INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 193

Stings of some species of Lordomynna and Mayriella (Formicidae: )

Charles Kugler Biology Department, Radford University, Radford, VA 24142

Abstract: The sting apparatus and pygidium are described for eight of20 species and one of five Mayriella species. The apparatus of L. epinotaiis is distinctly different from that of other Lordomyrma species. Comparisons with other genera suggest affinities of species of Lordomymw to species of and , while Mayriella abstinens Forel shares unusual features with those of P/'Oattct butteli.

Introduction into two halves and a separate sting. The stings were mounted in glycerin jelly for ease of precise This paper describes the sting apparatus in positioning and repositioning for different views. eight species of Lordomyrma that were once mem- The other sclerites were usually mounted in Cana- bers of four different genera. The stings of five da balsam. Lordomyrma species were partially described by Voucher specimens identified with the label Kugler (1978), but at the time three were consid- "Kugler 1995 Dissection voucher" or "Voucher spec- ered to be in the Prodicroaspis or Promera­ imen, Kugler study 1976" are deposited in the noplus (Promeranoplus rouxi Emery, one an unde- Museum of Comparative Zoology, Cambridge, Mas- termined species of Promeranoplus, and Prodi­ sachusetts. croaspis sarasini Emery). These genera are now Most preparations were drawn and measured considered synonyms of Lordomyrma (Bolldobler using a Zeiss KF-2 phase contrast microscope with and Wilson 1990, p. 14; Bolton 1994, p. 106). In an ocular grid. Accuracy is estimated at plus or addi tion, during a revision of Rogeria (Kugler 1994) minus O.OOlmm at 400X magnification. I transferred L. tortuosa Mann, L. leuifrons Mann, Definitions of terms (See also figure labels and L. striatella Mann from Rogeria to Lordomyr­ and Kugler 1978,1991,1992). The sensilla formula rna partly on the basis of sting apparatus anatomy. of the oblong plates is the range in number of On the other hand, L. epinota1is Mann was trans- intervalvifer sensilla, ramal sensilla, and fulcral ferred to Lordornynna on the basis of external arm sensilla found on oblong plates. Intervalvifer characters. Bere I extend and summarize what is sensilla are located near the articulation with the known of the sting apparatus in this expanded triangular plate. Ramal sensilla detect movement concept of Lordomyrrna. of the slender rami on the anterior edge of the Mayriella is a small genus from Australian and oblong plate. Fulcral arm sensilla are located in the eastern Oriental zoogeographic regions (Brown base of the fulcral arm. The sen sill a formula of the 197:3). Its phylogenetic relationships are unclear. gonostyli is the range in number on the proximal Holldobler and Wilson (1990) did not assign it to a segment followed by the range on the distal seg- tribe; Bolton (1994 p. 106) "dubiously included" it ment. The regions of the sting are defined by with Lordomynna in the . internal structure (Fig. 7). Moving anteriorly from tip, boundary between the The sting apparatus is a complex character the sting the sting shaft and the valve chamber is the point where the inner system that can be used for phylogenetic analysis of wall of the sting shaft meets the outer sting walL genera (Kugler 1978,1991; Bolton 1976; Baroni Continuing anteriorly, the boundary between the Urbani et al. 1992). Here I use it to search for valve chamber and the sting bulb is the point where Lordomyrma A1ayriella. genera related to and the inner and outer walls of the sting separate Methods again. The StingL is the sum of the lengths of the three sting regions. The Index of Reduction is a Sting apparatuses were dissected, cleared in measure of the length of the sting relative to the hot lactophenol solution, then further dissected size of the ant (sting shaft length / pronotal width). 194 Volume 11, Nos. 3-4, September-December, 1997, INSECTA MUNDI

Lordomyrma sclerotized, but often the two segments are not very well delimited. In L. epinota1is the segments are Specimens examined. L. caledonica Andre indistinct, with the proximal portion longer than (:3 workers) New Caledonia, Mt. Mou, E. O. Wilson, the distal, which has an acute membranous apex leg. L. epinotalis Mann (1 worker) British Solomon (Fig. :3). Sensilla formulae similar in all species: :3- Islands, Star Harbor, W. M. Mann, leg. L. levifrons 7, 1-4. Mann (1 worker) Fiji, Viti Levu, Tholo-I-Suva, N. L. Triangular plate (Figs. 1,2). Dorsoapical and H. Krauss, leg. L. punctiventris Wheeler (1 worker) ventroapical processes sometimes look short and Australia, Queensland, Broken River, Gkm S. Eun- truncate (Fig. 1), but sometimes appear subacute gella, W. L. & D. E. Brown, leg. L. rouxi (2 workers) (Fig. 2). Since both shapes are seen in preparations New Caledonia, Ciu, E. O. Wilson, leg. L. sarasini of L. punctiventris, L. rOlJ,xi, and L. sarasini, the (2 workers) New Caledonia, Ciu, E. O. Wilson, leg. apparent shape may depend on the orientation of L. striatella Mann (1 worker) Vanua Ava, W. M. this thick plate on the slide. Medial tubercle visible Mann, leg. L. tortuosa Mann (1 worker) Fiji, Viti in at least one preparation of all species except L. Levu, N. L. H. Krauss, leg. levifrons andL. epinotalis (may be present in these Spiracular plate (Figs. 2, :3). Rectangular in also, but just not visible in my preparations). Only most species (Fig 2); with an obvious posterodorsal L. rOlJ,xihas a dorsal tubercle. lobe only in L. rouxi (Kugler 1978 Fig. 120), L. Lancets (Figs. 5, G). Base of each lancet with a caledonica, and L. sarasini. In L. epinotalis, the single well developed valve. Lancet terminus is plate is less rectangular because the anterior apo- quite variable: sclerotized, cuneiform, and possibly deme has a large process (Fig. :3). Medial connection able to pierce in L. caledonica, L. rouxi, and L. membranous in all but L. tortuosa, where it is a sarasini (Fig. 5); filamentous in L. levifrons, L. narrow, weakly sclerotized band. Spiracle small. punctiventris, L. stn:atella, and L. tortuosa (Fig. G); Quadrate plate (Figs. 1-:3). Most species with weak and spatulate in L. epinotalis. a tapered anterodorsal corner (Fig. 2), but some- Sting (Figs. 7-11). Most species: wedge-shaped times pollicate. In a L. tortuosa specimen the plate in both lateral and ventral views; sting bulb and on one side has a pollicate corner, but the other has valve chamber little differentiated in external view a tapered corner (Fig. 1); neither looks distorted by (Figs. 7-10); internal ridge of sting base vestigial. the preparation. In some species, the dorsal edge of Variation occurs in the height of the sting base and the apodeme is not thickened (Fig. 2), but it is size of the basal notch and in the shape of the sting thickened in L. rouxi and L. sarasini, and produces apex. Sting apex is strong and evenly tapered in L. a small lateral lobe in L. epinotalis (Fig. :3) and L. rouxi (Fig. 7 and Kugler 1978, Fig. 127), strong with punctiventris. The body of the plate projects below slightly reduced sides in L. sarasini (Fig. 9), weak the level of the apodeme. The body is larger or with very reduced sides in L. levifrons, L. punc­ subequal in area with the apodeme in most species, tiventris, L. striatella, and L. tortuosa (Fig. 10), or but distinctly smaller in L. epinotalis. strong with flared sides in L. caledonica (Fig. 8 and Anal plate. Very weakly sclerotized so that Kugler 1978, Fig. 129). The sting of L. epinota1is edges of the "plate" are not visible. With 0-2 setae . (Fig. 11) is quite different, with a larger, convex . Oblong plate (Figs. 1-:3). Anterior apodeme sting bulb and slender sting shaft. Its sting shaft short, blun t. Posterior arm short, with thick, strong- seems weak and unable to pierce. The sting shaft of ly curved dorsal ridge and little or no subterminal all species examined ranges from 49% (L. caledon­ tubercle. No postincision separates the posterior ica, Fig. 8) to 5G% (L. sarasini, Fig. 9) ofStingL. The and ventral arms. Ventral arm short; fulcral arm sting bulb is 22-24% of StingL in most, but 29% in usually fusiform (Fig. 1), but with a thick dorsal L. ca1edonica (Fig. 8) and 28% in L. epinotalis (Fig. ridge in L. striatella (Fig. 2), and linear in L. 11). The Index of Reduction runs from 0.24 (L. epinotalis (Fig. :3). Sensillaformulae are similar (:3- caledonica) to 0.:31 (L. striatella,), indicating the 5, :3-8, 0-2), with only the largest apparatus (L. sting is quite small relative to the size of the ant. rouxi) having more than 5 ramal sensilla). Furcula (Figs 7, 8). In all species a simple arc Gonostylus (Figs. 3,4). In most species, clear- or V shape, with no dorsal arm. ly two segmented, with distal segment longer and Pygidium (Figs. 12, 1:3). In L. caledonica, L. narrower than the proximal; setae few and with no tortuosa, L. levifrons, L. punctiventris, L. rouxi, clear dorsoterminal chaeta or companion seta (Fig. and L. striatella with a pair of lateral depressions 4). In L. levifrons, the distal segment is heavily that may contain weak reticulation near the ante- INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 195

1

AnteriorApodeme

Gonostylus PosteriorArm

Ventral~

\ ~Fulcra/Arm

O.10mm 3

Figs. 1-3. Lateral views of Lordomyrma spp. sting apparatus sclerites. 1. L. tortuosa anterodorsal corner of the left quadrate plate; right quadrate plate, triangular plate, and oblong plate. 2. L. striatella spiracular plate, quadrate plate, triangular plate, oblong plate, and base of lancet. 3. L. epinotalis spiracular plate, quadrate plate, oblong plate, and gonostylus. 196 Volume 11, Nos. 3-4, September-December, 1997, INSECTA MUNDI

5

6

Furcula

Valve Chamber ~v

, -- .....,""":.....-- ---~------~~~~~~~~------

9 .... - .... Sting Base ------

Figs. 4-13. Lordomyrma spp. sting sclerites and pygidia. All scale lines =0.10 mm. 4. L. ronxi lateral gonostylus. 5. L. ronxi lateral lancet apex. 6. L. tortnosalaterallancet apex. 7. L. ronxilateral sting and furcula. 8. L. caledonicalateral sting and furcula. 9. L. samsini lateral sting. 10. L. striatella lateral sting. 11. L. epinotalis lateral sting and furcula. 12. L. caledollica dorsal view of pygidium. 13. L. epillotalis dorsal view of pygidium. INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 197 rior edge (Fig. 12). In striking contrast, L. epinota­ Rocks. 1200m. 35° 33'S, 148° 59'E. lG-IIl-1992 S. lis has an enlarged median reticulate-striate area Shattuck #2665. on the anterior edge of the plate (Fig. 13). No clear reservoirs were seen in any species, but they could Spiracular plate (Fig. 14). Spiracle small. have been lost in preparation. Anterior apodeme narrow. Plate narrows mesad then abruptly narrowing to a membranous medial Mayriella connection. Quadrate plate (Fig. 14). Body nearly uni- Specimens examined. Six workers of M. ab­ form in width. Apodeme becomes wider dorsad and stinens Forel, from Australia: ACT: Booroomba its anterior ridge becom~swider and weaker. An-

JDorsoterminal Chaeta

. ~ Companion Seta

(\, , I 14

O.10mm :;;,.-:------\ :;.-- \ -_...----...... rr --_~ _:::::~:==.._-.._-- __ _ Dorsal Arm~{/{/v___ ------____ ~~----.o:---_ vi ...... -- --...... , ------J , ....,,;;;::- " ...... 15 -....,\""--

\\

I I I /.i-'::-;.::-;.;_'--_-

Figs. 14-16. Mariella abstinens sting apparatus. 14. Lateral view ofspiracular plate, quadrate plate, triangular plate, oblong plate, gonostylus, and lancet. 15. Lateral view of sting and furcula. IG. Ventral view of sting and furcula. 198 Volume 11, Nos. 3-4, September-December, 1997, INSECTA MUNDI terodorsal corner blunt, not prominent. Dorsal edge other species in its sting apparatus: 1) spiracular of apodeme without medial or lateral lobes. ridge with a thick prominence on its anterior apo- Anal plate. Oval, very weakly sclerotized, its deme, 2) linear fulcral arm of the oblong plate, 8) edges not always distinct from surrounding mem- distal segment of gonostylus not so elongate, 4) brane. Posterior edge with 4-6 long setae. spatulate lancets, 5) sting bulb longer and sting Oblong plate (Fig. 14). Anterior apodeme long shaft more differentiated from the rest of the sting, and tapered. Posterior arm slender, with weak and 6) single patch of anterior pygidial sculpture. dorsal ridge, and no terminal tubercle; not separat- This is now the fourth case in which quite ed from ventral arm by a postincision. Ventral arm different sting apparatuses are found among spe- with rather long, narrow fulcral arm that is well cies of the same genus. Two kinds of stings have sclerotized only in the ventral half. Sensilla formu- been found also in species of A1onomoriwn (subgen- la: 2-8, 4-5, 0-2. era Holcomyrmex andMonomoriwn (Kugler 1978», Gonostylus (Fig. 14). Short, nearly membra- Pheidologeton (Kugler 1986), and Rogeria (Kugler nous, single-segmented. With two separate patches 1994). On the other hand, other genera presently of sensilla: 5-6 proximal plus a dorsoterminal cha- seem quite uniform, e.g. Gnamptogenys (Kugler eta and companion seta. 1991), Tetramorium (Kugler 1978, Bolton 1976), Triangular plate (Fig. 14). Body wide basally, and Crematogaster (Kugler 1978). then abruptly narrowed to the ramus. No dorsal or Bolton (1994) placed Lordomyrma in the Tribe medial tubercles. Stenammini, while Bolldobler and Wilson (1990) Lancet (Fig. 14). Valve single, small. Lancet placed it in the Pheidolini. I have examined the apex needle-like; no barbs. sting apparatus in 18 of Holldobler and Wilson's 19 Sting (Figs. 15, 16). StingL = 0.20-0.23. Sting Pheidolini genera and9 of Bolton's 18 Stenammini neither highly developed nor strongly reduced (In- genera. Lordomyrma species other than epinotalis dex of Reduction = 0.52-0.57). Sting shaft short, do have some similarities with , Cy­ about 45% to 47% of StingL; well sclerotized; apex phoidris, and Lachnomyrmex, which are members acute, without dorsal flange. Valve chamber low, of Holldobler and Wilson's Pheidolini as well as 16%-17% of StingL. Roof of valve chamber held Bolton's Stenammini. All three genera have elon- below dorsal surface of sting by a strong, wide gate an terodorsal corners of the quadrate plate like transverse ridge and median tubercle, so the cham- those of Lordomynna spp. In addition, species of ber is very weakly differentiated from sting shaft Cyphoidris an d Laduwmyrmex have gonosty li with and sting bulb externally. Sting bulb moderate in elongate distal portions (no clear segments), few height and length, about 87%-38% of StingL. Sting setae, and no dorsoterminal chaeta. But, on the base not transversely arched and with weak basal other hand, species of Cyphoidris and Lachno­ ridge; anterolateral processes reduced to corners. myrmex differ greatly from those of Lordomyr rna in Furcula (Figs. 15, 16). Shaped like an inverted sting and lancet shape. Moreover, quadrate plates Y. Dorsal arm prominent. like those in species of Lordomynna are also found Pygidium. Anterior edge with two patches of in the Pheidologetini and Cephalotini (Kugler 1978, microreticulate sculpture that meet at midline. Kugler 1986). Finally, the sting of Lordomyrma This sculpture suggests the presence of pygidial epinotalis is most similar to those of species of glands, but the sculpture is effaced and no convinc- A1onomorium subgenus fl,1onomorium (Solenopsi- ing reservoirs appear in any of the five pygidia dini). Thus, while suggesting an affinity of Lor­ examined. dornyrrna with Cyphoidris and Lachnomyrmex, (Stenammini or Pheiclolini, depending on one's Discussion classification), the sting apparatus does not provide unequivocal support. With the exception of L. epinotalis, all of the A1ayriella abstinens has an unusual feature in Lordomyrma species examined have several dis- its sting: a thick transverse ridge in the valve tinctive characters within the myrmicines: 1) gono- chamber plus a median tubercle. Of the specimens styli two segmented, with distal segment longer I have examined, a large median tubercle is found and narrower than the proximal; setae few and in species of Cardiocondyla, Tranopelta, and Lo­ with no terminal sensilla (Fig. 4), and 2) cuneiform phomyrmex. A thick, wide ridge is present in spe- sting with little or no basal ridge (Figs. 7-10). ButL. cies of Terataner, Oligomyrmex, and Pheidologe­ epinotalis, which looks unequivocally Lordornyrma ton. Tranopelta is also most similar in overall sting in external anatomy, differs markedly from the INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 199 shape. But only butteli has the com bination thrix Fore!' Bulletin of the British Museum (Natu- of ridge and tubercle. Another unusual character is ral History), Entomology 34(5):281-379. the shape of the end of the lancet. This is found in Bolton, B. 1994. Identification Guide to the Ant Genera some Tetrarnoriwn species, Cardiocondylaelegans, of the World. Harvard University Press Cambridge MA. 222 pp. . and in Proatta butteli. Thus, these characters, Brown, William L., Jr. 1973. A comparison of the likely to be apomorphic, are shared only by species Hylean and Congo-West African Rain Forest Ant of Mayriella and Proatta. In addition, Proatta but­ Faunas. In B. J. Meggers, E. S. Ayensu, and W. D. teli has a similar Y-shaped furcula (likely symple- Duckworth, eds., Tropical Forest Ecosystems in siomorphy), gonostyli (possible reduction conver- Africa and South America: A Comparative Review. gence), triangular plate, and a short but piercing pp. 161-185. Smithsonian Institution Press, Wash- sting. Bolton (1994) placed Mayriella and Proatta, ington, D. C. 350 pp. in the Stenammini. My analysis of the sting appa- Holldobler, B., and E. O. Wilson. 1990. The . ratus also indicates a relationship between those Belknap Press of Harvard University Press, Cam- bridge, MA. 732 pp. genera. Kugler, C. 1978. A comparative study of the myrmicine Acknowledgments sting apparatus (, Formicidae). Stu- dia Entomologica 20:413-548. I thank Steven Shattuck for sending the spec- Kugler, C. 1986. Stings of ants of the tribe Pheidologe- tini (Myrmicinae). Insecta Mundi 1(4):221-230. imens of Mayriella abstinens for study. Kugler, C. 1991. Stings of ants of the Tribe Ectatommi- References ni (Formicidae: Ponerinae). Insecta Mundi 5(3- 4):153-166. Baroni Urbani, C., B. Bolton, and P. S. Ward. 1992. Kugler, C. 1992. Stings of ants of the Leptanillinae The internal phylogeny of ants (Hy,menoptera: For- (Hymenoptera: Formicidae). Psyche 99(1): 103-ll5. Rogeria micidae). Systematic Entomology 17:301-329. Kugler, C. 1994. Revision of the ant genus Bolton, B. 1976. The ant tribe Tetramoriini (Hy- (Hymenoptera: Formicidae) with descriptions of menoptera: Formicidae). Constituent genera, re- the sting apparatus. Journal of Hymenoptera Re- view of smaller genera and revision of Triglypho- search 3: 17 -89.