Revision of the myrmicine genus Lophomyrmex, with a review of its taxonomic position (Hymenoptera: Formicidae).

14 Pages·1994·0.56 MB·English

by  Rigato, F.

Systematic Entomology (1994) 19, 47-60
Revision of the myrmicine ant genus Lophomyrmex,
with a review of its taxonomic position (Hymenoptera:
Formicidae)
FABRIZIO RIGATO
c/o Department of Entomology, The Natural History Museum, London
Abstract. The myrmicine ant genus Lophomyrmex is fully revised and trans-
ferred from tribe Pheidologetonini to Pheidolini (tribal transfer: Pheidolini,
seniorsynonym of Lophomyrmecini syn.n.). Ten species are recognized, which
are placed into two species groups. The quadrispinosm group, characterized by
thepresence ofpronotal teethorspines, includes birmanus, kalisp.n., opaciceps
stat.n. (=iavana syn.n.), quadrispinosus {=taprobanae syn.n.) and taivanae
stat.n. The bedoti group, in which the pronotum is dorsolaterally marginate,
includes ambiguus sp.n., bedoti, longicomis sp.n., lucidus stat.n. and striatulus
A
sp.n. key to the worker caste is presented and descriptions of known sexual
forms are given.
Introduction basalmargin, byanuninterruptedseriesofblunt, irregular
denticles; (b) anterior clypeal margin with a median
Lophomyrmex isasmall, well-defined. Oriental and Indo- anterioriy protruding point. Further, workers have the
Australian genus. These ants have been generally over- pronotumeitherwithdorsolateral,irregular,sharpborders
looked,althoughsomeLophomyrmexspeciesarecommon or with anterolaterally directed dorsal teeth, which can
ground dwellers and surface scavengers in secondary also be interpreted as an autapomorphic trait.
forest. Moffett (1986) briefly surveyed the foraging be- Few taxonomistshave treated thisgenus in the past and
haviour of L.bedoti and L.opaciceps in Malaysia and some subspecific forms {L.bedoti var. lucidus Menozzi,
Indonesia. Nests of Lophomyrmex are usually located L.quadrispinosus var. opaciceps Viehmeyer and L.
near the base of trees and seem moderately populous. quadrispinosus subsp. taivanae Forel) are actually good
Lophomyrmex workers forage following narrow trunk species. Others (L.quadrispinosus var. taprobanae Forel
trails from which they depart singly to collect food. The and L.quadrispinosus var. javana Karavaiev) are treated
more persistent pathways are surrounded by walls built as junior synonyms. The genus is divided in two equally-
with soil or sand particles; sometimes the trails are wholly sized groups; the quadrispinosus-group (birmanus, kali,
subterranean and undetectable. The diet is heterogeneous opaciceps, quadrispinosus and taivanae), whose species
and includes many kinds ofdead and living invertebrates; have two anterodorsal pronotal teeth, and the bedoti-
isopods. arachnids, termites, cockroaches, flies, larvae group{ambiguus, bedoti, longicomis,lucidusandstriatulus),
of various insect groups, as well as other ants. Moreover where the pronotum is sharply and irregularly dorsolater-
Lophomyrmex workers are enticed by sugar baits and ally marginated. Besides this major diagnostic character,
vegetable oil, but not at all byseeds. The peculiar mandibu- Lophomyrmex seems rather constant in all the other
lardentition ofLophomyrmexappearswell suited forcut- features.
tingbothpreyandthelimbsofforeignants; Lophomyrmex Some species (ambiguus, longicomis, lucidus, opaciceps
are well adapted for hunting other invertebrates notwith- and taivanae) seem tohave a ratherrestricteddistribution,
standing their reduced sting. but others (bedoti, birmanus and quadrispinosus) are
Lophomyrmex workers are small, slender, yellowish widespread over large areas. L.kali and L.striatulus were
or brownish ants; both workers and females are easily collected injustone localityand theiractual range can not
identifiable by means of the presence of the following be confidently established.
autapomorphies: (a) mandibles armed with an apical and Lophomyrmex has been placed in Pheidologetonini
one preapical tooth, followed, on both masticatory and since Emery (1922) and Wheeler (1922), but both authors
feltthistribalplacementwasprovisional. Neverthelessthe
Correspondence: Dr F. Rigato. viale Romagna 11, 20092 taxonomic position of the genus has remained unchal-
Cinisello Balsamo (Milano), Italy. lenged. In myopinion this treatment isveryunsatisfactory
47
48 Fabrizio Rigato
and, asIwilldiscusslater, Lophomyrmexis more properly (1) Palp formula 2,2.
placed in the tribe Pheidolini, close to Pheidole. (2) Mandibles with the following dentition: masticatory
marginwithapicalandone preapicaltooth, followedbyan
enlarged denticle; these then followed by a continuous
Measurements and indices series of irregular small denticles, usually with a larger
one at about the midlength of the row; basal margin
TL, HW, HL, CI. SL, SI, PW and AL are standard always finely serrated, continuing the denticulation of the
measurements in ant taxonomy (see Bolton 1987, for masticatory margin (Fig. 2).
definitions). (3) Clypeus vaulted in the middle, with a protruding
Two more measurements and one index, which are blunt tooth at the midpoint ofitsanteriormargin (Fig. 3).
useful for defining some species, have been added: At each side ofthis tooth there is a strongseta, anteriorly
Propodeal spine length (SpL): with the spine in full directed and slightly curved downward. The clypeus is
lateral view (slightly oblique to the body axis), it is the posteriorly broadly inserted between the frontal lobes
straight dorsal distance from the base of the spine to its (wider than a single lobe at this point).
apex. The base of the spine was arbitrarily considered as (4) Frontal lobes weakly converging anteriorly; frontal
the midpoint of the concavity where propodeal dorsum carinae and antennal scrobes absent; frontal triangle
and spine meet (Fig. 1). rather well defined.
Hind tibia length (HTL): the length of the hind tibia (5) Antenna 11-segmented with a distinct 3-segmented
excluding its more proximal portion, i.e. the 'knee' which club, which is longer than the rest of the funiculus.
isconcealed by the apex ofthe femur when the leg is fully (6) Eyes at about the midlength of the head in full-face
straightened. view; oval in profile, with a slight antero-ventral point.
Tibial index (TI): (HTL/HW) x 100. (7) Pronotum domed in profile, with the dorsum quite
All the measurementsare in millimetres andwere taken flattened and the humerus bearing a single erect hair. In
by means of a Wild M8 stereomicroscope with an ocular dorsal view the sides of the upper face may be either
graticule. irregularlymarginateorarmedanteriorlywithahorizontal
flat tooth or spine formed by a lateral and an anterior
edge. When present the teeth are outward and forward-
Depositories directed.
(8) Promesonotal suture vestigial. Mesonotum in lateral
ANIC: AustralianNational InsectCollection,C.S.I.R.O., view gradually sloping toward the propodeum, it bears
Canberra, A.C.T., Australia. around its midlength a prominence, sometimes very low.
BMNH: The Natural History Museum, London, U.K. (9) Metanotal groove well defined, slightly to deeply
DZTAU: DepartmentofZoology, Tel Aviv University, impressed in profile.
Tel Aviv, Israel. (10) Metapleural gland present.
lEB: Istituto di Entomologia 'Guido Grandi', Bologna, (11) Propodeum bispinose, spines well developed
Italy. and long.
IZUAS: Institute of Zoology, Ukrainian Academy of (12) Propodeal spiracle in side viewcircularand located
Sciences, Kiev, Ukraine. near the base of the spine.
MCSNG: Museo Civico di Storia Naturale, Genoa, (13) Propodeal lobes indistinct.
Italy. (14) Metasternal process present, low and rounded.
MCZ: Museum ofComparative Zoology, Harvard Uni- (15) Posterior portion of the metasternum closed,
versity, Cambridge, Mass., U.S.A. membraneous ligament at the articulation of the petiole
MNHG: Museum d'Histoire Naturelle, Geneva, not visible (Fig. 4).
Switzerland. (16) Tibial spur of middle and hind legs present and
MNHU; Museum fiir Naturkunde an der Humboldt simple, not clearly visible.
Universitat zu Berlin, Germany. (17) Petiole with a distinct, fairly long, peduncle, and
UCD: University of California, Davis, Calif., U.S.A. with a high node which isnot triangularinshape in profile.
(18) Petiolar spiracle placed at the midlength of the
peduncle.
Lophomyrmex Emery (Figs 1-28) (19) Postpetiole wider and usually slightly higher than
the petiole; the tergite much more developed than the
Lophomyrmex Emery, 1892: 114. Type-species: Oeco- sternite.
doma quadrispinosa Jerdon, 1851: HI, by monotypy. (20) First gastral tergite large, but ventrally not exten-
sively overlapping the sternite. Anterior corners of the
gaster. at the sides of the postpetiolar insertion, obtuse,
Diagnosis not protruding anteriorly.
(21) Sting reduced.
Worker (Fxp. 1-4. 9-28) (22) Sculpture usually poorly developed, mostly reticu-
Monomorphic, terrestrial myrmicine ants, with the late; head, pronotum andgaster usuallyshining, at least in
following combination ofcharacters. part. Mandiblesstriate at leaston theirbasal half. Clypeus
.
Revision ofLophomyrmex 49
finely shagreened or reticulate with posteriormost median (12) Forewings with one cubital cell and m-cu vein
portion smooth and shining. Colour varying from pale (Fig. 6); not hyaline: either uniformly infuscated (quadri-
brownish yellow to medium brown. spinosus,bedoti)orweaklyyellowishthroughout(lucidus).
(23) Long erect hairs present throughout the body
excluding the dorsum of the propodeum. Pubescence and Male (based on quadrispinosus, bedoti and lucidus
oblique tiny hairs sparse on the body; appendages with (Figs 7-8).
abundant, long, subdecumbent to oblique pubescence. (1) Palp formula 2,2.
Eyes covered with very small erect hairs, usually hooked (2) Head (including the eyes) wider than long (Fig. 7).
at their tip. (3) Mandibles moderately developed, but not touching
each other when fully closed; armed with a single apical
Female (basedonquadrispinosus, bedoti, lucidusandan tooth, which maybesometimesworn out (quadrispinosus,
unassociated specimen) (Figs 5-6). bedoti) or serrated on its masticatory and basal margin
Much larger, darker and more sculptured than the (lucidus) (Fig. 8). Theanglebetweenthe margins iswidely
worker; mandibles, clypeus and antenna as in the worker. rounded.
The other characters as follows: (4) Clypeusvaulted in the middle, verypoorlyangled at
(1) Palp formula 2,2. the midpoint of the anterior border.
(2) Head either trapezoidal (quadrispinosus, bedoti, (5) Frontal lobes absent. Antennal sockets visible in
lucidus), broader than long and anteriorly narrowed dorsal view.
(Fig. 5), or approximately square in shape (unassociated (6) Frontal triangle large, heavilysculptured and opaque,
female). not welldefined, sometimesdistinctly transversally rugose
(3) Eyes at the midlength of the head in full-face view. (quadrispinosus)
(4) Ocelli well developed, separated from one an- (7) Antenna 13-segmented without a club. Scape short
other by a distance more than their maximum diameter and thick, shorter than any funicular segment excluding
(quadrispinosus, bedoti, lucidus) or approximately equal the first (quadrispinosus) or about as long as these (bedoti
to it (unassociated female). and lucidus). First funicular segment approximately half
(5) Alitrunk much longer than wide. Scutum covering as long as the scape, but not swollen. Other funicular
the pronotum in dorsal view. Parapsidal furrows weak. segments very elongated and of similar length.
Axillae well separated, but linked by a strip that may be (8) Eyes at the midlength of the head sides in full-face
eitherthin orquite broad. Metanotum visible fromabove. view, well separated from the mandibular insertion and
Propodeum bidentate or bispinose. from the occipital head surface.
(6) Propodealspiracle approximately elliptical orcircular; (9) Ocelli not surmounting a turret and separated from
in lateral view its opening is posteriorly directed and one another by much more than their diameter.
sometimes slightly ventrally directed. The spiracle is (10) Alitrunk elongated. Scutum well above the pro-
anteriorly placed: a little above the midheight of the notum, without notauli and with weak but distinct parap-
propodeum and much nearer the anterior suture than the sidal furrows. Axillae small and well separated, linked by
f)Osterior propodeal surface. a very narrow strip (quadrispinosus) or a quite wide one
(7) Petiole pedunculate with the node triangular and (bedoti and lucidus). Metanotum visible from above.
rounded in profile. A weak anterior subpetiolar promi- Propodeum bidentate or weakly bituberculate.
nence present. (11) Propodeal spiracle as in the female.
(8) Postpetiole massive, wider than the petiole in dorsal (12) Petiole pedunculate. Its node broadly triangular
view and broadly articulated to the gaster. with a rounded top in profile, sometimes very low (bedoti
(9) Sculpture as follows: and lucidus).
Mandibles longitudinally striate. Clypeus smooth and (13) Postpetiole in dorsal view much wider than the
shining at least posteriody. Head dull, strongly reticulate, petiole,^thick and broadly attached to the gaster. The
with some overlapping fine, approximately longitudinal, tergite much more developed than the sternite.
rugulation, especially on the genae and frons. Alitrunk (14) Sculpture as follows:
mostly punctured and reticulate. Scutum approximately Headwell sculpturedandopaque, onlythecentre ofthe
subopaque, less smooth than the scutellum. .Mesopleuron clypeussmooth and shining. Alitrunk not as sculptured as
mostly unsculptured and shining. Petiole and postpetiole the head, only the propodeum fullyopaque. Middle ofthe
microreticulate and opaque. Gaster generally shining, scutum slightly shining, scutellum fairly smooth in the
with tiny, dense punctures from which pubescence and middle. Petiole and postpetiole dull, mostly reticulate.
pilosity rise. Gaster smooth.
(10) Bodycolourvarying from light (bedoti. lucidus and (15) Colour from brownish yellow (bedoti and lucidus)
unassociated specimen) to dark brown (quadrispinosus). to dark brown.
(11) Pubescence abundant and dense all over the body (16) Pubescence long and rich throughout the body.
and the appendages, usuallydecumbent tosubdecumbent. Pilosity as in the female. Eyes covered with dense, very
Erect hairs sparse on the dorsum of the body and on the short erect hairs.
ventral surface ofthe head. Eyescovered with dense hairs (17) Wings as in the female.
as in the worker.
50 Fabrizio Rigato
Figs 1-8. 1: Spine length measurement (see text): 2: mandible oiquadrispinosus worker; 3: head o(quadrispinosus worker: 4: ventral
surface of the alitrunk ofbedoti worker; dotted area represents the posterior portion of the metasternum (see text); 5; head ofbedoti
female: 6: wingofbedoti female; 7: head ofquadrispinosus male: 8: mandible oflucidus male. (Pilosity and sculpture omitted).
Taxonomicposition of a single cubital cell in the forewing. Yet Emery recog-
nized Lophomyrmex as a quite distinct genus among
Emery (1922) and Wheeler (1922) thought that Lopho- Pheidologetonini. He placed it in a subtribe Lophomyr-
myrmex was related to Pheidologeton. The main reasons mecini, along with Recurvidris (originally Trigonogaster,
theygaveforthe placementofLophomyrmex in Pheidolo- see Bolton, 1992), because of its 3-segmented antennal
getonini were the 11-segmented antenna and the presence club, in contrast with the other Pheidologetonini which
.
Revision ofLophomyrmex 51
have a 2-segmented club. Both Emery and Wheeler con- petiolarspiracleaboutatthe midlengthofthepeduncle, as
sidered this arrangement quite artificial, but in their fol- in many unrelated Myrmicinae (very anterioriy placed in
lowing papers neither author tried to change the tribal Pheidole, near the base of the peduncle, where it might
posLiattieorn,oWfhLeeolpehrom&yrWmheeexler (1954) studied the larvae of cruebpirteaslenctella (pFlieg.sio6)mo(rpthwioccustbaittea)l;cfelolrsewiinngPhewiidtohlea).siTnghlee
the genera included in Pheidologetonini but theyfailed to latter character is very feeble because the presence of a
gain anyevidenceofstrongaffinitiesbetweenthe larvaeof single cubital cell occurs in many unrelated Myrmicinae;
Lophomyrmex and Pheidologeton. also, Bolton (1982) stated it is present among Pheidolini
Ettershank (1966) was the last author who gave a full and both conditions (one or two cubital cells), linked by
diagnosis of the Pheidologeton genus group. He defined intermediate ones, may occur in the same genus.
this using a series of characters that were rather incon- Lophomyrmex shares with Pheidole, but not with
sistent among the genera: clypeus variable in shape and Pheidologeton, a series of features, mostly of difficult
weakly or not bicarinate; number of antennal segments interpretation: (4) frontal triangle well defined (poorly so
8-12 (with 2-3 segmented club); palp formula from in Pheidologeton): (5) antenna with a 3-segmented anten-
3,2to 1,2;workerspolymorphic,dimorphicorevenmono- nal club (2-segmented in Pheidologetonini); (23) pube-
morphic. He did not furnish any reliable synapomorphies scence on the appendages long, abundant and more or
for a better definition ofthe whole group. less obUque, although some Pheidole have a different
More recent comparisons among pheidologetonine condition(inPheidologetonthepubescenceontheappend-
generawerethoseofKugler(1986),basedonthestructure agesissparseorveryreducedand accompanied byat least
of the sting, and Moffett (1986) based on foraging be- afewlong, standinghairs). MoreoverLophomyrmexhasa
haviour. Kugler (1986) found only some widespread peculiar, autapomorphic mandibular dentition (Fig. 2),
superficial resemblances in the sting structure between which is somewhat comparable with the multidentate
Pheidologeton and Lophomyrmex, due tothe reduction of condition of Pheidole, but not at all with the reduced
the apparatus; moreover Lophomyrmex shares some number of teeth of Pheidologetonini (4-6 teeth, without
pecuUarfeatureswith apparently unrelated genera, above serration), which represents a fully divergent apomorphic
all Recurvidris. Moffett (1986) discovered some weak trait. Otherfeatures oflittle value recall a Pheidole minor
similarities between Pheidologeton and Lophomyrmex in worker: head shape and eye size (eyes are mostly second-
foraging behaviour, but these can easily be interpreted arily reduced in Pheidologetonini).
as convergence. Yet, besides these feeble similarities, there is a strong
Although former authors recognized the weakness of synapomorphyshowingthecloserelationofLophomyrmex
the arguments about the placement of Lophomyrmex with Pheidole(see points7 and8underworkerdiagnosis):
within Pheidologetonini, no one tried seriously to correct pronotum domed and marked posteriorly by a vestigial
its taxonomic position. The tribe Pheidologetonini, as promesonotal suture, and mesonotum gradually sloping
currently interpreted, contains a mixture ofonly superfici- downward and bearingaprominence nearits midlength in
ally related genera. In particular Lophomyrmex seems a profile. The domed pronotum with at least a trace of the
good candidate for being removed from the tribe. promesonotal suture is the major synapomorphic trait
AtthebeginningofhisgenericdiagnosisofLophomyrmex defining Pheidole and its allies and separating them from
Emery (1922) stated: 'appearance of a Pheidole worker'. the other Myrmicinae (Bolton, pers. comm.).
Infact, IthinkthatthecorrectplacementofLophomyrmex Moreover, Lophomyrmex and Pheidole have armed
is near Pheidole and quite far from Pheidologeton. or somewhat marked pronotal sides near the humeri
These three genera share some derived features, which (unarmed, unmargined and without any prominence in
are widespread among Myrmicinae and can also be inter- Pheidologeton and related taxa); but this is perhaps a
preted as convergent traits (numbers are those reported convergent trait, fortheirshape is ratherdifferent, although
under the worker diagnosis): (1) palp formula 2,2, it is itcould^e derivedfrom unpeculiarsmallprotuberancesas
constantinPheidologetonandLophomyrmex,anditoccurs it occurs in most Pheidole species.
also in some Pheidole groups (Bolton, pers. comm.); (5) Thus, Lophomyrmexisadistinctandeasilyrecognizable
antenna 11-segmented, constantly in Lophomyrmex and genus belonging to the Pheidolini (tribal transfer), with
Pheidologeton, rarely in Pheidole: (12)propodealspiracles autapomorphic clypeus, mandibular dentition and dorso-
near the base of the propodeal spine in profile (this con- lateral pronotal sides. Probably, its past placement within
ditionmaybethoughtofasanapomorphywhencompared the Pheidologetonini was due to the widespread use of
with the primitively more anterioriy placed propodeal placingsome moreorless peculiargenerain large 'parking
spiracleofthe Ponerinae; Bolton, pers. comm.); (21) sting tribes', hopingthat someone, sooneror later, could better
reduced. Further, there is another character common to explain their affinities.
thesethreegenera, butwhichat presentcannotbedefined
either as primitive or as derived: (15) in ventral view
posterior metasternal portion closed, obscuring the mem- Key to workers
braneous ligament between alitrunk and petiole (Fig. 4).
I foundjust two weak characters linking Lophomyrmex I Indorsalviewthepronotumwithlateralirregularmarginations
to Pheidologeton and separating them from Pheidole: (18) only(bedotigroup) 2
52 Fabrizio Rigato
- In dorsal view the anterior pronotum with two distinct flat- - SI>84, HL<0.80. Pronotal teeth blunt in dorsal view (Fig.
tened teeth or spines, horizontal and anterolaterally directed 21). Mesonotum, propodeum, petiole and postpetiole faintly
(quadrispinosusgroup)(Figs 19,21,23,25,28) 6 sculptured. Propodeal spines straight (Fig. 20). Postpetiole
2 SI& 100, SpL3^0.34. Pronotum in frontal viewwithout a pair m(Faisgs.iv2e0),.a(pTpariowxainm)ately as long as the petiole in latertaalivvaineawe
ofanteriorerect setae near the midline (Fig. 12); mesonotum
inprofileatmostwithsomethin,short,obliquehairs(Fig. 11). 8 SI and TI= 100. Propodeal spinesslightly upcurved in profile
(Brunei,E.Malaysia) longicornis (Fig.26). (India) kali
- SI< 100, SpL<0.34. Pronotum in frontal view with a pairof - SI and TI< 100. Propodeal spines straight or feebly down-
anteriorerect setae near the midline (Fig. 10); mesonotum in curvedinprofile 9
profilewithstandingstiffhairs(Figs9, 13, 14, 16) 3 9 Pronotal spines strongly diverging anteriorly and with longer,
3 Smoothandshining;onlytheupperportionofthemesopleuron sharpertips (Fig. 25). Moresculptured species: at least meso-
clearlyreticulate.TL<2.8.(W.Malaysia) lucidus thorax, propodeum, petiole and postpetiole clearlyreticulate;
often dorsum of head and pronotum longitudinally rugulose.
- Alattiloena.stTwLhooflteenme>s3othoraxandpropodeumwithdistinctreticu4- Postpetioledistinctlyshorterthanthepetiole(Fig.24)o.p(aJcaivcae)ps.
4 Sculpture less developed: pronotal sides and head surface - Pronotal spines neither so divergent nor with such long and
behindandbelowtheeyesshining, at most faintlysculptured. sharptips(Fig. 23). Lesssculptured: mesonotumand nodesof
Eyessmaller: in lateral view the distance between theiranter- petiole and postpetiole not wholly and clearly reticulate;
ior margin and the mandibular insertion is &1.5 times the head always shining. Postpetiole nearly as long as the petiole
maximumeyelength. Propodealspinesinlateralviewthinner, (Fig.22). (SriLanka,Burma,Thailand) birmanus
appearing longer and very straight (Fig. 9). Petiolar node
clearlyobliquely truncate: the steep anterior face and the flat
dorsummeetatadistinct,sometimesslightlyprotruding,angle
(Fig. 9). (Sri Lanka, India, Burma, Malaysia, Singapore,
Brunei,Indonesia,Philippines) bedoti Lbedoti group
- Sculpturemoredeveloped: pronotalsidesand/orheadsurface
behind and below the eyes always at least superficially sculp- Lophomyrmex amblguus sp.n. (Figs 14, 15)
tured,notfullyshining. Eyeslarger:inlateralviewthedistance
between their anterior point and the mandibular insertion is Holotype worker: TL 3.1, HL 0.80, HW, 0.75, CI 94,
usuallylessthan 1.5 timesthemaximumeyelength. Propodeal SL0.68, SI91, PW0.49, AL0.95, SpL0.23, HTL0.69,TI
spines in lateral view more gradually narrowing towards their
tipsand appearingshorterand not verystraight (Figs 14, 16). 92.
Petiolarnodesomewhatrounded(Figs14, 16) 5 In lateral view the distance between the mandibular
insertion and the anterior point of the eye is about 1.3
5 In lateral view the pronotum costulate with at least one lon- timesthe maximumeyelength. Mesonotum in profile with
gitudinal anterior, short costula running backward from the
feeble prominence and posterior step; both, as usual,
insertion of the humeral seta (Fig. 17). In full-face view head
usually more clearly rugulose in the space contained between marked by a couple of erect hairs. Metanotal groove
the inner margin of the eyes and the frontal lobes. (The rug- distinct in profile. Propodeal spines in profile straight, not
ulaearemoreeasilyvisibleinobliqueview)(Thailand) very long and with a quite wide base from which they
slriatulus gradually taper toward the tips. Petiolar node in profile
- cIonsltautlearealrvuinenwintghebparckownaortdumfrisonmotthleonignisteurdtiinoanlloyfctohsetuhlautmee:ranlo fhaicgehs amnedetbraiteflay rsouubtnrduendcataen.gleI;tspoasntteerriioorrlyantdherseupeisriaonr
seta (Fig. 15). Head usually devoid of rugulae in the space evenly rounded, well-defined, step. Postpetiole not mas-
containedbetweentheinnermarginoftheeyesandthefrontal sive, clearly shorter than the petiole and about as high as
lobes. (India,Nepal) ambigum that. In dorsal view the postpetiole is about 1.6 times
'wider than the petiole.
6 Pronotum with teeth that only slightly protrude (Figs 19. 21).
Clypeus finely reticulate, sublucid except for its pos-
In dorsal view their anterior margins are approximately at
a right-angle to the body axis or even faintly convergent teriormost median portion which is smooth. Frontal
anteriorly; their outer edges are straight or very weakly con- trianglemostlyshagreened,onlypartiallyshining.Cephalic
cave; the teeth do not look suddenly narrowed near their dorsum finely reticulate throughout and quite dull, chiefly
apices 7 on the portions contained between the level of the inner
- Pronotum with spiniform teeth that distinctly protrude (Figs margin ofthe eyes and the frontal lobes: frons sublucid in
23, 25. 28). In dorsal view their anterior margins are mod- the middle. Genae and frontal lobeswith tiny longitudinal
erately to markedly divergent anteriorly; if moderately, then rugulae; those on the frontal lobes look long and frag-
theirouteredgesareoftenclearlyconcaveandtheteethhavea mented and reach the vertex (they are more visible in
narrowerapex(Fig.23) 8 oblique view). Ventral head surface with some anteriorly
7 SI<84, HL often >0.80. Pronotal teeth more or less sharp convergent faint rugulae. Anterior slope of the pronotum
in dorsal view (Fig. 19). Mesonotum, propodeum, petiole and reticulate and sublucid. Pronotal dorsum between the
postpetiole distinctly reticulate. Propodeal spines slightly down- lateral margins quite shining and irregularly sculptured,
curved (Fig. 18). Postpetiole shorterthan the petiole in lateral feebly reticulate with few short faint rugulae anteriorly;
view(Fig. 18). (SriLanka,India.Burma) quadrispinosus a blunt poorly defined longitudinal median carina also
Revision ofLophomyrmex 53
vFtiegwso9f-1th7e.pbreondootuurgnro(u1p2.)9o-Uo0n:g.bcoodrnyipsr:of1i3l:ea(l9)itarnundkfprroonftiallevoifelwucoifdtuhs:ep1r4o-1n5o:tubmod(1y0)prooifibleed(o1n4:) 1a1n-d1c2l:osaeliltartuenrkalprvo,feiwleo(f11t)heanpdrofnroonttuaml
a(1n5d)s(csuelepttuexrte)oomiitatmebd^;8mUusl3.ia(n,d-V17: btohedyscpurlopftiulere(1a6r)ouannddctlhoesehulmaeterraall vhaeiwr ionfsetrhteiopnroisnoshtouwmn)(.17) (see text) oistnatulus. (Minute pilosuy
occurs. Pronotal sides shining, with very superficial longi- faintly reticulate and moderately shining. Gaster shining,
tsuudbilnuaclildy; omreiseonptledeurroenticumloartieon.strMoengsloynorteutimculraettei,culdautlle., vtienruyoussupoernfitchiealwlyhorleetiscuurlaftaec;e.the reticulation is not con-
spPoprsaotcppeeotdbieeoltuewmecerlneetairtclhuyelartseetp,iicnsuelusabtoaepnaodqnuttehh,eedesexiccdlelisuv;dititynh.gePtnehoteidoeslsmeoooantnlhdy sheuaPbdidl.eocsiutmybaesntusuoanl itnhethaeligtemnnuks,. Pmuobreesceanbcuendvaenrty ospnartshee,
54 Fabrizio Rigato
Colour chiefly light brown throughout; appendages sharp tip. Some specimens may have more proximally
slightly lighter. enlargedspines notsothin asusual. Petiole in profile with
a rather narrow peduncle and a distinct high node. This
Additional workers: TL 2.8-3.4, HL 0.70-0.85, HW has a steep anterior face meeting the superior one at a
0.64-0.81, CI 91-96, SL 0.59-0.71, SI 86-92, PW distinct angle; posteriorly the node often has an evident
0.44-0.54,AL0.84-1.03, SpL0.20-0.28,HTL0.61-0.75, rounded step toward the postpetiole.
TI 89-% (7 paratypes and 6 non-paratypes measured). Clypeus finely sculptured, sublucid except for its pos-
As the holotype but varying as follows: teriormost median portion which is smooth. Frontal
Eye distance from the mandibular insertion often 1.4 triangle sublucid. Cephalicdorsumsuperficially reticulate,
times the maximum eye length. Propodeal spines may be rather shining on the frons and vertex, but, mostly in
shghtlybentdownwardinprofile. Sometimesthesculpture largest specimens, sublucid on the lateral portions con-
islessdeveloped, chieflyinsmallestspecimens; the headis tained between the level of the inner margin of the eyes
a little more superficially reticulate and its ventral surface andthe frontal lobes. Genae and frontal lobes longitudin-
maybeshiningandwithoutanyrecognizablefaintrugulae. ally rugulose. Anterior slope of the pronotum clearly
The pronotum may be devoid ofshort rugulae. reticulate and sublucid. Pronotal dorsum between the
Colour often lighter on the sides ofhead and aUtrunk. lateral margins and pronotal sides shining, very superfici-
allysculptured, chiefly reticulate; avery faint longitudinal
Holotype worker, INDIA: Uttar Pradesh, Kuamun median carina also occurs on the pronotal dorsum. Meso-
District, Kathgodam, 600m, 6.x.1979, sifted leaf litter notumsuperficiallyreticulate, more shiningonitsanterior
(/. Lobf) (ANIC). half. Mesopleurae and propodeum, excluding the decliv-
ity, finely reticulate and more or less opaque. Petiole
Paratype workers, 2 specimens with the same data as and postpetiole reticulate on the sides, usually sublucid;
theholotype(BMNH: ANIC);2specimens,INDIA: Uttar top of petiolar node faintly reticulate; dorsum of post-
Pradesh, Dehra Dun, 30.viii.l978 (H. Imai) ('Indian petiole quite smooth anteriorly. Gaster mostly glassy
ants cytology', voucher specimens n°.78—60) (ANIC) smooth, shining.
[misidentified as L.bedoti]; 3 specimens, INDIA: NE Colour usually brownish yellow, darker on the dorsum,
India, without locality and date (Kurr) (BMNH; MCZ). to light brown.
Non-paratypic workers, NEPAL: 16km ENE Baglung, Female: TL about 9.5-10. Ocelli quite wide, but sep-
1100m[p. S. Ward) [with abradeddorsalpilosity] (BMNH; arate from one another by a distance much larger than
UCD). their maximum diameter. Axillae distinctly separate but
Unkedby a narrowstrip. Propodeal teeth short and strong
Comments. L.ambiguus may be confused with L. or spiniform and blunt. Propodeal spiracle directed a
striatulus (see there) and L.bedoti, butit ismore distinctly little backward.
sculptured than the latter. Moreover L.bedoti has slightly Clypeus punctured and sublucid. Head distinctly reticu-
smaller eyes, which are about 1.5 times their maximum late and opaque with superimposed longitudinal rugulae
length from the mandibular insertion in lateral view. The more evidenton the anterior half. Scutumopaque, reticu-
propodeal spines of L.ambiguus look shorter and thicker late. Scutellumclearlyshiningandpoorlysculpturedin the
inprofile. The petiolarnodeofL.ambiguus(Fig. 14)isnot middle. Gaster finely punctured, sublucid.
so clearly obliquely truncate as in L.bedoti (Fig. 9). Also Colour mostly brown, reddish brown on the sides, legs
L.ambiguus is usuallydarker and with adeepermetanotal lighter. Wings uniformly infuscated.
groove.
Male [previously undescribed]: TL about 5.6-6. Man-
.dibles not serrated. Propodeum bituberculate, tubercles
Lophomyrmex bedoti Emery (Figs 4-6, 9-10) widelyrounded. Petioleinprofilewith averylow, rounded
node. Postpetiole a little higher than the petiole.
Lophomyrmex bedoti Emery, 1893: 192. Syntype workers Mandibles finely striate. Clypeus feebly scuptured
and females, INDONESIA: Sumatra, Deli (Bedot) and sublucid. Head finely reticulate, opaque. Alitrunk
(MCSNG) [workers examined]. sculptured about as in the female; but scutellum less
shining and with some longitudinal sculpture pattern.
Worker: TL 2.5-3.5, HL 0.66-0.89, HW 0.60-0.87, Gaster finely punctured and quite lucid.
CI 88-99, SL 0.58-0.75, SI 85-98, PW 0.38-0.58, AL Colour mostly brownish yellow, brown on the posterior
0.75-1.09, SpL 0.21-0.33, HTL 0.55-0.75, TI 83-95 halfofthe head and scutellum, scutum and dorsum ofthe
(162 measured). gaster with an intermediate tinge. Wing slightly lighter
With the characters given in the key and the following: than in the female.
Mesonotum in profile with a very low, barely evident
prominence. Metanotal groove in profile very shallow Comments. A widespread species which sometimes may
without clear anterior and posterior borders. Propodeal be confused with L.ambiguus (see there). The original
spinesinprofile usuallyverystraight, quite thinandwith a locality was reported as Sumatra: Deli; yet Deli is a small
Revision ofLophomyrmex 55
island near the southern coast of the westernmost point notal slope distinctly reticulate and sublucid. Pronotal
ofJava. dorsum between the lateral margins neariyglassy smooth,
withonlyatrace ofa median longitudinalcarina. Pronotal
Material examined. SRI LANKA: Peradeniya (C. T. sides shining, very weakly reticulate. Mesonotum, meso-
ABBiYUnSRgIhMAaA:m):G.nM.IaNyGDmerIyaAoi:,(3GS.0i0kH0k.iftmL,(oCw.Teu)Tk;.vaBKriend(gaChh.a,m)GT..unWBu.inngMghJAeaLrma-)i arpenltediucpruoolsanttpeaetnaidnodlperswooipmtohedwweheuaamtk,sreeutxbicoclupuladaiqtnuigoenioatsrndsshusibunlbiulncugicdi.dde,cPclehitivieioftllyey,
(R. W. Taylor and R. A. Barrett); Georgetown (R. W. on the nodes. Gaster shining with only very superficial
Taylor and R. A. Barrett); Selangor Gombak (D. H. reticulation.
TMUaluurppahGhyo)(m;Db.aJokHh.oFroMeru.rJpaRsheoysn).s;(BRSa.eylCar(naDgz.oireHr).(;C.MSueBrlepatthnsgy)o);r;,Se1Kl0eanpmgoionrEg, atrriuPsniikln.ogsiItanytfartsohnetuasbluoavrlideoewrnttwhhieethphreotahndeo,taubnumttelraricoekrdsuscalenodypeoernencettahresetatlhaiee-
WaterworksRes. {R. Crozier);G. Lawit(T Clay);Penang midline. In profile mesonotum with only tiny hairs or
(M. W. Moffett and D. R. Fletcher); Bukit Rengit nr oblique pubescence marking the prominence and the
Lanchang (//. Imai and Kubota); Bukit Fraser, Jerian posterior step.
W. Fall (//. Imai and Kubota); Pasoh Forest (Watanabe); Colour brownish yellow, head slightly darker.
Negeri Sembilan, Pasoh For. Res. {K. Rosciszewski).
E. MALAYSIA: Sabah, Sepilok For. Res. nr Sandakan Paratype workers: TL 3-3.5, HL 0.75-0.86, HW,
{R. W. Taylor); Sabah, Gn. Silam, 810m {R. Leakey); 0.68-0.81, CI 91-99, SL 0.74-0.84, SI 100-110, PW
Sabah, Tuaran; Sabah, Poring Hot Springs, 500m (/. 0.46-0.55, AL 0.91-1.07, SpL 0.34-0.42, HTL 0.69-
Lobl and D. Burckhardt); Sarawak, Mulu Nat. Park 0.80, TI 92-103 (32 measured).
(M. Collins; P. M. HammondandJ. E. Marshall); Sarawak, As the holotype but varying as follows:
FMnMoiotr.4lSo3RcaeanLlstia.tu,byb,uoIk9In9gRm-di2n9reS4xWK(SuDacKrnhudicHanhakgivaningl(Ra(.(nRRd..)W;.WW.S.aTrTaTaayaywlylaolorkor)r,););;SSSeSaamarreraanawwgwaaaokkk,,h duproporwAfsaialrlefd.a,viPnitreoowrppooaesdnvtedeearnliionsdrpopiwrmnoneefswsioalnenroodttt.haaellyAwasmytmsaeopysrteirbsoeneogvfsllityidegnedhnittvlveyirsmigcebeuldnretivaeiindnn
r38'N-113°36'E, 600ft (A. Emerson). SINGAPO- longitudinalcarinamaybepresentonthepronotaldorsum.
MRaEn:daBiuki(tD.TiHm.ahM(uMr.phyW)..MoBfRfeUtNtEaIn:d DK.uaRl.a FBleetlcahleorn)g; Colour varying from brownish yellow to light brown.
Res.. Temburong (R. Levy). INDONESIA: Kalimantan, Comments. L.longicornis is a very distinct species. The
Pleihari Martapura Res. (A/. W. Moffett and D. R. more constant diagnostic characters are the high SI, the
(F;lVe.tcJhoehrn);sonK)a;liSmuamnattarna,, Lthiomb'eNrgacCaamnptabvia[=BLahloikknpgaap?a]n vonelryyblyonsgompreobpeoddoetailspsepciniemsen(sw)h(oFsieg.l1e1n)g;ththiessahpappreoaofchtehde
A(D/g..osDtBaiuv,ricsI.k)h;LaoSrbudlmt)aa;tnrdaS,uDm.PaatBnrutari,,ck2hn5arr0dmPt)a{;yDaJ.kauvAgamob[sustehie,,'/.C6o0Lmo0mbmelnta(sDn.'d ocpenotmitpohalearraelndiotwdriuetnhkin(aplFlriogtfshiel1e1o,atnh1d2e),r.asTbpheoecvieelsaatltile,nrttfhheeeartgueerdneuucissedu(snpeiieqlouFseiigtisyf
above]. Deli (Bedot). PHILIPPINES: Palawan, 14km S 9, 10 for comparisons).
Puerto Princesa (B. B. Lowery).
Holotype worker, EAST MALAYSIA: Sarawak, 4th
Lophomyrmex longicornis sp.n. (Figs 11, 12) rDaiivn.foGr.estMulleaufNliattt.er,Pa6r.xk.,1R97G7S(SE.xpBno.ltLoon)ng(BPaMlNa,H)l.owland
T0.I7H79o,9l.oStIyp1e03w.orPkWer:0.T4L9.3.A1L, H0L.980,.80S,pHLW0.309.,75,HCTIL94 7S4L thePahroaltoyjpyepe;wo6rksepresc,im1e9nss,peEc.iMmeAnLsAwYitShItAh:e sSaarmaewadka,ta4tahs
Mesonotum in profile with a faint prominence and MDi.v.ColSlaiwnasi);F3osr.pecRiesm.e,ns4,.iEi.i.Ml9A78L,ArYaSinIfAor:estSasboailh,coTraewa(uN,.
withouta posteriorstep. Metanotalgroove veryshallow in 13.xii.l981 (J. Waage); 2 specimens, Sabah, Gn. Silam
aprnodfilaec.uteP,rorpeoadcehailngsptihneeslevienl porfoftihlee pveetriyolelo-npgo,stpstertaiioglhet 2Su5l0amng1n9r83La(mRu.niLne,akBeyM);19382sp-e3c8i8m,en2s0.,viBiiR/U10N.EixI.:l98B2uTk9it/
articulation; they are also strongly divergent in dorsal 7, fogging (A^. E. Stork) (BMNH; MCZ: ANIC; UCD).
view. Petiolar node in profile quite domed, with a mod-
eratelydistinctangle between itsanterioranddorsal faces. Non-paratypic material: 2 damaged specimens with the
PosCtlpyepteiuoslewneoatklmyassreitviec,ulcalteea,rlsyubslhuocritderextcheapntthfeorpeittsioploes.- sSaarmaewadka,taBoasngtoheMth.ol(oJt.ypHee;wit1t)w.orker, E. MALAYSIA:
teriormost median portion which is smooth and shining.
Frontal triangle irregulariy sculptured and sublucid. Head
srehtiinciunlga,tec.hiGefelnyaoenwtihtehftrionnyslaonndgivteurdtienxa,lvreurgyulsauep.erfFircoinatlally Lophomyrmex lucidus Menozzi stat.n. (Figs 8, 13)
lobes with faint longitudinal rugulae only. Anterior pro- Lophomyrmex bedoti var. lucidus Menozzi, 1930: 328.
56 Fabrizio Rigato
Syntype workers, W. MALAYSIA: Penang, Georgetown Lophomyrmex strlatulus sp.n. (Figs 16, 17)
(H. Eidmann) (lEB) [examined]. Holotypeworker:TL2.8, HL0.73, HW0.69, CI95, SL
Worker: TL 2.5-2.7, HL 0.65-0.70, HW 0.61-0.65, 0.61, SI 88, PW 0.45, AL 0.83, SpL 0.24, HTL 0.60, TI,
CI 92-97, SL 0.54-0.59, SI 86-90, PW 0.39-0.44, AL 87.
0.75-0.82, SpL 0.23-0.26, HTL 0.53-0.58, TI 84-89 In lateral view the distance between the mandibular
(10 measured). insertion and the anterior point of the eye is about 1.3
With the characters given in the key and the following: timesthe maximumeyelength. Mesonotum inprofilewith
Mesonotum in profile with a feeble or faint prominence. a low prominence and a faint posterior step. Metanotal
Metanotal groove shallow in profile. Propodeal spines in groove very shallow in profile. Propodeal spines in profile
profile straight or slightly curved downward, quite thin rather straight and sharp; their superior border weakly
and very sharp. Petiolar node in profile high and usually convex in the basal half. Petiolar node in profile very
subtruncate. briefly subtruncate and with a rounded angle between its
Clypeus somewhat shagreened and subopaque or sub- anterior and dorsal faces; posterior slope distinct. Post-
lucid, exceptforitsposteriormost medianportion whichis petiole not massive and shorter than the petiole.
smooth. Frontal triangle shining. Head shining, with only Clypeus feebly shagreened and sublucid, except for its
a very superficial, quite indistinct, reticulation. Genae posteriormost median portion which is smooth. Frontal
with some short rugae. Frontal lobes finely longitudinally triangle weakly sculptured and sublucid. Head distinctly
irregularlyrugulose.Wholedorsumofthealitrunkshining, reticulate throughout, subopaque; frons and vertex more
at most very superficially reticulate. Sides of the alitrunk superficially sculptured and sublucid. Genae and frontal
with few sculptured areas: anterior upper part of meso- lobeswithweaklongitudinalrugulae. Ventralheadsurface
pleuron and metanotal groove; the rest is smooth with at with anteriorly convergent rugulae. Anterior pronotal
most a wide-meshed faint reticulation. Petiole and post- slopeclearlyreticulate,withsomeshorttransverserugulae
petiole chiefly shining and superficially reticulate mostly on the sides. Pronotal dorsum with a weak longitudinal
on the sides. Gaster glassy smooth, shining. median carina and some irregular short rugae on the rest
Colour very light brown, head with a reddish tinge. of the surface (more visible in oblique view). Pronotal
sides faintly longitudinally striolate. Two weak short
Female [previously undescribed]: TL about 8.5. Pro- rugulae rise backward from the insertion of the humeral
podeal spines stout, straight and quite sharp in lateral hair. Mesonotum,mesopleuronandpropodeum,excluding
view. Clypeus shagreened and subopaque, except for its its smooth decUvity, clearly reticulate and quite dull;
posteriormost median portion which is smooth. Frontal mesonotum sUghtly more superficially sculptured. Petiole
triangle quite shining. Head finely reticulate, opaque, and postpetiole wholly reticulate and mostly subopaque.
rugulose chiefly in its anterior half. Scutum and scutellum Gaster very faintly reticulate and shining.
finely punctulate-reticulate, subopaque. Axillae more Pilosity and pubescence as usual in the genus.
densely sculptured, opaque, linked by a narrow strip. Colourlightyellowish brown, the whole dorsum slightly
Gaster rather lucid. darker.
Colourbrown, with asomewhat reddish tinge chieflyon HW
the gaster and on the sides of the alitrunk. Wing nearly Paratype workers: TL 2.6-3.1, HL 0.65-0.77, PW
hyahne, very feebly yellowish throughout. 0.59-0.73, CI 91-96, SL 0.55-0.62, SI 85-93,
0.40-0.48, AL0.75-0.89, SpL0.22-0.26, HTL0.54-0.63,
Male[previouslyundescribed]:TLabout5.5. Mandibles TI 84-92 (29 measured).
withone apicalandone preapical tooth, followedbyafine As the holotype but varying as follows: Eye distance
serration on both masticatory and basal margins. Petiole fromthe mandibularinsertionrangesfromabout 1.2to 1.3
robust in profile, with a low and evenly rounded node. times the maximum eye length. Propodeal spines, meso-
Clypeusshagreened and sublucid. Head finely reticulate notum and alitrunk rather constant in shape.
and opaque; some evident oblique rugulae occur at the Smallest specimens are more superficially sculptured
posterior comers. Scutum finely punctulate-reticulate, with clearly shining frons and vertex, although still dis-
sublucid. Scutellum less sculptured and more shining in tinctly reticulate. Some rugulae always occur on the pro-
the middle. Axillae dull, densely reticulate. Sides of the notum. In largest specimens rugulae are often more
alitrunkmoreshining. Propodeumaboutaswellsculptured numerous and developed on the pronotum.
as the scutum and with distinct blunt teeth. Petiole and Colour from brownish yellow to yellowish brown.
postpetiole somewhat sublucid. Gaster rather shining.
Colour: head dark brown, alitrunk slightly lighter, Holotype worker, THAILAND: Khao Sabap Nat. Park
petiole, postpetiole and gaster light yellowish brown. 24.ix.1985 (/. Lobl and D. Burckhardt) (MHNG).
Wings as in female.
Paratype worker, 29 specimens with the same data as
Material examined. W. MALAYSIA: Penang, George- the holotype (BMNH; MCZ; MHNG).
town (H. Eidmann); Kelantan,Temangah (D. H. Murphy):
Sungei Petani (G. H. Lowe). Comments. ItsclosestrelativeappearstobeL.ambiguus.