni- form light brown, appendages often lighter, more yellowish brown. Etymology. The new species is named after the type locality, Karthala Forest on Mount Karthala volcano on the island of Grand Comore. The species epithet is a noun in apposition, and thus invariable. Distribution and biology. This new species is only known from the type locality, which is a montane rainforest on the Comorian island Grande Comore (Fig. 63). At present, it is the only described Tetramorium species endemic to any Comorian island. All other Tetramorium species (except one undescribed species from the 7. ranarum spe- cies group from Anjouan) found on the Comoros Islands are either introduced tramp species or species shared between Africa, the Comoros and Madagascar, or between the Comoros and Madagascar. Intriguingly, the type locality appears to have a quite unique fauna. Just recently Fischer and Fisher (2013) described a new Pheidole species (Phei- dole vulcan Fischer & Fisher) from this locality, which is also endemic to the island of Grande Comore. Tetramorium karthala is at elevations between 1000 to 1125 m where it was mainly collected from leaf litter, under moss, or within rotten logs. Discussion. 7etramorium karthala is easily identifiable in the Malagasy region since it is the only species of the 7: cognatum species complex and the whole T° schaufussii species group found on the Comores. Even without considering geography the new spe- cies can be well distinguished from the remainder of its species complex. In general, it is morphologically close to T: aspis, T. camelliae, T: cognatum, and T. rumo, whereas it Taxonomy of Malagasy Tetramorium 85 Figure 35. 7 karthala paratype worker (CASENT0136774). A Body in profile B Body in dorsal view C Head in full-face view. 86 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) differs more strongly from the rest of the complex. Since T: karthala has several pairs of long, standing hairs on the dorsum of the mesosoma and much larger eyes (OI 29-30), it is unlikely to be mistaken for T° freya, which has no standing pilosity on the mesoso- mal dorsum, or for 7: gladius, which has the smallest eyes in the complex (OI 19-20). Also, T: karthala cannot be confused with 7. myrmidon, T. proximum, or T. tenuinode as the latter three have very well developed, noticeably raised, and long frontal carinae compared to the much weaker frontal carinae seen in 7: karthala. The separation from the other five species, which appear closer to T: karthala, is also straightforward. Tetramorium camelliae has a strongly squamiform petiolar node which is around 2.8 to 3.0 times higher than long (LPel 33-36) and around 2.3 to 2.4 times wider than long (DPel 228-238), while T° karthala has a high nodiform petiolar node which is around 1.8 to 2.0 times higher than long (LPel 50-54) and around 1.5 to 1.6 times wider than long (DPel 148-158). The smallest species of the complex, T. rumo (HW 0.43-0.49; WL 0.56-0.67), differs from 7: karthala (HW 0.51-0.55; WL 0.72-0.79) not only in body size but also in the shape of the petiolar node. The node of 7. rumo is thinly cuneiform and moderately squamiform, in profile 2.3 to 2.7 times higher than long (LPel 37—43) while, as noted above, the node of 7. karthala is high nodiform and thicker, only 1.8 to 2.0 times higher than long (LPel 50-54). Fur- thermore, in 7: aspis the propodeal spines and lobes are strongly inclined towards each other, whereas the spines and lobes of T: karthala are not. The latter species also has two to four pairs of long, standing hairs on the dorsal pronotum and mesonotum while T. aspis has at least six on the pronotum and mesonotum and usually one or two on the anterior propodeum. ‘The last species in the complex, T° cognatum, appears mor- phologically very close to T: karthala, but has noticeably shorter antennal scapes (SI 61-67) and propodeal spines (PSLI 12-16) than T: karthala (SI 70-74; PSLI 20-22). To our knowledge, there is no significant intraspecific variation in T° karthala. Tetramorium myrmidon Hita Garcia & Fisher, sp. n. http://zoobank.org/11BD4CF2-149D-4A6A-9B9C-C59C0030B466 http://species-id.net/wiki/Tetramorium_myrmidon Figs 23C, 25B, 26C, 36, 64 Type material. Holotype, pinned worker, MADAGASCAR, Toliara, Réserve Spé- ciale d’Ambohijanahary, Forét d’Ankazotsihitafototra, 35.2 km 312° NW Ambara- varanala, 18.26667°S, 45.40667°E, 1050 m, montane rainforest, sifted litter (leaf mold, rotten wood), collection code BLF07020, 13.-17.1.2003 (B.L. Fisher et al.) (CAS: CASENT0028635). Paratypes, two workers with same data as holotype (CAS: CASENT0028642; CASENT0028643); and three workers MADAGASCAR, Toli- ara, Réserve Spéciale d’Ambohijanahary, Forét d’Ankazotsihitafototra, 34.6 km 314° NW Ambaravaranala, 18.26°S, 45.41833°E, 1100 m, montane rainforest, sifted litter (leaf mold, rotten wood), collection code BLF07086, 16.1.2003 (B.L. Fisher et al.) (CAS: CASENT0029810; CASENT0029821; CASENT0030099). Taxonomy of Malagasy Tetramorium 87 Diagnosis. Tetramorium myrmidon can be easily distinguished from the other members of the group by the following combination of characters: eyes relatively large (OI 24-25); antennal scapes short (SI 74-76); frontal carinae well developed, notice- ably raised, ending shortly before posterior head margin; petiolar node high rounded nodiform, in profile around 1.7 times higher than long (LPel 58-60), in dorsal view around 1.2 to 1.3 times wider than long (DPel 121-129); in dorsal view postpetiole around 1.3 to 1.5 times broader than petiolar node (PPI 133-141); dorsum of meso- soma with two pairs of long, standing hairs on pronotum and mesonotum. Worker measurements (N=6). HL 0.70-0.76 (0.73); HW 0.62—0.69 (0.65); SL 0.47—-0.52 (0.49); EL 0.16—0.18 (0.16); PH 0.33—0.37 (0.35); PW 0.45—0.50 (0.47); WL 0.85—0.94 (0.89); PSL 0.13—-0.16 (0.14); PTL 0.14—0.16 (0.15); PTH 0.24—0.27 (0.25); PTW 0.18-0.20 (0.19); PPL 0.18—0.21 (0.19); PPH 0.24—0.28 (0.26); PPW 0.24—0.28 (0.26); CI 88-91 (90); SI 74-76 (75); OI 24-25 (25); DMI 52-54 (53); LMI 38-40 (39); PSLI 18-20 (19); PeNI 39-40 (39); LPeI 58-60 (59); DPel 121-129 (125); PpNI 52-55 (54); LPpI 73-77 (76); DPpI 129-136 (133); PPI 133-141 (138). Worker description. Head much longer than wide (CI 88-91); in full-face view posterior head margin weakly concave. Anterior clypeal margin with distinct median impression. Frontal carinae well developed, noticeably raised, diverging posteriorly, merging with surrounding sculpture shortly before posterior head margin. Antennal scrobes very weakly developed, almost absent, shallow and without clear and distinct posterior and ventral margins. Antennal scapes short, not reaching posterior head mar- gin (SI 74—76). Eyes relatively large (OI 24-25). Mesosomal outline in profile flat to weakly convex, moderately low and long (LMI 38-40), moderately marginate from lateral to dorsal mesosoma; promesonotal suture absent; metanotal groove present, but weakly developed. Propodeal spines short to moderate, elongate-triangular, and acute (PSLI 18-20), propodeal lobes triangular and short, always much shorter than propodeal spines. Petiolar node in profile high nodiform with well rounded margins, around 1.7 times higher than long (LPel 58-60), anterior and posterior faces approxi- mately parallel, anterodorsal and posterodorsal margins situated at about same height, anterodorsal margin slightly more angulate than posterodorsal, more rounded margin, petiolar dorsum relatively flat to weakly convex; node in dorsal view 1.2 to 1.3 times wider than long (DPel 121-129), in dorsal view pronotum between 2.5 to 2.6 times wider than petiolar node (PeNI 39-40). Postpetiole in profile globular, between 1.3 to 1.4 times higher than long (LPp!I 73-77); in dorsal view around 1.3 to 1.4 times wider than long (DPpI 129-136), pronotum around 1.8 to 1.9 times wider than postpetiole (PpNI 52-55). Postpetiole in profile appearing more voluminous than petiolar node, postpetiole in dorsal view around 1.3 to 1.4 times wider than petiolar node (PPI 133-— 141). Mandibles completely unsculptured, smooth, and shiny; clypeus irregularly lon- gitudinally rugulose with three to seven rugulae, median ruga usually fully developed, one to three mostly unbroken lateral rugulae present on each side; cephalic dorsum between frontal carinae longitudinally rugulose with seven to nine rugulae, rugulae usually running from posterior clypeal margin to posterior head margin, often inter- rupted or with cross-meshes, especially posteriorly; scrobal area mostly unsculptured 88 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) with ground sculpture only, lateral head reticulate-rugose to longitudinally rugulose. Ground sculpture on head well developed and distinct, mostly reticulate-punctate. Dorsum and sides of mesosoma irregularly longitudinally rugulose to reticulate-ru- gulose. Forecoxae dorsally weakly rugulose, but mostly unsculptured, smooth, and shining. Ground sculpture on mesosoma weakly to moderately punctate. Both waist segments and gaster fully unsculptured, smooth, and shining. Dorsum of head with several pairs of standing, long, fine hairs; dorsum of mesosoma with two pairs only, one on anterior pronotum and one on anterior mesonotum; propodeum, waist seg- ments, and first gastral tergite without any standing hairs; first gastral tergite with very short, scarce, appressed pubescence. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae with appressed hairs. Head, mesosoma, waist segments, and gaster uniformly brown to dark brown, appendages yellowish to light brown. Etymology. The name of the new species is inspired by the ancient Greek “myr- midons”, which were skilled warriors and known as the legendary “ant-people” who inhabited the Greek island of Aegina. The species epithet is a noun in apposition, and thus invariant. Distribution and biology. The new species is so far only known from Ambohija- nahary, which is an isolated montane rainforest located in the midwest of Madagascar (Fig. 64), where it was collected only twice. With only six known specimens it repre- sents an extremely rare species. Tetramorium myrmidon was collected from leaf litter at elevations of 1050 to 1100 m. Discussion. Based on its larger body size and very well developed frontal cari- nae, T. myrmidon differs strongly from the smaller species T. aspis, T. camelliae, T: cognatum, T. karthala and T. rumo, all with more weakly developed frontal carinae, while being presumably morphologically closer to T. gladius, T. proximum and T. tenuinode. Of these, T. gladius possesses very small eyes (OI 19-20), while the eyes of T. myrmidon are much larger (OI 24-25). Tetramorium tenuinode has shorter anten- nal scapes (SI 66-70) and a thinner petiolar node, in profile 1.8 to 2.2 times higher than long (LPel 45-54), in contrast to T. myrmidon with its longer scapes (SI 74-76) and lower and thicker petiolar node, which is in profile around 1.7 times higher than long (LPel 58-60). The widespread 7: proximum, however, appears to be the species morphologically closest to 7: myrmidon and most of their morphometric ranges and characters overlap. Nonetheless, we consider both sufhciently demarcated from each other since they are found to co-occur in sympatry in Ambohijanahary. The specimens of T. proximum from this locality differ from T: myrmidon by having five to six pairs of long, standing hairs on the mesosoma and a generally thinner and higher petiolar node. Finally, 7. myrmidon is unlikely to be confused with the last species of the com- plex, T° freya, since the latter lacks long, standing pilosity on the mesosomal dorsum (present in 7. myrmidon). The small number of specimens from just two collection events in the same locality does not permit proper assessment of intraspecific variation. Taxonomy of Malagasy Tetramorium 89 gi ART och ¥ ’ Es Figure 36. 7° myrmidon holotype worker (CASENT0028635). A Body in profile B Body in dorsal view C Head in full-face view. 90 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Tetramorium proximum Bolton, 1979 http://species-id.net/wiki/Tetramorium_proximum Figs 20B, 22D, 24A, 25A, 37, 64 Letramorium proximum Bolton, 1979:137. Type material. Holotype, pinned worker, MADAGASCAR, Toamasina, Périnet & vicinity, 18.82667°S, 48.44778°E, rainforest, rotten wood, 18.IJI.1969 (W. L. Brown) (MCZ: MCZ_Holotype_32264) [examined]. Paratypes, ten pinned work- ers and one pinned queen with same data as holotype (BMNH: CASENT0102341; CASENT0102342; CASENT0235211; MCZ: MCZ_Paratype_32264). [Note: the GPS data of the type locality was not provided either by the locality label or the original description. The data presented above is based on our own geo- referencing of the Périnet=Andasibe area and should be considered an approximation and not the exact position of the type locality.] Non-type material MADAGASCAR: Antananarivo, Tsimbazaza, 18.928°S, 47.527°E, 1300 m, park/garden, 16.XII.2006 (B.L. Fisher et al.); Antsiranana, Parc National Montagne d’Ambre, 3.6 km 235° SW Joffreville, 12.53444°S, 49.1795°E, 925 m, montane rainforest, 20.—26.1.2001 (B.L. Fisher et al.); Antsiranana, Parc Na- tional Montagne d’Ambre, 12.2 km 211° SSW Joffreville, 12.59639°S, 49.1595°E, 1300 m, montane rainforest, 7.11.2001 (G.D. Alpert); Antsiranana, Parc National Montagne d’Ambre [1st campsite], 12.51444°S, 49.18139°E, 960 m, rainforest, 5.— 21.1V.2001 (R. Harin’Hala); Antsiranana, Parc National Montagne d’Ambre, 12.51389°S, 49.17784°E, 984 m, montane rainforest, 23.1].2001 (B.L. Fisher et al.); Antsiranana, Parc National Montagne d’Ambre, 12.51778°S, 49.17957°E, 1000 m, montane rainforest, 3.—6.I]I.2001 (B.L. Fisher et al.); Antsiranana, Parc National Montagne d’Ambre, 12.52306°S, 49.17901°E, 1100 m, montane rainforest, 11. III.2001 (B.L. Fisher et al.); Antsiranana, Parc National Montagne d’Ambre, Lac mau- dit, 12.58502°S, 49.15147°E, 1250 m, montane rainforest, 13.X1.2007 (B.L. Fisher et al.); Antsiranana, Parc National Montagne d’Ambre, Roussettes, 12.52574°S, 49.17238°E, 1025 m, montane rainforest, 15.XI.2007 (B.L. Fisher et al.); Antsiranana, Parc National Montagne d’Ambre, Petit lac, 12.53664°S, 49.17412°E, 1130 m, mon- tane rainforest, 17.X1.2007 (B.L. Fisher et al.); Antsiranana, Parc National Montagne d’Ambre, Antomboka, 12.51269°S, 49.17807°E, 970 m, montane rainforest, 17. X1.2007 (B.L. Fisher et al.); Antsiranana, Parc National Montagne d’Ambre, Ma- hasarika, 12.53176°S, 49.17662°E, 1135 m, montane rainforest, 17.XI.2007 (BL. Fisher et al.); Antsiranana, Rés. Anjanaharibe-Sud, 6.5 km SSW Befingotra, 14.75°S, 49.5°E, 875 m, rainforest, 19.X.1994 (B.L. Fisher); Antsiranana, Forét de Binara, 9.1 km 233° SW Daraina, 13.26333°S, 49.60333°E, 800 m, rainforest, 3.XII.2003 (B.L. Fisher); Antsiranana, Forét de Binara, 9.4 km 235° SW Daraina, 13.26333°S, 49.6°E, 1100 m, montane rainforest, 5.XII.2003 (B.L. Fisher); Antsiranana, Betaolana Forest, along Bekona River, 14.52996°S, 49.44039°E, 880 m, rainforest, 4.-5.III.2009 (B.L. Fisher et al.); Antsiranana, R.S. Manongarivo, 10.8 km 229° SW Antanambao, Taxonomy of Malagasy Tetramorium 91 13.96167°S, 48.43333°E, 400 m, rainforest, 8.-13.X.1998 (B.L. Fisher); Antsiranana, R.S. Manongarivo, 12.8 km 228° SW Antanambao, 13.97667°S, 48.42333°E, 780 m, rainforest, 11.X.1998 (B.L. Fisher); Antsiranana, R.S. Manongarivo, 14.5 km 220° SW Antanambao, 13.99833°S, 48.42833°E, 1175 m, montane rainforest, 20.X.1998 (B.L. Fisher); Antsiranana, Parc National de Marojejy, Manantenina River, 28.0 km 38° NE Andapa, 8.2 km 333° NNW Manantenina, 14.43667°S, 49.775°E, 450 m, rainforest, 12.-15.X1.2003 (B.L. Fisher et al.); Antsiranana, Parc National de Maroje- jy, Manantenina River, 27.6 km 35° NE Andapa, 9.6 km 327° NNW Manantenina, 14.435°S, 49.76°E, 775 m, rainforest, 15.-18.X1.2003 (B.L. Fisher et al.); Fianarant- soa, 45 km S Ambalavao, 22.21667°S, 47.01667°E, 785 m, rainforest, 25.—30.[X.1993 (B.L. Fisher); Fianarantsoa, 2 km W Andrambovato, along river Tatamaly, 21.51167°S, 47.41°E, 1075 m, montane rainforest, 3.—5.VI.2005 (B.L. Fisher et al.); Fianarantsoa, Rés. Andringitra, 43 km S Ambalavao, 22.23333°S, 47°E, 825 m, rainforest, 5.X.1993 (B.L. Fisher); Fianarantsoa, Rés. Andringitra, 40 km S Ambalavao, 22.21667°S, 46.96667°E, 1275 m, montane rainforest, 15.-19.X.1993 (B.L. Fisher); Fianarantsoa, Rés. Andringitra, 38 km S Ambalavao, 22.2°S, 46.96667°E, 1680 m, montane rainfor- est, 23.X.1993 (B.L. Fisher); Fianarantsoa, Parc National Befotaka-Midongy, Papango 28.5 km S Midongy-Sud, Mount Papango, 23.84083°S, 46.9575°E, 1250 m, mon- tane rainforest, 17.-18.X1.2006 (B.L. Fisher et al.); Fianarantsoa, 900 m E of Isalo National Park Interpretive Center, stream area, 22.62667°S, 45.35817°E, 750 m, open area near stream, 3.-10.11.2002 (Rk. Harin’Hala); Fianarantsoa, R.S. Ivohibe, 7.5 km ENE Ivohibe, 22.47°S, 46.96°E, 900 m, rainforest, 7.-12.X.1997 (B.L. Fish- er); Fianarantsoa, R.S. Ivohibe, 8.0 km E Ivohibe, 22.48333°S, 46.96833°E, 1200 m, montane rainforest, 15.-21.X.1997 (B.L. Fisher); Fianarantsoa, R.S. Ivohibe, 6.5 km ESE Ivohibe, 22.49667°S, 46.955°E, 1575 m, montane rainforest, 24.—30.X.1997 (B.L. Fisher); Fianarantsoa, 8.0 km NE Ivohibe, 22.42167°S, 46.89833°E, 1200 m, montane rainforest, 3.-9.X1.1997 (B.L. Fisher); Fianarantsoa, 9.0 km NE Ivohibe, 22.42667°S, 46.93833°E, 900 m, rainforest, 12.-17.X1.1997 (B.L. Fisher); Fianarant- soa, Parc Nationale Ranomafana, Talatakely, 21.24833°S, 47.42667°E, in guava for- est, 9.-26.1V.1998 (CE. Griswold et al.); Fianarantsoa, Ranomafana National Park, Vohiparara Kidonavo 2, 21.22603°S, 47.36963°E, 1100 m, 13.II.2003 (V.C. Clark); Fianarantsoa, Parc National de Ranomafana, Vatoharanana, 21.28955°S, 47.4304, 1100 m, 30.11.2003 (V.C. Clark); Fianarantsoa, Parc National de Ranomafana, Vato- haranana River, 4.1 km 231° SW Ranomafana, 21.29°S, 47.43333°E, 1100 m, mon- tane rainforest, 27.—31.II1.2003 (B.L. Fisher et al.); Fianarantsoa, Forét Classée Vato- vavy, 7.6 km 122° Kianjavato, 21.4°S, 47.94°E, 175 m, rainforest, 6.-8.V1.2005 (B.L. Fisher et al.); Fianarantsoa, Forét de Vevembe, 66.6 km 293° Farafangana, 22.791°S, 47.18183°E, 600 m, rainforest, transition to montane forest, 23.—24.IV.2006 (B.L. Fisher et al.); Mahajanga, Réserve Spéciale Marotandrano, Marotandrano 48.3 km S Mandritsara, 16.28322°S, 48.81443°E, 865 m, transition humid forest, 7.XII.2007 (B.L. Fisher et al.); Toamasina, 6.3 km S Ambanizana, Andranobe, 15.6813°S, 49.958°E, 25 m, rainforest, 14.X1.1993 (B.L. Fisher); Toamasina, Forét Ambatovy, 14.3 km 57° Moramanga, 18.85083°S, 48.32°E, 1075 m, montane rainforest, 21. 92 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) III.2004 (Malagasy ant team); Toamasina, Ambatovy, 12.4 km NE Moramanga, 18.84773°S, 48.29568°E, 1000-1010 m, montane rainforest, 3.—8.III.2007 (B.L. Fisher et al.); Toamasina, Amparihibe, 15°2’ S, 49°34 E, I.—III.2003 (K-A. Jackson & D. Carpenter); Toamasina, Station Forestiére Analamazaotra, Analamazaotra 1.3 km S Andasibe, 18.38466°S, 48.41271°E, 980 m, montane rainforest, 11.—13.XII.2007 (B.L. Fisher et al.); Toamasina, Montagne d’Anjanaharibe, 18.0 km 21° NNE Ambi- nanitelo, 15.18833°S, 49.615°E, 470 m, rainforest, 8.—12.III.2003 (B.L. Fisher et al.); Toamasina, Ankerana, 18.40062°S, 48.81311°E, 865 m, rainforest, 17.-18.1.2012 (B.L. Fisher et al.); Toamasina, Ankerana, 18.40829°S, 48.82107°E, 750 m, rainforest, 21.-22.1.2012 (B.L. Fisher et al.); Toamasina, Ankerana, 18.40636°S, 48.80254°E, 1108 m, montane rainforest, 29.1.2012 (B.L. Fisher et al.); Toamasina, Reserve Betam- pona, Camp Rendrirendry, 34.1 km 332° Toamasina, 17.924°S, 49.19967°E, 390 m, rainforest, 28.X1.2005 (B.L. Fisher et al.); ‘Toamasina, Réserve Nationale Intégrale Bet- ampona, Betampona 35.1 km NW Toamasina, 17.91801°S, 49.20074°E, 500 m, rainforest, 16.XII.2007 (B.L. Fisher et al.); Toamasina, Réserve Naturelle Betampona, 34.08 km 332° Toamasina, 17.91977°S, 49.20039°F, 525 m,_ rainforest, 17.11.—-5.X.2008 (B.L. Fisher); Toamasina, Réserve Naturelle Betampona, 34.1 km 332° Toamasina, 17.916136°S, 49.20185°E, 550 m, rainforest, 5.X.-14.XII.2008 (B.L. Fisher); Toamasina, F.C. Didy, 18.19833°S, 48.57833°E, 960 m, rainforest, 16.—23.XI11.1998 (HJ. Ratsirarson); Toamasina, Parc National Mananara-Nord, 7.1 km 261° Antanambe, 16.455°S, 49.7875°E, 225 m, rainforest, 14.X1.2005 (B.L. Fish- er et al.); Toamasina, P.N. Mantadia, 18.79167°S, 48.42667°E, 895 m, rainforest, 28.X1.—-1.XII.1998 (AJ. Ratsirarson); Toamasina, F.C. Sandranantitra, 18.04833°S, 49.09167°E, 450 m, rainforest, 21.1.1999 (ALJ. Ratsirarson); Toamasina, Torotoro- fotsy, 18.87082°S, 48.34737°E, 1070 m, montane rainforest, marsh edge, 24.—27. III.2004 (Malagasy ant team); Toamasina, Parc National de Zahamena, Tetezambat- ana forest, near junction of Nosivola and Manakambahiny rivers, 17.74298°S, 48.72936°E, 860 m, rainforest, 18.-19.II.2009 (B.L. Fisher et al.); Toamasina, Parc National de Zahamena, 17.73359°S, 48.72625°E, 950 m, rainforest, 19.I[.2009 (B.L. Fisher et al.); Toamasina, Parc National de Zahamena, Onibe River, 17.75908°S, 48.85468°E, 780 m, rainforest, 21.11.2009 (B.L. Fisher et al.); Toamasina, Parc Na- tional de Zahamena, Besaky River, 17.75244°S, 48.85321°E, 760 m, rainforest, 22. 11.2009 (BL. Fisher et al.); Yoliara, Réserve Spéciale d’Ambohijanahary, Forét d’Ankazotsihitafototra, 35.2 km 312° NW Ambaravaranala, 18.26667°S, 45.40667°E, 1050 m, montane rainforest, 13.-17.1.2003 (B.L. Fisher et al.); Toliara, Réserve Spé- ciale d’Ambohijanahary, Forét d’Ankazotsihitafototra, 34.6 km 314° NW Ambara- varanala, 18.26°S, 45.41833°E, 1100 m, montane rainforest, 16.1.2003 (B.L. Fisher et al.); Toliara, Forét Classée d’Analavelona, 29.2 km 343° NNW Mahaboboka, 22.675°S, 44.19°E, 1100 m, montane rainforest, 18.—22.I]].2003 (B.L. Fisher et al.); Toliara, Forét Classée d’Analavelona, 29.4 km 343° NNW Mahaboboka, 22.675°S, 44.18667°E, 1050 m, montane rainforest, 21.11.2003 (B.L. Fisher et al.); Toliara, Rés. Andohahela, 13 km NW Enakara, 24.55°S, 46.8°E, 1250 m, montane rainforest, 30.X1.1992 (B.L. Fisher); Toliara, Rés. Andohahela, 11 km NW Enakara, 24.5667°S, Taxonomy of Malagasy Tetramorium 93 46.83333°E, 800 m, rainforest, 17.-20.X1.1992 (B.L. Fisher); Toliara, Parc National d’Andohahela, Col du Sedro, 3.8 km 113° ESE Mahamavo, 37.6 km 341° NNW Tolagnaro, 24.76389°S, 46.75167°E, 900 m, montane rainforest, 21.—25.1.2002 (B.L. Fisher et al.); Toliara, Forét Ivohibe 55.0 km N Tolagnaro, 24.569°S, 47.204°E, 200 m, rainforest, 2.—4.XII.2006 (B.L. Fisher et al.); Toliara, Réserve Spéciale Kalambatri- tra, Ampanihy, 23.463°S, 46.47057°E, 1269 m, montane rainforest, 10.11.2009 (B.Z. Fisher et al.). Diagnosis. Tetramorium proximum can be distinguished from the other members of the species complex on the basis of the following character combination: eyes rela- tively large (OI 24-28); antennal scapes short (SI 74—76); frontal carinae strongly de- veloped, noticeably raised, and reaching or ending shortly before posterior head mar- gin; propodeal spines short to medium-sized, elongate-triangular, and usually acute (PSLI 17-21); petiolar node high rounded nodiform, in profile around 1.7 to 1.9 times higher than long (LPel 52-60), and in dorsal view around 1.3 to 1.5 times wider than long (DPpI 135-153); mesosoma with four to nine pairs of long, fine, standing hairs, usually five or six ranging from anterior pronotum to metanotal groove. Worker measurements (N=12). HL 0.72-0.88 (0.78); HW 0.65—0.81 (0.72); SL 0.49-0.60 (0.54); EL 0.18-0.20 (0.19); PH 0.34-0.40 (0.37); PW 0.49-0.60 (0.54); WL 0.92-1.07 (0.98); PSL 0.12-0.18 (0.15); PTL 0.15-0.19 (0.18); PTH 0.29-0.35 (0.32); PTW 0.18—0.23 (0.21); PPL 0.19-0.23 (0.22); PPH 0.29-0.35 (0.31); PPW 0.28-0.34 (0.31); CI 89-94 (92); SI 74-76 (75); OI 24-28 (26); DMI 53-58 (55); LMI 37-40 (38); PSLI 17—21 (19); PeNI 37-42 (39); LPeI 52-60 (56); DPelI 111-130 (120); PpNI 52-61 (57); LPpI 65-74 (69); DPpI 135-153 (145); PPI 135-160 (148). Worker description. Head much longer than wide (CI 89-94); posterior head margin weakly concave. Anterior clypeal margin with distinct median impression. Frontal carinae strongly developed, noticeably raised, diverging posteriorly, and either reaching or ending shortly before posterior head margin. Antennal scrobes present and distinct but weak, shallow and without clear and distinct posterior and ventral margins. Antennal scapes short, not reaching posterior head margin (SI 74-76). Eyes relatively large (24-28). Mesosomal outline in profile flat to weakly convex, relatively low and long (LMI 37-40), moderately marginate from lateral to dorsal mesosoma; promesono- tal suture absent; metanotal groove weakly to moderately developed. Propodeal spines short to medium-sized, elongate-triangular, and usually acute (PSLI 17—21), propodeal lobes short, triangular, and blunt or acute, always much shorter than propodeal spines. Petiolar node in profile high nodiform with well-rounded antero- and posterodorsal margins and strongly convex dorsum, around 1.7 to 1.9 times higher than long (LPel 52-60), anterior and posterior faces approximately parallel, antero- and posterodorsal margins situated at about same height; petiolar node in dorsal view around 1.1 to 1.3 times wider than long (DPel 111-130), in dorsal view pronotum around 2.4 to 2.7 times wider than petiolar node (PeNI 37-42). Postpetiole in profile subglobular and weakly anteroposteriorly compressed, around 1.4 to 1.5 times higher than long (LPpI 65-74); in dorsal view around 1.3 to 1.5 times wider than long (DPpI 135-153), pro- 94 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) notum around 1.7 to 1.9 times wider than postpetiole (PpNI 52-61). Postpetiole in profile appearing approximately of same volume as petiolar node, and both nodes of approximately similar height; postpetiole in dorsal view between 1.3 to 1.6 times wider than petiolar node (PPI 135-160). Mandibles completely unsculptured, smooth, and shiny; clypeus longitudinally rugose with usually three to five rugae, median ruga usually distinct and fully developed, sometimes partly reduced or broken, one or two weaker ru- gae present on each side; cephalic dorsum between frontal carinae longitudinally rugose with six to nine rugae, rugae running from posterior clypeal margin to posterior head margin, but often interrupted, splitting up or with cross-meshes; scrobal area mostly unsculptured; lateral head reticulate-rugose to longitudinally rugose. Ground sculpture on head weakly to moderately punctate. Dorsum of mesosoma mostly reticulate-rugose with smaller proportions of irregularly longitudinally rugose sculpture medially; lateral mesosoma mostly irregularly longitudinally rugose. Forecoxae mostly unsculptured, smooth, and shining, sometimes with traces of ground sculpture. Ground sculpture on mesosoma usually weak to absent. Both waist segments and gaster fully unsculptured, smooth, and shining. Dorsum of head with several pairs of long, fine, standing hairs; mesosoma with four to nine pairs of long, fine, standing hairs, usually with five or six ranging from anterior pronotum to metanotal groove, rarely propodeum with one pair anteriorly; waist segments and first gastral tergite without any standing hairs; appressed pubescence on first gastral tergite highly variable: ranging from short to very short and relatively scarce to relatively long and dense. Anterior edges of antennal scapes and dor- sal (outer) surfaces of hind tibiae usually with appressed, rarely decumbent, hairs. Head, mesosoma, waist segments, and gaster light brown to reddish brown, rarely of darker brown; mandibles, antennae, and legs lighter, usually light yellowish brown. Distribution and biology. Tetramorium proximum has a very broad distribution in Madagascar, and is indeed found in most sampled rainforests or montane rainforests at elevations from 25 to 1680 m (Fig. 64). The southernmost locality is Andohahela in the southeast; from there the species ranges with few interruptions up to Montagne d’Ambre at the northern tip of Madagascar. In addition to the eastern and northern forest belt, 7. proximum is also found in the few isolated humid forests located further west, such as Analavelona, Isalo, and Ambohijanahary. Tetramorium proximum ap- pears to live in leaf litter. Discussion. Tetramorium proximum is a very distinct species within the species complex. Due to its large body size (HW 0.65—0.81; WL 0.92-1.07) and strongly developed frontal carinae it cannot be mistaken for the five smaller species T: aspis, T: camelliae, T: cognatum, T. karthala, and T: rumo (HW 0.43-0.57; WL 0.56-0.81), all of which have weakly to moderately developed frontal carinae. Nor can 7. proximum with its larger eyes (OI 24—28) and its long, standing pilosity on the dorsal mesosoma be confused with the smaller-eyed T: gladius (OI 19-20) or T. freya, which lacks any standing pilosity on the mesosoma. The remaining two species, 7. myrmidon and T: tenuinode, are both morphologically close and relatively similar to T. proximum. The separation of these three requires more attention. Taxonomy of Malagasy Tetramorium 95 Tetramorium tenuinode, despite sharing a superficially similar habitus, differs from T. proximum in a number of characters, and co-occurs with the latter species throughout most of its distribution range while both species maintain their species- specific differences. The main diagnostic character is the pilosity on the dorsal meso- soma, which consists of two pairs of long, standing hairs (one on anterior pronotum and one on anterior mesonotum) in T: tenuinode, while in T. proximum there are usually five to six pairs (rarely four or more than six) from anterior pronotum to posterior mesonotum. Furthermore, 7. proximum has significantly longer antennal scapes (SI 74-77) than T. tenuinode (SI 66-70), and is normally a darker colour. Also, T. tenuinode generally has a thinner petiolar node in lateral view that is around 1.8 to 2.2 times higher than long (LPel 45—54), while the node of 7. proximum is around 1.7 to 1.9 times higher than long (LPel 52-60). There is some overlap in these morphometric ranges, but there is a strong general trend towards a much thin- ner node in 7! tenuinode, which together with the other characters mentioned above works very well in distinguishing both species. The remaining species, 7. myrmidon, might be confused with 7: proximum since they share most morphological characters and their morphometric ranges overlap. Nevertheless, 7. myrmidon is only found in Ambohijanahary where it co-occurs with 7. proximum, and there they are both easily separable based on mesosomal pilosity (two pairs in 7: myrmidon vs. five or six in T- proximum) and petiolar node height. Tetramorium proximum is a relatively variable species with respect to certain char- acters. [he greatest variation can be seen in gastral pubescence, which is always strong- ly appressed, but can vary from very short and scarce to relatively long and dense. Often this variation can be observed within the same locality or series, sometimes all specimens from one locality have short pubescence while populations from other localities have long pubescence. However, it seems that there is also a geographical component to this variation. The material from the southeast has predominantly long and dense pubescence, and the form with short pubescence is relatively rare, but still present. The material from central eastern Madagascar is a well-balanced mix of short and long pubescence. Interestingly, the material from the northeast, north, and the isolated forests in the west possesses almost exclusively short and scarce pubescence. Another, even though less variable character, is the shape of the petiolar node, which is high nodiform with a rounded dorsum, but can vary in height or thickness from population to population. Furthermore, some populations have fewer or more pairs of long, standing hairs on the mesosomal dorsum. ‘The most common count is five to six pairs ranging throughout most of the distribution range, but in some localities one can see only four pairs while in others six to seven, and in a few series seven to eight pairs are common. The variation observed in 7. proximum is not surprising, however, due to its wide distribution and commonness. Compared to other widely distributed and common Malagasy or Afrotropical Tetramorium species, this species actually shows much less variation. 96 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Figure 37. 7° proximum holotype worker (CASENT0906146). A Body in profile B Body in dorsal view C Head in full-face view. Taxonomy of Malagasy Tetramorium 97 Tetramorium rumo Hita Garcia & Fisher, sp. n. http://zoobank.org/555CD002-455A-4AE0-A404-99 F38EEFF755 http://species-id.net/wiki/Tetramorium_rumo Figs 22B, 23D, 24D, 27A, 38, 64 Type material. Holotype, pinned worker, MADAGASCAR, Fianarantsoa, Réserve Speciale Manombo 24.5 km 228° Farafangana, 23.01583°S, 47.719°E, 30 m, low- land rainforest, collection code BLF13963, 20.IV.2006 (B.L. Fisher et al.) (CAS: CASENT0073025). Paratypes, seven pinned workers with same data as holo- type (BMNH: CASENT0071823; CAS: CASENT0071827; CASENT0072469; CASENT0073028; CASENT0073033; CASENT0073038; CASENT0073039). Non-type material MADAGASCAR: Fianarantsoa, Réserve Forestiére d’Agnalazaha, Mahabo, 42.9 km 215° Farafangana, 23.19383°S, 47.723°E, 20 m, littoral rainforest, 19.IV.2006 (B.L. Fisher et al.); Fianarantsoa, P.N. Ranomafana, Tolongoina-Ampasimpotsy 3, 21.47412°S, 47.55742°E, 520 m, stomach contents of Mantella bernhardi Vences et al., 11.1V.2003 (V.C. Clark); Fianarantsoa, Forét de Vevembe, 66.6 km 293° Farafangana, 22.791°S, 47.18183°E, 600 m, rainforest, transition to montane forest, 23.-24.IV.2006 (B.L. Fisher et al.); Toamasina, 6 km ESE Andasibe (=Perinet), 18.95°S, 48.46667°E, 900 m, rainforest, 17.XI.1990 (2.S. Ward); Toamasina, F.C. Andriantantely, 18.695°S, 48.81333°E, 530 m, rainforest, 7.X11.1998 (Hj. Ratsirarson); Toamasina, Ankerana, 18.4061S, 48.82029°E, 725 m, rainforest, 16.—-21.1.2012 (B.L. Fisher et al.); Toamasina, Ankerana, 18.40829°S, 48.82107°E, 750 m, rainforest, 21.—-26.1.2012 (B.L. Fisher et al.); Toamasina, Reserve Betampona, Camp Vohitsivalana, 37.1 km 338° Toamasina, 17.88667°S, 49.2025°E, 520 m, rainforest, 1.—3.XII.2005 (B.L. Fisher et al.); Toamasina, F.C. Sandrananti- tra, 18.04833°S, 49.09167°E, 450 m, rainforest, 21.1.1999 (H./. Ratsirarson); Toli- ara, Rés. Andohahela, 10 km NW Enakara, 24.56667°S, 46.81667°E, 420-430 m, rainforest, 15.-22.X1.1992 (B.L. Fisher); Toliara, Rés. Andohahela, 13 km NW Ena- kara, 24.55°S, 46.8°E, 1250 m, montane rainforest, 30.X1.1992 (B.L. Fisher); Toli- ara, Rés. Andohahela, 6 km SSW Eminiminy, 24.73333°S, 46.8°E, 330 m, rainforest, 411.1993 (P.S. Ward); Toliara, Parc National Andohahela, Col de Tanatana, 33.3 km NW Tolagnaro, 24.7585°S, 46.85367°E, 275 m, rainforest, 28.X1.2006 (B.L. Fisher et al.); Toliara, Andohahela, 24.77639°S, 46.70528°E, 320 m, 9.XII.2007 (A. Ballerio); Toliara, Forét Ivohibe, 55.0 km N Tolagnaro, 24.569°S, 47.204°E, 200 m, rainforest, 2.—4.X11.2006 (B.L. Fisher et al.). Diagnosis. Tetramorium rumo can be well recognised within the T. cognatum spe- cies complex on the basis of the following character combination: very large eyes (Ol 28-31); antennal scapes very short (SI 60-66); propodeal spines moderately long, elongate-triangular to spinose, and usually acute (PSLI 22—26); propodeal spines and lobes not strongly inclined towards each other; petiolar node thinly cuneiform and moderately squamiform, in profile around 2.3 to 2.7 times higher than long (LPel 37-43), in dorsal view between 1.5 to 1.7 times wider than long (DPel 156-171); 98 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) mesosoma either with at least seven pairs of long, standing hairs on pronotum and mesonotum and propodeum sometimes with one pair anteriorly, or with just two pairs of long, standing hairs, one on anterior pronotum and one on anterior mesonotum. Worker measurements (N=12). HL 0.47-0.55 (0.51); HW 0.43—0.49 (0.45); SL 0.26—0.32 (0.29); EL 0.12—0.14 (0.13); PH 0.21-0.24 (0.23); PW 0.34-0.39 (0.36); WL 0.56-0.67 (0.60); PSL 0.11-0.13 (0.12); PTL 0.07—-0.09 (0.08); PTH 0.19-0.23 (0.21); PTW 0.12-0.15 (0.14); PPL 0.12-0.15 (0.13); PPH 0.17—0.20 (0.19); PPW 0.18-0.22 (0.20); CI 88-91 (90); SI 60-66 (64); OI 28-31 (29); DMI 58-62 (60); LMI 36-38 (38); PSLI 22—26 (23); PeNI 35-39 (38); LPel 37-43 (41); DPel 156-171 (162); PpNI 51-56 (54); LPpI 68-75 (72); DPpI 138-152 (146); PPI 141-152 (145). Worker description. Head much longer than wide (CI 88-91); in full-face view posterior head margin weakly concave. Anterior clypeal margin with distinct median impression. Frontal carinae very weakly developed, only feebly raised, usually ending shortly after posterior eye margin or merging with cephalic sculpture halfway between posterior eye margin and posterior head margin. Antennal scrobes very weak to absent, very shallow and without clear and distinct posterior and ventral margins. Antennal scapes very short, not reaching posterior head margin (SI 60-66). Eyes very large (Ol 28-31). Mesosomal outline in profile flat to weakly convex, comparatively low and long (LMI 36-38), moderately marginate from lateral to dorsal mesosoma; prome- sonotal suture absent; metanotal groove very weak to absent. Propodeal spines moder- ately long, elongate-triangular to spinose, and usually acute (PSLI 22—26), propodeal lobes triangular and short, always much shorter than propodeal spines, in profile spines and lobes not strongly inclined towards each other. Petiolar node thinly cuneiform and moderately squamiform, in profile around 2.3 to 2.7 times higher than long (LPel 37-43), anterior and posterior faces not parallel, anterodorsal margin usually situated higher and more strongly angled than posterodorsal margin, petiolar dorsum relatively flat to weakly convex and tapering backwards posteriorly; node in dorsal view between 1.5 to 1.7 times wider than long (DPel 156-171), in dorsal view pronotum around 2.6 to 2.8 times wider than petiolar node (PeNI 35-39). Postpetiole in profile globu- lar, between 1.3 to 1.5 times higher than long (LPpI 68-75); in dorsal view around 1.4 to 1.5 times wider than long (DPpI 138-152), pronotum around 1.8 to 1.9 times wider than postpetiole (PpNI 51-56). Postpetiole in profile appearing shorter and thicker than petiolar node, postpetiole in dorsal view around 1.4 to 1.5 times wider than petiolar node (PPI 141-152). Mandibles completely unsculptured, smooth, and shiny; clypeus weakly irregularly longitudinally rugulose, median rugula usually pre- sent but rarely fully developed, one or two mostly broken lateral rugulae present on each side; cephalic dorsum between frontal carinae longitudinally rugulose with eight to eleven fine rugulae, rugulae usually running from posterior clypeal margin to pos- terior head margin, often irregularly shaped, interrupted or with cross-meshes; scrobal area mostly unsculptured; lateral head mainly longitudinally rugulose to reticulate- rugulose, but larger areas often only weakly sculptured and appearing fairly smooth and shiny. Ground sculpture on head variable, ranging from weakly developed or absent to moderately punctate. Dorsum of mesosoma mostly longitudinally rugulose; Taxonomy of Malagasy Tetramorium 99 lateral mesosoma mostly unsculptured with smaller, irregularly longitudinally rugu- lose or reticulate-rugulose areas. Ground sculpture on mesosoma very weak to absent. Forecoxae usually unsculptured, smooth, and shining. Both waist segments and gaster fully unsculptured, smooth, and shining. Dorsum of head with several pairs of long, fine, standing hairs; pilosity on dorsal mesosoma variable: southern populations with at least seven pairs on pronotum and mesonotum, propodeum sometimes with one pair anteriorly, and northern populations with only two pairs, one on anterior prono- tum and one on anterior mesonotum; waist segments and first gastral tergite without any standing hairs; first gastral tergite with short, dense, appressed (rarely decumbent) pubescence. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae with appressed to decumbent hairs. Body usually uniformly whitish yellow to light brown, very rarely darker. Etymology. The new species is named after the fictional character “Rumo” from Walter Moers’ fantasy novel “Rumo and His Miraculous Adventures”. Tetramorium rumo is a very bright species, almost white, with distinct propodeal spines reminiscent of “Rumo”, who is a white wolperting with short but acute horns. ‘The species epithet is an arbitrary combination of letter, thus invariable. Distribution and biology. The new species is another rainforest inhabitant mainly found in eastern Madagascar (Fig. 64). However, the distribution range is somewhat unusual since 7° rumo is found in the southeast from Andohahela to Ranomafana, and then much further north from Perinet to Betampona. ‘The reasons for this patchy distribution are unclear. Tetramorium rumo was mainly sampled in rainforests, rarely montane rainforests or littoral rainforests, at elevations ranging from 20 to 1250 m. In addition, 7: rumo appears to live in leaf litter. Discussion. Tetramorium rumo is very unlikely to be confused with the much larger species T. freya, T. gladius, T. myrmidon, T. proximum, and T. tenuinode. All the latter species, except T° freya, also have very well developed frontal carinae, which contrast with the reduced and very weak frontal carinae seen in 7. rumo. Tetramorium freya has weaker frontal carinae than the other four species, but in contrast to 7? rumo this species does not have any standing pilosity on the mesosomal dorsum. Further- more, 7: gladius has very small eyes (OI 19-20) compared to T. rumo, which has very large eyes (OI 28-31). The remaining four species of the 7: cognatum complex are morphologically much closer to 7: rumo than the five species mentioned above. However, 7: camelliae is easily separable from T: rumo on the basis of the petiolar node shape, which is strongly squamiform and transverse in the former (LPel 33-36; DPel 228-238), contrasting with the highly nodiform to thinly cuneiform node of the latter (LPel 37-43; DPel 156-171). Tetramorium aspis and T. karthala both have longer antennal scapes (SI 68—74), shorter propodeal spines (PSLI 18-22), thicker and lower petiolar nodes in profile (LPel 46-54), and less transverse nodes in dorsal view (DPel 146-161) compared to 7. rumo (SI 60-66; LPel 37-43; DPelI 156-171). In addition, T. aspis has propodeal spines and lobes strongly inclined towards each other, an ar- rangement absent in 7. rumo, and T. karthala is only found on the Comorian island of Grand Comore while 7. rumo is distributed in Madagascar. 100 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Figure 38. 7) rumo holotype worker (CASENT0073025). A Body in profile B Body in dorsal view C Head in full-face view. Taxonomy of Malagasy Tetramorium 101 The last species of the T° cognatum complex, T: cognatum itself, is probably the closest relative of 7. rumo within the complex, and without careful examination they could be confused in some cases. However, both differ clearly in propodeal spine length and petiolar node shape. Tetramorium rumo has relatively long and spinose propodeal spines (PSLI 22-26) which contrast with the much more reduced, short, and triangular teeth of 7: cognatum (PSLI 12-16). Also, the thinly cuneiform petiolar node of 7! rumo, which is 2.3 to 2.7 times higher than long (LPel 37-43) and around 1.6 to 1.7 times wider than long (DPel 156-171), discriminates it from T° cognatum, in which the node is 1.8 to 2.0 times higher than long (LPel 49-55) and around 1.3 to 1.4 times wider than long (DPel 129-142). Nonetheless, the species most similar to 7: rumo is likely T: rala from the T) schau- Jussi complex. Both species share a very similar habitus. Compared to all other species of the 7: schaufussi species group, they are smaller in size, have relatively longer propodeal spines, a high nodiform to thinly cuneiform petiolar node, lack standing pilosity on the waist segments, and possess very bright body colouration. We have placed them in different species complexes on the basis of the presence (7! ra/a) or absence (T. rumo) of standing pilosity on the first gastral tergite, but otherwise these two species could be easily confused. However, despite the very strong similarities, substantial differences separate these species from each other. Most obviously, the petiolar node of 7: rumo is thinner and stronger anteroposteriorly compressed, in profile around 2.3 to 2.7 times higher than long (LPel 37-43), and in dorsal view between 1.5 to 1.7 times wider than long (DPel 156-171). By contrast, T° rala has a node in profile is 2.0 to 2.2 times higher than long (LPel 45-50), and in dorsal view around 1.3 to 1.5 times broader than long (DPel 129-145). Also, T° rumo has larger eyes (OI 28-31) than 7! rala (Ol 26-28). In addition, their distribution ranges strongly overlap in central-eastern Mada- gascar and both species maintain their species identities without intermediate forms. As mentioned above, there is some noteworthy variation in mesosomal pilosity in T. rumo. The populations in the southeast from Andohahela to Vevembe and the type locality Manombo all have at least seven pairs of long, standing hairs on the pronotum and mesonotum and the propodeum sometimes has one pair anteriorly, whereas the populations further north from Perinet to Betampona all have just two pairs of long, standing hairs (one on the anterior pronotum and one on the anterior mesonotum). Nevertheless, since both population groups are widely separated geographically and other character divides them, we consider this a clear case of geographical variation. Tetramorium tenuinode Hita Garcia & Fisher, sp. n. http://zoobank.org/4856B8A4-FBBA-4629-8619-21F62101BAA7 http://species-id.net/wiki/Tetramorium_tenuinode Figs 24B, 25C, 26A, 26B, 39, 64 Type material. Holotype, pinned worker, MADAGASCAR, Fianarantsoa, Parc Na- tional de Ranomafana, Vatoharanana River, 4.1 km 231° SW Ranomafana, 21.29°S, 102 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) 47 43333°E, 1100 m, montane rainforest, sifted litter (leaf mold, rotten wood), collec- tion code BLF08400, 27.—31.III.2003 (B.L. Fisher et al.) (CAS: CASENT0040115). Paratypes, 41 pinned workers with same data as holotype (CAS: CASENT0039644; CASENT0039646; CASENT0039651; CASENT0039655; CASENT0039660; CASENT0039664; CASENT0039668; CASENT0039671; CASENT0039673; CASENT0039737; CASENT0039739; CASENT0039745; CASENT0039749; CASENT0039750; CASENT0039754; CASENT0039759; CASENT0039761; CASENT0039809; CASENT0040000; CASENT0040020; CASENT0040023; CASENT0040035; CASENT0040036; CASENT0040090; CASENT0040092; CASENT0040096; CASENT0040099; CASENT0040105; CASENT0040106; CASENT0040112; CASENT0040123; CASENT0040181; CASENT0040279; CASENT0040284; CASENT0040285; CASENT0040293; CASENT0040296; CASENT0040304; CASENT0040306; CASENT0040311; CASENT0040317); and nine pinned workers with same data as holotype except collected ex rotten log and collection code BLF08488 (BMNH: CASENT0497623; CAS: CASENT0497621; MCZ: CASENT0497622). Non-type material. MADAGASCAR: Antsiranana, Forét Ambanitaza, 26.1 km 347° Antalaha, 14.67933°S, 50.18367°E, 240 m, rainforest, 27.X1.1994 (B.L. Fish- er); Antsiranana, 1 km W Andampibe, Cap Masoala, 15.69361°S, 50.18139°E, 125 m, lowland rainforest, 1.XII.1993 (G.D. Alpert); Fianarantsoa, 45 km S Ambalavao, 22.21667°S, 47.01667°E, 785 m, rainforest, 25.[X.1993 (B.L. Fisher); Fianarantsoa, 45 km S Ambalavao, 22.21667°S, 47.01667°E, 720 m, rainforest edge, 31.X.1993 (B.L. Fisher); Fianarantsoa, 2 km W Andrambovato, along river Tatamaly, 21.51167°S, 47.41°E, 1075 m, montane rainforest, 3.—5.VI.2005 (B.L. Fisher et al.); Fianarantsoa, Rés. Andringitra, 43 km S Ambalavao, 22.23333°S, 47°E, 825 m, rainforest, 5.X.1993 (B.L. Fisher); Fianarantsoa, R.S. Ivohibe, 7.5 km ENE Ivohibe, 22.47°S, 46.96°E, 900 m, rainforest, 7.-12.X.1997 (B.L. Fisher); Fianarantsoa, R.S. Ivohibe, 8.0 km E Ivo- hibe, 22.48333°S, 46.96833°E, 1200 m, montane rainforest, 15.-21.X.1997 (BL. Fisher); Fianarantsoa, 9.0 km NE Ivohibe, 22.42667°S, 46.93833°E, 900 m, rainforest, 17.X1.1997 (B.L. Fisher); Fianarantsoa, Réserve Speciale Manombo 24.5 km 228° Far- afangana, 23.01583°S, 47.719°E, 30 m, lowland rainforest, 20.1V.2006 (B.L. Fisher et al.); Fianarantsoa, Ranomafana Nat. Park, Maharira forest, 21°S, 47°E, 1190 m, mon- tane forest, 11.X.1992 (E. Rajeriarison); Fianarantsoa, Parc National de Ranomafana, Vatoharanana River, 4.1 km 231° SW Ranomafana, 21.29°S, 47.43333°E, 1100 m, montane rainforest, 27.—31.[.2003 (B.L. Fisher et al.); Fianarantsoa, Forét de Vevem- be, 66.6 km 293° Farafangana, 22.791°S, 47.18183°E, 600 m, rainforest, transition to montane forest, 23.1V.2006 (B.L. Fisher et al.); Toamasina, Montagne d’Akirindro 7.6 km 341° NNW Ambinanitelo, 15.28833°S, 49.54833°E, 600 m, rainforest, 17.-21.111.2003 (B.L. Fisher et al); Toamasina, 6.3 km S Ambanizana, Andranobe, 15.6813°S, 49.958°E, 25 m, rainforest, 14.X1.1993 (B.L. Fisher); Toamasina, Réserve Spéciale Ambatovaky, Sandrangato river, 16.77274°S, 49.26551°E, 450 m, rainfor- est, 20.—22.11.2010 (B.L. Fisher et al.); Toamasina, Réserve Spéciale Ambatovaky, San- drangato river, 16.7633°S, 49.26692°E, 520 m, rainforest, 22.—24.11.2010 (B.L. Fisher Taxonomy of Malagasy Tetramorium 103 et al.); Toamasina, Réserve Spéciale Ambatovaky, Sandrangato river, 16.81753°S, 49.29498°E, 360 m, rainforest, 25.—27.11.2010 (B.L. Fisher et al.); Toamasina, Forét Ambatovy, 14.3 km 57° Moramanga, 18.85083°S, 48.32°E, 1075 m, montane rainfor- est, 21.11.2004 (Malagasy ant team); Toamasina, Ambatovy, 12.4 km NE Moramanga, 18.83937°S, 48.30842°E, 1080 m, montane rainforest, 4.-8.]]].2007 (B.L. Fisher et al.); Toamasina, Analamay, 18.80623°S, 48.33707°E, 1068 m, montane rainforest, 21.10.2004 (Malagasy ant team); Toamasina, Corridor Forestier Analamay-Mantadia, Ambatoharanana, 18.80388°S, 48.40506°E, 1013 m, rainforest, 12.—19.XII.2012 (B.L. Fisher et al.); Toamasina, Corridor Forestier Analamay-Mantadia, Ambohibolake- ly, 18.76087°S, 48.37128°E, 1044 m, rainforest, 23.-28.X1.2012 (B.L. Fisher et al.); Toamasina, Corridor Forestier Analamay-Mantadia, Ambohibolakely, 18.77898°S, 48.36375°E, 918 m, rainforest, 23.-28.X1.2012 (B.L.Fisher et al.); Toamasina, Cor- ridor Forestier Analamay-Mantadia, Ambohibolakely, 18.76131°S, 48.36437°E, 983 m, rainforest, 26.X1.2012 (B.L.Fisher et al.); Toamasina, Corridor Forestier Analamay- Mantadia, Tsaravoniana, 18.76124°S, 48.42134°E, 939 m, rainforest, 2.-7.XII.2012 (B.L.Fisher et al.); Yoamasina, Corridor Forestier Analamay-Mantadia, Tsaravoni- ana, 18.76465°S, 48.41938°E, 1039 m, rainforest, 2.-7.X1I.2012 (B.L. Fisher et al.); Toamasina, Station Forestiére Analamazaotra, Analamazaotra 1.3 km S Andasibe, 18.38466°S, 48.41271°E, 980 m, montane rainforest, 11.-13.XII.2007 (B.L. Fisher et al.); Toamasina, F.C. Andriantantely, 18.695°S, 48.81333°E, 530 m, rainforest, 4 —7.X11.1998 (HJ. Ratsirarson); Toamasina, Montagne d’Anjanaharibe, 18.0 km 21° NNE Ambinanitelo, 15.18833°S, 49.615°E, 470 m, rainforest, 8.—12.III.2003 (B.L. Fisher et al.); Toamasina, Ankerana, 18.40829°S, 48.82107°E, 750 m, rainforest, 21.— 26.1.2012 (BL. Fisher et al.); Toamasina, Ankerana, 18.4104°S, 48.8189°E, 855 m, rainforest, 22.—27.1.2012 (B.L. Fisher et al.); Toamasina, Reserve Betampona, Camp Vohitsivalana, 37.1 km 338° Toamasina, 17.88667°S, 49.2025°E, 520 m, rainforest, 1.—3.X11.2005 (B.L. Fisher et al.); Toamasina, Reserve Betampona, Camp Rendriren- dry, 34.1 km 332° Toamasina, 17.924°S, 49.19967°E, 390 m, rainforest, 28.X1.2005 (B.L. Fisher et al.); TYoamasina, F.C. Didy, 18.19833°S, 48.57833°E, 960 m, rainforest, 16.—23.XIJ.1998 (HJ. Ratsirarson); Toamasina, Parc National Mananara-Nord, 7.1 km 261° Antanambe, 16.455°S, 49.7875°E, 225 m, rainforest, 14.X1.2005 (B.L. Fisher et al.); Toamasina, 19 km ESE Maroantsetra, 15.48333°S, 49.9°E, 350 m, rainforest, 22.1V.1989 (P.S. Ward); Toamasina, 16 km S Moramanga, 19.08333°S, 48.23333°E, 950 m, rainforest, 18.X1.1990 (P.S. Ward); Toamasina, Moramanga, 18.94417°S, 48.23067°E, 922 m, urban/garden, 14.11.2007 (B.L. Fisher et al.); Toamasina, F.C. San- dranantitra, 18.04833°S, 49.09167°E, 450 m, rainforest, 18.—24.1.1999 (HJ. Ratsirar- son); Toamasina, Parc National de Zahamena, Onibe River, 17.75908°S, 48.85468°E, 780 m, rainforest, 21.—23.11.2009 (B.L. Fisher et al.); Toamasina, Parc National de Zahamena, Tetezambatana forest, near junction of Nosivola and Manakambahiny riv- ers, 17.74298°S, 48.72936°E, 860 m, rainforest, 18.-19.I].2009 (BL. Fisher et al.); Toliara, Rés. Andohahela, 10 km NW Enakara, 24.56667°S, 46.81667°E, 430 m, rain- forest, 22.X1.1992 (B.L. Fisher); Toliara, Rés. Andohahela, 6 km SSW Eminiminy, 24.73333°S, 46.8°E, 330 m, rainforest, 4.11.1993 (G.D. Alpert & P.S. Ward); Toliara, 104 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Parc National d’Andohahela, Col du Sedro, 3.8 km 113° ESE Mahamavo, 37.6 km 341° NNW Tolagnaro, 24.76389°S, 46.75167°E, 900 m, montane rainforest, 21.— 25.1.2002 (B.L. Fisher et al); Toliara, Andohahela, 24.77639°S, 46.70528°E, 320 m, 9.XII.2007 (A. Ballerio); Toliara, Grand Lavasoa, 25.9 km W Tolagnaro, 25.08767°S, 46.749°E, 450 m, rainforest, 30.XI.-2.XII.2006 (B.L. Fisher et al.). Diagnosis. The following character set clearly distinguishes T. tenuinode from the remainder of the 7: cognatum species complex: eyes relatively large (OI 25—27); anten- nal scapes very short (SI 66—70); frontal carinae well developed, noticeably raised, and approaching or ending at posterior head margin; petiolar node high rounded nodiform, in profile around 1.8 to 2.2 times higher than long (LPel 45-54), in dorsal view around 1.3 to 1.4 times wider than long (DPel 125-143); mesosoma with only two pairs of long, standing hairs, one on anterior pronotum and one on anterior mesonotum. Worker measurements (N=12). HL 0.62-0.75 (0.71); HW 0.58-—0.70 (0.66); SL 0.40-0.47 (0.45); EL 0.16-0.18 (0.17); PH 0.30-0.36 (0.34); PW 0.42-0.54 (0.50); WL 0.76-0.94 (0.88); PSL 0.11-0.18 (0.14); PTL 0.13-0.17 (0.15); PTH 0.26-0.32 (0.30); PTW 0.16—-0.21 (0.20); PPL 0.15—0.22 (0.20); PPH 0.25-0.32 (0.30); PPW 0.24—0.33 (0.30); CI 91-93 (92); SI 66-70 (68); OI 25-27 (26); DMI 55-59 (57); LMI 38-40 (39); PSLI 17-24 (20); PeNI 36-42 (39); LPel 45-54 (49); DPel 125-143 (133); PpNI 57-64 (61); LPpI 60-72 (67); DPpI 143-168 (153); PPI 148-167 (155). Worker description. Head longer than wide (CI 91-93); in full-face view pos- terior head margin weakly to moderately concave. Anterior clypeal margin with dis- tinct median impression. Frontal carinae well developed, noticeably raised, diverg- ing posteriorly, approaching or ending at posterior head margin. Antennal scrobes weakly developed, shallow and without clear and distinct posterior and ventral mar- gins. Antennal scapes very short, not reaching posterior head margin (SI 66-70). Eyes relatively large (OI 25-27). Mesosomal outline in profile flat to weakly convex, moderately low and long (LMI 38-40), moderately marginate from lateral to dor- sal mesosoma; promesonotal suture absent; metanotal groove very weak to absent. Propodeal spines short to moderately long, usually elongate-triangular, and acute (PSLI 17-24), propodeal lobes short and triangular, always much shorter than pro- podeal spines. Petiolar node in profile high rounded nodiform and relatively thin, around 1.8 to 2.2 times higher than long (LPel 45-54), anterior and posterior faces approximately parallel, anterodorsal and posterodorsal margins usually situated at about same height and moderately rounded, petiolar dorsum distinctly convex; peti- olar node in dorsal view around 1.2 to 1.4 times wider than long (DPel 125-143), in dorsal view pronotum between 2.4 to 2.8 times wider than petiolar node (PeNI 36-42). Postpetiole in profile subglobular and weakly anteroposteriorly compressed, around 1.4 to 1.7 times higher than long (LPpI 60-72); in dorsal view between 1.4 to 1.7 times wider than long (DPpI 143-168), pronotum around 1.6 to 1.7 times wider than postpetiole (PpNI 57-64). Postpetiole in profile appearing more volu- minous than petiolar node, postpetiole in dorsal view around 1.5 to 1.7 times wider Taxonomy of Malagasy Tetramorium 105 than petiolar node (PPI 148-167). Mandibles completely unsculptured, smooth, and shiny; clypeus irregularly longitudinally rugose/rugulose with two to seven ru- gae/rugulae, median ruga rarely fully developed, usually broken, most other rugu- lae usually broken, sometimes merging with each other; cephalic dorsum between frontal carinae longitudinally rugose, usually with six to seven, rarely eight or nine rugae, rugae running from posterior clypeal margin to posterior head margin, often interrupted, splitting up or with cross-meshes, especially posteriorly; scrobal area partly unsculptured, but mostly merging with surrounding longitudinally rugose to reticulate-rugose sculpture present on lateral head. Ground sculpture on head weak- ly to moderately punctate. Dorsum and sides of mesosoma reticulate-rugose/rugu- lose to irregularly longitudinally rugose/rugulose, lateral pronotum sometimes only weakly sculptured and relatively smooth and shining. Forecoxae mostly unsculp- tured, smooth and shining, sometimes with traces of ground sculpture. Ground sculpture on mesosoma usually weak to absent. Both waist segments and gaster fully unsculptured, smooth, and shining. Dorsum of head with several pairs of long, fine, standing hairs; dorsum of mesosoma with two pairs only, one on anterior prono- tum and one on anterior mesonotum; propodeum, waist segments and first gastral tergite without any standing hairs at all; first gastral tergite with very short, scarce, appressed pubescence. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae usually with appressed, rarely decumbent, hairs. Head, mesosoma, waist segments and gaster usually uniformly chestnut brown to very dark brown, sometimes of lighter reddish brown; appendages usually of slightly lighter brown than remainder of body. Etymology. The name of the new species is a combination of the Latin adjective “tenuis”, meaning thin, and the Latin noun “nodus”, meaning node. Distribution and biology. Tetramorium tenuinode is widely distributed in eastern Madagascar (Fig. 64). It is found with few interruptions in an almost straight line from Grand Lavasoa and Andohahela in the southeast to Ambanitaza in the northeast. The new species clearly prefers rainforests and montane rainforests at elevations from 25 to 1200 m, even though it was also collected several times from guava forest and urban gardens. In addition, 7. tenuinode appears to be a leaf litter inhabitant since almost all of the material was collected from this microhabitat. Discussion. This new species is easily identifiable within its species complex. ‘The presence of well developed, raised, and long frontal carinae separates 7. tenuinode from the species 7. aspis, T. camelliae, T. cognatum, T. karthala, and T. rumo. These five species are generally also much smaller (WL 0.56-0.81) than T: tenuinode (WL 0.76—0.94). Also, T° tenuinode possesses much larger eyes (OI 25-27) than T: gladius (OI 19-20) and much shorter antennal scapes (SI 66-70) than 7: myrmidon and T. proximum (SI 74-77). Lastly, the presence of two pairs of long, standing hairs on the dorsal mesosoma distinguishes 7: tenuinode from T. freya, which lacks any standing hairs on the dorsal mesosoma. The latter species also displays weaker sculpture on the head and mesosoma than T° tenuinode. 106 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) C Head in full-face view. Taxonomy of Malagasy Tetramorium 107 It should be noted, however, that 7° tenuinode is morphologically still very close to T: proximum, and on very superficial observation it is possible to confuse both. At the initial stage of the revision we considered 7. tenuinode conspecific with T. proximum, a very variable species. However, after the detailed examination of more than five hundred pinned workers, it became apparent that the material consisted of these two fairly distinct species. Indeed, even though 7°. proximum is more broadly distributed than 7. tenuinode, both co-occur in sympatry throughout most of their respective distribution ranges, and both are always very well distinguishable. The most obvious character to separate both species is pilosity on the dorsal mesosoma. In 7. tenuinode this consists of two pairs of long, standing hairs only, one on anterior pronotum and one on anterior mesonotum, which contrasts with the usually five to six (rarely four or more than six) pairs found from anterior pronotum to posterior mesonotum in 7. proximum. In addition to mesosomal pilosity and the shorter an- tennal scapes mentioned above, T° tenuinode usually also has a thinner petiolar node in profile, which is around 1.8 to 2.2 times higher than long (LPel 45-54), while the node of 7. proximum is around 1.7 to 1.9 times higher than long (LPel 52-60). The identification key presented above might suggest that T° tenuinode can be easily confused with 7: myrmidon, but this is not the case since the two species differ in a number of characters. Among others, 7. tenuinode has a broader head (CI 91-93), shorter antennal scapes (SI 66—70), and a higher petiolar node (LPel 45—54) than T- myrmidon (CI 88-91; SI 74-76; LPel 58-60). In contrast to the more variable 7: proximum, T. tenuinode has little observed intraspecific variation. Some populations vary very slightly in propodeal spine length, thickness of the petiolar node, or body colouration. Tetramorium schaufussii species complex Comments. This complex contains ten species, which are characterised and distin- guishable from the T° cognatum species complex by their presence of standing pilosity on the first gastral tergite. All species are endemic to Madagascar, except 7: schaufussii, which is also known from Reunion. Also, they inhabit rainforests or montane rainfor- ests and are only occasionally found in other habitats. The taxonomy of this species complex is challenging. Some species like 7. nassonowii, T. pseudogladius, T. rala or T: scutum are relatively easily recognisable due to species-specific character states, but this is not true for the rest of the complex. Tetramorium schaufussii and T: xanthogaster are especially widespread species with high degrees of intraspecific variation, which causes a number of problems for the delimitation of species boundaries for most member of the complex. Intriguingly, only two species of this complex have very limited distribution ranges: T: scutum is only known from Ivohibe and T: pseudogladius from Zahamena. The remainder are all much more widespread. 108 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Identification key to species of the T) schaufussii species complex (workers) 1 Petiolar node relatively long and low, in profile around 1.2 to 1.4 times high- er than long (LPel 72-81) and in dorsal view between 1.0 to 1.2 times longer than wide (DPel 87-98) (Fig. 40A) [Madagascar] «0.0... T. nassonowti Petiolar node shorter and higher than above, in profile around 1.5 to 2.2 times higher than long (LPel 45-67) and in dorsal view between 1.1 to 1.5 times wider than long (DPelI 109-154) (Fig. 40B, C) oo. ceeeeeeeeeees 2 Eyes relatively small (OI 20) (Fig. 41A) [Madagascar]. ......T. pseudogladius Eyes usually much larger than above (OI 25-28, rarely OI 22—24) (Fig. 41A, TieS@)) Sehuceert Mee, rede nae ty Di es Naan a bs J eae ee Ler Een Be Gee 3 Propodeal spines moderate to long (PSLI 22—24) and propodeal lobes rela- tively well developed, spines and lobes strongly inclined towards each other ie 34270)s [Wia dae asear lic tnc. scutdbaseidan seule tessth, ruteta sesso eaartonscoe T. scutum Character combination never as above, propodeal spines usually shorter than PSLI 18, if longer than PSLI 21, then propodeal spines and lobes not in- clinedrtowatdsteach other (Rigs 42 BAC HID) esa. s ses swssnens eeenseisbenea robes Sieanes 4, Smaller species (HW 0.46-0.49; WL 0.58-0.64); propodeal spines relatively long (PSLI 21-25); petiolar node thinly cuneiform, in profile 2.0 to 2.2 times higher than long (LPel 45-50); waist segments without long, standing pilosity; body colour whitish yellow to light brown (Fig. 43A) [Madagascar]....... T. rala Character combination never as above, larger species (HW 0.53-0.84; WL 0.70—1.20) with relatively short propodeal spines/teeth, which are always shorter than above (PSLI 8-18); petiolar node variably shaped but always lower than above, in profile around 1.5 to 1.9 times higher than long (LPel 52-67); waist segments with or without long, standing pilosity; body colour EYE Of OTe aioe 575 ele Yas OP en ame ein ean oa een ge Re 5 Larger species (HW 0.71—0.84; WL 1.00—1.20); antennal scapes relatively long (SI. 77-82); dorsum of promesonotum with seven to ten pairs of long, standing hairs; waist segments usually without any long, standing pilosity, rarely one pair present on petiole or postpetiole (Fig. 44A) [Madagascar]............... T. obiwan Character combination never as above; antennal scapes always shorter (SI 66-75); usually smaller species, but if body size in range above HW 0.70 and WL 1.00, then both waist segments with several pairs of long, standing hairs (Vis Ail AC Me) be coc cue set asa aeet eran Peace ean aa cea ea ec aed aonieeen ene area naan 6 Dorsum of promesonotum usually with two to four pairs of long, stand- ing hairs; propodeum and waist segments without any standing pilosity (Fig. ASN) 3 NAGA CAS CAE] SRE: cident sceaiws oveaetentedsensttaee Si cielpbwctor ath se pBoanioin T. sikorae Dorsum of promesonotum always with more than four pairs of long, standing hairs; waist segments always with standing pilosity (Fig. 45B, C, D)........... 7. Bicoloured species, head and/or mesosoma very dark brown to black and strong- ly contrasting with yellow to light brown waist segments and gaster (rarely also mesosoma) (Fig. 46A, B) [Madagascar] ....... eee T. xanthogaster (in parts) Taxonomy of Malagasy Tetramorium 109 — More or less uniformly coloured species, sometimes gaster darker than rest of | srayek ral Wes reseacs 161 Cel BO) a) Ra Re ae Sti SRNR Diem AR IR isc ina Ian ee PPR ETO RIN 8 8 Head relatively long and thin (CI 86-90); clypeus always with well-developed and regular median longitudinal ruga (Fig. 47A, B) [Madagascar, Reunion]. shed edt amar i rh srdan ok aR NR Sr hai Re At ARR a T. schaufussii - Head relatively shorter and thicker (CI 92-95); clypeus usually with either median area unsculptured or traces of median ruga present but weakly devel- oped and irregularly shaped (Fig. 47C, D), very rarely with a well-developed Tone icucinal tug te A 7) dda snes teceecae Mecegeease Pare poperte ears tetaapepec cece edees 9 9 Frontal carinae relatively better developed; cephalic dorsum between frontal carinae with nine to thirteen relatively regularly shaped, mostly unbroken rugae; propodeal spines short to medium-sized (PSLI 18-22) (Fig. 48A, B) [Madagascar] bx, ..rata. scat ane teak dia g.cncse cies Aataecasnns MAsaRaare anata Ses T. monticola — Frontal carinae relatively more weakly developed; cephalic dorsum between frontal carinae with six to ten relatively irregularly shaped, often meandering or broken rugulae; propodeal spines reduced to short to very short teeth/ denticles HP SEE SiG). (BigeA SC, SOM. tosi.choascth extn taeSiache tes naoSen hs MSeaascan 10 10 Relatively smaller species (HW 0.54-0.69; WL 0.72—0.92); dorsum of pro- podeum usually with long, standing pilosity (Fig. 49A) [Madagascar] .......... Pec Bas tin eS OL en, eee eS, A T. xanthogaster (in parts) = Relatively larger species (HW 0.69-0.79; WL 1.01-1.10); dorsum of propodeum without any long, standing pilosity (Fig. 49B) [Madagascar] ... ck ARCO | Fe tI HR NORA 1 RR FR 1 T. merina Figure 40. Waist segments in dorsal view. A 7 nassonowii (CASENT0195504) B 7 merina (CASENT0437232) C 7 xanthogaster (CASENT0218041). : UE ia ; iL 4, . oP Figure 41. Head in profile. A 7) pseudogladius (CASENT0153605) BT’ scutum (CASENT0189116) C 7. xanthogaster (CASENT0189134) D T sikorae (CASENT0102402). 110 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Figure 42. Mesosoma in profile. A 7’ scutum (CASENT0189116) B 7’ monticola (CASENT0077172) C 7. schaufussii (CASENT0217760) D 7 merina (CASENT0437226). . 2, i. 4 ‘ /] zl ii ik . ai 4 . a Aga. } - 4 ne . t F ae = " - | . ; a a *| + 7 4 ‘ | 1" . 4 airy a a Fr Lo 1 tage ; ! = A i ‘ - iy 2 Figure 43. Head, mesosoma, and petiole in profile. A 7 rala (CASENT0162115) B Z obiwan (CASENT0447245) C 7 sikorae (CASENT0051946) D T xanthogaster (CASENT0006330). J pet } Z a _— a bee Se aed | P ag | A f he: . at \ a t = ae Z cs La 4 ' 7 7 & ‘ ‘eels | < [a fie : a 2 7 i << } a é sa _ ‘OS ome mm Figure 45. Head, mesosoma, and waist segments in profile. A 7’ sikorae (CASENT0102402) B T schau- Jussii (CASENT0474409) C T° monticola (CASENT0152401) D 7’ xanthogaster (CASENT0189134). Figure 46. Body in profile. A 7’ xanthogaster (CASENT0485100) B 7 xanthogaster (CASENT0189134) C 7. schaufussii (CASENT0152490) D 7 merina (CASENT0437226). Taxonomy of Malagasy Tetramorium 1a Figure 47. Head in full-face view. A JT whan AGENT A7LEOD) B TZ schaufussii (CASENT0497951) © T merina (CASENT0437232) D TZ monticola (CASENT0152427) E 7. xan- thogaster (CASENT0491748). Figure 48. Head in full-face view. A 7’ monticola(CASENT0152427) B 7 monticola(CASENT0497970) C 7. merina (CASENT0437232) D T. xanthogaster (CASENT0189134). pied 49. Body in profile. A 7’ ae TehseRe ean B 7. merina exeEN aoe Tetramorium merina Hita Garcia & Fisher, sp. n. http://zoobank.org/EDBCDB39-4EDB-439C-98 1 2-7675585170BD http://species-id.net/wiki/Tetramorium_merina Figs 20C, 40B, 42D, 44D, 46D, 47C, 48C, 49B, 50, 65 Type material. Holotype, pinned worker, MADAGASCAR, Antananarivo, Réserve Spé- ciale d’Ambohitantely, Forét d’Ambohitantely, 20.9 km 72° NE Ankazobe, 18.22528°S, 47.28683°E, 1410 m, montane rainforest, ex rotten log, collection code BLF03710, 17.— 22.1V.2001 (B.L. Fisher et al.) (CAS: CASENT0437226). Paratypes, 13 pinned work- ers with same data as holotype (BMNH: CASENT0437223; CAS: CASENT0437222; CASENT0437228; CASENT0437229; CASENT0437230; CASENT0437231; CASENT0437232; CASENT0437233; MCZ: CASENT0437225). 112 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Non-type material. MADAGASCAR: no locality (Sikora); Antananarivo, Ré- serve Spéciale dAmbohitantely, Forét d’Ambohitantely, 20.9 km 72° NE d Anka- zobe, 18.22528°S, 47.28683°E, 1410 m, montane rainforest, 17.-22.IV.2001 (BL. Fisher et al.); Antananarivo, Réserve Spéciale d Ambohitantely, Forét d’Ambohitantely, Jardin Botanique, 24.1 km 59° NE d Ankazobe, 18.17139°S, 47.28182°E, 1620 m, montane rainforest, 17.-22.IV.2001 (B.L. Fisher et al.); Antananarivo, Réserve Spéciale d’Ambohitantely, Forét d’ Ambohitantely, 18.18762°S, 47.28576°E, 1580 m, montane forest, 8.[II.2012 (B.L. Fisher et al.); Antananarivo, Réserve Spéciale d Ambohitantely, Porét d’Ambohitantely, 18.22444°S, 47.2774°E, 1490 m, montane forest, 9.III.2012 (B.L. Fisher et al.); Antananarivo, 3 km 41° NE Andranomay, 11.5 km 147° SSE Anjo- zorobe, 18.47333°S, 47.96°E, 1300 m, montane rainforest, 5.—13.XII.2000 (B.L. Fisher et al.); Antananarivo, Stn. Forestiére Manjakatompo, 19.35°S, 47.31667°E, 1600 m, montane rainforest, 20.11.1993 (P.S. Ward); Antananarivo, Réserve Naturelle Sohisika, Sohisika 24.6 km NNE Ankazobe, 18.10322°S, 47.18692°E, 1464 m, gallery montane forest, 1.—2.VI.2008 (B.L. Fisher et al.); Fianarantsoa, 29 km SSW Ambositra, Anka- zomivady, 20.77667°S, 47.165°E, 1700 m, disturbed montane forest, 10.—15.1.1998 (H.G. Robertson); Fianarantsoa, Forét d’Atsirakambiaty, 7.6 km 285° WNW Itremo, 20.59333°S, 46.56333°E, 1550 m, montane rainforest, 22.—26.1.2003 (B.L. Fisher et al). Diagnosis. The following character combination separates T: merina from the remaining species of the 7: schaufussii complex: larger species (HW 0.69-0.79; WL 1.01—1.10); head longer than wide (CI 92-95); eyes relatively large (OI 26-28); fron- tal carinae usually weakly developed, only slightly raised, often not better developed than cephalic rugulae, and usually fading out halfway between posterior eye margin and posterior head margin; propodeal spines reduced to very short triangular teeth/ denticles (PSLI 8-11), propodeal lobes short and triangular, always appearing more voluminous than propodeal teeth, and spines and lobes not strongly inclined towards each other; petiolar node rounded nodiform to rounded high nodiform, in profile around 1.6 to 1.8 times higher than long (LPel 57—64), in dorsal view around 1.2 times wider than long (DPel 117-124); dorsum of promesonotum with at least ten pairs of long, fine, standing hairs, propodeum without standing pilosity, and both waist segments with few long pairs each; body uniformly light to dark brown, gaster often darker, appendages usually lighter. Worker measurements (N=12). HL 0.75—0.84 (0.79); HW 0.69-0.79 (0.73); SL 0.51-0.57 (0.54); EL 0.18—0.21 (0.19); PH 0.37-0.40 (0.39); PW 0.52—0.57 (0.54); WL 1.01-1.10 (1.05); PSL 0.06—0.10 (0.08); PTL 0.16—0.20 (0.18); PTH 0.28—0.31 (0.30); PTW 0.19-0.24 (0.22); PPL 0.21-0.24 (0.22); PPH 0.28-—0.30 (0.29); PPW 0.27-0.31 (0.29); CI 92—95 (93); SI 72-75 (74); OI 26—28 (26); DMI 50-53 (52); LMI 36-38 (37); PSLI 8-11 (10); PeNI 36-43 (40); LPel 57-64 (61); DPel 117-124 (120); PpNI 50-56 (53); LPpI 71-80 (75); DPpI 128-136 (131); PPI 125=142-(133). Worker description. Head longer than wide (CI 92-95); posterior head mar- gin weakly concave. Anterior clypeal margin with distinct median impression. Frontal carinae usually weakly developed, only slightly raised, often not better developed than Taxonomy of Malagasy Tetramorium ila cephalic rugulae, and usually fading out halfway between posterior eye margin and pos- terior head margin. Antennal scrobes present but weak, shallow and without clear and distinct posterior and ventral margins. Antennal scapes short, not reaching posterior head margin (SI 72-75). Eyes relatively large (OI 26-28). Mesosomal outline in pro- file flat to weakly convex, comparatively low and elongate (LMI 36-38), weakly mar- ginate from lateral to dorsal mesosoma; promesonotal suture absent; metanotal groove very weakly developed to absent. Propodeal spines reduced to very short, triangular, and acute or blunt teeth/denticles (PSLI 8-11), propodeal lobes short, triangular, and acute or blunt, always appearing more voluminous than propodeal teeth, spines and lobes not strongly inclined towards each other. Petiolar node in profile nodiform to high nodiform with well-rounded antero- and posterodorsal margins, around 1.6 to 1.8 times higher than long (LPel 57-64), anterior and posterior faces approximately parallel, anterodorsal and posterodorsal margins situated at about same height, petiolar dorsum weakly to moderately convex; petiolar node in dorsal view around 1.2 times wider than long (DPel 117-124), in dorsal view pronotum between 2.3 to 2.7 times wider than petiolar node (PeNI 36-43). Postpetiole in profile globular, approximately 1.3 to 1.4 times higher than long (LPpI 71-80); in dorsal view around 1.3 to 1.4 times wider than long (DPpI 128-136), pronotum between 1.8 to 2.0 times wider than postpetiole (PpNI 50-56). Postpetiole in profile appearing slightly lower or of same height but thicker than petiolar node, postpetiole in dorsal view around 1.3 to 1.4 times wider than petiolar node (PPI 125-142). Mandibles unsculptured, smooth, and shiny; clypeus weakly longitudinally rugulose with two to five lateral rugulae, median area usually completely unsculptured or only weakly sculptured, median ruga usually absent, at most traces present, lateral rugulae often interrupted or irregularly shaped, but at least one lateral rugula well developed on each side; cephalic dorsum between frontal carinae irregularly longitudinally rugulose with eight to ten rugulae; rugulae running from posterior clypeal margin to posterior head margin, often meandering, broken or with cross-meshes; scrobal area mostly unsculptured and laterally merging with surrounding reticulate-rugose sculpture present on lateral head; ground sculpture on head weakly to moderately punctate. Dorsum of mesosoma reticulate-rugose, espe- cially anteriorly, to irregularly longitudinally rugose, lateral mesosoma mostly irregu- larly longitudinally rugose; ground sculpture on mesosoma weak to absent. Forecoxae mainly unsculptured, smooth, and shining. Waist segments and gaster unsculptured, smooth, and shining. Dorsum of head with numerous pairs of long, fine, standing hairs; dorsum of mesosoma with at least eleven or twelve pairs of long, standing hairs ranging from anterior pronotum to posterior mesonotum, propodeum without long, standing pilosity; petiole usually with two pairs and postpetiole with three to four pairs; first gastral tergite with short, scarce, appressed pubescence in combination with abundant, long, standing hairs. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae with decumbent to suberect hairs. Body uniformly light brown to dark brown, gaster often of darker brown, appendages usually lighter. Etymology. The new species is named for the Merina people of Madagascar, in honor of their culture and language. The distribution range of T° merina fits the 114 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) boundaries of the traditional Merina kingdom very well. ‘The species epithet is an arbi- trary combination of letters, thus invariant. Distribution and biology. Tetramorium merina is only known from the Central Highlands of Madagascar (Fig. 65). Its distribution ranges from the two southern- most localities Atsirakambiaty and Ankazomivady through Manjakatompo north to Andranomay, Ambohitantely and Sohisika. All localities are montane rainforests situ- ated at elevations of 1410 to 1700 m, where 7. merina appears to live in leaf litter. Discussion. 7etramorium merina is only moderately distinct within the T° schau- fussii complex, and can be confused with several other species. The possession of large eyes (OI 26-28) separates T° merina from T. pseudogladius (OI 20), and the petiolar node, which in dorsal view is around 1.2 times wider than long (DPel 117-124) dis- tinguishes it from T° nassonowii since the petiolar node of the latter in dorsal view is between 1.0 to 1.2 times longer than wide (DPel 87-98). Tetramorium merina with its very short propodeal teeth (PSLI 8-11) is also not likely to be mistaken for 7! scutum, which possesses much longer propodeal spines (PSLI 22—24) and relatively well-devel- oped propodeal lobes. Also, in 7! scutum the propodeal spines and lobes are strongly inclined towards each other, a condition absent in 7: merina. Tetramorium rala, which is also the smallest species of the complex (HW 0.46-0.49; WL 0.58-0.64), has rela- tively long propodeal spines (PSLI 21-25), and the body is very bright yellowish to light brown, and thus not confusable with 7: merina. In addition, the latter has nu- merous pairs of long, fine, standing hairs on each waist segment distinguishing it from T. sikorae, which lacks any standing pilosity on the waist segments. The species closest to 7: merina are T. monticola, T. obiwan, T: schaufussii, and T. xanthogaster, and the differentiation between these is sometimes challenging. Te- tramorium merina is a relatively large species (HW 0.69-0.79; WL 1.01-1.10) and could be confused with 7: obiwan, which is in the same size range or even larger (HW 0.71-0.84; WL 1.00-1.20), or T° xanthogaster, which is usually smaller but reaches its upper size limit in the range of 7: merina (HW 0.54-0.75; WL 0.72-1.05). However, T. obiwan has longer antennal scapes (SI 77-82), better-developed frontal carinae, and usually no standing pilosity on the waist segments contrasting with the shorter anten- nal scapes (SI 72-75), much weaker frontal carinae and the presence of several pairs of standing pilosity on the waist segments of 7. merina. Furthermore, T: xanthogaster, de- spite being a very variable species throughout its range, can be easily separated from T: merina in the central Highlands where they are usually found occurring in sympatry. In this region 7! xanthogaster is always strongly bicoloured with head and mesosoma very dark brown to black, strongly contrasting with the yellow to light brown waist segments and gaster. Also, 7° xanthogaster usually has long, fine, standing pilosity on the propodeal dorsum while 7: merina lacks any standing pilosity on the propodeum. Few other diagnostic characters separate both species, but the fact that they co-occur in sympatry in several localities without any intermediate forms is a good indication of their heterospecificity. Tetramorium monticola, which is found in the northeast and north of Madagascar, does not co-occur with 7: merina, and both are relatively easy to differentiate. In 7. monticola the frontal carinae are relatively better developed, the Taxonomy of Malagasy Tetramorium Figure 50. 7) merina holotype worker (CASENT0437226). A Body in profile B Body in dorsal view C Head in full-face view. 116 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) cephalic dorsum between them has nine to thirteen relatively regularly shaped, most- ly unbroken rugae, and the propodeum is armed with short to medium-sized spines (PSLI 18-22) while 7° merina has much weaker frontal carinae with eight to ten, of- ten broken and irregular rugulae, and the propodeum is only armed with small teeth/ denticles (PSLI 8-11). The most difficult species to separate from T° merina is certainly T\ schaufussii. As is the case with 7: xanthogaster, T: merina is also found regularly in sympatry with 7: schaufussii, and again, they are separable based on a few characters. Tetramorium me- rina is larger in size (HW 0.69-0.79; WL 1.01-1.10), has a broader head (CI 92-95), and longer (though not significantly so) antennal scapes (SI 72—75) than T° schaufussii (HW 0.51-0.68; WL 0.73—0.93; CI 86-90; SI 66-73). These differences seem dif- ficult to assess without measuring, except for body size, which however, can vary from one population to another and therefore should only be used with caution. Neverthe- less, where the two species are found in sympatry (often they are found within the same litter sample) they can be easily discriminated based on body size. In contrast to T: schaufussii and T. xanthogaster, which are both very variable species, 7. merina is very stable in its morphological appearance. Tetramorium monticola Hita Garcia & Fisher, sp. n. http://zoobank.org/F1CDB914-B817-44BA-8825-013D6BIDFD96 http://species-id.net/wiki/Tetramorium_monticola Figs 42B, 44C, 45C, 47D, 48A, 48B, 51, 65 Type material. Holotype, pinned worker, MADAGASCAR, Antsiranana, Betaolana Forest, along Bekona River, 14.52996°S, 49.44039°E, 880 m, rainforest, ex rotten log, collection code BLF22473, 4.II].2009 (B.L. Fisher et al.) (CAS: CASENT0152401). Paratypes, 1 pinned worker with same data as holotype (CAS: CASENT0152402); three pinned workers with same data as holotype except collection codes BLF22465, BLF22486, and BLF22504 (CAS: CASENT0151864; CASENT0151868; CASENT0152427); and four pinned workers with same data as holotype except col- lection date 5.III.2009 and collection codes BLF22612, BLF22644, and BLF22667 (BMNH: CASENT0151589; CAS: CASENT0151590; CASENT0151949; MCZ: CASENT0152285). Non-type material MADAGASCAR: Antsiranana, Rés. Anjanaharibe-Sud, 9.2 km WSW Befingotra, 14.75°S, 49.46667°E, 1200-1280 m, montane rainfor- est, 6.-10.X1.1994 (B.L. Fisher); Antsiranana, Betaolana Forest, along Bekona River, 14.52996°S, 49.44039°E, 880 m, rainforest, 4.-5.III.2009 (B.L. Fisher et al.); Ant- siranana, Makirovana forest, 14.17066°S, 49.95409°E, 415 m, rainforest, 29.IV.2011 (B.L. Fisher et al.); Antsiranana, Makirovana forest, 14.16666°S, 49.95°E, 715 m, rain- forest, 2.V.2011 (B.L. Fisher et al.); Antsiranana, R.S. Manongarivo, 14.5 km 220° SW Antanambao, 13.99833°S, 48.42833°E, 1175 m, montane rainforest, 20.X.1998 (B.L. Fisher); Antsiranana, RNI Marojejy, 10.5km NW Manantenina, 14.43333°S, Taxonomy of Malagasy Tetramorium inies 49.75°E, 1625 m, montane rainforest, 6.-12.X1.1996 (E.L. Quinter); Antsiranana, Parc National de Marojejy, Manantenina River, 28.0 km 38° NE Andapa, 8.2 km 333° NNW Manantenina, 14.43667, S, 49.775°E, 450 m, rainforest, 12.-15.X1.2003 (B.L. Fisher et al.); Antsiranana, Parc National de Marojejy, Manantenina River, 27.6 km 35° NE Andapa, 9.6 km 327° NNW Manantenina, 14.435°S, 49.76°E, 775 m, rainforest, 17.X1.2003 (B.L. Fisher); Antsiranana, Parc National de Marojejy, An- tranohofa, 26.6 km 31° NNE Andapa, 10.7 km 318° NW Manantenina, 14.44333°S, 49 .74333°E, 1325 m, montane rainforest, 18.-21.X1.2003 (B.L. Fisher); Antsira- nana, Parc National de Marojejy, 25.7 km 32° NNE Andapa, 10.3 km 314° NW Manantenina, 14.445°S, 49.74167°E, 1575 m, montane rainforest, 21.—25.XI.2003 (B.L. Fisher); Antsiranana, Parc National de Marojejy, 25.4 km 30° NNE Andapa, 10.9 km 311° NW Manantenina, 14.445°S, 49.735°E, 2000 m, montane rainforest, 23.X1.2003 (B.L. Fisher); Toamasina, Montagne d’Akirindro 7.6 km 341° NNW Am- binanitelo, 15.28833°S, 49.54833°E, 600 m, rainforest, 17.—21.II1.2003 (B.L. Fisher et al.); Toamasina, 6.9 km NE Ambanizana, Ambohitsitondroina, 15.56667°S, 50°E, 1000 m, montane rainforest, 8.XII.1993 (B.L. Fisher); Toamasina, Ambanizana, Parc National Masoala, 15.57167°S, 50.00611°E, 848-925 m, montane rainforest, 26.II.- 2.11.2003 (D. Andriamalala, D. Silva, et al.); Toamasina, Montagne d’Anjanaharibe, 18 km 21° NNE Ambinanitelo, 15.18833°S, 49.615°E, 470 m, rainforest, 8.—12. III.2003 (B.L. Fisher et al.); Toamasina, Montagne d’Anjanaharibe, 19.5 km 27° NNE Ambinanitelo, 15.17833°S, 49.635°E, 1100 m, montane rainforest, 12.—16.III.2003 (B.L. Fisher et al.); Toamasina, F.C. Sandranantitra, 18.04833°S, 49.09167°E, 450 m, rainforest, 18.—21.1.1999 (HL/. Ratsirarson); Toamasina, Parc National de Zaha- mena, Tetezambatana forest, near junction of Nosivola and Manakambahiny Rivers, 17.74298°S, 48.72936°E, 860 m, rainforest, 18.11.2009 (B.L. Fisher et al.); Toamasina, Parc National de Zahamena, 17.73359°S, 48.72625°E, 950 m, rainforest, 19.II.2009 (B.L. Fisher et al.); Toamasina, Parc National de Zahamena, Onibe River, 17.75908°S, 48.85468°E, 780 m, rainforest, 21.]].2009 (B.L. Fisher et al.). Diagnosis. The following character combination distinguishes 7: monticola from the other species of the T° schaufussii complex: head longer than wide (CI 92-94); eyes relatively large (OI 24-28); antennal scapes very short (SI 67—74); propodeal spines of moderate length, elongate-triangular to spinose, and usually acute (PSLI 8-11), pro- podeal lobes short, triangular, and usually blunt, always much smaller than propodeal spines, spines and lobes not strongly inclined towards each other; petiolar node high nodiform to high cuneiform, in profile around 1.7 to 2.0 times higher than long (LPel 50-60), node in dorsal view around 1.2 to 1.3 times wider than long (DPel 124-133); both waist segments with long, standing pilosity; body uniformly dark yellow to or- ange light brown. Worker measurements (N=12). HL 0.55-0.72 (0.62); HW 0.51—0.67 (0.58); SL 0.34—0.46 (0.40); EL 0.13—0.16 (0.15); PH 0.26—0.33 (0.29); PW 0.35—0.48 (0.41); WL 0.67-0.88 (0.77); PSL 0.10—0.15 (0.12); PTL 0.11-0.15 (0.12); PTH 0.20-0.26 (0.23); PTW 0.13—0.19 (0.16); PPL 0.14—0.18 (0.16); PPH 0.18-0.24 (0.21); PPW 0.19-0.26 (0.23); CI 92-94 (93); SI 67-74 (69); OI 24—28 (25); DMI 118 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) 51-55 (54); LMI 35-39 (37); PSLI 18-22 (20); PeNI 37-42 (39); LPelI 50-60 (54); DPel 124-133 (129); PpNI 54-60 (56); LPpI 71-78 (76); DPpI 131-153 (142); PPI 138-150 (143). Worker description. Head longer than wide (CI 92—94); posterior head margin weakly concave. Anterior clypeal margin with distinct median impression. Frontal cari- nae usually well developed, moderately raised on anterior half, strongly diverging pos- teriorly, and usually approaching or ending at posterior head margin. Antennal scrobes usually weakly developed, shallow and without clear and distinct posterior and ventral margins, sometimes scrobe better developed, slightly deeper and with weak posterior, and rarely even ventral margin. Antennal scapes very short, not reaching posterior head margin (SI 67-74). Eyes relatively large (O] 24-28). Mesosomal outline in profile flat to weakly convex, comparatively low and elongate (LMI 35-39), weakly margin- ate from lateral to dorsal mesosoma; promesonotal suture absent; metanotal groove weakly to moderately developed, sometimes slightly impressed, but mostly moderately deep. Propodeal spines of moderate length, elongate-triangular to spinose, and usually acute (PSLI 8-11), propodeal lobes short, triangular, and usually blunt, always much smaller than propodeal spines, spines and lobes not strongly inclined towards each oth- er. Petiolar node in profile high nodiform to high cuneiform, around 1.7 to 2.0 times higher than long (LPeI 50-60), anterior and posterior faces approximately parallel, an- terodorsal margin always higher and more angled than posterodorsal margin, petiolar dorsum usually weakly convex and tapering backwards posteriorly; petiolar node in dorsal view around 1.2 to 1.3 times wider than long (DPel 124-133), in dorsal view pronotum between 2.4 to 2.7 times wider than petiolar node (PeNI 37-42). Postpeti- ole in profile globular, approximately 1.3 to 1.4 times higher than long (LPpI 71-78); in dorsal view around 1.3 to 1.5 times wider than long (DPpI 131-153), pronotum between 1.7 to 1.9 times wider than postpetiole (PpNI 54-60). Postpetiole in profile appearing always lower and having less volume than petiolar node, postpetiole in dor- sal view around 1.4 to 1.5 times wider than petiolar node (PPI 138-150). Mandibles unsculptured, smooth, and shiny; clypeus usually longitudinally rugulose with two to six rugulae, median area usually either completely unsculptured without median rugula or only weakly sculptured with traces of median rugula, very rarely median ru- gula fully developed, lateral rugulae usually well developed and unbroken, sometimes irregularly shaped or broken; cephalic dorsum between frontal carinae longitudinally rugose with nine to thirteen rugae; rugae running from posterior clypeal margin to posterior head margin, generally very regularly shaped and only rarely broken or with cross-meshes; scrobal area partly unsculptured and laterally merging with surrounding longitudinally rugulose to reticulate-rugose sculpture present on lateral head; ground sculpture on head usually moderately punctate. Dorsum of mesosoma usually longi- tudinally rugose, sometimes irregularly so; lateral mesosoma irregularly longitudinally rugose to reticulate-rugose, sometimes lateral pronotum with less sculpture; ground sculpture on mesosoma weak to absent. Forecoxae always unsculptured, smooth and shining. Waist segments and gaster unsculptured, smooth, and shining. Dorsum of head with numerous pairs of long, fine, standing hairs; dorsum of promesonotum with Taxonomy of Malagasy Tetramorium 119 at least ten pairs of long, standing hairs ranging from anterior pronotum to posterior mesonotum, propodeum usually without long, standing pilosity, sometimes with one or two shorter pairs of hairs; petiole usually with at least two pairs and postpetiole with at least three to four pairs; first gastral tergite with short, scarce to abundant, decum- bent to appressed pubescence in combination with abundant, long, standing hairs. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae usually with decumbent to suberect hairs. Body uniformly dark yellow to orange light brown, gaster sometimes of darker brown, appendages usually slightly lighter. Etymology. The name of the new species is a Latin noun and means “inhabitant of the mountains” referring to the fact that 7: monticola is predominantly found in higher elevation montane forests. The species epithet is a nominative noun, and thus invariant. Distribution and biology. Tetramorium monticola is distributed in the northeast and north of Madagascar (Fig. 65). The distribution range is outlined by Manongarivo in the west, the northernmost locality Makirovana, and the easternmost Ambanizana on the Masoala Peninsula. Most of the material available was collected in that triangle, but 7: monticola is also known from Zahamena and Sandranantitra, which are located much further south. All localities are rainforests or montane rainforests situated at el- evation ranging from 415 to 2000 m. Also, it seems that 7° monticola lives in leaf litter. Discussion. Tetramorium monticola is well recognisable within the 7! schaufussii complex. It differs from 7! pseudogladius (Ol 20) by its much larger eye size (OI 24— 28), and from T° scutum by having propodeal spines and lobes not strongly inclined towards each other. Also, 7: monticola (DPel 124-133; LPel 50-GO) is very unlikely to be confused with 7. nassonowii, which has a much longer and lower petiolar node (DPel 87-98; LPel 72-81). In addition, 7. monticola is a relatively hairy species with long pilosity all over the body, which separates it from the few species with partly reduced pilosity, such as T: rala, T! sikorae, and T. obiwan, which all usually lack standing pilosity on the propodeum and waist segments, except in 7: obiwan, where the petiole or postpetiole occasionally have a few long hairs. Nevertheless, 7. monticola cannot be mistaken for 7: obiwan. The latter is of larger body size (HW 0.71—0.84; WL 1.00-1.20), has longer antennal scapes (SI 77-82), and has a petiolar node with the antero- and posterodorsal margins situated at about the same height and well rounded. Tetramorium monticola is much smaller than that (HW 0.51—0.67; WL 0.67—0.88), has shorter antennal scapes (SI 67—74), and its petiolar node has the anterodorsal mar- gin always higher and a little bit more angled than the posterodorsal margin with the petiolar dorsum tapering backwards. The remaining three species, 7: merina, T. schaufussii, and T. xanthogaster, are morphologically closer to T° monticola, and their separation requires more attention. Tetramorium schaufussii can be easily mistaken for 7. monticola in northern Mada- gascar where both are sympatric, but 7° schaufussii differs from T. monticola (and the other two) by having a much longer and thinner head in full-face view (CI 86-90). The other two species, 7: merina and T. xanthogaster, usually have shorter, and in the case of 7. merina much shorter, propodeal spines (PSLI 8-16) than 7. monticola (PSLI 18-22). However, since the spine length is somewhat variable in the latter species, 120 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Figure 51. 7: monticola holotype worker (CASENT0152401). A Body in profile B Body in dorsal view C Head in full-face view. Taxonomy of Malagasy Tetramorium 12a this character cannot be the sole distinguishing feature. In addition, 7: monticola also differs from the other two by having relatively better developed frontal carinae and the cephalic dorsum between them with nine to thirteen relatively regularly shaped, mostly unbroken rugae. Tetramorium merina and T. xanthogaster have relatively weak- er developed, and often shorter, frontal carinae and a cephalic dorsum with six to ten relatively irregularly shaped, often meandering or broken rugulae. Tetramorium nassonowii Forel, 1892, stat. n. Figs 40A, 52, 65 Tetramorium nassonowii Forel, 1892:521. [Synonymy with 7. schaufussii by Bolton, 1979 1:37] Type material. Lectotype [designated here], pinned worker, MADAGASCAR, An- tananarivo (Province central de Madagascar), Andrangoloaca, 47.27729°E, 18.22198 S (Sikora) (MHNG: CASENT0101289) [examined]. Paralectotype [designated here], pinned worker with same data as lectotype (MHNG: CASENT0101290). [Note 1: the GPS data of the type locality was not provided by the locality label or the original description. The data presented above is based on our own geo-referencing of the Foret d’ Andrangoloaca and should be considered as an approximation and not the exact position of the type locality.] [Note 2: one specimen from the MHNG collection (CASENT0101556) with the label data “Tetramorium nassonowii Forel var., no. 14, Moramanga (Sikora)” also has a red type label. However, we do not consider this as a real name-bearing type speci- men since the original description of Forel (1892) only mentioned the type material from Andrangoloaca and nothing from Moramanga. ‘This specimen from Moramanga (which is also partly broken) is actually not conspecific with T° nassonowii, but belongs to the newly described species 7! obiwan.| Non-type material. MADAGASCAR: Antananarivo, 3 km 41° NE Andranomay, 11.5 km 147° SSE Anjozorobe, 18.47333°S, 47.96°E, 1300 m, montane rainforest, 5.-13.X11.2003 (B.L. Fisher et al); Antsiranana, Rés. Anjanaharibe-Sud, 9.2 km WSW Befingotra, 14.75°S, 49.46667°E, 1260 m, 11.X1.1994 (B.L. Fisher); Antsiranana, Rés. Anjanaharibe-Sud, 11.0 km WSW Befingotra, 14.75°S, 49.45°E, 1565 m, mon- tane rainforest, 16.XI.1994 (BL. Fisher); Fianarantsoa, 29 km SSW Ambositra, An- kazomivady, 20.77667°S, 47.165°E, 1700 m, disturbed montane rainforest, 7.1.1998 (B.L. Fisher); Fianarantsoa, Ranomafana National Park, 21.21667°S, 47.41667°E, 706 m, montane rainforest, 1.X.1992 (E. Rajeriarison); Toamasina, 5.3 km SSE Am- banizana, Andranobe, 15.67133°S, 49.97395°E, 425 m, rainforest, 21.XI.1993 (B.L. Fisher); Toamasina, F.C. Andriantantely, 18.695°S, 48.81333°E, 530 m, rainforest, 4—10.X11.1998 (HJ. Ratsirarson); Toamasina, Montagne d’Anjanaharibe, 19.5 km 27° NNE Ambinanitelo, 15.17833°S, 49.635°E, 1100 m, montane rainforest, 12.-16. [1.2003 (B.L. Fisher et al.); Toamasina, F.C. Didy, 18.19833°S, 48.57833°E, 960 m, 122 Francisco Hita Garcia & Brian L. Fisher | ZooKeys 413: 1-170 (2014) rainforest, 16.—23.XI1.1998 (4./. Ratsirarson); Toliara, Parc National d’Andohahela, Manampanihy River, 5.4 km 113° ESE Mahamavo, 36.7 km 343° NNW Tolagnaro, 24.76389°S, 46.76683°E, 650 m, rainforest, 24.1.2002 (B.L. Fisher et al.); Toliara, Réserve Spéciale Kalambatritra, Befarara, 23.4178°S, 46.4478°E, 1390 m, montane rainforest, 7.11.2009 (B.L. Fisher et al.); Toliara, Réserve Spéciale Kalambatritra, Befa- rara, 23.4144°S, 46.459°E, 1360 m, montane rainforest, 8.11.2009 (B.L. Fisher et al.); Toliara, Réserve Spéciale Kalambatritra, Befarara, 23.4502°S, 46.45658°E, 1325 m, montane rainforest, 11.11.2009 (B.L. Fisher et al.). Diagnosis. Tetramorium nassonowii is clearly recognisable within the 7! schaufussii complex on the basis of its petiolar node shape, which is relatively long and low, in profile around 1.2 to 1.4 times higher than long (LPel 72—81) and in dorsal view be- tween 1.0 to 1.2 times longer than wide (DPel 87-98). Worker measurements (N=12). HL 0.82—0.96 (0.87); HW 0.74—0.86 (0.78); SL 0.52-0.64 (0.57); EL 0.18-0.21 (0.19); PH 0.38—0.45 (0.41); PW 0.54—0.62 (0.57); WL 1.02-1.16 (1.08); PSL 0.06—0.09 (0.08); PTL 0.21—0.26 (0.23); PTH 0.28-0.34 (0.30); PTW 0.20—0.25 (0.22); PPL 0.21—0.26 (0.23); PPH 0.28—0.34 (0.30); PPW 0.27—0.34 (0.30); CI 90-92 (91); SI 70-74 (72); OI 23-25 (24); DMI 51-54 (53); LMI 36-39 (38); PSLI 7-11 (9); PeNI 36-40 (38); LPel 72-81 (77); DPel 87-98 (95); PpNI 50-55 (52); LPpI 75-82 (78); DPpI 122-134 (128); PPI 130-145 (137). Worker description. Head clearly longer than wide (CI 90-92); posterior head margin weakly concave. Anterior clypeal margin with distinct median impression. Frontal carinae weakly to moderately developed, moderately raised, diverging posteri- orly, and usually fading out halfway between posterior eye margin and posterior head margin or approaching posterior head margin. Antennal scrobes present but weak, shallow and without clear and distinct posterior and ventral margins. Antennal scapes short, not reaching posterior head margin (SI 70-74). Eyes moderate to large (Ol 23-25). Mesosomal outline in profile flat to weakly convex, comparatively low and long (LMI 36-39), weakly to moderately marginate from lateral to dorsal mesosoma; promesonotal suture absent; metanotal groove weakly developed or absent. Propodeal spines reduced to very short teeth (PSLI 7-11), propodeal lobes short, triangular, and blunt or acute, usually longer than propodeal spines, rarely as long as propodeal spines, spines and lobes not strongly inclined towards each other. Petiolar node rounded nodi- form, in profile around 1.2 to 1.4 times higher than long (LPel 72-81), anterior and posterior faces not parallel, anterodorsal and posterodorsal margins situated at about same height, petiolar dorsum distinctly convex; node in dorsal view weakly longer than wide (DPel 92-96), in dorsal view pronotum between 2.5 to 2.8 times wider than petiolar node (PeNI 36-40). Postpetiole in profile globular, approximately 1.2 to 1.3 times higher than long (LPpI 75-82); in dorsal view around 1.2 to 1.3 times wider than long (DPpI 122-134), pronotum between 1.8 to 2.0 times wider than postpetiole (PpNI 50-55). Postpetiole in profile appearing more or less of same vol- ume as petiolar node, postpetiole in dorsal view around 1.3 to 1.5 times wider than petiolar node (PPI 130-145). Mandibles unsculptured, smooth, and shiny; clypeus weakly longitudinally rugulose with three to seven rugulae, rugulae often interrupted Taxonomy of Malagasy Tetramorium 123 or irregularly shaped, median area often weakly sculptured, median ruga usually absent or mostly reduced, very rarely fully developed; cephalic dorsum between frontal cari- nae irregularly longitudinally rugose/rugulose with six to nine rugae/rugulae; rugae/ rugulae running from posterior clypeal margin to posterior head margin, often me- andering, broken or with cross-meshes; scrobal area mostly unsculptured and laterally merging with surrounding reticulate-rugose to longitudinally rugose sculpture present on lateral head; ground sculpture on head weak to absent. Dorsum of mesosoma ir- regularly longitudinally rugose to reticulate-rugose, lateral mesosoma mostly irregu- larly longitudinally rugose; ground sculpture on mesosoma weak to absent. Forecoxae mainly unsculptured, smooth and shining. Waist segments and gaster unsculptured, smooth, and shining. Dorsum of head with several pairs of long, fine, standing hairs; dorsum of mesosoma with at least six or seven pairs of long, standing hairs ranging from anterior pronotum to posterior mesonotum, propodeum without long, standing pilosity; petiole with one pair and postpetiole with one or two pairs; first gastral tergite with short, scarce, appressed pubescence in combination with scattered, long, standing hairs. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae with appressed to decumbent hairs. Body uniformly light brown to dark brown colour, ap- pendages often lighter. Distribution and biology. Tetramorium nassonowii is distributed in the montane rainforests and rainforest belt of eastern Madagascar (Fig. 65) at altitudes ranging from 425 to 1700 m, although it is predominantly found in montane rainforests situated higher than 1000 m. Also, based on the available collection data, it seems that T. nas- sonowii inhabits leaf litter or the ground. Discussion. In this study we propose to raise 7. nassonowii, which was first de- scribed by Forel (1892), to species rank. In the original description Forel (1892) com- pared 7! nassonowii with T. schaufussii, and found them to be different in petiolar node shape and propodeal spine length. Much later, in his revision of the Malagasy Tetramorium, Bolton (1979) synonymised T: nassonowii under T. schaufussii, likely because the limited material available for both species suggested their conspecificity. Indeed, the only genuine specimens of 7. nassonowii available to Forel and Bolton were the two syntypes from MHNG. Based on the examination of a few thousand specimens from the whole 7° schaufussii complex, we believe the material listed as T. schaufussii by Bolton (1979) to consist of the species T: merina, T. nassonowii, T. obiwan, and T. schaufussi. Tetramorium nassonowii is especially distinctive within the complex due to its large body size and characteristic petiolar node shape. As noted in the diagnosis, the lower and longer petiolar node shape of T° nassonowii, which in profile is around 1.2 to 1.4 times higher than long (LPel 72-81) and in dorsal view between 1.0 to 1.2 times longer than wide (DPel 87—98), clearly distinguishes it from the remainder of the T° schaufussii species complex. The other species all have a higher and broader petiolar node, in profile around 1.5 to 2.2 times higher than long (LPel 45-67) and in dorsal view between 1.1 to 1.5 times wider than long (DPel 109-154). Despite the relatively wide distribution, we find 7) nassonowii without much ob- servable intraspecific variation. 124 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Figure 52. 7° nassonowii syntype worker (CASENT0101289). A Body in profile B Body in dorsal view C Head in full-face view. Taxonomy of Malagasy Tetramorium 125 Tetramorium obiwan Hita Garcia & Fisher, sp. n. http://zoobank.org/8EE7AACC-8 18A-4B99-AB95-75CD 1969ECA7 http://species-id.net/wiki/Tetramorium_obiwan Figs 20D, 43B, 44A, 53, 65 Type material. Holotype, pinned worker, MADAGASCAR, Toliara, Parc Na- tional d’Andohahela, Col du Sedro, 3.8 km 113° ESE Mahamavo, 37.6 km 341° NNW Tolagnaro, 24.76389°S, 46.75167°E, 900 m, montane rainforest, beating low vegetation, collection code BLF05014, 21.—25.1.2002 (B.L. Fisher et al.) (CAS: CASENT0447245). Paratypes, three pinned workers with same data as holotype (CAS: CASENT0447199; CASENT0447207; MCZ: CASENT0447237); eight pinned workers with same data as holotype except sampled from canopy moss and leaf litter and collection codes BLF05036 and BLF05136 (CAS: CASENT0451274; CASENT045 1275; CASENT045 1276; CASENT0455961); four pinned workers with same data as holotype except sampled from sifted litter and collection code BLF05010 (BMNH: CASENT0484542; CAS: CASENT0484425; CASENT0484536; CASENT0484554). Non-type material. MADAGASCAR: Antananarivo, 3 km 41° NE Andranomay, 11.5 km 147° SSE Anjozorobe, 18.47333°S, 47.96°E, 1300 m, montane rainforest, 5.-13.X11.2000 (B.L. Fisher et al.); Antsiranana, Rés. Anjanaharibe-Sud, 9.2 km WSW Befingotra, 14.75°S, 49.46667°E, 1260 m, montane rainforest, 11.X1.1994 (B.L. Fish- er); Antsiranana, Betaolana Forest, along Bekona River, 14.52996°S, 49.44039°E, 880 m, rainforest, 4.[11.2009 (B.L. Fisher et al.); Fianarantsoa, Rés. Andringitra, 40 km S Ambalavao, 22.21667°S, 46.96667°E, 1225 m, montane rainforest, 19.X.1993 (B.L. Fisher); Fianarantsoa, Parc National Befotaka-Midongy, Papango 28.5 km S Midongy- Sud, Mount Papango, 23.84083°S, 46.9575°E, 1250 m, montane rainforest, 17-18. XI.2006 (B.L. Fisher et al.); Fianarantsoa, Ranomafana National Park, radio tower, 21.25083°S, 47.40717°E, 1130 m, forest edge, mixed tropical forest, open area, 4.— 21.1.2002 (VM. Irwin & R. Harin’Hala); Fianarantsoa, Parc National de Ranomafana, Vatoharanana River, 4.1 km 231° SW Ranomafana, 21.29°S, 47.43333°E, 1100 m, montane rainforest, 27.—31.II1.2003 (B.L. Fisher et al.); Fianarantsoa, Parc National de Ranomafana, Sahamalaotra River, 6.6 km 310° NW Ranomafana, 21.23667°S, 47 .39667°E, 1150 m, montane rainforest, 31.11.2003 (B.L. Fisher et al.); Toamasina, 6 km ESE Andasibe (=Perinet), 18.95°S, 48.46667°E, 900 m, rainforest, 17.XI.1990 (P.S.. Ward); Toamasina, Corridor Forestier Analamay-Mantadia, Ambatohara- nana, 18.80388°S, 48.40506°E, 1013 m, rainforest, 12.-19.2012 (B.L. Fisher et al.); Toamasina, Corridor Forestier Analamay-Mantadia, Ambatoharanana, 18.79956°S, 48.4028°E, 1058 m, rainforest, 12.-19.2012 (BL. Fisher et al.); Toamasina, Corri- dor Forestier Analamay-Mantadia, Ambatoharanana, 18.80398°S, 48.40358°E, 1064 m, rainforest, 12.-19.2012 (BL. Fisher et al.); Toamasina, Ankerana, 18.4017°S, 48.80605°E, 1035 m, montane forest, 26.1.2012 (B.L. Fisher et al.); Toliara, Parc Na- tional d’Andohahela, Col du Sedro, 3.8 km 113° ESE Mahamavo, 37.6 km 341° NNW 126 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Tolagnaro, 24.76389°S, 46.75167°E, 900 m, montane rainforest, 21.—25.1.2002 (B.L. Fisher et al.); Toliara, Forét Ivohibe 55.6 km N Tolagnaro, 24.56167°S, 47.20017°E, 650 m, rainforest, 4.X1I.2006 (B.L. Fisher et al.). Diagnosis. The following character combination distinguishes 7: obiwan from the other species of the 7° schaufussii complex: relatively larger species (HW 0.71-0.84; WL 1.00—1.20); frontal carinae moderately to very well developed, diverging posteriorly, becoming weaker halfway between posterior eye margin and posterior head margin, and ending at or shortly before posterior head margin; antennal scapes short to moderate (SI 77-82); eyes relatively large (OI 25—27); propodeal spines short, triangular to elon- gate-triangular, and acute (PSLI 10-16), propodeal lobes short, triangular, and blunt or acute, always much shorter than propodeal spines, spines and lobes never strongly inclined towards each other; petiolar node high rounded nodiform, in profile around 1.6 to 1.9 times higher than long (LPel 54-64), and in dorsal view around 1.1 to 1.3 times wider than long (DPel 109-129); waist segments usually without any standing hairs, sometimes one pair of long, standing hairs present on petiole and/or postpetiole. Worker measurements (N=15). HL 0.78—-0.93 (0.86); HW 0.71-0.84 (0.78); SL 0.56-0.69 (0.62); EL 0.18—0.22 (0.20); PH 0.36—0.44 (0.41); PW 0.50-0.60 (0.57); WL 1.00-1.20 (1.10); PSL 0.08-0.14 (0.12); PTL 0.15—-0.20 (0.18); PTH 0.27-0.33 (0.30); PIW 0.18—-0.23 (0.21); PPL 0.19-0.24 (0.23); PPH 0.27-0.32 (0.31); PPW 0.25—0.32 (0.29); CI 89-92 (90); SI 77-82 (80); OI 25-27 (26); DMI 50-53 (51); LMI 36-38 (37); PSLI 10-16 (14); PeNI 35-39 (36); LPel 54-64 (58); DPel 109-129 (117); PpNI 50-53 (51); LPpI 70-77 (74); DPpI 124-135 (129); PPI 135-148 (142). Worker description. Head clearly longer than wide (89-92); posterior head mar- gin weakly concave. Anterior clypeal margin with distinct median impression. Frontal carinae moderately to very well developed, diverging posteriorly, becoming weaker half- way between posterior eye margin and posterior head margin, and ending at or shortly before posterior head margin. Antennal scrobes very weak to absent, shallow, and with- out any posterior and ventral margins. Antennal scapes short to moderate, not reaching posterior head margin (SI 77-82). Eyes relatively large (OI 25-27). Mesosomal outline in profile relatively flat to weakly convex, comparatively low and elongate (LMI 36-38), weakly to moderately marginate from lateral to dorsal mesosoma; promesonotal suture absent; metanotal groove either weakly developed or absent. Propodeal spines short, triangular to elongate-triangular, and acute (PSLI 10-16), propodeal lobes short, tri- angular, and blunt or acute, always much shorter than propodeal spines, spines and lobes not strongly inclined towards each other. Petiolar node in profile high rounded nodiform, around 1.6 to 1.9 times higher than long (LPel 54-64), anterior and pos- terior faces approximately parallel, anterodorsal and posterodorsal margins situated at about same height and well rounded, petiolar dorsum weakly to moderately convex; node in dorsal view around 1.1 to 1.3 times wider than long (DPel 109-129); in dorsal view pronotum around 2.6 to 2.9 times wider than petiolar node (PeNI 35-39). Post- petiole in profile globular, around 1.3 to 1.4 times higher than long (LPpI 70-77); in dorsal view around 1.2 to 1.4 times wider than long (DPpI 124-135); in dorsal view Taxonomy of Malagasy Tetramorium 127 pronotum around 1.9 to 2.0 times wider than postpetiole (PpNI 50-53). Postpetiole in profile appearing more or less of same volume as petiolar node, postpetiole in dorsal view between 1.3 to 1.5 times wider than petiolar node (PPI 135-148). Mandibles unsculptured, smooth, and shiny; clypeus weakly longitudinally rugulose with three to five, rarely more, rugulae, rugulae often interrupted or irregularly shaped, median area usually weakly sculptured without median rugula, median rugula sometimes partly and rarely fully developed; cephalic dorsum between frontal carinae longitudinally rugose/ rugulose with seven to ten rugae/rugulae; rugae/rugulae running from posterior clypeal margin to posterior head margin, often interrupted, or with cross-meshes; scrobal area mostly unsculptured and laterally merging with surrounding reticulate-rugose to longi- tudinally rugose sculpture present on lateral head, sometimes head posterolaterally with very little sculpture, very smooth, and shining. Dorsum of mesosoma irregularly lon- gitudinally rugose to reticulate-rugose, anterior pronotum especially reticulate-rugose; lateral mesosoma mostly irregularly longitudinally rugose with few unsculptured areas on lateral pronotum and/or katepisternum. Forecoxae mainly unsculptured, smooth, and shining. Waist segments and gaster unsculptured, smooth, and shining. Ground sculpture everywhere on body weak to absent. Dorsum of head with numerous pairs of long, fine, standing hairs; dorsum of mesosoma with seven to ten pairs, hairs present from anterior pronotum to posterior mesonotum, propodeum without standing pilos- ity; usually waist segments without any standing hairs, sometimes one pair of long, standing hairs present on petiole and/or postpetiole; first gastral tergite with very short, scarce, appressed pubescence in combination with few scarce, long, standing hairs. An- terior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae usually with ap- pressed, rarely decumbent hairs. Head, mesosoma, waist segments and gaster uniformly light reddish, orange-brown to dark brown contrasting with lighter yellowish to light brown mandibles, antennae, and legs. Etymology. The name of the new species is inspired by a fictional character from George Lucas’ “Star Wars” Universe: the wise Jedi master Obi-Wan Kenobi. ‘The spe- cies epithet is an arbitrary combination of letters, thus invariant. Distribution and biology. Tetramorium obiwan is widely but relatively patchily distributed throughout the rainforests and montane rainforests of eastern Madagascar (Fig. 65). Its known distribution encompasses nine localities, ranging from Ando- hahela in the southeast to Anjanaharibe-Sud in the northeast. A likely explanation for this patchy distribution record may be that 7. obiwan lives in the vegetation and is consequently less often sampled by ground or leaf litter methods. ‘The available collec- tion data supports this, as 7: obiwan was mainly collected by beating low vegetation or Malaise traps, and rarely from pitfall traps or leaf litter. If true, then 7. obiwan is possibly much more widespread than currently known, and more collecting in lower vegetation will likely provide additional material of this species. Furthermore, T° obi- wan seems to prefer higher elevations. It can be found from 675 to 1300 m, but most collections were from the upper altitudinal range limit. Discussion. Tetramorium obiwan is a fairly conspicuous member of the 7. schau- Jussii complex, thus easily separable from the other species. As mentioned in the diag- 128 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Figure 53. 7. obiwan holotype worker (CASENT0447245). A Body in profile B Body in dorsal view C Head in full-face view. Taxonomy of Malagasy Tetramorium 129 nosis above, its larger body size (HW 0.71-0.84; WL 1.00-—1.20), strongly developed frontal carinae, relatively long antennal scapes (SI 77-82), relatively large eyes (Ol 25-27), lack of median clypeal ruga, and (usually) lack of long, standing pilosity on the waist segments will separate it from the other species of the complex. Except for the relatively long antennal scapes, none of the characters listed is unique to T° obiwan, but the combination renders its identification very straightforward. Still, in a few cases it is possible to mistake 7’ obiwan at first glance for T. merina or T. nassonowii, which are also larger species. Nevertheless, 7: merina has noticeably much weaker frontal carinae than T: obiwan. Tetramorium nassonowii is likely the species closest to T. obiwan but they differ significantly in the shape of the petiolar node. The node of T. obiwan is higher and broader, in profile around 1.6 to 1.9 times higher than long (LPel 54-64), and in dorsal view around 1.1 to 1.3 times wider than long (DPel 109-129), whereas the node of 7! nassonowii is relatively long and low, in profile around 1.2 to 1.4 times higher than long (LPel 72-81), and in dorsal view between 1.0 to 1.2 times longer than wide (DPel 87-98). Despite being that distinctive in its own species complex, T: obiwan is comparatively similar to 7: proximum from the T. cognatum species complex in that they are both large, reddish-brown species with relatively well-developed fron- tal carinae and a high rounded nodiform petiolar node. Nevertheless, the presence of long, standing pilosity on the first gastral tergite easily distinguishes T° obiwan from T: proximum that lacks any standing pilosity on the first gastral tergite. Intraspecific variation in 7° obiwan seems relatively low, especially considering that the species is fairly widely distributed. Tetramorium pseudogladius Hita Garcia & Fisher, sp. n. http://zoobank.org/4C29B150-F567-4F4D-ABOF-062EA2D2EDCD http://species-id.net/wiki/Tetramorium_pseudogladius Figs 41A, 54, 65 Type material. Holotype, pinned worker, MADAGASCAR, Toamasina, Parc Na- tional de Zahamena, Tetezambatana forest, near junction of Nosivola and Manakam- bahiny Rivers, 17.74298°S, 48.72936°E, 860 m, rainforest, sifted litter (leaf mold, rotten wood), collection code BLF21974, 18.—19.II.2009 (B.L. Fisher et al.) (CAS: CASENT0153605). Diagnosis. Tetramorium pseudogladius can be easily separated from the remainder of the species complex by its relatively small eyes (OI 20). Worker measurements (N=1). HL 0.74; HW 0.68; SL 0.55; EL 0.14; PH 0.36; PW 0.533 WL O96? PSI 05 GP TE0:14;, PTH 0:27- P EW 20219:8PPL0:21;; PRP 0.28; PPW 0.27; CI 91; SI 80; OI 20; DMI 55; LMI 38; PSLI 22; PeNI 35; LPel 52; DPel132; PoNi5i;-L Ppl -75;-DPpl 129;-PPl 146. Worker description. Head longer than wide (CI 91); in full-face view posterior head margin weakly to moderately concave. Anterior clypeal margin with distinct median im- 130 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) pression. Frontal carinae well developed, diverging posteriorly, approaching posterolateral corners of head. Antennal scrobes very weak, shallow and without clear and distinct poste- rior and ventral margins. Antennal scapes of moderate length, not reaching posterior head margin (SI 80). Eyes relatively small (OI 20). Mesosomal outline in profile relatively flat, moderately low and long (LMI 38), moderately marginate from lateral to dorsal meso- soma; promesonotal suture absent; metanotal groove mostly reduced. Propodeal spines moderately long, elongate-triangular to spinose, acute (PSLI 22), propodeal lobes short and triangular, much shorter than propodeal spines, spines and lobes not strongly inclined towards each other. Petiolar node in profile high rounded nodiform, around 1.9 times higher than long (LPel 52), anterior and posterior faces approximately parallel, anterodor- sal and posterodorsal margins situated at about same height and weakly rounded, petiolar dorsum weakly convex; node in dorsal view around 1.3 times wider than long (DPel 132); in dorsal view pronotum around 2.8 to 2.9 times wider than petiolar node (PeNI 35). Postpetiole in profile globular, around 1.3 times higher than long (LP pI 75); in dorsal view around 1.3 times wider than long (DPpI 129); in dorsal view pronotum around 1.9 to 2.0 times wider than postpetiole (PpNI 51). Postpetiole in profile appearing more or less of same volume as petiolar node, postpetiole in dorsal view around 1.4 to 1.5 times wider than petiolar node (PPI 146). Mandibles completely unsculptured, smooth, and shiny; clypeus longitudinally rugulose with three similarly weak but unbroken rugulae; cephalic dorsum between frontal carinae longitudinally rugose with six or seven rugae, rugae run- ning mostly unbroken from posterior clypeal margin to posterior head margin, a few rugae interrupted or with cross-meshes; scrobal area partly unsculptured, but mostly merging with surrounding reticulate-rugose to longitudinally rugose sculpture present on lateral head; ground sculpture on head weakly to moderately punctate. Dorsum and sides of mes- osoma mostly irregularly longitudinally rugose; forecoxae mostly unsculptured, smooth, and shining; ground sculpture on mesosoma very weak to absent. Both waist segments and gaster completely unsculptured, smooth, and shining. Dorsum of head with several pairs of long, fine, standing hairs; dorsum of mesosoma with six pairs restricted to pronotum and mesonotum, propodeum without standing pilosity; waist segments and first gastral tergite without any standing hairs; first gastral tergite with very short, scarce, appressed pu- bescence in combination with a few scarce, long, standing hairs. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae with appressed to decumbent hairs. Head, mesosoma, waist segments and gaster uniformly reddish, orange-brown contrasting with lighter yellowish to light brown mandibles, antennae, and legs. Etymology. The name of the new species is a combination of the ancient Greek word “pseudes”, which means “false” or “lying”, and the species name of T° gladius from the 7. cognatum complex. This combined name takes account for the fact that both species are almost identical in morphology. The species epithet is treated as a nominative noun, and is thus invariant. Distribution and biology. At present, 7. pseudogladius is known only from the type locality, Parc National de Zahamena (Fig. 65) where it was collected in lowland rainfor- est at an altitude of 860 m. In addition, the new species was sampled from leaf litter. Taxonomy of Malagasy Tetramorium feel Figure 54. 7: pseudogladius holotype worker (CASENT0153605). A Body in profile B Body in dorsal view C Head in full-face view. 132 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Discussion. Like 7: gladius in the T. cognatum complex, T: pseudogladius is also immediately recognisable on the basis of its much smaller eyes (OI 20). The other spe- cies of the T: schaufussii complex all have much larger eyes (OI 22—28). In addition, T. pseudogladius lacks the long, standing pilosity on the waist segments present in most other members of the species complex, and has relatively long antennal scapes (SI 80). Generally, 7. pseudogladius looks very similar to T. gladius in the T. cognatum spe- cies complex and they also share most of their morphometric range. Indeed, if not for the few long, standing hairs on its first gastral tergite, the holotype of T: pseudogladius could be easily confused with 7! gladius. They also differ in antennal scape length, however, which is longer in 7° pseudogladius (SI 80) than in T: gladius (SI 71-74). Until more material from the type locality becomes available, we consider these differ- ences as diagnostic to delimit the species boundary between these two species. Tetramorium rala Hita Garcia & Fisher, sp. n. http://zoobank.org/60FD 1630-677 1-49 E0-8C47-4FFCE28FA02C http://species-id.net/wiki/Tetramorium_trala Figs 43A, 55, 65 Type material. Holotype, pinned worker, MADAGASCAR, Antsiranana, Parc National de Marojejy, Manantenina River, 28.0 km 38° NE Andapa, 8.2 km 333° NNW Manantenina, 14.43667°S, 49.775°E, 450 m, rainforest, sifted litter (leaf mold, rotten wood), collection code BLF08722, 12.-15.X1.2003 (B.L. Fisher et al.) (CAS: CASENT0046163). Paratypes, five pinned workers with same data as holo- type (BMNH: CASENT0046035; CAS: CASENT0045975; CASENT0046039; CASENT0046049; MCZ: CASENT0046176); and nine pinned workers with same data as holotype except sampled from rotten log and collection code BLF08772 (CAS: CASENT0077304; CASENT0077305; CASENT0077306). Non-type material. MADAGASCAR: Antsiranana, Cap Masoala, 19 km W Andampibe, 15.69361°S, 50.18167°E, 125 m, lowland rainforest, 2.XII.1992 (G.A. Alpert); Antsiranana, Makirovana forest, 14.17066°S, 49.95409°E, 415 m, rainfor- est, 28.-29.1V.2011 (B.L. Fisher et al.); Antsiranana, Makirovana forest, 14.16044°S, 49.95216°E, 550 m, rainforest, 1.V.2011 (B.L. Fisher et al.); Antsiranana, Parc Na- tional de Marojejy, Manantenina River, 28.0 km 38° NE Andapa, 8.2 km 333° NNW Manantenina, 14.43667°S, 49.775°E, 450 m, rainforest, 12.—15.XI.2003 (B.L. Fisher et al.); Antsiranana, 15 km S Sambava, 10 m, 19.XII.1972 (J.-M. Bentsch); Fianarant- soa, Réserve Speciale Manombo 24.5 km 228° Farafangana, 23.01583°S, 47.719°E, 30 m, rainforest, 21.IV.2005 (BL. Fisher et al.); Toamasina, 6.3 km S Ambaniza- na, Andranobe, 15.6813°S, 49.958°E, 25 m, rainforest, 14.X1.1993 (BL. Fisher); Toamasina, 5.3 km SSE Ambanizana, Andranobe, 15.67133°S, 49.97395°E, 425 m, rainforest, 21.X1.1993 (B.L. Fisher); Toamasina, Réserve Spéciale Ambatovaky, San- drangato river, 16.77274°S, 49.26551°E, 450 m, rainforest, 20.—22.11.2010 (B.L. Fish- er et al.); Toamasina, Réserve Spéciale Ambatovaky, Sandrangato river, 16.76912°S, Taxonomy of Malagasy Tetramorium 133 49.26704°E, 475 m, rainforest, 21.11.2010 (BL. Fisher et al.); Toamasina, Réserve Spéciale Ambatovaky, Sandrangato river, 16.7633°S, 49.26692°E, 520 m, rainforest, 22.11.2010 (B.L. Fisher et al.); Toamasina, 19 km ESE Maroantsetra, 15.48333°S, 49.9°E, 300 m, rainforest, 22.1V.1989 (P.S. Ward); Toamasina, Nosy Mangabe, 15.5°S, 49.76667°E, 20—50 m, rainforest, 20.1V.1989 (P.S. Ward); Toamasina, F.C. Sandranantitra, 18.04833°S, 49.09167°E, 450 m, rainforest, 18.—21.1.1999 (4_/. Ratsirarson); Toamasina, S.F. Tampolo, 10 km NNE Fenoarivo Atn., 17.2825°S, 49 .43°E, 10 m, littoral rainforest, 4.1V.1997 (B.L. Fisher). Diagnosis. The following character combination distinguishes 7: rala from the remainder of the T° schaufussii species complex: relatively small species (HW 0.46—-0.49; WL 0.58-—0.64); eyes relatively large (OI 26—28); antennal scapes very short (SI 61-68); frontal carinae usually very weakly developed, only feebly raised, usually ending shortly after posterior eye margin or merging with cephalic sculpture halfway between posterior eye margin and posterior head margin; propodeal spines medium-sized to long, elon- gate-triangular to spinose, and acute (PSLI 21—25), propodeal lobes short, triangular, and acute or blunt, but always much shorter than propodeal spines, spines and lobes not strongly inclined towards each other; petiolar node in profile high rounded nodiform to thinly cuneiform, in profile 2.0 to 2.2 times higher than long (LPel 45-50), in dorsal view around 1.3 to 1.5 times broader than long (DPel 129-145); mesosoma with three to four pairs on dorsal promesonotum, propodeum and waist segments without any standing pilosity; body uniformly whitish yellow to light brown. Worker measurements (N=15). HL 0.49-0.53 (0.51); HW 0.46—0.49 (0.47); SL 0.28-0.33 (0.31); EL 0.12-0.13 (0.13); PH 0.22—0.24 (0.23); PW 0.32-0.37 (0.35); WL 0.58-0.64 (0.61); PSL 0.11-0.13 (0.12); PTL 0.09-0.11 (0.10); PTH 0.20-0.23 (0.21); PTW 0.13-0.16 (0.14); PPL 0.12—-0.14 (0.13); PPH 0.18-0.21 (0.19); PPW 0.18—0.22 (0.19); CI 90-94 (92); SI 61-68 (65); OI 26-28 (27); DMI 52-60 (57); LMI 36-39 (37); PSLI 21-25 (23); PeNI 37-43 (40); LPel 45-50 (49); DPel 129-145 (136); PpNI 52-59 (54); LPpI 67-75 (70); DPpI 141-157 (148); PPI 129-142 (138). Worker description. Head clearly longer than wide (CI 90-94); posterior head margin weakly concave, almost straight. Anterior clypeal margin with distinct me- dian impression. Frontal carinae usually very weakly developed, only feebly raised, usually ending shortly after posterior eye margin or merging with cephalic sculpture halfway between posterior eye margin and posterior head margin. Antennal scrobes very weak to absent, very shallow and without clear and distinct posterior and ventral margins. Antennal scapes very short, not reaching posterior head margin (SI 61-68). Eyes relatively large (OI 26-28). Mesosomal outline in profile flat to weakly convex, comparatively low and long (LMI 36-39), moderately marginate from lateral to dorsal mesosoma; promesonotal suture absent; metanotal groove very weak to absent. Pro- podeal spines medium-sized, elongate-triangular to spinose, and acute (PSLI 21-25), propodeal lobes short, triangular, and acute or blunt, but always much shorter than propodeal spines, spines and lobes not strongly inclined towards each other. Petiolar node in profile high rounded nodiform to thinly cuneiform, around 2.0 to 2.2 times 134 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) higher than long (LPel 45—50), anterior and posterior faces not parallel, anterodorsal margin usually situated higher and more strongly angled than posterodorsal margin, petiolar dorsum relatively flat to weakly convex and tapering backwards posteriorly; node in dorsal view around 1.3 to 1.5 times broader than long (DPel 129-145), in dorsal view pronotum between 2.3 to 2.7 times wider than petiolar node (PeNI 37— 43). Postpetiole in profile globular to subglobular, approximately 1.3 to 1.5 times higher than long (LPpI 67-75); in dorsal view around 1.4 to 1.6 times wider than long (DPpI 141-157), pronotum between 1.7 to 1.9 times wider than postpetiole (PpNI 52-59). Postpetiole in profile lower, thicker, and more rounded than petiolar node, postpetiole in dorsal view around 1.3 to 1.4 times wider than petiolar node (PPI 129-142). Mandibles completely unsculptured, smooth, and shiny; clypeus longitu- dinally rugulose with three to five rugulae, median rugula always present and usually fully developed, one or two mostly entire, rarely broken, lateral rugulae present on each side; cephalic dorsum between frontal carinae irregularly longitudinally rugulose with seven to ten fine rugulae, rugulae usually running from posterior clypeal margin to posterior head margin, but mostly irregularly shaped, interrupted or with cross- meshes; scrobal area mostly unsculptured; lateral head mainly longitudinally rugulose to reticulate-rugulose, but larger areas often only weakly sculptured and appearing fairly smooth and shiny; ground sculpture on head weakly to moderately punctate. Dorsum of mesosoma ranging from weakly longitudinally rugulose with larger areas with almost completely reduced sculpture to longitudinally rugose with well developed rugae; lateral mesosoma weakly to moderately irregularly longitudinally rugulose or reticulate-rugulose, often with larger areas of almost completely reduced sculpture; ground sculpture on mesosoma usually weak to absent, sometimes moderately punc- tate. Forecoxae , both waist segments, and gaster fully unsculptured, smooth, and shin- ing. Dorsum of head with several pairs of long, fine, standing hairs; mesosoma with three to four pairs on promesonotum; propodeum and waist segments without any standing pilosity; first gastral tergite with short, moderately dense, appressed (rarely decumbent) pubescence combined with several scattered, long, fine erect to suberect hairs; anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae with appressed to decumbent hairs. Body uniformly whitish yellow to light brown. Etymology. The new species is named after the fictional character “Rala” from Walter Moers’ fantasy novel “Rumo and His Miraculous Adventures”. The species epithet is an arbitrary combination of letter, thus invariable. Distribution and biology. ‘The distribution range of 7° rala is relatively disjunctive (Fig. 65). Its main distribution seems to be in the northeast in the area around the Bay of Antongil and the Masoala Peninsula north to Marojejy and Makirovana. Further south of this main cluster of localities 7° rala is only known from three additional places: Ambato- vaky, Tampolo, and Manombo. One explanation for this patchy distribution would be a preferense for lowland rainforests. Tetramorium rala lives in rainforests and littoral forests at lower elevations ranging from 10 to 550 m where it is found in leaf litter. Very few intact lowland rainforests remain south of its main distribution in the northeast of Madagascar. Taxonomy of Malagasy Tetramorium 135 Figure 55. 7: rala holotype worker (CASENT0046163). A Body in profile B Body in dorsal view C Head in full-face view. 136 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Discussion. Tetramorium rala is a very distinctive member of the T°. schaufussii complex due to its smaller size, relatively long propodeal spines, high nodiform to thinly cuneiform petiolar node, lack of standing pilosity on the waist segments, and very bright body colouration. Consequently, it is very unlikely to be confused with any other species. Most species are much larger in body size, have shorter propodeal spines, and/or possess standing pilosity on the waist segments. Only some smaller specimens of T° schaufussii could at first glance be mistaken with 7! rala, but these two species are effortlessly separable. Tetramorium schaufussii has much shorter propodeal spines (PSLI 11-18), a lower petiolar node, which in profile is only 1.6 to 1.9 times higher than long (LPel 52-63), and long, standing pilosity on the waist segments. By con- trast, in 7. rala the propodeal spines are much longer (PSLI 21-25), the petiolar node is higher, in profile around 2.0 to 2.2 higher than long (LPel 45—50), and the waist segments lack standing pilosity. However, even though 7° ralz is easily identifiable within the T° schaufussii com- plex, it is morphologically very close to T° rumo from the T: cognatum complex since they share a very similar gestalt. Both species are smaller in size, have relatively longer propodeal spines (compared to most other species of the species group), a high nodi- form to thinly cuneiform petiolar node, lack standing pilosity on the waist segments, and possess very bright body colouration. The main separating character, which also places them in separate species complexes, is the lack of standing pilosity on the first gastral tergite in 7: rumo versus the presence of standing pilosity on the first gastral tergite in 7° rala. Both species are morphologically closer to each other than to any other species of the 7: schaufussii species group, but there are highly diagnostic differ- ences to support their heterospecificity. The petiolar node of 7: rumo is thinner and stronger anteroposteriorly compressed, in profile around 2.3 to 2.7 times higher than long (LPel 37-43), and in dorsal view between 1.5 to 1.7 times wider than long (DPel 156-171), whereas T° rala has a node which in profile is 2.0 to 2.2 times higher than long (LPel 45-50), and in dorsal view around 1.3 to 1.5 times broader than long (DPel 129-145). In addition, T° rumo also has larger eyes (OI 28-31) than 7: rala (OI 26-28). Further evidence of their heterospecifity can be deduced from their distribu- tion ranges, which overlap in central-eastern Madagascar, but both species maintain their species-specific characteristic without intermediate forms. In Manombo both species were also found living in sympatry in close proximity. Tetramorium schaufussii Forel, 1891 http://species-id.net/wiki/Tetramorium_schaufussii Figs 42C, 45B, 46C, 47A, B, 56, 66 Tetramorium schaufussii Forel, 1891:158. Type material. Holotype, pinned worker, MADAGASCAR, Central Madagascar (C. Schaufuss) (MHNG: CASENT0101697). Taxonomy of Malagasy Tetramorium L357 Non-type material MADAGASCAR: Antananarivo, Réserve Spéciale d’Ambohitantely, Forét d’Ambohitantely, Jardin Botanique, 24.1 km 59° NE Anka- zobe, 18.17139°S, 47.28182°E, 1620 m, montane rainforest, 17.-22.1V.2001 (BL. Fisher et al.); Antananarivo, 3 km 41° NE Andranomay, 11.5 km 147° SSE Anjo- zorobe, 18.47333°S, 47.96°E, 1300 m, montane rainforest, 5.—13.XII.2000 (B.L. Fisher et al.); Antananarivo, Andrangoloaca (Sikora); Antananarivo, Stn. Forestiére Manjakatompo, 19.35°S, 47.31667°E, 1600 m, montane rainforest, 20.11.1993 (PS. Ward); Antsiranana, Réserve Spéciale de ’Ankarana, 13.6 km 192° SSW Anivorano Nord, 12.86361°S, 49.22583°E, 210 m, tropical dry forest, 19.—20.11.2001 (G.D. Alp- ert); Antsiranana, Parc National Montagne d’Ambre, Ambre grand lac, 12.59656°S, 49.15932°E, 1350 m, montane rainforest, 13.X1.2007 (B.L. Fisher et al.); Antsiranana, Parc National Montagne d’Ambre, Lac Maudit, 12.58502°S, 49.15147°E, 1250 m, montane rainforest, 13.-14.X1.2007 (B.L. Fisher et al.); Antsiranana, Parc National Montagne d’Ambre, 12.2 km 211° SSW Joffreville, 12.59639°S, 49.1595°E, 1300 m, montane rainforest, 2.—7.1].2001 (B.L. Fisher et al.); Antsiranana, Parc National Mon- tagne d’Ambre, 12.53417°S, 49.17607°E, 1325 m, montane rainforest, 12.11.2011 (B.L. Fisher et al.); Antsiranana, Rés. Anjanaharibe-Sud, 9.2 km WSW Befingotra, 14.75°S, 49.46667°E, 1200 m, montane rainforest, 9.-11.X1.1994 (B.L. Fisher); Antsiranana, Rés. Anjanaharibe-Sud, 11.0 km WSW Befingotra, 14.75°S, 49.45°E, 1550-1565 m, montane rainforest, 15.—21.X1.1994 (B.L. Fisher); Antsiranana, Forét de Binara, 9.4 km 235° SW Daraina, 13.26333°S, 49.6°E, 1100 m, montane rainfor- est, 5.X1I.2003 (B.L. Fisher); Antsiranana, Makirovana forest, 14.16666°S, 49.95°E, 715 m, rainforest, 2.V.2011 (BL. Fisher et al); Antsiranana, R.S. Manongarivo, 12.8 km 228° SW Antanambao, 13.97667°S, 48.42333°E, 780 m, rainforest, 11.— 17.X.1998 (B.L. Fisher); Antsiranana, R.S. Manongarivo, 14.5 km 220° SW Antan- ambao, 13.99833°S, 48.42833°E, 1175 m, montane rainforest, 19-25.X.1998 (BL. Fisher); Antsiranana, R.S. Manongarivo, 17.3 km 218° SW Antanambao, 14.02167°S, 48.41833°E, 1580 m, montane rainforest, 27.X.1998 (B.L. Fisher); Antsiranana, R.S. Manongarivo, 20.4 km 219° SW Antanambao, 14.04667°S, 48.40167°E, 1860 m, montane rainforest, 3.X1.1998 (B.L. Fisher); Antsiranana, R.N.I. Marojejy, 11 km NW Manantenina, 14.45°S, 49.73333°E, 1875 m, montane rainforest, 13.-19.X1.1996 (E.L. Quinter); Antsiranana, R.N.I. Marojejy, 11 km NW Manantenina, 14.43333, 49.75°E, 1225 m, montane rainforest, 25.X.—19.X1.1996 (E.L. Quinter); Antsiranana, Parc National de Marojejy, Manantenina River, 28.0 km 38° NE Andapa, 8.2 km 333° NNW Manantenina, 14.43667°S, 49.775°E, 450 m, rainforest, 12.-15.X1.2003 (B.L. Fisher et al.); Antsiranana, Parc National de Marojejy, Antranohofa, 26.6 km 31° NNE Andapa, 10.7 km 318° NW Manantenina, 14.44333°S, 49.74333°E, 1325 m, montane rainforest, 19.—20.XI.2003 (B.L. Fisher); Antsiranana, Parc National de Marojejy, 25.7 km 32° NNE Andapa, 10.3 km 314° NW Manantenina, 14.445°S, 49.74167°E, 1575 m, montane rainforest, 21.—22.X1.2003 (B.L. Fisher); Fianarant- soa, 28 km SSW Ambositra, 20.76667°S, 47.18333°E, 1660 m, rainforest, 29.[V.1989 (P.S. Ward); Fianarantsoa, 27.4 km SSW Ambositra, 20.77°S, 47.18667°E, 1600 m, montane rainforest, 15.1.1998 (B.L. Fisher); Fianarantsoa, 26.8 km SW Ambositra, 138 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Parc naturel communautaire, 20.775°S, 47.18362°E, 1755 m, disturbed montane rain- forest, 20.V.2008 (B.L. Fisher et al.); Fianarantsoa, Rés. Andringitra, 43 km S Amba- lavao, 22.23333°S, 47°E, 825 m, rainforest, 5.X.1993 (B.L. Fisher); Fianarantsoa, Rés. Andringitra, 38 km S Ambalavao, 22.2°S, 46.96667°E, 1680 m, montane rainforest, 23.X.1993 (B.L. Fisher); Fianarantsoa, P. N. Andringitra, Forét Ravaro 12.5 km SW Antanifotsy, 22.21167°S, 46.845°E, 1650 m, montane rainforest, 10.—15.1.2000 (S. Razafimandimby); Fianarantsoa, Forét d’Atsirakambiaty, 7.6 km 285° WNW Itremo, 20.59333°S, 46.56333°E, 150 m, montane rainforest, 22.—-26.1.2003 (B.L. Fisher et al.); Fianarantsoa, Parc National Befotaka-Midongy, Papango 27.7 km S Midongy- Sud, Mount Papango, 23.83517°S, 46.96367°E, 940 m, rainforest, 13.-19.XI.2006 (B.L. Fisher et al.); Fianarantsoa, Parc National Befotaka-Midongy, Papango 28.5 km S Midongy-Sud, Mount Papango, 23.84083°S, 46.9575°E, 1250 m, montane rainfor- est, 17.-18.2006 (B.L. Fisher et al.); Fianarantsoa, R.S. Ivohibe, 6.5 km ESE Ivohibe, 22.49667°S, 46.955°E, 1575 m, montane rainforest, 24.-30.X.1997 (B.L. Fisher); Fianarantsoa, 7 km W Ranomafana, 1100 m, montane rainforest, 1.—7.1.1988 (WE. Steiner); Fianarantsoa, Ranomafana National Park, Maharira forest, 21°S, 47°E, 1375 m, montane rainforest, 12.X.1992 (E. Rajeriarison); Fianarantsoa, Ranomafana Na- tional Park, Maharira forest, 21°S, 47°E, 1130 m, montane rainforest, 29.XII.1992 (E. Rajeriarison); Fianarantsoa, Parc National de Ranomafana, Vatoharanana River, 4.1 km 231° SW Ranomafana, 21.29°S, 47.43333°E, 1100 m, montane rainforest, 27.-31.111.2003 (BL. Fisher et al); Fianarantsoa, Parc National de Ranomafana, Sa- hamalaotra River, 6.6 km 310° NW Ranomafana, m, 21.23667°S, 47.39667°E, 1150 m, montane rainforest, 31.11.2003 (B.L. Fisher et al.); Mahajanga, Réserve Spéciale Marotandrano, Marotandrano 48.3 km S Mandritsara, 16.28322°S, 48.81443°E, 865 m, transition humid forest, 7.XII.2007 (B.L. Fisher et al.); Toamasina, 6.9 km NE Ambanizana, Ambohitsitondroina, 15.58506°S, 50.00952°E, 825 m, rainforest, 2.X11.1992 (B.L. Fisher); Toamasina, Réserve Spéciale Ambatovaky, Sandrangato river, 16.76912°S, 49.26704°E, 475 m, rainforest, 21.11.2010 (BL. Fisher et al.); Toamasi- na, Réserve Spéciale Ambatovaky, Sandrangato river, 16.7674°S, 49.26813°E, 500 m, rainforest, 23.11.2010 (B.L. Fisher et al.); Toamasina, Réserve Spéciale Ambatovaky, Sandrangato river, 16.81745°S, 49.2925°E, 400 m, rainforest, 26..2010(B.L. Fisher et al.); Toamasina, Analamay, 18.80623°S, 48.33707°E, 1068 m, montane rainfor- est, 23.11.2004 (Malagasy ant team); Toamasina, 6 km ESE Andasibe (=Perinet), 18.95°S, 48.46667°E, 900 m, rainforest, 17.XI.1990 (P.S. Ward); Toamasina, Mon- tagne d’Anjanaharibe, 19.5 km 27° NNE Ambinanitelo, 15.17833°S, 49.635°E, 1100 m, montane rainforest, 12.—16.II.2003 (B.L. Fisher et al.); Toamasina, Reserve Bet- ampona, Camp Rendrirendry 34.1 km 332° Toamasina, 17.924°S, 49.19967°E, 390 m, rainforest, 29.—30.X1.2005 (B.L. Fisher et al.); Toamasina, Parc National de Zaha- mena, Tetezambatana forest, near junction of Nosivola and Manakambahiny rivers, 17.74298°S, 48.72936°E, 860 m, rainforest, 19.11.2009 (B.L. Fisher et al.); Toamasina, Parc National de Zahamena, Onibe River, 17.75908°S, 48.85468°E, 780 m, rainforest, 22.11.2009 (B.L. Fisher et al.); Toliara, Parc National d’Andohahela, Col du Sedro, 3.8 km 113° ESE Mahamavo, 37.6 km 341° NNW Tolagnaro, 24.76389°S, 46.75167°E, Taxonomy of Malagasy Tetramorium 139 900 m, montane rainforest, 21.-25.I. (B.L. Fisher et al.); Toliara, Réserve Spéciale Kal- ambatritra, Betanana, 23.4144°S, 46.459°E, 1360 m, montane rainforest, 8.[1.2010 (B.L. Fisher et al.); Toliara, Réserve Spéciale Kalambatritra, Ambinanitelo, 23.4502°S, 46.45658°E, 1325 m, montane rainforest, 11.11.2010 (B.L. Fisher et al.); Toliara, Ré- serve Spéciale Kalambatritra, Ampanihy, 23.4635°S, 46.4631°E, 1270 m, montane rainforest, 9.—10.11.2010 (B.L. Fisher et al); REUNION: no locality data (G. Mayr). Diagnosis. The following character combination separates 7: schaufussii from the remaining species of the 7° schaufussii complex: head much longer than wide (CI 86-90); antennal scapes short to very short (SI 66-73); eyes moderate to large (OI 24—28); frontal carinae usually moderately well developed, slightly diverging posteri- orly, and typically fading out or merging with surrounding sculpture halfway between posterior eye and posterior head margin; propodeal spines usually short, triangular to elongate-triangular (PSLI 11-18), propodeal lobes short and triangular, usually of approximately same length as propodeal spines, sometimes lobes longer, rarely much shorter than spines, but spines and lobes never strongly inclined towards each other; petiolar node in profile high rounded nodiform, around 1.6 to 1.9 times higher than long (LPeI 52-63), in dorsal view between 1.1 to 1.4 times wider than long (DPel 112-136); propodeum without standing pilosity; petiole and postpetiole with long, standing pilosity. Worker measurements (N=60). HL 0.59-0.76 (0.67); HW 0.51—0.68 (0.60); SL 0.36-0.50 (0.42); EL 0.14-0.19 (0.16); PH 0.26—0.35 (0.31); PW 0.38-0.50 (0.45); WL 0.70-0.93 (0.82); PSL 0.07—0.14 (0.10); PTL 0.12-0.17 (0.14); PTH 0.21-0.29 (0.25); PTW 0.15-0.20 (0.17); PPL 0.15-0.22 (0.18); PPH 0.20—0.28 (0.24); PPW 0.21—0.28 (0.25); CI 86-90 (88); SI 66-73 (71); OI 24-28 (26); DMI 52-59 (54); LMI 35-39 (37); PSLI 11-18 (15); PeNI 35-41 (39); LPel 52-63 (57); DPel 112-136 (124); PpNI 53-61 (56); LPpI 69-83 (75); DPpI 125-150 (137); PPI 131-153 (142). Worker description. Head much longer than wide (CI 86-90); posterior head mar- gin weakly concave. Anterior clypeal margin with distinct median impression. Frontal carinae usually moderately well developed, slightly diverging posteriorly, and typically fading out or merging with surrounding sculpture halfway between posterior eye and posterior head margin. Antennal scrobes weak to absent, shallow and without clear and distinct posterior and ventral margins. Antennal scapes short to very short, not reach- ing posterior head margin (SI 66-73). Eyes moderate to large (OI 24-28). Mesosomal outline in profile flat to weakly convex, comparatively low and long (LMI 35-39), moderately marginate from lateral to dorsal mesosoma; promesonotal suture absent; metanotal groove either weakly developed or absent. Propodeal spines usually short, triangular to elongate-triangular (PSLI 11-18), propodeal lobes short and triangular, usually of approximately same length as propodeal spines, sometimes lobes longer, rare- ly much shorter than spines, but never strongly inclined towards each other. Petiolar node in profile high rounded nodiform, around 1.6 to 1.9 times higher than long (LPel 52-63), anterior and posterior faces approximately parallel, usually anterodorsal and posterodorsal margins situated at about same height and equally angled, sometimes an- 140 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) terodorsal margin slightly higher, petiolar dorsum generally weakly convex, sometimes flat; node in dorsal view between 1.1 to 1.4 times wider than long (DPel 112-136), in dorsal view pronotum between 2.4 to 2.8 times wider than petiolar node (PeNI 35-41). Postpetiole in profile globular, around 1.2 to 1.4 times higher than long (LPpl 69-83); in dorsal view between 1.2 to 1.5 times wider than long (DPpI 125-150), pro- notum between 1.6 to 1.9 times wider than postpetiole (PpNI 53-61). Postpetiole in profile appearing more or less of similar volume as petiolar node, postpetiole in dorsal view around 1.3 to 1.5 times wider than petiolar node (PPI 131-153). Mandibles com- pletely unsculptured, smooth, and shiny; clypeus longitudinally rugulose/rugose with three to six usually regularly shaped and unbroken rugulae/rugae, median ruga usu- ally fully developed and distinct, very rarely broken, one or two lateral rugulae/rugae present on each side; cephalic dorsum between frontal carinae longitudinally rugulose/ rugose with seven to ten rugae/rugulae, rugae/rugulae usually running from posterior clypeal margin to posterior head margin, often irregularly shaped, interrupted, or with cross-meshes; scrobal area partly unsculptured and merging with surrounding sculp- ture; lateral head reticulate-rugose to longitudinally rugose, often posteriorly mostly unsculptured. Ground sculpture on head usually well developed, moderately reticulate- punctate, especially on cephalic dorsum and scrobal area, sometimes ground sculpture much weaker, almost absent. Dorsum of mesosoma usually irregularly longitudinally rugose to reticulate-rugose, sometimes almost completely reticulate-rugose with few longitudinally rugose elements; lateral mesosoma mostly irregularly longitudinally ru- gose to reticulate-rugose, often lateral pronotum weaker sculptured to almost unsculp- tured. Ground sculpture on mesosoma variably developed, usually weakly to moder- ately punctate, sometimes very weak or absent. Forecoxae either unsculptured, smooth, and shining or with longitudinally rugulose or reticulate-rugose sculpture on upper half. Both waist segments and gaster fully unsculptured, smooth, and shining. Dorsum of head with several pairs of long, fine, standing hairs; mesosoma with six or more pairs on promesonotum, propodeum without standing pilosity; petiole usually with one or two and postpetiole with two to three pairs of long, standing hairs; first gastral tergite with short, moderately dense, appressed pubescence in combination with several scat- tered to numerous, long, standing hairs. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae with appressed to decumbent hairs. Head, mesosoma, waist segments, and gaster uniformly light yellowish brown to very dark brown, almost black contrasting with lighter yellowish to light brown mandibles, antennae, and legs. Distribution and biology. Tetramorium schaufussii is broadly distributed through- out most of the humid forest zones from the southeast to the north of Madagascar (Fig. 66). Surprisingly, 7. schaufussii is also known from one very old collection event from the island of Reunion, but apart from that no modern collection exists from that island. Despite that we list Reunion as record for 7. schaufussii we consider that record as problematic and highly questionable. In Madagascar T° schaufussii is almost always found in rainforests or montane rainforests at elevations from 210 to 1875 m, even though most of the material was collected at elevations higher than 1000 m. In addi- tion, 7: schaufussii seems to be an inhabitant of the leaf litter stratum. Taxonomy of Malagasy Tetramorium 141 Discussion. Tetramorium schaufussi is the most widespread, common, and abun- dant species of the T° schaufussi complex. Also, T. schaufussii co-occurs with all other members of the complex throughout its range. It can be well separated from most species, but its separation from a few morphologically close forms is more challenging. Due to its larger eyes (OI 24-28) T° schaufussii is very unlikely to be mistaken for T- pseudogladius (OI! 20). Additionally, its petiolar node shape, which is always broader than long (DPel 112-136), distinguishes it from 7. nassonowii, which has a longer than broad node (DPel 87-98). ‘The species T. scutum, despite being morphologically relatively similar, has longer propodeal spines (PSLI 22—24) and the spines and pro- podeal lobes are strongly inclined towards each other. Tetramorium schaufussii always has shorter spines (PSLI 11-18) and the spines and lobes are never strongly inclined towards each other. Tetramorium rala and T. sikorae \ack long, standing pilosity on the waist segments, separating them easily from 7. schaufussii, but is should be noted that without consideration of the pilosity on the waist segments, 7: sikorae is generally very similar to T° schaufussii. The difference, however, is very constant and both are usually found in sympatry, leading us to the decision to retain them as two species. Further- more, most of the material of 7: obiwan also lacks pilosity on the waist segments, distinguishing it from 7! schaufussii, and, if pilosity is present, it is reduced to one pair on the petiole or postpetiole. Additionally, 7: obiwan possesses longer antennal scapes (SI 77-82) and is much larger (HW 0.71-0.84; WL 1.00-1.20) than T° schaufussii (SI 66-73; HW 0.51-0.68; WL 0.70—0.93). In Tetramorium body size is usually not use- ful and often even misleading due to high intraspecific variation in many species (Hita Garcia et al. 2010; Hita Garcia and Fisher 2012a), but in the case of these two species we can confidently separate them by size. The three species 7’ merina, T. monticola, and T: xanthogaster are often difficult to separate from T! schaufussii. The most important discriminating difference between these three and T° schaufussii is the shape of the head. In T° schaufussii the head is usually much longer and thinner (CI 86—90 vs. CI 91-95 in the other three species). Additionally, in T. schaufussii the clypeus almost always has a very distinct and unbro- ken median longitudinal ruga, whereas T: merina, T: monticola, and T. xanthogaster normally have the median area fully unsculptured without any median ruga/rugula at all, or the median ruga/rugula is present, but then it is usually interrupted, very ir- regularly shaped, or only present as traces. This character has to be treated with caution however, as there are a few specimens in 7: merina, T. monticola, and T. xanthogaster, in which the median ruga/rugula is present. The delimitations of T° merina and T: schaufussii can be difficult, but in areas where they co-occur in the Central Highlands of Madagascar they can be easily separated by body size. In that area T. merina is always much larger in size and has shorter propodeal spines (PSLI 7-11) than 7! schaufussii (PSLI 11-18). It must be pointed out that 7! schaufussii is highly variable, likely the most vari- able species of the T. schaufussii complex and the whole species group. ‘There are several important characters varying significantly throughout the distribution range that need to be addressed here. ‘The frontal carinae are moderately well developed in most of the 142 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) 1 Figure 56. 7° schaufussii holotype worker (CASENT0101697). A Body in profile B Body in dorsal view C Head in full-face view. material, but in one series from Mt. Anjanaharibe they are reduced and much weaker. As do several other species of the group, T° schaufussii possesses a comparatively variable petiolar node shape. It is generally high rounded nodiform with the anterodorsal and posterodorsal margins situated at about the same height and both equally rounded or an- Taxonomy of Malagasy Tetramorium 143 gled. However, the node is often lower and thicker, and this variation can be seen within the same series or population. Sometimes the node also has a slightly higher and more angled anterodorsal margin in comparison to the posterodorsal margin, even though only weakly so. In addition, even though mostly stable, the propodeal spines do occasionally vary. [hey are usually short to very short (PSLI 11-15), but some specimens from An- dranomay, Binara, and Marojejy have longer spines (PSLI 16-18). Also highly variable is colouration that also contains a geographic component. Most of the material from the southeast to the central east is much darker in colour, usually dark brown to almost black. ‘The material from the north is very variable in colour ranging from very bright yellow (Mt. Anjanaharibe) to almost black (Marojejy), but usually constant on a local scale. Parallel to the colouration, there is also geographic variation observable in the sculpture on the forecoxae. Most of the darker material has the upper part of the forecoxae dis- tinctly sculptured, whereas in most of the brighter material the forecoxae are completely unsculptured. However, this is not always the case, and thus not used for diagnostics. The series from Mt. Anjanaharibe mentioned above also requires some comments. The specimens are much brighter in color, smaller in size, and have much weaker frontal carinae than the rest of the 7. schaufussii material. These are not unlikely to turn out to be a distinct species, but for the moment we keep it as a smaller, brighter geographical variation of T° schaufussii. Tetramorium scutum Hita Garcia & Fisher, sp. n. http://zoobank.org/7D34D957-AAD0-4C22-8 1 C5-AIDEACC35303 http://species-id.net/wiki/Tetramorium_scutum Figs 41B, 42A, 57, 66 Type material. Holotype, pinned worker, MADAGASCAR, Fianarantsoa, R.S. Ivo- hibe 8.0 km E Ivohibe, 22.48333°S, 46.96833°E, 1200 m, montane rainforest, sift- ed litter (leaf mold, rotten wood), collection code BLF0O1747, 15.—-21.X.1997 (BL. Fisher) (CAS: CASENT0189116). Paratypes, five pinned workers with same data as holotype (BMNH: CASENT0218026; CAS: CASENT0218027; CASENT0248303; CASENT0248304; CASENT0248305); five pinned workers from Fianarantsoa, 8.0 km NE Ivohibe, 22.42167°S, 46.89833°E, 1200 m, montane rainforest, sifted litter (leaf mold, rotten wood), collection code BLF01753, 3.—9.X1.1997 (B.L. Fisher) (CAS: CASENT0218022; CASENT0218023; CASENT0218028; CASENT0248309; MCZ: CASENT0248308); seven pinned workers from Fianarantsoa, R.S. Ivohibe, 6.5 km ESE Ivohibe, 22.49667°S, 46.955°E, 1575 m, montane rainforest, sifted litter (leaf mold, rotten wood), collection code BLF0O1751, 24.-30.X.1997 (B.L. Fisher) (CAS: CASENT0189154; CASENT0218021; CASENT0218024; CASENT0218025; CASENT0248306; CASENT0248307). Diagnosis. Tetramorium scutum can be separated from the remainder of the spe- cies complex by the following character combination: moderate to large eyes (OI 24— 25); short antennal scapes (SI 71-74); frontal carinae moderately to well developed, 144 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) slightly diverging posteriorly, ending at or shortly before posterior head margin; pro- podeal spines moderate to long (PSLI 22-24), propodeal lobes elongate-triangular, always weakly shorter than propodeal spines, in profile spines and lobes strongly in- clined towards each other; petiolar node in profile around 2.0 to 2.1 times higher than long (LPel 48-50) and in dorsal view around 1.4 to 1.5 times wider than long (DPel 144-154); dorsum of mesosoma with six or more pairs of long, standing hairs from anterior pronotum to posterior mesonotum, propodeum without any standing pilos- ity; waist segments with long, standing pilosity. Worker measurements (N=10). HL 0.65—0.70 (0.67); HW 0.60-—0.64 (0.61); SL 0.43-0.45 (0.44); EL 0.15—0.16 (0.15); PH 0.31—0.33 (0.32); PW 0.44-0.48 (0.45); WL 0.79-0.86 (0.83); PSL 0.15—-0.17 (0.16); PTL 0.12—0.14 (0.13); PTH 0.25—0.28 (0.27); PTW 0.19-0.21 (0.19); PPL 0.18—-0.21 (0.19); PPH 0.25—0.29 (0.XX); PPW 0.25—0.29 (0.27); CI 90-92 (91); SI 71-74 (72); OI 24-25 (24); DMI 54-59 (55); LMI 38-39 (38); PSLI 22—24 (23); PeNI 41-44 (43); LPeI 48-50 (49); DPel 144-154 (148); PpNI 57-62 (59); LPpI 70-74 (72); DPpI 134-140 (138); PPI 133-142 (138). Worker description. Head longer than wide (CI 90-92); in full-face view poste- rior head margin weakly concave. Anterior clypeal margin with distinct median im- pression. Frontal carinae moderately to well developed, slightly diverging posteriorly, ending at or shortly before posterior head margin. Antennal scrobes weak to absent, shallow and without clear and distinct posterior and ventral margins. Antennal scapes short, not reaching posterior head margin (SI 71-74). Eyes moderate to relatively large (OI 24-25). Mesosomal outline in profile flat to weakly convex, comparatively low and long (LMI 38-39), moderately marginate from lateral to dorsal mesosoma; promesonotal suture absent; metanotal groove weak or absent. Propodeal spines mod- erate to long, elongate-triangular to spinose, and acute (PSLI 22-24); propodeal lobes elongate-triangular, always weakly shorter than propodeal spines, in profile spines and lobes strongly inclined towards each other. Petiolar node in profile high rounded nodi- form to weakly squamiform, around 2.0 to 2.1 times higher than long (LPel 48-50), anterior and posterior faces approximately parallel, anterodorsal and posterodorsal margins situated at about same height, petiolar dorsum relatively flat; node in dorsal view between 1.4 to 1.6 times wider than long (DPel 144-154), in dorsal view prono- tum between 2.2 to 2.5 times wider than petiolar node (PeNI 41-44). Postpetiole in profile subglobular, around 1.3 to 1.4 times higher than long (LPpI 70-74); in dorsal view around 1.3 to 1.4 times wider than long (DPpI 134-140), pronotum between 1.6 to 1.8 times wider than postpetiole (PpNI 57-62). Postpetiole in profile appearing slightly more voluminous than petiolar node, postpetiole in dorsal view around 1.3 to 1.4 times wider than petiolar node (PPI 133-142). Mandibles completely unsculp- tured, smooth, and shiny; clypeus weakly, irregularly, longitudinally rugulose with me- dian area mostly unsculptured, smooth, and shiny, sometimes median rugula present, but then weakly developed and/or interrupted, one or two weak, irregular and broken rugulae present on each side; cephalic dorsum between frontal carinae longitudinally rugulose with six to nine rugae/rugulae, rugae/rugulae usually running from poste- rior clypeal margin to posterior head margin, often irregularly shaped, interrupted or Taxonomy of Malagasy Tetramorium 145 Figure 57. 7° scutum holotype worker (CASENT0189116). A Body in profile B Body in dorsal view C Head in full-face view. 146 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) with cross-meshes; scrobal area partly unsculptured and merging with surrounding sculpture; lateral head anteriorly reticulate-rugose to longitudinally rugose and posteri- orly mostly unsculptured. Ground sculpture on head weakly to moderately reticulate- punctate, especially laterally. Dorsum of mesosoma irregularly longitudinally rugose; lateral mesosoma mostly irregularly longitudinally rugose with some reticulate-rugose elements. Ground sculpture on mesosoma weak to absent. Forecoxae unsculptured, smooth, and shining. Both waist segments and gaster fully unsculptured, smooth, and shining. Dorsum of head with several pairs of long, fine, standing hairs; mesosoma with six or more pairs from anterior pronotum to posterior mesonotum, propodeum without standing pilosity; petiole with one and postpetiole with one or two pairs of long, standing hairs; first gastral tergite with short, moderately dense, appressed pu- bescence in combination with several scattered, long, standing hairs. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae with appressed hairs. Head, mesosoma, waist segments and gaster uniformly brown to dark brown contrasting with yellowish to light brown mandibles, antennae, and legs. Etymology. The name of the new species is Latin and means “shield”, referring to the weakly squamiform condition of the petiolar node. ‘The species epithet is a nomi- native noun, and thus invariant. Distribution and biology. This new species is only known from the area around Ivohibe (Fig. 66) where it was collected from a few montane rainforest localities situ- ated at elevations of 1200 to 1575 m. All specimens were sampled from leaf litter. Discussion. 7 etramorium scutum is relatively easy to distinguish within the species complex since it is the only species in which the moderately long spines (PSLI 22-22) and the comparatively long propodeal lobes are strongly inclined towards each other. In all the other group members the spines are either much shorter or if they are of ap- proximately similar length as in T° scutum, then the lobes are much smaller and spines and lobes are never strongly inclined towards each other. Interestingly, the species most similar to 7! scutum are T. aspis and T. camelliae from the T: cognatum complex, and we suspect that they are also more closely related to each other than to the rest of the group. These three species all have strongly inclined spines and lobes and are also found in the same small area in the southeast of Madagascar. The new species is only known from a relatively small area and consequently shows very little intraspecific variation. Tetramorium sikorae Forel, 1892 http://species-id.net/wiki/Tetramorium_sikorae Figs 41D, 43C, 44B, 45A, 58, 66 Tetramorium (Xiphomyrmex) sikorae Forel, 1892:522. [Combination in Xiphomyrmex: Emery 1895:343; in Tetramorium: Bolton 1979:138] Xiphomyrmex latior Santschi, 1926:243. [Synonymy with 7’ sikorae by Bolton 1979:138; here confirmed] Taxonomy of Malagasy Tetramorium 147 Type material. Of 7: sikorae: lectotype [designated here], pinned worker, MAD- AGASCAR, Toamasina, Amparafaravantsiv, Mangoro Ufer 984, 20°S, 47.08333 E (Sikora) (MHNG: CASENT0101260) [examined]. Paralectotypes [designated here], three pinned workers with same data as holotype (MHNG: CASENT0101259; CASENT0101898) [examined]. Of 7: Jatior: syntypes, six pinned workers, MADAGASCAR, Fianarantsoa, 21.45°S, 47.083333 E (Descarpentries) (NHMB: CASENT0101140; CASENT0101141; CASENT0101142; CASENT0101143; CASENT0101144) [examined]. [Note: the GPS data of both type localities was not provided by either locality labels or the original descriptions. The GPS data presented above is based on our own geo-referencing of the locality Amparafara, which is close to the Mangoro River, and the city Fianarantsoa. The data for both locliaties should be considered as approxima- tions and not the exact positions of the type localities. ] Non-type material. MADAGASCAR: no locality, 1.11.1977 (W.L. & D.E. Brown); Antsiranana, Forét Ambanitaza, 26.1 km 347° Antalaha, 14.67933°S, 50.18367°E, 240 m, rainforest, 27.X1.2004 (B.L. Fisher); Antsiranana, Res. d’ Anjanaharibe-Sud, 17 km W Andapa, 14.75775°S, 49.51019°E, 875 m, primary rainforest, 7.XI.1994 (G.D. Alpert); Antsiranana, Forét de Binara, 9.4 km 235° SW Daraina, 13.26333°S, 49.6°E, 1100 m, montane rainforest, 6.XII.2003 (B.L. Fisher); Antsiranana, Parc National de Marojejy, Manantenina River, 28.0 km 38° NE Andapa, 8.2 km 333° NNW Manantenina, 14.43667°S, 49.775°E, 450 m, rainforest, 12.-15.XI.2003 (B.L. Fisher et al.); Antsiranana, Parc National de Marojejy, Manantenina River, 27.6 km 35° NE Andapa, 9.6 km 327° NNW Manantenina, 14.435°S, 49.76°E, 775 m, rain- forest, 16.X1.2003 (B.L. Fisher); Fianarantsoa, 2 km W Andrambovato, along riv- er Tatamaly, 21.51167°S, 47.41°E, 1075 m, cultivated land (tavy), recent one- to two-year-old tavy, 3.-5.V1.2005 (B.L. Fisher et al.); Toamasina, Corridor Forestier Analamay-Mantadia, Ambatoharanana, 18.80388°S, 48.40506°E, 1013 m, rainfor- est, 12.-19.XII.2012 (BL. Fisher et al.); Toamasina, Parc National d’ Andasibe- Mantadia, Forét de Mantadia, 25.7 km 248° Moramanga, 18.81402°S, 48.43028°E, 1040 m, rainforest, 13.VI.2006 (F.N. Raharimalala & B. Blaimer); Toamasina, Ankerana, 18.4061°S, 48.82029°E, 725 m, rainforest, 20.1.2012 (B.L. Fisher et al.); Toamasina, Morarano-Chrome, 25 km W, forest, 15.VI.1991 (A. Pauly); Toamasina, Torotorofotsy, 18.87082°S, 48.34737°E, 1070 m, montane rainforest, marsh edge, 27.111.2004 (Malagasy ant team); Toliara, Réserve Spéciale d’Ambohijanahary, Forét d’Ankazotsihitafototra, 35.2 km 312° NW Ambaravaranala, 18.26667°S, 45.40667°E, 1050 m, montane rainforest, 13.—17.III.2003 (B.L. Fisher et al). Diagnosis. The following character combination distinguishes 7: sikorae from the remainder of the 7. schaufussii species complex: smaller species (HW 0.53-0.61; WL 0.70-0.82); head much longer than wide (CI 86-89); antennal scapes short (SI 70-74); eyes relatively large (OI 26-28); frontal carinae moderately developed, slightly diverging posteriorly, becoming weaker halfway between posterior eye margin and posterior head margin, ending at or shortly before posterior head margin; propodeal spines reduced to short, triangular teeth (PSLI 11-14), propodeal lobes short, trian- 148 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) gular, and usually blunt, approximately of same size as propodeal teeth, and in profile teeth and lobes not strongly inclined towards each other; petiolar node high rounded nodiform, in profile around 1.7 to 1.9 times higher than long (LPel 53-59), in dorsal view around 1.2 to 1.3 times wider than long (DPeI 120-133); dorsum of mesosoma usually with two, sometimes four, pairs of long, standing hairs on promesonotum; propodeum and waist segments without any long, standing pilosity. Worker measurements (N=15). HL 0.60—0.69 (0.62); HW 0.53-0.61 (0.55); SL 0.38-0.43 (0.39); EL 0.14-0.16 (0.15); PH 0.26-0.31 (0.28); PW 0.39-0.47 (0.42); WL 0.70-0.82 (0.74); PSL 0.07—0.09 (0.08); PTL 0.12-0.15 (0.13); PTH 0.21-0.26 (0.23); PTW 0.16-0.19 (0.17); PPL 0.14-0.17 (0.16); PPH 0.21—0.25 (0.23); PPW 0.21—0.26 (0.23); CI 86-89 (88); SI 70-74 (72); OI 26-28 (27); DMI 53-63 (57); LMI 36-39 (38); PSLI 11-14 (12); PeNI 38-43 (40); LPeI 53-59 (57); DPel 120-133 (129); PpNI 51-58 (55); LPpI 67-72 (69); DPpI 142-157 (149); PPI 131-144 (137). Worker description. Head much longer than wide (CI 86-89); in full-face view posterior head margin weakly to moderately concave. Anterior clypeal margin with distinct median impression. Frontal carinae moderately developed, slightly diverging posteriorly, becoming weaker halfway between posterior eye margin and posterior head margin, ending at or shortly before posterior head margin. Antennal scrobes weak to absent, shallow and without clear and distinct posterior and ventral margins. Antennal scapes short, not reaching posterior head margin (SI 70-74). Eyes relatively large (OI 26-28). Mesosomal outline in profile flat to weakly con- vex, comparatively low and long (LMI 36-39), moderately marginate from lateral to dorsal mesosoma; promesonotal suture absent; metanotal groove weak or absent. Propodeal spines reduced to short, triangular, and usually blunt, rarely acute, teeth (PSLI 11-14); propodeal lobes short, triangular, and usually blunt, approximately of same size as propodeal teeth, in profile teeth and lobes not strongly inclined towards each other. Petiolar node in profile high rounded nodiform, around 1.7 to 1.9 times higher than long (LPel 53-59), anterior and posterior faces approximate- ly parallel, usually anterodorsal and posterodorsal margins situated at about same height and equally rounded, petiolar dorsum weakly to moderately convex; node in dorsal view around 1.2 to 1.3 times wider than long (DPel 120-133), in dorsal view pronotum between 2.3 to 2.6 times wider than petiolar node (PeNI 38-43). Postpetiole in profile subglobular, around 1.4 to 1.5 times higher than long (LPpl 67-72); in dorsal view around 1.4 to 1.6 times wider than long (DPpI 142-157), pronotum between 1.7 to 2.0 times wider than postpetiole (PpNI 51-58). Postpe- tiole in profile usually appearing less voluminous than petiolar node, postpetiole in dorsal view around 1.3 to 1.4 times wider than petiolar node (PPI 131-144). Man- dibles completely unsculptured, smooth, and shiny; clypeus longitudinally rugulose with three to seven rugulae, median rugula always present and usually fully devel- oped, one or two mostly entire, rarely broken, lateral rugulae present on each side; cephalic dorsum between frontal carinae irregularly longitudinally rugulose with Taxonomy of Malagasy Tetramorium 149 six to ten rugulae, rugulae usually running from posterior clypeal margin to poste- rior head margin, but mostly irregularly shaped, meandering, interrupted or with cross-meshes; scrobal area mostly unsculptured; lateral head reticulate-rugulose to longitudinally rugulose, posteriorly often partly unsculptured; ground sculpture on head weakly to moderately punctate. Dorsum and sides of mesosoma irregu- larly longitudinally rugulose to reticulate-rugulose, sometimes in parts only very weakly sculptured and relatively smooth; ground sculpture on mesosoma weak to absent. Forecoxae, both waist segments, and gaster fully unsculptured, smooth, and shining. Dorsum of head with numerous pairs of long, fine, standing hairs; dorsal promesonotum usually with two pairs, rarely three or four; propodeum and waist segments without long, standing pilosity; first gastral tergite with short, moderately dense, appressed pubescence in combination with several scattered, long, standing hairs. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae with appressed to decumbent hairs. Head, mesosoma, waist segments and gaster uniformly orange brown contrasting with yellowish to light brown mandibles, an- tennae, and legs. Distribution and biology. 7etramorium sikorae is widely distributed in Madagas- car from Andrambovato in Fianarantsoa in the southeast to Binara in the north, and is also found in the isolated montane forest Ambohijanahary in western Madagascar (Fig. 66). Its main distribution is centered in central-eastern Madagascar in the area from Torotorofotsy and Andasibe-Mantadia north to Betampona and Zahamena. Tetramorium sikorae clearly prefers montane rainforests, rarely lowland rainforests, and the elevational range of 240 m to 1100 m must be taken with caution since it is predominantly found at the higher range limit. Also, 7: sikorae seems to be a leaf litter inhabitant. Discussion. Tetramorium sikorae is easily identifiable within the T° schaufussii spe- cies complex. The character that best separates it from most species is the lack of long, standing pilosity on the waist segments since such pilosity is present in T° merina, T. monticola, T: nassonowii, T. pseudogladius, T. schaufussii, T: scutum, and T. xan- thogaster. Only T. rala and T. obiwan share the lack of pilosity on the waist segments, and sometimes pilosity is present in the latter species. However, 7° sikorae is unlikely to be confused with 7: obiwan for several reasons. First, the latter species is much larger in body size (HW 0.71-0.84; WL 1.00-1.20) than 7! sikorae (HW 0.53-0.61; WL 0.70—0.82). Second, T: obiwan has longer antennal scapes (SI 77 -82) than T. sikorae (SI 70-74). Third, both species appear to live in different microhabitats with T. obi- wan in the lower vegetation and 7: sikorae in leaf litter. Beyond the differing pilosity on the waist segments, 7: sikorae is morphologically very close to T! schaufussii. Since the presence or absence of this pilosity seems fairly stable at species levelwe retain these as separate species. Tetramorium sikorae is also relatively similar to T. cognatum in the T. cognatum complex, but apart from the gastral pilosity diagnostic for the species complexes, both species also differ in antennal scape length. Currently, there seems to be no obvious intraspecific variation in T° sikorae. 150 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Figure 58. 7. sikorae syntype worker of junior synonym T: latior (CASENT0101141). A Body in profile B Body in dorsal view C Head in full-face view. Tetramorium xanthogaster Santschi, 1911 http://species-id.net/wiki/Tetramorium_xanthogaster Figs 40C, 41C, 43D, 45D, 46A, 46B, 47E, 48D, 49A, 59, 66 Tetramorium (Xyphomyrmex) sikorae var. xanthogaster Santschi, 1911:124. [Original spelling of Yetramorium xantogaster justifiably emended to Tetramorium xan- thogaster by Wheeler 1922:1032] [Raised to species by Bolton 1979:139] Taxonomy of Malagasy Tetramorium 151 Type material. Holotype, pinned worker, MADAGASCAR (MJ. de Gaulle) (NHMB: CASENT0101146) [examined]. Non-type material MADAGASCAR: Antananarivo, Réserve Spéciale d’Ambohitantely, Forét d Ambohitantely, 20.9 km 72° NE Ankazobe, 18.22528°S, 47 .28683°E, 1410 m, montane rainforest, 17.-22.1V.2001 (B.L. Fisher et al.); Antana- narivo, Ankokoy Forest, 3 km E Ibity, 20.0675°S, 46.9995°E, 1700 m, Uapaca forest, 4.-14.X1.2008 (ME. Irwin & Rin ha); Antsiranana, Parc National Montagne d’ Ambre, 3.6 km 235° SW Joffreville, 12.53444°S, 49.1795°E, 925 m, montane rainforest, 20.—26.1.2001 (B.L. Fisher et al.); Antsiranana, Parc National Montagne d’Ambre [1st campsite], 12.51444°S, 49.18139°E, 960 m, rainforest, 21.-26.1.2001 (WE. Irwin et al.); Antsiranana, Parc National Montagne d’Ambre, 12.2 km 211° SSW Joffreville, 12.59639°S, 49.1595°E, 1300 m, montane rainforest, 2.-7.1].2001 (B.L. Fisher et al.); Antsiranana, Parc National Montagne d’Ambre, 12.52306°S, 49.17901°E, 1100 m, montane rainforest, 11.[1J.2011 (B.L. Fisher et al.); Antsiranana, Forét de Binara, 9.4 km 235° SW Daraina, 13.26333°S, 49.6°E, 1100 m, montane rainforest, 5.XII.2003 (B.L. Fisher); Antsiranana, Galoko chain, Mont Galoko, 13.59358°S, 48.73157°E, 1100 m, montane forest, 22.I].2013 (B.L. Fisher et al.); Antsiranana, 7 km N Jof- freville [camp 2 of Fisher], 12.33333°S, 49.25°E, 360 m, dry forest, 13.-16.V.2001 (R. Harin’Hala); Antsiranana, R.S. Manongarivo, 14.5 km 220° SW Antanambao, 13.99833°S, 48.42833°E, 1175 m, montane rainforest, 21.X.1998 (B.L. Fisher); Ant- siranana, R.S. Manongarivo 17.3 km 218° SW Antanambao, 14.02167°S, 48.41833°E, 1580 m, montane rainforest, 27.X.1998 (B.L. Fisher); Antsiranana, Parc National de Marojejy, Manantenina River, 28.0 km 38° NE Andapa, 8.2 km 333° NNW Manan- tenina, 14.43667°S, 49.775°E, 450 m, rainforest, 12.-15.X1.2003 (B.L. Fisher et al.); Mahajanga, Réserve Spéciale Marotandrano, Marotandrano 48.3 km S Mandritsara, 16.28322°S, 48.81443°E, 865 m, transition humid forest, 7.XII.2007 (B.L. Fisher et al.); Toamasina, Andranobe, 5.3 km SSE Ambanizana, 15.67133°S, 49.97395°E, 425 m, rainforest, 21.X1.1993 (B.L. Fisher); Toliara, Forét Classée d’Analavelona, 29.2 km 343° NNW Mahaboboka, 22.675°S, 44.19°E, 1100 m, montane rainforest, 18.— 22.11.2003 (B.L. Fisher et al.); Toliara, FForét Classée d’Analavelona, 33.2 km 344° NNW Mahaboboka, 22.64333°S, 44.17167°E, 1300 m, montane rainforest, 22.—25. 11.2003 (B.L. Fisher et al). Diagnosis. The following character set discriminates 7. xanthogaster from the oth- er species of the T° schaufussii complex: moderate to large eyes (OI 22—25); short an- tennal scapes (SI 70-75); frontal carinae weakly developed, only faintly raised, usually becoming much weaker around eye level and fading out halfway between posterior eye margin and posterior head margin; propodeal spines/teeth very short to short (PSLI 10-16), propodeal spines and lobes not strongly inclined towards each other; petiolar node in profile around 1.5 to 1.7 times higher than long (LPeI 59-67) and in dorsal view around 1.2 to 1.3 times wider than long (DPeI 119-133); dorsum of promesono- tum with numerous pairs of long, standing hairs (10+), propodeum usually with one or two pairs, sometimes with up to five; waist segments with long, standing pilosity. 152 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Worker measurements (N=25). HL 0.54—-0.75 (0.64); HW 0.59-0.80 (0.70); SL 0.38-0.53 (0.46); EL 0.13—-0.18 (0.16); PH 0.27—0.41 (0.32); PW 0.39-0.56 (0.46); WL 0.72-1.05 (0.86); PSL 0.06—0.13 (0.09); PTL 0.12-0.19 (0.15); PTH 0.21-0.29 (0.25); PTW 0.16—-0.24 (0.19); PPL 0.14-0.21 (0.18); PPH 0.19-0.29 (0.24); PPW 0.19-0.29 (0.24); CI 92-94 (93); SI 70-75 (72); OI 22-25 (24); DMI 51-55 (54); LMI 36-39 (38); PSLI 10-16 (13); PeNI 37-46 (41); LPel 59-67 (62); DPel 119-133 (126); PpNI 48-56 (51); LPpI 68-80 (75); DPpI 125-140 (135); PPI 117-129 (125). Worker description. Head clearly longer than wide (CI 92-94); posterior head margin weakly concave. Anterior clypeal margin with distinct median impression. Frontal carinae weakly developed, only faintly raised, usually becoming much weaker around eye level and fading out halfway between posterior eye margin and posterior head margin. Antennal scrobes very weak, shallow and without clear and distinct pos- terior and ventral margins. Antennal scapes short, not reaching posterior head margin (SI 70-75). Eyes moderate to relatively large (OI 22—25). Mesosomal outline in profile flat to weakly convex, comparatively low and long (LMI 36-39), weakly to moderately marginate from lateral to dorsal mesosoma; promesonotal suture absent; metanotal groove weakly developed or absent. Propodeal spines/teeth very short to short, vary- ing from triangular and blunt to elongate-triangular and acute (PSLI 10-16), propo- deal lobes short, triangular, and blunt, always much shorter than propodeal spines, spines and lobes not strongly inclined towards each other. Petiolar node in profile high nodiform, nodiform or weakly cuneiform, always with relatively well rounded antero- and posterodorsal margins, around 1.5 to 1.7 times higher than long (LPel 59-67), anterior and posterior faces often approximately parallel and often not, anterodorsal and posterodorsal margins usually at about same height, sometimes anterodorsal mar- gin situated slightly higher than posterodorsal, petiolar dorsum weakly to moderately convex; petiolar node in dorsal view around 1.2 to 1.3 times wider than long (DPel 119-133), in dorsal view pronotum between 2.2 to 2.7 times wider than petiolar node (PeNI 37-46). Postpetiole in profile globular to subglobular, approximately 1.3 to 1.5 times higher than long (LPpI 68-80); in dorsal view around 1.3 to 1.4 times wider than long (DPpI 125-140), pronotum between 1.8 to 2.1 times wider than postpetiole (PpNI 48-56). Postpetiole in profile appearing more or less as voluminous as petiolar node, postpetiole in dorsal view around 1.2 to 1.3 times wider than peti- olar node (PPI 117-129). Mandibles unsculptured, smooth, and shining; generally sculpture on clypeus very much reduced, median area usually unsculptured with one or two weak, irregular, and broken rugulae laterally, very rarely median rugula present but then weak and broken; cephalic dorsum between frontal carinae irregularly lon- gitudinally rugulose with six to eight widely separated rugulae, rugulae running from posterior clypeal margin to posterior head margin, but irregularly shaped, often bro- ken or with cross-meshes, and becoming weaker or fading out towards posterior head margin; scrobal area partly unsculptured, but mostly merging laterally with surround- ing reticulate-rugose to longitudinally rugose sculpture present around eyes, most of lateral head predominantly unsculptured, smooth, and shiny; ground sculpture on Taxonomy of Malagasy Tetramorium 153 head weakly to moderately punctate. Dorsum of mesosoma weakly to moderately lon- gitudinally rugulose, sometimes irregularly so, often rugulae very weak and parts of dorsal mesosoma smooth; lateral mesosoma anteriorly often only weakly sculptured and shiny, katepisternum and lateral propodeum usually irregularly longitudinally ru- gulose to reticulate-rugose, sometimes almost completely unsculptured, smooth, and shiny; ground sculpture on mesosoma usually only weakly developed, mostly absent. Forecoxae unsculptured, smooth, and shining. Waist segments and gaster unsculp- tured, smooth, and shining. Dorsum of head with several pairs of long, fine, standing hairs; dorsum of promesonotum always with more than ten pairs of long, standing hairs, propodeum usually with one or two pairs, sometimes with up to five, rarely without any standing pilosity; waist segments each with several pairs; first gastral ter- gite with short, scarce to moderately abundant, appressed to decumbent pubescence in combination with scattered, long, standing hairs. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae with decumbent to subdecumbent hairs. Body colour variable, usually bicoloured with head and mesosoma of dark brown to blackish colour contrasting with yellowish or light brown appendages, waist segments, and gaster; very rarely only head of dark brown contrasting with yellowish remainder of body, sometimes body uniformly coloured, ranging from yellow to dark brown. Distribution and biology. Tetramorium xanthogaster has a relatively patchy dis- tribution range (Fig. 66). The southernmost locality is Analavelona in the southwest. The next known localities are located in the Central Highlands much further north and east (Ambohitantely, Ankokoy, and Marotandrano). The remaining localities are situated in the northern part of Madagascar (Andranobe, Marojejy, Binara, Montagne d’Ambre, and Manongarivo). Many of these localities are widely separated from each other. Furthermore, it seems that 7. xanthogaster prefers montane rainforests since most collections are from elevations around or above 1000 m, whereas it was only occasionally collected from lower elevations. An explanation for the currently patchy distribution records could be that 7: xanthogaster nests and/or forages in the vegeta- tion since almost all of the available material was collected either from beating low vegetation or Malaise traps. Consequently, we expect that more collecting in the lower vegetation stratum will likely yield more material of this species. Discussion. Despite its pronounced variability in colouration and petiolar node shape, the determination of 7! xanthogaster is fairly straightforward. ‘The species was de- scribed by Santschi (1911) and later redescribed by Bolton (1979) as a bicoloured species with dark head and mesosoma contrasting with the yellowish to light brown remainder of the body. Indeed, the material from the southwest and the Central Highlands is con- sistently strongly bicoloured, and even though one should not overly rely too heavily on body colouration, no other species of the T: schaufussii species group is similarly bicol- oured. However, material from the northern localities shows remarkable diversity in col- ouration. The populations from Andranobe, Marojejy, and Binara are of a fairly bright, yellowish brown. By contrast, the material from Manongarivo and Montagne d’Ambre is in parts bicoloured like the southern populations, partly bicoloured with only the head or mesosoma darker than the rest of the body, or just uniformly light brown to dark 154 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) brown. But even misregarding colouration, T° xanthogaster cannot be misidentified with another member of the complex. Due to the presence of long, standing pilosity on its waist segments it cannot be mistaken for T° obiwan (partly), T: pseudogladius, T: sikorae, or 7! rala, and the broader than long petiolar node (DPeI 119-133) distinguishes it from T. nassonowii (DPel 87—98). In addition, the petiolar node of T: xanthogaster in profile is around 1.5 to 1.7 times higher than long (LPel 59-67), which separates it from T° rala and 7? scutum that have nodes which are 2.0 to 2.2 times higher than long (LPel 45-50). The petiolar nodes of the latter two are also thinly cuneiform to weakly squamiform while the node of T° xanthogaster is variably shaped, but lower and less angled. Tetramorium obiwan possesses well-developed frontal carinae and cephalic sculpture, as well as relative- ly long antennal scapes (SI 77-82), whereas T: xanthogaster has much weaker frontal cari- nae and cephalic sculpture and shorter antennal scapes (SI 70-75). The remaining three species, T: merina, T. monticola, and T: schaufussii, are more difficult to separate from 7! xanthogaster. Tetramorium merina is only found in the central Highlands and co-occurs there with T° xanthogaster. In this region T. xanthogaster is always strongly bicoloured with head and mesosoma very dark brown to black, which contrasts with the yellow to light brown waist segments and gaster. In addition, 7° merina lacks any standing pilosity on the propodeum while 7! xanthogaster usually has long, fine, standing pilosity on the propodeal dorsum. As mentioned above, there are not too many other diagnostic charac- ters to separate these species, but the fact that they co-occur in sympatry in several locali- ties without any intermediate forms supports their heterospecificity. Tetramorium monti- cola is also sympatric with T° xanthogaster, this time in the northern part of Madagascar, where the colouration of the latter species is not reliable. However, in 7: monticola the frontal carinae are more strongly developed and the cephalic dorsum between the frontal carinae has nine to thirteen relatively regularly shaped, mostly unbroken rugae while the frontal carinae of 7: xanthogaster are weakly developed and the cephalic dorsum has only six to eight widely separated, often irregularly shaped rugulae. Also, the propodeal spines of the latter are shorter (PSLI 10-16) than in 7: monticola (PSLI 18—22). The last and most common and abundant species of the complex, T° schaufussii, is also usually found in sympatry with 7° xanthogaster, but both species can be well separated by head shape. The head of T! schaufussii is much longer and thinner (CI 86—90) than in T° xanthogaster (CI 92-94). Additional differentiating characters are the long, standing pilosity on the propodeal dorsum (usually present in 7° xanthogaster but absent in T° schaufussii) and the median clypeal ruga (present in 7° schaufussii but absent in 7: xanthogaster). It has to be noted that 7: xanthogaster is relatively variable in the shape of the petiolar node, which can be high nodiform, nodiform, cuneiform, or in intermediate stages. Ihe node shape varies strongly even within material from the same collection event. Another noteworthy variation can be seen in the material from Analavelona. In contrast to the rest of the material from all other localities it seems that the sculpture on the cephalic dorsum and the mesosomal dorsum is much better developed, and does not differ significantly from other species like 7. schaufussii. However, the frontal carinae are still weaker than in the latter. Taxonomy of Malagasy Tetramorium 155 Ss Figure 59. 7: xanthogaster holotype worker (CASENT0101146). A Body in profile B Body in dorsal view C Head in full-face view. 156 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Revision of the Tetramorium severini species group Tetramorium severini species group Diagnosis. Eleven-segmented antennae; anterior clypeal margin with distinct median impression; frontal carinae strongly developed and long, usually ending shortly before posterior head margin; antennal scrobes present, but weak and without well-devel- oped posterior and ventral margins; anterior face of mesosoma weakly developed; mesosomal outline in profile flat and relatively elongated, only very weakly marginate from lateral to dorsal mesosoma, sides usually rounding onto dorsum; mesosoma rela- tively low (LMI 35-37); propodeal spines long to very long, spinose and acute (PSLI 38-43); propodeal lobes triangular and short; petiolar node in profile high rounded nodiform, in profile around 1.5 to 1.7 times higher than long (LPel 57-69), node in dorsal view around 1.1 to 1.2 times wider than long (DPel 104-121), anterior and posterior faces approximately parallel; postpetiole in profile globular to subglobular; mandibles unsculptured, smooth, and shining; cephalic dorsum with distinct longi- tudinally rugose sculpture; mesosoma laterally, distinctly, irregularly, and longitu- dinally rugose to reticulate-rugose, sculpture on mesosomal dorsum relatively weak, usually consisting of feeble, irregular longitudinal rugulae; waist segments and gaster unsculptured, smooth, and shiny; head with numerous standing, long hairs, meso- soma with one or two long hairs, waist segments and first gastral tergite without any standing pilosity; first gastral tergite with very short and appressed pubescence; sting appendage spatulate. Comments. This group only holds the species 7. severini, which is a common faunal element in most rainforests in eastern Madagascar. Previous to Hita Garcia and Fisher (2011), 7. severini was placed in the T! schaufussii group by Bolton (1979). Despite strong similarities in the shape of the mesosoma and waist seg- ments, we still consider 7. severini distinct enough from all members of the 7. schaufussii group to justify its placement in its own species group. All species of the T. schaufussii group are comparatively small species with very short to moderately long propodeal spines, whereas 7. severini is one of the largest species found in Madagascar outside the 7. tortuosum, T. kelleri, and T. tosii species groups, and possesses very long propodeal spines. Admittedly few more factors argue for the separation, and future reexaminations may come to a different conclusion. How- ever, at present, we believe the similarities between T° severini and the T: schaufussii group are convergent in nature, an idea also supported by unpublished mtDNA data (FHG & BLF, unpublished data). Furthermore, the 7° severini group can be easily distinguished from all other Mala- gasy Tetramorium species and groups by its 11-segmented antennae, long and slen- der mesosoma (LMI 35-37) without distinct margination between lateral and dorsal mesosoma, very long propodeal spines (PSLI 38-43), rounded high nodiform petiolar node, and completely unsculptured waist segments. Taxonomy of Malagasy Tetramorium ise Tetramorium severini (Emery, 1895) http://species-id.net/wiki/Tetramorium_severini Figs 60, 66 Xiphomyrmex severini Emery, 1895:343. [Combination in Tetramorium by Bolton 1979:138] Type material. Holotype, pinned worker, MADAGASCAR, Diego Suarez, 1893 (C. Allaud) (MCSN: CASENT0102078) [examined]. [Note: Emery (1895) did not note the number of specimens in the primary de- scription of the species. He only stated the type locality to be Diego Suarez. Bolton (1979) noted in his redescription that the species was described based on a syntype se- ries located in MCSN and MHNG. However, examination of the material from both collections indicates that only the single type from MCSN is a primary name-bearing type from the type locality mentioned in the primary description (Emery 1895). The material from MHNG consists of one cotype only, which was collected from the Bay of Antongil, and is thus neither part of the type series nor a valid type at all. To in- crease the confusion, the type from MCSN was labelled as type and also as syntype. Nevertheless, since there does not seem to be another primary type in either MCSN or MHNG, we consider the single type from MCSN as holotype.] Non-type material. MADAGASCAR: Antsiranana, Ambato, 14 km W Cap Est, 15°17'28.6"S, 50°20'16.9"E, 150 m, secondary rainforest, (G.D. Alpert et al.); Ant- siranana, Parc National Montagne d’Ambre, 3.6 km 235° SW Joffreville, 12.5344°S, 49.1795°E, 925 m, montane rainforest, 20.—26.1.2001 (B.L. Fisher et al.); Antsiranana, Réserve Spéciale d’Ambre, 3.5 km 235° SW Sakaramy, 12.4689°S, 49.2422°E, 325 m, tropical dry forest, 26.-31.1.2001 (B.L. Fisher et al.); Antsiranana, Parc National Montagne d’Ambre (Petit Lac road), 12.5203°S, 49.1792°E, 1125, rainforest, 12.— 14.V.2001 (R. Harin’Hala); Antsiranana, Parc National Montagne d’Ambre, Créte, 12.5813°S, 49.1337°E, 1110 m, 13.X1.2007 (B.L. Fisher et al.); Antsiranana, Parc Na- tional Montagne d’Ambre, Roussettes, 12.5257°S, 49.1724°E, 1025 m, montane rain- forest, 15.X1.2007 (B.L. Fisher et al.); Antsiranana, Parc National Montagne d’Ambre, Antomboka, 12.5003°S, 49.175°E, 885 m, montane rainforest, 16.X1.2007 (B.L. Fisher et al.); Antsiranana, Parc National Montagne d’Ambre, Antomboka, 12.5127°S, 49.1781°E, 970 m, montane rainforest, 16.-17.X1.2007 (B.L. Fisher et al.); Ant- siranana, Parc National Montagne d’Ambre, Pic Bades, 12.5186°S, 49.1862°E, 900 m, 20.X1.2007 (B.L. Fisher et al.); Antsiranana, Parc National Montagne d’Ambre, 12.5139°S, 49.1778°E, 984 m, montane rainforest, 23.-27.II].2011 (BL. Fisher et al.); Antsiranana, 1 km W Andampibe, Cap Masoala, 15°41'37"S, 50°10'53.4"E, 125 m, rainforest, 30.X1.1993 (G.D. Alpert); Antsiranana, 2.0 km S Andrakata, 14.65°S, 49.7167°E, 520 m, disturbed rainforest, 2.XII.1994 (B.L. Fisher); Antsiranana, Rés. Anjanaharibe-Sud, 14°45'27.9"S, 49°30'36.7"E, 875 m, rainforest, 4.X1.1994 (G.D. Alpert); Antsiranana, Rés. Anjanaharibe-Sud, 6.5 km SSW Befingotra, 14.75°S, 49.5°E, 158 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) 875 m, rainforest, 20.X.1994 (B.L. Fisher); Antsiranana, Betaolana Forest, along Bekona River, 14.53°S, 49.4404°E, 880 m, rainforest, 4.-5.III.2009 (B.L. Fisher); Antsiranana, Betaolana forest, Ambodihazovolabe village along Ambolokopatrika river, 14.5448°S, 49.4516°E, 740 m, disturbed forest patch, 6.[II.2009 (B.L. Fisher); Antsiranana, Forét de Binara, 9.1 km 233° SW Daraina, 13.2633°S, 49.6033°E, 800 m, rainfor- est, 3.X1I.2003 (B.L. Fisher); Antsiranana, Forét de Binara, 9.1 km 233° SW Daraina, 13.2633°S, 49.6033°E, 725 m, rainforest, 4.XII.2003 (B.L. Fisher); Antsiranana, Forét de Binara, 9.4 km 235° SW Daraina, 13.2633°S, 49.6°E, 1100 m, montane rainforest, 5.-6.X1I.2003 (B.L. Fisher); Antsiranana, Forét de Binara, 9.1 km 233° SW Daraina, 13.2633°S, 49.6033°E, 725 m, rainforest, 19.1V.2004 (B.L. Fisher); Antsiranana, Forét de Binara, 9.1 km 233° SW Daraina, 13.2633°S, 49.6033°E, 800 m, rainforest, 20-21. IV.2004 (B.L. Fisher); Antsiranana, Makirovana forest, 14.1707°S, 49.9541°E, 415 m, rainforest, 28.—29.IV.2011 (B.L. Fisher et al.); Antsiranana, R.S. Manongarivo, 10.8 km 229° SW Antanambao, 13.9617°S, 48.4333°E, 400 m, rainforest, 8.—13.XI.1998 (B.L. Fisher); Antsiranana, R.S. Manongarivo, 12.8 km 228° SW Antanambao, 13.9767°S, 48.4233°E, 760-780 m, rainforest, 11.-17.X.1998 (B.L. Fisher); Antsira- nana, Marojejy R.N.I. #12, 14°26'43.2"S, 49°47'8.3"E, 375 m, rainforest, 23.X1.1993 (G.D. Alpert); Antsiranana, Parc National de Marojejy, Manantenina River, 28.0 km 38° NE Andapa, 8.2 km 333° NNW Manantenina, 14.4367°S, 49.775°E, 450 m, rainforest, 12.XI.—12.X11.2003 (B.L. Fisher et al.); Antsiranana, Marojejy National Park, 5 km W Manantenina village, 1st Camp site (Mantella), 14.4382°S, 49.774°E, 487 m, rainforest, 20.XII.2004—7.V.2005 (VM. Rin’Ha); Antsiranana, Parc Nation- al de Marojejy, Manantenina River, 27.6 km 35° NE Andapa, 9.6 km 327° NNW Manantenina, 14.435°S, 49.76°E, 775 m, rainforest, 11.XII.2005 (B.L. Fisher et al.); Antsiranana, Nosy Be, 4 km ESE Andoany (=Hellville), 13.4167°S, 48.3°E, 100 m, rainforest, 1.V.1989 (P.S. Ward); Antsiranana, Ampasindava, Forét d’Ambilanivy, 3.9 km 181° S Ambaliha, 13.7986°S, 48.1617°E, 600 m, rainforest, 4.-9.IJI.2001 (B.L. Fisher et al.); Antsiranana, Nosy Be, Réserve Naturelle Intégrale de Lokobe, 6.3 km 112° ESE Hellville, 13.4193°S, 48.3312°E, 30 m, rainforest, 19.-24.1]].2001 (B.L. Fisher et al.); Antsiranana, 84 km SW Sambava on road to Andapa, degraded forest, 115 m, 17.11.1977 (WL. & D.E. Brown); Antsiranana, 84 km SW Sambava, second- ary forest, 26.1.1991 (G.D. Alpert); Fianarantsoa, Réserve Speciale Manombo 24.5 km 228° Farafangana, 23.0158°S, 47.719°E, 30 m, rainforest, 21.1V.2006 (B.L. Fisher et al.); Fianarantsoa, Ranomafana Nat. Park, Miaranony Forest, 21.25°S, 47.41667°E, 1050 m, montane rainforest, 24.X.1992 (E. Rajeriarison); Fianarantsoa, Forét Classée Vatovavy, 7.6 km 122° Kianjavato, 21.4°S, 47.94°E, 175 m, rainforest, 6.-8.VI.2005 (B.L. Fisher et al.); Fianarantsoa, Forét de Vevembe, 66.6 km 293° Farafangana, 22.791°S, 47.1818°E, 600 m, rainforest, transition to montane forest, 23.—24.IV.2006 (B.L. Fisher et al.); Toamasina, Montagne d’Akirindro 7.6 km 341° NNW Ambinan- itelo, 15.2883°S, 49.5483°E, 600 m, rainforest, 17.-21.I11.2003 (B.L. Fisher et al.); Toamasina, Andranobe, 6.3 km S Ambanizana, 15.6813°S, 49.958°E, 25 m, rainfor- est, 14.1993 (B.L. Fisher); Toamasina, Andranobe, 6.3 km S Ambanizana, 15.6813°S, 49.958°E, 150 m, 15.1993 (B.L. Fisher); Toamasina, F.C. Andriantantely, 530 m, Taxonomy of Malagasy Tetramorium 159 18.695°S, 48.8133°E, rainforest, 4.-7.XJI.1998 (HJ. Ratsirarson); Toamasina, Mon- tagne d’Anjanaharibe, 18.0 km 21° NNE Ambinanitelo, 15.1883°S, 49.615°E, 470 m, rainforest, 8.—12.H1.2003 (B.L. Fisher et al.); Toamasina, Baie d’Antongil, no addition- al collection data; Toamasina, P.N. Mantadia, 18.7917°S, 48.4267°E, 895 m, rain- forest, 25.XI.—1.XII.1998 (AJ. Ratsirarson); Toamasina, Sahafina forest, 11.4 km W Brickaville, 18.8144°S, 48.9621°E, 140 m, rainforest, 14.XII.2007 (B.L. Fisher et al.); Toamasina, Parc National de Zahamena, Tetezambatana forest, near junction of Nos- ivola and Manakambahiny Rivers, 17.743°S, 48.7294°E, 860 m, rainforest, 18.11.2009 (B.L. Fisher et al.); Toamasina, Parc National de Zahamena, 17.7336°S, 48.7262°E, 950 m, rainforest, 19.11.2009 (B.L. Fisher et al.); Toamasina, Parc National de Zaha- mena, Sahavorondrano River, 17.7526°S, 48.8573°E, 765 m, rainforest, 23.11.2009 (B.L. Fisher et al.); Toliara, Res. Andohahela, 5 km NNW Isaka-Ivondro, 24°45'32"S, 46°51'15.2"E, 235 m, 3.111992 (E. Rajeriarison); Toliara, Res. Andohahela, 6 km SSW Eminimimy, 24°44'35"S, 46°48'19"E, 330 m, 4.11.1992 (£. Rajeriarison); Toli- ara, Res. Andohahela, 10 km NW Enakara, 24.5667°S, 46.8167°E, 400 m, rainforest, 25.X1.1992 (B.L. Fisher); Toliara, Parc National d’Andohahela, Manampanihy River, 5.4 km 113° ESE Mahamavo, 36.7 km 343° NNW Tolagnaro, 24.7639°S, 46.7668°E, 650 m, rainforest, 24.1.2004 (B.L. Fisher et al.); Toliara, Parc National Andohahela, Col de Tanatana, 33.3 km NW Tolagnaro, 24.7585°S, 46.8537°E, 275 m, rainforest, 23.X1.2006 (B.L. Fisher et al.); Toliara, Forét lvohibe 55.0 km N Tolagnaro, 24.569°S, 47.204°E, 200 m, rainforest, 3.XII.2006 (B.L. Fisher et al). Diagnosis. Tetramorium severini can be easily distinguished from all other Mala- gasy Tetramorium by the following character combination: 11-segmented antennae; long and slender mesosoma (LMI 35-37) without distinct margination between lateral and dorsal mesosoma; very long propodeal spines (PSLI 38—43); rounded high nodi- form petiolar node; weakly sculptured mesosomal dorsum; completely unsculptured waist segments; very dark brown to black colouration. Worker measurements (N=12). HL 0.87—1.08 (0.95); HW 0.82—1.03 (0.89); SL 0.58-0.80 (0.66); EL 0.19-0.24 (0.21); PH 0.40-—0.55 (0.45); PW 0.61-0.75 (0.65); WL 1.14-1.48 (1.24); PSL 0.35—-0.47 (0.39); PTL 0.20-0.24 (0.22); PTH 0:34-0742+(0.36) PTW 023-029 :(0:25)«PPE 0:27-0:35: (0:30) PPH 0:33-0:42 (0.36); PPW 0.30-0.39 (0.33); CI 92-96 (94); SI 69-78 (73); OI 22—24 (23); DMI 51-54 (52); LMI 35-37 (36); PSLI 38-43 (41); PeNI 35-41 (38); LPel 57-69 (62); DPel 104-121 (111); PpNI 47—54 (50); LPpI 79-91 (83); DPpI 103-117 (110); PPI 125-3145: (132). Worker description. Head distinctly longer than wide (CI 92—96); posterior head margin weakly concave. Anterior clypeal margin with distinct median impression. Frontal carinae strongly developed, diverging posteriorly, and usually ending shortly before posterior head margin; antennal scrobe present, but weak, shallow, and without defined posterior or ventral margins. Antennal scapes short, not reaching posterior head margin (SI 69-78). Eyes of moderate size (O] 22—24). Mesosomal outline in profile flat, relatively low and elongated (LMI 35-37), only weakly marginate from lateral to dorsal mesosoma; promesonotal suture and metanotal groove usually present, 160 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) but relatively weak. Propodeal spines very long, spinose, and acute (PSLI 38-43); pro- podeal lobes short, triangular, and blunt, always much shorter than propodeal spines. Petiolar node in profile high, rounded nodiform, with well-rounded antero- and pos- terodorsal margins, around 1.5 to 1.7 times higher than long (LPel 57-69), anterior and posterior faces approximately parallel, generally anterodorsal and posterodorsal margins situated at about same height (sometimes anterodorsal margin higher than posterodorsal margin), petiolar dorsum always noticeably rounded and convex; node in dorsal view around 1.1 to 1.2 times wider than long (DPel 104-121), in dorsal view pronotum between 2.4 to 2.8 times wider than petiolar node (PeNI 35-41). Postpe- tiole in profile globular to subglobular, approximately 1.1 to 1.3 times higher than long (LPpI 79-91); in dorsal view between 1.0 to 1.2 times wider than long (DPpl 103-117), pronotum around 1.8 to 2.1 times wider than postpetiole (PpNI 47-54). Postpetiole in profile usually appearing thicker and slightly lower than petiolar node, postpetiole in dorsal view between 1.2 to 1.4 times wider than petiolar node (PPI 125-143). Mandibles variably sculptured, ranging from fully unsculptured, smooth, and shining, through partially striate, especially basally, to fully covered by fine striae; clypeus longitudinally rugose/rugulose, with four to eight rugae/rugulae, median ruga always very well developed and distinct, lateral rugulae usually much weaker and usu- ally broken; cephalic dorsum between frontal carinae with seven to nine thick, longi- tudinal rugae, rugae running mostly unbroken from posterior clypeal margin to poste- rior head margin, rarely interrupted or with cross-meshes; scrobal area usually mostly unsculptured, rarely longitudinally rugulose to reticulate- rugulose; lateral head longi- tudinally rugulose to reticulate- rugulose, often with unsculptured and smooth areas. Mesosoma laterally irregularly longitudinally rugose to reticulate-rugose; sculpture on mesosomal dorsum relatively weak, usually consisting of feeble, irregular longitudinal rugulae, often with unsculptured and smoother areas. Ground sculpture on head and mesosoma weakly to moderately punctate Forecoxae rugulose to reticulate-rugulose. Waist segments and gaster completely unsculptured, smooth. and shining. Head with numerous long, standing hairs, mesosoma with one or two long hairs, waist segments and first gastral tergite without any standing hairs at all; first gastral tergite with very short, moderately scattered, and appressed pubescence. Anterior edges of antennal scapes and dorsal (outer) surfaces of hind tibiae with appressed hairs only. Body very dark brown to black, appendages often slightly lighter. Distribution and biology. Tetramorium severini is widely distributed in the rainforests and montane rainforests of eastern and northern Madagascar (Fig. 66). The distribution ranges from Andohahela in the southeast to Montagne d’Ambre on the northern tip of the island, and from there south to Nosy Be, Ampasindava, and Manongarivo on the northwestern side. Throughout its distribution, T° severini is found at altitudes ranging from 25 to 1125 m. In addition, it seems that the species lives in leaf litter and/or the ground. Discussion. As mentioned above, 7. severini is the only member of its species group, and a prominent element of the Malagasy Tetramorium fauna. ‘The large body Taxonomy of Malagasy Tetramorium 161 Figure 60. 7° severini holotype worker (CASENT0102078). A Body in profile B Body in dorsal view C Head in full-face view. size, very dark colour, very long propodeal spines, reduced sculpture on the mesosoma, and complete lack of sculpture on the waist segments render it immediately recognis- able. Considering its wide distribution and large number of specimens examined, T° severini displays very little intraspecific variation. 162 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Tetramorium alperti Tetramorium dalek Tetramorium enkidu Tetramorium gilgamesh Tetramonum naganum Figure 61. Geographic distribution maps for the species of the 7: naganum species group. Star symbols represent type localities while circles represent non-type localities. Taxonomy of Malagasy Tetramorium 163 Tetramorium bressleri Tetramorium gollum Téetramonum hobbit fetramorium plesiarum Figure 62. Geographic distribution maps for the species of the 7: plesiarum species group. Star symbols represent type localities while circles represent non-type localities. 164 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Tetramorium aspis Tetramorium camelliae Tetramonum cognatum Tetramorium freya Tetramorium gladius Tetramorium karthala Figure 63. Geographic distribution maps for the species of the T° cognatum species complex I. Star sym- bols represent type localities while circles represent non-type localities. Note that 7: karthala is endemic to the island of Grand Comore and not found on Madagascar. Taxonomy of Malagasy Tetramorium 165 Tetramonum myrmidon Tetramorium proximum Tetramonum rumo Tetramorium tenuinode Figure 64. Geographic distribution maps for the species of the 7: cognatum species complex II. Star symbols represent type localities while circles represent non-type localities. 166 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Tetramorium merina Tetramorium monticola Tetramorium nassonowii Tetramonum obiwan Tetramorium pseudogladius Figure 65. Geographic distribution maps for the species of the 7. schaufussii species complex I. Star symbols represent type localities while circles represent non-type localities. Taxonomy of Malagasy Tetramorium 167 Tetramorium schaufussii Tetramorium scutum Tetramorium sikorae Tetramorium severini Figure 66. Geographic distribution maps for the species of the T: schaufussii species complex II and the T. severini species group. Star symbols represent type localities while circles represent non-type localities. Note that T° schaufussii is also recorded from the island of Reunion, but we omit it from the map due to questionable locality data. 168 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Acknowledgements First, we would like to thank Michele Esposito from CAS for her enduring support with databasing, imaging processing, proofreading, and her overall support in the lab. We are also very thankful to the current and past members of the CAS imaging crew involved in creating the montage images used in this study: April Nobile, Michele Esposito, Erin Prado, Estella Ortega, Shannon Hartman, Zachary Lieberman, Wil- liam Ericson, Ryan Perry, and Cerise Chen. Zachary Lieberman also helped by provid- ing important label and collection information for several specimens from the Forel collection at MHNG. We also appreciate the support from Ms Suzanne Ryder, Ms Natalie Dale-Skye Papilloud, and Dr. Gavin Broad from BMNH, Dr. Stefan Cover from MCZ, Dr. Daniel Burckhardt and Isabell Ziircher-Pfander from NHMB, and Dr. Bernhard Merz from MHNG, who either loaned important type material or wel- comed us to their collections. We also want to thank Dr. Gary D. Alpert from MCZ and Prof. Phil S. Ward from the University of California, Davis, for providing impor- tant additional material collected in Madagascar prior to the Malagasy ant inventory. In addition, we want to express our gratitude to Marek Borowiec and two anonymous reviewers for editing and reviewing the manuscript. Moreover, the fieldwork on which this study is based could not have been completed without the gracious support of the Malagasy people and the Arthropod Inventory Team (Balsama Rajemison, Jean- Claude Rakotonirina, Jean-Jacques Rafanomezantsoa, Chrislain Ranaivo, Hanitrini- ana Rasoazanamavo, Nicole Rasoamanana, Clavier Randrianandrasana, Dimby Ra- harinjanahary). In Comoros, we benefited from the generous help of Yahaya Ibrahim from the CNDRS, Grande Comore. We thank the following institutions for deliver- ing authorizations for the capture, collection and exportation of ants: COMOROS: Centre National de Documentation et de Recherche Scientifique (CNDRS); MADA- GASCAR: Ministére de Environnement et des Foréts and the Madagascar National Parks. This study was supported by the National Geographic grant No. 8429-08 and the National Science Foundation under Grant No. DEB-0072713, DEB-0344731, and DEB-0842395. Finally, FHG was granted two Ernst Mayr Travel Grants from the MCZ to visit the ant collections at BMNH and MCZ. References Bharti H, Kumar R (2012) Taxonomic studies on genus Tetramorium Mayr (Hymenoptera, Formicidae) with report of two new species and three new records including a tramp spe- cies from India with a revised key. Zookeys 207: 11-35. doi: 10.3897/zookeys.207.3040 Blard F, Dorow WHO, Delabie JHC (2003) Les fourmis de Pile de la Reunion (Hymenoptera: Formicidae). Bulletin de la Société Entomologique de France 108: 127-137. Bolton B (1975) A revision of the ant genus Leptogenys Roger (Hymenoptera: Formicidae) in the Ethiopian region. Bulletin of the British Museum (Natural History) Entomology 31: 237-305. doi: 10.1080/00222938500770191 Taxonomy of Malagasy Tetramorium 169 Bolton B (1976) The ant tribe Tetramoriini (Hymenoptera: Formicidae). Constituent genera, review of smaller genera and revision of Triglyphothrix Forel. Bulletin of the British Museum (Natural History) Entomology 34: 281-379. Bolton B (1977) The ant tribe Tetramoriini (Hymenoptera: Formicidae). The genus Tetramorium Mayr in the Oriental and Indo-Australian regions, and in Australia. Bulletin of the British Museum (Natural History) Entomology 36: 67-151. Bolton B (1979) The ant tribe Tetramoriini (Hymenoptera: Formicidae). The genus Tetramorium Mayr in the Malagasy region and in the New World. Bulletin of the British Museum (Natural History) Entomology 38: 129-181. Bolton B (1980) The ant tribe Tetramoriini (Hymenoptera: Formicidae). The genus Tetramo- rium Mayr in the Ethiopian zoogeographical region. Bulletin of the British Museum (Nat- ural History) Entomology 40: 193-384. Bolton B (1985) The ant genus Triglyphothrix Forel a synonym of Tetramorium Mayr. (Hyme- noptera: Formicidae). Journal of Natural History 19: 243-248. Bolton B (2014) An online catalog of the ants of the world. http://antcat.org [accessed 27 January 2014] Cs6sz S, Radchenko A, Schulz A (2007) Taxonomic revision of the Palaearctic Tetramorium chefketi species complex (Hymenoptera: Formicidae). Zootaxa 1405: 1-38. Csdsz S$, Schulz A (2010) A taxonomic review of the Palaearctic Tetramorium ferox species complex (Hymenoptera, Formicidae). Zootaxa 2401: 1-29. Emery C (1895) Mission scientifique de M. Ch. Alluaud dans le territoire de Diego-Suarez (Madagascar-Nord). (Avril-aodt 1893). Annales de la Société entomologique de Belgique 39: 336-345. Evenhuis NL (2013) The insect and spider collections of the world website [online]. http://hbs. bishopmuseum.org/codens [accessed 12 December 2013] Fischer G, Fisher BL (2013) A revision of Pheidole Westwood (Hymenoptera: Formicidae) in the islands of the Southwest Indian Ocean and designation of a neotype for the invasive Pheidole megacephala. Zootaxa 3683: 301-356. doi: 10.11646/zootaxa.3683.4. 1 Fisher BL (2005) A model for a global inventory of ants: A case study in Madagascar. Proceed- ings of the California Academy of Sciences 56: 86-97. Forel A (1891) Histoire naturelle des Hyménoptéres. Deuxiéme partie: Les Formicides. In: Grandidier A (Ed) Histoire Physique, Naturelle et Politique de Madagascar. L’Imprimerie Nationale, Paris, 1-280. Forel A (1892) Nouvelles espéces de formicides de Madagascar. (Récoltées par M. Sikora). Annales de la Société entomologique de Belgique 36: 516-535. Giisten R, Schulz A, Sanetra M (2006) Redescription of Tetramorium forte Forel, 1904 (Insec- ta: Hymenoptera: Formicidae), a western Mediterranean ant species. Zootaxa 1310: 1-35. Hita Garcia F, Fischer G, Peters MK (2010) Taxonomy of the Tetramorium weitzeckeri species group (Hymenoptera: Formicidae) in the Afrotropical zoogeographical region. Zootaxa 2704: 1-90. Hita Garcia F, Fisher BL (2011) The ant genus Tetramorium Mayr (Hymenoptera: Formici- dae) in the Malagasy region—introduction, definition of species groups, and revision of the T. bicarinatum, T. obesum, T. sericeiventre and T. tosii species groups. Zootaxa 3039: 1-72. 170 Francisco Hita Garcia & Brian L. Fisher / ZooKeys 413: 1-170 (2014) Hita Garcia F, Fisher BL (2012a) The ant genus Tetramorium Mayr (Hymenoptera: Formici- dae) in the Malagasy region—taxonomic revision of the T: kelleri and T. tortuosum species groups. Zootaxa 3592: 1-85. Hita Garcia F, Fisher BL (2012b) The ant genus Tetramorium Mayr (Hymenoptera: Formicidae) in the Malagasy region—taxonomy of the T: bessonii, T. bonibony, T. dysalum, T. margina- tum, T. tsingy, and T. weitzeckeri species groups. Zootaxa 3365: 1-123. Hita Garcia F, Fisher BL (2013) The Tetramorium tortuosum species group (Hymenoptera, Formicidae, Myrmicinae) revisited—taxonomic revision of the Afrotropical T: capillosum species complex. Zookeys 299: 77-99. doi: 10.3897/zookeys.299.5063 R Core Team (2014) R: A Language and Environment for Statistical Computing. R Founda- tion for Statistical Computing, Vienna, Austria. http://www.R-project.org [accessed 28 January 2014] Roberts DL, McGlynn TP (2004) Tetramorium insolens Smith (Hymenoptera: Formicidae): a new record for Mauritius, Indian Ocean. African Entomology 12: 265-267. Santschi F (1911) Nouvelles fourmis de Madagascar. Revue Suisse de Zoologie 19: 117-134. Santschi F (1926) Description de nouveaux Formicides éthiopiens (II[me partie). Revue de Zoologie Africaine (Bruxelles) 13: 207-267. Sharaf M, Aldawood SA, Taylor B (2012) A new ant species of the genus Tetramorium Mayr, 1855 (Hymenoptera: Formicidae) from Saudi Arabia, with a revised key to the Arabian species. PLoS ONE 7: e0030811. doi:10.1371/journal.pone.003081 1 Wheeler WM (1922) Ants collected by the American Museum Congo Expedition. A con- tribution to the myrmecology of Africa. Bulletin of the American Museum of Natural History 45: 1-1055. Wilson EO (1955) A monographic revision of the ant genus Lasius. Bulletin of the Museum of Comparative Zoology 113: 1-201.