HARVARD UNIVERSITY Library of the Museum of Comparative Zoology BULLETIN OP THE MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD COLLEGE, IN CAMBRIDGE VOL. 118 CAMBRIDGE, MASS., U. S. A. 1958 The Cosmos Pkess, Inc. Cambkidgk, Mass., V S..\. CONTENTS PAGE No. 1. — Observations on the Egg-Capsules of Skates of THE Family Ra.tidae, found in Japan and its Adjacent Waters. By Reizo Ishiyama. March, 1958 1 No. 2. — A GENERIC Review of the Plovers (Charadriinae, AvEs). By Walter J. Bock. March, 1958 . . 25 No. 3. — Studies on the Ant Fauna of Melanesia. I. The Tribe Leptogenyini. II. The Tribes Amblyop- onini and platytiiyreini. By E. 0. Wilson. April, 1958 . . . . ' 99 No. 4. — The Fossil Carnivore Amphicyon Intermedius from the Thomas Farm Miocene. Part I: Skull and Dentition. By Stanley J. Olsen. May, 1958 . 155 No. 5. — Contribttions toward a Reclassification of the FoRMiciDAE. II. Tribe Ectatommini (IIymenop- tera). By William L. Brown, Jr. June, 1958 . 173 No. 6. — The Spider Subfamily Clubioninae of the United States, Canada and Alaska (Araneae: Clubionidae). By Robert J. Edwards. (24 plates). June, 1958 . ' 363 No. 7. — The Nearctic Members of the Genus Entomob- RYA (CollexMbola). By Kenneth Christiansen. (24 iilates). June. 1958 437 Bulletin of the Museum of Comparative Zoology AT HARVARD COLLEGE Vol. 118, No. 1 OBSERVATIONS ON THE EGG-CAPSULES OF SKATES OF THE FAMILY RAJIDAB, FOUND IN JAPAN AND ITS ADJACENT WATERS By Reizo Ishiyama CAMBRIDGE, MASS., U.S.A. PRINTED FOR THE :MUSEUM March, 1958 Publications Issued by or in Connection WITH THE MUSEUM OF COMPARATIVE ZOOLOGY AT HAPvVARD COLLEGE Bulletin (octavo) 1863 — The current volmne is Vol. 118. Breviora (octavo) 1952 — No. 85 is current. Memoirs (quarto) 1864-1938 — Pul)lication was tei-minated with Vol. 55. Johnsonia (quarto) 1941 — A publication of the Department of Mollusks. ^^ol. 3, no. 35 is current. Occasional Papers of the Department of Mollusks ((^-tavo) 1945 — Vol. 2, no. 21 is current. Proceedings op the New England Zoological Club (octavo) 1899-1948 — ■ Published in connection with the Museum. Publication terminated with Vol. 24. The continuing publications are issued at irregular intervals in num- bers which may be purchased separately. Prices and lists may be obtained on application to the Director of the Museum of Comparative Zoology, Cambridge 38, Massachusetts. Of the Peters "Check List of Birds of the World," volumes 1-3 are out of print ; volumes 4 and 6 may be obtained from the Harvard Uni- versity Press; volumes 5 and 7 are sold by the Museum, and future volumes will be published under Museum auspices. Bulletin of the Museum of Comparative Zoology AT HARVARD COLLEGE Vol. 118, Xo. 1 OBSERVATIONS ON THE EGG-CAPSULES OF SKATES OP THE FAMILY RAJIDAE, FOUND IN JAPAN AND ITS ADJACENT AVATERS By Reizo Ishiyama CAMBEIDGE, MASS., U.S.A. PRINTED FOR THE :\r U S E U M March. 1958 Xo. 1 — Obscrvdfioiis on tli( I'Jf/fi-CapsKirs of Skates of the Fdinihi li(tj;(hi(. found ni ■Inpon (ni:l its fi(lj(icsen1 investigation liave b(H'n based in part on these works, as well as on various reports given bj' other authors. In the present study, it has been found tliat the egg-capsules of Japanese rajid fishes have certain features that aid in their identification besides establishing possible relationships between genera and even between species ; also these features may throw some light on adaptive differentiation of various species. It is a pleasure to record here a debt of gratitude to Dr. K. Matsubara, Professor of the Kyoto X^niversity, for his kindness in giving valuable suggestions and in contributing a number of specimens for study. I am especially indebted to Dr. Henry B. Bigelow and to Mr. William C. Schroeder of the Harvard Mu- seum of Comi')arative Zoology and the Woods Hole Oceano- graphic Institution for much assistance in the i)reparation of the manuscript. Further, many colleagues Avorking on fisheries biology at various stations in Japan helped me with collections of specimens and with important literature. To them I am grate- ful. MATERIALS AND METHODS The specimens used in the ])rese]it study were collected at various localities (Tabh^ 1 and Fig. 1), where the egg-capsules were obtained directly li-om parent females of three genera, involving twenty-one species and two subspecies, which repre- sent almost all those uKMiihers that are known in the seas ai-onnd Ja])an. In addition to thes(% some capsules Avere removed from both ovi])arous an;l o\-ovivii)arous adults, and the rest were gathered in the field. All the specimens are preserved in a formalin solution (lOo;-) in our college. The examinations were first made on the external features and followed by histological study, the specimens being cut in sections, 20-r)() micra thick, with the aid of a cvlinder microtome. ISHIYAMA: EGG-CAPSULES OF JAPANESE SKATES Table 1 Tlie number of egg-eapsules examined of 21 species and 2 subspecies of Japanese skates, together with k^calities, depths, and temperatures where parent females were captured. The six species and two subspecies which are abbreviated by the let- ters C, J, N, P, Q, Q', S and W, respectively, are thought to be new to science. Abbreviations within brackets represent the seas where the specimens were collected: [C], East China Sea; [J], Sea of Japan; [0], Okhotsk Sea; [P], Pacific Ocean. Xo. of egg-capsules Depth (m) 1 'ollowed by Scieutiflc name ami i ts I'xamiiicil and temperature (° C) where the letter symbol localities parent females were captured* Bieviraja tobitukai A 5 [P] 300-400 9-10 B. isotrachys B 4[J;0J 90-230 3-4 - B. sp. C 5 [P] .500-700 3 ^ B. diplotaenia E 5 [P] 300-500 2-3 ^-1 B. inatsubarai F 2 [P] 800-1000 o s B. aleutiea G 12 [P] 300-700 2 z> B. paimifera H 2 [P] ? 1 +- B. smirnovi I i 0] 100-200 5 3 Rhiiioraja sp. J •2 [P] 230-400 9-10 r i Rli. kujiensis K 6 [P] 600-800 2 Rh. longicauda L 4 [P] 300-600 5 Raja kenojei M 10 JJ 80-90 17 R. sp. N 6 [J] 50-90 21 R. hollandi 0 32 [C;J] 70-90 14 I R. sp. P 4 [J] 80-90 17 "-4— R. subsp. Q 5 [J] 80-150 8-10 - R. subsp. Q' 8 LP] 30-50 21 c; R. fusea R 4 [P] 20-50 12 ^ R. sp. S 4 [P] 20-40 11 o R. tengu T 2 [P] 150-200 14 CC R. pulchia U 25 [J;0] 50-100 8 R. inaeioeaudi I V 7 [P] 300-400 10 R. sp. W 2 [P] 300-400 10 Description and Comparison of the Egg-capsules A. External features : Shape. The egg-capsule in raj ids is usually rectangular in shape with a horny process at each corner. The anterior end of * Data from the Annual Uepi'rt on the BMsh Resources in Oct. lSl.".l-Sept. lOoli, Section 4: Bottom Fishes. pul)lishe(l by Tohoku Regional Fisheru-s Laboratory, 1954. and frnm the Semi-Anuual Report on Oceauographical Investigation Xos. 70-74, compiled bv the Tokai Regional Fisheries Laboratory and published b.v the Investigation of Research Division of Fisheries Agency, 194.:!, litol and TJo.i. 6 niLLP^Tix : ^ri^sErM of comparative zoologv \\\e capsule is tightly fused with a luorc or less tiattenetl niargiu wliicli is directed downwards towards the head of the parent fish, hut posterioi'ly it is loosely closed with a thinner ])ellicle, from wliich tlu^ (Mubryo emerges at hatching, in most cases. Tlie sides are in general finely keeled lengthwise, to a greatcM- oi- lesser degree, for almost the entire leugtli excejiting ihe capsule of species T'l (Fig. 3, U). On the basis of shape, the egg-capsules luay be classified into six forms (Figs. 2, 3) which will be described in detail. Both dorsal and ventral sides of the capsules are more or less convex, but the dorsal side is in general decidedly more domed than the ventral and forms the dorsal border which is orientetl towards the dorsal side of the body. Size. It should be ])articularly noticed that the egg-capsule in the rajids is relatively large in size, although rather considerable intras])ecific variations are found to exist in some cases. As a mattei- of fact, the size of the capsule may serve as a means for grouping the species. Measurements showing the variations in tw^o examples follow. Tabl.- 2 Variations in the dimensions of the capsules of two species, P and U. The numbers in parentheses represent the number of species examined. Sppcios Main portion Horns longtli. mm. 1 width, mm. anterior, mm. posterior, mm. P (32) 53.2-65.9 31.5-39.2 15.0-22.0 11.0-22.0 U (25) 140-185 70-94 The sizes of the egg-capsules were found, in general, to bear a i-elationship to the sizes of the parents, as illustrated in Figure 4, which would seem to be different from that given for birds (Hux- ley, 1932, p. 225). Horns. At each of the four corners of the capsule there is shown a projection, the horn, one pair of which is usually longer than the other. IJoth the length and shape of the liorn are im- 1 For the sake of hrevit.v in the description, the specific names of all the ra.iids treated here are abbreviateil h.v letters as shown in Table 1. ISIIIYAMA: EO(i-(AI'SlTLES OK . JAPANESE SKATES i portaiit as a means of specific identification. Usnally, the honi oil the cajisuh^ of the nortliern form is lonn of the surface. There is a marked difference in the configuration of the surface of the main portion of the capsule between the northern and the southern forms. The capsule is more or less roughened with minute prickles or tubercles develop- ing over the entire surface in that of the northern form, l)ut it is, in general, very smooth in the southern one. It may be convenient to compare the features of the northern and southern forms with the structural evidence of the capsules CAamined histologically as Avill be mentioned later on. Fibroid tendril. The egg-capsule in the rajids is usually covered with a tight-fitting felty mass of fibres immediately after it is extruded from the parent female, but the fibres are gradually reduced after it is deposited in the sea. The lateral margin of the capsule is also provided with a mass of loose filjres, well devel- oped in the species belonging to the northern form, especially in those found in deep water (Fig. 2, F, I), although the fibres are found to occur in greater or lesser degrees in all examples dealt with here. The degree of development of the felty mass of fibres in some cases is a character of vakic in distinguishing one species from another. 1). Histological structure of the egg-capsule: Tlie main ])ortion of the capsule in tlie rajitls is made of two or more kinds of tissue, which are referred to as an inner, pulpy, layer and an outer layer (Ishiyama, l!)5()). In general, the inner hiycr forming the lining of the capsule-wall is relatively soft and colorless, but the tissue l)ecom(N hard and the color changes from ISHIYAMA : EGG-CAPSULES OF JAPANESE SKATES 9 yellow to ])rowii in the outer layer -whieh covers the surface of the capsiUe. (ienerally the eapsiUe in the northern form is charac- terized hy having- a mueli thickened outer layer which reveals, ill different species, various features on the surface hy the de- velopment of either tubercles or prickles of a hard, horny, brown substance. But the inner layer is rather thin, and without a noticeable difference in histological structure in the different species. In the southern members, on the other hand, a special- ization is found to exist not only in the outer layer but in the inner one as well. Based on certain characteristics the writer classifies the capsules of the Japanese raj ids into seven groups, ab!)reviated herein as Group Eg^ to Eg'^. Group Eg^ (Fig. 5, A) is represented only by species A. The capsule-wall is made of two layers, the inner one being relatively thick with parenchymatous structure, whereas the outer one is very thin and faintly colored. A capsule of this type is smooth externally, translucent, and tlie yolk mass is visible in situ. This ty])e may be the most ])rimitive with respect to its histological structure. Group Eg- (Fig. 5, I) may be exemplified by the cap- sules of two species, H and I, in which the tissue of the wall con- sists (if Uw layers, the outer one being very thick and bearing many dark-colored tubercles so that the surface of the capsule is i-ather i-ough. With regard to the degree of development and the shape of the tubercles two forms may be recognized, referred to here as Subgroups Eg--^ and Eg-". In Subgroup Eg-^ the char- acter of the wall is as described above and shown in Figure 5, I in which the tubercles are sparsely developed in lengthwise series, and bear domed apexes when cut in cross section. Subgroup Eg-" (Fig. 6, ID, which corresponds to the Parmif era-type in my previous report (Ishiyama, 1!)5U), is exemplified by that of species H ; the tubercles occur more densely than in the preceding, and reveal a rather rough surface because of the many minute ridges running in lengthwise series. Group Eg^ (Fig. 5, L) is exemplified hy the capsule of two species (J and L), which cor- responds to the Isotraehys-subtype in a previous paper (Ishi- yama, 1952 ) . The wall of this type also consists of the two layers, the outer being provided with some stalks and prickles as well, both of which are in a lengthwise series forming many obscure, minute longitudinal ridges. Group Eg^ (Fig. 5, B) includes 10 ijii.Lirnx : :\rusEUM of comparative zoology capsules belonging to six species (B, V, E, F, G and K), these capsules having in common a thorny, very rough surface. The tissue consists of the two layers, of which the outer one is very thick and characteristic. The surface is densely armed with numerous series of minute velvety prickles. Here, the fact should be mentioned that there exist tliree different forms of the spina- tion which are referred to as Subgroup Eg^^ to Eg**^ Subgroup Eg^-^ (P^ig. 5, B; Fig. G, C, E) includes the capsules of three species (B, C and E) and corresponds exactly to the Isotrachys- type in my previous report (Ishiyama, 1950). In this subgroup, the minute i)rickles covering tlie capsule are perpendicular to the surface, so tliat it feels very thorny when touched. The de- gree of development of the prickles, however, has progressed more in the capsule of species E than in that of species B and (' (Fig. 7, B, E). Subgroup Eg'" (Fig. 6, G, K) is represented by si)ecies (t and K, whose capsules are characterized l)y having numerous, elongate;! ju-ickles bent nearly horizontally forward on the surface of the capsule (Fig. 7, G). The length of the ])rickies is somewhat greater in species G than in K. Subgroup Eg^e (Fig (j^ Xh"') involves only the capsule of species F, and is (|uite ])eculiar in the manner of its armature, being referred to as the Isotrachys-superty]X' in my previous report (Ishiyama, 1952). The general features of tlie proper capsule and its his- tological structure are rather similar to Subgroup Eg^\ but Subgroup Eg^'' diff'ers from the latter in having many prickles wlucli form a velvety texture on the surface densely beset with small stiff rods serrated at the tip, over which many strong Hbroid masses of hair entirely cover the surface. Such being the case, the capsule has a rather smooth surface ifi situ, owing to its tibi-oits covering. It is l)elieved, therefore, that this type of capsule might have evolve(l from th(> i-elated fontis. foi- it seems to be the most specialized. Capsules of the southern species may be separated into three groups although they fall into but two groups as regards their features //; f>itu. Group Eg'' ( Fig. f), R ) is characteristic of the capsules of species R and subspecies Q and Q', which have a smooth surface and a wall composed of two layers, a condition connnonly found among the northei-n members. The inner layei' is relativelv thick instead of being thin, while tlie outer one has ISHIYAMA: EGG-CAPSULES OK JAPANESE SKATES 11 IK) (lilVcrtMitiation on tlu> (mtcrniost tissue. A cairsiilc of lliis Lironp socnis to have llie most ronp appear much the same in external appearance as the preceding', hut the two are sliai'ply defined in histological structure. The tissue of the inner hivei- of ({roup Eg'' is characterized hy having a pulpy layer inserted in greater or lesser degree. Therefore, the wall is made of three layers, of wliich tlie outer one is usually very thin and fui-nislit^l with mimite gianules which correspond to the basal portion of the fibroid bairs, covering the surface of the capsule. If nuist be pointed out here that both Groups Eg"^ and Eg'' were included in the Kenojei-type. which was divided into two groups, as mentioned above in a foregoing report ( Ishiyama. 1950), being based on histological evidence. Further, it is of con- siderable imjiortanee to record here that the relative thickness of both the inner and the jndpy layers differs among species (Fig. •"). T: Fig. 8, M. X. O, P, S, \' '.. Thus the pulpy layer in species T is very thin as compared with tlu> relatively thicker inner one, but both layers have moderate thickness in species M, S. and V, and ihe judjiy layer is quite remarkable in thickness as revealed in sjiecies X. O and P. Based on these facts it may be possible to use the above char- acteristics in ideniifymg those capsules which are quite similar in external appearance and thus difficult to distinguish one from anothei'. (iroup Eg" (Fig. o. \V ; is represented liy the capsule of species r and W. The wall referable to this group is also composed of three layer.s. as is the case with the preceding, but these are greatly ciitferent in respect to their histological structure and surface configuration as well. The pulpy layer of this group is reticulat(Ml to form a porous structure which is backed internally iiy a vei-y Thin inner layer; and besides, the outer layer becomes very thick, containing externally several zones of fibrous frame, transformed from the outermost tissue of the layer, and the sur- face is roughened by having longitudinal tubular fibres which can t^asily fall out by themselves Avithout any strong friction. This group was referred to previously as the Pulchra-ty])e in marking 12 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY the eliaraetei-istics observed on the capsule of species U, but the fibres whicli form the outermost tissue are different in the capsules of species V and W, the tissue in that of species U having very thickened fibres (Fig. 8, IT) rather than fine thread-lilce ones as ill species AV (Fig. 5, W). On the basis of these facts it may be said that this grouji (Eg') seems to be the most higlily specialized among those found in the southern forms, and further, the structure revealed in the caj)- sule-wall in si)ecies U may be more advanced than that in species W. On account of the evidence mentioned above, it will be as- sumed that the two extremes in the course of specialization in capsules of the southern form may be represented by (Groups Eg"' (Fig. 8, Q) and Eg' (Fig. 8, U)! Thus, the examination of the features of the capsules reveals much diversification in botli the external and histological char- acteristics in the Japanese rajids. DISCUSSION 1. Systematic significance of the egg-capsule Based on the facts found in the external as well as in the his- tological characteristics of the egg-capsules of Japanese rajids, the present writer gives an analytical key to species and groups of species. From this key we may be able to recognize that the char- acteristics of the capsules are not only of much importance as a means for their identification but also that they seem to be of taxonomic importance. la. Very large in size, iiieasuriiig nioie tlian 100 mm. in length exclusive of horns. 2a. Surface very rough being velvety in texture, Group Eg*. 3a. Without strong fibroid hairs entirely covering the surface; prickles de- void of small stiff rods at their tips. 4a. I'rickles rather long being perpendicular to surface, Subgroup Eg-*-'^. 5a. Posterior horns longer than width of capsule; capsules 106 mm. to 132 mm. in length and 7(i mm. to 86 mm. in width Bnviraja isotrdclni.^ (B, in Figs.) ; Breviraja sp. C (C, in Figs.). .lb. Posterior horns shorter than width of capsule; capsules 106 mm. to 120 mm. in ieiigtli and 76 mm. to 78 mm. in width Breviraja diplotaenia (E, in Figs.). -Irb. Piicklcs ver\- long, liending anteriorly, Subgroup Eg'*^. ISHIYAMA: EGG-CAPSULES OF JAPANESE SKATES 18 (in. Capsules imicli Inrjier than tliosi' of other northern forms, ahout 120 nun. to 13(3 mill, ill h>iiasuriiig about 0.2 mm. in thickness exclusive of prickles Breviraja matsnharai (F, in Figs.). 2b. Surface smooth or somewhat rough, without any velvety texture, Groups Eg-, Eg", and Eg". 7a. Surface somewhat rough; horns tape-like in shape or reduced to rudi- ments. Groups H-, H-'', and H'^. 8a. Horns rather long, growing slenderer toward tips, pointed at their tips, -nath a longitudinal slit located at their basal parts. Group H- ; capsule-wall composed of two layers, the outer one furnished with many tubercles over the surface. Group Eg-. 9a. Length and width of capsule about 150 mm. and 90 mm., respectively; tubercles covering outermost layer sparsely developed with domed apexes. Subgroup Eg-^^ Breviraja smirjiovi (I, in Figs.). 9b. Length and width of capsule about 120 mm. and 90 mm., respectively; the tubercles densely developed with ridged apexes. Subgroup Eg-^ Breviraja parmifera (H, in Figs.). 8b. Horns very short or rudimentary, with a round opening located at their tips or at each of the four corners. Groups IP and H'' ; capsule- Avall composed of three layers, the outermost composed of many fibres running lengthwise in series. Group Eg'. lUa. Horns very short, with the opening at their tips, Group H-J ; broad notches present on sides; length and width of capsules from 140 mm. to 185 mm. and from 70 mm. to 9-4 mm., respectively Eaja pulchra (U, in Figs.). 10b. Horns reduced to rudiments, with an opening at each of the four corners. Group H'^; notches absent on sides; fibres composing outer- most tissue less in diameter than those in 10a; greatest in size of all capsules examined, measuring about 235 mm. in length and 145 mm. in width Baja sp. W (W, in Figs.). 7b. Surface smooth; horns short, tube-like in shape. Group H"*. 11a. Length of capsule about 100 mm.; capsule-wall very thick, measuring more than 1.0 mm., with a very thin pulpy layer which comprises about 13% of the total thickness of the wall Eaja iengu (T, in Figs.). 14 lU LLETIX : MT'SEIM OF COMPARATIVE ZOOLOGY llli. Length of i-apsulcs about loO nun. to 140 nun.; capsule-wall relatively thin, measuring about 0.4 nini., with a pulpy layer which comprises ;ili(uit 42% of the total thickness of the wall Baja macroeauda (V. in Figs. ). 111. I'sually small in size, measuring less than 70 nun. in length exclusive of hoins. rJa. Horns long, wliijilike in shajie, l>earing a longitudinal fissure; surface cither rather rouuli or very smooth, (ii'oups IP and H-'. 18a. Surface rather rough, liearing minute prickles with ridged apexes: capsule-wall divisible into two thickened layers, being compact in structure. Group Eg^; moderate in size, measuring iil)out ~u mm. to 68 mm. long, and 33 mm. to 46 mm. wide Rlnnornjii sp. J (J, in Figs.) ; Fli. Innciicauda (L, in Figs.). Kill. Suiface smooth, without any sculpture; cai)sule-wall translucent, com- posed of two layers, the outer one very thin, instead of being re'atively thick, the inner one iiarcndiymatous in structuic, (iroup Egi ; wvy ^^niall in size, mc;isuring about 38 nnn. long, and 'I'l mm. wide Bnviraja tohitul-oi (A, in F'igs. ». ll!li. Horns short, tube-like in Nliapc, bearing a I'ound opening at the liji. Group rH. 14a. Capsule-wall composed of two layers, Groui) Eg''. l"ia. Capsule-wall about 0.25 mm. thick; length and width of capsule about 51 mm. to 'u mm. and 33 mm. to 37 mm., resjiectivcly Unja fused (R, in Figs.). 151). Caiisule-wall thin, aliout ii.l5 nun. thick: length and width of capsules about 43 mm. to 59 mm. and 29 mm. to 37 mm., respectively Ji'ajd subsj). Q (Q, in Figs.); N. subsp. Q' ( ()' . in Figs.). 14b. Capsule wall composed of three layers, (Jrouj) Eg*'. 16a. Capsule-wall 0.25 mm. to 0.4 mm. thick, with a puljiy la\er which i.-s thinner than 50' < of the total thickness. 17a. ruliiy layer coniinises about 211% to 30% of the total thicknes.s of the wall; length and width of ca])sules 5(i mm. to (i(i mm. and 34 mm. to 37 mm., respectively; tiliroid hair exceedingly well develo])e(l over the surface Raja J:enojei (INI, in Figs.). 171i. Pulpy layer comprises up to 50% of the total thickness of the wall; length and width of capsules 53 mm. to 65 mm. and 27 mm. to 30 mm., respectively; filiroid h;ii)- less developed than in 17a . . . ti'djd lidlhindi (O, in Figs.) ; L'ajo sp. 8 (S. in Figs.). lOa. Capsule-wall less than 0.2 mm. in thickness, with a puljiy layer comiuis- ing as much as 60% of the total thickness of the wall. ISa. Capsules measure from 49 mm. to 54 mm. in length; surface densely covered with tine fibres L'ajo sp. X (X, in Figs.). ixb. Cajtsules the smallest in size, nuasuiing only from 39 mm. to 42 mm. in length: fine filues covering the surface less developed than in ISa Raja sp. P (P, in Figs.). ISllIVAMA: i;(;(;-(Al'Sl'LKS OI' .IAI'ANESE skatks 1"i 2. Adaptive differentiation of the egg-capsules As ren'cirds the present item tln' wi-iter directs attention to the following points : a.) The characteristics of the capsules differ between the northern and southern members : As already mentioned, the horns of the capsules of the northern members are as a whole nuich longer than those of the southern ones, and moreover the location of the respiratory perforation on the horn also differs l)etween the two forms. Another remarkal)le difference between them is found to exist in the surface confignration and in the histological character as well. In the northern form, the outer layer is in general clad in an armour of either tubercles or prickles, which is useful as a means for identifying the capsule of this group of rajids. In the southern form, however, it is the internal tissue of the capsule-wall. remarkal)ly modified in most cases and espe- cially recognizable within the inner layer, that serves as a useful means of identifying the capsules of this group. Wherefore, it may be said that a modification of the outer layer is the more usual condition in the northern form, while it is the inner layer that has become modified in the southern one. This difference in characteristics may have resulted, in part at least, from differ- ences in their breeding haliits inasmuch as the capsules of the northern form, with their long horns or filirous tendrils, are laid so as to be entwined around some ol)ject above the sea floor; but it is supposed, according to Clark (,!!'--. P- 582), that the capsules of the southern form are attached to the bottom using the fibroid hair as ;iii aui-hor. b.) The diff'erentiation in the characteristics of the capsule l)ears a close relationshiji to the geographical distribution of the adult skates: (ienerally speaking, there is a tendency in the •lapanesc i-ajids that tlie more northward the fish ranges, the greater the differentiation of the capsules. The capsule of species A, which is found in the southernmost part of the geographical distribution occupied by the northern forms, has the least degree of specialization in its characters as a whole. The cajisules of species .1 and L. which are found farther northward next to the preceding species, have characteristics which seem to show greater progress in diff'erentiation than that of the formei-. The same view may l)e expressed by the capsules of sjjccies 1. which 16 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY has a raiiiie of distribution froin the deep sea in the southern region of the Sea of Japan to the far north and has the character of the capsule-Avall of Group Eg-. Further, the capsule-wall of (irouj) Eg"* is possessed exclusively by skates which reach still farther northward in distribution or inhabit dcM'per waters than Ihose having the capsules of Groups Eg' and Eg-. A case which exactly conilrnis this opinion may be given by the most specialized Group Eg^*^' represented by the capsule of species F, which lives in waters of great depth (Ishiyama, 1952). Among the southern forms, on the other liand, though there are some cxam})l('s which do not fit tlie above conditions, it may be pointed out that in general the degree of develojiment of the pulpy layer Avithin the inner one seems to be correlated in the iiKiin with the distribution of the tish. For example, species Q and K were obtained in the coastal waters ranging from the ex- trcnic norlh lo the southerly region of the main island of Japan, and 1he eai)sule-wall in these species is characterized by having only iwo layers ((iroup Eg"'), whereas species X and !S, which are 1 bought to be closely related to the preceding species, were found to inhabit the waters ranging from the middle region of the main island to I'arlher soutli, and their capside-wjdl is eom])osed of three layers as in Group P]g''. Another example of this is to be found in the ca])sules of species U and AV. These two si)ecies not only have many s])ecialized features in the capsule-wall (Group Eg' ), but also cover either an extremely wide liorizontal distril.ution or the deepest region of the range. It should be men- tionel hei-e that the capsules of these two si)eeies are I'.elieved to be laid on the sea floor. Such being the case their capsules are often gather(Hl by the trawl net. Taking the above into consideration, we conclude that tliere seems to l.e a close relationship between the differentiation of the capsule and the geograiihicrd distril.ution as well as the breeding habits of the adult skates, though there are some exam|)les thai do not fit this conclusion. 3. Consideration of the differentiation of the capsule The of:!:^^ capsule, of course, serves essentially for the protection of the egg, which is destined to form the young fish, during its l)eriod of development (Wilson, i;)2S. p. 274). Thus the caiisules might have uiulergone successive chanues in their characteristics ISHIYAMA : EGG-CAPSULES OF JAPANESE SKATES 17 ill order to adapt to their environment. On this hasis the writer lias eiuh'avorod to (dassify the capsnU^s of the dapanese rajids into eiirht types (Table 3). Table 8 Characters of the eg^-eapsules in Japanese rajid fishes classi- fied into 8 types. For abbreviations, see text. Horn, Group Capsult'-wiill Type ContiKuratioii of surface Tissup Group E<; and its Subgroup Species 1 HI smooth 2 layers Eg A H-' rather rough 2 layers Eg^ Eg2A I Eg2B H 3 H3 rather rough 2 layers Eg^^ J; L H3 rough 2 layers Eg-i Eg4A B; C; E 4 Eg4B G; K Eg-ic F _ H-i smooth 2 layers Eg5 Q; Q'; R 6 H^ smooth 3 layers Eg« M; N; 0; P; S; T; V 7 H>"> rather rough 3 layers Eg" U / 8 H6 rather smooth 3 layers Eg- W Questions arise as to the possible causes of these differentiations in the capsules. Differential roles of the capsule which serve in the development of the embryos would seem to be present. This hypothesis, however, is based principally upon morphological evidence. Although various opinions have been given in regard to the aeration of the capsule, they should agree with the view of Clark (1922. p. 586) in that the respiration of the embryo enclosed in the capsule takes place by means of general osmosis through the capsule-wall and by the circulation of Avater through the openings 18 mTLLETIN: MUSEUM OF COMPARATIVE ZOOLOGY of tlic horns as well. Such being the case, it may he assumed that the eapsiiies have a role divided into two main functions, me- chanical and physiological. According to Clark (1922) the period of incubation of the eml)ryo rajid is very long, ranging from four to about fifteen months, even under artificial conditions. Therefore, it can !-eadily he inferred that a similarly long period would be needed for the development of the embryo of the deep-sea forms inhabit- ing low temperatures (cf. Table 1). This is probably what might have caused the capsule to be clad in the armour which protects the egg mechanically from natural dangers during the long period of incubation. This hypothesis seems to justify the differentia- tion, at least, of the outer layer in the capsule-wall observed in the northern members, in which the successive changes of the layer, to some extent, hear a I'elation to their geographical distri- bution from the south to the northward. Tiius the trend of dif- ferentiation in the cai)sules of the northern form may be easily traced, if we adopt the hypothesis mentioned above. It seems reasonable to believe, therefore, that the nuich smoother surface and undifferentiated structure of the capsule of Type 1 (Table ■i) has resulted from its adaptation to the environment of the ex- fT-eme southern area where the temperature is relativelv hiiz-b (cf. Tal)lel). In the ca])sules in the southern form, the histological struc- ture differs so much from that of the northern one, that the mechanical theoi-y alone cannot be accepted for this case, but another hypothesis must be developed. The existence of tissue I'omposed of such a delicate structure as the pnlp.y layer which develops in the wall in many of the forms would seem to off'ei- but small resistance towai-d mechanical shock. On the other hand, the puli)y layer should assist the osmosis of the wall thus providing for the needs of a fast-growing endoryo developing under a relatively high temperature. Thus, is it reasonable to assume that the differentiation in the structure of the capsule of the southern form nngiit have been caused by the physiological needs of the embyro? Hypothetical as this may seem, it is logical to trace the trend of differentiation of the cai:)sules in the south- ern form in this way. For example, the capsule of Type 5 has oidy two layers, and its i-eprescntatives ai'e found to iidiabit ISIIIYAIMA: EGG-CAPSUIiES OF JAPANESE SKATES 19 even tlic shallow waters towards the extreme northern regions of our main ishnul. whereas the skates of Type 6, having a eapsule-wall t'onij)ose(.l of a somewhat more specialized structure than the preceding, inhabit areas mostly limited to the southern waters of the land. Since Types 7 and 8 are rather aberrant in the structure of their capsules, it is difficult to develoji a i)lausil)l(' hypothesis based either on the mechanical or on the physiological functiini of the capsule. The likelihood is that both of these processes take place. It may not be unreasonable to assume that the highly speciali7.ed features of these capsules might be linked to the special breeding habits of these species (U and W), the cap- sules of which usually contain more than one egg (ranging up to five in number) and are laid on the sea tioor, as mentioned above. White (1937, pp. 96-97) stated that "the raja-like forms have l)een reported from the Jurassic, so it is possible that this grou]) branched from the squaloid line before the viviparous habit had been established. In all the oviparous groups there is shown some tendency toward ovoviviparity. . ." Thus, oviparous re- production is thought to have occurred independently within groups of the sharks. However, there exist some tendencies in common among the elasmobranchs regarding the structure of the capsules of the oviparous forms. Upon examination and comparison of specimens and of de- scriptions of various kinds of egg-capsules of oviparous sharks, the characters which are shared in common are as follows : 1. The capsules with smooth surface and devoid of any sculp- ture on the surface appear to be reminiscent of the tem- porary capsule of the viviparous fish. 2. The tendrils or the horns are often elongated in a varied manner in both groups. S. The eapsule-wall consists of two layers. Considering these eonnnon characteristics in the capsules, it is credible to assume that the features in a capsule like that of sjiecies A may be expected to display the most primitive con- dition among the Japanese specimens, and this type of capsule appears to have been differentiated in two divergent directions as the specialization of the rajids ])r()gressed (Fig. 9), in contra- diction to White's suggestion. 20 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY At any rate, jud5^'in. Rays and skates, a revision of the European species. Sei. Invest. Fisher. Pxl. Scothmd, no. 1, (>(> iiji. Athis. 3(i pis. Clkjiens, W. a. and (i. V. Wilby 1949. Fishes of the Pacific coast of Canada. Fisher. Res. Bd. Canada, Bull. 68, 368 pp., 253 tigs. DwiKi,, .1. F. 1!»2S. 'I'he elasnioliraiuh lishes. (2nd ed.), xi + 332 i>|>.. I^niv. I'alif. Press. Berkele\', C.-ilifornia. Dkan, Basufokh 1904. Kvolutiou in a deltrniinate line as illustrated by the egg-cases (if cliiniaeroid fishes. Midi. P>ull., vol. 7, pp. 105-112. 19(Mi. Cliiniaeroid fishes and their development. Carnegie Inst. Wash- ington, Pulil. no. 32, 172 pii., 144 figs., 11 pis. 11M2. Oithogenesis in tlu' egg capsules of Cliimai ra. Bull. Anier. Mus. .Xat. Mist., vol. 3], art. 3, i)p. 35-40, 2 figs. Dklacv, .\. ('. and \V. M. Ciiap.max I9.'15. Notes on some elasniobranchs of Puget Sound, with descriptions of their egg cases. Cojieia, 1935, no. 2, ])p. 63-(i7. EvERMANN, B. VV. and E. L. GoLDSBOKorGU 1907. The fishes of Alaska. Hull. U. 8. Bur. Fisher., vol. 26. pp. 219- :'.6(l, 144 ligs., ].ls. 14 42. liAK.MAN, yAiVirEL 1899. Reports on the expedition of the west coasts of Mexico, Central and South America, and oft' the Galapagos Islands, in charge of Alexander Agassiz, by the U. S. Fish Conini. Str. "Albatross," during 1891. Mem. Mus. Comp. Zool., vol. 24, 431 pp. Atlas, 97 pis. 1913). The Plagiostomia. ^Mem. Mus. Comp. Zool., v(d. 36, xiii + 515 pp. Atlas, 75 ]ds. IIOHSOX. A. 1). 1930. A note on the toiniation of the egg case of the skate. Jour. Mai-. Biol. Assoc. T". K., n. s.. vol. 16, no. 2, pp. 577-581, 2 figs. llrs.sAKoK, L. and W. 11. Welkek IIMIS. Notes on flic chemical nature of egg-eases of two s]iecies of sliaiks. ■lour. IJiol. ('hem., vcd. 4, jip. xliv-xh. I8IIIYAMA : HOG-CAPSULES OK JAPANESE SKATES 28 Ht-XLEY, J. S. 1932. Problems of relative growth, xix + -"•' VP-- T^^ tig^'-. 1 I'l. London nnrl Now York. 1SH1Y.\MA, K. 19.^0. Studies on llic r;iys ;uul f^kjites lielonginy to the t';niiil\ Hajidae, found in .l,ip;in and adjacent regions. 1. Rgg-eapside of ten species, .hip.-m. .lour, li-lithyol., \(il. 1, no. 1. pji. .'!ii ;!(). i! figs. ( Tn Japanese with English imuniary. i i;t.")ll. Studies on the rays and skates I>elonging to the family JAajidae, found in .Japan and adjacent regions. 4. A re\isi(in of three genera of .)ai)anese rajids with desciiiitions of one new genus and four new species mostly occurring in northern Jajian. Joui'. Shimonoseki Coll. Fish., vol. 2, no. 2, pp. 1-.S4, 7 tigs., 4 pis. I!t5."). Studies on the rays and skates hehmging to tlie family Rajidae, fouiul in .lapaii and adjacent regions. (>. Hdja uuf-roat uda, a new skati'. .lour. Shimonoscki (dll. Fisli., vol. 4, no. 1, pp. 4o--')l, '1 figs. Jensen, Ad. S. 1914. The selachians of (Greenland. Mindeskr. .lap. Steen trap, vol. _, no. 30. 40 pp.. 1 pi. im^. Contributions to the ichthyorauna of tlreenland. Spolia Zool. Mus. Hauniensis, Skrifter . . . rnivcisitetets Zoolog'ske Museum Kobenhavn. vol. 9, 182 pp.. 4 pis. Jordan. D. S. and E. C. Starks 189."). The fishes of Puget Sound. I'roc. Calif. Acid. S;i., sor. 2, vol. 5, pp. 784-8.'>5, i.ls. 7(vl04. LtDWIG, II. 1874. Uelier die Kibildung ini Thienciclic. Voili. Wiir/bui-giT Phys. nied. Gtsell., vol. 7, 224 \)\k, .'> pN. SCATTERGOOD, L. W. 1951. The occurrence of egg capsules in the winter skate {Eaja (liop)ianr.s) in Maine waters. Copeia, 1951, no. 2. p. 1(39. Springer, S. 1939. The egg ca.se of the Texas skate. Copeia, 1939, no. 4. \>. 23 7. Steven, G. A. 1936. Migrations and growth of the thornback ray {Haia clavata L. ). Jour. Mar. Biol. Assoc. U. K., n. s., vol. 20. iip. (i05-H14, 2 figs. 24 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Vladykov, V. D. 1936. Capsules d'oeufs de Raies de I'atlantique Canadien apparteiiaiit au genre Unjd. Nat. Canad., vol. (iS, pp. 211-231, 7 figs. White, E. G. 1937. Interrelationshiiis of the elasiaol)iaiicl)s with a key to tlie ordei- Galea. Bull. Amer. Mus. Nat. Hist., vol. 74, art. 2, pp. 25-138, ()6 figs., 51 pis. Whitley, G. P. 1938. The eggs of Australian sharks and rays. Austr. Mus. Mag., vol. 6, no. 11, pp. 372-382, 28 figs. Wilson, E. B. 1928. The cell in diveloimuMit and heredity. 3rd ed., xxxvii + 1232 pp.. New York. Fig. 1. Map showing localities where the specimens (egg-capsules) exani- in?d were taken. For alibreviations, see Table 1. I'^iji. 12. Dorsal aspects of the t'g'K-capsulos of the northern forms, A, F, (i and I. Abbreviations as in Table 1. Wliite and black bars represent 5 and 2.5 cm., respectively. Fig:. •">■ Dorsnl aspects of the egg-capsnles of the soutbei'Ti t'oims. T, T'. and \V. Alilireviations as in Talile 1. Wliite bars inclii-ate ."i cm. 200 . 150 100 50 t M '- 1 1 500 1000 Length of body, mm. 15O0 Fig. 4. Size-relation between lengths of the body of jiarent species and of the egg-capsules exclusive of horns, excepting W. Bar shows range in dimensions, and cross shows the arithmetic mean. Al»breviations as in Table 1. A ^^^ T W ^^^i^^g^^^aa^^a^i^^ 4-++V+++V"4+++++4.+4+x+4.+-(.+ Fig. 5. Semidiagrammatic representation of the cross sections of capsule- walls in seven groups Egi-Eg". In these, solid lines represent inner layer; dashes, parenchymatous inner layer; broken ones, pulpy layer; crosses, reticulated pulpy layer; black covering upper border of each section with or without projections (protuberances or prickles) and dots, outer horny layer. A, group EgT^; I, Group Eg-; L, Group Eg^ ; B, Group Eg-*; E, Group Eg'"'; T, Group Eg*'; W, Group Eg''. Magnification, A, x 100; B, I and L, x 24; R, T and W, x 15. Fi<>'. 6. SemidintjiamniMlic icinescntatidii of tlie cross sections of capsule- walls showing varieties as in Fiouic 5. .Magnification, (', E, F, G and K, X la: TI, X 24. Fig. 7. Compnrisou of tlu' ariangoment of priekles developing over tlic surface of the egg-capsulo in three .species, B, E and (!. P> and E, longi- tudinal sections; G, dorsal aspect. Magnification, x 15. U N M O Fig. S. Semidiagrauiniatic representation of the cross sections of capsule- walls, showing varieties as in Figure 5. Groups represented are Eg-*, Eg-"" and Kg'. M. Q, T' and V. x 15; X, O, P and S, x (56. Fig. 9. Dorsal aspects of the egg-capsules, showing the probable phyletic relationship. The filiroid haiis covering the capsule in species F are partly re- moved so as to show the prickles underlying them. Scales accompanied, 5 cm. Northern form Southern form Fig. 10. Suggested phylogeny liased im tlie egg-eapsuleis of Japanese rajid fishes. Each number found in the figure represents the type of the capsule. Bulletin of the Museum of Comparative Zoology AT HARVARD COLLEGE Vol. 118, No. 2 A GENERIC REVIEW OF THE PLOVERS (CHARADRIINAE, AVES) By Walter J. Bock CAMBRIDGE, MASS., U. S. A. PRINTED FOR THE MUSEUM March, 1958 Publications Issued by or in Connection WITH THE MUSETTM OF COMPARATIVE ZOOLOGY AT HARVARD COLLEGE Bulletin (octavo) 1863 — The current volume is ^"ol. 118. Breviora (octavo) 1952- — ■ No. 85 is current. Memoirs (quarto) 1864-1938 — ^Publication was terminated witli Vol. 55. Johnsonia (quarto) 1941 — ^A publication of the Department of Mollusks. Vol. 3, no. 35 is current. Occasional Papers op the Department op Mollusks (octavo") 1945 — Vol. 2, no. 21 is current. Proceedings of the New England Zoological Club (octavo) 1899-1948 — Published in connection with the Museum. Publication terminated with Vol. 24. The continuing publications are issued at irregular intervals in num- bers which may be purchased separately. Prices and lists may be obtained on application to the Director of the Museum of Comparative Zoology, Cambridge 38, Massachusetts. Of the Peters "Check List of Birds of the World," volumes 1-3 are out of print ; volumes 4 and 6 may be obtained from the Harvard Uni- versity Press ; volumes 5 and 7 are sold by the Museum, and future volumes will be published under Museum auspices. Bulletin of the Museum of Comparative Zoology AT HARVARD COLLEGE Vol. 118, No. 2 A GENERIC REVIEW OF THE PLOVERS (CHARADRIINAE, AVES) By Walter J. Bock PRINTED FOR THE MUSEUM CAMBRIDGE, MASS., U. S. A. March, 1958 No. 2 — A Generic Review of the Plovers (Charadriinae, Aves) By Walter J. Bock Biological Laboratories Harvard University For many years the relationships between the grey and golden plovers have been argued about with little agreement between the opposing schools of opinoin. While there has been much discus- sion of the problem, a critical evaluation of the evidence support- ing the maintenance of the genus "Squatarola" as distinct from Pliivialis has never been presented. With this in mind, Dr. Ernst Mayr sugested that I undertake a study of the skull morphology of the large plovers {Pliivialis) so that the earlier works of Lowe could be better evaluated and so that our understanding of the relationships of these species could be further clarified. Prelimi- nary examination of some specimens and study of Lowe's papers on the anatomy and classification of the shorebirds revealed that the variations in the skull morphology and the plumage color and pattern as outlined by Lowe were not limited to Pluvialis, but were common to the entire subfamily. Further study of Lowe's and Peters' classification of the Charadriinae sensu strict a focused attention on the need for a revision of the existing generic arrangement. This need has already been pointed out by Stein- bacher (1932) in his review of Lowe's major paper (1931b), and is reflected by the dissatisfaction of many workers with Peters' classification as indicated by the various, but conflicting proposals to modify his system. Plovers have always held the interest of ornithologists from which it can be said almost ipso facto that many different classi- fications have been advanced for them. Before 1800 the species of plovers were placed in one of two large inclusive genera, Charadrius or Yanellus. The next century was characterized by the proposal of many new genera, almost to the extreme of having only one species to each genus. Seebohm, in his monumental work on the classification and distribution of the shorebirds (1888), objected to this trend toward what he considered a monotypic and impractical generic concept. In his classification, the plovers were placed in three genera, Charadrius (== the Charadriinae of Peters), Vanellus and Lobivanellus (^ the Vanellinae of Peters). 28 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY This arrangement, although it is conservative, and the genera Vanellus and Lohivanellus are artificial, is far more acceptable than the classification in use today. With the publication of volume 24 of the "Catalogue of the Birds in the British Museum" (Sharpe, 1896), the plovers were once again divided into many genera. More importantly, Sharpe 's arrangement of these genera is so unnatural that a clear idea of the generic relationships can- not be gained from study of his work. Lowe's papers on the anatomj', relationships and classification of the shorebirds, in- cluding the plovers, serve as the basis for much of the current classification of the Charadriidae sensu lata. Unfortunately, al- though Lowe did much work on the anatomy of plovers, most of his interpretations are, at best, questionable and have led to an unacceptable taxonomic arrangement. Eensch (1923), in his re- view of Lowe (1922), had suggested that the variation in the skull maj^ well be modified by variations in the jaw muscles or some other factor and that there had been much parallel evolu- tion of the skull within the plovers. The clue to a more reasonable interpretation of the skull variation has been subsequently pointed out by several German workers, but no one has yet done a complete job of checking Lowe's papers and aligning the skel- etal and plumage variations with an acceptable classification of the Charadriinae sensu lato. Peters (1934) corrected some of Lowe's errors, mainly by shifting several misplaced genera from the Vanellinae to the Charadriinae sensu stricto, but in general used Lowe's conclusions as the basis for his classification, which thus still contains most of Lowe's misinterpretations. Peters' two subfamilies are natural (monophyletic) groups but they are sub- divided into far too many genera. In recent years some genera, especially in the charadriine plovers, have been merged — a trend leading back to the classification of Seebohm. However, the merging has been erratic, with little agreement in the delimi- tation of genera, as most clearly shown in the case of Charadrius whose limits differ with almost every author. The merging of the charadriine genera reached its extreme limit with the recent action of the Nomenclature Committee of the British Ornitholo- gists Union (Anonymous, 1949). This committee placed all British plovers, with the exception of Vanellus vanellus, into Charadrius without giving reasons for their action or taking BOCK : GENERIC KEVIEVV OF THE PLOVERS 29 the non-British species into consideration. Even if it were correct, this type of work is unsatisfactory^ for only the opinion of the several workers is presented, without the supporting evidence. Lastly, there has been no recent attempt to understand the posi- tion of the more aberrant species found in the Southern Hemi- sphere or the course of evolution within the subfamily. The aims of this paper are several. Firstly, Lowe's studies on the morphology of plovers are reviewed and a new interpretation is presented in the hope that it will be in agreement with the classification of plovers here presented. Secondly, a study of the relationships and a generic classification of the plovers is ad- vanced. Tliis classification is based mainly on a study of external characters, of habits, habitat, and some features of the internal anatomy, chiefly the osteology. Behavior will not be used in spite of the fine work that has been done on a few species, largely be- cause the behavior of most species is still unknown. This, however, is not to be interpreted as an attitude of underevaluation of the usefulness of comparative ethology in understanding the rela- tionships between species of plovers, for I believe that a compara- tive study of their behavior may prove to be the key to clarifica- tion of the phylogeny within the large genera. Most of the characters used in this work are those that can be seen in study skins. The original survey was done in the Museum of Comparative Zoology and supplemented by study in the Ameri- can Museum of Natural History. I was able to examine all known species of plovers and most of the major plumage variations. Skeletons of a number of species were available, and those studied are listed below : Vanellxi^ vanellus 4 specimens • ' coranattus 2 specimens ' ' gregarius - 1 specimen ' ' chilensis 3 specimens ' ' indicus 2 specimens ' ' tricolor 3 specimens " miles 2 specimens Pluvialis apricariti 1 specimen " dominica 13 specimens ' ' squatarola 29 specimens Charadrins hiaticula 26 specimens " wilsonia 3 specimens ' ' vociferus 16 specimens 30 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Charadrius " , melodus 10 specimens " alexandrinus 4 specimens " montanus 3 specimens These specimens were examined in the collections of the Ameri- can Museum of Natural History, Cornell University, and the Museum of Comparative Zoology, or borrowed from the United States National Museum, the Museum of Vertebrate Zoology at the University of California and the Museum of Natural History at the University of Kansas. In addition, a few alcoholic speci- mens were examined, some specimens of Pluvialis dominica, of P. squatarola and a few species of Charadrius. They were checked for the size and position of the nasal glands. I am deeply indebted to Dr. Ernst Mayr who suggested the original problem, helped and guided the entire study. The offi- cials of the Museum of Comparative Zoology and the American Museum of Natural History were most cooperative and helpful to me in the course of my work. Drs. Friedmann, Pitelka, Tordoff and Sibley kindly made available skeletons that were of great value in the study of the skull morphology. Drs. Ernst Mayr, Dean Amadon, Ernest Williams, Karl Koopmann, Daniel Marien, Robert Dressier, and Mr. Robert Risebrough have read the manu- script and offered many useful comments and suggestions for which I am most grateful. Mr. William Partridge must be thanked in particular for his help in providing information about South American plovers and for translating some important papers written in Spanish. Mr. Terrell Hamilton kindly trans- lated von Boetticher's revision of the lapwings from the French. Miss Patricia Washer is to be credited with the fine drawings of the skull and palate. In any taxonomic paper it is of the greatest importance to state the principles on which the proposed classification is based, but it is not necessary to outline these principles in every paper. The principles followed in this study are the same as those used in my revision of the herons (Bock, 1956). In brief, a broad con- cept of the genus and family is used for this is in closer agree- ment with the present-day species concept and results in a sounder, more easily comprehended classification. BOCK : GENERIC REVIEW OF THE PLOVERS 81 Characters Used Understanding of a taxonomic study depends almost entirely upon a clear presentation of the characters upon which the study is based. It is not enough to give complete and accurate diagnoses of the proposed groups because, unless he is a specialist in the group, the reader usually cannot separate the significant from the non-significant characters or understand how the characters vary. Futhermore, merely to discuss the variation of the characters is still not sufficient. To insure a full understanding of a char- acter, it is necessary to describe and discuss its variation, its function, and how it is correlated with other structures to form character complexes. Character complexes must be treated as units, not as separate entities, for the same selection forces act on all and thus fuse them into a single evolutionary unit. When studied in this manner the selection forces acting on the char- acter and its evolution can be more accurately examined. After the functional and phylogenetic aspects of a character have been separated and analyzed, its taxonomic value can be evaluated on a much sounder basis. The value of a systematic study is greatly increased if the taxonomic characters are evaluated in this way and, although I realize that I fall short of the goal, the characters used in classifying the plovers are presented with these ideas in mind. The Skull Of the several characters used by Lowe in his classification of the plovers, the skull and the color of the back were considered by him to be the most important. Eight skull characters were listed ; however, only the first two were of any importance. These two characters show the most striking variation, which was "correlated" with the color of the back and upon which the classification of plovers was largely based. Since the skull char- acters played so large a role in Lowe's writing, I shall cite them in full and then give a brief summary of his interpretations. In separating the genera Pluvialis and ''Squatarola," Lowe lists the following skeletal characters (Lowe, 1922, pp. 478-482) : ' ' Turning to the skull we find : "(a) That the lacrymals in Squatarola are strikingly different, being 32 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY prominent out-jutting processes, almost Larine or Tringine in appearance; while in Pluvialis their outer margin is rounded and merged into the line of the orbital rim, being continued forwards and inwards in a smooth and somewhat noticeable convexity in a manner somewhat reminiscent of Vanellus (text-figs. 10b & lib). "(b) The interorbital space presents very distinct differences in the two forms. In Squatarola it is narrower both actually and relatively, while the raised corniced and everted orbital rim so characteristic of Pluvialis is not present; moreover, the grooves for the supra-orbital glands are not nearly so deep or defined as in Pluvialis, and the general arrangement here is Larine or Tringine (Text-figs. 10b & lib). In Squatarola there are no an- terior foramina caudad of the lacrymals. They are well marked in Pluvialis, and this seems to be a Charadrijne character. In Squatarola the inner mar- gins of the grooves for the supra-orbital glands meet in the middle line of the vertex, forming a prominent sagittal ridge down the centre. In Pluvialis there is a fairly broad and clearly-marked smooth medial depression down the centre of the interorbital space, which is not encroached by the supra- orbital grooves. "(e) Turning to the palatal plates, we find in Squatarola that the postero-external angle is rounded off (in some specimens much cut away). In Pluvialis the angle is squarer. "(d) In Squatarola the ectethmoid or antorbital plate is somewhat triangular in form, the extero-inferior angle representing the apex. In Pluvialis the antorbital plate has a quadrilateral form. ' ' Adding in a foot- note, ' ' This, at any rate, is evident in perfectly ossified examples. ' ' "(e) In Sqxiatarola the descending process of the lacrymal falls per [lendicularly to just touch the apex of the antorbital plate. In Pluvialis it runs along the outer margin but does not fuse with it. "(f) Turning to a comparison of the maxillo-palatines, we find that in the two forms under discussion these are not identical. In Squatarola they appear to be more closely applied to the pre-palatines, their posterior of free points being little separated from the palatal plate. In Pluvialis the free ends converge towards the middle line and underline the vomer, so that that part of the vomerine process is hidden when these structures are viewed from the palatal aspect. The maxillo-palatines in Pluvialis are also more shell-like concavo-convex structures (or more scroll-like). The attach- ment to the palatal process of the premaxilla is less than is Squatarola. " (g) In Squatarola I have noticed that the dentary margin of the pre maxilla is not completely fused with the corresponding portion of the maxillo-palatine as it is in Pluvialis. This is a Larine as opposed to Plu- vialine character. "(h) In Squatarola the postero-external angles of the basitemporal plate end in two fairly conspicuous downwardly projecting processes of BOCK : GENERIC REVIEW OF THE PLOVERS 33 bone. These processes are but little evident in Pluvialis, but are quite char- acteristic of the Laridae and Sternidae. If well-prepared skeletons of the skulls of the two genera under discussion are compared, these differences are generally apparent. A similar distinction is noted between Larus and Stercorarius." Later in the same paper (p. 483), the species of Charadrius were divided, on the basis of the same characteristics, into two groups, "Leucopolius" (resembling " Squat arola") and Charad- rius (resembling Pluvialis). It should be noted that not all of the species of Charadrius as recognized in this paper or by Peters were included by Lowe in " Leucopolius" or Charadrius; some were placed in other genera. The color pattern and color of the back of these genera were given [op. cit., pp. 483-485) and the latter "correlated" Avith the skull. The light versus dark back color was said to be correlated with the degree of ossification of the supra-orbital rims (= char- acters "a" and "b"). Lowe considered the less ossified skull and light dorsal color primitive ("adumbrated") and the more ossified skull and dark dorsal color advanced. As he put it, the former condition was the initial attempt by nature to produce these characters and the latter was the more complete (finished) product. Thus, relationship on the horizontal level (in the same taxonomic group) is shown by skull type and back color. On the vertical axis (between ancestral and derived groups) rela- tionship is indicated by color pattern. " Squat arola" and "Leu- copolius," in addition to a number of other forms, were combined as the "Pre-Charadriinae, " a primitive group that was con- sidered a subfamily, but never given formal status by Lowe or any subsequent author. In a later paper, Lowe (1933a) again discussed the problem of color and color pattern and here pre- sented a list of eight "pairs" of species, set "A" being pale colored dorsally and having the skull type of " Squat arola," and set "B" resembling Pluvialis in these characters. The relation- ship betw'een the 16 species is as outlined above. That is, each species or genus in set "A" (= primitive subfamily) gave rise to the corresponding species or genus in set "B" (== advanced subfamily), which assumes parallel evolution on a grand scale. Lowe speaks of some forms (his "Pre-Charadriinae") as "living fossils" (1922, p. 488; 1933a, p. 120), and believes that various groups of birds are maintained as they were in past 34 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY geological ages thus allowing ornithologists to establish phyloge- netic series based on living forms. He states further (1933a, p. 114) that in the ontogenetic development of the skull of the advanced group there is a stage in the immature bird when the skull resembles that of the adult "pre-charadriine" plover. He is quite correct in his observations and indeed for a time I be- lieved, misled by an incorrectly identified skull, that the differ- ences between the two skull types were mainly age variations, the "pre-charadriine" condition representing the immature and the Table 1 Character complex 1 Least ossified skull Light dorsal color Ancestral (primitive) Character complex 2 More ossified skull Dark dorsal color Descendant (advanced) ' ' Pre-Charadriinae ' ' Charadriinae ■ ' Sqxiatarola" Pluvialis ■ ' Leucopolhis ' ' Charadrius Set "A" (1933a) Each species or genus of the eight groups listed Set"B" (1933a) Corresponding species or genus The vertical columns represent the two charadriine subfamilies as de- limited by Lowe. They are characterized by having a similar skull and back color. The horizontal levels show ancestral and descendant groups which are bound together by a common color pattern, the pattern being different for each group. BOCK : GENERIC REVIEW OF THE PLOVERS 35 a charadriine" condition representing the fully ossified adult skull. This is not the case, as I discovered later. Lowe, in stating that the skull of the advanced type passed through the "primi- tive" stage in its ontogeny, claimed that (1933a, p. 114) : "This would appear to support my conclusion that the Grey Plover and Kentish Plover are members of a group which may be regarded as antecedent in origin to, or at least more generalized than the more specialized group of which the Golden and Ringed Plovers are representatives." This is a direct application of the theory of recapitulation and as in so many other cases has led to an erroneous conclusion. Lowe always argued very strongly that these characters were not directly affected by the present day environment, but represent the condition inherited unchanged from an ancestral form. Finally, he never stated whether he considered color pattern or back color and skull type as the more important in showing relationships between genera of plovers. Table 1 summarizes Lowe's interpretations of the relationships within the plovers. The subfamilies Vanellinae and Lobivanellinae were established by Lowe in 1922, only to be merged by him in a later paper (1931b). The main difference cited by Lowe between the Vanel- linae and the Charadriinae (including the "Pre-Charadriinae") is the condition of the supraorbital rims, which in the Vanellinae are simply more ossified than in Pliivialis. The use of the more completely ossified nature of the supraorbital rims as the major distinguishing feature of the Vanellinae necessitated placing many obvious charadriine plovers, such as Charadrius vociferus, in the Vanellinae, a move to which many workers objected. The main object in briefly suunnarizing Lowe's interpretations of the variations in the skull and back color in the plovers is to show that any classification based on them would be artificial. Unfortunately space does not permit a clear explanation of all the disputed points which has made the above discussion some- what confusing. Lowe may well be right in some of his conclu- sions (for example, placing the turnstones in the Scolopacinae), but as so much of his work on classification and phylogeny is unsound, all of it must be reviewed before being accepted. In regard to the plovers, I was unable to accept any of Lowe's 36 BULLETIN : MUSEUM OF COMPARA'HVE ZOOLOGY conclusions after a careful consideration of both the evidence and other possible interpretations. The results of the present study force me to suggest that Lowe's conclusions dealing with the anatomy and phylogeny of plovers be ignored in future considerations of the relationships within the Charadriinae sensu lato. Figure 1. Dorsal view of the skull of a) immature golden plover (Pliivialis dominica) , b) intermediate stage golden plover, c) adult golden plover, d) adult lapwing (Fanellus vaneUus), e) adult turnstone {Arenaria interpres) , and f) adult grey plover (Pluvialis squatarola) to illustrate the variation in the degree of ossification of the supraorbital rims. The labels are, from posterior to anterior, the supraorbital rims (S), the groove or foramen for the duct of the nasal gland (G), and the lacrimal bone (L). Figures are approximately life size. BOCK : GENERIC REVIEW OF THE PLOVERS 37 Characters "a" and "b." Lowe did not describe these char- acters with sufficient clarity. Therefore the nature of the varia- tion and correlation of these characters will have to be more clearly outlined before an alternative interpretation can be of- fered. Examination of the differences between " Sqiwtarola" and Pluvial is in these characters reveals that the degree of ossification of the supraorbital rims is the most important factor. In '^Squa- farola," the rims are only slightly ossified, hence the interorbital space is narrower, there is a groove, not a foramen, for the duct of the nasal gland at the anterior end of the groove in which the gland lies, and the lacrimals jut out to the sides. In the adult skull of Pluvialis dominica, the supraorbital rims are more fully ossified, hence the interorbital space is wider, there is a foramen, not a groove, for the duct of the nasal gland at the anterior end of the groove in which the gland lies, and the lacrimals do not jut out to the sides, but merge with the edge of the supraorbital rims in an even curve. In the lapwings, the supraorbital rims are still more ossified with small, but definite grooves for the nasal glands. Thus the interorbital space is very wide, a foramen is present for the duct of the nasal glands, and the edge of the supraorbital rims and the lacrimals merge with one another in a very smooth curve. See Figures If, Ic, and Id which illustrate these structures in Pluvialis squatarola, P. dominica, and Vanel- lus vanellus respectively. Lowe (1933a, p. 114) reported that the skull of the immature J'luvialis doyninica ("the advanced type") passes through a stage that resembles the adult skull of the "pre-charadriine" group. My series of dominica fully supports this observation. The skulls of a very immature, an intermediate, and an adult golden plover are illustrated in Figures la, lb, and Ic. These show an increase in the ossification of the supraorbital rims and with this, a change from the ' ' pre-charadriine " to the " advanced ' ' condition. Deter- mination of the age of these skeletons is based on the total degree of ossification of the skeleton including the supraorbital rims ; no skulls of known age were available. (I have only one specimen of known age, a piping plover [Charadrius melodus] W.B. 432 de- posited at Cornell University, a bird banded as a chick and col- lected seven years later. The bones of this specimen, including the supraorbital rims (see Figure 2e), were completely ossified.) 38 BUXiLETIN : MUSEUM OF COMPARATIVE ZOOLOGY When this ontogenetic change became apparent it was necessary to determine the exact nature of the fully adult (^ossified) skull of P. squatarola. In my series of P. squatarola which contained both immature and adult birds, the skulls of all specimens re- sembled that of the immature golden plover. It is doubtful that Figure 2. Dorsal view of tlie skull of a) adult snowy plover (Charadrius alexandrinus) , b) adult Wilson's plover {Charadrius wilsonia), c) adult piping plover {Charadrius melodus), d) immature killdeer {Charadrius vociferus), e) adult killdeer, and f) adult mountain plover {Cliaradrius montanus) to illustrate the variation in the degree of ossification of the supraorbital rims. Abbreviations as in Figure 1. Figures are approximately life size. a series of almost thirty birds collected at random over all of North America would be composed entirely of immature birds, and indeed, some specimens are certainly adults as shown by the total degree of ossification of the skeleton. It can be concluded that the adult skull of squatarola is similar to the very immature skull of dominica (see Figures la and If). The rims of the im- BOCK : GENERIC REVIEW OF THE PLOVERS 39 mature killdeer (Charadrins vociferus) are less ossified than those of the adult and resemble those of the adult snowy plover (C alexandrinus) or Wilson's plover (C wilsonia) (see Figures 2a, 2b, 2d, and 2e). However there is no basis, as we will see moi-e clearly later, to conclude that the skull of squatarola rep- resents an ancesti;al type ; it merely has less ossified supraorbital rims and if a species such as P. dominica has more fully ossified rims, it has had to pass through a squatarola-like stage sometime in its ontogeny — there is no alternative. The elimination of the possibility that the differences observed are the result of comparing an immature with an adult bird necessitated an investigation of other possible factors that could influence the degree of ossification of the supraorbital rims. Since the roof of the skull is so intimately associated with the nasal (or supraorbital) glands, it would seem reasonable to try and determine whether there is a correlation between the size of these glands and the degree of ossification of the supraorbital rims. This suggestion is not new, but has been previously advanced by several German workers, who in fact have given the best possible answer to the problem of the variation in size of the nasal glands and the correlation between the size of the gland and the degree of ossification of the supraorbital rims, but the value of their work has never been fully recognized. Schildmacher (1932), on the earlier suggestion of Heinroth, showed that in Anas plaiyrhynchus the salt content of the en- vironment directly affected the size of the nasal glands during the life of the individual and hence the morphology of the roof of the skull. In general, the saltier the water, the larger are the nasal glands and the less ossified are the supraorbital rims and the lacrimal bones. The reason for this correlation is of no importance to us at this time, but will be discussed later; the important thing is that an inverse correlation between the size of the nasal glands and the ossification of the supraorbital rims does exist. To show this, Schildmacher conducted a simple, but conclusive experiment. He took ducklings from the same brood and reared half of them with fresh water for drinking, while the other half had salt water. At the end of a year he killed half of each group and prepared the skulls. The birds reared on fresh water had well ossified supraorbital rims and small nasal glands while the salt-water birds had poorly ossified supra- orbital rims and well developed glands. These changes are clearly shown in Stresemann (1927-34, p. 52) who illustrates, after 40 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY Schioler (1925, Danmarks Fugle, Bd. 1), the differences in the skull of the Continental European race and the Greenland race oi* the mallard. The remaining ducks were placed together on fresh water and at the end of the second year they were killed and their skulls prepared. This time all of the skulls were alike and resembled the skulls of the birds raised on fresh water. The changes found by Schildmacher are phenotypic and can be brought about by simply changing the salt content of the water. Schildmacher 's experiments were carefully conducted with all of the necessary controls and there is no reason to doubt his results or interpretations. He points out that the supraorbital rims of the salt-water-dwelling race Anas platyrhynchus con- hosacs of Greenland are less ossified than those of the fresh-water European race platyrhynchus, the differences being comparable to those he obtained in his experiment. Lastly, Schildmacher reported on several eiders {Somateria mollissima) and a marine merganser {Mergus serrator) which were held on fresh water for several years. While the nasal glands did not change as much as in the case of the mallard, they did degenerate slowly in both species. It is not surprising to have a smaller change in the gland of a salt-water bird, for the nasal glands are more important to salt-water species and hence it would be advantageous to have the size of the organ more completely determined genetically and less susceptible to changes in the environment. Technau (1936a, 1936b) studied the nasal gland in the entire class of birds. He showed that one of the functions of the secre- tion of the nasal glands is to protect the mucous membrane of the nasal cavity against the action of salt water.^ With this he con- cluded that if, of two races of the same species or of two closely ^ While this paper was in press, I learned of the studies of Schmidt-Nielsen and his collaborators on the function of the nasal glands of marine birds (Federation Proc, vol. 16 (1): 113-114, 1957; Amer. Journ. Physiologj', April, 1958). They have shown that the nasal glands secrete (excrete) salt thereby enabling marine birds to be independent of fresh water. Schmidt- Nielsen told me (personal communication) that they have not discovered any other function of the nasal glands so that my statement of its function would be incorrect. However, the following argument of the evolution and taxonomic value of the nasal gland and associated structures is still perfectly correct with this newly discovered function of the nasal gland. Indeed, it is easier to see how the size of the gland will alter with changes in the salinity of the environment for as the amount of salt increases, the glands will have (o increase in order to remove the excess salt from the body and vice versa. *" BOCK : GENERIC REVIEW OF THE PLOVERS 41 related species, one were found on salt and the other on fresh water, the size of the nasal gland would differ between the two. Several cases were cited to support this conclusion, as for instance the salt-water species Charadrius hiaticula and the fresh-water C. diihius (1936b, pp. 601-603). The difference in size of the nasal glands in this species agrees with his conclu- sion though not as clearly as would an extreme salt-water species such as C. alexandrinus compared to C. duhius. However, the special problem of variation in the size of the nasal gland in any family was outside the scope of his study, and while Technau presented all the necessary evidence, it remained for another worker to utilize his results to solve the problem of the variation in the ossification of the supraorbital rims in the plovers. Stegmann (1937) in a short note discussed the relationship between "Eupoda" {=Charadrius) a. asiatica and "E." a. veredus which he points out are conspecific, as concluded earlier by Hartert. Yet Lowe had placed these forms in the "Pre-Cha- radriinae" and the Vanellinae, respectively, on the basis of skull morphology. These forms, I should add, constitute one of the pairs of species listed by Lowe in his 1933a paper. Charadrius a. asmticus breeds in areas of salt deserts, veredus in areas of fresh water, and both winter in the interior of Africa. On the basis of this and the results of Technau 's studj^, Stegmann con- cluded that the dissimilarity in the degree of ossification of the supraorbital rims was caused by a difference in the size of the nasal glands resulting from the dift'erence in the salinity of the environment of the two species. Lowe knew of the earlier papers on the nasal glands including Marples' (1932) discussion, but discounted the nasal glands as a possible explanation in favor of his earlier interpretation. However, Lowe's interpretation (1933a, pp. 119-129) has no factual basis and is best rejected in favor of the interpretation outlined by Technau and Stegmann. If our hypothesis is correct, then a tabulation of the habitat and the degree of ossification of the supraorbital rims (or the shape of the skull) should show a definite correlation. What is actually being compared is the salinity of the habitat and the size of the nasal glands. The glands and the supraorbital rims change together (see below). Unfortunately a complete survey of all species cannot be made at this time for the skulls of many species are lacking in our collections as well as the much needed data on the habitat. I shall, however, outline several cases for which the necessary evidence is available. 42 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY Since this study originally started with a consideration of the grey and golden plovers, it would seem fitting to discuss them first. The golden plovers, dominica and apricaria, are predomi- nantly fresh-water birds and have well developed supraorbital rims (Figure Ic). On the other hand, the grey plover {squat a- rola) which is predominantly a salt-water bird, has very poorly ossified rims (Figure If). This agrees with our hypothesis. Of the North American species of Charadrius, montanus and vociferiis are fresh-water birds, wilsonia and alexandrinus are the most extreme salt-water forms, and melodus and hiaticula are intermediate, but are found more on salt water than on fresh water. If the skulls of these species are compared (Figures 2a, 2b, 2c, 2e, and 2f) the close correlation between the degree of ossification of the supraorbital rims and habitat is readily ap- parent. The skull of hiaticula, which is not figured, is almost identical with that of melodus. The lapwings are all strictly fresh-water birds and are even found on dry grasslands. They have the smallest nasal glands and the most ossified supraorbital rims (see Figure Id). The close resemblance between the lapwings and some of the chara- driine species such as Charadrius vociferiis, C. montanus, C. asiaticus veredus, Eudromias morinellus and E. ruficolUs (see Figures 2e and 2f ) , which were placed in the Vanellinae by Lowe, is due to the fact that these species are also strictly fresh- water forms and not because of any close relationship between these species and the lapwings. The relative difference in size of the nasal glands in a fresh- water species {Charadrius vociferiis) and a moderately salt- water species {Charadrius hiaticula) is shown in Figure 3. It is thus safe to conclude that, in general, there is a strong correlation between the habitat and the shape of the skull. The species listed by Lowe in his "Pre-Charadriinae" (those with a squatarola-\ike skull) are generally salt-water birds while the species included in his "Charadriinae" (those with a doniinica-\ike skull) are mainly fresh-water birds. All marine (or salt desert) species do not have one type of skull and all fresh-water-dwelling species a second type, but rather within a group, the coastal (or salt desert) species have less ossified rims BOCK: GENERIC REVIEW OF THE PLOVERS 43 than the fresh- water-living species of that group. This point must be made because some marine species, such as Charadrius melodus or C. hiaticula, have more ossified supraorbital rims than some other marine forms such as Pluvialis squatarola or Charadrius alexandrinus. Figure 3. Diagrammatic sketch of the head of a) fresh-water plover, the killdeer, (Charadrius vociferus) and b) salt-water plover, ringed plover, {Churadrius hiaticula) to show the difference in the size of the nasal glands (G) and their relationship to the eyes (E). One problem remains. What is the basis of the correlation between the size of the nasal glands and the degree of ossification of the supraorbital rims? Support for the glands is easily pro- vided for by the heavy membranes present in this region in those species with poorly ossified rims — a support of bone is not needed. A number of experiments that have a direct bearing on this problem have been reported by Murray (1936) in his general treatise on bone. In the treatment of the functional changes in bone, he discusses (p. 78) the yet unexplained fact that "other pressures can cause either atrophy or at any rate limitation of growth of bone in the direction of the pressure." Blood vessels, tendons and muscles can press against the surface of a bone and restrict growth of the bone at that point. Several experi- ments were cited in which the bone grew after the overlying muscle had been removed. A large nasal gland would exert a similar pressure on the bone of the supraorbital rim and in this way prevent development of bone in this region. This could be experimentally verified by a unilateral removal of the gland and examination of the skull for changes after an appropriate period 44 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY of time. This has not yet been undertaken and the basis for the correlation is still an open question. In summary, the hypothesis is offered that the characters "a" and " b " of Lowe are affected by the size of the nasal glands and hence by the saltiness of the environment. There is a considerable mass of supporting data for this hypothesis, but it is by no means absolutely proven. A complete survey of all species to determine how well the correlation holds remains to be done. As mentioned above, this is impossible at this time because of a great lack of both skulls and habitat information for each species. A listing of the breeding and wintering grounds as well as the migration routes of each species is needed. It is also necessary to know what per- centage of each species or subspecies lives on fresh or salt water, as well as whether the species breeds on fresh water and winters on the coast. An excellent example of the kind of data needed is shown by the map of the breeding and wintering grounds of the European races of Charadrkis hiaticula in Salomonsen (1955, p. 45), who stresses the importance of the wintering area as well as the breeding grounds in understanding the selection forces that are acting on a species. Habits are important, for a bird may be strictly coastal, yet seldom go near the water, as for example Charadrius melodns. Knowledge of the age of the specimen is absolutely necessary because only fully adult skulls may be com- pared. Needless to say, a mixture of adult and immature skulls could completely obscure the picture as it did in the early stages of this investigation. It is impossible at this time to determine exactly how easily a change in the environment could change the size of the nasal gland (and also how closely the size of the nasal gland and degree of ossification of the rims are correlated), but the available evidence indicates that the selection pressure of the environment is quite strong, and if altered, it could readily change the size of the gland. This change may be genetic, non- genetic, or very likely a combination of both. It would be inter- esting and of the greatest value to perform the simple experiment of Schildmacher on several species of plovers. This would at least show whether the nasal glands and supraorbital rims can be phenotypically modified by the environment. Other yet un- known factors may act on this region of the skull, and must not be discounted at this time. BOCK : GENERIC REVIEW OF THE PLOVERS 45 The variation in the amount of ossification of the supraorbital rims has provided an excellent example of an osteological feature that exhibits a very marked change as the result of a relatively minor change in the environment. The character is so readily modified that the variation in the rims has little or no value in showing generic relationships and cannot and will not be used at all in establishing the classification of plovers to be presented later in this paper. It certainly does not have the phylogenetic importance that has been ascribed to it by Lowe. Characters "c" to "h." The remaining skull characters de- scribed above do not seem to be correlated with characters "a" and "b" or w4th each other and therefore will be discussed separately. A careful comparison was made only between Plu- vialis dominica and squat arola which serves as the basis of the following discussions. The ''postero-external" angle of the palatines is highly variable in both species, this variation being in part natural and in part artificial, as the result of preparation. The range of variation appears, however, to overlap completely in the two species. The ectethmoid of squatarola is, in general, triangular, while that of fully ossified skulls of dominica is quadrilateral in shape. If in squatarola there were a slight increase in ossification in the space between the apex of the ectethmoid and the descending process of the lacrimal, then the difference between the two species would disappear. This difference is so slight, and so well within the range of variation of the ectethmoid in Charadrius that it is doubtless of no generic importance. I was unable to see any difference in the descending process of the lacrimal. In each species this process reaches the apex of the ectethmoid to touch or fuse with that bone in a similar way. There is much variation, both individual and artificial (due to preparation), in the structure of the maxillo-palatines, but I was unable to see any constant difference between the two species. The dentary process of the premaxillary is completely fused with the mlkxillo-palatine in both species. However it is entirely possible that a suture may be present in very young individuals. Any difference that may exist is most likely due strictly to varia- tion in age. 46 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY The small process found at the "postero-external" angle of the basitemporal plate is present in both species and may be some- what larger in squafarola. There is, however, a cap of tissue, probably cartilage or collagenous fibers, covering the process, which if lacking in a specimen would produce a large difference in the size of the process. The interesting feature of this process is that the internal process of the articular (medial process) of the lower jaw "articulates" on it. As the lower jaw opens, it apparently rotates on the basitemporal process as well as on the articular surfaces of the condyles of the quadrate. A full discus- sion of this ' ' articulation ' ' will be presented in a separate paper. I have carefully checked these characters in Pluvialis but only briefly in Charadrius. However, there is no evidence to indicate that the differences as given by Lowe hold up in the latter genus. To conclude, these characters show very little difference between groups of species in Pluvialis or Charadrius and do not appear to be of value in showing relationships in those genera. Hence they will not be used in this study. Hind Toe The presence or absence of the hind toe has been considered !iy some authors to be one of the important characteristics in delimiting genera of plovers. The best example is the use of the presence of the hind toe in " Squatarola" as one of the major reasons for separating that genus from Pluvialis. It is commonly believed that, with few exceptions, all species of plovers lack the hind toe, and hence the presence of the hallux in P. squatarola was regarded as a very important feature. However, about half i)f the species of lapwings and a few charadriine plovers also have a hind toe, which fact reduces its value as a taxonomic char- acter. When present, the hind toe is usually very short and somewhat elevated above the plane of the remaining toes and is clearly functionless. The metatarsal for the hallux of squatarola is very small and free floating in the fascia of the rear of the tarsus and thus is usually lost during preparation of the skeleton. In addi- tion, the musculature for this toe appears to have degenerated. 1 have dissected one specimen of this species and found what seem to be the tendons leading to the hallux. They were much BOCK : GENERIC REVIEW OF THE PLOVER.^ 47 reduced but because the reand is given. and reddish brown breast lose much of their importance. This is supported by the condition in Charadrius where the color of the breast bands may be black or rufous in closely allied species or even in the same species. The habits of ohscnra are much like those of dominica (Robson, 1884). The two species of golden plovers {dominica and apricaria) are very similar to one another and their ranges are almost com- pletely allopatric. All of the major works on Palearetic birds state that the ranges of the two overlap in the region of the Yenisei River in Western Siberia (Popham, 1897, p. 192, De- mentiev and Clladkov, 1951, pp. 40, 47) ; hence the two forms must be considered as distinct species. As the amount of overlap is so slight and the two species are so similar, their relationship to one another is best shown if they are placed in the same super- species, as has been concluded by Delacour and Mayr (1945. p. 106). The closeness of this relationship is further supported by the report of an apparent hybrid lietween the two species (Popham. 1900). Charadrius Linneaus (hiaticula) 1758 Hynonymy: Erythrogonys Gould, 1838 (cinctus) ; Eupoda J. F. Brandt, 1845 (asMticus) ; Thinarnis G. E. Gray, 1845 (novaeseelandi-ae) ; Znnihjix Keichenbacli, 1852 (moil eat us) ; Podasocys Coues, 186fi ( mnnfn- BOCK: GENERIC REVIEW OF THE PLOVERS 69 )iius) ; Eupodella Mathews, 1913 {veredus = asiaticus) ; Elseifo Mathews, 1913 (melanops) ; Elseyornis Mathews, 1914 (mclanops) ; See Peters, 1934, pp. 245-246 for complete synonymy. Included species: hiaticiUa, placidiis, cliibius, ivilsonia, vocif- erus, melodus, tJioracicus, pecuarius, tricollaris, alexandrinus, peronii, ve7iustiis, collaris, hicinctus, falklandicus, mongolus. leschenault a , asiaticus, modestus, montanus, melanops, cinctus. luhicollis, and novaeseelandiae . Diagnosis: Small to mefliuni sized plovers, usually with a heavy breast band or a black forehead and black line connecting the bill and the eye, or both. This very characteristic breast and head pattern, so well illustrated in hiaticula, is found in a more or less developed state throughout the genus. In the superspecies asiaticus-nwdestus these markings have largeh^ disappeared, but the last remnants can still be seen. The pectoral bands may be single or double ; often they are incomplete around the breast and exist only as a vertical bar on the shoulder or may be com- pletely absent. The breast band is usually black but may be reddish or rufous. In one species, peronii, the male has a com- plete black band while in the female the band is rufous and incomplete. In a few species the l)lack breast band is complete around the back. More commonly, however, there is a complete white collar on the hind neck which separates the brown or grey crown from the back. The back is dark brown to pale grey in color. Underparts are usually white except for the breast bands, but in a few species, such as modestus, the breast may be reddish in color. A white wing stripe may be present or absent. The central tail feathers are dark brown or grey according to the color of the back while the lateral feathers are white. In a few species such as vociferus the tail pattern has become elaborate. Most species have little or no sexual or seasonal variation in plumage and the immature is similar to the adult. Range: World-Avide. Remarks: Except for the addition of several somewhat aber- rant species, Peters' delimitation of the genus Charadrius is followed in this paper. Like Vanellus, the genus is large and complex and the path to understanding the relationships be- tween the species is full of pitfalls. ]\Iy attempts to arrange the species in a natural order and to discover the relationships be- tween them have met with onlv limited success because of the 70 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY nature of the characters used. The color of the back is strongly selected for as concealing coloration and, contrary to Lowe's claims, is of little use in showing relationships. The number, development and color of the breast bands and head markings a a o 2 ID .2 O -4-> •r-3 § 3 1— < -« S rfi o 3 ■o > S a> I OJ g> O •^-l 't: 13 o « Cj g .M s: 'I K -*-^ S ■^ '% oT 05 C CO -«-' 03 c3 o a 00 b 1 ' o cS •1 a P< r^ 53 • ^ CO o o T3 "U -i.A a> 0) ^ y-^ _cS t3 ■♦-' SO 13 O n3 o •8 £> 2 t3 ,SS (» fl « '*-' .~^ rH o j~ H CC Tt o bp rt in CQ fs; ^H , 13 S •♦* o 2 03 .2 e3 -73 05 ?! o 2 s ■8 •S TS 5 ■K» s o iri ^ • 1— t =0 00 -1-^ S 5S OJ q; •iH •^ •«^ 3 ^ '2 be s 55 ^ 05 >- i. 0, i^ li o J ^ 4~i v> o O Ci c: Q v^ BOCK : GENERIC REVIEW OF THE PLOVERS 71 vary greatly and probably serve as releasers associated with both "courtship" displaj's and species recognition. In proposing the relationships outlined below, I have used mainly the pattern of the head and breast and to a slight extent the color of the back, but always mindful of the many dangers that exist. As in the lapwings, I doubt that it will be possible to discover the course of evolution in Charadrius by a study of museum skins. Nor is internal anatomy likely to provide the answer to the problem. Most probably, the solution will be supplied by a comparative study of their behavior and perhaps other techniques such as serology. It has been impossible to determine which species or char- acteristics are primitive or specialized. However it is useful to designate one species as the basis for comparative purposes, and hiafinda has been selected for this mainly because it is so well known and not because it is considered primitive. The relation- ships within Charadrius are illustrated in Figure 5. Group A. The typical species of the ringed plover group are hiaticula. placidus, duhius, wilsonia, vociferus, and mclodus. Aberrant members are pecuarius, thoracicus and tricollaris. These species are characterized by a rather well developed head and breast pattern. The African pecuarius and thoracicus have the head markings as in hiaticula, but a pectoral band is present only in thoracicus. The most aberrant member of the ringed plovers is tricollaris which has two breast bands but a grey throat and a somewhat different head pattern. Charadrius pecuarius of Africa is very similar to the larger sanctaehelenae of St. Helena. The major differences between the two forms are the larger size and the lack of the tan color on the breast in sanctaehelenae. These two forms are similar in all other respects and there is no reason to consider them as distinct species. Thus it is proposed that they be regarded as eonspecific as they generally were before Peters gave sanctae- helenae specific rank in his "Check-list." The Madagascan thoracicus is also very close to pecuarius and may represent an earlier invasion of Madagascar by a -pre-pecu- arius stock. Later pecuarius invaded Madagascar for the second time so that today the tw^o species are sympatrie. The interesting fact is that thoracicus has a breast band which is a "primitive" 72 BULLETIN : MUSEUM OB^ COMPARATIVE ZOOLOGY trait in this genus ; its loss in pec'uariiis represents a more ad- vanced condition. If Africa is the original home of the species, then this is a case oL" a perij^heral population of a species retaining a primitive characteristic. There is, however, an equally good alternative hypothesis, namely that Madagascar is the ancestral home of the species which invaded Africa and gave rise to pecu- (irius which in turn reinvaded Madagascar. If this were true, it would be the "central" population that retained the primitive character. The ringed plovers hiaticula and sernipalmatus are considered conspecific for the purposes of this paper. A fuller discussion of the status of these forms will be presented in a separate paper. To this complex belongs placidus Avhich is perhaps best regarded for the present as a distinct species, but belonging to the same superspecies as hiaticvla. Group B. The sand plovers embrace the species alexandrinus, peronii, veniistus, collaris, hicinctus and falklandicus. The sand plovers have in general a lighter colored back than that of the ringed plovers and commonly have rufous on the crown or breast. Considering hiaticula as our reference species, alex- andrinus can be derived from it by a regression of the breast bands and a change from a dark to a light-brown back. Char- adriiis alexandriyius, in turn, became a world-wide species and seemed to have given rise to peronii in the East Indies, venustiis in Africa and collaris in South America. These species are so similar to alexandrinus that Avere it not for the fact that each one is sympatric with some race of alexandrinus, they would be considered conspecific with it. The relationships between alex- andrinus, falklandicus (including alticoJa) and hicinctus are more complex and will have to be discussed with some detail. Charadrius falhla7idicus is found from the southern tip of South America north through Patagonia to northern Argentina. The closely related alticola ranges in the high Andes from northern Argentina to Peru. There is no overlap in the ranges of these two forms as given in the latest catalogues (Steullet and Deautier, 1939, pp. 565-566, 567; Ilellmayr and Conover, 1048, pp. 61-64). They are very similar in size and plumage except that there are two very heavy breast bands in falklandicus as compared to the very faint ones of alticola, and that falk- landicus loses its reddisli cfowii and head and breast pattern in BOCK : GENERIC REVIEW OF THE PLOVERS 73 the winter while the winter plumage of alticola is similar to its breeding plumage. These differences are slight compared to the overall similarities between the two birds and as there is no overlap in their ranges, there is no reason why they should not be placed in the same species. Mr. William Partridge of Buenos Aires tells me the distribution pattern of falklandicus and alticola (that is, ranging from the lowlands of Patagonia north into the Andes as far as Peru) is a common pattern of many Patagonian birds. The mountain alticola is similar to collaris which is found in the lowlands of South America and north to Mexico. Except for a difference in size (alticola is larger), and the presence of a single heavy breast band in collaris, the two species are similar. It is possible that alticola is a highland representative of collaris that gave rise to falklandicus, but this is highly unlikely. The closest relative of the subspecies falklandicus is hicinctus of New Zealand. They are almost identical except for the color of the lower breast band which is red in hicinctus and black in falklandicus. The color of the breast bands varies greatly in this group of Charadrius so that the contrast of a black versus a reddish band is not a very important difference. Both species lose the breast bands and head markings in the fall molt. In order to express their great similarity and as they have with little doubt descended from the same common ancestor, falk- landicus and hicinctus will be considered as members of the same superspecies. The problem of dispersal over the water gap between South America and New Zealand will be discussed later. Charadrius hicinctus has probably evolved from an alex- andrinus-like form as shown by its similarity to that species in plumage color and pattern and by the fact that it has recently been reported to have hybridized with the Australian subspecies alexandrinus ruficapillus. A full discussion of the hybrid and its history can be found in Oliver (1955, p. 263). The following account has been abstracted from his discussion. Firstly, it must be mentioned that hicinctus breeds only in New Zealand and that part of the population migrates to Australia each winter. This could be interpreted as an indication that hicinctus invaded New Zealand from Australia. The migration of several European birds now breeding in Greenland and Baffinland offers 74 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY some support for this interpretation, but there is no basis for accepting it as a general hypothesis. One year at the beginning of the breeding season, a female alexandrinus was seen with a male hicinctus in New Zealand. It was assumed that it had flown to New Zealand with a returning flock of hicinctus. The female alexandrinus paired with the hicinctus male and nested. Both birds were seen incubating. After the first set of eggs was washed away by a flood, a second set of two eggs was laid which were later collected. One egg was infertile, the other contained a dead, partly developed embryo. Three years later another pair of female alexandrinus and male hicinctus was seen in the same area. It was implied that the female was the same one that had nested there three years before. Two chicks were raised, one of which was collected when it was a year old, and described. All facts indicate a close relationship between the two species which while they are able to interbreed, are distant enough so that the hybrids are not very viable. It is possible that C. alexandrinus rnficapillus has given rise to hicinctus which in turn reached South America and gave rise to falklandicus and hence to alticola. On the other hand, hicinc- tus and falklandicus may have nothing to do with each other and the similarity between them may be due to convergence. This is entirely possible, but as they are the only species of sand plovers with two heavy breast bands, and unless a similar selec- tion force is shown to exist to explain this convergence, it is far more likely that the two species are related. Considering all of the facts, I would prefer to read the series as alexandrinus rnficapillus — hicinctus — falklandictis - — alticola, and regard the resemblance between alticola and collar-is as the result of parallelism. Peters combined the formerly accepted species alexandrinus, rufi,capillus, marginatus and nivosus into a single species, an arrangement that has been generally accepted. However, there has been some doubt as to whether or not the ranges of alex- andrinus and marginatus overlap. Mackworth-Praed and Grant (1952, pp. 340-342) claim that the two forms are distinct species on the grounds that their ranges overlap in the region of British Somaliland. Meinertzhagen (1954, pp. 478-479) and Chapin (1939, p. 67) agree with Peters and state that there is no over- BOCK: GENERIC REVIEW OF THE PLOVERS 75 lap between alexandrinus and marginatus in either East or West Africa. Further study of the distrilnition of these forms is needed before we can be sure of their status, but for the present the best evidence is that there is no overlap in range and there- fore Peters' classification will be followed. Group C. The mountain or plains plovers, composed of mon- goUis, leschenaidtii, asiaticus, modestus and montanus, have probably evolved from an alexandrinus-Uke ancestor. The rufous crown and head markings of mongolus are similar to those seen in some of the Far Eastern races of alexandrinus. The rufous breast of mongolus is foreshadowed in peronii. Charadrius lesch- enaidtii is almost identical to mongolus and would be regarded as conspecific with that form if they were not sympatric. How- ever, as their ranges are almost allopatric (Dementiev and Glad- kov, 1951, pp. 81, 85), they are placed in the same superspecies to express their close relationship. Closely allied to this super- species is asiaticus which differs from mongolus in its sharp white superciliary line, the black border to the posterior edge of the reddish breast and the faintness of the black line between the bill and the eye, all of which are modifications of the 7non- golus pattern. I have followed Hartert (1912-1921) and De- mentiev and Gladkov (1951, p. 88) in placing asiaticus and vere- dus in the same species. They are extremely similar to one another in size and plumage color and as their ranges do not overlap at all, there is no basis for maintaining them as distinct species. The South American modestus resembles asiaticus except that its throat is grey, not white, and the markings on the head and breast are sharper. I have placed it in the same super- species as asiaticus, in spite of the great gap between the ranges of these species, to show their relationship. The mountain plover, montanus, although it is a plain colored bird, shows its affinities to asiaticus by its white superciliary line, white forehead, and faint black line between its bill and eye. The anterior part of the crown is black as in many of its congeners. The plains plovers are the largest and chunkiest species of Charadrius as well as being the species in which the head and breast pattern is developed the least. In these respects they are similar to Pluvialis and may be the species "connecting" the two genera. 76 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Group D. The Australian melanops appears to be an aberrant offshoot of the ringed plover group for it has a black pectoral band and head markings similar to those of hiaticiUa. Its mottled back and wings are unusual for this genus and set it apart from the other species. Group E. Charadrius cinctus seems to be another aberrant derivation of the ringed plovers. It has a broad black breast band that extends down the flanks to end in a series of red markings on the thighs. The solid brown of the head is continu- ous with the brown of the back which is very unlike the hiaticula pattern of a white collar separating the brown color of the head from that of the back. Groups F and G. The two black-headed species, ruhricollis and novaeseelandiae, are similar in some respects, but probably are not very closely allied. The head and foreneck of ruhricollis are black and contrast with the white hindneck. A black band on the upper back delimits the posterior border of the hindneck. There is no breast band. Instead, a short ventral bar is present on each side of the breast. The black on the head and back is lost in the winter plumage which makes the bird look very much like a nondescript ringed plover. The forehead, sides of the face and foreneck of novaeseelandiae are black and separated from the brown crown hy a narrow white line. There is no breast band ; however, there is a thin black band about the upper back. The bill is slender and is the chief feature separating novaesee- landiae from the other species of Charadrius. Yet the difference between the bill of novaeseelandiae and hiaticula is largely bridged by some species as melanops, fricollaris and thoracicus. Anarhynchus Quoy and Gaimard (frontalis) 1830 Included species: frontalis. Diagnosis: The outstanding feature of this monotypic genus is its unusual bill which bends sharply to the right at its mid- point. The angle of the bend is about 20 degrees and is already present in the chick. The dorsal surface is grey ; underparts are white with a black breast band of even width throughout. For a time it was believed that the band was wider on the left side ; however, this is not so. The tail is grey, sometimes with lighter edges. The flight feathers are dark grey with a faint wing bar ; BOCK : GENERIC REVIEW OF THE PLOVERS 77 the rest of the whig is lighter grey similar to the back. In the fall molt the black breast band and head marks are lost. The immature bird is similar to the adult winter plumage. The \rry- bill breeds inland in the shingly (rockj") river beds; the nest is placed among the rocks (Oliver, 1955, p. 269). During the rest of the 3'ear it is found on mud and sand flats along the coast. Habits and behavior are in all respects like those of Chamdrius (Stead, 1932). Range. Resident in New Zealand; breeds on South Island and winters along the coast of North Island. Remarks: In spite of its remarkable bill, Anarhynchus is a poorly marked genus. In fact, save for structure of the bill which is unique among birds, there would be no basis for separ- ating A)wrhynchu.s from Charadrius. Because of the importance of the bill in the differentiation of Anarhynchus, an inquiry into the feeding habits of the wrybill and the functional significance of the bend in the bill would be most desirable. The habits of the wrybill have been discussed in a number of papers (Potts, 1871, pp. 93-97; Hutton and Drummond, 1923, pp. 216-218 ; Smith, 1926, p. 41 ; Stead, 1932 ; Oliver, 1937 ; and summarized in Oliver, 1955, p. 269). The habitat and distribu- tion of the wrybill which are vital to the problem of the function of its bill are described in the above papers, especially by Stead, and also by Sibson (1943), and Urquhart and Sibson (1952). Yet the feeding habits have never been adequately described. According to Potts (p. 96) the bend in the bill would aid the bird in capturing insects that are found abundantly under the Avater-woru rocks of the river beds of its breeding grounds. Stead's conclusions (pp. 91-92) are somewhat colored by his beliefs, so that, although his evidence supports Pott's earlier statement, he does not believe that the wrybill gains any advan- tage from its deflected bill. Smith (p. 41) says: "in North Island, where the bird migrates in the winter he had observed it sweeping the wet sands with a remarkable scythe-like action of its bill for some minute food supply." Despite the fact that the wrybill is a common and easily observed bird, this coixstitutes our entire knowledge of its feeding habits. A complete descrip- tion of its feeding habits on both the breeding and wintering 78 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY grounds is sorely needed. The use of motion pictures and a statistical approach to the problem would be desirable. Even though the evidence is poor, there is enough to indicate that Anarhynchus utilizes the bend in the bill in two ways. When it is on its rocky breeding grounds, the bend is advantage- ous in obtaining insects found under the rocks. On the mud flats of its wintering grounds, it may make use of the crook in the sweeping motions described hy Smith. Until we have care- ful observations, these suggestion are the best that can be offered. However, it is certain that the bill is used in some special way (s) ; there had to be some selection force (s) responsible for the evolu- tion of this peculiar bill. The second aspect concerns the anatomical features of the skull and how they became modified with the change in the shape of the bill and feeding habits. It would be most interesting to see if the asj^mmetry of the anterior part of the bill is reflected in the hind part of the skull. A thorough study of the functional anatomy and evolution of the deflection in the bill of Anarhyn- chus should provide a most fascinating study of adaptation in the bill of birds. Related to the structure of the bill is the problem of whether this species should be given generic rank. Aside from its bill, the wrybill agrees with Charadrius in all respects. It has with little doubt evolved from some member of Charadrius, and except for the shape of its bill, would be placed in that genus without hesitation. The handling of cases in which a species differs from its nearest relatives in a single character, no matter how remark- able, was discussed in connection with Cochleariu^ in my revision of the herons (Bock, 1956, pp. 31-35). Anarhynchus has not yet given rise to any new radiation of forms and may well represent an evolutionary dead-end. I do not consider frontalis markedly different from Charadrius and it is with much hesitation and reluctance that it is kept in a separate genus, but done only to point out the truly unique structure of its bill. However if the generic limits in the plovers are further broadened, this genus will almost automatically have to be merged with Charadrius. EuDROMiAS C. L. Brehm (morinellus) 1830 Synonymy: Oreophohts Jardine and Selby, 1835 iruficollis) ; Morinellus BOCK : GENERIC REVIEW OF THE PLOVER.S 79 Bonaparte, 1856 (morinellus) . Included species: morinellus and ruficollis. Diagnosis: Medium sized plovers with medium to long bills. The back is mottled, but unlike that of Pluvialis. The crown of the adult is solid brownish and bordered. by a white superciliary line. The breast is uniformally colored, reddish or tan. The wing- is similar to the back in color and pattern and without a wing stripe. Winter plumage {morinellus only ?) lacks the color of the underparts of the breeding plumage. The immature is similar to the winter plumage of the adult. Range: E. morinellus breeds in the tundra and mountains of northern Eurasia and winters in the Mediterranean region and southern Asia. E. ruficollis is a permanent resident in the moun- tains and plains of southern South America. Remarks: The grouping of these species into one genus may well be artificial. However the two species agree in many points of color pattern that are not seen in any other plover. The large gap between the ranges of the two species is a problem, but not an insurmountable one when compared to the many disjunct ranges in other genera. The number of similarities that exist in these two species makes it reasonable to regard tliem as eon- generic unless additional evidence should prove otherwise. Pluvianellus G. R. Gray (socialis) 1846 Included species: socialis. Diagnosis: This medium-sized plover has a solid grey back and white underparts with a broad grey breast band in the female while the breast of the male is mottled grey. The wings are dark grey with a broad white wing stripe much like the wing stripe of Yanellus. The central tail feathers are dark grey; the lateral ones are dirty white. The bill is rather flattened laterally for a plover and is sharply pointed. In some ways the bill resembles that of the turnstones. The habits of this species are given in Goodall et al. (1951, pp. 216-217) and seem to be like those of the rest of the charadriine plovers. Range: Found only in Tierra del Fuego. Remarks: Pluvianellus is a nondescript and rather strange plover. Nothing that can be seen in a museum skin gives any clue to its relationships. I have not seen any anatomical material of 80 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY this species, nor has its anatomy ever been described. Most of its features, except for the broad wing stripe, are more charadriine- like than vanelline-like, but are still not very convincing. I have considered it as allied to the Charadrius group on the basis of past usage rather than on any strong evidence, and should like to emphasize that much more must be known about the anatomy and behavior of this plover before we can be rea- sonably sure of its position. Genera Incertae Sedis Phegornis G. R. Gray (mitchellii) 1846 Included species : mitchellii. Diagnosis: A small bird, about the size of C. hiaticula. The head is dark brown with a narrow white band across the fore- head, continuing around the sides as a superciliary line and completing the circuit about the rear of the head. The back of the neck is reddish-brown while the back is dark brown. Chin and throat are black, the rest of the underparts are barred with black and white transverse strips. The tail is dark brown except for the lateral feathers which are white with dark bars as seen in the tail of many sandpipers {e.g. Tringa solitaria). "Wings are dark broAvn with the secondaries tipped with white. The bill is quite long and thin compared to that of the plovers. Range: High Andes from Peru south to central or southern Chile. Remarks: The relationships of this genus are still obscure and there are good reasons to doubt that it is even a plover. Seebohm placed it with the sandpipers (Scolopacinae) and included can- cellatus {^Aechmorhynclius cancellatus and A. parvirostris of Peters) and leucopterus {=Prosohonia leucoptera of Peters) in the same genus. Sharpe kept the three species in the Scolo- pacidae sensu stricto, but separated them into three genera. In his first paper on plovers, Lowe (1922, p. 491) stated that he did not have anatomical material of mitchellii and hence did not commit himself as to its systematic position.. However in his major work (1931b, p. 743) he placed Phegornis in the Char- adriidae sensu stricto on the basis of its color pattern (no elabora- tion given) and the nature of its maxillo-palatine strut which BOCK : GENERIC REVIEW OF THE PLOVERS 81 is illustrated on page 769 (see Figure 6). The difference between the plovers and the sandpipers in this structure, according to Lowe, is that in the plovers the maxillo-palatine strut meets the jugal bar at right angles, while in the sandpipers the strut runs forward from the maxillo-palatines to meet the jugal bar at a rather sharp angle. In addition to the differences given by Lowe, the maxillo-palatine strut of the plovers fuses to the jugal bar Figure 6. Ventral view of the left side of the palate of a) Eroiia, h) Tringa, c) Phegornis (after Lowe, 1931b, p. 769), d) Arenaria, and e) Pluvialis to show the nature of the maxillo-palatine strut. The anterior end of the palate is at the top, midline is to the left. The labeled structures are : X) the point of junction between the jugal bar and the lateral ramus of the nasal bone (not shown in the drawing;, B) the maxillo-palatine strut, and C) an unnamed strut anterior to the maxillo-palatine strut. Figures arc twice life size. 82 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY at a point posterior to the junction of the jiigal bar and the lateral ramus of the nasal. In the sandpipers, on the other hand, the junction of the strut is at a point where the lateral ramus of the nasal fuses to the jugal bar. This is quite evident in Lowe's drawings but is not mentioned by him in the text. In the plovers, the maxillo-palatine strut is always as described by Lowe. The sandpipers, however, exhibit a considerable amount of variation which was appreciated and described by Lowe. Usually the strut is as described above or a minor variation of it. But in some genera, notably Tringa and its allies, the strut meets the jugal bar almost at a right angle at a point slightly posterior to the junction of the lateral process of the nasal bone and the jugal bar. In spite of its variation in the sandpipers, the nature of the maxillo-palatine strut seems to be a good means of separating the plovers from the sandpipers. In Lowe's draw- ing of Phcgornis on page 769, the process labeled as the maxillo- palatine strut, while it does meet the jugal bar at right angles, is anterior to the junction of the jugal bar and the lateral ramus of the nasal, not i)osterior as in all other plovers. Also the more dorsal parts of the palate included in the drawings on pages 785 and 736 illustrating the strut in other plovers and sandpipers seem to be omitted in this plate. If the drawing of Phegornis is compared to the one showing the palate of Tringa (Figure 6. and see also Lowe, 1931b, p. 375, fig. b), the bone marked as the maxillo-palatine strut in Phegornis seems to correspond to an unnamed process in Tringa which is anterior to the maxillo- ])alatine strut and which meets the jugal bar at right angles, but which is just anterior to the junction of the jugal bar and the lateral bar of the nasal bone. Because of these differences, I would hesitate to definitely label the strut shown by Lowe in his draw- ing of Phegornis as the maxillo-palatine strut seen in other species of plovers and sandpipers, but instead suggest that it corres^oonds to the above mentioned, but unnamed strut in Tringa. I have not seen any anatomical material of this species and until I do, 1 cannot make a more definite statement about the condition of the maxillo-palatine strut in Phegornis. Lowe (1927; 1931 a, b) studied the anatomy of Aechmor- hynchus cancellatus. one of the species considered closest to Phe- gornis by Seebohm. He concluded that Aechmorhynchus was a BOCK : GENERIC REVIEW OP THE PLOVERS 83 sandpiper and most eloselj^ related to the group that he called the Limosinae {Bartramia, Numenius, Limosa, etc.)- In regard 10 the maxillo-palatine strut in this species, Lowe said onl}^ that : "the maxillo-palatine region seems to conform to the arrange- ment seen in the curlews" (1927, p. 129), but further on he says that this region was badly decalcified, thus making the deter- mination of the morphological features very difficult. According to Lowe, the color of Phegornis agrees with that of the plovers, but did not cite any definite points of resemblances. The barred underparts and tail (underside and edges) of Phc- (jornis match the plumage of some sandpipers and are quite unlike any plover. The pattern of the head is, however, similar to that found in many charadriine plovers. It should be stressed that as a general rule, it is unsafe to determine the family status of a bird on the basis of its color pattern. In Phegornis the taxonomic implications of the plumage color and pattern are (certainly unclear. Phegornis is found along mountain streams in pairs or singly where it walks on the rocks looking for aquatic animals under the algae that cover the rocks. It is protectively colored, silent and tame so that it is difficult to see until it flushes at the last possible moment and flies ofi' with strong wing beats. The nest is a depression in the grass, not far from water. This account taken from Goodall et al. (1951, p. 218) is the extent of our knowledge of this species and is of no help in discovering its true position. Peters and other recent workers follow Lowe and assign Phe- gornis to the Charadriinae sensu stricto. No one has studied the anatomy of this species since Lowe and indeed we are still com- pletely ignorant of its morphology. Wliile I believe that future work will prove that Phegornis belongs to the Scolopaeinae, perhaps allied to Aechmorhynchus and Prosohonia, there is no evidence at present to support this belief. I must also empha- size that there is at present no reason other than past usage to retain Phegornis in the plovers. However, the most practical solution is to keep Phegornis in the Charadriinae until evidence proves otherwise, but to remember that its true affinities are still unknown. 84 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Peltohyas Sharpe (australis) 1896 Included species: australis. Diagnosis: The upper parts of this medium-sized bird are mottled Ijrown, much like those of Eudromias. The throat, breast and belly are tannish, the lower breast is reddish brown, and the under tail coverts whitish. The white forehead is bordered be- liind by a black bar extending between the eyes and continued below the eye as a short vertical bar. There is a black breast band that is continuous about the back. The breast band extends down the mid-ventral line as a narrow streak as far as the lower breast. The wings are similar to the pattern of the back with no wing bar. The tail is brownish with lighter outer tail feathers. The immature is like the adult, but lacks the black markings on the head and breast. The hind toe is lacking. The bill has an expanded distal portion, and while it is slightly pointed, it is no more so than the bill of Pluvianellus. Range: Australia. Remarks: This puzzling genus was originally described as a species of Eudromias by Gould and placed in that genus or Charadrius until Sharpe placed it in a separate genus and sub- family of the Charadriidae scnsu stricto. Seebohm included australis in Charadrius near C. {^Eudromias) morinellus, but was not sure of its proper position as indicated by his remark (1888, p. 110) : "It is difficult to say which it most resembles, Charadrius hiaticula, Charadrius riiorineUus or Cursorius hicinc- fus, but its resemblance to the latter is probably an example of analog}' rather than affinity." Mathews (1913-1914, pp. 335-336) placed Peltohyas in the Glareolidae on the I)asis of the shape of the bill, the scutellation of the tarsus, and the flattened nature of the claws. Lowe (1931b, 1). 771) listed several anatomical characters such as the thigh muscle formula, the patagial wing muscles, the feather tracts of the neck, and some aspects of the skull and vertebral morphology in which Peltohyas agrees Avith the glareolids and not with the charadriids. A number of these characters are those given by Gadow (1893, pp. 195-203) and Beddard (1898, pp. 336-350") to separate the two groups and thus may be of considerable tax- onomic importance. BOCK: GENERIC REVIEW OF THE PLOVERS 85 Externally, Peltohyas in all respects resembles the plovers. Tn plumage it is closest to Charadn'us which it resembles in the head markings, breast band and tail pattern, and does not match in any way the plumage of any species of the Glareolidae. The l)ill is that of a plover and is completely unlike the arched, pointed bill of the glareolids. While the scutes of botli the an- terior and posterior surfaces of the tarsus are rectangular, they are not like those of the glareolids, especially the scutes of the l)lantar surface, but rather more like those of some species of Vanellus or Eudromias ruficolUs. A number of species of plovers have rectangular scutes on the front surface of the tarsus, but at best the scutes of the plantar surface are hexagonal. The middle claw of most genera of the Glareolidae is pectinate (lack- ing in Stiltia and rudimentary in Rhinoptilus) , but while the claws of Peltohyos are flattened, the middle claw is not pectinate. Tn view of the strongly conflicting evidence — the external features being charadriine-like while some of the features of the internal anatomy (as reported by Lowe) agree very closely with the Glareolidae — Peltohyas must be placed with the other genera of uncertain position. A careful comparative anatomical study of Peltohyas and the Charadriinae and the Glareolidae is needed before it can be assigned to the proper family. The Turnstones ( " Arenariini " ) The genera Arenaria and Aphriza may be thought of as a tribe of shorebirds of uncertain affinities, " Arenariini." They have been placed either in the Charadriinae sensu lato or in the Scolo- pacinae sensu lato, but a convincing argument for either pro- posal has never been given. Most authors include the turnstones in the plovers, basing their action on such external features as the shape of the bill and the plumage pattern. At present, this is the most widespread opinion. Lowe, on the other hand, cites several features of the skull in which the turnstones agree with the Scolopacinae sensu lato and not with the Charadriinae (see Lowe, 1931b, p. 747). These characters are : the type of maxillo- palatine strut (see Figure 6, and for a discussion of this struc- ture, see above, p. 81), the nature of the articulation of the (piadrate with the skull, and lasth' the shape of the mandibular articular surfaces of the quadrate. I have compared skulls of 86 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Arenaria and Aphriza with skulls of a number of genera of plovers and sandpipers and thus far have confirmed Lov^e's earlier findings and conclusions. The condition of the supraorbi- tal rims in Arenaria is illustrated in Figure le. The rims do not i-esemble those of any of the plovers and are similar to those of sandpipers in the narrow shelves of bone. Within the sandpipers, Lowe places the turnstones closest to the eroliine group and pro- poses the following "evolutionary sequence": Erolia (perhaps more definitely E. maritima or E. ptilocnemis) — Aphriza — Arenaria, a reasonable hypothesis that deserves serious attention. However, Lowe's works need to be checked before they can be accepted and until the anatomy of the turnstones is further studied and compared with that of the plovers and the sand- pipers, they must be regarded as a group of doubtful affinities. Yet for the present there is better evidence for placing the turn- stones in the Scolopacinae sensu lato and they will be considered as members of this group for the purposes of this paper. If future work shows that the turnstones are in reality related to the ]ilovers rather than to the sandpipers, the present evidence indi- cates that they probably would have to be considered as a group distinct from the species studied in this paper; perhaps the two groups would be best regarded as separate tribes. Zoogeographic Considerations A careful examination of the ranges of closely related species of plovers reveals a number of interesting zoogeographic prob- lems. For example, some members of the same species or super- species are separated by ocean gaps of 1000 to 5000 miles. Equally interesting are the species that are confined to a small area some- where in the southern tips of the southern land masses. There are enough problems of general zoogeographic interest to warrant a full discussion of some aspects of the distribution and dispersal of the plovers. In a recent paper, Larson (1957) discusses the past and present distribution of the North Temperate and Arctic shorebirds, including the plovers, to which the interested reader is referred. The present day center of distribution for the lapwings is Africa and for the charadriine plovers, the Holarctic region. All Arctic species migrate south in the winter often as far as BOCK : GENERIC REVIEW OP THE PLOVERS 87 the southern tips of South America, Africa and Australia. Many of these species make extremeh' long flights (1000 miles or more) over the ocean ; an excellent example is the flight of the golden plover {Pluvialis dominica) from Nova Scotia to South America or from Alaska to Hawaii. They may rest on long flights by settling on driftwood (Nicholson, 1928, pp. 126-127), or by swim- ming if they are forced to (Cottam, 1928, and numerous other reports). Except for Phcgornis, which may not even be a plover, most species are highly gregarious or at least occur in small flocks. Lastly, there are a few records of northern species breeding in their winter range : P. dominica in New Zealand (Robson, 1884), which apparentlj^ bred in its winter plumage, nest found on January 9, 1883, eggs hatched two days later ; and the chick of C. leschenaidtii has been found in the region of the Red Sea (Archer and Godman, 1937, pp. 384-385, and Meinertz- hagen, 1954, pp. 482-483). It should be pointed out that .some species of plovers are wintering in the temperate areas of the Southern Hemisphere at a time when the day length is increasing and reaching a maximum in these regions. Nothing is known of the annual cycle of these species and its correlation to day length, and especially the possible effects of wintering in the south temperate zone. Apparently for plovers, ocean gaps are not important barriers to successful colonization of new areas. Two examples of recent invasions may be cited. In 1927, during the winter, large flocks of the European lapwing, V. vanellus, flew from England to Newfoundland and Labrador Avhen they missed their course in a storm (Spencer, 1953, p. 88). Vanellus miles novaeJiollandiae has recently successfully invaded New Zealand from Australia (Oliver, 1955, p. 270). Thus the three species of lapwings found in South America had with little doubt come from the west coast of Africa. The two species chilcnsis and resplendens represent one invasion, and the third species, cay anus, represents a separate invasion. Charadrius alexandrinus has probably also travelled over this route. The close relationship between Pluvialis ohscura of New Zea- land and the Arctic apricaria-doniinica and squatarola can best he explained by regarding ohscura as descended from a gTOup 88 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY of individuals of some species of PUcvialis that remained in New Zealand to breed. The old nesting record of dominica in New Zealand offers some support for this hypothesis. Charadrius hicinctiis (New Zealand) and falklandicics (South America) belong to the same superspecies, but their ranges are separated by many thousands of miles of open ocean. It is possi- ble that the invasion was direct from New Zealand to South America. There is, however, another possibility that may be more likely. It is well known that at times in the past, Antarctica was not alwaj^s covered with ice (Axelrod, 1952 a, b). If the ice at the edges of the Antarctic Continent melted and a tundra-like vegetation developed, there is no reason why plovers should not have bred there and migrated north in the fall. If this is true, then ticinctns could have reached South America by way of Antarctica. More likely falhlandicus and hicinctus differentiated from each other in Antarctica, one migrating north to South America and the other to New Zealand and Australia. This may well be the explanation of the relict nature of the species (ten in number) of Southern Hemisphere plovers that are found today breeding in a small area in the very southernmost tips of the southern land masses. It is of interest that the relict plovers are all related to the present-day Arctic species and that there are no plovers confined to the Cape of Good Hope region of Africa. The gaps between the ranges of Charadrius asiaticus and modestus or between Eudromias morinellus and ruficollis are the largest of any that exist in the plovers. In both cases it is a gap between northern Asia and the southern Andes. But even here, it could be explained by invasion and perhaps a partly relict nature of the southern species. It is hoped that these considerations of the migration and dis- persal habits of the plovers have shown that there is nothing in the proposed classification of the Charadriinae that is in conflict with currently accepted principles of zoogeography. History and Future Studies The shorebirds, including the plovers, being conspicuous birds became well known early in the history of ornithology. A brief survey of the dates of the original descriptions shows that less BOCK : GENERIC REVIEW OP THE PLOVERS 8!) than half a dozen species were discovered in the past century and all were known before the turn of the century. It is quite safe to say that no species of plovers remains to be discovered, but much must still be done before we understand the geographic variation and status of many species {e.g. Charadrius hiaticula). Much has been published on the generic and suprageneric rela- tionships of the Charadriiriae ; however, few of the conclusions have stood the test of time. The same is true of the past anatomi- cal work. "While it is hoped that the delimitation of genera pro- posed in the present study is reasonable, little can be said of their relationships and evolution. We cannot even set limits to the Charadriinae or determine whether several genera such as Phe- gornis, Peltohyas, Aphriza and Arenaria belong to this subfamily or to some other group. Our knowledge of the anatomy of these groups is almost nonexistant so that a good comparative study of the anatomy of the entire shorebird group is sorely needed. Perhaps after this is done, we may gain some understanding of the evolution of the Charadriinae and their position in the Charadriidae. Behavior was briefly mentioned several times in the discus- sions, but never gone into fully. Despite the fine work that has been done on the behavior of several species (Rinkel, 1940 ; Laven, 1940; Laven, 1941; Deane, 1944; Williamson, 1948; Simmons, 1952, 1953, 1955; and Smith and Hosking, 1955), the comparative ethology of the plovers is still in its beginnings and of no help to our understanding of the specific relationships of the plovers at this time. Yet all indications point to the fact that the rela- tionships within the large genera, Charadrius and Vanellus. and perhaps even between the genera will be understood only after their behavior is well known, so that the need and desirabil- ity of behavioral studies comparable to those done on the ducks, gulls and terns cannot be urged too strongly. A knowledge of the ecologj^ even a rough indication of their habitat, is necessary for a proper understanding of several fea- tures of the anatomy and plumage and here again careful studies are not available and are much needed. It can be seen that while much work has already been done, our knowledge of the biology of the plovers must be greatly in- creased if we hope to understand the relationships and evolution 90 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY of this group. This review must then be thought of, not as a definitive study, but rather as a preliminary work with the hope of clearing the path for future studies. Summary The structure and variation of the skull of the plovers were studied. It was shown that the degree of ossification of the supra- orbital rims is strongly correlated with the size of the nasal glands and hence with the salinity of the water, and is of no taxonomio value. Some other features of the skull were also studied. The important earlier works on the classification and anatomy uf the Charadriinae were discussed, especially those of Lowe upon which much of the current accepted classification is based. It is shown that Lowe's interpretations of the morphology of the skull, color and color pattern are at variance with many of the observed facts and with many of the ideas and principles of evolution and classification. Any classification of the plovers resulting from these interpretations would, therefore, be highly artificial. In addition to the skull, the major characters studied were the hind toe, the wattles, the wing spur, the color and color pattern of the plumage, and the osteology. Each character is described and its variation and possible evolution within the subfamily, and its value in the proposed classification is discussed. The im- portance of the habitat is mentioned. A new classification of the plovers, based on a comparative study of the above mentioned characters, is presented. The plovers, following Mayr and Amadou, are considered as a sub- family Charadriinae of the Charadriidae. The subfamilies Chara- driinae and Vanellinae of Peters are dropped and the 56 recog- nized species are placed in 6 genera as compared to the 61 species and 32 genera of Peters. No new species or genera are proposed. A diagnostic description is given for each group within the Charadriinae, but not for the subfamily as a whole. The status of several genera remains uncertain. Phcgornis is retained in the plovers only on the basis of past usage, but it is believed that more study will prove it to be a member of the Scolopacinae. Peltohyas may belong either to the Charadriinae or to the Glareo- lidae ; at present its status is very uncertain. The turnstones and surf-birds, Aretiaria and Aphriza, are for the purposes of this BOCK : GENERIC REVIEW OF THE PLOVERS 91 paper considered as sandpipers, but their position is still doubt ful. A discussion of subspecies lies outside the scope of this review. A brief mention of the zoogeographic implications of the pro- posed classification is given. Lastly, a brief summary of past investigations of the group including an outline of the largest gaps in our knowledge of the biology of the plovers is presented. LITERATURE CITED Anonymous 1949. Twentieth report of the Committee on the Nomenclature . . . Ibis, 91: 508-513. Archer, G. and E. M. Godman 1937. The birds of British Somaliland and Gulf of Aden. 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Catalogue of the birds in the collections of the British Museum, volume 24, London. Sibley, C. G. 1957. The evolutionary and taxonomic significance of sexual dimorphism and hybridization in birds. Condor, 59: 166-191. 96 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY SiBSON, E. B. 1943. Observations on the distribution of the Wry bill in the North Island, New Zealand. Emu, 43: 49-62. Simmons, K. E. L. 1952. The nature of the predator-reactions of breeding birds. Be- haviour, 4: 161-171. 1953. Some aspects of the aggressive behaviour of three closely related plovers (Charadrius) . Ibis, 95: 115-127. 1955. The nature of the predator-reactions of waders towards humans ; with special reference to the role of the aggressive, -escape, - and brooding drives. Behaviour, 8: 130-173. Smith, H. G. 1926. The Wry-billed plover {Anarhynchus frontalis). Bull. Brit. Om. Club, 47: 41. Smith, S. and E. Hosking 1955. Birds fighting. London, 128 pp. VAN SOMEREN, V. G. L. 1956. Days with birds. Studies of habits of some East African species. Fieldiana: Zoology, 38: 1-520. Spencer, K. G. 1953. The lapwing in Britain. London, xii •+- 166 pp. Stead, E. R. 1932. The life histories of New Zealand birds. London, xvi -f 162 pp. Stegmann, B. 1937. Die Nasendriisen von Charadrius asiaticus und Charadrius vere- dus. Orn. Monatsber., 45: 80-81. Steinbacher, F. 1932. Review of Lowe "An anatomical review of the "Waders" ..." Orn. Monatsber., 45: 92-94. Steullet, a. B. and E. A. Deautier 1935-46. Catalogo sistematico de las aves de la Republica Argentina. Obra Cine. Mus. La. Plata, 1: (entr. 1-5), 1-1006. Stresemann, E. 1927-34. Aves. In, Kiikenthal, Willy and Thilo Krumbach, Handbuch der Zoology. Berlin und Leipzig, vol. 7, second half, 899 pp. BOCK : GENERIC REVIEW OF THE PLOVERS 97 Technau, G. 1936a. Der Lage- und Formwechsel der Nasendruse bei den Vogeln und seine niogliche funktionelle Erklarung. Nebst Bemerkungen zur Morphologie der Nasenhohle. Sitz-Ber. Gesell. Nat. Freunde Berlin for 1935: 251-256. 1936b. Die Nasendruse der Vogel. Zugleich ein Beitrag zur Morphologie der Nasenhohle. Journ. f. Orn., 84: 511-617. Urquhart, D. a. and R. B. Sibson 1952. Observations on Wrybill Plover at Karaka. Notornis, 4: 170-172. Vaubie, C. 1951. A study of Asiatic larks. Bull. Amer. Mus. Nat. Hist., 97: 431- 526. White, C. M. N. 1952. Systematic notes on African birds. Ostrich, 23: 43. Williamson, K. 1948. Field-notes on nidifieation and distraction-displays in the golden plover. Ibis, 90: 90-98. MJ Bulletin of the Museum of Comparative Zoology AT HARVARD COLLEGE Vol. 118, No. 3 STUDIES ON THE ANT FAUNA OF MELANESIA. I. THE TRIBE LEPTOGENYINI. II. THE TRIBES AMBLYOPONINI AND PLATYTHYREINI By E. 0. AViLSON CAMBRIDGE, MASS., U. S. A. PRINTED FOR THE :MUSEUM April, 1958 Publications Issued by or in Connection WITH THE MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD COLLEGE Bulletin (octavo) 1863 — The current volume is Vo]. 118. Breviora (octavo) 1952 — No. 85 is current. Memoirs (quarto) 1864-1938 — Puhlication was terminated witli Vol. 55. JoHNSONiA (quarto) 1941 — A ]niblication of the Department of Mollusks. \"ol. 3, no. 35 is current. Occasional Papers op the Department op Mollusks (octavo) 1945 — Vol. 2, no. 21 is current. Proceedings op the New England Zoological Club (octavo) 1899-1948 — Published in connection with the Museum. Publication terminated with Vol. 24. The continuing publications are issued at irregular intervals in num- bers which may be purchased separately. Prices and lists may be obtained on application to the Director of the Museum of Comparative Zoology, Cambridge 38, Massachusetts. Of the Peters "Check List of Birds of the World," volumes 1-3 are out of print ; volumes 4 and 6 may be obtained from the Harvard Uni- versity Press; volumes 5 and 7 are sold by the Museum, and future volumes will be published under Museum auspices. Bulletin of the Museiim of Comparative Zoology AT HARVARD COLLEGE Vol. 118, No. 3 STUDIES ON THE ANT FAUNA OP MELANESIA. I. THE TRIBE LEPTOGENYINI. II. THE TRIBES AMBLYOPONINI AND PLATYTHYREINI By E. 0. Wilson CAMBRIDGE, MASS., U. S. A. PRINTED FOR THE MUSEUM April, 1958 No. 3. — Sttcdies on the Ant Fauna of Melanesia I. The Tribe Leptogenyini II. The Tribes Aniblyoponini and Platythyreini By E. 0. Wilson Biological Laboratories, Harvard University I. THE TRIBE LEPTOGENYINI The tribe Leptogenyini, principally composed of the large genus Leptogenys, is exceptionally well developed in the Mela- nesian region and in terms of individual species and colonies must be counted among the dominant ant groups there. During the present revisionary study three zoogeographic divisions have been recognized as feasible at the species or species-group level within Melanesia : (1) The Leptogenys of western and central Melanesia, from New Guinea to the New Hebrides, containing at least six distinct species groups, most of which are precinctive or of recent Indo- malayan origin. (2) The Leptogenys of the Fiji Islands, composed of a homog- eneous group of six species, apparently all derived from the Papuan-based L. bituberculata stem. (3) The Leptogenys and Prionogenys of New Caledonia, with four species representing at least three phyletic groups, two of which are apparently of Australian origin and the third with uncertain relationships. These three faunal divisions have been accorded separate sys- tematic treatment in the present paper. The fauna of the Moluc- cas has also been reviewed and an account of it included with the first section (western and central Melanesia). There are two reasons for this procedure. First, the Moluccan fauna is closely allied to that of New Guinea, so close in fact as to represent little more than a depauperate extension of it. Second, by taking all of the Moluccan species into account, there is a better chance of obtaining a complete coverage of the New Guinea fauna, since the Moluccas have been more intensivelv collected, and there are 102 BULLETIN' : MUSEUM OF COMPARATIVE ZOOLOGY undoubtedly many species and even genera of ants known at present only from the Moluccas that will eventually be found on New Guinea also.' DESCRIPTION OF COLLECTING STATIONS Most of the material on which this revision is based was col- lected bv the author during field work in Melanesia in 1954-55. In order to document more fully the distributions presented in this and future parts of a proposed review of the Melanesian ant fauna, a gazetteer of collecting stations is given below, including a brief ecological characterization of each locality. The classifica- tion of New Guinea forest tj-pes follows that of Lane-Poole (in P. W. Richards, The Tropical Rain Forest, Cambridge University Press, 1953). NEW GUINEA Bandong, upper Bunhok Valley, Northeast New Guinea {1100 meters). May 26, 1955. Disturbed foothills rain forest near the village, which is located approximately 12 kilometers nortli of Boana. Bisiavumu, near Sogeri, Papua {500 meters). March 15-20, 1954. Field work at Bisianumu was conducted almost entirely in the second-growth foothills rain forest partly surrounding the Government experimental rubber plantation. A few acces- sions were made in similar forest several kilometers farther inland along the Kokoda Trail. Boana, Bunbok Valley, Northeast New Guinea {1100 meters). May 25, 1955. Partly disturbed foothills rain forest on the steep western slope of a hill rising just east of the Boana air- strip. Brown River Road, 1.5 km. south of Karema, Papua. March 8- 11, 1955. Most of the collecting at this locality was conducted in undisturbed lowland rain forest. Several accessions were made in an enclave of eucalyptus savanna several kilometers to the south. 1 To cite an example, the aberrant formiclne genus Meaoxena was previeusly known only from a single specimen of M. mistura Fr. Smith from Batjan, collected iiv Wallace more than a century ago. A second species was recently discovered by the present author at Bisianumu, Papua. WILSON: ANTS OF MELANESIA 1, II 103 Bubia, 12.5 km. northwest of Lae, Northeast New Guinea. March 26-27, May 18-19, 1955. Lowland rain forest, moderately dis- turbed. Upper Bunhok Valley, southern slope of Saruwaged Range. May 27-June 1, 1955. The southern slope of the central portion of the Saruwaged Range was climbed to a point on the crest at approximately 4000 meters. Insect collections were made all along the altitudinal transect, and in all of the major vege- tational belts, including mid-mountain forest, mossy forest, high mountain forest ("tree-line" zone), and alpine savanna. Ants diminished rapidly in the mid-mountain forest belt, above 1500 meters, and appeared to be altogether absent above the lowermost levels of the mossy forest, the la.st colonies being encountered in a clearing at approximately 2500 meters. In this connection it is interesting to note that Dr. J. J. H. Szent- Ivany and a party of native assistants were unable to turn up a single ant during several hours search in mossy forest at 2600 meters on the Asoro-Chimbu Divide, in the Central High- lands (Szent-Ivany, pers. commun.). All present evidence points to the complete absence of a high-alpine ant fauna in New Guinea. Lower Biisu River, near Lae {100-150 meters). March 27, April 28-May 17, 1955. Collecting was conducted principally between the Busu and Bupu Rivers in the area indicated in Figure 1. At the time of my visit this area was covered for the most part by primary lowland rain forest, which was in the process of being high-graded by the South Pacific Lumber Company. Through the courtesy of officials of the Company, I was able to reside for a period of three weeks in a camp on the forest border, and was thus presented with a superb opportunity to collect in the tops of the large trees as they were felled, in forest clearings at various stages of overgrowth, and in and around logs at all stages of decomposition. The ant fauna of the area proved exceptionally rich, and in one section of less than two square kilometers an estimated minimum of 170 species w^as collected. Several short excursions w^ere also made into the extensive grassland fringing the western banks of the Busu River. 104 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Didiman Creek, Lae, Northeast New Guinea. March 27, 29, May 8, June 3, 1955. Moderately disturbed lowland rain forest adjacent to the Government Botanical Gardens at the head of Didiman Creek. (See Figure 1.) LAE 3 KM. Fig. 1. The Lae area in 1955, showing the Didiman Creek and lower liiisii River collecting stations. WILSON: ANTS OF MELANESIA 1. 11 105 Karema. See Brown River Road. Laloki River, 16 km. northeast of Port Moresby, near Little Alt. Lawes, Papua. March 8, 1955. Collecting was limited to the terraced, forested floodplain of the Laloki. Mongi-Mape Watersheds, eastern Iluon Peninsula, Northeast New Guinea. April 2-23, 1955. In the company of Mr. Robert Curtis, of the Territorial Department of Agriculture, Stock, and Fisheries, the author made a three-week collecting triji through the mountainous country of the Dedua and Hube dis- tricts, embracing the headwaters of the Mongi and Mape (Go") rivers. The route of this trip is shown in Figure 2 ; the localities indicated are villages, the elevations of which range from sea level to 1800 meters. Collecting was conducted in the forests and clearings along native foot trails. Nadzah, Markliam Valley, Northeast New Guinea. May 20-22. 1955. The collecting station was a plantation managed by Mr. Keith Smith, located at the southwestern corner of the old wartime Nadzab airstrip, 1.5 km. east of the junction of the Markham and Erap rivers. Most accessions were made in dry broadleaf evergreen forest, marked by dense under- growth, open canopy, and prevalence of palms in all stories. At the time of the author's visit, the area was suffering from a record drought, not having received rain in over two months. On the forest floor leaf litter and mold were thin and dry. while the interiors of the larger rotting logs were generally moist. Port Moresby, Papua. March 5-7, 1955. Collecting was e(jn- ducted in open Eucalyptus woodland in and around Port Moresby, and in Eucalyptus woodland and dense second- growth monsoon forest on the summit of Kini-Kini, a 250- meter-high hill on the southeastern outskirts of the town Saruwaged Range. See Bunbok Valley. NEW HEBRIDES Eight kilometers north of LuganviUe, Espiritu Santo {100-130 meters). January 10, 1955. Second-growth rain forest. Auhert Ratard Plantation, 8 km. southwest of LuganvUle. Janu- ary 7-13, 1955. Collecting was limited to the border and in- 106 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY terior of primary lowland rain forest in the western section of the Ratard Plantation. FIJI ISLANDS Eight miles west of Korovau, on King's Road, Viti Levu. Decem- ber 3, 1954. Second-growth rain forest. Nadala, near Nadarivatu, Viti Levu. December 1-2, 1954. Partly disturbed rain forest on the slope of a steep hill just north of Nadala. The soil was clayey, with numerous fragments of volcanic rock at the surface, and covered by fairly thick leaf 20 KM HELOSBACH FINSCHHAFEN OAGIOU Fig. 2. Southeastern part of the Huon Peninsula, showing the itinerary of a collecting trip in the region of the Mongi-Mape watersheds in April. 1955. litter. The canopy was open, due to native lumbering opera- tions, and the undergrowth was moderately to very dense. Despite the obviously disturbed condition of the forest, the WILSON: ANTS OF MELANESIA I, n 107 local ant fauna was almost entirely endemic in composition, and a large percentage of the known Viti Levu species was encountered during the course of only two days' work. NEW CALEDONIA Anse Vata, Noumea. December 4, 15, 1954. Most collecting was conducted in open Melaleuca and Santalum woods within several hundred meters distance of the Anse Vata beach. A few accessions were made in Ficus groves on the beach itself. Le Chapcau Gendarme, near Yahoue. December 5-7, 1954. Most of the hills in the area immediately to the north of Noumea are covered by open Melaleuca woodland, occupied chiefly by intro- duced ant species, but along the stream running down the southern slope of Chapeau Gendarme there was during the time of the author's visit a section of rich, semi-deciduous angiosperm forest that yielded many of the New Caledonian endemic ants. The forest contained two strata — an upper, discontinuous stratum exceeding twelve meters in height, and a lower, denser stratum of small trees and shrubs ranging mostly three to seven meters in height. Much of the forest floor was covered by bracken, and the leaf litter was generally thin, patchy, and dry. (See Figure 3.) Ciu, near Mt. Canala. December 21, 1954; December 31, 1954- January 3, 1955. Field work was conducted at three stations in the vicinity of Ciu. (1) On the property of Mr. D. Fere, un the west bank of the Canala River at 300 meters elevation, collections were made in an isolated section of broadleaf evergreen forest completely surrounded by pastures. The floor of this woodlot had been badly disturbed by cattle, but many of the endemic New Caledonian ant species were found nesting under deep-set rocks and in the occasional stable ac- cumulations of leaf litter. (2) On the east bank of the Canala River, only several hundred meters from the Fere woodlot, was the start of an extensive stretch of relatively undisturbed broadleaf evergreen forest that was made the second site of intensive collecting. The more remote sections of the forest probably approached a mature (primary?) condition and resembled the Chapeau Gendarme forest in being two-storied. Epiphytes and lianas were uncommon. The floor was well 108 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY insolated and covered in most parts by moist leaf litter up to a meter in depth. Rocks and rotting logs were everywhere common on the ground. (3) The third collecting station was two kilometers southwest of the Ciu Palls, on the trail to MONTAGNE DES SOURCES MT MOU DUMBEA NOUMEA Fig. 3. The Noumea area, southwestern New Caledonia, showing the route t'ollov.ed during collecting in December, 1954, and January, 195,'. Koinde, at an elevation of 500 meters. The forest here was limited to deep stream ^'alleys and was similar in form and composition to that at the second station, apparently differing only in being somewhat denser and moister. Koh {ca. 500 meters). December 20, 1954. This locality is on the eastern slope of the central New Caledonian massif. Col- lections were made in moist broadleaf evergreen forest along a steep stream course crossing the main La Foa-Canala road. Montagne des Sources. December 17, 1954. Collections were made at two stations in this montane area : at 800 meters, mixed Araiicaria-Agathis-angiospeTm forest on the steep val- WILSON : ANTS OF MELANESIA I, II lOJt ley walls at the head of the Dumbea River ; at 1000 meters, a pure stand of second-growth Araucaria forest. Mt. Mou {180-1200 meters). December 10-12, 1955. At lower elevations collecting was conducted near the Bourdinat resi- dence in valley-pocket forest very similar to that on Chapeau 10 KM. I 1 KOH • f SARRAMEA Fig. 4. The Canala area, north-central New Caledonia, showing the route followed during collecting in 1954-55. Gendarme. On December 12 an ascent was made of Mt. Mou to a point on the summit ridge at about 1200 meters. Between approximately 300 and 700 meters the trail led through dense bracken scrub, toward the latter elevation passing into a belt of scattered, low, shrubby trees. Beyond this marginal zone no BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY the summit cloud forest was encountered, dominated by large Araucaria and Podocarpus, and with abundant ground moss and epiphytes. Somewhat higher the cloud forest gave way to true mossy forest, consisting of gnarled, twisted trees averaging only eight to ten meters in height and bearing thick layers of mosses and other epiphytes. Ants were present but relatively scarce in both the cloud and mossy forest zones. Noumea. See Anse Vata. Sarramea {160 meters). December 22, 1954. Most of the acces- sions came from under rocks in moist, well shaded pastureland in the rich farming country around Sarramea. Several stands of seeond-groM'th broadleaf evergreen forest were also visited and proved to contain most of the same aut species as the surrounding cultivated land. Yahoue. See Chapeau Gendarme. REFERENCE COLLECTIONS Most of the available study material of Melanesian ants, both in original and duplicated series, is currently housed in the Museum of Comparative Zoology, Harvard LTniversity (MCZ). Below are listed other principal collections from which material has been drawn for the present study ; these are preceded in each case by the abbreviation by which they are cited in the formal synonymies of the systematic section to follow. (Bishop Museum ! B. P. Bishop Museum, Honolulu (BMNH) British Museum (Natural History), Lon- don (CAS) California Academy of Sciences, San Francisco (Emery Coll.) Emery Collection, Museo Civico di Storia Naturale, Genoa (ForelColl.) Porel Collection, Museum d'Histoire Naturelle, Geneva (OUM) Hope Department of Entomology, Ox- ford University Museum, Oxford, Eng- land (Paris Museum) Museum National d'Histoire Naturelle. Paris WILSON : ANTS OF MELANESIA 1,11 111 iUSNM) United States National Museum, Wash- ington, D. C. ( Yasumatsu Coll.) Collection of Dr. Keizo Yasumatsu, Uni- versity of Kyushu, Japan ACKNOWLEDGEMENTS The author is indebted to Dr. W. L. Brown and Mrs. Eleanor Lowenthal for preparing a large part of his Melanesian collec- tions and shipping them to him for advance study in Europe on his return from the Pacific in 1955. Dr. Brown also supplied a set of preliminary taxonomic notes and inquiries that proved most helpful during examination of type material in the major European collections. As a result of this cooperation, the author was able to firmly identify many of the little known and critical Melanesian forms, as well as settle a number of pressing nomen- clatural problems pertaining to other parts of the world fauna. Appreciation is also expressed to the following individuals for their help in assembling the study collection and granting access to type material: Mr. J. Auber (Paris Museum), Dr. Ch. Fer- riere (Forel Coll.), Dr. H. Gisin (Forel Coll.), Dr. J. L. Gressitt (Bishop Mus.), Dott. Delfa Guiglia (Emery Coll.), Mr. G. E. J. Nixon (BMNH), Dr. E. S. Ross (CAS), Dr. M. R. Smith (USNM), Dr. K. Yasumatsu (Yasumatsu Coll.). Finally, the author wishes to express his gratitude to the following persons for material aid supplied him during the course of field work in Melanesia : Rev. G. Bergmann, Mr. Francois Cohic, Mr. Robert Curtis, Mr. L. J. Dumbleton, Mr. Arnold Himson, Mr. Jacques Rageau, Mr. Aubert Ratard, Mrs. Suzanne Ratard, Mr. Keith Smith, Mr. G. A. V. Stanley, Dr. J. J. H. Szent-Ivany. Their help greatly increased the amount of collect- ing and field study that could be accomplished during my brief stay in the South Pacific. MEASUREMENTS Measurements in the present study follow the conventions already adopted bj' the author and W. L. Brown in previous taxonomic work.^ The dimension head length, over which there 1 See W. L. Brown, 1953, Amer. Midi. Xat., 50, p. 7 : or E. O. Wilson, 1955. Bull. Mus. Comp. Zool., U3. p 22. 112 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY lias been some disagreement in the past, is taken as follows ; maximum length of the head, measured from a transverse through the posteriormost point or points along the occipital border to a transverse through the anteriormost point or points on the anterior clypeal border (after Brown, 1953). THE LEPTOGENYINI OF THE MOLUCCAS AND WESTERN MELANESIA In the area extending from the Moluccas to the New Hebrides, six species groups of Leptogenys are currently recognizable. These are listed below, along with their constituent species and attendant new synonymy. Group of L. caeciliae Viehmejer caeciliae Viehmeyer optica Viehmeyer Group of L. chinensis Mayr bituberculata Emery drepanon Wilson hehrideana Wilson indagatrix Wilson papuana Emery triloba Emery Group of L. diminuta (Fr. Smith) diminuta (Fr. Smith) =: Ponera simillima Fr. Smith = Ponera ferox Fr. Smith = Leptogenys dimimita var. bismarekensis Forel = Leptogenys diminuta subsp. santsohii Mann =1 Leptogenys diminuta var. stitzi Viehmeyer =3 Leptogenys diminuta fruhstorferi var. amboinensi-s Karawa.iew = Leptogenys diminuta frulistorferi var. buruensis Karawajew nitens Donisthorpe oreshia Wilson purpurea Emery violac-ea Donisthorpt WILSON: ANTS OF MELANESIA 1, 11 113 Oroup of L. emeryi Forel nneryi Forel foreli Mann ;= walkeri Donisthorpe trvncatus Mann Group of [.. keyssn'i Viehmeyer keysseri Viehmeyer Group of L. proces^onalis (Jerdon) breviceps Viehmeyer = Pscudoponera lubbocki Donisthorpe =: Euponera niger Donisthorpe Key to the species, based on the worker caste 1 . When the mandibles are in a closed position, there is a gap between the point where their inner borders overlap and the midpoint of the anterior I'lypeal border which is at least as wide as the width of the individual mandibles at their insertions; mandibles linear and with no more than a single tooth besides the apical. '2 When the mandibles are in a closed position, there is little or no gap between the point of overlap of their inner borders and the midpoint of the anterior clypeal border; mandibles varying in shape among species and frequently, but not always, bearing numerous teeth in addition to the apical 9 2. Alitrunk completely lacking standing hairs ; entire body surface opaquely sculptured and covered with pruinose pubescence (PoljTiesia; possibly occurring also in Melanesia as a tramp species) . . . insularis Fr. Smith Alitrunk with abundant standing hairs; body surface smooth and shin- ing at least in part, and lacking pruinose pubescence ?■• 3. Smaller species, the pronotal width 1.14 mm or less; almost all of the head and alitrunk densely f oveate-punctate and subopaque to opaque . 4 Larger species, the pronotal width 1.30 mm or more; almost all of the head and a large part of the alitrunk smooth and shining 6 4. Dorsal face of propodeum only slightly longer than the posterior face (Xew Britain) emeryi Forel Dorsal face of propodeum about twice as long as the posterior face o 114 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 5. Median anterior clypeal projection nearly twice as broad as long (Santa Cruz Islands) truncata Mann Median anterior clypeal projection as long as broad or longer (widely distributed, New Guinea to New Hebrides) foreli Mann (i. When the mandibles are in a closed position, the gap between the point of intersection of their inner borders and the midpoint of the anterior clypeal border is approximately three times the maximum mandible width triloba Emery When the mandibles are in a closed position, the gap described above is about equal to the maximum mandible width or a little less 7 7. The petiolar node about 1.1 X longer than broad, its dorsal surface distinctly tuberculate indagatrix Wilson The petiolar node at least 1.3 X longer than broad, its dorsal surface smooth or at most bearing shallow f oveae 8 8. Petiolar node about 1.3 X longer than broad ; its width measured just behind the anterior margin 0.7-0.8 X the maximum width, which is located just in front of the posterior border; body surface showing bluish reflections papuana Emery Petiolar node about 1.5 X longer than broad; its width measured just behind the anterior border only about 0.5 X the maximum width; body lacking bluish reflections drepanon Wilson 9. Petiolar node compressed antero-posteriorly, forming a narrow, trans verse dorsal crest; head nearly as broad as long; dark brown, robust, small-eyed species breviceps Viehmeyer Petiolar node not compressed antero-posteriorly, its crest longitudinally oriented ; head much longer than broad ; mostly, but not all, jet-black, slender species with moderately large eyes 10 10. Very small, slender species, the pronotal width not exceeding 0.79 mm; the petiolar node strongly compressed laterally and seen from above strongly tapered in an anterior direction 11 Larger species, the pronotal width 0.82 mm or greater, and if approach- ing the smaller size class (pronotal width 0.82-1.17 mm) then the petiolar node is neither markedly compressed laterally nor strongly tapered in an anterior direction 12 11. Head almost entirely smooth and shining (New Guinea) bituberculata Emery Entire head foveate-punctate and opaque (New Hebrides) hebrideana Wilson 12. Showing the following combination of characters: medium-sized (pro- notal width 1.06-1.23 mm), most of the body surface showing strong WILSON : ANTS OF MELANESIA I, II 115 steel-blue or violaceous reflections, the entire head surface finely and densely striate 13 Not showing all of the above characters 14 13. Entire alitrunk covered with long erect hairs ; bluish-violaceous reflec- tions absent or very weakly developed on the gaster and appendages (New Guinea mainland) purpurea Emery Alitruncal pilosity limited to a few short hairs on the declivitous faces of the propodeum; bluish-violaceous reflections well developed on the first several gastric tergites and on the appendages (Waigeo) violacea Donisthorpe 14. Smaller species, the pronotal width not exceeding 1.02 mm. Showing in addition the following combination of characters : petiolar node seen from above shaped like a half -ellipse; sculpturing of the head limited almost entirely to longitudinal striation; the lower halves of the sides of the propodeum longitudinally striate, the upper halves mostly or entirely smooth 15 Larger species, the pronotal width at least 1.17 nun. Not showing all of the additional characters given above 17 15. The area between the posterior margin of the eye and the occipital corner at least partly striate and feebly shining to subopaque diminuta (Ft. Smith) Cephalic area described above lacking striae, completely smooth and shining 16 16. Longitudinal striae just mesad of the eye extending for a short distance posterior to the level of the posterior margin of the eye; head propor- tionately longer, cephalic index 71 ; scapes proportionately longer, scape index 138-145 (Netherlands New Guinea and Waigeo) nitens Donisthorpe Longitudinal striae just mesad of the eye not surpassing the level of the posterior margin of the eye; cephalic index 76; scape index 121- 125 (Solomons) oreshia Wilson I". Distance from the posterior margin of the eye to the posterior margin of the head only slightly greater than the maximum eye length; cir- cumocular sulcus lacking ; entire body smooth and shining Iceysseri Viehmeyer Distance from the posterior margin of the eye to the posterior margin of the head more than twice the maximum eye length; the eye mostly or entirely surrounded by a shallow but distinct sulcus; body in large part sculptured and opaque .18 ll(j BULLETIN; MUSEUM OF COMPARATH'^ ZOOLOGY 18. Ill full face view, the eyes do not quite reach the lateral margins of the head ; the eyes are completely surrounded by the circumocular sulcus . . cacoiliae Viehmeyer In full face view, the eyes slightly surpass the lateral margins of the head ; the circumocular sulcus does not extend around the anterior margin of the eye . optica Viehmeyer Leptogenys bituberculata Emery (Figure 5) Leptogenys bituberculata Emery, 1902, Termeszetr. Fiiz., 25: 160, worker. Original localities: Tamara I., Oudemaine I., and Sattelberg, N.-E. New Guinea. (Syntype examined — Emery Coll.) Material examined. PAPUA: Dobodura (P. J. Darlington); Bisianumu, 500 m. (Wilson, nos. 637, 640, 663). Taxonomic notes. The Dobodura specimens were compared with a syntype in the Emery Collection and found to be nearly identical. The series from Bisianumu, however, differ in their slightly larger size and more rounded propodeum, and may prove to belong to a distinct species. For descriptive data on hituhercvlata, see under the comparative descriptions of L. hehrideana Wilson and L. sagaris Wilson. Ecological note. At Bisianumu, a colony containing about 300 workers and a quantity of brood was found nesting in and under loose leaf litter on the floor of second-growth foothills rain forest. Leptogenys breviceps Viehmeyer (Figure 5) Leptogenys (Lobopelta) breviceps Viehmeyer, 1914, Arch. Naturgesch., 79A(12): 30, fig. 4, worker. Type locality: Wareo, N.-E. New Guinea. Pseudoponera lubbocM Donisthorpe, 1938, Ann. Mag. Nat. Hist., (11)1: 593, 596, fig., worker. Type locality: Mt. Lina, Cyclops Mts., Neth. New Guinea. (Syntype examined — MCZ). NEW SYNONYMY. Euponera (Brachyponera) niger Donisthorpe, 1949, Ann. Mag. Nat. Hist., (12)2: 405, male. Type locality: Maffin Bay, Neth. New Guinea. (Holo type examined — CAS). NEW SYNONYMY. Material examined. N.-E. NEW GUINEA: lower Busu River (Wilson, no. 873) ; Boana, upper Bunbok Valley, 1100 m. (Wil- WILSON : ANTS OF MELANESIA I, H 117 son, no. 1116). NETH. NEW GUINEA : Mt. Lina, Cyclops Mts., 160 m. {Pseudoponera luhhocki Donisthorpe sj^ntype) ; Maffin Bay {Euponera niger Donisthorpe holotype). Taxonomic notes. This is the only known Papuan member of the L. proccssionalis group, the species of which are predom- inantly Indomalayan in distribution. The series collected by myself and a single syntype of Pseudoponera luhhocki Donis- thorpe agree well with Viehmeyer's description of hreviceps, except in slight color characters. Viehmeyer states that hreviceps is colored similarly' to L. processionalis, whereas the series exam- ined by me are slightly darker than proccssionalis, as I conceive that species, and show metallescent reflections not exhibited by processionalis. There is a good possibility that this difficulty may be resolved by the fact that Viehmeyer was using as his standard some species of the taxonomieally difficult processionalis group other than processionalis, e.g. L. iridescens (Fr. Smith). The holotype of Euponera niger Donisthorpe, on loan from the Cali- fornia Academy of Sciences, has been compared with a male of hreviceps from the Busu River and found to be nearly identical. Ecological notes. At Boana a large colony was found nesting in open soil in a partial clearing in second-growth rain forest. The nest was marked externalh^ hy a low, irregular pile of excavated earth about a meter in diameter and bearing multiple entrance holes. At the Busu River a group of workers were found foraging in leaf litter during the day. The workers are un- usually aggressive when disturbed and capable of delivering a powerful, shocking sting. No observations were made to deter- mine whether hreviceps conducts raids on termite nests as do other members of the processionalis group. A male determined as hreviceps was taken at light at the Busu River on May 13, 1955. Leptogenys caeciliae Viehmeyer Leptogenys (Lobopelta) caeciliae Viehmeyer, 1912, Abh. Zool.-anthrop.-ethn. Mus. Dresden, 14: 6, figs. 5, 5a, worker. Type locality: Torricelli Mts., N.-E. New Guinea. This large and striking species is known only from the single type worker. As indicated in the key, it is closely allied to L. optica Viehmeyer. 118 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Leptogenys diminuta (Fr. Smith) Ponera diminuta Fr. Smith, 1857, J. Linn. Soc. Zool., 2: 69, worker.* Tj^pe locality: Sarawak. Leptogenys diviinuta, Mayr, 1867, Tijdschr. Ent., 10: 57, worker, distribu- tion. Ponera simillim-a Fr. Smith, 1860, J. Linn. Soc. Zool., 5 (suppl.) : 104, worker. Type locality: Batjan. NEW SYNONYMY (provisional). Nee Ponera simillima Fr. Smith, op. cit., p. 105 (^ Prionopelta inajmoula Emery). Leptogenys (Loiopelta) simillima, Donisthorpe, 1932, Ann. Mag. Nat. Hist., (10)10: 462. Ponera fcrox Fr. Smith, 1865, J. Linn. Soc. Zool., 8: 70, worker. Type locality: Salawati. NEW SYNONYMY (provisional). Leptogenys (Lobopelta) ferox, Donisthorpe, 1932, Ann. Mag. Nat. Hist., (10)10: 472. Leptogenys (Lobopelta) diminuta var. hismardceiufis Forel, 1901, Mitt. Zool. Mus. Berlin, 2(l,b): 7, worker. Type locality: Muarlin, New Britain. (Syntype examined — Emery Coll.). NEW SYNONYMY. Leptogenys (Lobopelta) diminuta var. papuana Stitz, 1912, Sitzber. Ges. Naturf . Freunde Berlin, no. 9 : 498, fig. 1, worker. Nee Leptogenys papuana Emery 1897; see L. diminuta var. stitzi Viehmeyer 1924. Leptogenys (Lobopelta) diminuta subsp. santschi [!] Mann, 1919, Bull. Mus. Comp. Zool., 63: 299, worker, ergatogyne, male. Type locality: Malapaina I., Three Sisters Group, Solomons. (Sj'ntypes examined — MCZ). NEW SYNONYMY. Leptogenys (Lobopelta) diminuta var. stitzi Viehmeyer, 1924, Ent. Mitt., 13: 310, nom. pro L. diminuta var. papuana Stitz. NEW SYNONYMY (provisional). Leptogenys (Lobopelta) diminuta subsp. fruhstorferi, Karawajew, 1925, Konowia, 4: 276. Leptogenys (Lobopelta) diminuta fruhstorferi var. amboinensis Karawajew, 1925, ibid., p. 277, worker. Type locality: Amboina. NEW SYNONY- MY. Leptogenys (Lobopelta) diminuta fruhstorferi var. huruensis [!] Karawa jew, 1925, ibid., p. 278, worker. Type locality: Tifu, Burn. NEW SYNONYMY. Material examined. This abundant ponerine species ranges from Ceylon and India east to Botel Tobago and south through Indonesia to New Guinea, the Solomons, and Queensland. Below are listed the records from Melanesia that have been verified during the present study. WAIGEO: Camp Nok, 800 m. (L. E. Cheesman). N.-E. NEW GUINEA: Nadzab, Markham Valley (Wilson, nos. 1087, 1106, WILSON: ANTS OF MELANESIA I, II 110 1107) ; Bubia (Wilson, no. 1059) ; lower Busu River (Wilson, nos. 934, 940) ; Mongi River at Sambeang (Wilson). PAPUA: Karema, Brown River (Wilson, no. 539) ; Bisianumn, 500 m. (Wilson, nos. 608, 644, 659). SOLOMONS: Wai-ai, San Cristo- val (W. M. Manni. Significant literature records include Amboina and Burn (^Karawajew, 1925) ; Muarlin, New Britain (Forel, 1901) ; and Malapaina and Ugi, Solomons (Mann, 1919). Taxonomic notes. The status of diminuta is complicated by the existence of at least four closely related species that occupy restricted distributions within its range. On Negros Oriental and Luzon, Philippines, there is a species, apparently unde- scribed, which is distinguished from diminuta by its denser, heavier body sculpturing. A second species occurs in northern Borneo and Sarawak and is distinguished from sympatric sam- ples of diminuta by much feebler body sculpturing and smaller size ; this species may well be Frederick Smith 's Ponera laeviceps. A third species, L. nitens Donisthorpe, is widespread in Nether- lands New Guinea, while a fourth, L. orcshia Wilson, occurs in the Solomons. The possibility exists that sibling species addi- tional to L. nitens occur with diminuta on New Guinea. A single stray worker (or ergatogyne?) from Bisianumn, Papua (ace. no. 644) appears identical with other workers from the same locality except for a much thinner petiolar node ; it has been treated tentatively as a pathological specimen of diminuta but may represent a distinct species. The single definitely associated ergatogyne collected in New Guinea (Nadzab; ace. no. 1106) is distinguished from connidal workers by its slightly larger size, proportionately thinner petio- lar node, and proportionately much larger gaster. The alitrunk is worker-like. Lateral ocelli are developed but the median ocellus is represented only by an empty pit. Geographic variation in the worker caste of L. diminuta is most conspicuous in body sculpturing. In this character at least three separate surface areas show independent variation. In the accompanying table I have indicated the records of the species that I have personally verified or that are adequately described in the literature, and I have attempted to indicate by means of a crude classification of sculpturing form and relative sculpturing density the geographic variation in the three body areas. Most 120 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY GEOGRAPHIC VARIATION IN LEPTOGBNYS DIMINUTA Postocular Lateral Cephalic Pronotal Mesopleural Locality Surface Surface Surface Lengoo, Kwan- lieavily striate feebly heavily striate tung Prov., punctate-striate China Botel Tobago heavily striate completely smooth heavily striate Misamari, Assam feebly striate smooth to feebly • feebly striate shagreened Rangoon, Burma heavily striate feebly feebly punctate-striate punetate-striate Walajanagar, feebly striate completely feebly E. Madras Prov., smooth shagreened India Kandy, Ceylon :eebly striate completely feebly smooth shagreened Batu Caves, heavily striate feebly striate feebly striate Kuala Lumpur, Malaya Pang Mop, heavily striate feebly striate feebly striate Sumatra North Borneo lieavily striate feebly striate moderately and Sarawak with smooth patches striate Pemalang, Java iieavily striate feebly striate with smooth patches feebly striatic Bali lieavily striate moderately striate heavily striate WILSON: ANTS OF MELANESIA I. U 121 Locality Binaluan, n. Palawan Zamboanga, Mindinao, P. I. Cebu and Liniay, Luzon Buru (after Kaiawajew. 1925) Waigeo New Guinea Wai-ai, Solomons Kuranda, Queensland Postocular Cephalic Surface heavily striate heavily striate heavily striate heavily striate heavily striate heavily striate feebly striate heavily striate Lateral Pronotal Surface feebly punctate-striate feebly punctate-striate feebly punctate striate smooth to feebly striate completely smooth smooth to feebly striate completely smooth Mesopleural Surface moderately moderately punctate-striate striate feebly striate moderately striate heavily striate heavily striate moderately to heavily striate heavily striate of the records involve single nest series. This is only a preliminary sketch and will serve merely to indicate the extensive nature of the variation and the marked discordance between the inde- pendent areas. L. diminuia should prove in the future one of the most satis- factory of all ant species for the detailed study of geographic variation. However, the unfortunate circumstance prevails that even the most elementary analysis will be hampered by the large number of infraspecific names that have been attached to this species during its long taxonomic history. In the present treat- ment I have accounted for only those names applied to material 122 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY originating from Melanesia and the Moluccas. Some of the names placed in synonymy, such as Karawajew's Moluccan varieties and subsp. santschii Mann from the Solomons, are clearly geographic variants or less and do not need further comment. Several other names, however, present more complex problems and, in the absence of type material, have been placed in provisional syn- onymy. Their cases are treated individually below. Ponera simillima Fr. Smith. In the original description this species was compared with Ponera laeviceps Fr. Smith, in such a way as to suggest its probable identity with L. diminuta. There is no clue in the descriptions of either laeviceps or simil- lima as to any character by which the latter can be separated from diminuta. Ponera ferox Fr. Smith. In the original description this species was stated to differ from diminuta by its larger size and weaker sculpturing. The present author made a special attempt to locate the type of ferox during a recent visit to the British Museum (Natural History) and Hope Department of Entomology, but was not successful. Lcptogenys diminuta var. stitzi Viehmeyer. This form was described as differing from "typical" diminuta by (1) larger size, (2) somewhat more elongate alitruuk, and (3) less flattened petiolar node. In my opinion the stitzi types probably fall within the extreme range of the New Guinea diminuta population in the first two characters. I have seen no specimens with the precise petiolar node form shown by Stitz in his figure, but an excep- tionally large worker from Sambeang approaches it, and if the node of this specimen is turned slightly so as to be seen from a posterior-oblique view, the outline actually corresponds well to that in Stitz 's figure. Until a species is defined which corre- sponds with Stitz 's characterization, it will probably be best to leave this form in provisional synonymy. Ecological notes. Lepiogenys diminuta has been found in a variety of habitats in New Guinea, from dry evergreen forest at Nadzab to primary lowland rain forest at the Busu River and foothills rain forest at Bisianumu. It sems to favor forest borders and partial forest clearings. Nest sites are variable, from logs oi' various degrees of decomposition to cavities in open leaf litter. WILSON : ANTS OF MELANESIA I, n 123 The following estimates of colony size and composition were made : 1. Ace. no. 1087. May 20-22, 1955. 70 to 80 workers, 10 males, and an undetermined amount of brood, consisting of larvae, one- (juarter to full grown, and cocoons, the latter predominating. 2. Ace. no. 1106. May 20-22, 1955. One ergatogyne, about 150 workers, 10 males, and an undetermined amount of brood, consisting of larvae, one-half to full grown, and cocoons, the latter predominating. 3. Ace. no. 539. March 9, 1955. Somewhat in excess of 300 workers; brood not examined. 4. Ace. no. 60S. March 15-20, 1955. About 200 workers and several males ; brood not examined. The ethology of L. diminuia has already been described in some detail in another paper. ^ The species shows what I have considered to be true legionary behavior, involving botli frequent colony movement between temporary nests and the tendency of the workers to forage in groups. The following prey were recorded in New Guinea: a large adult millipede (Karema), 2 small pentatomid bugs (Nadzab), and a large passalid larva (Nadzab). Leptogenys drepanon Wilson, n. sp. (Figure 5) Diagnosis (ivorker). A large member of the chinensis group, with almost completely smooth and shining body surface and elongate, non-tuberculate petiolar node. Holohjpe ivorker. HW 1.49 mm, HL 2.15 mm, SL 2.68 mm, ML 0.50 mm, CI 69, SI 179, EL 0.38 mm, PW 1.30 mm, petiolar node length 1.03 mm, petiolar node height 0.94 mm, dorsal petiole width 0.69 mm. Mandibles linear, their maximum width (at the level of the junction of the apical and basal segments of the masticatory border) 0.29 mm. Apical tooth narrow but blunt ; apical segment of masticatory border 0.40 mm in length, concave, meeting the basal segment in an obtu.se, rounded angle. When the mandibles are in a closed position the distance between the point of overlap of the masticatory borders of the two mandibles 1 Wilson, E. O., 19-58. The beginnings of nomadic and group-predatory behavior ill the poueriue ants. Evolution, in press. 124 BULLETIN : MUSETIM OP COMPARATIVE ZOOLOGY 8REVICEPS DREPANON BITUBERCULATA SA6ARIS Fig. 5. Heads and petiolar nodes of the workers of selected species of Leptogenys. L. triloba Emery, worker from Bandong, N.-E. New Guinea : L. drepanon Wilson, worker holotype; L. breviceps Viehmeyer, drawn from syntype worker of the synonymous Pseudoponera lubbocki Donisthorpe; L. bituberculata Emery, worker from Dobodura, Papua : L. sagans Wilson, holotype worker. WILSON: ANTS OF MELANESIA 1, U 125 and the anterior clypeal projection is 0.32 mm. Eyes well developed, completely surrounded by a narrow sulcus, their outer margins extending- well beyond the lateral margins of the head when the latter is viewed in full face. Median anterior clypeal i)rojeetion 0.19 mm in length, trapezoidal, tapering slightly anteriorly, bearing three stout bristles 0.04 to 0.05 mm in length on its anterior border. Anterior clypeal border bearing on each side of the median projection a smaller, triangular projection the apex of which forms an angle of slightly more than 90°. Between the median and lateral projections the clypeal border is strongly convex. Posterior margin of head evenly rounded, grading into the lateral margins through a continuous curve. Head broad- ening anteriorly, so that its maximum width is just posterior to the mandibular insertions. Petiolar node as shown in Figure 5. Entire body, including gula and appendages, covered by abundant, long, coarse, predominantly erect hairs. Scapes cov- ered by predominantly oblique hairs, the longest of which are 0.14 mm in length, or more than 0.7 X the maximum scape width, which is 0.19 mm. The longest hairs of the posterior lateral mar- gins of the head are 0.23 mm ; those on the gula are 0.40 mm ; those on the pronotum are 0.44 mm; and those on the petiolar node are 0.22 mm in length. As in L. triloba Emery, the body surface is almost entirely free of underlying pubescence ; the .scapes are also bare, but the funiculi and remainder of the ap- pendages bear variably abundant appressed pubescence. Mandibles feebly and irregularly striate; occiput bearing scattered small, low tubercles ; remainder of the head almost en- tirely smooth and moderately to strongly shining. Entire pro- notum, mesonotum, and propodeum (except the posterior pro- podeal face) smooth and shining. Upper one-fifth of episternum covered by rugae oriented transversely to the long axis of the Ijody ; lower four-fifths of episternum and entire metapleuron covered by obliqueh^ oriented, anteriorly descending rugae. Basal two-thirds of posterior propodeal face covered by trans- verse costulae. Petiolar node nearly smooth and moderately shining, bearing broad, extremely shallow, indistinct, contigu- ous depressions which give it a very feebly wavy outline when viewed from the side. Gastric tergites completely smooth and shining. 126 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY Entire body, except mandibles and apical gastric segments, jet-black. Coxae also jet-black. Antennae, apical gastric seg- ments, and leg segments distal to the coxae medium to dark brown. Material examined. N.-E. NEW GUINEA : Bialowat, Morobe District (II. Stevens) ; a single worker. Taxonomic notes. L. drepanon is closely related to L. indaga- trix AVilson and can be distinguished by the characters given in the comparative description of that species. It is also close to L. papuana Emery, from which it can be distinguished by its more slender, tapering petiolar node, and lack of bluish surface reflections. Prom the more distantly related L. triloba Emery it differs by its smaller size, shorter mandibles, different clypeal outline, and longer and denser pilosity. Leptogenys emery'i Forel Leptogenys {Leptogenys) Emeryi Forel, 1901, Mitt. Zool. Mus. Berlin, 2(l,b): 7, worker. Type locality: Lowon Valley, near Ralum, New Britain. Known from the holotype worker only. This species is closely related to L. foreli Mann, a species widely distributed in western and central Melanesia, but should be easily distinguished by the distinctive propodeal proportions as given by Forel in his orig- inal description. Leptogeny^s FORELI Mann Leptogenys {Leptogenys) foreli Mann, 1919, Bull. Mus. Comp. Zool., 63: 297, tig. 9 {nee fig. 10, which is labelled as this species but is actually L. tnincata), worker, male. Type locality: Malapaina I., Solomons. ( Syntypes examined — USNM ) . Leptogenys {Leptogenys) walkeri Donisthorpe, 1942, Ann. Mag. Nat. Hist., (11)9: 704, worker. Type locality: Y^ila, Efate, New Hebrides. (Holo type examined — BMNH). NEW SYNONYMY. Material examined. NETII. NEW GUINEA: Maffin Bay (E. S. Ro&s). SOLOMONS: Malapaina (type locality). NEW HEBRIDES: Vila, Efate {walkeri holotype); Aubert Ratard Plantation, Espiritu Santo (Wilson, no. 237). Mann (1919^ also records foreli from Auki and Simoli, in the Solomons. WILSON: ANTS OF MELANESIA I, IT 127 Taxonomic notes. The Maffin Bay series differs from the foreli types and New Hebrides series in having slightlj- longer anterior genal teeth and sparser appendage pilosity. Ecological notes. Ross (unpublished notes) found the Maffin Ba.y colony nesting in a sago pahn trunk. The colony that 1 found on Espiritu Santo was nesting in a large rotting log, partly elevated off the ground, in primary lowland rain forest. It con- tained approximately 25 workers, which were extremely timid and agile. Leptogenys hebrideana Wilson, n. sp. Diagnosis {worker). Closely resembling L. bituberculata Emery of New Guinea, showing the following principal differ- ences : (1) The entire surface of the head is coarsely sculptured and opaque (as opposed to completely smooth and shining in bituber- culata), with the sculpturing showing the distinctive pattern described herewith : All of the clypeus but the central raised portion is longitudinally costulate. Posteriorly the costulae ex- tend onto the frons, where they turn into rugae, which in turn form into a rugoreticulum in the area between the eye and anten- nal insertion. At about the level of the posterior margin of the eye, the rugoreticulum gives way to flat, shallow foveae, 0.02 to 0.05 mm in diameter and separated from one another by distances of about the same magnitude as the diameters. The bottoms of the foveae are densely punctate, whereas the inter- foveal spaces are sparsely punctate to smooth. The entire gular area is covered by similar foveae. At the dorsal occipital zone the foveae are crowded together and the interfoveal spaces form a rugoreticulum. (2) In hebrideana the anterior half of the pronotum and entire dorsal surfaces of the mesonotum and propodeum are irregularly pitted and furrowed, while in bituberculata the same area is completely smooth and shining. In hebrideana the entire lateral surface of the alitrunk is coarsely sculptured, with oblique rugae predominating, whereas in biturberciilata sculptur- ing is limited to the immediate vicinities of the metapleural- propodeal suture and metapleural gland. 128 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY (3) Both appendage and body pilosity are much denser in hebrideana. (4) The petiolar node is proportionately shorter in hebride- ana. Holotype worker. HW 0.86 mm, HL 1.40 mm, SL 1.58 mm, CI 61, SI 184, EL 0.34 mm, PW 0.75 mm, petiole node length 0.74 mm, petiole node height 0.45 mm, dorsal petiole width 0.41 mm. Paratype worker. HW 0.90 mm, HL 1.43 mm, SL 1.61 mm, CI 03, SI 179, PW 0.76 mm. Material examined. NEW HEBRIDES: A. Ratard Planta- tion, near Luganville, Espiritu Santo (Wilson, no. 348 ; 2 work- ers). Taxonomic notes. L. hebrideana also bears a close resemblance to L. foveo punctata Mann, L. humiliata Mann, and L. navua Mann, all of the Fiji Islands. The Fijian species have foveate head surfaces as in hebrideana, but humiliata and navua are considerably smaller in size and have nearly smooth alitruncal dorsa, while fovcopiinctata is larger than hebrideana and is further distinguished from all other members of the bitubercu- lata complex by its coarsely sculptured petiolar node. There are other characters but the above should suffice for diagnostic pur- poses. Ecological note. The two type workers of hebrideana were taken as strays foraging on the floor of undisturbed lowland rain forest. Leptogenys indagatrix Wilson, n. sp. Diagnosis {worker). Closely related to L. papnana Emery and L. drepanon Wilson, but easily distinguished from these two species by the following characters : (1) Smaller size (see measurements of holotype worker). (2) The petiolar node is proportionately shorter. (3) The surface of the petiole is covered by relatively small, non-contiguous, rounded tubercles ; the intermediate areas are feebly and irregularly shagreened and their surfaces only feebly shining. WILSON: ANTS OF MELANESIA I, II 129 (4) The head surface, but no other part of the body, bears very faint bluish reflections. L. drepanon completely lacks sur- face reflections, while they are prominently developed on the body (and head also?) of papuana. The following- additional difference between indagatrix and drepanon is noteworthy : in indagatrix the pilosity of the dorsal propodeal and petiolar node surfaces is predominantly oblique, the individual hairs set at an angle at about 45° from the cuticular surface, whereas in drepanon the pilosity is pre- dominantly erect, the individual hairs set at an angle of about 70° ; the comparison cannot be extended to papuana in the absence of material belonging to that species. Holotype worker. HW 1.44 mm, HL 2.02 mm, SL 2.48 mm, (,'I 71, SI 172, EL 0.41 mm, PW 1.26 mm, petiolar node lenglh 1.06 mm, petiolar node height 1.00 mm, dorsal petiole width 0.86 mm. Worker paratype variation. HW 1.38-1.43 mm, HL 1.94-2.00 mm, SL 2.40-2.46 mm, CI 71-72, SI 171-175. Material examined. PAPUA: Bisianumu, 500 m. (Wilson, no. 661) ; holotype and six paratype workers. Ecological note. The types were collected together from the floor of somewhat disturbed foothills rain forest. Leptogenys insularis Fr. Smith Leptogenys insvlaris Fr. Smith, 1879, J. Linn. Soc. Zool., 14: 675, worker. Type locality: Oahu, Hawaii. Although this distinctive tropicopolitan tramp species has not yet been recorded from Melanesia, it has been included in the key and mention made of it here on the basis of the high order of probability that it does occur somewhere in the area. Leptogenys keysseri Viehmeyer Leptogenys (Lobopelta) Tceysseri Viehmeyer, 1913, Arch. Naturgesch., 79 A (12): 29, fig. 3, worker. Type locality: Sattelberg, N.-E. New Guinea. Known from type material only 1 30 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Leptogenys nitens Donisthorpe Leptogenys (Lobopelta) nitens Donisthorpe, 1943, Ann. Mag. Nat. Hist., (11)9: 169, worker. Type locality: Camp Nok, Waigeo, Neth. New Guinea. (Syntypes examined — MCZ). Ibid., pp. 444-445, male, doubt- fully associated. Material examined. WAIGEO : Camp Nok (lectotype). NETH. NEW GUINEA: Maffin Bay (E. S. Ross). Taxonomic notes. One of two syntype workers deposited in the Museum of Comparative Zoology belongs to the typical Papuan form of L. diminuia. The other belongs to a distinct, related species to which the name L. nitens is correctly applied. In order to stabilize the nomenclature, the latter specimen is hereby designated the lectotype of nitens. This species resembles L. oreshia Wilson of the Solomons in its reduced cephalic sculpturing but is easily distinguished by the fact that the longi- tudinal striae mesad of the eye extend well beyond the level of the posterior margin of the eye, whereas in oreshia they just attain the level of the posterior margin. Nitens further differs from oreshia, and resembles diminuta, in its smaller size, dis- tinctly lower cephalic index, and proportionately longer scapes. The lectotype has the following cephalic proportions: HW 1.23 mm, HL 1.72 mm, SL 1.70 mm, CI 71, SI 138. The largest of four workers examined from the Maffin Bay series has the fol- lowing cephalic proportions: HW 1.26 mm, HL 1.78 mm, SL 1.83 mm, CI 71, SI 145. Leptogenys optica Viehmeyer, new status Leptogenys (Lobopelta) caeciliae var. optica Viehmeyer, 1914, Zool. Jahr. Syst., 37: 609, worker. Type locality: Wareo, N.-E. New Guinea. (Syntype examined — Forel Coll.). Material examined. N.-E. NEW GUINEA: Wareo (syntype) ; lower Busu River (Wilson, no. 927) ; Zingzingu, Mongi Water- shed, 1200 m. (Wilson, no. 761). Taxonomic notes. Workers from the Busu River differ from the syntype in the Forel Collection in the following manner : in the type, the sculpturing of the occipital region consists of transverse rugae, whereas in the Busu specimens it consists of an irregular rugoreticulum with no tendency toward a transverse orientation. WILSON : ANTS OF MELANESIA I, 11 131 In addition to the characters given in couplet 18 of the key, L. optica can be distinguished from the closely related L. caeciliae by the following characters : eyes larger and more convex, petiolar node lower, first gastric tergite more densely punctate. Ecological notes. At the Busu River a small colony of this remarkable ant was found nesting in and under a large, partly buried "Zoraptera-stage" log on the floor of primary lowland rain forest. Workers and brood occupied several large galleries in the lower part of the log itself, as well as several smaller gal- leries that passed laterally from the bottom of the log into the soil. At Zingzingu a single worker was found in late afternoon crossing a foot trail that led through a clearing in second-growth midmountain forest. The workers are unusually sluggish and timid for Leptogenys. Leptogenys oresbia Wilson, n. sp. Leptogenys (Lobopelta) diminuta var. laeviceps, Mann, 1919, Bull. Mus. Comp. Zool., 63 : 300. Nee Ponera laeviceps Fr. Smith. Diagnosis (ivorker). Closelj- related to L. diminuta (Fr. Smith), L. nitens Donisthorpe, and the writer's present concep- tion of L. laeviceps (Fr. Smith ),^ but differing in the following characters : (1) Larger size. The HW of a series of diminuta from Wai-ai, San Cristoval, ranges 1.10-1.18 mm ; the maximum HW of a large number of individuals of diminuta measured from all over its range is 1.26 mm. The HW of the tw^o available series of nitens varies 1.23-1.26 mm, while the HW of the putative Bornean laeviceps varies 1.02-1.08 mm. (2) The body sculpturing is finer and less extensive. Dorsal cephalic sculpturing consists almost entirely of longitudinal striae, and these are limited to the lateral portions of the elypeus, to the area bordei'ed by the eyes, antennal insertion, and man- dibular bases, to a restricted area just ventral to the eye, and to the gula. In the gular area the striae are relatively feebly developed and wavy. All of the remainder of the head is com- pletely smooth and shining. Sculpturing is distributed on the 1 Two series from northern Borneo and Sarawak in the Museum of Comparative Zoology have been tentatively determined as laeviceps. As noted under the dis- cussion of diminuta, these specimens evidently represent a distinct sibling species that occurs sympatrically with diminuta on Borneo. 132 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY alitrunk as follows : a thin area at the base of the pronotal ■'neck" is covered Vtv transverse striae ; the episternum is covered by arcuate striae running parallel to the long axis of the sclerite ; the metapleuron and lateral face of the propodeum below the level of the propodeal spiracle are covered by moderately wavy. hnigitudiually oriented striae ; and the posterior face of the jMopodeum below the level of the dorsal margin of the meta- pieural gland bulla is covered by transverse striae. All the rest of the doj-sal and lateral surfaces of the alitrunk, as well as the entire surfaces of the petiolar node and gastric tergites, are r<>iri[)letely smooth and shining. Holoiype worker. HW 1.29 mm, IIL 1.69 mm, SL 1.56 ram, CI 76, SI 121, PW 0.95 mm. Parafype worker. HW 1.29 nun, IIL 1.67 mm, SL 1.59 mm, CI 77, SI 123, PW 0.95 mm. Paratypc ergatoyyne. HW 1.18 mm, HL 1.56 mm, SL 1.48 mm. CI 7G, SI 125, PW 0.83 mm. This specimen is distinguished ex- ternally from the two associated w'orkers by its smaller head and alitrunk, thinner petiolar node, and much larger gaster. The ali- trunk is workerdike in form. Material examined. SOLOMONS : near Fourafi, in the moun- tainous interior of Malaita (W. M. Mann) ; 2 workers, 1 erga- togyne. Nidotypes are on deposit in the United States National Museum. Ecological note. According to Mann (1919), the types were taken running in file across a foot trail in montane forest. The presence of an ergatogyne in this series suggests that the colonj' may have been in the process of migration. Leptogenys papuana Emery Leptogenys papuana Emery, 1897, Ann. Mus. Civ. Stor. Nat. Genova, 38: 556, pi. 1, figs. 5, 6, worker. Type locality: "N. Guinea Mer. " Known from the holotype only. Leptogexys purpurea Emery, new status (Figure 6) Leptogenys Eitteli var. purpurea Emery, 1887, Ann. Mus. Civ. Stor. Nat. Genova, (2)0: 433, worker. Type locality: .\ndai, Neth. New Guinea. (Holotype examined — Emery Coll.). WILSON : ANTS OF MELANESIA I, II 133 Material examined. NETH. NEW GUINEA: Andai, near Manokwari (holotype) ; Maffin Bay (E. S. Ross). N.-E. NEW GUINEA : Gemeheng, Mongi Watershed, 1300 m. (Wilson, no. 781) ; Tumnang, Mongi Watershed, 1500 m. (Wilson, no. 803) ; Ebabaang, Mongi Watershed, 1300-1400 m. (Wilson, no. 828) ; Wamuki, Mongi Watershed, 800 m. (Wilson, no. 849) ; Finsch Harbor (N. G. L. Wagner). In eastern New Guinea, at least, this species is most abundant at intermediate elevations in the Fig. 6. Worker of Leptogenys purpurea Emery from Gemeheng, N.-E. New Guinea. mountains. It was never encountered by the author during intensive collecting in the lowlands around Lae and Gagidu. Taxonomic notes. Three species, L. kitteli Forel of the Asian mainland and Indonesia, L. violacea Donisthorpe of Waigeo, and L. purpurea Emerj-, together form what may be provisionally termed the kitteli superspecies. They are obviously closely inter- related, replace one another geographically, and yet seem too well marked to be considered as mere geographic variants of the same species. L. purpurea can be distinguished from L. kitteli on the basis of the following set of worker characters : ( 1 ) purpurescent reflections common over most of the body surface of purpurea, 134 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY as opposed to none in kitteli; (2) cephalic striae finer and denser in purpurea than in kitteli; (3) most of the lateral surfaces of the pronotum of purpurea smooth and shining, as opposed to this surface striate and subopaque in kitteli; (4) petiolar node some- what thicker in side view in purpurea. L. purpurea can be sep- arated from L. violacea by means of the characters given in couplet 13 of the key. Ecological notes. In the mountains of the Huon Peninsula, L. purpurea inhabits both clearings and the interior of undisturbed forests. Its behavior has already been described in some detail in another paper (cited on p. 123). Like the related species L. diminuta, it has marked legionary habits, employing group- foraging to prey on large arthropods not ordinarily vulnerable to ants that forage solitarily. Leptogenys triloba Emery (Figure 5) Leptogenys triloba Emery, 1902, Termeszetr. Fiiz., 25 : 159, worker. Type locality: Sattelberg, N.-E. New Guinea. Material examined. N.-E. NEW GUINEA: Tumnang, Mongi Watershed, 1500-1600 m. (Wilson, no. 811) ; 1.5 km. north of Bandong, upper Bunbok Valley, 1300 m. (Wilson, no. 1131). Taxonomic note^i. The Bandong worker agrees with Emery's description in having the gaster reddish and contrasting with the remainder of the body, which is jet black. The Tumnang worker differs, in that its gaster is as darkly colored as the remainder of the body. Both specimens agree well with the remainder of Emery's description. 1 have tentatively determined as this species a single male collected at light during the April, 1955, trip in the mountainous portion of the Huon Peninsula. Ecological notes. Near Tumnang, in undisturbed midmountain forest, a Avorker was found at about 9 a.m. running along the top of a large rotting log, with a dead oniscoid isopod in its mandi- bles. Near Bandong another lone worker was found during the morning foraging on a native foot trail in midmountain forest. Leptogents truncata Mann Leptogenys (Leptogenys) truncatus Mann, 1919, Bull. Mus. Comp. ZooL, WILSON : ANTS OF MELANESIA I, IT 135 63: 296, fig. 10 (nee fig. 9, which is labelled as this species but is actually L. foreli), worker. Type locality: Graciosa Bay, Santa Cruz I. Known from the holotj^pe worker only. Leptogenys violacea Donisthorpe Leptogenys (Lobopelta) violacea Donisthorpe, 1942, Ami. Mag. Nat. Hist., (11)9: 705, worker. Type locality: Camp Nok, Waigeo, 800 m. (Syn- type examined — MCZ ) . Known from type material only. THE LEPTOGENYINI OF THE FIJI ISLANDS Nothing has been published on the Fijian Leptogenys since Mann's pioneering- monograph on the Fijian ant fauna (W. M. Mann, 1921, Bull. Mus. Comp. Zool., 64: 401-499). In this publi- cation all six of the currently known species were described for the fir.st time: foveopunctata, fugax, humiliata, letilae, navuu. and vitiensis. A recent re-examination of the types of all of these .species, with the exception of L. foveopunctata, has led me to the tentative conclusion that all are closely interrelated and even possibh- derived from the same parental species. They are mem- bers of the L. chlncnsis Mayr group, which is represented by a large number of species in the Indo-Australian region, and their closest Melanesian relatives within the chiyieyisis group are hitiiherculata Emery of New Guinea, hehrideana "Wilson of the New Hebrides, and sagaris Wilson of New Caledonia. On the basis of morphological evidence it appears likely that hitiihercu- lata, hehrideana, and most or all of the Fijian species were de- rived from a single ancestral stock, while sagaris has been derived independently from near the related, possibly cognate Australian species anitae Forel. Bitiiherculata, hehrideana, sagaris, navua, and humiliata have undergone little change, while foveopunctata, fugax, letilae, and vitiensis have diverged along several inde- pendent morphoclines. The reason why this species complex has radiated so extensively on the proportionately tiny land mass of the Fiji Islands, while remaining restricted and conservative elsewhere, is perhaps that in the Fijis it faced relatively little competition from the depauperate endemic ponerine-myrmicine fauna and hence had more evolutionary "opportunity" than elsewhere. 136 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY THE LEPTOGENYINI OF NEW CALEDONIA The four known New Caledonian species represent at least three, and possiblj^ four, distinct species groups. Three of the species have obvious affinities with the fauna of tropical Queens- land, while the fourth (Leptogenys puni'tata) is of uncertain position. Key to the species, hosed on the worker caste 1. Large species, head width exceeding 1.5 mm ; mandibles exceptionally long, their length measured from insertion to tip exceeding the head widtli Prionogenys ron^i Emery Smaller species, head width never exceeding 1.2 mm ; mandibles shorter, their length much less than the head width 2 2. Head entirely covered by coarse punctures, subopaque to opaque Leptogenys punctata Emery Head entirely smooth and shining 3 ?>. Larger species, head width not less than 0.90 mm ; posterior corners of petiolar node seen from directly above forming distinct angles of 45-50° Leptogenys acutangula Emery Smaller species, head width not exceeding 0.75 mm ; posterior corners of petiolar node seen from directly above thick and broadly' rounded Leptogenys sagaris Wilson Leptogenys acutangula Emery Leptogenys {Lobopelta) acutangula Emery, 1914, Nova Caledonia, 1: 398- 399, pi. 13, fig. 2a, worker. Original localities: Valine de la Ngoi, 200 m. ; Yatc. (Syntype examined — Emery Coll.) Leptogenys (Lobopelta) acutangula var. brevinoda Emery, ibid., p. 399, pi. 13, fig. 2b, ergatogyne. Type locality: Valine de la Ngoi. NEW SYNONYMY. Material examined. NEW CALEDONIA: Ciu, 300 m. (Wil- .son, ace. nos. 196, 197, 198, 199, 227, 230, 237, 282). Taxonomic notes. As Emery suggested in his original descrip- tion of var. hrevinoda, this form is nothing more than the erga- togyne of acutangula. Individuals were taken by the present author on three occasions in association with typical acutangula workers (ace. nos. 198, 199, 282). L. acutangula is clearly a member of the Australian conigera group. Of all the Tndo-Australian species examined during the WILSON: ANTS OF MELANESIA 1, II K"! present study, it is most closely approached by an undescribed species of the conigera group represented by a single series in the Museum of Comparative Zoology from Mt. Carbine, Queensland (Jr*. J. Darlington). The two species share, among other charac- ters, the angulate posterior petiolar corners in the worker caste. Ecological notes. This species was one of the most abundant ants encountered in the isolated woodlot on the Fere property at Ciu. Colonies were found occupying well formed chambers and galleries in the soil beneath rocks. Three colonies removed entire and examined consisted in each case of a single ergatogyne and approximately fifty workers. Callow males were found in one nest, on December 21, 1954. In the less disturbed forest near the Ciu Falls only a single colony was encountered ; this was nesting in a small rotting log buried in thick leaf litter. Leptogenys punctata Emery Leptogenys {Lobopdta) punctata Emery, 1914, Nova Caledonia, 1 : 398, worker. Type locality: Cone, New Caledonia. (Syntype examined — Emery Coll.) Material examined. NEW CALEDONIA: Cone (syntype); Ciu, 300 m. (Wilson, ace. nos. 228, 262). Taxonomic notes. The Ciu specimens have distinctly lighter body sculpturing than the syntype but are otherwise nearly identical to this specimen. L. punctata is not closely related to any other Australian or Melanesian species known to me. Its worker caste is very similar to that of L. punctiventris Mayr, a widespread Indomalayan species, but the males of the two species are quite different, particularly in genitalic structure. Ecological notes. One colony was collected in the Fere wood- lot and one in the forest east of the Canala River. Each w-as nesting in the soil beneath a rock and contained probably less than a hundred workers. Males were found in the Canala River nest, on December 31, 1954. Leptogenys sagaris Wilson, n. sp. (Figure 5) Diagnosis {worker). Closely resembling L. anitae Forel of Queensland, differing principalh' in the following characters : 138 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY (1) Distinctly smaller in size. Head width of sagaris types 0.64-0.69 mm; head width in two nest series of anitae examined not less than 0.82 mm. (2) Eyes proportionately smaller in size. In sagaris there are only five to six ommatidia along the maximum length of the eye, while in anitae there are approximately twelve to sixteen ommatidia along the same line of measurement. (3) Petiolar node proportionately shorter in sagaris. The above diagnosis will also serve to distinguish L. sagaris from its nearest Melanesian relative, L. hituhcrculata Emery. The latter species is further distinguished from both sagaris and anitae by the following characters : body pilosity proportionately longer ; head tapering more posteriorly when viewed in full face ; dorsal border of petiolar node viewed from the side descending more abruptly in an anterior direction. Holotijpc worker. HW 0.67 mm, HL 0.97 mm, SL 0.83 mm. CI 69, SI 124, EL 0.12 mm, PW 0.53 mm, petiolar node height 0.69 mm, petiolar node length 0.56 mm, dorsal petiole width 0.35 mm. Paratype worker variation. HW 0.64-0.69 mm (encompassed by a single nest series, ace. no. 240). The paratype series shows very little variation in external morphology. Ergatogijne. HW 0.64 mm, HL 0.89 mm, SL 0.73 mm, CI 72, SI 114, EL 0.12 mm, PW 0.48 mm, petiolar node height 0.41 mm, petiolar node length 0.38 mm, dorsal petiole width 0.40 mm. The single individual examined (ace. no. 272) is very worker- like. The head is very similar to that of a small worker, lacking t)celli and showing no enlargement of the compound eyes. The alitrunk is smaller proportionate to the head than in the worker. A distinct metanotal groove is present which is lacking in the worker. The petiolar node is much shorter and broader than in the worker ; seen from the side it approximates an isosceles tri- cingle in outline, with a broadh^ rounded dorsal border. The gaster is large, its volume exceeding by 1.5 X that of a large worker. Material examined. NEW CALEDONIA : Ciu, 300 m. ; holo- type nest series, AVilson ace. no. 272, one ergatogyne and 9 workers ; ace. no. 230, one worker ; ace. no. 240. 9 workers, (hapeau Gendarme, one worker. WILSON: ANTS OF MELANESIA I, n 139 Ecological notes. My accession no. 240 consisted of a colony found nesting under a rock on the floor of the Fere woodlot. It contained an estimated 50-75 adult workers. The holotype nest series (no. 272) was taken from a colony nesting in a small rot- ting log buried in leaf litter in the Canala River forest. It con- tained an ergatogyne and an undetermined number of workers. As in the related species L. hituherciilata and L. hehrideana, the workers of sagaris are relatively timid and fast, and scatter (luickly when the nest is opened. Prionogenys rouxi Emery I'riotioffenys rouxi Emery, 1914, Nova Caledonia, 1: 399-400, pi. \?<, fig. 3, worker. Type locality: Mt. Canala, 700 m.. New Caledonia. Material examined. NEW CALEDONIA : Fere woodlot, Ciu, .300 m. (Wilson, ace. nos. 189, 303, 304). Taxonomic notes. This species appears to be distinguished from the species of Leptogeni/s solely by its aberrant head form. In particular, the mandibles are extremely elongate, their length measured from insertion to tip nearly as great as the head length (standard measure) ; the masticatory borders are lined with distinctive stiff sensory hairs that project out at 60° and criss-cross when the mandibles are closed ; and the eyes are placed far forward on the head, their anterior margins being less than maximum-eye-length distance from the mandibular insertions. As far as I have been able to ascertain, P. rouxi is not marked by any other characters, in external morphology, that might be construed to be of generic magnitude. In fact, it bears a particu- lar resemblance in body structure to Leptogcnys acutangula and other members of the L. conigera group. The ergatogyne, col- lected for the first time during the present study, differs from the worker caste in the same manner already indicated by Emery for L. acutangula, i.e., ocelli lacking, head size about that of a small worker, alitrunk proportionately smaller relative to head size than in the worker, gaster approaching twice the volume of that of connidal workers, metanotal groove better developed, the petiolar node much shorter and broader. The male of P. rouxi, also collected for the first time in 1954- 55, shows no truly remarkable characteristics. Compared with Lepfogenys acutangula, only two differences seem noteworthy : in 140 BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY /'. rouxi, the notaulices are better developed, and the subgenital plate is much shorter and lacks the paired posterior marginal hooks that are present in L. acutangula. Both of these structures, however, are extremely variable within Leptogenys and do not at the present time seem to offer much opportunity for generic divisions. The precise relationships of P. rouxi are in need of further study. As noted already, the species bears a distinct resemblance to L. acutangula, appearing closer to this species than to any other known Melanesian species, and indeed may have been derived phylogenetically from acutangula, a common ancestral species, or a related Australian member of the conigera group. Account must also be taken of the relationship of P. rouxi to the generitype P. podenzanai Emery of Queensland. The two species resemble each other, at least superficially, in several im- portant features, including the peculiarly elongate mandibles, forwai-d position of the eyes, and conigera-type structure of the petiolar node. But podenzanai differs strikingly from rouxi in the possession of well developed mandibular teeth, which are completely lacking in rouxi. There is an excellent possibility that further study will show that the Prionogenys diagnostic characters were evolved independently in the two species, in which case Prio7iogenys will have to be placed in the synonymj' of Leptogenys, or else a separate monotypic genus erected to receive rouxi. Ecological notes. All three of the colonies of P. rouxi found at Ciu were in the Fere woodlot. Each was nesting under a flat rock embedded in the soil. The nest of colony no. 189, discovered on December 21, 1954, consisted principally of a flat, irregular L-hamber about 10 centimeters in maximum diameter; a lateral gallery led for a short distance to the base of a small stump adjacent to the covering rock, and then descended vertically to a depth of approximately 12 centimeters. The colony contained an estimated 55 Avorkers, 25 worker cocoons, 20 larvae at various stages of development, and an undetermined number of eggs. The ergatogyne, if present, w^as not recovered. Colony no. 303, collected on January 3, 1955, was lodged in a single gallery, approximately 2.5 cm. in diameter and 10 cm. in length, that led from beneath a rock into the soil at one side. WILSON: ANTS OF MELANESIA I, IT 141 It contained a single ergatogyne and an estimated 30 workers, 6 callow males, 20 cocoons of undetermined caste, and an unde- termined number of larvae at various stages of development. Colony no. 304, also collected on January 3, occupied a nest verj- similar to that of no. 303. It contained an ergatogyne and an estimated 40 workers, 6 callow males, and an undetermined quantity of brood. Prionogcnys rouxi is a very shy, timid species, and the colonies scatter swiftly when their nest is exposed. When captured and handled, the workers open their mandibles to a 90° angle in a threatening posture and are capable of inflicting a painful sting. The central nest chamber of colony no. 189 was partially filled with the chitinous remains of isopods and earwigs. Both types of arthropods are a prominent part of the cryptofauna at Ciu and are assumed to serve as the chief, if not exclusive, prey of the Prionogenys. It is probable that the workers are primarily noc- turnal in habit, since they were never found foraging above irround in the vicinity of the nests during the day. II THE TRIBES AMBLYOPONINI AND PLATYTHYREINI This section deals with five genera, Amblyopone, Myopopone, Prionopelta, Mystrium, and Platythyrea, represented by a small number of species mostly limited within Melanesia to New Guinea, the Bismarck Archipelago, and Solomon Islands. No member of these genera has yet been recorded from the Fiji Islands, while New Caledonia is known to harbor only two en- demic species: Amhlyopone australis Erichson, which is wide- spread in other parts of Melanesia as well as in Australia and Tasmania, and Prionopelta hrocha n. sp., a remarkable relict form with primitive features reminiscent of Amhlyopone. Western Melanesia contains a zoogeographic mixture, including old en- demics (Amhlyopone celata, Prionopelta majuscnla,), probable Indo-Malayan immigrants {Myopopone castanea, Platythyrea spp.), and a single probable Australian immigrant {Amhlyo- pone australis). Certain species, e.g. Myopopone castanea and Platythyrea parallela, are widely distributed over the islands and are among the dominant ants in the lowland rain forests there. 142 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Amblyopone Erichson Key to the species, based on the worker caste Pontal carinae separated by a wide space; larger species, head width never less than 1.32 mm (widespread in Melanesia) australis Erichson Frontal carinae contiguous; smaller species, head widtli not exceeding 0.64 mm (Solomons) celata Mann Amblyopone australis Erichson Amblyopone australis Erichson, 1841, Arch. Naturgesch., 8: 261, pi. 5, fig. 7, woi'ker. Type locality: Woolnorth, Tasmania. Amblyopone laevidens Emery, 1887, Ann. Mus. Civ. Stor. Nat. Geneva, 25: 447 worker. Type locality: Hatam, Arfak Mts., Neth. New Guinea. NEW SYNONYMY. Amblyopone levidcns, Mann, 1919, Bull. Mus. Comp. Zool. 63: 281. Amblyopone australis subsp. levidetis, Wheeler, 1927, Proc. Amer. Acad. Arts Sci., 62 : 12. Amblyopone nana Emery, 1914, Nova Caledonia, 1: 394-395, worker, queen. Original localities: Mt. Panic, 500 m., and Mt. Canala, 700 m.. New Caledonia. NEW SYNONYMY^ Amblyopone australis subsp. nana, Wheeler, 1927, op. cit., 16. Material examined. N.-E. NEW GUINEA: Joangeng, 1500 m. (E. 0. Wilson, no. 752) ; Ebabaang, 1400 m. (Wilson, no. 819). NEW HEBRIDES: Tanna (L. E. Cheesraan). Mann (1919) records this species from near Fourafi, in the monntainons in- terior of Malaita, Solomon Islands. Outside western and central Melanesia, A. australis is widespread in Australia and occurs on Tasmania, New Caledonia, Norfolk I., Lord Howe I., and New Zealand. According to W. L. Brown (pers. commun.), the New Zealand population was proljably introduced by man from Australia and is currently limited to the vicinity of cities and towns on North Island. Taxonomic notes. In the present review I have followed Wheeler (1927) in considering laevidens and 7iana as no more than geographic forms of the widespread and highlj^ variable australis. The New Guinea workers examined (laevidens) are relatively large, darkly colored, and with smooth dorsal mandibu- lar surfaces. In these characters they most closely resemble ma- terial from Queensland and New Zealand. In contrast, the Tanna, New Hebrides, workers (nana) are small, light colored, and with beavily striate dor.sal mandibular surfaces. They most WILSON: ANTS OF MELANESIA I, II 143 closely resemble specimens from New Caledonia (fide Emery's ilescription of the iiann types) and southeastern Australia. Ecological notes. At Ebabaang workers were found scattered under the bark of a large "Zoraptera-stage" rotting log in second-growth midmountain rain forest. At Joangeng a lone dealate queen was found under the bark of a log in a forest clearing. It is noteworthy that the western Melanesian collec- tions, including that of the laevidens type and Mann's Solomons collection, were all made at higher elevations in cool forest zones. If this is indicative of its true distribution, A. australis makes only limited contact with its closest ecological equivalent in this area, Myopopone castanea, a predominantly lowland species. Amblyopone celata (Mann) Sfifpnatomma (Fulakora) celata Mann, 1919, Bull. Mus. Conip. Zool., 63: 279-281, fig. 2, worker. Type locality: Fulakora, Santa Isabel, Solo- mons. (Syntypes examined — MCZ.) Known from type material only. In his original description Mann records this species from additional Solomons localities: Auki, Malaita ; Tulagi, Florida ; Wai-ai, San Cristoval. He found it nesting in small colonies under rocks on the floor of lowland rain forests. Myopopone Roger Key to the species, based on the ivorker caste Dorsal surface of petiole bearing only scattered piligerous foveae, the extensive interspaces completely smooth and shining (Moluccas to Solomons) castanea (Fr. Smith) Dorsal surface of petiole longitudinally rugulose and subopaque (Moluccas only) beccarii Emery Myopopone beccarh Emery Myopopone Beccarii Emery, 1887, Ann. Mus. Civ. Stor. Nat. Geneva, (2)5: 447-448, worker. Type locality: Ternate. Known from type material only. According to Emerj^ this species differs from M. castanea principally in the following characters : mesonotum and petiole longitudinally rugulose, ce- phalic ''striation" more extensive, and propodeal and dorsal petiolar foveae elongated into weak sulci. 144 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Myopopone castanea (Fr. Smith) Amblyopone castaneus Fr. Smith, 1860, J. Linn. Soc. Zool., 5 (suppl.) : 105, pi. 1, fig. 6, worker. Type locality: Batjan. (Holotype examined — Oxford University Museum). Myopopone rufula Roger, 1861, Berl. Ent. Z., 5 : 52, worker. Tj-pe locality : Batjan. Myopopone castanea, Forel, 1901, Mitt. Zool. Mus. Berl., 2 : 5. Myopopone castanea, Mann, 1919, Bull. Mus. Comp. Zool. 63: 281, distribu- tion. Myopopone moelleri, Santschi, 1932, Mem. Mus. Nat. Hist. Belg., 4: 11. Nee M. moelleri Bingham. Myopopone picea Donisthorpe, 1938, Ann. Mag. Nat. Hist., (11)2: 493, worker. Type locality: Mt. Dulit, Sarawak. (Holotype examined — BMNH). NEW SYNONYMY. Myopopone woUastoni Donisthorpe, 1942, Ent. Mon. Mag., 78: 29, queen. Type locality: Mimika River, Neth. New Guinea. (Holotype examined — BMNH). NEW SYNONYMY. Myopopone smithi Donisthorpe, 1946, Ann. Mag. Nat. Hist., (11)13: 577, queen. Type locality: Nadzab, Markham Valley, N.-E. New Guinea. NEW SYNONYMY (provisional). Myopopone rossi Donisthorpe, 1947, Ann. Mag. Nat. Hist., (11)14: 297-299, worker. Type locality: Finschhafen, N.-E. New Guinea. (Holotype examined — CAS). NEW SYNONYMY. Myopopone siinilis Donisthorpe, 1949, Ann. Mag. Nat. Hist., (12)1: 488, queen. Type locality: Maffin Bay, Neth. New Guinea. (Holotype exam- ined—CAS). NEW SYNONYiMY. Material examined. MOLUCCAS: Batjan {castanea holo- type). NETH. NEW GUINEA: Maffin Bay {similis holotype). N.-E. NEW GUINEA: Finschhafen (rossi holotype) ; Tor River. PAPUA: Karema, Brown River (Wilson, no. 566) ; Bisianumu, 500 m. (Wilson, no. 652). SOLOMONS: Tenarii River, Guadal- canal (G. E. Bohart) ; Simba Mission, Bougainville (E. J. Ford, Jr.) ; Tulagi, Florida (W. M. Mann) ; Siota, Florida (Bohart) ; Fulakora, Santa Isabel (Mann) ; Lambeti, New Georgia (Mann) : Star Harbor, Wai-ai, San Cristoval (Mann). In addition to the material listed above, I have seen series from Hainan, Sumatra. Borneo, Philippines, and North Queensland. Taxonomic notes. This species shows considerable geographic variation in at least four independent characters, as indicated in the accompanying table. It is the author's present view that WILSON : ANTS OF MELANESIA I, 11 145 (GEOGRAPHIC VARIATION IN MYOPOPONE CASTANEA bocality Subpetiolar ' Process (Side View) Cephalic Sculpturing Sculpturing of Dorsal Surface of Petiolar Node Sculpturing of First Two Gastric Tergites Hainan Labuan I., .v. Borneo Sindanglaja. Java l::ingano I., near Sumatra N. Palawan 1 vii r. proaima Stitz syntype) Philippine Islands — vari- ous localities Bisianumu, Papua Finschhafen, X.-E. New Guinea Tor R., Neth. New Guinea X'arious Solomons localities Alcll wraith Range, N". Queensland about as long as broad; apex posteriorly acute twice as long as broad; apex rounded about as long I as broad; [ apex rounded about 1.5 X longer than broad ; apex rounded about 1.5 X longer than broad; apex rounded distinctly shorter than broad; apex rounded distinctly shorter than broad; apex rounded j distinctly shorter than broad; apex rounded distinctly shorter than broad; apex rounded distinctly shorter than broad; apex rounded feebly rugose shallowly furrowed and punctate moderately rugose moderately rugose shallowly furrowed and punctate shallowly furrowed and ijunctate moderately rugose moderately rugose moderately rugose moderately rugose moderately rug-ose sparsely punctate; post, central area non-striolate sparsely punctate; post, central area non-striolate moderately punctate; post, central area non-striolate moderately punctate; post, central area non-striolate moderately punctate; post, central area non-striolate moderately punctate; post, central area non-striolate sparsely foveate; post, central area non-striolate sparsely foveate; pest, central area striolate sparsely foveate; post, central area striolate sparsely foveate; post, central area striolate sparsely foveate; post- central area striolate finely punctate smooth and shining densely punctate densely punctate feebly shagreened first tergite feebly shagreened, second smooth smooth and shining feebly shagreened feebly shagreened feebly shagreened feebly shagreened 146 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY all of the Melanesia!! material examined, including the types of the several included Donisthorpe species, belongs to the single species M. castanea. The Philippine series,^ on the other hand, appear sufficiently divergent to rank as a separate species and may be considered so by future revisers of Myopopone. The queen of castanea differs strikingly from the worker in its much larger size, darker color, and more extensive and deeper cephalic sculpturing. This circumstance has led to taxonomic confusion in the past and was undoubtedly the principal source of inspira- tion for the numerous new names proposed by Donisthorpe. Ecological notes. Both the Karema and Bisianumu collections consisted of workers found under the thick bark of large rotting logs on the floor of rain forests. At Bisianumu workers were clustered with larvae around two large, freshly killed cerambycid larvae on the same log. Since the beetle larvae were well sepa- rated from one another, and appeared to be too large for the ants to transport through the preformed galleries under the bark, it is inferred that the ants had transferred their own larvae to feed on the prey after the latter had been attacked and killed. The Myopopone are singularly clumsy and sh}^ ants, and immediately commence searching for cover when exposed to light, abandoning their brood in the process. When handled, however, they arc capable of inflicting a painful sting. Prionopelta Mayr Key to the species, based on the worker caste 1. Larger species, head width of single known specimen 0.64 mm; genal teeth strongly developed, at least 0.03 mm in length (New Caledonia) brocha Wilson Smaller species, head width not exceeding 0.54 mm; genal teeth feebly developed, not exceeding 0.01 mm in length (New Guinea) 2 2. Cephalic and thoracic dorsa finely punctate and feebly shining; larger species, head width at least 0.48 mm majuscula Emerv Cephalic and thoracic dorsa densely and coarsely punctate and opaque; smaller species, head width not exceeding 0.42 mm opaca. Emery r 1 Including a syntype worker of M. castanea var. proxima Stits (MCZ) . WILSON : ANTS OF MELANESIA 147 I'RIOXOPEILTA BROCHA WilsOll, 11. s]) ■ Figure 7) Hiagvo^ns. Distinguished in the worker caste from all other known members of the genus 1)3' several apparently primitive characters, including exceptionally large size, strongly developed genal teeth, and Amhlyopone-Yike head shape. Two features of head shape are considered to be more typical of Avihlyopoiie than of Prionopelia — the head is broadest near its anterior end, and the lateral borders are relatively straight. Brocha is thus seen to be a species intermediate in position between the I'eniainder of Prionopelia and the presumably more primitive Arnhlyopone. It has been placed in Prionopelia primarily because of its mandible form, which is typical for that genus. Fig. 7. Prionopelia brocha n. sp., head of holotype worker. Drawing by Mrs. Nancy Buffler. Holotype worker. HW (exclusive of genal teeth) 0.64 mm, HL 0.79 mm, ML 0.16 mm, SL 0.45 mm. CT 81, SI 70. PW 0.43 148 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY mm, alitrunk length (maximum) 1.05 mm, dorsal petiole width 0.38 mm, petiolar node length (exclusive of ventral lobe) 0.42 mm, width of first gastric tergite 0.52 mm. Head as shown in Figure 7. Remainder of body similar to that of other Priono- pdta, including the relatively large maj^iscula Emery. Dorsal surfaces of mandibles finely and longitudinally striolate, and shining. Central portion of clypeas smooth and shining. Remainder of head covered by coarse, contiguous punctures, its surface for the most part completely opaque. Body posterior to the head sparsely punctate, its surface almost entirely feebly shining. Pilosity very similar to that of most other Prionopelta, in- I'luding majnscula. Abundant standing hairs present on most body and appendage surfaces, nearly or completely absent from posterior propodeal face, anterior and ventral surfaces of petiolar node, and most of extensor surfaces of legs ; everywhere grading into equally abundant, predominantly oblique underlying pu- bescence. Body uniformly light ferruginous, appendages clear to ver^ light reddish yellow. Material examined. NEW CALEDONIA: Mt. Mou, 180 m.. December 12, 1954, a single worker (E. 0. Wilson). This speci- men was collected by means of a Berlese funnel from leaf litter on the floor of dry, semi-deciduous native forest. Prionopelta majuscula Emery Piynera simillima Ft. Smith, 1860, J. Linn. Soc. Zool., 5 (suppL; : 105, worker. Neo Fonera simillima Fr. Smith, ibid., p. 104 (z= Ponera diminuta Fr. Smith). Prionopelta majuscula Emery, 1897, Termeszetr. Fiiz., 20: 595, worker, queen. Type locality: Beliao I., near Friedrich-Wilhelmshafen ( = Madang), N.-E. New Guinea. "! RJwpalopone simillima,, Emery, 1911, Gen. Ins., 118: 35. (Generic re-alloca- tion for Ponera simillima Fr. Smith, 1860, p. 105). Prionopelta poultoni Donisthorj)e, 1932, Ann. Mag. Nat. Hist., (10)10: 462, nom. pro Ponera simillima Fr. Smith, 1860, p. 105. Synonymy bv Brown, 1953, Breviora, no. 11: 12. WILSON : ANTS OF MELANESIA 1, II 149 Ilenea testacea Donisthorpe, 1947, Ann. Mag. Nat. Hist., (11)14: 183-186, fig., worker, queen. Type locality: MafBn Bay, Neth. New Guinea. Synonymy by Brown, loo. cit. (SjTitypes examined — CAS). Examblyopone churchilli Donisthorpe, 1949, Ann. Mag. Nat. Hist., (12)2: 401-402, queen. Type localtiy: Maflfin Bay, Neth. New Guinea. Syn- onymy by Brown, loc cit. (Holotype e.xamined — CAS). Taxonomic note. Through the courtesy of Dr. E. S. Ross, 1 have been able to re-exaraine the type series of Renea testacea Donisthorpe and Examblyopone churchilli Donisthorpe and to confirm the earlier opinion of Brown that these two forms are unqualified synonyms of P. majuscula. Prionopelta opaca Emery I'rionopeUa opaca Emery, 1897, Termeszetr. Fiiz., 20: 596, pi. 15, figs. 44- 45, worker, queen, male. Type locality: N.-E. New Guinea. Prionopelta Mocsdryi Forel, 1907, Ann. Mus. Nat. Hungar., 5: 1, worker. Type locality: Asuncion, Paraguay (Anisits leg.). NEW SYNONYMY (provisional; see below). Material examined. N.-E. NEW GUINEA: Nadzab (Wilson, no. 1089) ; Bubia (Wilson, no. 687) ; lower Busu River (Wilson, nos 899, 963, 978) ; Wamuki, 800 m. (Wilson, no. 846). PAPUA: Karema (Wilson, nos. 563, 573, 575) ; Bisianumu, 500 m. (Wil- son, no. 636). This species also occurs in ]\Iicronesia. Taxonomic notes. A single worker collected in the mountains near Wamuki is slightly larger than all of the other specimens examined, including those from nearby localities in the lowlands (head width 0.39 mm as opposed to maximum head width in other material measured of 0.3S mm). Dr. W. L. Brown has transmitted the following unpublished note concerning the status of Prionopelta mocsaryi: **A worker type of P. mocsaryi received from the Hungarian National Mu- seum can not be distinguished from P. opaca samples from New Guinea (Wilson, Biro leg.) in any character, after long and careful examination and comparison. Since the original Biro New Guinea collections were housed in the Hungarian National Museum, and since some of this New Guinea material is known to have been included in the series sent Forel for his 1907 study, it seems likely that the label ' Paraguay /Anisits ' 150 15ULLET1N : MUSEUM OF COMPARATIVE ZOOLOGY is misplaced. No specimen of Prionopelta anything like this one has been reported a second time from the Americas, al- though other species of the genus have all been collected repeat- edly in tropical America. My conclusion is that P. inocsaryi sliould be added to the synonymy of P. opaca." Ecological notes. This species was found in a wide range of major habitats in New Guinea: dr}', open tropical evergreen forest at Nadzab, foothills rain forest at Bisianumu and Wamuki, and primary lowland rain forest at the Busu River. Stray work- ers were very common on the forest floor, and could easily be secured by tapping loose material from the bottoms of small pieces of rotting wood buried in leaf litter. They also turned up frequently in leaf -litter berlesates. A single colony found at Karema was nesting in a rotting section of tree branch on the forest floor. It contained between 15 and 20 workers and a small (juantity of brood. Mystrium Roger Mystrium camillae Emery Mystrium Camillae Emery, 1889, Ann. Mus. Civ. Stor. Nat. Genova, 27 : 491, pi. 10, figs. 1-3, worker, queen. Type locality: Bhamo, Burma. Wheeler and Chapmaxi, 1925, Philippine J. Sci., 28: 55, dist. Karawa- .iew, 1925, Konowia, 4: 73, dist. Brown, 1952, Psyche, 59: 25, dist. A single male, collected by E. S. Ross at Maffin Bay, Neth. New Guinea, in July, 1944, has been tentatively determined as this species. Although this is the first time Mystrium has been recorded from Melanesia, its occurrence there is not surprising, since M. camillae was already known from the Philippines (, Wheeler and Chapman, 1925)^ Java (Karawajew, 1925), and Northern Territory, Australia (Brown, 1952). Platythyrea Roger Seen from above, the posterior border of the petiolar node is deeply concave, and its posterior corners are drawn out into long, flattened, blunt processes quadruienta Donisthorpe Seen from above, the posterior border of the petiolar node is sinuate, while its posterior corners arc not drawn out into processes parallela (Fr. Smith "> WILSON: ANTS OF MELANESIA I, 11 151 Platythrea pakallela (Fr. Smith) Ponera parallela Fr. Smith, 1859, J. Linn. Soc. Zool., 3 : 143, worker. Type locality: Aru. Donisthorpe, 1932, Ann. Mag. Nat. Hist., (10)10: 454, worker. Ibid., 1943, (11)10: 434-435, male, doubtfully associated. (Holotype examined — Oxford University Museum). Pachycundyla melancholica Fr. Smith, 1865, J. Linn. Soc. Zool., S: 71, worker. Type locality : Morotai. NEW SYNONYMY (provisional). Platythyrea pitsilla Emery, 1893, Rev. Suisse Zool., 1 : 188-189, worker. Type locality: Amboina. (Holotype examined — Emery Coll.). NEW SYNONYMY. Platythyrea coxalis Emery, 1893, ihid., p. 189, nota, worker. Type locality: Perak, Malaya. (Holotype examined — Emery Coll.). NEW SYN ONY'MY. Platythyrea melancholica var. aruana Karawajew, 1925, Konowia, 4: 75, worker. Type locality: Wammar I., Aru Archipelago. NEW SYN- ONYMY (provisional). Material examined. MOLUCCAS: Kalara I., Ilalmahera (C S. Banks) ; Amboina {pusilla holotype). NETII. NEW GUINEA: Maffin Bay (E. S. Ross). N.-E. NEAV GUINEA: Nadzab (Wil- son, nos. 1086, 1090, 1097) ; Lae (N. L. H. Krauss) ; Bnbia (Wilson, no. 683); Finschhafen (Ross). PAPUA: Bisianumn. 500 m. (Wilson, nos. 617, 644, 655, 659, 660). ARU: (holotype). Taxonomic notes. This species shows non-geographic variation in several characters, notably total size, proportions of the peti- ular node, form and placement of the petiolar teeth, sculpturing, and color, which is of similar magnitude to the differences sepa- rating many related species in other ponerine groups. Neverthe- less, in an analysis of the sizable collections of material in the Museum of Comparative Zoology and Museo Civico di Storia Naturale, Genoa, I have been unable to detect any constant dif- ferences that might be construed as partitions along species lines. Variation in tv;o of the characters, total size and relative thickness of the petiolar node, is indicated in the plot of petiolar length times width given in Figure 8. It can be seen that these data are distributed along a single regression zone. The other characters examined show a similar pattern of variation. The form melancholica is placed in provisional synonymy here liecause of its inadequate definition in previous literature. The holotype is probably lost, since neither Donisthorpe (1932) nor the present author (in 1955) were able to find it among the 152 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY extant Frederick Smith types. The chances are very remote that this specimen represented anything more than a variant of P. parallela. Ecological notes. All of the author's collections consisted of stray workers found in leaf litter and rotting logs. At Nadzab a worker was found carrying a small moth larva in its mandibles. 0.80 z UJ Q o < 0.70 - 0.60 - 0.40 0.50 PETIOLAR NODE WIDTH 0.60 Fig. 8. Variation in petiolar node proportions in Indo-Papuan material of Platythyrea parallela. No more than two workers were measured from each nest series. Platythyrea quadridenta Donisthorpe Platythyrea quadridenta Donisthorpe, 1941, Ann. Mag. Nat. Hist., (11)7: 134, worker. Type locality: Wharton Range, Kokoda, Papua. (Syn- type examined — BMNH). Material examined. PAPUA : Wharton Range (syntype) ; WILSON: ANTS OP MELANESIA I, II 153 Karema, Brown River (Wilson, nos. 550, 570) ; Bisianumu, 500 ra. (Wilson, no. 642). Taxonomic note. P. quadridenta is closely related to the Indo- Malayan species P. sagei Forel, but can be easily separated by its smaller size, thicker petiolar node, and unique possession of abundant, short, erect hairs over most of the dorsal surface of the body. Ecological note. At Karema a colony consisting of approxi- mately 50 workers and an undetermined quantity of brood was found nesting under the bark of a large, "Zoraptera-stage" log on the forest floor. Bulletin of the Museum of Comparative Zoology AT HARVARD COLLEGE Vol. 118, No. 4 THE FOSSIL CARNIVORE AMPHICYON INTERIMEDIUS FROM THE THOMAS FARM MIOCENE, PART I: SKULL AND DENTITION By Stanley J. Olsen Florida Geological Survey CAMBRIDGE, MASS., U.S.A. PRINTED FOR THE MUSEUM May, 1958 Publications Issued by or in Connection WITH THE MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD C^OLLEGE Bulletin (octavo) 1863 — The current volume is Vol. 118. Breviora (octavo) 1952 — No. 85 is current. Memoirs (quarto) 1864-1938 — Publication was terminated with Vol. 55. JoHNSONiA (quarto) 1941 — A publication of the Department of Mollusks. Yol. 3, no. 35 is current. Occasional Papers op the Department op Mollusks (octavo) 1945 — Vol. 2, no. 21 is current. Proceedings op the New England Zoological Club (octavo) 1899-1948 — Published in connection with the Museum. Publication terminated with Vol. 24. The continuing- publications are issued at irregular intervals in num- bers which may be purchased se]iarately. Prices and lists may be obtained on application to the Director of the Museum of Comparative Zoology, Cambridge 38, Massachusetts. Of the Peters ' ' Check List of Birds of the World, ' ' volumes 1-3 are out of print ; volumes 4 and 6 may be obtained from the Harvard Uni- versity Press ; volumes 5 and 7 are sold by the Museum, and future volumes will be pulilished under Museum auspices. Bulletin of the Museum of Comparative Zoology AT HARVARD COLLEGE Vol. us, No. 4 THE FOSSIL CARNIVORE AMPHICYON INTERMEDIUS FROM THE THOMAS FARM MIOCENE, PART I: SKULL AND DENTITION By Stanley J. Olsen Florida Geological Survey CAMBRIDGE, MASS., U.S.A. PRINTED FOR THE MUSEUM May, 1958 No. 4 — The Fossil Carnivore Amphicyon intermedius from the Thomas Farm Miocene, Part I: Skull and Dentition^ By Stanley J. Olsen INTRODUCTION The lower Miocene beds of the Thomas Farm quarry, located in north central Florida, have in recent years yielded numerous \-ertebrates that are complete enough to fill in many of the gaps in our knowledge of species originally described from fragmentary material. Among the rarer remains that fall in this category are those of the large carnivore Amphicyon, which was first recorded from the Gilchrist County deposit by Simpson (1932), who briefly described some specifically undetermined fragments. White (1940) described Amphicyon intermedius on a left mandible with incom- plete dentition, and later (1942) described a second species, A. longiramus, based on a right mandible with Po - M2. In the time that has elapsed since these species were described, much new evidence has come to light. The present restudy was undertaken to determine if the characters given as diagnostic would hold up when subjected to analysis with a larger sample of specimens for comparison. Variation in Canis lupus and Ursus americanus, which have to some degree a similar form and size in the skull and mandible, has also been taken into consideration. All material used by White in his type descriptions has been examined, together with all specimens since found. Casts of the types of the majority of described North American species of Amphicyon have been available for comparison. A discussion of the broader affinities of Amphicyon will be given in the paper describing the postcranial skeleton. CANIDAE AMPHICYONINAE AMPHICYON Lartet Amphicyon intermedius White Figures 1 to 5 Amphicyon sp. Simpson, 1932, pp. 20, 21, fig. 5. Amphicyon intermedius White, 1940, p. 32, pis. 3, 4. Amphicyon longiramus White, 1942, pp. 6, 7, pi. 5. Horizon and locality. Lower Miocene, Thomas Farm, Gilchrist County, Florida. 'The postcranial skeleton will be treated in a subsequent contribution. 158 bulletin: museum of comparative zoology Material examined. M.C.Z. 3630, right mandible with P3 - Mj, alveoh for M2-3; 3631, left mandible with P3-M2, alveoli for C, Pi-2, M3 (type) ; 3632, left mandible with dm4, and detached molars, alveoli for P1-3, (original hypodigm); 3919, right mandible with P2 - M2, alveoli for Ii - Pi, M3 (type of A. longi ramus); 4060, palate with left and right P^-M-; 4424, partial braincase and sagittal crest; 5832, right Mi-M.; 5833, left M\ M-; 7140, re- stored skull; 7141, 16 miscellaneous teeth; 7142, left dm4; 7143, left yi\ F.G.S. V-5257, right P4; V-5258, trigonid of right M.; V-5648, right Mi; V-5649, right M.; V-5650, lower canine; V-5659, lower canine; V-5663, right M2. U.F. 6501, nearly complete skull. White distinguished this species as follows: ". . . intermediate in size between A. frendens Matthew and A. superbus- Peterson; diastema between P2 and P3 the longest yet recorded, . . . cusps of the molars resembling those of A. sinapius Matthew, but dis- tinguished from that species by the smaller size and the greater length of the diastema between P2 and P3. It differs from A. pon- toni Simpson in that M2 is longer for its width, and that the basin between the hypocone and entocone^ is shallower." A. longiramus was characterized by: "Pi and M3 single-rooted, M3 with groove on outer side of root but not on the inner side, P4 with heel and ac- cessory cusp, two mental foramina . . . about one-fourth larger than the preceding, and differs from it also in the double-rooted P2 and in the single-rooted M3. In dental characters, in size, and in the relative proportions this form is difficult to distinguish from the jaws from the Snake Creek beds referred by Matthew to A. sinapius (Matthew, 1924) and perhaps does not merit specific designation." Synonymy. Simpson (1932) referred a partial right M2 (F.G.S. V-5258) and a right P4 (F.G.S. V-5257) to Amphicyon sp. These specimens are morphologically indistinguishable from A. inter- medins and must be referred to this species. The stated differences between the two species of Amphicyon described bj^ White are such that they can be attributed to indi- vidual variation and to age and do not appear to warrant taxo- nomic separation; examples of comparable dental variation are known in other carnivores. ^This specirs was made the type of Daphoenodon (Peterson, 1909). ^Hypoconid and entoconid in this paper. olsen: amphicyon intermedius . 159 A wide range of variation in premolars, especially as regards numbers of roots, is of rather common occurrence in carnivores. Colbert for Hemicyon teilhardi (1939), Merriam for Canis dims (1912), Scott (1890), and Matthew and Gidley (1904) for Aeluro- don haydeni, Peterson (1910, pp. 213, 214) for Da-phoenodon super- bus, Hall (1928) and Simpson (1949) for Ursus americanus, Kurten (1955) for Ursus spelaeus have all recorded variations of a similar nature to those used by W hite to establish A . longiramus, and have shown that they are individual in nature. Furthermore, on comparing P2 in the types of the two species, it is noticeable that P2 in the type of A. intermedius is not fully single-rooted. The dividing wall in the alveolus is incipient but distinct. In an immature specimen (M.C.Z. 3632) referred by White to A. intermedius, the alveolus for P2 is definitely for a double-rooted tooth. IVIeasurements of the Thomas Farm rami show that, proportionately, the types of A. intermedius and A. longiramus are almost exactly comparable in dimensions to M.C.Z. 46547, the longest (and the oldest) and M.C.Z. 46553, the shortest (and the youngest) of a series of Canis lupus rami from Alaska*. The morphological differences, such as tooth cusp arrangement and pattern between intermedius and longiramus, are within the range of variation found in comparable groups of Recent carnivores. I, therefore, believe that the species longiramus should be re- duced to the synonymy of A. intermedius. Amphicyon intermedius is now known also from the Garvin Gui- le}^ of Texas. The material will be described by Dr. J. A. Wilson. It should be pointed out that caution must be exercised in com- paring the type of A. longiramus (M.C.Z. 3919) with specimens of other members of this genus. It is an aberrant individual, at least in regard to the premolar region (Fig. lA, and White, 1942, p. 5), having a fully formed P4 that never erupted. I have figured this specimen (Fig. lA), inasmuch as it is the only complete lower jaw of the Thomas Farm Amphicyon known to date. All measure- ments, with the exception of those anterior to P2, are comparable to those of other specimens of the species. ♦Measurements taken by Dr. E. E. Williams and Prof. Bryan Patterson. 160 bulletin: museum of comparative zoology MORPHOLOGY Skull and mandible (Figs. 1, 2 and 3). Although the skull of Amyhicyon intermedius (Fig. 2) has been crushed and distorted to a certain extent, nevertheless the form is sufRciently well pre- served that its characters may be readily seen and compared, without recourse to paper reconstructions or other means of interpretation usually resorted to in interpreting buckled or mis- shapen material. The general form and proportions of the skull are more canid than ursid-like. One of the most prominent characters of the skull is the very high and strongly developed sagittal crest. This crest arises on the interparietal apex and terminates at a point which is midway along the median sagittal plane, and is much larger than the crest found in the Recent members of the Canidae and Ursidae or that which is present in the extinct genus Hemicyon, to which Ampkicyon has been compared (Colbert, 1939; Frick, 1926). The base of this attachment for the powerful temporal muscle is formed by two compact layers of bone, separated by cancellous tissue, which tend to fuse into a denser structure at the sagittal suture. The structure of this crest was disclosed by a break. The smaller occipital crest runs laterally downward on either side to meet the heavy, expanded posterior roots of the zygomatic arches. Since the described individual (U.F. 6501) is of advanced age, as evidenced by the worn condition of the dentition, it is difficult to determine the synostosed sutures or to distinguish them from the post-fossilization cracks, and thus nearly impossible to define the limits of the different bones of the skull roof and palate. The braincase is quite small, in proportion to the rest of the skull, for a carnivore of this size. This observation is confirmed by an- other skull fragment (M.C.Z. 4424) which has a portion of the parietal and sagittal crest preserved in an undistorted condition. The supraoccipital extends far back, to overhang the occipital condyles. The occiput is narrow and triangular as seen from the posterior view, having a central, vertical, occipital ridge much the same as Hemicyon teilhardi or the Recent members of Canis. olsen: amphicyon intermedius 161 Although this ridge is also found in members of the Ursidae, the same region in these animals has less overhang and the occipital area tends to be more square in shape when viewed posteriorly. Proportions of the frontals are much the same as those found in Canis dims or Canis lupus; these bones have the "dished" ap- pearance in the area of their union with the maxillaries and nasals — a condition not usually encountered in the bears. The palate is long and narrow, expanding sharply posterior to P^, much more so than in Ursus or Hemicyon, but to nearly the same degree when compared with that of Canis. The anterior palatine foramina have been destroyed but a remaining partial border of one of these openings locates them as being between the posterior margin of the canine alveolus and the anterior face of P-. The posterior palatine foramina are present opposite M^. The glenoid fossa of A. intermedius is shallower than that of Canis. There is a small postglenoid process, as in the bears and Hemicyon, rather than the definite recurved or hook-like process of the dog and wolf. The palatine-pterygoid area has been crushed and broken so that wevy little in the way of comparisons can be made with these elements. Auditory region (Fig. 2C). The auditory region as a whole is of an ursid rather than of a canid type. The bullae in A. intermedius are partially destroyed in the figured specimen, but this structure is known in A. sinapius (A.M.N.H. 18257), where it is flattened and flask-shaped and not highly inflated as in the bullae of the dog and wolf. As pointed out by Van der Klaauw, the name "bulla" is not very appropriate, in the literal sense, when applied to this region in the Ursidae (Van der Klaauw, 1931). This statement also holds true for the Amphicy oninae . The external meatus, as in Ursus americanus, is quite long and exhibits the form of a gutter rather than that of a cylinder or closed tube as found in the Canidae. The superior roof of the meatus is formed by the squamosal (Segall, 1943). The basioccipital in A. intermedius possesses wings or crests similar to those found on this element in the bears and Daphoenus but not in the dogs or wolves. l^^-Z bulletin: museum of comparative zoology By far the largest single element of the auditory region is the paroccipital process which is composed of heavy, thickened bone having a deflected plane nearly parallel to that of the palate and forming the posterior border of the foramen lacerum posterius and the stylomastoid foramen. In most cases, the courses of the various basicranial foramina cannot be followed to completion, due to the fragmentary nature of this area, and the fact that this entire region is supported by sand that has been artificially hardened in order to make the skull hold together, makes further preparation hazardous. The condylar foramen is situated in much the same position as that found in Ursus. Immediately anterior to this opening is the foramen lacerum posterius which is the largest external opening in the auditory region. Located on the anterior and anteroexternal margins of this large opening are, respectively, the carotid canal (in part) and the stylomastoid foramen. This latter foramen forms a groove along the inner face of the mastoid process. The eustachian opening and foramen lacerum medium are lo- cated at the junction of the postglenoid process with the squamosal, and are separated by a well defined wall into two surface openings instead of one as is present in Ursus americanus. The foramen ovale and the foramen rotundum are located to- gether, slightly anteromedial to the postglenoid process. Laterally and slightly anterior to these openings, there seems to be a part of a margin of what might be the alisphenoid canal but due to the damage in this particular area it is too doubtful to record as such with any degree of confidence. The auditory region in Amphicyon and other primitive carni- vores has been used as a clue to relationships for some time. How- ever, this area does not give us a necromantic insight into the relationships of these animals. Hough, in speaking of the ties between primitive carnivores, has this to say: "The structure of the auditory region leaves no doubt of the affi.nity of Daphoenodon and Amphicyon or the relationship of these two genera to Dapho- enus It seems probable, therefore, that Amphicyon and Daphoenodon represent different branches of the same subfamily rather than having a directly ancestral relationship and that neither olsen: amphicyon intermedius 163 was in the direct line of the bears" (Hough, 1948a, pp. 598-600). Speaking of the same animals in a subsequent contribution (1948b, pp. 108, 109), Hough placed Amyhicyon in the Ursidae and came to the following conclusions concerning this carnivore: "A dissec- tion of the auditory region shows this region to be ursid. . . . There is no basis in the auditory region for the association of Amphicyon and Daphoenus ... a close relationship with Daphoenus seems improbable." Hough sums up the significance of the auditory region as a means of classification by pointing out that this area does preserve an- cestral characters and that it would be a mistake to depend wholly upon it, or upon any one feature, in determining relationships. The lower jaw is canid-like with the ascending ramus describing a gentle curve from Ms to the apex of the coronoid process, as in the wolf and dog. In the bear and Hemicyon this same margin of the ascending ramus takes a vertical rise preceding the curve and ends in a decided, posteriorly deflected hook at the terminus of the coronoid process. A single masseteric fossa occupies the greater portion of the as- cending ramus in the Thomas Farm carnivore as compared to an additional premasseteric fossa in the mandible of Hemicyon. The dental foramina in carnivores in general are subject to con- siderable variation both as to number and location and the use of these openings as specific keys is not practicable. Dentiiion. The dental formula is I - 1, C - 1, P - 5, M - 1. The upper incisors are missing in the nearly complete skull but the alveolus for P indicates that this was a heavier and larger tooth than the other two. The canines show the wear of an old adult, that of the right side being worn to a level of the gum line in life and the other to one-half of its original length. As noted by Matthew (1924, p. 105) for Amphicyon, it has a double cutting edge, and is similar in form to the corresponding lower tooth (Fig. IC). Kurten (1955) has found that the relative proportions of the canines in Ursus spelaeus show a stronger dimorphism than that found in the cheek teeth. Whether or not this will hold for Amphicyon is still to be determined. P^ is missing on both sides but is a single-rooted tooth. P- is a simple, double-rooted structure 164 bulletin: museum of comparative zoology with a slightly expanded heel, having no accessory cusp present in the known examples. P^ has the same features as the preceding premolar with the exception of being larger and more expanded in the heel. P^ is of a definite canid pattern. The protocone ( = deuterocone) is at the antero-lingual end of the tooth instead of being more centrally located along the inner margin as it is in Hemicyon and the modern bears. M^ is canid-like, being similar in proportion to the corresponding tooth in Cynodesmus and other primitive dogs, rather than having the more square outline found in Hemicyon and Ursus americanus. The metaconule found on M^ and M- in A . intermedins is a variable character in other mem- bers of this genus. M^ is highly variable, both in size and form^. The examples found in the described skull differ but slightly from M^ in general proportions, although the size and outline of this tooth do not adhere to a typical shape (Fig. 5). Both M^ and M- are triple- rooted and exhibit considerable wear in the figured specimen (Figs. 2C, 4B). The heel in the upper molars of unworn specimens has fine crenulations along the cingulum, not unlike those found on the surface of these same teeth in the bears. The alveoli for the third molars are perfectly preserved in the skull and furnish us with more accurate information concerning the location of these teeth than has been heretofore known. M^ in A. intennedius, known only from a single specimen (M.C.Z. 7143), is a strongly developed single-rooted tooth, roughly oval in shape, having a single principal cusp, the paracone, and an internal cingulum of large proportions. The presence of M' has been the cause of much confusion in assigning specimens to the genus Am- phicyon. In some cases it was the only reason given for the generic assignment. Although M^ is typical of this genus, it is also pres- ent in several other canids, such as Daphoenodon superbus (origin- ally described as Aynphicyon superbus) and Daphoenus. The proportions of the area occupied by the lower premolars in relation to the total jaw length are nearer to those of Canis than to those of Ursus. There is a wide variable diastema between each of the premolars, with no crowding evident in any of the known specimens. The canine as mentioned earlier is double-edged with ^For similar \-ariation in Recent Canidae see plates in Young and Jackson (1951), and Young and Goldman (1944). olsen: amphicyon intermedius 165 fine serrations along these edges (Fig. IC). The first premolar is single-rooted, and is absent in the known specimens of the jaw of this animal but its position is located by the alveolus. P2 varies from a double-rooted tooth to one having a single root with an incipient division. The first three premolars have the same shape and cusp pattern as those found in the upper series and are pro- gressively larger from Pi to P3. P4 has a strong accessory cusp posterior to the principal cusp, and the cingulum is well defined. Ml, as in other members of this genus, has a high prominent pro- toconid with the metaconid posterior to it, with little or no separa- tion between the two. The heel is large with the hypoconid and entoconid forming ridges on the labial and lingual edges. The talonid is expanded lingually being wider than the trigonid. The paraconid is a shearing cusp and is placed, in relation to the other cusps, as in Cams. The general structure of this lower carnassial is of a dog-like form. M2 has three principal cusps; the metaconid and protoconid are equidistant from the anterior face of the tooth and of the same height. The hypoconid is prominent and is similar in form to that of Mi. The entoconid varies from being well de- fined to being subordinated in the cingulum. Mr, may be either single or double-rooted, judging from alveoli. Only one specimen of this tooth has been collected (M.C.Z. 7141) ; this is well preserved and single-rooted. As in the corresponding upper molar, it is roughly oval in shape with the principal cusp, the protoconid, situated on the anteroexternal margin. The heel has a cingulum which terminates in a small but distinct cusp just posterior to the protoconid. The upper and lower dental series are not ursid-like and more nearly approach the canids in shape, form, and proportions. Deciduous dentition (Fig. 3A, B). Very little is known of the immature dentition in Amphicyon. However, a note concerning the lower sectorial of A. intermedius is inserted here, if only as a warning to students of the possibility of confusing this deciduous tooth with an adult carnassial of one of the smaller carnivores found in the same quarrj' as A. intermedius. Two specimens of dm4 are known. One is associated with a nearly complete lower jaw (M.C.Z. 3632, Fig. 3A) and was re- 166 bull:ztin: museum of comparative zoology f erred to in White's description of A. intermedius (1940). The other is an isolated tooth (JVi.C.Z. 7142, Fig. 3B). The roots of both are spread in anteroposterior direction due to the pressure of the permanent tooth pushing up from below. This spreading of the roots in the milk molars, not present in the permanent denti- tion, is also obvious in many Recent carnivores. As may be seen from the figures, both teeth are well preserved and little worn. During this study, I had occasion to refer to the type of A. fren- dens (A.M.N.H. 18913) which was described more than two dec- ades ago (Matthew, 1924). Mrs. Rachel Nichols of the American Museum of Natural History brought to my attention the fact that this specimen has a supernumerary M'4 which has never erupted. This tooth was mentioned by M^atthew but no detailed description was given. The tooth is circular and button-shaped with a princi- pal, central cusp and two cuspules situated close together along the outer margin. The undersurface of the tooth is concave im- parting a caplike structure to the whole. It is difficult to de- termine how the tooth was oriented in life, due to its circular structure and lack of identifj-ing characters. It was situated well up in the ascending ramus, the walls of which are very thin around the crypt. Comparison with other American species of Amphicyon Amphicyon americanus Wortman, 1901. Type, Yale Peabod}' Museum 10061, upper Miocene, Nebraska. This is a much smaller species than A. intermedius. The upper teeth, and the molars in particular, have quite different propor- tions from those of A. iritermedius. The heel of M^ is quite con- stricted with no noticeable metaconule. M^ is very narrow com- pared to its length, with the metaconule absent. M' has the general form of A. intermedius but is, in the. type at least, triple- rooted. Amphicyon sinapius Matthew, 1902. A.M.N.H. 9358, upper Miocene, Colorado. Also A.M.N.H. 18258, 18257, 18259B; California Institute of Technology 207 (Stock, 1930), Mio- cene, Oregon; and California Institute of Technology 1621, 1622, 1616, 1619, 1620 (Bode, 1935), upper Miocene, Cali- fornia. olsen: amphicyon intermedius 167 The type of this species, a right M2, agrees fairly well with the Thomas Farm species, with the exception of being rather wider on the anterolabial surface. This is not a profound difference in such a variable tooth. The remaining teeth of the lower dentition (A.M.N.H. 18258) agree closely with those of A. intermedius (M.C.Z. 5832) except for the heavier heel and greater width of Ml in the former. The paraconid of Mi in A . intermedius tends to curve inward rather than maintain a straight line with the proto- conid and hypoconid as it does in A. sinayius. The upper denti- tion in A. si7iapius shows but little difference from that in A. intermedius. Matthew found nothing to separate A. amnicola (Matthew and Cook, 1909; Cook, 1915) from A. sinapius and my comparisons of these species uphold Matthew's observations relating to this synonymy. Amphicyon idoneus Matthew, 1924. Type, A.M.N.H. 18912, middle Miocene, Nebraska. The skull and dentition (A.M.N.H. 20495) differ very little, if at all, from A. sinapius; M'atthew noted that the only essential difference in this species from A. sinapius was its smaller size. If more material of this animal were available, I believe it would be found to be synonymous with sinapius. Amphicyon frendens Matthew, 1924. Type, A.M.N.H. 18913, middle Miocene, Nebraska. California Institute of Tech- nology 376, 377 (Gazin, 1932), Miocene, Oregon; California Institute of Technology 3192 (Wallace, 1946), Miocene, Oregon. This species has a much shorter, heavier jaw proportionally than that of A. intermedius with nearly the same general form as Osteohorus and its relatives. P2 has the same transverse crowding of the alveolus that is found in the borophagines, due to the short space occupied by these teeth. Mi is a much heavier tooth with a higher hj^poconid than that which is found in the other members of the genus Amphicyon. The upper dentition is also of a more robust nature than that possessed by A. intermedius but has the same cusp arrangement as those teeth in the Florida form. 16S bulletin: museum or comparative zoology Amphicyon pontoni Simpson, 1930. Tj^pe, Florida Geological Survey V-4112, lower Miocene of Florida. The type, a left M2 is morphologically indistinguishable from A . intermedius, but is much larger in size. There is a difference of 42 percent between it and the largest known M2 of A. intermedius. Amphicyon riggsi McGrew, 1939. Type, Chicago Natural History Museum P12029, upper Miocene, Montana. A. riggsi has much shorter tooth rows than A. intermedius, with little or no diastema between the lower premolars. The heel of Ml has a distinct swelling in the postero-external margin of the tooth that is peculiar to this species alone. Otherwise, this molar has the same general shape as in A. sinapius. The general form and shape of the upper dentition is the same as in other species of Amphicyon, with the exception of a concavity on the anterior base of the protocone of P^ at its junction with the paracone. ACKNOWLEDGMENTS I wish to express my thanks to Professors A. S. Romer and Bryan Patterson for the full use of all of the Amphicyon material in the collections of the Museum of Comparative Zoology at Harvard University. Thanks are also due Dr. Herman Gunter, Director of the Florida Geological Survey, for permission to use the speci- mens which are a part of the collections in that institution, and to Dr. W . Auffenberg of the Biology Department of the University of Florida, for the loan of the fine skull of A . intermedius, which is a part of the collections of that university, and for permission to describe and figure the specimen. The following people have aided me by furnishing casts of types or other related material in the collections under their care: Dr. G. G. Simpson and Mrs. R. H. Nichols of the American Museum of Natural Historv, Dr. H. J. Cook of Agate, Nebraska, Mr. W. D. Turnbull and Dr. E. C. Olson of the Chicago Natural History Museum and the University of Chicago, Dr. J. T. Gregory of Yale Peabody Museum and Dr. J. A. Wilson of the University of Texas. The figures are the work of Mr. Andrew Janson, artist for the Florida Geological Survey. olsen: amphicyon intermedius 169 TABLE OF MEASUREMENTS All Measurements in Millimeters REFERRED SPECIMENS AND SYNONYMS A. intermedius F.G.S. V5648 A. intermedius M.C.Z. 5832 A. intermedius F.G.S. V5649 A. longiramus (Type) M.C.Z. 3919 A. intermedins M.C.Z. 3632 A. intermedius F.G.S. V5663 A. intermedius (Type) M.C.Z. 3631 A. intermedius M.C.Z. 3630 A. intermedius U.F. 6501 A. intermedius M.C.Z. 4060 A. intermedius M.C.Z. 5833 A. intermedius M.C.Z. 7140 A. intermedius M.C.Z. 7141 (in part) Antero- Greatest Posterior Transverse Diameter Diameter Antero- Greatest Posterior Transverse Diameter Diameter Ml Ml Mj M2 33 17 32.5 17.5 26 19 25.5 17 32.5 17 23.5: 16.5 .... .... 23 15.5 .... .... 21 15.5 30 15.5 20.5 16 29 14.5 . . . . M' Ml M2 M2 27.5 32.5 20.5 29.5 26.5 30.5 21.5 29 26 32 23 31.5 24 29 19 26 .... 19 26.5 170 bulletin: museum of compail\tive zoology REFERENCES Bode, F. D. 1935. The fauna of the Merychippus zone, north Coalings District, California. Carnegie Inst. Washington, Contrib. Paleo., no. 453: 65-96. Colbert, E. H. 1939. Carnivora of the Tung Gur formation of Mongoha. Bull. Amer. Mus. Nat. Hist., 76: 47-81. Cook, H. J. 1915. Note on the dentition of Amphicyon amnicola a gigantic fossil dog. Nebraska Geol. Surv., 7: 57-59. Frick, C. 1926. The Hemicyoninae and an American Tertiary bear. Bull. Amer. Mus. Nat. Hist., 56: 1-119. Gazin, C. L. 1932. A Miocene mammalian fauna from southeastern Oregon. Carne- gie Inst. Washington, Contrib. Paleo., no. 418; 37-86. Hall, E. R. 1928. Records of supernumerary teeth in bears. Univ. Calif. Pub. in Zool., 30: 243-250. Hough, J. R. 1948a. A systematic revision of Daphoenus and some allied genera. Jour. Paleont., 22: 573-600. 1948b. The auditory region in some members of the Procyonidae, Cani- dae, and Ursidae. Bull. Amer. Mus. Nat. Hist., 92: 71-118. KtJRTEN, B. 1955. Sex dimorphism and size trends in the cave bear Ursus spelaeus Rosenmiiller and Heinroth. Acta Zool. Fennica, 90: 1-48. Matthew, W. D. 1902. New Canidae from the Miocene of Colorado. Bull. Amer. Mus. Nat. Hist., 16: 28-290. 1924. Third contribution to the Snake Creek fauna. Bull. Amer. Mus. Nat. Hist., 50: 59-210. 1909. (and Cook, H. J.) A Pliocene fauna from western Nebraska. Bull Amer. Mus. Nat. Hist., 26: 361-414. olsen: amphicyon intermedius 171 1904. (and Gidlej^ J. W.) New or little known mammals from the Mio- cene of South Dakota. Bull. Amer. Mus. Nat. Hist., 3: 241-268. McGrew, p. O. 1939. A new Amphicyon from the Deep River Miocene. Geol. Ser., Field Mus. Nat. Hist., 6: 341-350. Merbiam, J. C. 1912. The fauna of Rancho La Brea, Part Il-Canidae. Mem. Univ. Calif., 1: 215-272. Peterson, O. A. - 1909. A new genus of carnivores from the Miocene of western Nebraska. Science (n. s.), 29: 620-621. 1910. Description of new carnivores from the Miocene of western Ne- braska. Mem. Carnegie Mus., 4: 205-278. Scott, W. B. 1890. Preliminary account of the fossil mammals from the White River and Loup Fork formations contained in the Museum of Compara- tive Zoology. Part Il-Carnivora and Artiodactyla. Bull. Mus. Comp. Zool., 20: 65-100. 1936. (and Jepsen, G. L.) The mammalian fauna of the White River Oligocene, Part I-lnsectivora and Carnivora. Trans. Amer. Phil. Soc. (n. s.), 28: 1-153. Segall, W. 1943. The auditory region of the arctoid carnivores. Zool. Ser., Field Mus. Nat. Hist., 29: 33-59. Simpson, G. G. 1930. Tertiary land mammals of Florida. Bull. Amer. Mus. Nat. Hist., 59: 149-211. 1932. Miocene land mammals from Florida. Bull. Florida Geol. Survey, 10; 11-41. 1944. Tempo and mode in evolution. Columbia Univ. Press, New York: 1-237. 1945. The principles of classification and a classification of mammals. Bull. Amer. Mus. Nat. Hist., 85: 1-272. 1949. A fossil deposit from a cave in St. Louis. Amer. Mus. Novitates, no. 1408: 1-46. 172 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY Stock, C. 1930. Carnivora new to the Mascall Miocene fauna of eastern Oregon. Carnegie Inst. Washington Contrib. Paleo., no. 404: 45-76. Van der Klaauw 1931. The auditory bulla in some fossil mammals with a general intro- duction to this region of the skull. Bull. Amer. Mus. Nat. Hist., 62: 1-352. Wallace, R. E. 1946. A Miocene mammalian fauna from Beatty Buttes, Oregon. Car- negie Inst. Washington Contrib. Paleo., no. 551: 115-134. White, T. E. 1940. New Miocene vertebrates from Florida. Proc. New England Zool. Club, 18: 31-38. 1942. The lower Miocene mammal fauna of Florida. Bull. Mus. Comp. Zool., 92: 1-49. Wortman, J. L. 1899. (and Matthew, W. D.) The ancestry of certain members of the Canidae, the Viverridae, and Procyonidae. Bull. Amer. Mus. Nat. Hist., 12: 109-139. 1901. A new American species of Amphicyon. Amer. Jour. Sci., (4) 11: 200-204. Young, S. P. 1944. (and Goldman, E. A.) The wolves of North America. Stackpole Company, Harrisburg, Pa.: 1-588. 1951. (and Jackson, H. H. T.) The clever coyote. Stackpole Company, Harrisburg, Pa.: 1-411. B A-(l) (2) B C Figure 1 Lower jaw and dentition of Aniphicyon intermedius Lower jaw and dentition, labial and occlusal views showing wear pattern of old adult, M.C.Z. 3919. Adult P4 - Ms, labial and occlusal views showing no perceptible wear; P4, F.G.S. V-5257; Mi - M2, M.C.Z. 5832; M3, M.C.Z. 7141. , , Same as 2, lingual view. Cusp nomenclature for lower dentition. Left lower canine; (1) lingual view, with detail of serrations. (2) Posterior view with sections. (3) Enc(nil Audlrory PartocctpitBl -Practss^. Occipitml Coadylc , \ tlondyUt Foramen ^ Fotmtaea Lacenun Pesie u> Scylomastsitl Fommea— ^-^ E.,„hi„ ' Po»[|i1eAoitl Foramra Figure 2 Skull of Amphicyon intermedius U. F. 6501 A — Dorsal view. B — Lateral view. C — Palatal view with detail of auditory region — crushed and restored areas as indicated by dashed lines. A — Figure 3 Immature jaw and deciduous carnassial Immature jaw with dm4 (replaced bv P4\ AI.C.Z. 3632. B — Dm4, M.C.Z. 7142. ProtocoQc Metaconr c Hypocone B A 0 5 I) to mm Figure 4 Upper dentition of Amphicyon intermedius A — Occlusal and lingual views of adult showing no perceptible wear F^ - I\P. PS M.C.Z. 4060; AP, AP, M.C.Z. 5833; M\ M.C.Z. 7143. B — Occlusal and lingual views of P* - AP of old adult showing pattern of wear. U. F. 6501. C — Cusp nomenclature of upper dentition. LEFT M- MCZ 7140 V-5647 MCZ 5833 Amphicyon intermedius LEFT M- MCZ 4655 I MCZ 46548 MCZ 45646 Canis lupis INDIVIDUAL TOOTH VARIATION Figure Bulletin of the Museum of Comparative Zoology AT HARVARD COLLEGE Vol. nS, No. 5 CONTRIBUTIONS TOWARD A RECLASSIFICATION OF THE FORMICIDAE. II. TRIBE ECTATOMMENI (HYMENOPTERA) By William L. Brown, Jr. CAMBRIDGE, MASS., U. S. A. PRINTED FOR THE MUSEUM June, 1958 Publications Issued by or in Connection WITH THE MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD COLLEGE Bulletin (octavo) 1863 — The current volume is Vol. 118. Breviora (octavo) 1952 — No. 86 is current. Memoirs (quarto) 1864-1938 — Publication was terminated with Vol. 55. Johnsonia (quarto) 1941 — -A publication of the Department of Mollusks. Vol. 3, no. 35 is current. Occasional Papers of the Department op Mollusks (octavo) 1945 — Vol. 2, no. 21 is current. Proceedings op the New England Zoological Club (octavo) 1899-1948 — Published in connection with the Museum. Publication terminated with Vol. 24. The continuing publications are issued at irregular intervals in num- bers which may be purchased separately. Prices and lists may be obtained on application to the Director of the Museum of Comparative Zoology, Cambridge 38, Massachusetts. Of the Peters ' ' Check List of Birds of the World, ' ' volumes 1-3 are out of print ; volumes 4 and 6 may l)e obtained from the Harvard Uni- versity Press; volumes 5 and 7 are sold by the Museum, and future volumes will be published under Museum auspices. Bulletin of the Museum of Comparative Zoology AT HAEVAED COLLEGE Vol. 118, No. 5 CONTRIBUTIONS TOWARD A RECLASSIFICATION OF THE FORMICIDAE. II. TRIBE ECTATOMMINI (HYMENOPTERA) By William L. Brown, Jr. CAMBEIDGE, MASS., U. S. A. FEINTED FOE THE MUSEUM June, 1958 No. 5 — Contrih lit ions toward a Reclassification of the Formi- cidae. II. Tribe Ectatommini (Hymenoptera) By William L. Brown, Jr. CONTENTS Page Introduction 175 Emery -Wheeler classification, with a summary of proposed changes ... 17G Characters and relationships among the genera 178 Geographical distrilmtion of the genera 181 Treatment of species-level taxonomy 184 Key to the genera of Ectatommini (workers) 185 The genera 188 Acanthoponera Mayr 188 Heteroponera Mayr 194 Rhytidoponera Mayr 197 Paraponera Fr. Smith 205 Aulacopone Arnoldi 206 Eetatomma Fr. Smith 206 Gnamptogenys Roger 211 Proceratium Roger 241 Discothyrea Roger 248 Appendix 254 INTRODUCTION This section deals with a large group of species in the sub- family Ponerinae, including most genera formerly placed in the old tribes Ectatommini, Paraponerini and Proceratiini. These genera are now grouped in a single, broadened tribe Ectatom- mini. The previous "standard" classification of the subfamily Pon- erinae, stabilized by Emery in his 1911 fascicle of the Genera 176 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Insectorum, was altered l)ut little when Wheeler made additions to it in his generic key of 1922. In 1911, Emery's system was a great advance over older arrangements in many respects, but the flood of description in more recent years has swamped it completely. Since basic classification has been neglected for so long a time, we are now faced with the necessity of constructing a completely new system. I have undertaken to make this replacement through the revision of tribes or similar-sized taxa, one at a time, and the publication of the results of tribal units. Thus, at least some of the results will be available for the neces- sary use and testing by other specialists long before a full sub- family revision could possibly be completed. It is hoped, however, that after the series of tribal sections is complete, there will be an opportunity to make final adjustments and to construct a grand key to the genera of the subfamily. The Emery- Wheeler classification of the Ectatommini, Paraponerini and Proceratiini, with a summary of the clianges here proposed Emery recognized the Ectatommini, Paraponerini and Procera- tiini as separate tribes within the Ponerinae, but the present evidence of their close relationship seems to make any such tribal division unnecessary and unrealistic. The Paraponerini include a single genus with one species, and the conclusion reached after a close consideration of the characters of all castes of Paraponera clavata is tliat trilial rank was granted in this case almost solely on the basis of the extraordinarily large size of the individuals of the species. Admittedly, Paraponera is unique in a couple of other respects as well, but its ectatommine relationships are so clear in its habitus in all adult castes, in its behavior, and in details of morphology such as the male genitalia (Weber, 1946) that it can no longer be considered as more than a slightly aberrant ectatommine. The Proceratiini have even less claim to tribal separation from the Ectatommini, at least on adult characters. In fact, there is grave difficulty to be encountered in separating Proceratium of the Proceratiini from Heteroponera of the Ectatommini even at the generic level. The only doubts here rest on the unusual BROWN : ANT TRIBE ECTATOMMINI 177 nature of the proceratiine larvae, but then these are known for only a few species in one genus, rroccratium, and for none of the Iletcroponera species. The Ectatommini appear to make up one reasonably clearcut tribe with these inclusions. Emery divided his tribe Ectatommini into two subtribes : Ecta- tommini, scnsu stricto and Typhlomyrmicini. It has already been shown that two of the three genera of the Typhlomyrmicini {Typlilomyrmex Mayr and Prionopelta Mayr) are members of different ponerine tribes, and that they should be excluded from consideration under the Ectatommini (Brown, 1950, 1953a, 1953b). The third genus in this subtribe is Rhopalopone Emery, the species of w^hich have been consistently characterized, and wrongly so, as lacking median teeth on the tarsal claws. In fact, such teeth are present in most, if not all species that have been placed in Rhopalopone, and a separation between these species and others now placed in Gnanipiogenys is considered impossible. The sj'nonymy of Rhopalopone with Gnamptogenys removes the last genus from subtribe Typhlomyrmicini, and thus removes all necessity for recognizing this subdivision any longer. I had considered retaining Paraponerini and Proceratiini as subtribes in the Ectatommini, sensii lato, but in the end this subdivision seemed misleading and an unnecessary complication to the sys- tem, and it was rejected. Among the genera of the ectatommines, Acanthoponera is worthy of generic rank on present evidence. The erstwhile sub- genus Anacanthopo7iera is a straight synonym of genus Hetero- ponera Mayr (Brown, 1952c), and Paranomopone is also con- sidered to belong to Heteroponera in this revision. Aulacopone is retained as a genus, at least until more is known about it. Rhyticloponera is retained as a separate genus, also, and Chalco- ponera is included as a synonym (Brown, 1953b). Ectatomma is considered to be a genus apart from Gnamptogenys and the remainder of its erstwhile subgenera. Gnamptogenys is pre- served as a separate genus and. despite long and earnest attempts to find some basis for separation of two or more generic or sub- generic groups, I have been forced to place in its synonymy a number of generic and subgeneric names, including some much- used and familiar ones : Ilolcoponera, Stictoponera, Emeryella, Parectatomma, Rhopalopone and Alfaria, as well as some less 178 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY well known names: Wheeleripone, Spaniopone, Poneracantha, Barhourella, Opisthoscyphus, Tamnioteca and Commateta. Of the genera formerly included in the Proceratiini, Proholomyrmcx Mayr and its probable synonym Escherichia Forel have been transferred to tribe Platythyreini (Brown, 1952a). Proceratium has been found to grade into Sysphi7igta {Sysphincta of authors), so synonymy is indicated here, with Proceraiium the prior name. For similar reasons, Discothyrea becomes the senior synonym of Prodiscothyrea and Pscudosysphincta. The Ectatommini come in this way to contain nine reasonably distinct genera : Acanthoponera, Hcteropo7iera, Bhytidoponera, Paraponera, Ectatomma, Aulacopone, Gnampfof/enys, Procera- tium and Discothyrea. Characters and Relationships among the Genera of Ectatommini Taxonomy in the Ectatommini is chiefly dependent upon the characters of the adult worker, for this is the caste most com- monly collected and described, and the one with the clearest differentiation of external morphological features. Queens follow workers in generic and specific characters for the most part, and many of them are present only as wingless ergatoids scarcely distinguishable from workers, or perhaps not distinct at all [Rhytidoponera spp.). The degree of caste divergence in the female sex is moderate, therefore, to nearly or quite non-existent. The males show marlted divergence from the females, but sexual dimorphism is not as strong as among some other formicid genera, even in the Ponerinae. The size is usually a little smaller than the corresponding female, and the sexual differences are otherwise much as seen in other ponerines. Males are rare in collections, and often are not in association with workers or females in the cabinet. Within the tribe, the males show greater conservatism in morpliology than do the workers and queens, and, with few exceptions, the male characters are not known to furnish clear generic distinctions. The males of a species may often be distinguished by characters reflecting tliose shown more strongly by its workers and queens, so that males in associa- tion with workers are normally determined through the workers. BROWN : ANT TRIBE ECTATOMMINl 179 Perhaps a study of such characters as wing venation, genitalia, palpal segmentation, antennal form, and so on, will eventually furnish good male distinctions independent of those seen in the workers, but such a study must await much more complete collections. Some male characters will be mentioned below for the genera in which this caste is known. The larvae of the ectatommines are also relatively incompletely known, and before the pioneering work of the Wheelers on the ponerine larvae (1952a: 117-139, pis. 2-5; 1952b, 657-661), there was virtually nothing in print of any value for morphological or systematic purposes. The larval findings agree in most respects at the generic level with the new classification adopted here. The Wheelers have been hampered in their work by a lack of material in some of the rare, but taxonomically critical genera. It is hoped that those specialists having access to the material needed to fill these gaps will see that larval specimens are forwarded to colleagues ready to study them. At the present time, the relationships of the Ectatommini are not entirely clear, but it does seem likely that tribes Ponerini and Platythyreini are approximately cognate members of the Poneroid Complex (Brown, 1954a) . There is evidence in the form of the male alitrunk, wings and genitalia, in the integumental consistency, proventricular structure, propodeal form and arma- ment of all castes, and especially in the structure and propor- tions of the post-propodeal segments and their organs, that subfamily Myrmicinae arose from an ectatommine ancestor, and this supposition is strengthened by the characteristics of the fossil Agroecomijnnex Wheeler, which seems to provide a near- ideal intermediate form. These speculations on ectatommine con- nections seem reasonably safe to put forward at this time, but beyond this, nothing is gained by suggesting detailed relation- ships in the absence of the necessary comparative morphologi- cal data. Within the tribe Ectatommini, the genus surviving with the greatest number of primitive characters appears to be Acanfho- ponera. This genus retains (in the female castes) the primitive palpal formula for ants : 6 maxillary, 4 labial segments. Acan- ihoponera also bears the strong extra tooth on each tarsal claw found in the most primitive ponerines and myrmeciines, and 180 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY the alitrunk has a complete and possibly flexible articulation between pro- and mesonotnm. It is interesting to note that this genus, like some other genera primitive within their respective tribes, is fitted with the large eyes and strong spinous armament marking an arboreal forager, and it is now known that Acantho- potiera is in fact a nocturnal arboreal forager. All of the charac- ters mentioned above, which happen to be generalized conditions for the ants as a family, oppose the tendency toward ankylosis of antennae, palps, and alitrunk. Other trends that cause the body to become more compact, such as reduction of spinous arma- ment, and also the reduction of the eyes, accompany the aiikylotic tendencies more or less consistently, apparently in correlation with a change from arboreal through ground-foraging to hypogaeic or otherwise cryptobiotic habits of the ants. This trend is clearly observable within the single myrmicine tribe Dacetini, as it is within the ectatommines. The frequent association of arboreal, tropical forms with characters generally considered to be the generalized ones for ants indicates a greater likelihood that the forested tropics were a more prolific evolutionary source in terms of major groupings than Avere the more arid regions usually cited as important to formicid major-group evolution (see also Ber- nard, 1948). AcantJwponera is advanced in one respect, however ; the known winged specimens of both sexes lack the anal lobe of the hind wing. All other ectatommines lack this lobe, too, except Para- ponera and Ectatomma, each of which has well-developed lobes in all of the species so far seen. The loss of the anal lobe may mark a monophyletic lineage within the ectatommines, since there are additional grounds for linking most of the genera without it. Heteroponera, for instance, can be derived directly from Acanthoponera, and RJiytidoponera is very closely related to Heteroponera, especially through the species H. relict a. Judging from the original description and figures, Aulacopone also seems to be close to Heteroponera, but this needs to be confirmed by a new examination of the type or additional material that may have accumulated in collections. Proccratimn and DiscotJiyrca seem, on the basis of adult cliaracters, to be closest to Heteroponera, and the amber species Bradoponera mcieri (Mayr, 1868 ; Wheeler, 1914) looks like a reasonable intermediate step in this line. A BROWN : ANT TRIBE ECTATOMMINI 181 comparative study of the larvae in this group is needed in order to decide how real the link between the " proceratiines " and Iletcroponcra may be. The large and heterogeneous Gnamptogenys group is difficult to relate precisely to other genera, but it seems closer to the Acanthoponcra-IIeteroponera line than to Ectaiomma, despite previous placements to the contrary. This leaves Ectaiomma and Parapuncra as offshoots from near the base of the ectatommine stock. The poorly-known Baltic Amber genus Elcctroponera Wheeler may represent an intermediate link between these genera and the main line of the tribe. Two other fossil species, Ectaiomma europaeum Mayr (1868) and Archiponcra wheeleri Carpenter (1930; see also Wheeler, 1930) appear to belong either in or very close to Gnamptogenys in the present broad sense. Gnamptogenys europaea (Mayr) is the new combination for the Baltic Amber species, proposed tentatively' on the basis of the original description of the winged female and Wheeler's (1914) characterization of the supposed male of this species. As AVheeler suggests, Archiponera may be near the schmitti group of Gnamptogenys {= Emery ella), al- though the state of preservation of the fossil, from the Oligocene Florissant Shales, leaves something to be desired. All of the fossils are interesting chiefly in that they show a wddely diversified group of ectatommines to have existed at least as far back as the Oligocene, in northern areas from which the tribe is now absent completely or nearly so. Geographical Distribution of the Genera The pattern of ectatommine distribution is today essentially a peripheral one. The relatively large-bodied, epigaeically-forag- ing stocks are developed almost exclusively in the Neogaeic and Indo-Australian (especially Australian) Regions, where they are largely confined to tropical and warm temperate climates. Prom this, it will be clear that by "peripheral," I mean peri- pheral to the larger continental land masses of Eurasia, North America and Africa. However, since the presence of epigaeically- foraging stocks is clearly indicated in the fossil record of the Northern Hemisphere, we know that the distribution of ecta- tommines has not always been an essentially fringing one, and 182 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY we suspect that the group may have had its principal source of evolutionary change in the central regions during earlier times. In the same central regions now, we find only a thin and discontinuous representation of specialized cryptobiotic types, like the species of Proceratium and Discothyrea, managing to survive by remaining ecologically unobtrusive. Among the epigaeic genera, Paraponera, Ectatomma, and Acanthoponera are restricted to the New World in warm climates and are all clearly distinct from Old AVorld members of the tribe. Heteroponera, a rather conservative stock, is distributed rather widely in both the Australian and Neogaeic Regions, but shows no obvious tendency to split into two groups following the geographical separation. Ehytidoponera is a very successful and luxuriantly radiating stock of the Australian Region ; it halts at New Caledonia in the east, but ranges widely through Melanesia to the islands stretching from Timor to the southern Philippines on the west. Rhytidoponera looks as though it may have originated in the Australian region from some Hetero- ponera-like stock. Aulacopone has so far been reported only from the type locality in Russian Armenia ; obviously, it has cryptobiotic tendencies. Gnamptogenys is a large and wide-ranging genus, occurring both in the Indo- Australian Region (Ceylon and western China to Fiji, but not known from Australia, New Caledonia or New Zealand) and in the New World (Argentina to Texas, West Indies). This genus can be grouped into different lines, more or less equivalent to the genera and subgenera here synonymized, and reflecting in part the separation between New and Old World, as well as the Avater gap between archipelagic and con- tinental faunas. However, these lines have not lost their associa- tions through intergradient species, and these intergradient species, most of which exist in the New World, show that the lines still belong to the same genus at the present time level. It thus appears that Gnamptogenys must represent the last major radiative burst of the epigaeic eetatommines, already replaced in its center of origin, and unable to expand in the direction of Australia Ijecause of the powerfully-developed Rhytidoponera stock already present in that country. BROWN : ANT TRIBE ECTATOMMINI 183 Of the two predominantly cryptobiotic genera, Proceratium and Discothyrca, the former tends to have a more northerly range, while the latter is found more to the south. Both genera occur in the tropics, often in the same areas, and there are broad overlaps in range in both northern and southern temperate areas. Discothyrea is best developed and most abundant in tropical America and southward to Argentina, in the Australian region, and in central and southern Africa. Proceratium reaches a peak of abundance and diversity in the warmer parts of the llolarctic Region, wherever sufficient moisture exists, and this genus is well represented in the East Indies. In Central America there are three rare species, and two species have reached into the eastern tropical forest belt of Australia as far south as Brisbane. Another species has colonized Fiji. Two species are known from tropical Africa. This apparent tendency of the two genera to have complementary ranges outside the tropics is very interesting, but we know so little about the habits, particularly the food liabits, of these ants that it is not now possible to say whether the tendency is adaptive in promoting avoidance of competition between the groups. The presence of two endemic ectatommine species of different stocks {Hctcroponera and Discothyrca) in New Zealand (out of a total of at most ten ant species endemic in New Zealand), and of moderately to highly diverse native representations of the tribe in southern South America, southern Australia, the Solomons, Fiji and New Caledonia, speaks for the age of the group. Few ant tribes can match this " extralimital' ' range, and of those that do, the Amblyoponini share an archaic aspect and a poverty of representation in the Ethiopian and other "central" land masses. Bearing in mind the Oligocene fossil traces and the lack of good ant fossils in earlier levels, it seems likely that the Ectatommini go back to or slightly beyond the beginning of the Tertiary. It may be guessed that the present peripheral contrac- tion of the Ectatommini is due, at least in part, to the pressure of the rising Myrmieinae. The beginnings of a diversified myr- micine fauna are seen in the Baltic Ambei-, as well as in the Florissant Shales, but true members of the most potent myrmi- cine genera, Pheidolc and Crematogaster, do not appear in these formations, nor even in later (Miocene) strata. 184 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Treatment of Species-Level Taxonomy The primary purpose of this section is to revise the generic classification of the Ectatommini, but in the course of the neces- sary review of the literature and authentic material of nearly all of the named forms, many species-level situations were found to require synonymy, adjustment or amplification. Also, among a number of previously undescribed species seen, certain ones were found to affect generic concepts by broadening them or by bridging what were formerly thought to be natural gaps. For these reasons, descriptions of a few new species were neces- sary to the primary purposes of this work ; descriptions of still other species that do not affect generic concepts as significantly have been omitted here, though it is hoped that these can be treated elsewhere. The compilation of a work of this sort also inevitably includes a miscellany of new or under-appreciated data referring to morphology, distribution, behavior, ecology, and variation, of a kind lacking in the literature for most of the forms in this tribe. In order that the required documentation backing the pres- ent major conclusions, and also the subsidiary information, should neither clutter the presentation of the generic data nor become a lost by-product, I have relegated most of this matter to an appendix, where the items are listed consecutively against boldface numbers corresponding to the indicator numbers placed in brackets in the text. After synonymy and general remarks, there are listed for each genus the currently recognized species as far as known to me. Synonyms listed as such in Emery (1911) are omitted here. Each species is followed by author, date and page of original reference ; the sources are listed more fully at the end of the paper. Of the many forms treated until now as varieties or subspecies, some are rather obviously synonyms, and are treated as such with a minimum of discussion. Varieties and subspecies of unclear status are listed arbitrarily as species, since I feel that these names will eventually stand or fall as species or as infraspecific variants not worthy of formal nomenclatorial recognition (Wilson and Brown, 1953; Brown and Wilson, 1955). I should perhaps em- phasize that the listing of former infraspecies names as species BROWN : ANT TRIBE ECTATOMMINI 185 here implies no real conviction on my part that these entities are really worth the distinction; actually, I feel that most of such names will be revealed as synonyms when the proper study of their types has been made. The capital letters placed in parentheses before each specific name indicate the kind of evidence upon which the present generic placement is directly based. (T) indicates that type material, nidotypes, reliably type-com- pared material, or similarly authentic specimens have been ex- amined, in most cases by myself ; rarely, examination has been made by other myrmecologists. (P) means that material identified from reasonably good de- scriptions, or from other satisfactory evidence, has been examined and is thought to be correctly determined. In cases where the species is not seriously questioned, but no specimens referable to it have been seen, or if specimens seen cannot be satisfactorily verified as to identity, no entry has been made before the species name. (?) signifies that, in my opinion, the species is inadequately described for purposes of distinction and that its taxonomic status is doubtful. The species lists have been based on various myrmecological compendia and basic ]iapers, and were checked against Emery's Genera Insectorum list and the Zoological Record, 1908 through 1953. References through 1957 are included so far as I am aware of them. I shall consider it a great favor if readers will send me notice of the inevitable omissions for inclusion in a corrective supplement. Key to the genera of Ecfatomniini. based on the workers 1. Size very large, head width across eyes over 3.6 mm.; hypopygium with an upwardly-directed comb of fine teeth on each side (Neotropical) Paraponera Fr. Smith Size much smaller, head Avidth across eyes under 3.6 mm.; hypopygium without lateral combs 2. 2. Lobes of frontal carinae more or less sharply raised (sometimes vertical and fused together), leaving the condylar bulbs of the antennae com- pletely or nearly completely open to dorsal full-face view (Figs. 1, 23-25, 186 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 46, 48). 8mall, compact chiefly cryptobiotic forms, usually ferruginous in color and with reduced eyes (Figs. 45, 47) 3. Lobes of frontal carinae horizontal or at most feebly raised, covering or nearly covering the condylar bulbs of the antennae (Figs. 12, 36, 41, 43, 44). Mostly epigaeic foragers, but a few species are cryptobionts with characters to match 4. Figure 1. Proceralium arnoldi Forel, syntj'pe worker, dorsal view of anterior part of head, from a sketch by Dr. G. Arnold. Figure 2. Ecta- tomma ^confine Mnyr, worker (Lancetilla, Tela, Honduras), side view of alitrunk and petiole. Figure 3. Ehytidoponera laciniosa Viehmeyer, worker, side view of ali- trunk. Inferior pronotal tooth indicated bj' arrow. 3. Apical antennomere strongly bulbous, its length subequal to or > the combined lengths of the second through penultimate funicular segments (Fig. 48) ; mandibles small, edentate (though frequently with a masti- catory comb of short setae), largely or completely overhung by clypeus (Widespread in tropical and warm temperate regions) Discothyrea Roger BROWN : ANT TRIBE ECTATOMMINI 187 Apical antennomere at most moderately enlarged, much shorter than combined lengths of second through penultimate funicular segments; mandibles larger, armed with 3 or more teeth or denticulae, not or only in very small part overhung by clypeus (Figs. 1, 23-35, 45, 46) (Wide- spread except S. America) Proceratiwm Roger 4. Inferior pronotal margins just in front of each anterior coxa with a distinct, usually acute tooth, best seen from an oblique dorsolateral view (Fig. 3) (rarely missing on one or both sides in individual specimens); posterior tarsal claws always with a distinct median tooth; posterior coxae unarmed above (Australia; N. Guinea, to N. Caledonia, s. Philippines) Ehytidoponera Mayr Inferior pronotal angles unarmed or forming an obtuse angle, or, in the rare cases where the angle is present and more nearly toothlike, then the posterior tarsal claws lack a median tooth, or else the posterior coxae are toothed above 5. 5. Mesonotiim rounded and prominent, surrounded by deeply impressed sutural lines ; it and the propodeum forming distinct convexities separated by the deep transverse fissure of the metanotal remnant (Fig. 2) ; pos- terior coxae always unarmed (Neotropical) Ectatomma Fr. Smith Mesonotum and propodeum part of one continuous or near-continuous profile, interrupted at most by a suture-like groove at the position of the metanotum (rarely deep), or by an ill-defined impression in this region; posterior coxae armed above, or unarmed 6. 6. Clypeus much modified, the median portion raised and produced as a short, blunt triangular point forward from the antenna! insertions somewhat over the mandibles ; antennal scrobes lateral, long and very deep (Russian Armenia, rare) Aulacopone Arnoldi Clypeus of normal form, broad and in one piece, not or only to a small extent covering the mandibles (Figs. 12, 41, 43, 44) ; antennal scrobes distinct and deep in only a few species 7. 7. Dorsum of head without a median costa as distinct from other sculpture, or, if with a costa, it is short or not fully continuous (Figs. 41, 43, 44), and either the posterior coxae are toothed above (Figs. 17-20), or else the alitrunk and petiole are predominantly smooth and shining (Neo- tropical, Indo- Australian) Gnamptogenys Roger Dorsum of head with a distinct median costa from clypeus to vertex, con- tinuous across frontal triangle (Fig. 12) ; posterior coxae unarmed above (Fig. 13) ; head and alitrunk thickly sculptured, usually hairy and pubescent and in large part opaque 8. 8. Tarsal claws each with a prominent basal lobe and a large submedian tooth; propodeum with paired long teeth and petiole produced pos- terapically as a single long spiniform tooth (Figs. 7-11) ; palpal formula 188 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 6, 4. (Neotropical) Acanthoponera Mayr Tarsal claws without prominent basal lobes, submedian tooth often absent, particularly on claws of posterior legs; propodeum and petiolar summit unarmed or with short teeth (Fig. 13); palpal formula 4, 3 or less (S. and C. America to Mexico; Australia, N. Zealand) . . Heteroponera Mayr THE GENERA ACANTHOPONERA MaYR § Ectatomma subgenus Ac-anthoponcra Mayr, 1862:732. Type: Ectatomma (Acanthoponera) mucronatum = Po7iera viucronata Eoger, 1860, by desig- nation of Emery, 1911. < Acanthoponera, Emery, 1911, p. 35; nee male. ^Acanthoponera subgenus Acanthoponera, Wheeler, 1932b: 190. It has already been shown (Brown, 1952c) that Heteroponera Mayr is the prior name for a group of species included by Emery in Acanthoponera, and ranked as subgenus Anacanthoponera by Wheeler (see below). Heteroponera deserves to be regarded as a separate genus on present evidence. The species remaining in Acanthoponera as defined here are medium-sized ants (workers ranging from 5 to 10 mm. in out- stretched length), generally of a yellowish or tan color, with large, convex eyes and shallow aiitennal scrobes. The propodeum bears a pair of teeth or spines (long and slender in worker), and the apex of the petiolar node is produced dorsocaudad as a slender subconical spine with a more or less acute tip. The tarsal claws are very well developed, and each has not only a strong submedian tooth, but also a prominent, narrow lobe resembling a third tooth. The palpi of the female and worker are segmented maxillary 6, labial 4 (this seems to be the formula in the single male available, although the basalmost segment in each of the palps of this specimen is not properly visible under the partly- retracted maxillae and labrum) ; this formula is the primitive one for the Formicidae, and it is not known to hold in any other ectatommine genus. The palpi are long and slender as compared to those of other ectatommines. The characters above are those expected of an epigaeic, nocturnal forager that climbs trees or other vegetation, habits now confirmed by observations of Weber (1939) and Wilson (personal communication) on A. minor. BROWN: ANT TRIBE ECTATOMMESTI 189 The females of Acanfhoponcra, where known, are winged, slightly larger than workers from the same nest, and differ from the workers in the same ways that ponerines most often do. The female propodeal teeth are usually less well developed than in the worker. The color is the same or nearly the same as in the corresponding workers. The forewings have "complete" venation of the type of Ectatonima, l)Ut the hind wings differ from those of Ectatomyna in that they lack the anal lobe. In the single Acanfhoponera female available with wings, there is no trace of the first radial crossvein remaining, although such traces are present in the male specimen seen. The male of this genus is known to me only from a single speci- men sent by Father Borgmeier ; I believe that this is the first true AcantJioponera male to be recorded. It will suffice here to de- scribe this male as like a larger, more slender Heteroponera with tarsal claws as in worker-female Acanfhoponera. A further notable character is the very extensive ventral excavation of the gaster ; the cavity involves large parts of the second and following segments, and even the posteroventral border of the postpetiole. No such conformation exists in the few Heteroponera males I have seen, nor does it occur in other genera of ectatom- mines. More species must be examined in the male sex before we know whether or not this is a generic, or only a specific, character. The general color of this male is ferruginous yellow. The eyes are large, convex, and medially emarginate ; the ocelli are large and clear. As already mentioned above, the male specimen seen has traces remaining of the first radial crossvein, visible in good light as tenuous veinlets extending downward from R toward Rs, but not reaching Rs before they fade out. Acanthoponera is a rarely collected genus, less than a score of separate collections having reached myrmecologists to my knowledge. Nothing is known of its nesting habits or food, and the larva has never been seen, so far as I can learn. The genus is restricted to the Ncav "World, Avhere it is known to range from Veracruz south into northwestern Argentina. Apparently it is restricted to forested country, although information on this is scanty. Probably an increase in night collecting in the tropics will give us more insight into the biology of this most interesting generalized ectatommine. 190 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY Eight names have been proposed for forms in the genus Acan- thoponera, each based on one or two type specimens only. Ma- terial, whether determined or undetermined, is very scarce in collections, and the wide scattering of type depositions adds to the difficulty of revision at species level. Among the 21 w^orker and female specimens I have seen from 15 nests or individual collections, there appear to be only five distinct entities, each of which may represent a distinct species. It should be emphasized, however, that no real idea of species variability or species limits can be securely gained from such limited material. It is possible that all 21 specimens actually belong to one or two species only. Figures 4-6. Acanthoponera spp., workers, dorsal views of first two gastric segments. Fig. 4, A. minor. Fig. 5, A. mucronata. Fig. 6, A. pernvmna sp. uov., liolotypc. All drawn to same scale. Of the five kinds, two have the "unconstricted" type of gaster, in which the second gastric segment is narrower than the post- petiolar segment in which it is based, and from which it issues without notable constriction, tapering from this point toward the apex, as shown in Figure 4. Of the two kinds with uncon- stricted gaster, one averages a little smaller and has a petiole shaped as in Figure 9 ; to this kind belong the holotype of schwarzi, the cotypes of sphiinodis, and a specimen taken in Veracruz by Wilson. The type of spininodis before me has the BROWN : ANT TRIBE ECTATOMMINI 191 head distinctly broader than in the Guatemalan or Mexican speci- mens, but the diiferenee is not great enough to warrant continu- ing spininodis as a species apart from minor, which name I think should apply to the first unconstricted-gaster kind. In applying this name, I am relying on Forel's statement concerning the small size of minor, and also the fact that his type locality is Teapa, Tabasco, near the known localities for this species in Mexico and CTuatemala from which my specimens have come. Forel also states that the spine of the petiolar node in minor is less strongly elevated and forms less of an angle with the node, which, if I interpret the statement correctly, applies well enough to the specimens here assigned to minor. The second kind with unconstricted gaster is represented in my collection by a worker from Hamburg Farm, Santa Clara Prov., Costa Rica (F. Nevermann), and by three workers from 6 miles west of Santo Domingo de los Colorados, Pichincha, Ecuador (leg. E. I. Sehlinger and E. S. Ross). The average size is a little larger than in minor (average head width, without eyes, ca. 1.04 mm., as against a head width of 1.00 mm. or less in minor), and the node is thicker from front to rear, with the spine elevated a bit more strongly (Fig. 10). The Costa Rican specimen is light yellowish ferruginous in color (teneral ?), and the Ecuadorian workers are considerably darker, yellowish- brown. The metanotal groove is represented by a distinct im- pressed line, averaging more distinct than in minor. 1 doubt whether these specimens belong to minor, and their correspond- ence with other described species seems doubtful also ; perhaps they represent an undescribed species, and it seems best to de- scribe and name them formally, in view of the probabilities, as Acanthoponera crassa sp. nov. [2]. Among the three kinds with constricted gaster, the smallest is represented by a single, winged female taken at Agudos, Sao Paulo State, Brazil (December 15, 1955), at light by W. W. Kempf. This specimen has a peculiarly-formed petiolar node (Fig 11), and the gastric dorsum is shining, with small, sep- arated punctulae. The gastric constriction is distinct, but not especially strong. I cannot match this female with any known species, but it seems best to wait for material in which workers are associated before deciding whether it should be named. 192 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Of the kinds with constricted gaster, one of intermediate-to- large size corresponds well to the types examined of mucronata, wagneri and goyana (Figs. 5, 8), and is nndonbtedly the form described by Donisthorpe as plaumanni. This, the commonest form in collections, is known from Kio de Janeiro and Goias State in Brazil southward into northwestern Argentina and Bolivia. Through the kindness of Dr. G. Steinbach, of the Zoologisches Museum, Humboldt Universitat, Berlin, 1 have been able to see Fibres 7-11. Acanthoponera spp., side views of petiolar node. Fig. 7, A. peruviana sp. nov., holotype worker. Fig. 8, A. nucronata worker from Beni E., Bolivia. Fig. 9, A. minor worker from Trinidad (syntype of A. spinnodis Weber). Fig. 10, A. crassa sp. nov., paratype worker from Ecuador. Fig. 11, A. sp. indet., female from Agudos, Sao Paulo, Brazil. All drawn to same scale. Roger's mucronata type. Roger called the dorsa of postpetiole and succeeding segment, "dicht runzlig punktirt," which fits the next species discussed below better than it does this one. There is some size variation among different samples : wagneri type worker and three workers from Rurrenabaque, Beni, Bolivia (Mann leg.) average rather smaller (HW without eyes, 1.37-1.44 BROWN : ANT TRIBE ECTATOMMINI 193 mm.) than those from the vicinity of Rio, where HW may reach 1.63 mm. in the worker. Cephalic indices are much the same in all of these samples, ranging from 90-94. Donisthorpe cited directly only larger size as a distinction for his plaumanni, but, although he stated that he had checked previous descriptions, it seems that he did not realize that there was no significant dif- ference between his measurement and the one given originally bj- Roger. The plaumanni type is thus probably only an average specimen of mucronata. The third kind of Acanthoponera with constricted gaster is near the previous kind (mucronata) in size; its HW without eyes is about 1.55 mm., and its cephalic index is similar (93), but its gaster is even broader and more depressed, particularly the sec- ond segment (Fig. 6). The second segment is also distinctive in having a semicircular impression extending from the impressed band along the posterior border forward into the raised portion of the tergite (Fig. 6). The petiole is of a particular conforma- tion ( Fig. 7 ) , and the gastric dorsum is decidedly more coarsely, closely and irregularly punctured than in the preceding forms, perhaps conforming to Roger's "dicht runzlig punktirt, " based, of course, on a female. We know that the female of mucronata has gastric punctation much like that of its worker, so the sculptural character is not just a caste or allometric difference. I have seen only a single worker of this last kind, from Tingo-Maria, Peru, but this is so distinct from the previously described species, in- cluding the closely related mucronata, that I have little hesita- tion in describing it as a new species, A. peruviana sp. nov. [1]. There is one named form of the genus that we have yet to consider: A. goeldii Forel. Dr. Wilson has examined the A. goeldii type and has compared it directly with A. minor. He feels that the two are different species, although their size is about the same. His quick sketch of the profile of alitrunk, petiole and gastric dorsum as seen from the side, and his description of the coarse sculpture and pilosity, particularly the rough, con- tiguous punctation of the gastric dorsum, are reminiscent of these features in peruviana, just discussed above, although the latter would of course be much larger than goeldii. At present, we know nothing directly of the size variation of either peruviana 194 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY or goeldii, but judging from the size variation known for mucro- nata, minor and crassa, it does not seem likely that peruviana and goeldii are large and small forms of one and the same species. The possible relationship of these two species should, however, be kept in mind in the future. To summarize the present and provisional species-level taxon- omy of Aca.yithoponera, we may recognize five reasonably distinct species at present named : mncronata, peruviana, minor, crassa and goeldii. A stray female seen cannot be referred with safety to any described species. AcANTHOPONERA species (T) crassa Brown, sp. nov. C. America, Ecuador [2, Fig. 10] (T) goeldii Forel, 1912b: 34. n. status c. Brazil (P) minor Forel, 1899a :9. n. status s. Mexico to Triuidad [2, Figs. 4, 9] (T) —schwarzi Wheeler, 1923b: 188. n. syn. (T) ^ s^nMmodis Weber, 1939:97. ii. syn. (T) mucronata (Roger), 1860:299. s. Brazil, n. Argentina, Bolivia [1, Figs. 5, 8] (T) = wagneri Santschi, 1921 :84. n. syn. (P) = plaumanni Donisthorpe, 1938:140. n. syn. (T) ^ (701/ona Borgnieier, 1939:418. n. syn. (T) peruviana Brown, sp. nov. e. Peru [1, Figs. 6, 7] Heteroponera Mayr ^ Heteroponera Mayr, 1887:532. Type: Heteroponera carinifrons Mayr, 1887, monobasic. < Acanthoponera Mayr et auct. > Paranoinopone Wheeler, 1915b: 117. Type: Paranomopone reUcta Wheeler 1915, monobasic, n. syn. ^Acanthoponera subgenus Anacantlioponera Wheeler, 1923b: 176. Type: Ponera dolo Roger, 1860, by original designation. Synonymy by Brown, 1952c. > Heteroponera (reinstated), Brown, 1952c. See discussion of synonymy under Acanthoponera, above. Wheeler's Paranomopone was set up for the sole aberrant species relicta, segregated chiefly because of its well-marked scrobes and its angulate humeri. But other Heteroponera show lesser development of the scrobes, varying with the species, so that the difference is reduced to one of degree. Angulate humeri are found also in H. leae. BROWN : ANT TRIBE ECTATOMMINI 195 Essential characters of the workers are given in the generic key. IMost known females of Heteroponera {hroinii, carinifrons, dolo, imheUis, rclicta) are wingless and ergatoid, but those of dentinodis and schwebeli have normal wings before nuptial flight. The few known males of Heteroponera have the u.sual gastric shape of other ectatommines, with no excavation beneath. The relicta worker palpi are segmented 4, 3 ; hrouni and imhellis both have 3, 3 (my dissections) ; Kusnezov (1954) claims that the worker of H. dolo has 3, 2. In general, the species of the genus are small in size and form small colonies in soil, rotten wood or epiphyte masses. The distribution is discontinuous, with four species in Australia and New Zealand and six or seven in the New World tropics south into Chile. Figures 12 and 13. Heteroponera inca, sp. nov., paratype worker. Fig. 12, full-face view of head. Fig. 13, side view of petiolar node and adjacent structures. Heteroponera species (T) hrouni (Forel), 1892:335. n. comb. N. Zealand: North I. [5] (T) =fcirfci (Wheeler), 1923b: 184. n. syn. (P) fan?i!/ro7i.s Mayr, 1887:533. Chile [3] (P) denfijiodis (Mayr), 1887:541. n. comb. s. Brazil, Bolivia [3] (P) do?o (Eoger), 1860:293. n. comb. se. Brazil to n. Argentina [3] (P) =aurea (Forel), 1913c: 203. n. sjti. (P) f?nbe??!s (Emery), 1895b :346. n. comb. sw. and e. Australia [4] 196 BULLETIN : MUSEUM OF GOMPAEATIVE ZOOLOGY (P) —hilar is (Forel), 1895 -.421. n. syn. (T) =scai>ra (Wheeler), 1923b: 181. n. syn. (T) =occidentalis (Clark), 1926 -AT. n. syn. (P) =ni^ra (Clark), 1930:6. n. syn. (T) inca Brown, sp. uov. sw. Colombia [3, Figs. 12, 13] (?) mermfs (Emery), 1894:143. n. comb. se. Brazil [3] (T) Zeae (Wheeler), 1923b: 181. n. comb. N.S.Wales (T) mierops Borgmeier, 1957:112. se. Brazil, Colombia [3] (?) panamensis (¥ore\), 1899a -.9. n. comb. Panama [3] (T) relicta (Wheeler), 19ir)b:118. n. comb. n. Queensland (T) schwebeli (huedevwaldt), 1918:54:. n. comb. se. Brazil [3] Note : Among the new combinations, relicta is transferred from Paranomopone, and the rest ironi Acantlioponcra. Santsehi's Acanthoponera (Anacanthoponera) reichenspergeri is now put in Gnamptogenys [74]. Heteroponera niinuta Kusnezov (1954:34) is a nomen nudum so far as I can determine at present. Key to known Australian-New Zealand species of Heteroponera Mayr — workers 1. Petiole tapering dorsally to an acute upwardly-directed tooth (e. N. S. Wales) leae (Wheeler) Petiole rounded or subtruncate above, without an acute tootli at the summit 2 2. Humeri angulate as seen from above; antennal scrobes distinct and deep, each surrounded by a fine, sharp carina and divided for much of its length by another (u. Queensland, rainforest) relicta (Wheeler) Humeri evenly and gently rounded as seen from above; antennal scrobes at most shallow, not distinctly bounded or divided 3 3. Petiolar node seen in lateral view profile fully erect, both anterior and posterior faces sloping very slightly inward toward the apical face, which is horizontal (e., s., sw. Australia) imbellis (Emery) Petiolar node seen in lateral view profile slightly inclined posteriad; the anterior nodal face sloping posteriad, apical face sloping upwards toward the rear; posterior face feebly concave, overhung by the posterodorsal angle, which projects slightly posteriad (N. Z.) broiini (Forel) Key to known New World species of Heteroponera Mayr — workers 1. Eyes minute, with only a very few facets; body size small, color ferru- ginous yellow, petiolar node thick scale-like, unarmed above (se. Brazil, Colombia) microps Borgmeier TT^TT-cic! loTrpia Ttri+Vi momr fliG+i'n/>f ■filr^pfa 9. BROWN : ANT TRIBE ECTATOMMINI 197 2. Full adult color black or dark piceous; petiolar node unarmed above (Chile) carinifrons Mayr Full adult color testaceous to dark reddish-brown, but not black; petiolar node with or without a posterodorsal tooth-like process or angle 3. 3. Propodeal teeth well developed and acute, longer than half the distance between the centers of their bases and longer than broad at base; occi- pital angles each forming a prominent subtruncate lobe (best seen from side) (Figs. 12, 13); posterodorsal petiolar tooth present and acute; color deep reddish-brown, (sw. Colombia) inca sp. nov. Propodeal teeth reduced or obsolete, never half as long as the distance between the centers of their bases 4. 4. Large ferruginous yellow species, HL 1.20 mm. or more, with well devel- oped but blunt tooth on apex of petiolar node ; small but distinct extra (submedian) teeth on all tarsal claws (se. Brazil to n. Argentina) . . dolo (Eoger) Smaller species, HL 1.15 mm. or less; full adult color yellowish to reddish- brown; tarsal claws without distinct submedian teeth, at least on posterior legs 5. 5. Head relatively narrow (CI < 85) ; petiole in all sizes of workers ter- minating in a bluntly rounded posterodorsal border, without tooth or pointed process, petiolar node nearly or quite as long as high and as long as broad (se. Brazil) schweheli (Luederwaldt) Head broader (CI > 85) ; larger workers with a small, acute tooth on the posterodorsal petiolar border, this tooth vestigial in many smaller workers ; petiolar node compressed from front to rear, distinctly higher than long and much broader than long (s. Brazil, Bolivia) dentinodis (Mayr) Judging from the original description, Emery's inertnis would key to couplet 5, and it is probably the same as either schweheli or dentinodis. It is possible, but perhaps not likely, that the large and small workers of dentinodis represent different species. [3] H. panamensis apparently runs to couplet 5, but we do not know enough about it to kev it further. Rhytidoponera Mayr Ectatomma subgenus Ehytidoponera Mayr, 1862:731. Type: Rhytido- ponera araneoides = Ponera araneoides Le Guillou, by designation of Emery, 1911. 198 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY '^ Rhytidoponera subgenus Chalcoponera Emery, 1897a: 548. Type: Chalco- ponera mctallica z= Ponera metallica Fred. Smith, by designation of Emery, 1911. Synonymy by Brown, 1953b :2. = Rhytidoponera, Emery, 1911:36-39, including subgenera Bhytidoponera -f- Chalcoponera. 1914:397. = lihytidoponera -\- Chalcoponera, Wheeler, 1922:643-644. Clark, 1936:14- 15. 1941:71. This genus is distinguished by means of its dentiform inferior pronotal angles, each of which is situated, pointing downward, just in front of the fore coxa (Figure 3). These teeth are some- what variable in shape, and at times their apices are blunt, but the sides are usually more or less concave, so that in basic form, the w^hole tooth is acute. Viehmeyer (1912) claims that the teeth are occasionally absent from some of his New Guinea examples, but it may l)e that he merely overlooked them in specimens that had the sternal plate of the prothorax in the dropped position, masking the silhouette. In such specimens, the tooth can be seen, with difficulty, only if the alitrunk is viewed from above and to one side, obliquely. Out of several thousand specimens of this genus, representing nearly all of the species from all over the range, I have seen only two specimens in which the inferior pronotal angles w^ere absent or so reduced as to hardly deserve the term "tooth," and in one of the two cases this condition occurred on only one side of the insect. Clearly, absence of the pronotal teeth is an extremely rare occurrence of secondary nature, and perhaps represents a pathological condi- tion. Other characters of importance in generic diagnosis : well de- veloped tooth present near midlength on all tarsal claws; pos- terior coxae unarmed above ; promesonotal suture complete, ap- parently not ankylosed ; eyes always present and well developed. Palpal segments, worker and female, appear to be constant at 3 maxillary, 2 labial, as based on dissections of several specimens each of aurata, incisa, spoliaia, taurus, impressa, metallica. Males dissected {impressa, mayri) had 5 maxillary, 3 labial. The basal maxillary segment is very short in both sexes. Wing venation of both sexes as in Ectatomma, but anal lobe of hind wing absent. BROWN : ANT TRIBE ECTATOMMINI 199 A few species in related genera (e.g. Ilcteroponera) may bear a more or less angulate projection of the lower pronotal border corresponding to that of Rhytidoponera, but never so well marked as it usually is in the latter ; furthermore, these species differ in one or more of the other characters, i.e., the posterior tarsal claws lack a distinct median tooth, or there is a tooth on the dorsal surface of the coxa, etc. In a few of the medium-sized and smaller species of Rhytidoponera, presumably closest to the generalized stock, normal winged males and females occur together in a pro- portion of nests in the same season at a given locality, and ap- parently nuptial flight and nest-founding occur much as in other generalized ponerines. In most species, however, true females become very scarce or absent, and their place is taken in whole or in part by fertile workers or by very worker-like erga- toids (Brown, 1953b; C. P. Haskins, in Hit.) ; the males are pro- duced over a wide season, which may in some cases last virtualh' the year around, and mating is accomplished when the males fly to another nest singly. Details of mating, reproduction and nest-founding in these forms remain to be clarified. Larval morphology is discussed by the Wheelers (1952a: 124- 127). Pupae are enclosed in rather dark-colored cocoons in the nests I have seen; the color approaches black in some arid-land Australian species. The male genitalia, at least in dissections of R. impressa, R. metallica and R. mayri, show only slight interspecific variation, and there appear to be no significant variations in these organs from the general plan as seen in other ectatommines. In particular, the volsella is very similar in outline to those of Ectatomma and the myrmicine genus Myrmica. The hypupygium shows differences in outline of the posterior border, l)ut these are slight and in no case approach spectacular aberra- tions such as the forked member of Faraponcra males. Rhytidoponera ranges in Australia and adjacent islands. New (luinea. New Caledonia and neighboring parts of Melanesia in the east (but misses Fiji and New Zealand) ; westward it ranges to Timor, the Moluccas, and even to the southern Philippines fas based on a single series of the widespread form araneoides, collected by J. AV. Chapman near Dumaguete, Negros Oriental). Considerable adaptive radiation has aftVcted the genus, resulting 200 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY in around 100 known, validly-named species ; the total will un- doubtedly surpass 150 as collections become more complete. One large group of forms of increased size has taken up xeric niches in Australia (e.g., mayri, punctata), and has produced rain- forest-inhabiting species in Melanesia (e.g. strigosa, laciniosa) ; both of these ecological types tend to be dark, piceous or black in color. Members of the same group living in savannah or open- woodland areas of northern Australia are often reddish or ferru- ginous in color. Perhaps the most successful and abundant life- forms are represented by the smaller species of the metallica and victoriae groups, strongest in eastern and southern Australia. These are among the most plentiful and conspicuous of Austral- ian ants in many localities, and they seem in many ways to take the place of general-feeding myrmicines, particularly Myrmica, in the northern countries ; indeed, such species as the common greenhead ant, B. metallica, apparently play a role in the Australian environment that is relatively more important than that of the commonest of Myrmica species in most parts of the Northern Hemisphere. R. metallica and R. victoriae are common even in lawns and gardens in some Australian cities, and the former is respected for the potency of its sting, which produces a dull, lasting ache out of all proportion to the size of the insect. There exists very little detailed information on foraging and feeding habits of this genus. The species vary greatly among themselves, and according to season, in diurnation of foraging. R. metallica is, in general, a predominantly daytime-foraging species, while R. mayri and other large deserticolous species forage mostly at night, at least during warm weather. Foraging is mostly limited to the ground and low plants, except in the rain-forest, where some species may normally live and forage in arboreal situations. Even in southern Australia, however, there are some species {anceps, aspera) that have been found running on tree trunks well above the ground, and perhaps most species do this to some extent. Food consists of insect remains, and probably in some cases honeydew is taken (Clark, 1936) ; what else, if anything, is gathered we do not know. Wilson will pub- lish notes elsewhere on the foraging biology of the New Guinea species. BROWN: ANT TRIBE ECTATOMMINT 201 The nests of Rhytidoponcra species are usually made in the soil, except in rain forest, where many species live in rotten logs, and a few even in the "peat" gathered in epiphytic masses on the trees. Soil nests may be with or without a stone or log as cover, and if without cover, a crater or masonry dome may be present or absent, according to species and habitat. Nests under stones are more frequent in cooler upland districts, and masonry domes with a very wide apical entrance ( often ' ' decorated ' ' with pebbles or small lateritic concretions) are characteristic of some of the larger desert species. These desert nests are externally very similar to those built by certain arid-land Camponotus species. The genus Rhytidoponera in the present broad sense has never been comprehensively revised, although Clark dealt with a ma- jority of the larger Australian species in his review of 1936. Aside from its proposal of some fairly obvious synonymy, this work is largely unsatisfactory as a revision, the basic failure being a lack of proper emphasis on the constant differences sep- arating species, where such differences exist. Apart from the inadequacies of this revision and of the material upon which it was based, however, the larger Australian species are particularly difficult taxonomieally. Many of the entities (species or super- species) have vast, but strikingly discontinuous ranges, with more or less distinctly different populations completely isolated one from the next by great stretches of inhospitable country. The related questions of just how continuous this isolation is, and whether and to what extent such populations should be formally named, require much more study than Clark gave them, or than I can give them here. Emery (1912) offered a brief review and key covering the .smaller "Chalcoponera" species that is useful, but far out of date. Some revision of this group was completed by Brown (1954b), but basic revisionary work needs to be extended to the entire Australian fauna of the genus before a workable key will be possible. Wilson [6], in his manuscript revision of the ants of Melanesia, has covered the Rhytidoponcra species of that region and con- structed a key to the species, so I shall for the most part avoid mention of Melanesian forms here. As a result of his cooperation during and after his trip to Melanesia and his subsequent tour of 202 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY the European museums, I am able to list new synonymy confirmed by him, some of which is drawn from his manuscript study. I have revised and keyed the New Caledonian species in this work At the time of his death in 1956, Clark had completed part of a manuscript dealing with the Australian Rhytidoponera and other ectatommines, but the status of this work at present is not clear. It is to be hoped that Australian and other interested specialists will soon be able to undertake a full revision of this difficult but \evy important continental fauna of the genus [8]. Rhytidoponera species Excluded from this list are names placed in synonymy by Clark (1936) and Brown (1954b) ; Ponera ruginoda Fr. Smith was placed in Rhytidoponera by Emery (1911), but is omitted here since the name was based on the male of Myrmecia pilosula Fr. Smith (Brown, 1953c). (T) atdomiRoZis Viehmeyer, 1912 :4. ii. stntus. N. Guinea [6] (P) acanthoponeroicles Viehmeyer, 1924:227. N. Caledonia [33] (P) aciculata (Fr. Smith), 1858:104. e. Australia [9, 20] (P) =ZaeOTor Stitz, 1911:352. n. syn. (T) aenesoens Emery, 1900:312. N. Guinea [6] (T) fljiceps Emery, 1898:233. e. and sw. Australia [12] (T) araneoides (Le Guillou), 1842:317. Melanesia, etc. [6] (T) =rugosa (Fr. Smith), 1859:143. syn. Donisthorpe, 1932 : 454. (T) =/ro(7<7o«iForel, 1910:10. n. syn. Wilson = arcuata Stitz, 1911 :352. n. syn. (P) = impresshiodis Stitz, 1912:498, n. syn. Wilson (P) 1= ceramensis Viehmeyer, 1914b: 112. n. syn. Wilson (P) a^pera (Roger), 1860:308. se. Australia [7] atropurpurea Emery, 1914:396. ne. N. Caledonia [33] (P) owra^a (Roger), 1861b:169. n. Australia (T) fear?!ardi Clark, 1936:54. n. Queensland barreiti Clark, 1941 :81. c. Australia (P) ftoreaZis Crawley, 1918:88. n. status. Australia: Darwin dist. [11,28] (P) = brunnea (Clark), 1941:86, n. syn. (T) mrmaia Clark, 1936:54. (T) celtinodis Wilson, ms. sp. nov. N. Guinea [6] (T) cerastes Crawley, 1925:584. nw. Australia (P) chalybaealEimevj, 1901 -.51. se. Australia [7] (T) chnoopyx Brown, sp. nov. n. Queensland [16] BROWN : ANT TRIBE EOTATOMMINI 203 (T) clarki Donisthorpe, 1943:115, uoui. pro hilli (Clark) Queensland [32] (P) — obscura Forel, 19001) :60, nee Emery, 1896. n. s.ni. (T) = hilli (Clark), 1941 :8n, nee Crawley, 1915. n. syn. (T) convej-a (Mayr), 1876:92. o. Australia [13, 19] (P) cormita Emery, 1895b :347. n. Queensland (P) crassinodis Forel, 1907:270. W. and e. Australia (P) rr!.s?a^a (Mayr), 1876:91. N. S. Wales, s. Queensland (P) croesn^ Emery, 1901:50. e. N. S. Wales, se. Queensland [7] (T) douglasi Brown, 1952b: 137, nom. pro levior Crawley sw. Australia [17] (iubia Crawley, 1915:132. Australia: Darwin dist. (T) (rre/zHfa Clark, 1936:78. C.Australia (T) /e/TH^wieo Clark, 1936:48. nw. Queensland flavicorni^ Clark, 1936:64. W. Australia flavipes (Clark), 1941:84. S. Australia ( ?) flindersi Clark, 1936:60. R. Australia [17] (T) /oreZi Crawlej-, 1918:87. Australia: Darwin dist. (T) /oi'e?oZafa Crawley, 1925:581. sw. Australia (T) fitlgens (Emery), 1883:14:8. N. Caledonia [33] (T) ^ socrifZa Emery, 1914:395. n. syn. fuliginosa Clark, 1936:47. ne. S. Australia ffreavesi Clark, 1941:81. n. Queensland grcgoryi Clark, 1936:47. ne. S. Australia (T) 7(aert:e?i Forel, 1910:15. Queensland: C.York [30] lianieli Forel, 1913b: 660. Timor (T (P (P (P (T (P (T (T (T (P (T (T (T (T (T (P (P (T hilli Crawley, 1915:131. Australia: Darwin dist. [18, 32] impressa (Mayr), 1876:92. e. Queensland [7] incisa Crawley, 1915:132. c. Australia [17] inops Emery, 1900:312. N. Guinea [6] = striata Donisthorpe, 1949b: 744. n. sjti. Wilson inorna^a Crawley, 1922:436. n. status sw. Australia [14] = carhonaria Wheeler, 1934:139. n. syn. Tcurandensis Urowa, sp.-ROv. n. Queensland [15] laciniosa Viehmeyer, 1912:5. N. Guinea [6, Fig. 3] = petiolata Viehmeyer, 1912:5. n. syn. Wilson lamellinodis Santschi, 1919b :327. n. Queensland [30] ?a!iceps Forel, 1915b: 12. n. Queensland [25] maledieta Forel, 1915b:15. n. status n. Queensland [21, 31] maniae Forel, 1900b: 57. S. Australia, w. N. S. Wales [18, 19 1 z=z spatiata Forel, 1900b :58. n. syn. mayri (Emery), 1883:150. arid s. half of Australia [20] = glabrior Forel, 1907:268. n. syn. = quadriceps Clark, 1936 :30. n. syn. = stridulator Clark, 1936 :37. n. syn. 204 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY (P) =z occidentalis Clark, 1936:39. n. syn. (P) = petiolata Clark, 1936:41, nee Viehmeyer, 1912, n. syn. (P) = dixoni Clark, 1936:46. n. syn. (T) metallica (Fr. Smith), 1858:94. Australia, except far n. [14,21] (T) =: purpurasce)isWhee\ev,1915c:805. n. syn. (T) = varians Crawley, 1922:436. n. sjti. (T) = caeciJme Vielxmeyer, 1924:227. n. syn. =: pulchra (Clark), 1941:86. n. syn. (T) micons Clark, 1936:62. W.Australia (T) mw-aMZis Clark, 1936:29. c. Australia [20] (T) TieraStitz, 1912:500. n. status N. Guinea [6] (T) = major Stitz, 1912:501. n. syn. Wilson = gagates Donisthorpe, 1941 :51. n. syn. Wilson (T) = u'fli^ieMensis Donisthorpe, 1942:703. n. syn. Wilson (?) nigra Clark, 1936:81. S. Australia [19] (T) niftda Clark, 1936:45. w. N. S. Wales (T) n.orfi/era Emery, 1895b: 348. e. N. S. Wales, se. Queensland (P) riMfZa^a (Mayr), 1876:91. Queensland (T) riMmeensisE. Andre, 1889:221. N. Caledonia [33] (T) = act/pMWcto Emery, 1914:396. n. syn. (T) peninsularis Brown, sp. nov. C. York Peninsula [27] (T) pt7o5«Zo Clark, 1936:80. w. N. S. Wales (T) pulcliella (Emery), 1883:149. N. Caledonia [33] punctata (Fr. Smith), 1858:104. S. (and W.?) Australia [17] (T) pMWcti^/ero Crawley, 1925:582. sw. Australia (?) pwnctii;en^m Forel, 1900b: 56. n. status N.S.Wales (P) purpurea (Emery), 1887:444. N. Guinea, n. Queensland [6, 7] (P) reflexa Clark, 1936:76. Australia: Darwin dist. [22] (P) re/iew.iaia Forel, 1893:458. Australia: Darwin dist. [23] (P) rotundiceps Viehmeyer, 1914a :28. n. N. Guinea [6] (P) rufescens Forel, 1900b :58. n. status e. Queensland [13] riifithorax Clark, 1941:82. n. Australia (P) rufiventris Forel, 1915b: 11. n. and W. Australia [24] (T) rw/om>ra Clark, 1936:58. sw. Australia [17] (P) sca&erTOna Emery, 1895b: 347. n. Queensland [10] (T) = malandensis Forel, 1915b: 10. n. syn. (T) scabra (Mayr), 1876:90. e. Queensland [25] socrus Forel, 1894:236. w. N. S. Wales (T) spoZiaf a Emery, 1895b: 348. n. Queensland [25] (T) strigosa Emery, 1887:444. n. status N. Guinea, etc. [&] (T) = intricata Emery, 1910:533. n. syn. Wilson (T) =otfrt'aia Stitz, 1912:499. n. syn. Wilson = sclilaginhaufeni Viehmeyer, 1912 : 4. n. syn. Wilson (T (T (T (P (P (T (T (P (T (T (T (T (T (T (T (T (T (T (T (P (T BROWN : ANT TRIBE ECTATOMMINI 205 ^ 7ute7is Donisthorpe, 19-49e:403. n. syn. Wilson suhci/anea Emery, 1897a: 548. N. Guinea, etc. [6] = transversiriiga Emery, 1910:532. n. syn. Wilson = aruana Karawajew, 1925 :78. n. syn. Wilson = icaUacei Donisthorpe, 1932:474. n. syn. Wilson tasmaniensis Hmery, 1898:232. n. status se. Australia [21] = cristulata Forel, 1900b: 59, n. syn. taurus Forel, 1910:12. n. and c. Australia [20, 28] tenuis Forel, 1900b: 58. coastal n. Queensland [27] trachypyx Browai, sp. nov. Australia: c. N. Territory [28] fwraeri Forel, 1910:14. Queensland: C.York [30] tyloxys Brown and Douglas, sp. nov. W. Australia [29, Figs. 36, 37] versicolor Brown, sp. nov. mts. of N. Caledonia [33] victoriae E. Andre, 1896:261. e. Australia [15, 31] = modesta Emery, 1895b: 348. n. syn. = scrobiculata Forel, 1900b: 59. n. syn. = cedarensis Forel, 1915b: 15. n. syn. violacea Forel, 1907:269. n. status W. Australia [13] = opacior Crawley, 1925:583. n. syn. viridis (Clark), 1941:83. ne. S. Australia [26] wilsoni Brown, sp. nov. N. Caledonia [33] yorlccnsis Forel, 1915b: 12. Queensland: C. York Paraponeea Fr. Smith =^ Paraponera Fr. Smith, 1858:110. Tj^je: Paraponera clavata = Formica elavata Fabricius, monobasic. = Paraponera, Emery, 1911:27. This monotvpic genus, exclusively neotropical, is well known and easily recognized from its giant size. Smith correctly noted the palpal segmentation ; all castes of both sexes have a 5,3 formula. The female is winged, and both it and the worker have the hypopygium bordered on each side by an upwardly-directed comb of slender spinules. The male subgenital plate is in the form of a slender, upeurved biramous fork resembling that seen in the males of cerapachyines. The hind wings of both sexes have a well-developed anal lobe. Weber (1946) has discussed certain aspects of the morphology and habits, and has called attention to the resemblances Paraponera bears to Ectatoinma. Larval morphology is covered by G. C. and J. Wheeler (1952a :117, pi. 2, tigs. 1-9). 206 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY AuLACOPOXE Arnoldi = Aulacopone Arnoldi, 1930a: 139. Type: Aulacopone relicta Arnoldi, 1930a: 140, figs. 1-5, female, monobasic. All that is known to me about the remarkable species A. relicta is contained in the original description, based on a single dealate female — still the sole specimen known. The genns, if correctly described and figured, is apparently closest to Heteroponera, but differs wddely in the structure of the fronto-clypeal area (see key to genera, above). The type locality is in intermediate moun- tain forest at Alazapin, 40 km. southwest of Lenkoran in Russian Armenia. ECTATOMMA Fr. Slllitll <^ Ectatomma Fr. Smith, 1858:102. Type: Ectatomma tuherculatum ^ Formica tuherculata Olivier, by designation of Bingham, 1903. := Ectatomma subgenus Ectatomma^ Emery, 1911:42. ^Ectatomma subgenus Ectatomma, Wheeler, 1922:643. In the Emery-Wheeler classification, Eciaiomma included the nominate and three other subgenera : Gnmnpfogenys, Ponera- cantha, and Par ectatomma. Mann (1922) and Borgmeier (1929) proposed the elevation of Gnam.ptogenys to generic rank. Santschi followed Emery and Wheeler, and even added two more subgenera in 1929, each subgenus to include a single species. These two names, Tammoteca and Commateta, never gained wide acceptance, and it seems clear that both should be placed in the synonymy of Gnamptogenys (see below). When all the species placed under Ectatomma in the sense of Emery and Wheeler are assembled, one finds that the asseml)lage is divided into two distinct groups on the basis of botli adults and larvae. Concerning the larvae, the Wheelers (1952b) say, " Emery ella, Stictoponera, Ectatomma (Poneracantha) , E. (Parectatomma) , E. {Gnamptogenys) are so similar that they can be separated only by differences of a sort that distinguish species elsewhere. It is interesting to note that Emery in the Genera Insectorum regarded Emeryella as very close to Gnamptogenys.''^ The Wheelers regard Ectatomma sensu stricto larvae as quite differ- ent from those of the genera and subgenera just listed (1952b: 657,658). BROWN : ANT TRIBE ECTATOMMINI 207 My own studies, based on adult characters, are in good agree- ment with the larval findings. I find several good differences separating Eciatomma from its erstwhile sister subgenera, and when one gets used to the idea of separation, it is even a bit surprising that these two groups of species have been associated in one genus for so long. The workers of the true Eciatomma resemble in some ways those of the larger species of Bhyiidopon- era from Australia and New Guinea, and like Rhytidoponera, they always lack a tooth on the dorsal surface of the posterior coxa. The build of the worker alitrunk is distinctive, with its convex mesonotum (Fig. 2) set off on all sides by distinct sutural grooves, and its tendency (damped in two or three species) to develop three eminences on the pronotum ; also, several of the species have evolved slit-shaped propodeal spiracles. The antennal insertions are covered by translucent bullae, very prominent and situated just within the frontal lobes, and the petiolar node is erect and more or less angular in the worker and female. The sexes have a well developed anal lobe on the hind wing, a character shared with Paraponera. All tarsal claws preserve the distinct submedian tooth. Male genitalia are much like those of Paraponera (Weber, 1946), but the hypopygium is of the normal form. The wing venation of both sexes is of the "complete" type, the forewing having all of the primitive formicid elements except the first radial crossvein. The palpi are segmented 2, 2, in worker and female ; the basal segment of the maxillary palp is broadened and .strongly- compressed, but is not very short like those of Gnampto- genys and Rhytidoponera. The apical segment of the maxillary pulp is slender. In the male, the maxillary palp is normally 5-segmented, but occasionally the terminal segment is very short and fused to the preceding segment, so that there are only four movable segments. Relative lengths of segments variable, especially the last two. Male labial palpi 3-segmented. The species of Ectatomma are mostly widespread and rela- tively successful insects, frequently common over wide areas of ^lexico and Central and South America, southward into northern Argentina. Some species iquadridens, ruidum) are more usually encountered in forested areas, while others may occur in more open, even in arid areas (opaciventre) . E. tuberculatum is 208 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY found in rather arid districts as well as in rain forest. The genus is, however, rather strictly confined to those parts of the Americas with a tropical or warm subtropical climate. The biology of two of the common species {tuherculatHm and riiidum.) is reviewed in a full and informative account b}- Weber (1916). Among his more interesting observations, one maj^ note that both he and Cook found that the eggs laid by the females became very dark in color, whereas worker-laid eggs may in some cases remain light in color, as is usual for other ants ; also, the larvae are capable of some limited locomotion l)y crawling. The species-level taxonomy of Ectatomma has been more or less confused, partly by mixups involving muticum and edenta- tum, also ruidum and morgani, and partly because of excessive accumulation of infraspecific names. Most of this confusion is rather simply dissipated when sufficient material of all the variable species is considered at one time. However, there remain a number of problems that can be settled only when more material from critical areas reaches the proper collections. Outstanding among these problems are the status of confine and aztecum, and the two north-south species pairs in South America, lugens- permagnum and edentatum-morgani. \A^eber {op. cit.) has already suggested synonymy for the variants of E. tuberculatum, and hLs suggestions are mostly formally adopted here. Ectatomma species (P) ocri^to Forel, 1909:254. n. status. Paraguay [42] (T) astecum Emery, 1901:50. s.w. Mexico [34] (?) co7ifine Mayr, 1870a: 397. "New Grauada; " C. America? [35, Fig. 2] (P) edentatum Eoger, 1863:173. s. Brazil, n. Argentina [36, 41] (P) = ins Forel, 1909:253. n. syn. (P) = densestriata Forel, 1912b: 31. n. syn. (P) ^ TO^'e?"sa Santschi, 1912b: 521. n. syn. (T) ?!i5rens Emery, 1894:144. n. status. Amazon-Orinoco basins [37] (?) viacdonagJii Forel, 1915a •.Z51. n. status, n. Argentina [36] (P) flior^'ani Forel, 1912b: 31. Amazon-Orinoco basins [38] (P) imiticum Mayr, 1870b: 962. n.e. Brazil: Ceara, etc.; Mexico? [41] (P) =lobulifera Forel, 1909:254:. n. syn. (P) opacivenlre Tioger,18Qlh:169. s. Brazil, n. Argentina [39] (P) = concoZor Santschi, 1919a:37. n. syn. (T) pervmgnum Forel, 1908:342. n. status, c. Brazil to Bolivia and n. Argentina [37] BROWN : ANT TRIBE ECTATOMMINI 209 (P) = strigosum Emery, 1894:144, nee Emery, 1887. n. syn. (P) =coH/M«arorel, 1909:266. n. syn. (P) =aereaForel, 1912b: 32. n. syn. (T) p?o?i(f?en.sBorgmeier, 1939:418. s. Brazil [40] (P) quadridcns (Fabricius), 1793:362. Panama to n. Argentina [40] (P) ruidum Eoger, 1860:306. s. Mexico to Amazon Basin [34, 38] (P) tuberculatum (Olivier), 1791 :498. Mexico to n. Argentina [42] (P) z= punctigcrurn Emery, 1890b :56. syn. after Weber, 1946. (P) = irregularis Santschi, 1921 : 83. syn. after Weber, 1946. A key to the species of Ectatomma, based primarily on the workers Note : The postpetiole and first gastric segment are one and the same. Both first and second gastric segments usually bear coarse piligeroiLs punctures in addition to the sculpture discussed in this key. See qualifications marked with an asterisk (*) at end of key. 1. Second gastric segment very finely, superficially and evenly punctulo- reticulate and opaque, the reticulation not forming striolae except pos- sibly some very fine indistinct ones near the margins or sides, away from the center of the disc 2 Second gastric segment in large part distinctly striate or striolate over the center of the disc 3 2. Fine punctulo-reticulation of first gastric segment unrelieved on the disc (but often forming fine striolation across vertical anterior face of the tergite) (n. Argentina, s. Brazil) opaciventre Roger Fine sculpture of first gastric segment generally overlain loosely by fine, irregular rugulation, transversely arched in front (Amazon Basin to Venezuela) lugens Emery* 3. First and second gastric segments, as well as head, alitrunk and petiole, very regularly and evenly striate, similarly finely throughout 4 The second gastric segment, and often also the posterior part of the first, with sculpture finer or otherwise contrasted to that of the anterior half of the first segment 5 4. Individual costulae of sculpture all with rather smooth and even sur- faces; color of full adults approaching black (Venezuela to n. Argen- tina and Bolivia) quadridens (Fabricius) Individual costulae of sculpture on head and alitrunk, especially on the 210 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY pronotum, with eroded, rough surfaces, in extreme manifestation yield- ing a beaded effect at magnifications of 80X or more; usual full adult color dull yellowish-brown, with alitrunk and gaster patchily infuscated above (se Brazil) planidens Borgmeier 5. Eye moderate in size, its greatest diameter approximately equal to, or shorter than, the length of the apical antennal segment 6 Eye larger, its greatest diameter distinctly greater than the length of the apical antennal segment 8 6. Pronotum with an obtuse, but very high and prominent median eminence and small but distinct, subrectangular lateral (humeral) tubercles Fig. 2) (Colombia, C. America, rare) confine Mayr Pronotum without, or with only a very indistinctly differentiated median eminence ; lateral tubercles or angles obsolete 7 7. Full adult color ferruginous brown, with yellowish legs and antennae; sculpture rather coarse and irregular, wavj' or vermiculate on first gastric segment (Peru to Trinidad; see text) morgani Forel* Full adult color piceous to black, legs and antennae not or scarcely lighter; transversely arched stria tion of anterior dorsum of first gastric segment rather smooth and regular, not wavy (s. Brazil, n. Argentina) edentatum Roger* 8. Node of petiole seen from side high, narrow, distinctly constricted near the middle of its height; anterior half of first gastric segment coarsely vermiculate or ruggedly reticulate-rugose (c. Mexico to Trinidad and Ecuador) ruidum Eoger Node of petiole seen from side gradually increasing in thickness from apex to base; at least the anterior half of the first gastric segment with rather regular, parallel rugulation or costation, this not or only gently wavy 9 9. Median and lateral eminences of pronotum suppressed, usually quite obsolete, or the laterals represented by brief indistinct dorsolateral mar- gination; second gastric segment superficially transversely striate, the striation spaced and partly effaced anteriorly, so that this much of the disc is distinctly shining (Ceara, etc., in ne Brazil; fsw Mexico) muticum Mayr Median and (more particularly) lateral pronotal eminences developed as more or less salient eminences or angles ; second gastric segment finely, densely and regularly striolate (except in some samples of tuberculatum, but in these, the lateral pronotal eminences are very well developed and acute) 10 BROWN : ANT TRIBE ECTATOMMINI 211 10. All surfaces of body and appendages covered with very numerous short, erect hairs; in perfect full-face view, more than 15 erect hairs projecting beyond the straight outline of the cheek between eye and anterior corner of head (sw Mexico) aztecum Emery Erect hairs present, but longer and much less abundant; in perfect full- face view, fewer than 10 hairs (often none at all) visibly projecting beyond the straight outline of the cheek between eye and anterior corner of head 11 11. In perfect full-face view, the occipital border straight to feebly convex, usually with more or less rectangular occipital angles; head coarsely reticulate-rugose, the rugae enclosing foveae with more or less shining bottoms ; color ferruginous except in N. South America, where some populations are more brownish (Mexico to n. Argentina) tuberculatum (Olivier)* In perfect full-face view, the occipital border broadly concave; occipital angles narrowly rounded ; head finely longitudinally striate, with some V-like rugules across the vertex; color deep reddish to black (c. Brazil to n. Argentina and Bolivia) permagnum Forel* *The form acrista Forel, from Paraguay and vicinity, is probably only a southern extreme geographic variant of tuhercxdatum and is not keyed. The species pairs lugens-permagna and edentatum-morgani may indicate merely the ends of respective clines of single species. In both cases, material from tlie crucial intermediate zone in the Amazon Basin is lacking or insufficient. GrNAMPTOGENYs Roger ^ Gnamptogenys Eoger, 1863:174. Type: Ectatomma {Gnamptogenys) tornatum ^= Poncra tornata Eoger, 1861, by designation of Emery, 1911. ^Ectatomma subgenus Stictoponera Mayr, 1887:539. Type: Ectatomma coxale Eoger, 1860, by designation of Bingham, 1903. n. syn. '^Ectatomma subgenus Holcoponera Mayr, 1887:540. Type: EoJcoponera striatula = Gnamptogenys striatula Mayr, 1883, by designation of Emery, 1911. n. syn. > Alfaria Emery, 1896:41. Type: Alfaria simulans Emery, 1896, monobasic, n. syn. > Ectatomma subgenus PoneracantJia Emery, 1897a: 547. Type: Ectatomma (Poneracantha) Mspinosum = Ectatomma (Holcoponera?) iispinosum Emery, 1890, monobasic, n. syn. '^ Rlwpalopone Emery, 1897a:549. Type: Rhopalopone epinotalis Emery, 1897, monobasic, n. syn. > Emeryella Forel, 1901a: 334. Type: Emeryella schmitti Forel, 1901, mono- basic, n. syn. 212 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY ^ Ectatomma subgenus Mictoponera Forel, 1901a:372. Type: Ectatomma (Mictoponera) diehli Forel, 1901, monobasic, syn. Emery, 1911. '^Ectatomma subgenus Parectatomma Emery, 1911:44. Type: Ectatomma triangulare = Ectatomma {Gnamptogenys) triangulare Mayr, 1887, by original designation, n. syn. ^ Spaniopone Wheeler and Mann, 1914:11. Type: Spaniopone haytiana Wheeler and Mann, 1914, monobasic, n. syn. > Whcelcripane Mann, 1919:282. Type: Wheeleripone alhiclava Mann, 1919, by original designation, n. syn. ^ Opisthoseyphus Mann, 1922:4. Type: Opisthoscyphus scahrosus Mann, 1922, monobasic, n. syn. ^Ectatomma subgenus Commateta Santschi, 1929c: 476. Type: Ectatomma {Parectatomma) hruchi Santschi, 1922, by original designation, mono- basic, n. syn. '^Ectatomma subgenus Tamvioteca Santschi, 1929c: 476. Type: Ectatomma concinn.um ;= Ectatomma concinna Fred. Smitli, 1858, by original desig- nation, monobasic, n. syn. '^ Emery ella subgenus Barhourella Wheeler, 1930:10. Type: EmeryeUa (Barbourella) hanlcsi Wheeler, 1930, by original designation, monobasic, n. syn. The genera and subgenera here placed in synonymy have been the subjects of long and painstaking inquiry. It was clear from the beginning of the study that the old association of Gnmnpto- (jenys with Ectatomrua (q.v.) within one genus was incorrect, as Mann (1922) and Borgmeier (1929) already had indicated by their use of Gnamptogenys as a separate generic name. It now seems clear that Ectatomma {sensu stricto) is a phyletically "lower" ectatommine genus, whereas Gnamptogenys and the synonyms listed above constitute the ''upper" ectatommines. As the present study progressed, the "upper" genera fell into four groups, as follows : Gnamptogenys Group (New World) Gnamptogenys, Pnneracontha, EmeryeUa, Parectatomma, Commateta, Tair motrca. Bar hour ell a . IIOLCOPONERA Group (New World) /{tileopatiera, Spa)iiopone. Stictoponera Group (Old World) Stirtoponei-ti, Uhapahipone, Mictoponera, Wheeleripone. BROWN : ANT TRIBE ECTATOMMINI 21:1 Alfaria Group (New World) Alfaria, Opi.sthoscyphu^. Within each group, it is uow safe to consider the genera synonymous. The justification for this synonymy is now offered for each group in turn. Gnamptogenys Group The genera and subgenera in this group have been separated primarily' on the basis of mandibular structure, and secondarily on alitruncal suturation, plus propodeal armament. The man- dibular forms represented grade through from the extremes by way of a remarkably finely stepped array of intermediate species, assembled for the first time during the course of this study. This series of species can be divided only by the drawing of arbitrary lines, with no natural gaps to help in the drawing of these lines. The other characters that have been used to divide this assemblage of species prove useless because they are not in con- cordance with the more conspicuous mandibular tendencies. Of course, it is possible to eliminate the troublesome "intermediate" species, those carrying "diagnostic" characters discordantly, bj' relegating them to monotypic subgenera, as Santschi did with Tammoteca and Commateta. But Santschi 's approach, whether cynical or extremely naive, seems justly to have been shunned by other specialists in the Formicidae. Everything considered, it makes better biological sense to drop the artificial generic assortment of these species. This done, it becomes easier to recognize and describe the very interesting phylogenetic trends in development of the mandibles within the group, considered as one radiating unit. The primitive iuan(lil)ular form apparently is the triangular type more or less cliaracteristic of tlie Parecfatomnia species, and also found more or less faithfully copied in Poneracantha and in the species of the Holcoponera, Stictoponera, and Alfaria groups. In this type, apical and basal borders are approximately perpendicular to one another (Figs. 41, 43, 44), but, contrary to the impression given hy some older writings, these two l)orders most often meet through a distinct curve, even considering that this curve can in some cases be rather sharply rounded. 214 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY In the Gnaniptogenys group, variation away from the tri- angular condition follows a double morphocline {sensu Maslin 1952) . One of the two series of variants, expressed approximately in the consecutive arrangement acideaticoxae — > hartmani — >• nigrifrons — > hruchi — >■ haenschi — ► concinna — ► alfaroi — >• tornata — >• mordax, shows a tendency toward broadening the curve of the basal angle until, in extreme cases such as ynordax, 16 Figures 14-16. Gnamptogenys spp., dorsal views of left mandibles. Fig. 14, G. scmiferox, sp. nov., paratype worker. Fig. 15, G. mediatrix sp. nov., holotype worker. Fig. 16, G. vast rata (Mayr), holotype female. All drawn to same scale. the basal and apical borders have become continuous, and the mandible has achieved a linear shape. Accompanying this change of shape, the blade of the mandible is becoming more and strongly deflected ventrad in its apical half, so that the mandibles as seen from the side are strongly arched, while the dentition transforms into a vaguely double-ranked set of blunt tubercles. BROWN : ANT TRIBE ECTATOMMINI 215 If we follow the other morphoeline outward from aculeaiicoxac in the series represented roughly hy aculeaticoxae — >• iriangu- laris — >• rastrata — >• mediatrix — > semiferox — >> Emery ella schniitti, we see the blade first lengthen without losing its tri- angular shape, and then, beginning with rastrata and mediatrix, it becomes concave along the masticatory margin and more convex along the outer margin, with further increase in length (Figs. 14-16), until an extreme falciform shape is attained in schmitti. The mandible of E. (Barhoicrella) 'banksi belongs with the group just discussed, but it is slightly aberrant in its own right ; the ditt'erence is not great enough to support the separa- tion of hanksi in a monotypic genus or subgenus. The morpho- eline that obliterates the distinction between Gnamptogcnys and Emery ella as genera also brings Barbourella into the fold. It should be noted that this morphoeline has been completed by the two new species described in this contribution — G. mediatrix and G. semiferox [78, 77]. The other character previously utilized in making subgeneric and generic splits in this group, the alitruncal suturation, con- sists in its generalized condition of a weaklj' impressed, but fully ankylosed, semicircular line marking the promesonotal suture, and a straight, deeply cut transverse line at the site of the metano- tum (metanotal groove; most aculeaticoxae — >■ schmitti morpho- eline members). In the aculeaticoxae — > mordax morphoeline, eft'acement of one or both of these lines is erratic in its expression among species and species-groups, and intermediate stages exist in all possible combinations. There seems to be no practicable means of using this suturation for the definition of groupings \vorthy of formal generic or subgeneric names, and convergence here makes even the plotting of informal species-groups a more or less arbitrary matter. In this group, there remains one aberrant species, Q. hispinosa (Emery), that has previously been assigned to a separate sub- genus, Poneracantha. G. hispinosa has very long propodeal spines, but G. aculeaticoxae has small, acute teeth in the same position, and in this and other ways, aculeaticoxae and the new species, mecotyle, are intermediate between hispinosa and the other Gnamptogcnys. There remains the somewhat different form of the head in hispinosa; however, to anticipate the findings of 216 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY synonymy among the four generic groups for which evidence is offered below, the cephalic difference cannot well stand as a generic character when such a diverse group of other species is included in Gnamptogenys. In this, I only follow the evidence of the larvae, dealt with farther below. HoLcopoNEBA Group Holcoponera has seemed like a compact and well-defined genus up to the time of this study, and I treated it as a genus in early drafts of this revision. The workers and females are similar to those of Gnamptogenys in their widespread and uniform costulate sculpture, but the workers have a more compact, dorsally convex alitrunk, crossed by a single suture (the pro- mesonotal suture) which is deeply cut, and, at least to external appearances, completely separates the alitrunk into two parts. No such comjilete suturatioii is found in Gnamptogenys sensic stricto. The node of the petiole in worker and female is erect or inclined posteriad, usually more or less cupuliform to thick disciform. Added to the seemingly clearcut alitruncal difference in the worker there has previously been stressed a character in the forewing of the male, namely, the absence of the second and third abscissae of Rs ; but this character is now known to occur in some species of undoubted Gnamptogenys as well. G. C. and J. Wheeler found the larvae of Holcoponera distinct from the related genera in that the head hairs were branched or bifurcate, whereas those of the related genera examined had simple head hairs. This character may eventually prove useful in segregating a generic group that could bear the name Holcoponera, but at the present time, its usefulness is greatly restricted by the fact that only a very few species of "higher ectatommines" have been examined in the larval stage. Of these few species, none are particularly "critical" in problems involving generic limits, and among them are none of the species that seem to be intermediate between Holcoponera and the other "higher" genera. Such species do exist, as I found to my great inconvenience at a time when I thought I had this revision just about completed, except for minor details. In speaking of the connections I have discovered between Holcoponera and other groups, it should first be emphasized that BROWN : ANT TRIBE ECTATOMMINI 217 Holcoponera is not as large a genus in number of species, races and varieties as Santschi and others have considered it to be, nor is it quite as homogeneous a group as some, including myself, have thouglit. The first steps toward this realization were made in a recent paper (Brown, 1957) in which several forms were synonymized, two new species were described, and a species formerly placed in Bhopalopone {relicta Mann) was transferred into Holcoponera. Of the two new species added, one, H. acuta, was merely somewhat bizarre in possessing a petiole toothed at the summit ; the other, H. mina, was seen to form a bridge be- tween the more "typical" Holcoponera members (specifically, H. sfrigata) and relicta, hence the generic transfer. The species relicta was not transferred from Rhopalopone just because it fitted so very badly there, but because I felt that it was a little closer to Holcoponera, especially considering the fact that all other known Bhopalopone (synonym of Stictoponera, and there- fore of Gnamptofjenys, see below) species were Indo-Melanesian in distribution. It can be seen from this that the separation of Holcoponera from Stictoponera was more a matter of geography and convenience than morphological distinctness, but so long as no further difficulties arose, it seemed best to me to preserve the old and familiar generic arrangement for practical reasons. But furtlier difficulties have arisen. The species reichenspergeri, described by Santschi in Acan- thoponera, was viewed belatedly, and proved not to belong to the genus in which Santschi had placed it. Instead, it was found to have characters allying it with Holcoponera, with the "Rhopa- lopone" group of Stictoponera, and with Spaniopone. The gastric sculpture [74] does not have the regular costulation of the other New AVorld species, and all in all, I think we must realize that reichenspergeri is the last straw to be piled on the already considerable weight of evidence against the separation of the three major generic groups centered around Gnampto- genys, Holcoponera and Stictoponera. We can add to this evi- dence farther below in discussing the characters of new species in the Stictoponera Group, but before we leave the Holcoponera Group, it seems well to discuss briefly the monotypic genus Spaniopone. 218 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Wheeler and Mann described Spaniopone Jiaytiana as a pro- ceratiine because of superficial resemblances to Proceratium, but Emery (1919, p. 107) soon pointed out the characters that allied kS. Jiaytiana with what I now call the "higher" ectatommines. Although Emery's comments were ignored by Wheeler in his 1922 classification, it is clear that Emery was correct in stating the relationships of Spaniopone. Among former Holcoponera, two species, mina and relicta, make approaches to Spayiioponc in several respects, as does the omniparent reichenspergeri. If hayfiana had been discovered in New Guinea or Java, rather than in Haiti, the chances are that it would have been placed in Rhopalopone from the beginning; certainly, there is nothing striking to separate it from the described "Rhopalopone'' we al- ready know from the Old World. Instead of a monotypic genus, it seems we have here just another moderately modified descend- ant of an old faunal element that was neither Holcoponera nor Stictoponeim, but was ancestral to both stocks — in short, we have another intergradient species. In view of the above facts and interpretations placed on them, it can be seen that we have in Holcoponera a stock that has been born and has radiated within the New World tropics, and that has attained a certain degree of adaptive success while evolving along a certain morphological pathway. There are some species, among them the common species, that we can point to and say, "There is Holcoponera." But we must not forget that there are other species of which we must say, "That could be Holcoponera, or it could l)e Stictoponera." Now, the question arises, should we recognize the new adaptive penetration that the Holcoponera stock has apparently made, and upon which its radiation is based, by granting the group generic status? Or should we emphasize instead the continuing existence of inter- gradient species, albeit frequently exceedingly rare and local ones, that still link the Holcoponera stock with other, inde- pendently radiating sister groups? I take the second choice be- cause I believe that it is the most practical one. If we separate Holcoponera by drawing an arbitrary line around it, we shall never rest easy about distinguishing, or keying, or making bio- logical statements about, the groups sundered in this way. The claim is often made that genera with many species become BROWN : ANT TRIBE ECTATOMMINI 219 "unwieldy" — although it is never stated just how one gains by splitting such a genus on arbitrary lines into two or more genera, or, worse, into subgenera. We have only to look at the generic classification of the birds, or, within the ants, at the subgeneric classification of Cmnponotus, to see where such arbitrary splitting leads. I feel that it is best to recognize now, while there are still few enough species known in the ectatommine tribe for one man to see and assimilate in one revisionary effort, that the old generic limits are mostly fading under the accession of new species, rather than the reverse. There is every reason to believe that many more species exist, unknown, and it is from the ranks of these rare and local forms that so many of our intergradient species come. We shall do the future discoverers of these forms a service if we give them a reasonably clear choice among a few large- and medium-sized genera instead of presenting them with a welter of small genera, none of which are sharply defined. And, just as important on the theoretical level, we shall be defining our genera in the way that is closest to objectivity, by stressing the gaps left when the intergrades die out. This is not, however, meant to discourage the search for new characters that may lead to new phylogenetic concepts and to new and more objective generic splits. Stictoponera Group The genus Bhopalopone, already discussed under the Holcop- onera Group, above, is clearly a key group in ectatommine classifi- cation. Due to an extraordinary series of oversights, Emery, Porel, Wheeler and Mann all missed the extra tooth on the tarsal claws of (most) RJiopalopone species, and therefore continued to classify this genus in subtribe Typhlomyrmicini (see introduc- tory section), apart from Stictoponera and Wheeleripone — the genera with which it is actually so closely allied as to be in- separable on any rigorous basis. Another character supposedly distinguishing Bhopalopone was its possession of a fairly well defined funicular club ; but this character is rendered worthless by the fact that club distinctness grades through without regard for the old generic limits between 220 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Rhopalopone and Stictoponera. Mann's diagnosis of Wheeleri- pone and Wheeler's 1922 key to the eetatommine genera grant unwarranted value to the state of the alitruncal suturation and Figures 17-22. Gnamptogenys spp., workers, side views of petioles and adjoining structures. Fig. 17, G. hiroi (Emery) from near Sogeri, Papua (ex Stictoponera). Fig. 18, G. panda (Brown) from near Muping, Sikang Prov., Avestern China (ex Stictoponera). Fig. 19, G. chapmani sp. nov., holo- type. Fig. 20, G. epinotalis (Emery) from tlie Huon Peninsula, New Guinea, Wilson No. 931 (ex Ehopalopone) . Fig. 21, G. mina (Brown), paratype (ex Holcoponera). Fig. 22, G. acuta (Brown), paratype (ex Holcoponera) . Figs. 17-20 are all drawn to the same scale; Figs. 21 and 22 drawn to a very slightly larger scale. BROWN : ANT TRIBE ECTATOMMINI 221 to the degree of unclercurvature of the g-aster; actually, these traits are graded in degree, and differ by species rather than by genera. A review of almost all the Old World species shows clearly that the characters so far employed will not separate RhopaIopo7ie, Stictoponori and Wheeleripone as generic entities. If we look at all the species of this group together, we can see better whether a division is necessary. The first thing one notes is that there are two extremes of nodal form among the species. In coxalis and relatives, the petio- lar node is low and paniform (Figs. 17, 19), while in all the former Rhopalopone species examined, the node is erect and more or less anteroposteriorly compressed, even unto the form of a thick, blunt scale (Fig. 20). In addition, the Rhopalopone species are quite small, and the Stictoponera species described up to now have been considerably larger. However, we must note the following exceptions. The species taiuanensis and panda combine Rhopalopone nodes (Fig. 18) with Stictoponera size. The new species chapmani (Fig. 19) is typically Stictoponera in form of node, but is in the Rhopalopone size range. In Wheeleripone, alhiclava would fall into Stictoponera in all characters, crenaticeps and aterrima fit best in Rhopalopone, and lucida is in between. I think this summary of the status of the diagnostic characters should serve as convincing proof of the futility of trying to separate the Stictoponera Group species as genera or subgenera. It is an e(iually difficult matter to separate these Old World forms from the New World ones. In the matter of sculpture, for instance, one finds that the new species of the Stictoponera Group from New Guinea {G. grammodes, and certain of the Rhopalopone species) approach, in the extent of their costulate or striate sculpture, the New World Gnamptogenys. Then, how does one separate Spaniopone haytiana from the smaller Rhopalopone f Or the latter from Mann's relicta. of Brazil? Or Borgmeier's Alfaria striolata (Brazil) from the Old AVorld Stictoponera? I think that the answer must be that we cannot separate these species or groups of species into genera or subgenera on the presently recognized characters. 222 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Alfaria Group As I shall show in my speeies-level treatment of this group [65], Mann's genus Oijisfhosci/phus was founded on an artist's misrepresentation of the species formerly known as Alfaria miniita Emery. It is easier to eliminate this name than to ac- count for the position of Alfaria as a genus. Of the nine species described in Alfaria, only four appear to stand at present. A. siniulans, A. hufonis and A. minuta share as a characteristic the extremely inflated second gastric segment, but A. striolaia has a less strongly developed second gastric segment, in this respect coming much closer to some species of the former genera Stictop- onera, Rhopalopone and Parectatomma. In its sculpture also, A. striolaia, is more typical of Stictoponera, and, in fact, striolaia connects the other three valid Alfaria species so broadly to Gnamptofjenys that I am forced to still another generic syn- on.vmy. This is the way generic revisions in the ants seem often to end ; when I began this section, I would have bet on Alfaria as the most firmly founded and most distinct of the ectatommine genera, and I gave up the idea of generic distinctness for the group only when I first saw Father Borgmeier's striolaia. This completes the survey of the four groups of higher ectatommine genera. I think I have made a reasonable case for the synonymy of the genera and subgenera within each group, and it also now seems plain that no clear distinction exists among the groups, at least so far as we now know. Following the evidence as I see it, I have assigned all of these generic and subgeneric names to the synonymy of Gnamptogenys. This synonymy is set down with the greatest reluctance, and with the hope that some future study based on better material may establish a clear division of the species. The species now falling in Gnamptogenys are very various, as might be expected by the former large number of genera and subgenera now included in the concept; however, there are as many features of homogeneity among these species as there are of heterogeneity. The Gnamptogenys species run for the most part to compact, heavily sclerotized forms. The size is small to medium-large, and the eyes of the workers range from minute to large. The shapes of the head and body are rather conserva- tive, and never really bizarre, although the elongate mandibles BROWN: ANT TRIBE ECTATOMMINT 223 of the schmitti and mordax groups are unusual modifications of the triangular type found in most species. No distinct scrobes are formed to receive the antennae, and the clypeus is present and distinct as a broad, convex shield extending across the front of the head over the mandibles. Frontal carinae horizontal and with distinct, horizontal lobes roofing to some extent the antennal insertions. The eyes are usually placed near or behind the middle of the sides of the head, rarely absent. Antennae 12-segmented, with or without a club at the apex of the funiculus. Palpi of worker and female segmented 3,2 or 2,2. The 3,2 formula has been found in workers or females of aculeaticoxae, alfaroi, an- mdata, hispinosa, chapmam, co7icinna, coxalis, menadensis and schmitti, although the basal segment is short and very easily missed. A few species, notably mordax and close relatives, have lost one maxillary segment (perhaps the basal and second seg- ments have fused), so that they now have a formula of 2,2. The former Holcoponera species, including relicta, porcata and forms of the striatida complex, all actuallj^ dissected, have formulae of 2,2, with the basal segment of the maxillary palp compressed and triangular, the free angle bearing a stout sensillum ; the apical segment is long, slender and weakly curved. G. minuta (formerly Alfaria yn.) has a formula of 1,2, teste Borgmeier, 1957 :in. The alitrunk is compact, and usually has a simple dorsal out- line seen in profile, either convex or straight, rarely with a deeply impressed metanotal groove. Either the promesonotal suture or the metanotal groove, or more rarely both, may be developed to varying degrees, but the promesonotal suture, even when apparently completely separating the alitrunk into two parts (as in many former Holcoponera) , seems to confer little or no flexibility upon the alitrunk. The propodeum may have a distinct declivity, or it may merge into the dorsal surface. The propodeum usually is unarmed, but small paired teeth are present in a few species, and these reach large size in hispinosa. A charac- ter of importance occurs on the dorsal face of the posterior coxae, each of these having a distinct tooth, spine or tubercle. This armament of the hind coxae is exclusive to the genus Gnampto- genys, and is lacking in only a few species scattered through different groups, so that it seems very likely that it is a primitive 224 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY trait secondarily lost in these species. The tarsal claws of each leg usually bear each a median or snbbasal tooth, which may be difficult to detect except at very high magnifications in ideal silhouette against a brightly lighted background ; in epinotalis, the extra tooth appears to be absent from the middle and hind tarsi, but is present on each fore tarsal claw. The extra tooth is well developed in the larger species, but tends to be inconspicuous in the smaller, cryptobiotic forms. The petiolar node varies in form (Figs. 17-21, 42) from low panif orm to thick disciform ; a few species have the summit produced posterodorsally as a blunt tooth (Fig. 22). The form of the gaster varies with the species group ; usually the postpetiole is separated from the succeeding (second gastric) segment by a very distinct constriction, and the latter segment is more or less distinctly undercurved. The remaining apical segments are reduced and more or less retractile within the second gastric segment. A functional sting is always present in the castes of the female sex. Sculpture in Gnaniptogenys species is based on a thick, heavy integument, so that figures can be and often are deeply cut and bizarre. The New World species are mostly covered with costulate or striate sculpture of a very regular and striking kind, whereas in the Indo-Melanesian species, striate and costulate sculpture is either al)sent or present over only limited areas of the body, and deep-set, umbilicate foveolae are often substituted. In the smaller, cryptobiotic species of both continents, the sculpture tends to become shallower and finer, and punctation may become mixed with striation or rugulation on the same surfaces. Pilosity is normally simple and fine, the hairs being of uneven lengths, erect or suberect, and usually fairly abundant but not overly conspicuous. Smaller hairs may form a pubescence of the subappressed kind over parts of the appendages or other limited areas of the body. Color of the worker and female castes varies from jet black, sometimes with contrasting light-colored appendages or sections of appendages, to testaceous in some small cryptobiotic species. The adults take a long while to develop full pigmentation, so that a full range of yellow and reddish examples may occur in the nest of a species that is definitively' black. This color problem has been the cause of considerable synonymy at the species level. BROWN : ANT TRIBE EOTATOMMINI 225 It is perhaps risky to generalize concerning the sexual castes, since these are known for only a fraction of the species ; females are known for more species than are males, since the nests are frequently small and the females therefore not difficult to find and take. Most females are either known in the winged state or from apparently normally dealate queens, but a few species, particularly among the smaller ones, have ergatoid queens with- out wings or fully developed flight sclerites in the thorax. The males are quite distinct from the female castes, and are always winged. The size is usually slightly smaller than the workers of the same nest, and the head is much modified from the female-worker plan. The eyes are very large, occupying often a major portion of the sides of the head. Mandibles usually remaining fairly well developed, triangular and w4th serial den- tition, but possibly not importantly functional. The palpi usually have more segments than in the worker, and perhaps this is invariably true. The basic formula in the male is maxillary 5, labial 3, but in some species or individuals there may be variable reduction. Thus, the fourth and fifth maxillary palp segments are fused in concinna, but are still distinguishable, and Kusnezov (1954:34) claims that the triangularis male has a formula of 4, 2. The notauli and parapsidal furrows are well developed (so far as known), and the scape is shorter, usually very much shorter, than the funiculus, appearing much the same as one of the longer funicular segments. The genitalia are retractile, or largely so, but the genital capsule is usually extruded in the available material. The parts are heavily sclerotized and rather simple, resembling the corresponding pieces of Myrinica so far as investigation has gone. The subgenital plate is unremarkable, being usually' narrowly rounded and pubescent at its apex. How- ever, very few species have been given more than a very super- ficial examination. "Wings similar in male and female, usually of the "complete" type, with all basic formicid elements in the forewing except the first radial crossvein. In some species (e.g., G. mordax, Brown and Nutting, 1950: pi. 8, fig. 5) Mf2 is completely con- tracted, and in quite a few others (Holcoponera Group), Rsf2'3 has disappeared as well. 226 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY The male sculpture is usually much reduced as compared to that of the corresponding workers, and is often largely effaced to give smooth, shining areas of wide extent, particularly over alitrunk and gaster. Otherwise, the sculpture is often a feeble and much less regular version of the corresponding worker 's. The pilosity is little different from that of the female castes, but the color is often darker. Males are often taken separately at light, and in this case, it is usually impossible to determine to what worker-based species they belong, unless previous association has been made from nest material. Furthermore, we know very little about variation in the males, either individual or geographic, and it seems from the present evidence that this sex is not going to play a very important role in the taxonomy of the group. Final judgment on this matter, however, will have to rest on closer examination of a much more nearly representative sample of the species. The larvae of species now included in Gnamptogenys are de- scribed by G. C. and J. Wheeler in their "Ant larvae of the subfamily Ponerinae" (1952a, pp. 121-124, 127-128, 132-134, pis. 3-5), and the generic affinities discussed in Part II of the same work (1952b, pp. 657-660). As limited here, Gnamptogenys is known from two areas : one in the Old World, from Ceylon and western China to Fiji; and the other in the New World, from Texas to Tucuman and Buenos Aires, including Hispaniola in the Antilles. It seems safe to consider the genus a part of the "Old Endemic" fauna of South America, present there at the beginning of the Tertiary. Some of the imperfectly known fossils of the Tertiary (Oligocene) in Europe and North America are close to Gnamptogenys, and at least one does actually belong to the genus. Like Heteroponera, the Old and New World branches of Gnamptogenys show some separate radiation, but without loss of the linking species, particularly the small " Eliopalopone" forms. We know very little about the biology of Gnamptogenys species, since many of them are rare and secretive in habits. Most of the species are restricted to moist tropical forests, where they nest mostly in rotten wood or in the soil ; a few are semi- arboreal nesters in epiphyte "peat," at least facultatively so. Usually, the colonies consist of only a few individuals, probably BROWN : ANT TRIBE EOTATOMMINI 227 in most cases under 100, and frequently under 50. The queen is usually a dealate, more rarely an ergatogyne, and some colonies may have several reproductive females. The food probably con- sists mostly of small arthropods; Mann (193-1:177-178, pi. 6) found schmitfi, a long-mandibulate species, preying on millipeds in Haiti, and it seems likely that other specialized types of mandibles in the genus reflect prey preferences as yet unknown. Gnamptogenys species Only new synonymy is listed here. Old synonyms can be found in Emery, 1911; Santschi, 1929c; Brown, 1954c, 1957. The new combinations are marked with an asterisk (*). Following the grand list, the trivial names are listed again in the genera or subgenera in which they stood prior to this publication ; the order is that of the grand list, and all names are listed, whether syn- onj-ms or not. (P (P (T (T (P (P (P (T (T (P (P (P (P (T (T (T (P (T (P (T (P (P aculcatico.rae (Santschi), 1921:81. * Panama to Bolivia [45, 73] acuminata Emery, 1896:50. Amazon-Orinoco drainage [43] acuta (Brown), 1957:491. * Bolivia, Peru [46, Fig. 22] albicJava (Mann), 1919:283. * Solomons: Ysabel I. alfaroi Emery, 1894:145. Costa Rica [50] amiulata Mayr, 1887:543. se. Brazil, Boli\na n. to C. Amer. [44] arcuata (Santschi), 1929c: 457. * se. Brazil [79] = regularis (Santschi), 1929e:457, nee Mayr, 1870. * n. syn. aterrima (Mann), 1921:411. * Fiji banJcsi (Wheeler), 1930:10. * Panama hicolor (Emery), 1889:493. * se. Asia [47] binghami (Forel), 1900a :317. * Burma, Borneo, Philippines [47] bispinosa (Emery), 1890a: 40. * Panama, Costa Rica [49] biroi (Emery), 1902b: 154. * N. Guinea [47, fig. 17] bruchi (Santschi), 1922:241. * n. Argentina bufonis (Mann), 1926:101. * s. Mexico [65] chapmani Brown, sp. nov. Philippines [58, Figs. 19, 44] concinna (Fr. Smith), 1858:103. Bolivia to C. America [57] r= romani Wheeler, 1923a: 2. n. syn. = semicircularis Borgmeier, 1929:195. n. syn. := conica Borgmeier, 1929:196. n. syn. continua Mayr, 1887:544. s. Mexico to s. Brazil [56] = panamensis Santschi, 1931:265. n. syn. costata (Emery), 1889:494. * Tenasserim to Sumatra, Philippines [47] 228 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY (T) coxalis (Roger), 1860:308. * Ceylon [47] crassicornis (Forel), 1912a:51. * Sumatra [47] (T (T (P (P (T (T (T (T (T (T (T (T (T (T (T (T (P (T (T (? (T (T (T (T (P (T (T (T (T (T (P (T (T (P (T (P (P (T (T (T orenatieeps (Mann), 1919:285. * Solomons: Ysabel I. crihrata (Emery), 1900:311. * N. Guinea [53] curtula (Emery), 1896:44, 47. * s. Mexico, C. America [70, 79] = stolli (Forel), 1899a:7. * n. syn. dammermani (Wheeler), 1924:2. * E. Indies, Philippines [53] diehli (Forel), 1901a: 372. * Borneo [53] epinotalis (Emery), 1897a :550. * N. Guinea [53, Fig. 20] exarata Emery, 1901:50. Amazon-Orinoco drainage [54] gyacilis (Santschi), 1929c: 468. * British Guiana grammodes Brown, sp. nov. Papua [60, Fig. 41] liaensc'lii Emery, 1902a:27. Bolivia, Peru, Ecuador [51] hartmani (Wheeler), 1915a: 390. * Texas [52] haytiana (Wheeler and Mann), 1914:11. * Haiti [63] liorni Santschi, 1929c: 475. n. status Panama to Bolivia [62] interrupta Mayr, 1887:543. C. America, ?S. America [55] Tcalahit Brown, sp. nov. n. Borneo [59, Fig. 43] laevior (Forel), 1905:7. * Java [47] liu'ida (Mann), 1919:285. * Solomons: Malaita I. Ivzonensis (Wheeler), 1929:30. * Philippines [53] maoretes Brown, sp. nov. Papua [61] magnifica (Santschi), 1921:81. * sw. Brazil [71] major (Emery), 1902b:lo4. * N. Guinea [53] malacnsln (Mann), 1919:281. * Solomons: Malaita I. [53] mecotylc Brown, sp. nov. Bolivia [66, Fig. 42] mediatrix Brown, sp. nov. Brazil [78, Fig. 15] menadennis (Mayr), 1887:539. * E. Indies, Philippines [47] meiioszii (Borgmeier), 1928a :32. * se. Brazil [64, 73] mina (Brown), 1957:494. * Bolivia, Peru [46, 73, Fig. 21] mlnuta (Emery), 1896:106. * S. and C. America [65] = emcryi (Forel), 1901b :326. * n. syn. ^= scahrosus (JNIann), 1922 :5. * n. syn. =: mus (Santschi), 1931:265. * n. syn. ^^ panamensis (Weber), 1940:80, nee Santschi, 1931. * n. syn. = caririala (Weber), 1940:82. * n. syn. moclJeri (Forel), 1912b:34. * se. Brazil [81] = splendens (Santschi), 1929c :450. * n. syn. viordax (Fr. Smith), 1858:98. C. and S. America [69] =z nodosa (Latreille), 1802:217. n. syn. z= purensis Forel, 1912b :33. n. syn. = schastiani^ovgmeier, 1937:220. n. syn. nigrifrons Borgmeier, 1948:199. Peru pamda (Brown), 1948a: 263. * w. China [48, Fig. 18] BROWN : ANT TRIBE ECTATOMMINI 220 (P) pleiirodon (Emery), 1896:47. * Amazon drainage [71, 72] (P) =: emeryi (Santsehi), 1929c :463, nee Forel, 1901. * n. syn. (P) =reota (Santsehi), 1929c:465. * n. syn. (P) = vidua (Santsehi), 1929e:467. * n. syn. (P) porcata (Emery), 1896:48. * Costa Eiea [71] (T) posteropsu (E. Gregg), 1951:77. * Sumatra, Philippines [47] (T) rastrata (Mayr), 1866:890. * se. Brazil [73, Fig. 16] (T) regularis Mayr, 1870b :965. s. Mexico to Paraguay [68] (P) =: splendid a Pergande, 1895:871. n. syn. (P) —fiehrigiFovel, 1909:253. n. syn. (T) reichenspergeri (Santsehi), 1929b:274. * se. Brazil [74] (T) relicta (Mann), 1916:403. * Matto Grosso [46] (P) rimwiosa (Roger), lS61a:18. se. Brazil [75] (?) rustim (Santsehi), 1929c :446. * Paraguay [79] (P) sohmitti (Forel), 1901a :334. * Haiti [83] (T) = minor (Wheeler), 1936:195, nee Forel, 1900. * n. syn. (T) schuharti (Borgmeier), 1948:198. * se. Brazil [64, 73] (T) semiferox Brown, sp. nov. Dominican Eepublic [77, Fig. 14] (P) simplex (Emery), 1896:48. * C. America [46] (?) simplicoides (Forel), 1908:341. * se. Brazil [79] (T) simulans (Emery), 1896:42. * C. America [65] (?) spiralis (Karawajew), 1925:79. * Java [47] (P) striatula Mayr, 1883:32. S. America, Jamaica, etc. [79, 46] (P) = hrasilieihsis (Emery), 1902c :181. ^ n. syn. (P) ^angustiloba (Forel), 1908:341. * u. syn. (P) = angustipleura (Forel), 1908:342. * n. syn. (P) = paulina (Forel), 1908:342. * n. syn. (T) = p 1.0 mm., WL > 1.6 mm.; superficial sculpture consisting of extremely fine, even striolation, giving a silky luster to the integument 4. 4. Alitrunk with a well-defined and distinctly impressed metanotal groove (s. Mexico) hufonis (Mann) Metanotal groove obliterated on dorsum of alitrunk (C. America) .... simiilans (Emery) .J. Postpetiole completely transversely striolate both above and below ; very small, yellowish Proceratium-Wke species with minute eyes (Haiti) . . haytiana (Wheeler and Mann) Postpetiole predominantly longitudinally costulate or striate over the dorsum, or at least with some longitudinal striation or costulation in the central or posteromedian part of the disc 6. 6. Pronotum separated from remainder of alitrunk by a very distinct suture, completely breaking the sculpture .7. Promesonotal suture absent across the dorsum, or indicated by an impressed line that does not break the sculpture 18 7. Alitrunk with sculpture on dorsum partly replaced by extensive smooth, shining areas 8. Dorsum of alitrunk sculptured (usually costulate or striate) throughout 9. 8. Postpetiolar dorsum (and that of succeeding segment) coarsely and regularly costulate in a longitudinal direction; color (of types) red- brown (w. Brazil) relicta (Mann) Postpetiolar dorsum (and that of succeeding segment) with irregular striation arching over a shining, obscurely longitudinally striate postero- median area; color deep piceous to Ijlack (se. Brazil) . . reichenspergeri (Santschi) 232 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 9. Compound eye small, its maximum diameter < the greatest thickness of the apical funicular segment ; postpetiole truncate at base, the smooth anterior face separated from the costulate dorsal surface by a rim or margin (Fig. 21) (Bolivia, Peru) viina (Brown) Compound eye larger, its maximum diameter equalling or exceeding the greatest thickness of the apical funicular segment; postpetiole with anterior face costulate and curving directly into dorsal surface (Fig. 22) 10. 10. Petiolar node terminating in a distinct flattened posterapical tooth or process, grossly acute, especially in lateral view; ventral process of petiole triangular, tapering to a narrow, slightly recurved lobe or tooth (Fig. 22) (Bolivia, Peru) acuta (Brown) Petiolar node not terminating in a distinct posterapical tooth or process, though the posterior apex may project slightly backwards; ventral process of petiole differently shaped (not as in Fig. 22) 11. 11. Ventral surface of postpetiole predominantly transversely costulate (C. America) simplex (Emery) Ventral surface of postpetiole predominantly longitudinally or obliquely costulate 12. 12. Funicular segment I (basal segment) as long as or slightly longer than II and III taken together; small species (s, Mexico, C. America) . , strigata (Norton) Funicular segments II and III relatively longer (II particularly) ; II and III together longer than I 13. 13. Larger species, with long antennae, scape length (excluding basal neck) normally > 1.08 mm 14. Smaller species, with shorter antennae, scape length normally •< 1.08 mm 17. 14. Petiolar node moi'e or less tilted posteriad in side view, the postero- dorsal angle sharply rounded and slightly overhanging the posterior face; ventral process triangular or rounded, unilobed 15. Petiolar node erect, in side view the posterodorsal angle not very much more sharply rounded than anterodorsal angle, and not overhanging posterior face ; ventral process bilobed, or at least tending to form a posterior heel or corner 16. 15. Sculpture fine, more than 30 costulae between eyes; anterior % to ^/^ of ])ostpetiolar dorsum transversely-arched costulate ; head narrow, espe- cially across occiput, occipital angle as seen from side sharply rounded, subauriculate (Guianas) gracilis (Santschi) Sculpture coarser, less than 30 costulae between eyes; at most a very BROWN : ANT TRIBE ECTATOMMINI 233 narrow band of transverse costulae along the anterior margin of post- petiole; head broader, occipital angle as seen from side gently rounded, not auriculate (Costa Eica) porcata (Emery) (sw. Brazil) m-agnifica (Santschi) 16. Antennae (and individual segments) more slender, the penultimate seg- ment distinctly longer than broad (se. Brazil) moelleri (Forel) Antennae (and individual segments) thicker, the penultimate segment as thick as or thicker than long (Amazon-Orinoco drainage) teff ens-is (Santschi) 17. Petiolar node and ventral process usually more or less as described in first lug of couplet 14, ventral process usually a tapered triangle (Panama to Bolivia) pleurodon (Emery) wasmamni (Santschi) Petiolar node usually as described in second lug of couplet 14: ventral process bilobed, square-cut, or with a single rounded lobe group of striatula (Emery) (The striatula group includes six nominal species: arciiata, curtiila, rustica, simplicoides, striatula and whceJeri, of which perhaps two to four are good species. One extreme of variation is "typical" striatula, with fine costulation and a square-cut or emarginate ventral petiolar process [S. America, Antilles] ; another fairly well-marked form is curtnla, with posterior lobe of ventral process reduced to a broad con- vexity [s. Mexico, C. America]. These and other forms are linked by iiitergrades ; apparently character displacement is involved.) 18. Mandibles elongate and falcate or subfalcate, their inner margins in large part very distinctly concave, only their apical portions meeting or crossing when mandibles are closed (Figs. 14, 15) 19. Mandibles triangular, with distinct apical and basal angles separated by a rounded basal angle, and the apical margin straight or very feebly concave (Fig. 16), or else the apical and basal margins continuous with each other in one broad convexity bearing low, vaguely double-ranked tubercles 22. 19. Mandibles exceedingly long and slender, each with an exposed straight- line length > length of head proper (HL) (Haiti) . . .schmitti (Forel) Mandibles shorter and relatively broader, straightline length of each mandible < HL 20. 20. Inner borders of mandibles each with only a short triangular tooth to represent the basal angle (Panama) hanlcsi (Wheeler) Inner borders of mandibles each retaining the basal angle as a low, broadly rounded lobe or flange lying basad of midlength (Figs. 14, 15) 21. 234 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 21. Mandibles longer (Fig. 14) ; propodeum unarmed, its declivity trans- versely costulate; sternum of postpetiole predominantly smooth and shining (Dominican Eepublic) semiferox sp. nov. Mandibles shorter (Fig. 15) ; propodeum with a pair of short teeth, its declivity longitudinally costulate; sternum of postpetiole transversely to obliquely costulate throughout (se. Brazil, Para) . mediatrix sp. nov. 22. When head is seen in perfect full-face (dorsal) view, and the antenna! scapes are laid back as straight as possible from their insertions, they either fail to reach, or just barely reach, but do not distinctly surpass, the part of the occipital border nearest to which they fall 23. In same view, the antennal scapes when laid straight back from their insertions do distinctly surpass the posterior occipital border nearest to where they fall 33. 23. Dorsal surfaces of mandibles distinctly and continuously striate over the basal third or more of their length 24. Dorsal surfaces of mandibles smooth and shining to the base, though sometimes with scattered punctures 25. 24. Larger species, head width across eyes > 1.2 mm.; mandibular stria- tion extending from insertions to apices (Bolivia, Ecuador) haensohi Emery Smaller species, head width across eyes < 1.0 mm.; apical half of mandibles smooth and shining (Texas) hartmani (Wheeler) 25. Dorsal face of propodeum predominantly or wholly transversely striate, although the anteriormost portion may sometimes bear sharply diverging oblique striation 26. Dorsal face of propodeum predominantly longitudinally costulate or striate, continuing the longitudinal promesonotal sculpture straight back to at least the propodeal midlength, and usually beyond 27. 26. Large, long-headed species, head length (HL) > 1.7 mm.; when head is seen in perfect full-face view, occipital border evenly convex; blunt but distinct vestiges of paired teeth on the propodeal declivity (Costa Rica) alfaroi Emery Smaller, more short-headed species, HL < 1.5 mm. ; when head is seen in perfect full-face view, occipital border straight or feebly concave in the middle; propodeum absolutely unarmed (C. America to s. Brazil and Bolivia) annulata Mayr 27. Anterior margin of narrow anterior apron of clypeus (not to be con- fused with underlying labral outline) distinctly concave in the middle, and the lateral corners rounded 28. Anterior margin of anterior apron of clypeus straight, or if shallowly concave, then terminating laterally in sharply rectangular corners . . .31. BROWN : ANT TRIBE EOTATOMMINI 235 28. Compound eye of worker large, its greatest diameter > the maximum width of antennal scape 29. Compound eye of worker small, usually consisting of only about 6-15 pigmented facets; greatest diameter less than, or at least not exceed- ing, maximum width of antennal scape 30. 29. Larger species, size and sculpture very variable, width of head across and including compound eyes > 1.0 mm. (C. America, tropical S. America) mordax (Fr. Smith) Smaller species, width of head across and including eyes < 1.0 mm. (Peru, Guianas) exarata Emery 30. Medium-sized species, width of head across and including compound eyes > 0.7 mm. (C. America) interrupta Mayr Very small species, width of head across eyes < 0.7 mm. (s. Mexico to se. Brazil, Jamaica) continua Mayr 31. Metanotal groove distinct and impressed, visible in all lights ; sciilpture fine, 38 or more costulae between compound eyes, 19 or more between frontal carinae (se. Brazil) rimidosa (Eoger) Metanotal groove absent or very indistinct, at best visible only in certain lights and views; sculpture coarser, < 38 costulae between eyes, < 19 between frontal carinae 32. 32. Larger species (usually HW > 0.73, WL > 1.25 mm.) ; petiolar node longitudinally costulate, at most only the lower part of the anterior face transversely costulate (s. Mexico to Paraguay) regularis Mayr Smaller species (usually HW < 0.73, WL < 1.25 mm.) ; petiolar node with costulae arching transversely across entire anterior face, from top to bottom (Panama, Trinidad, Amazon-Orinoco drainage to Bolivia) Iwrni Santschi 33. Large species, head width without eyes > 2.0 mm. ; color usually fer- ruginous ; sculpture fine ; petiolar node produced to a blunt posterapical point or tooth (C. into tropical S. America) concinna (Fr. Smith) Small and medium species, head width without eyes < 2.0 mm 34. 34. Postpetiolar tergum with parallel chains of coarse punctures separated by irregular rugulae arranged in a pattern arching over a postero- median area of finer longitudinal costulation ; succeeding segment strongly recurved and longitudinally costulate; eyes reduced (se. Brazil) striolata (Borgmeier) Tergum of postpetiole (and of succeeding segment) evenly costulate, nearly always in a longitudinal direction; second postpetiolar segment more weakly downcurved 35. 236 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 35. Mandibles predominantly smooth and shining, with scattered punctures ; striae or costulae, if present, restricted to basal third or less of length; transverse dorsal sutures of alitrunk obliterated, except in G. nigri- frons 36. Mandibles predominantly longitudinally costulate or striate ; metanotal groove represented by a distinct narrow impressed line 41. 36. Size larger, head width Avithout eyes > 1.3 mm. (Amazon-Orinoco drainage to Ecuador) tortuolosa (Fr. Smith) Size smaller, head width without eyes < 1.3 mm 37. 37. Anterior clypeal apron with sharply angular free corners; mandibles sublinear, sharply downcurved, their basal and apical margins meeting through a single continuous convexity; petiolar node longer than broad to (rarely) very slightly broader than long 38. Anterior clypeal apron with rounded free corners; mandibles more or less triangular, the basal and apical margins distinct and separated by a distinct but rounded basal angle; petiolar node always distinctly broader than long 40. 38. Node of petiole seen from the side ending in a blunt point distinctly overhanging the posterior face (Amazon-Orinoco drainage) acuminata Emery Node of petiole seen from the side without a distinctly projecting posterodorsal point 39. 39. Head in back of antennal insertions black or nearly so ; mandibles straw yellow, contrasting sharply wdth black of cranium ; alitrunk, node and gaster varying from black to yellowish-ferruginous; costulae of pro- pddeal declivity longitudinal (C. and tropical S. America) sulcata (Fr. Smith) Head yellowish to brown in color; mandibles dull yellow to brownish, usually lighter than cranial color, but not markedly contrasting with it; head and rest of body nearly concolorous ; many workers from Central America and Mexico with costulae transverse across propodeal declivity (s. Mexico to Colombia) iornata (Roger) (G. acuminata, G. sulcata and G. tornata are often difficult to distin- guish, and further material could well show them to be just variants of a single plastic species.) 40. Transverse sutural lines effaced on alitrunk dorsum, so that the surface forms one unbroken, verj^ gentle convexity from pronotum to propodeal declivity; propodeal angles evenly rounded as seen from the side (n. Argentina) bruchi (Santschi) Metanotal groove impressed and distinct, separating propodeal dorsum into a convexity distinct from promesonotum ; propodeal angles salient and more nearly rectangular as seen from side (Peru) nigrifrons Borgmeier BROWX : ANT TRIBE ECTATOMMINI 237 41. Petiolar node markedly longer than broad (Fig. 42) ; anterior pro- notum covered by a broad band of transversely arching costulae (Bolivia) mecotyle sp. nov. Petiolar node broader than long, as broad as long, or very slightly longer than Inoad ; costulae longitudinal over the full pronotal length, only occasionally with one or two obscure transverse costulae at the junction of pronotum and cervix 42. 42. Size smaller, head width without compound eyes < 1.0 mm. (se. Brazil) rastrata (Mayr) Hrigona Emery Size larger, head width without compound eyes > 1.0 mm 43. 43. Sculpture finer, usually > 26 costulae between compound eyes; size smaller, head width without eyes 1.25 mm. or less 44. Sculpture coarse, usually 26 or less costulae between compound eyes; size larger, head width without eyes > 1.25 mm 45. 44. Sculpture finer, petiolar node with 13-16 transverse costulae visible from above; node usually wider than long (s. Brazil to n. Argentina) triangularis Mayr Sculpture coarser, petiolar node with 11-13 transverse costulae visible from above; node usually not distinctly broader than long (Panama to Bolivia) acnleaticoxae (Santschi) (G. triangularis and G. acnleaticoxae have mutually exclusive ranges so far as known ; they are both variable in the diagnostic characters, and may possibly be mere geographical variants of a single species.) 4"). "S'entral process of petiole with a blunt or rounded anterior free angle and a salient, dentiform or subdentiform posterior free angle (Rio Grande do Sul, Parana) menozzii (Borgmeier) Ventral process of petiole with only the rounded anterior free angle or lobe developed ; posterior angle absent or reduced to a simple low, broad convexity (S. Paulo, Eio de Janeiro) schuharti (Borgmeier) (G. menozzii and G. schuharti may be part of the geographical varia- tion of a single species.) Key to Old World species of Gnamptogenys — workers 1. Second postpetiolar (second gastric) segment with tergum predomi- nantly (usually longitudinally) costate or striate, this sculpture extend- ing over the basal third or more of the exposed portion of the segment. 2. Second postpetiolar segment smooth, punctate, or otherwise sculptured ; remnants of costation or striation, if present, peripherally distributed, and if found at base of segment, not extending beyond basal quarter . 7. 238 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 2. Compound eyes placed within the posterior quarter of the sides of the head, nearly at the posterior angles (Sumatra, Mindanao) postero2)sis (R. Gregg) Centers of compound eyes placed anterior to posterior quarter of sides of head 3. 3. Eyes small, each composed of 20 or less facets; petiolar node antero- posteriorly compressed, in the form of a thick scale, rounded above; body size smaller, head width (without eyes) < 0.80 mm. (N. Guinea) . . major (Emery) Eyes larger, each composed of many more than 20 facets ; petiolar node low, rounded, paniform; body size larger, head width (without eyes) > 0.80 mm 4. 4. Eyes very large, occupying fully % length of sides of head, their great- est diameter approximately = combined length of penultimate and ante- penultimate antennal segments (N. Guinea) biroi (Emery) Eyes not so large, occupying distinctly < % length of sides of head, their greatest diameter distinctly < combined length of penultimate and antepenultimate antennal segments 5. 5. Occipital angles as seen from the side each produced as a small, simple, narrowly rounded, lamellate lobe or margin, much narrower than com- pound eye ; petiolar node distinctly transversely costulate its length ; second postpetiolar tergum longitudinally finely costulate (or striate) over basal half to third, remainder smooth and shining (N. Guinea) . . . macretes sp. nov. Occipital angle as seen from the side produced as a large, more or less fan-like lobe or "ear" with a narrow translucent margin, usually angled at both ends, lobe as wide as or wider than compound eye ; petiolar node not transversely costulate, or with a few costules near posterior end ; second postpetiolar tergum coarsely longitudinally costate (or fluted) for nearly its entire length 6. 6. Sculpture finer, denser and more opaque, with noticeable development of longitudinal striation or costulation between punctures over head and alitrunk ; costulae of postpetiole fine, close and distinct ; funiculi slender, its segments III through VI longer than thick (Ceylon) .... coxalis (Roger) Sculpture coarser, striation absent or suppressed in favor of the coarse f oveolae over head and alitrunk ; costulae of postpetiole coarse and often indistinct, sometimes consisting merely of elongate welts between foveo- lae; funicular segments III through VI as thick as or thicker than long (Sumatra, Tenasserim to Mindanao) costata (Emery) BROWN: ANT TRIBE EOTATOMMINl 239 7. Postpetiole coarsely and very distinctly striate over tergum, intercostular punctures reduced in center of disc, larger on sides (N. Guinea) grammodes sp. nov. Postpetiole smooth, or punctate, or indefinitely rugulose ; striation, if present, fine, indistinct and interrupted, subordinate to coarser punc- tures or other sculpture 8. 8. Length of antennal scape (excluding basal neck) 0.87 mm. or more . . .9. Length of antennal scape (excluding basal neck) < 0.87 mm 14. 9. Petiolar node anteroposteriorly compressed, excluding ventral process, higher than long, with steep, high anterior and posterior faces (Fig. 18) 10. Petiolar node low, paniform, gently rounded above, longer than high, excluding ventral process (Fig. 19) 11. 10. Gastric dorsum with fine, irregular interpunctural sculpture developed (rendering the surface more opaque) and reaching even to the center of the second postpetiolar tergum; color red-brown; posterior occipital corner as seen from the side terminating in a distinct (but variable) point or lobe (w. China) panda (Brown) Gastric dorsum with interpunctural spaces smoother, more shining, espe- cially the disc of the second tergum; color blackish-brown; posterior occipital corner as seen from the side evenly rounded, without lobe or point (Formosa) taivanensis (Wheeler) 11. Alitrunk almost completely smooth and shining, with scattered incon- spicuous punctures ; mesonotum coarsely and evenly longitudinally cos- tate; four apical antennomeres ivory white, contrasting with rest of funiculus, which is ferruginous in color (Solomons) . alhiclava (Mann) Alitrunk coarsely foveolate, including mesonotum; antennal club not white 12. 12. Greatest diameter of eye subequal to greatest thickness of apical anten- nomere; body usually more or less concolorous brownish-red (Burma, Sumatra to Philippines) binghami (Forel) (G. crassicornis Forel and G. spiralis Karawajew probably would run to binghami, but in these, the eye is placed in front of the middle of the side of the head. They may both be only variants of binghami, in which eye position varies.) Greatest diameter of eye markedly > greatest thickness of apical antennomere; full adult color usually blackish or bicolored 13. 13. Full adult color piceous to black over body; a narrow median strip on mesonotum smooth and shining (Sumatra to Celebes and Philippines) . menadensis (Mayr) 240 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Full adult color with gaster (and often the head) piceous, alitrunk much lighter, orange-ferruginous ; median strip of mesonotum finely, indefi- nitely, longitudinally rugulose, more or less opaque (se. Asia to Java) bicolor (Emery) 14. Alitrunk smooth and shining dorsally (when clean), with separated punctures or f oveolae 15. Alitrunk dorsally in large part (at least the pronotum) densely sculp- tured and more or less opaque, with or without definite punctures or f oveolae 19. 15. Head with straight, parallel sides and rectangular occipital corners; eyes large and only gently convex, occupying aljout % length of sides of head (Figs. 43, 44) ; antennal scapes short and thick, surpassing occipital margin by much less than their apical thickness 16. Head with sides more or less convex, occipital corners rounded ; eyes smaller, occupying 1/5 or less of length of sides of head ; antennal scapes longer and more slender, surpassing occipital margin by their apical width or more 17. 16. Size larger, head width (without eyes) > 0.70 mm. (Java) laevior (Forel) Size smaller, head width (without eyes) < 0.70 mm. (Philippines) ... cJiapmani sp. nov. 17. Antennal funiculus bicolored, the four apical segments whitish in color, contrasting with the ferruginous color of the rest of the antenna (Solomons : Malaita I.) lucida (Mann) Antennae concolorous ferruginous 18. 18. Dorsum of head with coarse contiguous punctures, the median strip longitudinally costulate and subopaque (Fijis) aterrima (Mann) Dorsum of head with punctures mostly separated by smooth, shining intervals ; median strip almost free of punctures, smooth and shining (Solomons: Ysabel I.) crenaticeps (Mann) 19. Eyes large, occupying about y^ length of sides of head; their greatest diameter > greatest thickness of antennal scape (Fig. 43) (n. Borneo) Jcalabit sp. nov. Eyes small, occupying much < ^4 length of sides of head ; their greatest diameter < greatest thickness of antennal scape 20. 20. Second tergum of gaster (second postpetiolar segment) densely punctate, subopaque, the coarse punctures subcontiguous and separated only by vague longitudinal costulae (Solomons: Malaita I.) . malaensis (Mann) Second tergum of gaster with separated small punctures, the broad intervals smooth and shining 21. BROWN : ANT TRIBE ECTATOMMINI 241 21. A broad median area extending over the anterior half or more of the propodeum smooth and shining (N. Guinea) 22. Propodeum completely sculptured, or at most with a very narrow, in- definite smoothed area limited to the anterior third of the dorsum (E. Indies, Philippines) 23. 22. Punctures of postpetiole large and deep, relatively few in number, separated by broad shining spaces; full adult color of body deep brownish-red cribrata (Emery) Punctures of postpetiole small, superficial and densely arranged, sep- arated by narrow shining spaces, the total surface subopaque; full adult body color ferruginous yellow epmotalis (Emery) 23. Size larger, maximum pronotal width 0.3S mm. or more; full adult body color deep brownish-red, postpetiolar punctures large and deep da'mmei'7na7ii (Wheeler) Size smaller, maximum pronotal width < 0.38 mm., full adult color yellowish-ferruginous; postpetiolar punctures smaller and shallow luzonensis (Wheeler) (According to the original description, diehli (Forel) would run to couplet 23 ; Wilson has seen the type and finds it to differ from dammer- mani. Differences from luzonensis unclear.) Proceratium Roger ^ Proceratium Eoger, 1863:171. Type: Proceratium silaoeum Roger, 1863, monobasic. > Sysphingta Roger, 1863:175. Type: Sysphingta micrommata Roger, 1863, monobasic. Synonymy after Dalla Torre, 1893:18. ^ Sysphincta (!) Mayr, 1865:12, and most subsequent authors; emendation of Sysphingta Roger. '^Proceratium subgenus Sysphincta, Forel, 1913a: 212. The genera Proceratium. and Sysphincta have been relatively well known to specialists of the northern countries. Taking sep- arately the Nearctic, European and Japanese representations of these two genera, it has previously been considered possible to separate them on the Emery-Wheeler key characters, thus : Clypeus with an anterior median projection; petiole more or less nodi- form Sysphincta. Clypeus without an anterior median projection; petiole in the form of a thick erect scale Proceratium. However, it seems that the protests of Forel (1913a) against this generic distinction have been overlooked. Forel stated that 242 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY Figures 23-35. Proceratimn spp., workers, dorsal views of anteromedian part of head, all drawn to same scale. Pig. 23, P. pergandei (Missouri). Fig. 24, P. melimim (Italy). Fig. 25, P. toschii (holotype, Kenya). Figure 26. P. croceum (Alabama), detail of maxillary palpus, female. Figures 27-32. Proceratium spp., workers, side views of petioles and adjoining struc- tures, all drawn to same scale. Fig. 27, P. stictum, sp. nov. (holotype, n. Queensland). Fig. 28, P. micrommatum (Veracruz). Fig. 29, P. melinum (n. Italy). Fig. 30, P. toschii (holotype). Fig. 81, P. croceum (Alabama). Fig. 32, P. papuanrim (Malanda, n. Queensland). Fig. 33, P. relictum, female syntype (Fiji). Figures 34, 35. P. arnoldi, body, from sketches by Dr. G. Arnold, worker svntvwe CS. Ehodesia"). Fiff. 34. side view. Fisr. 35. BROWN : ANT TRIBE ECTATOMMINI 243 for both the clypeal projection and nodal form characters there existed "all possible intergrades." Since that time, the number of species described has doubled, and some of the forms added are further intermediates in the two crucial characters. Other forms show extreme development of characters of either genus; stictum (Fig. 27, 45, 46) is a ' ' su-per-Sysjjhincta/' while relictum is a ''su-per-Proceratium" (Fig. 33). It has been possible to assemble a representative series of species showing the gradual- ness of change from '^super-Sysj:)hi7icta" to "super-Procera- tiinn" in nodal form (Figs. 27-35), and also illustrating nicely the loss of the median projection of the clypeus. Figure 46 shows the broad median clypeal lobe, apparently the generalized condition, still preserved in stictum. Figures 23 to 25 show the degeneration of the lobe into a bicarinate median projection in pergandei and allies (Fig. 23), then through the stage of fusion of the carinae to a unipartite projection in the nielinum group (Fig. 24), and finally to the low vestigial flange of the African species toschii (Fig. 24) and arnoldi (Fig. 1), after which, in the remainder of the species, there is left no trace of the projec- tion. Other characters have been studied in the search for separatory features to distinguish two or more genera among the full array of species, but so far all characters appear to show gradation from one extreme to the other. In mandibular dentition, one begins with a distinctly four- dentate masticatory border in stictum and follows the gradation through a number of species to the condition where two strong apical teeth remain, followed basad by an indefinite number of low, irregular denticles or else a relatively smooth margin. In some of the Syspliincta species, the undercurving of the principal (second) gastric segment is extreme, but this character again grades over by easy steps to a less extreme condition. The same applies to the palpal segmentation, as far as known [84]. Too few larvae are yet known for these to be of any help with the classifi- cation, and this goes also for male genitalic characters. The workers of Proceratium are small to medium-small in size, generally ferruginous to reddish-brown in color, with greatly reduced eyes. The general habitus is distinctive (see Fig. 45) bocause of the under-turned gastric apex, which is a consequence 244 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY of the large, vaulted second segment of the gaster. The reduced apical segments bear a stout sting and are retractile to some extent within the second segment. The first gastric segment (postpetiole) is very variable in size with the species, and in some of the "Sysphincta" species it is reduced almost to the status of the myrmicine postpetiolar node (e.g., in P. watasei). In fact, these variations in the gastric proportions, and especially in the ratio of postpetiole to the succeeding segment, are of the type that must have led to the myrmicine gastric configuration from an ectatommine stock not far distant from Proceratium hegiiiniugs. Most of the remaining characteristics of the genus are, like the reduced eyes and depigmented coloration, connected with the cryptobiotic life led by the species. The antennae are well de- veloped, and their insertions have migrated anteromedially in order to give freer play to the funiculi straight ahead. The clypeus and "frontal" area are constricted by this migration of the antennal sockets, and the lobes of the frontal carinae are narrowed and tilted on edge so as not to hamper the movement of the scapes. The alitrunk is welded into one compact piece, with little or no trace left of the transverse dorsal sutures. The antennae are 12-segmented in worker and female, and 13-seginented in the male. The maxillary palpi are 2-, 3-, or 4-segmented in the workers and females so far dissected, while in tliese same specimens, the labial palpi are either 2- or 3-seg- mented (known palpal formulae are 2,2; 3,2 and 4,3) [84j. Sculpture in the worker and female usually consists of fine, dense reticulo-punctatioii, with sometimes a little superimposed rugulation, but in sticfum the ]ninctures are coarser and the entire sculpture rougher. The gaster is usually predominantly smooth and shining, with fine piligerous punctulation. The pilos- ity normally consists of abundant fine, short hairs, subappressed to inclined-erect, forming a kind of pile over large areas of trunk and appendages. Occasional longer fine hairs are more scattered. There is often a definite fine carina running down the middle of the cephalic dorsum, probably homologous to the similar feature of Acanthoponera and Hcteroponera. In the female, the compound eyes are much larger than the minute ones of the worker, and are similarly situated near the BROWN : ANT TRreE ECTATOMMINI 245 middle of the sides of the head. The female and male often (particularly in the North American species) have the metanotum produced into an acute median tooth or bladelike carina. The female may be either fully winged (and later dealate) as usual for ants, or she may appear in ergatoid form, inter- grading to the worker form. Both normal and ergatoid females may occur in the same nest. The wings in both sexes are a somewhat reduced version of the basic ponerine type ; in those I have seen of Proceratium, the forewing vein (Rsf2-3) formed of the second and third free abscissae of the radial sector is missing, leaving a single large polygonal cubital cell; occasionally, the distal part of this vein, equivalent to Rsf3, is retained, as in some Myrmica. The cross- vein m-cu is often lacking, too. Details of morpholog}^ are dealt Avith by Kennedy and Talbot (1939, all castes), by Kratochvil (1944, all castes) and by M. E. Smith (1943, male) for different species. The larvae, at least during later stages of development, develop large, smooth bosses that are arranged symmetrically over the body and are unlike those of any other ant genus. However, the larvae are known for only the North American species, and even then from very few specimens (G. C. and J. Wheeler, 1952a: 134-137, pi. 5, figs. 17-27). The male genitalia are figured by Kennedy and Talbot {loc. cit.), but I can find no close correspondence between my dissections of males of the same species (silaceum) and their drawings of these organs. The genitalia need to be restudied. Proceratium Ls widely distributed in the warmer half of the North Temperate Zone in areas with sufficient moisture in the soil, being found from Spain to Japan, and it ranges through southeastern Asia, the Malay Archipelago to New Guinea, reach- ing southeastern Queensland in Australia [96] and having a repre- sentative {relictum) in Fiji in the east. In eastern North Amer- ica, three species range widely in the forested areas from the (rulf of Mexico northwards, two of them reaching New England and the Great Lakes region [97, 98]. Another three species range widely in southern Mexico and Central America, and one of these has lieen taken on Cuba, to which country it may have beeu historically introduced. In addition to the North African species, 246 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY which are part of the southern Palaearctic fauna, there are two species, arnoldi (Southern Rhodesia) and toschii (Kenya), known from the African Continent. We have only fragmentary, and often contradictory, informa- tion on the biology of Proceratmni species. The nests are small, usually consisting of from 10-50 workers and one dealate female, though sometimes supernumerary queens, either dealate or erga- toid, are present in one nest. The nest consists of small rounded chambers hollowed out of soft rotten wood or in the soil ; toward the cooler limits of the range, particularly in North America, nests and forag'ing workers are found under deepset rocks instead of in rotten wood. The nest site is usually in forest shade, in old moist gardens, or similar habitats that are constantly moist. In the artificial nest, Proceratium species have been offered various foods. Fiala (in Kratochvil, 1944) states that his me- linum {^fialai) took bee honey; Haskins (1930) found that his croceum would not take honey, but that both adults and larvae would feed on the larvae, and sometimes on the pupae, of various genera of ants. Haskins also reported his croceum feeding on "meat" after long hesitation. L. G. and R. G. Wesson (1940) studied pergandei in Ohio, and noted that this species refused the brood of Camponotus and Formica as food, and also avoided all other insects offered, both living and dead. The perga7idei did, however, appear to feed on the gastric contents of several Formica and Camponotus worker adults. Observations made more recently by Wilson and myself and others (to be published elsewhere) on silaceum and pergandei indicate that these species may be specialized egg predators, eating the eggs of various other arthropods, especially of spiders. This may be the explanation also for the Wessons' observation of pergandei feeding on the gastric contents of dead worker ants, for these ants may well have been carrying eggs in their ovaries. Just how far egg predation extends as a general habit among Proceratium species is not known. Males and females are produced in small numbers, usually toward the end of the summer in the North Temperate Zone, where the nuptial flight occurs normally during the last half of August in many places. The flight takes place during after- noon {meliyium) , and both sexes climb to a distance from the nest BROWN: ANT TRIBE ECTATOMMENTI 247 entrance before taking flight. Workers issue from the nest also during tli£ nuptial flight, as is often the case with otherwise cryptobiotic ants. The flight has been observed by Fiala, (in Kratochvil, 1944) for melinum. Haskins (1930) mentions the curious habit in croceum workers of backing up to the brood of other ants to employ the sting. What, if any, function can be ascribed to this action is a matter for future observation to settle. The general activity of Procer- atium species is rather sluggish, although they can move quickly on occasion. The antennae are held porrect in front of the head and during foraging are vibrated ceaselessly, much in the man- ner of cerapachyines and dorylines. On the basis of external morphology, the species of Proceratium can be grouped conveniently in series from thick-noded C ^Us- phincta") to "super-Proceratium" with a scalelike petiole. One can see seven vague groups : Stictum Group: stictum. Pergandei Group: algiricum [85], mayri, pergandei, ivatasei. Melinum Group: itoi, melinum [94], Micrommatum Group : convexiceps, micrommatum. Arnoldi Group : arnoldi, foschii. Silaceum Group : carinifrons, crocevm., japonicum, lombokense, longigaster, mancum, normandi, numidi- cum, papuanum, silaceum. Relictum Group : relictum. Unas- signed : calif ornicum. Proceratium species The species marked with an asterisk (*) have previously been placed in Sysphincta bj' most authors, although those names not marked "n. comb." were either placed in Proceratium when originally described, or were transferred to Proceratium by some author (Mayr, Dalla Torre) subsequently. The names marked ■'n. comb." were all originally described in Sysphincta. ( ?) algiricum Forel, 1899b: 305. * n. Africa [85] (T) arnoldi Forel, 1913a:210. * S. Ehodesia [86, Figs. 1, 34, 35] (?) calif ornicum T. W. Cook, 1953 : 45. California [87] (P) carinifrons Menozzi, 1939:175. Sumatra, Philippines [88, 91] (T) co/M'e.rice^'s Borgmeier, 1957:120. Costa Rica [89] (P) croceum (Roger), 1860:288. se. N. America [Figs. 26, 31] (P) itoi (Forel), 1917:717. n. comb. * Japan [90] (P) ja^JonicMWi Santsehi, 1937:362. Japan 248 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY (T) lom'bo]censeEmevy,1897h:593. Lombok [91] longigaster Karawa jew, 1935:59. Indo-China [88] (P) manoum Mann, 1922:6. C. America, s. Mexico [92] (P) mayri Forel, 1899b: 306. * s. Europe [85] (P) melmum (Eoger), 1860:291. * s. and c. Europe [94, 84, Figs. 24, 29] (P) = europaea Forel, 1886:clxiii. * n. syn. = rossica Arnoldi, 1930b: 144. * n. syn. (T) = fialai Kratoclivil, 1944:54, 86. * n. syn. (P) micrommatum (Roger), 1863:176. * Panama, s. Mexico, Cuba [93, Fig. 28] (P) =: cavernicola Borgmeier, 1937:221. * synonymy by Borgmeier, 1957:118. (?) normandi Santsclii, 1929a: 138. n. Africa [95] (P) nwmidicum Santschi, 1912a: 172. n. Africa, Balkan Pen. [95] (T) papuanum Emery, 1897b: 592. N. Guinea, ?e. Australia [96, Fig. 32] (T) = PoJiera rrtffa Donisthorpe, 1949a : 491. n. syn. (P) pergandei (Emery), 1895a :264. n. comb. * se. N. America [97, Fig. 23] (T) reZic^Min Mann, 1921:413. Fiji. [Fig. 33] (P) si?aceMm Roger, 1863:172. se. N. America [98] =z crassicorne Emery, rugulosuin Wheeler and vestititm Emery, syn- onymy by Creighton, 1950:36-40. (T) stictum Brown sp. nov. n. Queensland [99, 84, Figs. 27, 45, 46] (T) loschii (Consani), 1951:167. n. comb. * Kenya [100, Figs. 25, 30] (P) watasei (Wheeler), 1906:303. n. comb. * Japan DiscoTHYREA Roger ^ Discothyrea Roger, 1863:176. Type: Discothyrea testacea Roger, 1863, monobasic. ^ PseudosyspJiincta Arnold, 1916:161. Type: Pseudosysphincta poweri Arnold, 1916, Ijy original designation, monobasic, n. syn. > ProdiscotJiyrea Wheeler, 1916:33. Type: Prodiscothyrea velutina Wheeler, 1916, monobasic, n. syn. ^ Pseudosphincta Wheeler, 1922:645, 762. Variant spelling of Pseudo- sysphincta. n. syn. The genus Discothyrea includes a modest number of small to minute species, very compact in form and extreme in the development of the " proceratiine traits," and especially notable for the exaggerated enlargement of the apical antennal segment. The raised median portion of the clypeus between the frontal lobes is fused with the frontal lobes and elevated with them to form, in some species, a flat-topped platform (Fig. 47, 48) BROWN: ANT TRIBE ECTATOMMENTI 249 l.etween the antenual insertions; in other species, the width of this structure is decreased, and finally, in several species, it is reduced to a simple vertical plate or flange that separates the novr very closely approximated antennal insertions. The lower part of the clypeus is reduced to a strongly projecting apron covering the middle parts of the mandibles (P"'ig. 48) and serving as the front rim of the platform upon which the antennae are inserted. The elypeal apron may be rounded or truncate as seen from above. With the enlargement of the apical antennal segment, the segments between it and the first funicular segment (pedicel) become axially compressed and tend to fuse one to another to reduce the number of segments. This fusion and reduction renders determination of segment numbers virtually impossible in some species, even when one uses cleared specimens under the higher powers of a compound microscope. Furthermore, it is clear that fusion is irregular in some species, and differs in degree among individuals, and even on different sides of the same specimen. In spite of this difficulty, the antennal segmentation has been granted value as a generic character. Discothyrea has been considered to be "normally" nine-segmented, including the scape, although actually there are a number of species in Africa, the New World, and Australia with lesser counts : eight, seven, and even as few as six antennal segments, including the scape ; for a discussion of this situation, see Borgmeier, 1954:191. Prodiscothyrea is distinguished primarily on the basis of its ten-segmented antennae ; the other characters of cephalic struc- ture cited by Wheeler as diagnostic are all found well developed in one or more nine-segmented species of Discothyrea (e.g., clavicornis, oculata, mixta), and therefore are useless as generic characters. We therefore have the anomalous situation in which one genus is distinguished by its having ten antennal segments, from another genus that has six, seven, eight, or nine. Clearly, the ten-segmented species belong in Discothyrea. But the same character does not rest its variation at ten segments, for the new species mixta [105], otherwise exceedingly similar to certain Discothyrea and Prodiscothyrea species, turns out to have eleven 250 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY segments. And there is Pseudosysphincta poweri to be consid- ered ; essentially, this last is a Discothyrca, but with twelve antennal segments. Now that we have an unbroken series of species with all possible segmentations from six to twelve, it is plainly not advantageous to recognize more than one genus for this group, at least until some better character can be found on which to base the separation. Except for the segmentation, Discothyrca in this new, collective sense is an exceptionally homogeneous and well-marked genus. Some other characters of the Discothyrca worker may be reviewed. Eyes small to minute, placed near or anterior to the middle of the sides of the head. Mandibles rather small, apex acute, followed by a usually edentate masticatory margin that is straight to concave and lined with a very even, close rank of minute peg-like bristles ; the masticatory border is terminated basally by a rounded or bluntly toothlike basal angle. The palpi of maxilla and labium are variously segmented, according to species; Wheeler (1916:33 and pi. 4, fig. 3) found a formula of 4,4 in vehitina, whereas Borgmeier counted 1,3 in sexarticulata (1954:193, figs. 8, 9). A comparative study of the maxillary palpi, very difficult in these small insects, remains to be made by the first reviser of the genus. (But see the recent study by Borg- meier, 1957:122-125.) The mouthparts are altogether peculiar in their reduction, and it would be interesting to know what the food is. The alitrunk is short, usually vaulted and sutureless dorsally, and terminates posteriorly in a declivity that is more or less distinctly set off from the dorsal surface by paired propodeal angles or vestigial teeth. Petiole reduced, anteroposteriorly com- pressed and not very high, usually in the form of a thick disc or scale, bluntly pointed above or with one or two low rims running around its free margin from one side to the other. The node is normally attached over a large proportion of its posterior surface to the postpetiole. Postpetiole and succeeding gastric segments are arranged much as in Proccratium (Fig. 47). Legs stout ; spur of posterior leg pectinate, spur of middle leg very small, perhaps absent in some species. Tarsal claw^s small, simple. Basic sculpture consisting of minute piligerous or tuberculate punctures, in most species crowded densely over most of the BROWX: ANT TRIBE EOTATOMMINI 251 body, rendering it opaque or subopaque ; rarely, however, the punctures are much smaller, spaced, and the surface generally shining. Usually, the sculpture tends to thin out more toward the gastric apex, so that shining interspaces appear in the species with this type of sculpture. Entire, or nearly the entire, body covered by a very dense, fine erect pubescence or pile, helping to render the surface more opaque, or else the pubescence reduced to virtually invisible pruinescence. Longer hairs are scarce or absent. Integument color ranges from testaceous to dark brown or mahogany, or combinations of these. The female is like the worker, but slightly to distinctly larger ; usually with well developed ocelli and flight sclerites of the thorax, and with larger compound eyes. Ergatoid females may occur in some species, as workerlike individuals with rather large compound eyes and a trace of one or more alitruncal sutures remaining on the dorsum. AVing venation is of the Myrmecina type (see M. R. Smith, 1943: fig. 5C), with Rs+M, Rsf2-3, and Rsf 4*5 lined up to form a single strong unit joining the anterior margin of the wing near its apex (i.e., "radial cell closed"). Mf2 is present, but not r-m, Mf3 or beyond, or m-cu. Of course, wings are known for only a few of the species so far. The male is smaller and more slender than the female, with a subglobose head, large compound eyes, small ocelli, and well developed, triangular mandibles shaped much as in the worker. Antennal insertions very close together, separated by a vertical carina representing the fused elypeus and frontal lobes in the female castes ; a more or less distinct vestige of the horizontal clypeal apron is also present. The antennae are composed of a short scape, approximately equalling in length the succeeding three segments, and twelve funicular segments, all but the first of which (pedicel) are longer than broad. The first segment of the funiculus is globular or subglobular and broader than long to slightly longer than broad, according to species. The funiculus as a whole is slender, with slight incrassation toward the apex. Mesonotum with or without notauli. Wings as in the female. Petiole as in female, but usually longer and lower. Gaster vari- able in shape and proportions, the postpetiole either larger than or smaller than the succeeding segment; apical segments only slightly deflected ventrad. Sculpture of fore part of body much 252 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY as in the corresponding workers, densely punctiilate, in the species so far known ; gaster more smooth and shining. Color dark brown or black. Genitalia as yet remaining unstudied ; hypopygial border rounded. Larva remaining undescribed. The range of Discothyrea lies predominantly within the tropics and warm temperate parts of the Southern Hemisphere, although we have records for the southeastern United States and for Hainan Island, southern China, and Formosa. The genus is represented by one (and possibly more) species on New Cale- donia, one on New Zealand, two or three in the southern Brazil- northern Argentina area, and two or so in South Africa. It is restricted to the moister nest sites, though it can be carried about by commerce in plant roots and humus, and therefore often turns up in gardens. It should be expected in greenhouses and plant propagating beds wherever the temperature and moisture are held sufficiently high. Considering the relative abundance of colonies as measured by number of collections to date, it seems that by and large, Proceratium and Discothyrea tend to have complementary distributions outside the tropics. Due to its chiefly tropical and south temperate distribution, as well as to its small size, inconspicuous nests, death-feigning habits and cryptobiotic tendencies, Discothyrea is relatively rarely collected, and little is known concerning its habits. Nests are found in rotten wood, rotting nuts, etc., in the leaf litter, and in large rotting logs. In the hilly parts of southeastern Australia, I have found them under stones rather frequently, especially in the galleries of other species of ants. Individual workers have been recovered from plant roots and similar situa- tions in the soil, and in moss covering logs and rocks in wet areas. It is probable that the colonies are more frequent than they seem to be to a collector working in the usual way to search for ants. The food is unknown ; from the aberrant nature of the mouth- parts, it must be something rather special. Several authors have thought that it "is almost a certainty that they are carniv- orous," and Moore (1940:308) gave it as his opinion that the food of Discothyrea in New Zealand might consist of mites taken in ant nests. Since the similarly specialized Proceratium is now BROWN : ANT TRIBE ECTATOMMINI 253 thought to be an arthropod egg predator, in part at least, a similar feeding' haint could possibly be followed by Discothyrea; at any rate, it is "worth checking- in the field. Colonies of Discothyrea thus far examined held ten or twenty workers plus a female and brood ; Avinged forms can be produced in equal or even greater numbers, and both sexes are normally produced in the same nest. Discothyrea species New combinations are marked with an asterisk (*). D. hryanti and D. vchitina are transferred from Prodiscothyrea, D. poweri from Pseudosysphincta. (P) antarctica Emery, 189oa:266. N. Zealand: North I. (T) ?)iV?fns Clark, 1928:38. Australia : s. c. Victoria [101] (T) hryanti (Wheeler), 1917:29. * Malaya, Java, Hainan I. (P) r/aiioorTiis Emery, 1897b: 593. Melanesia (T) cmssicorn i.s Clark, 1926:46. sw. Australia [101] (7t'n/iVM?a/« Weber, 1939:100. British Guiana [103] (P) ^rZoZ; us Forel, 1905:4. Java (?) hewitti Arnold, 1916 : 160. S. Africa [102] /ioniiMenozzi, 1927:270. Costa Eica [103] /^nofZ/.f Weber, 1939:100. Panama [103] ?V/a Weber, 1939:101. Trinidad [103] (?) tst7(mica Weber, 1940:78. Panama [103] (T) leae Clark, 1934a:29. S. Australia: Lofty Eange [101] (T) niiarfo Browni, sp. nov. Liberia [105, Figs. 47, 48] (P) neotropicaBvwQh, 1919 -AOO. [103] (P) ofM?fl/rt Emery, 1901:52. av. and c. Africa [102] (?) patrizii Weber (emended from "patrizzii"), 1949:2. [102] iwiueri (Arnold), 1916:162. * S. Ehodesia [102] (T) )Tm)?u//o«i Brown, 1948b: 38. jST. Caledonia [104J soMieri Forel, 1912:47. n. status Formosa ( ?) sculptior Santsehi, 1913 :302. n. status French Congo [102] (T) «ea-arfi>M?o/a Borgmeier, 1954:191. Brazil: S. Cat., S. Paulo [103] (P) testacea Eoger, 1863:177. se. United States [103] (P) traegaordhi Santsehi, 1914:3. w., e., s. Africa [102] , (T) fi/rfoni Clark, 1934b :53. Australia: Victoria [101] (T) velutina (Wheeler), 1916:34. * n. Queensland 254 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY APPENDIX The pages of this appendix are reserved for notes and descrip- tions dealing chiefly with species-level taxonomy and biology. In the descriptions, the abbreviations for measurements and indices are as follows : TL, total outstretched length of head and body, including mandibles. In ectatommines, many species have re- curved gasters, and in these, the length measurement is made to the posteriormost point of the gaster in its normal position, this point usually falling on the curved tergital surface of the vaulted second gastric segment. HL, maximum measurable length of head, including clypeus but not mandibles, as seen from full-face (dorsal) view. The occipital angles are normally included if these project. HW, maximum measurable width of head as seen from full-face view, eyes are usually excluded. CI, or cephalic index, is HW/HL X 100. In cases where places of deposition, often cited in brackets, are abbreviated, [MCZ] stands for the Museum of Comparative Zoology at Harvard College, Cambridge, Massachusetts; [USNM] stands for the United States National Museum at Washington; [CAS] stands for the California Academy of Sciences, San Francisco; [NM Vienna] stands for the Naturhistorisches Mu- seum at Vienna. [1] ACANTHOPONERA PERUVIANA sp. nOV, (Figures 6, 7) Holotype worker: TL 8.7, HL 1.73, HW excluding eyes 1.56 (CI 90), greatest diameter of eye 0.43, scape L 1.07, WL 2.60 mm. Similar to A. mucronata, but differing as follows : Head with sides more nearly straight and parallel, occipital angles rectangu- lar. Eyes larger. Sculpture of head looser, with wider, more shining spaces between the rugae, especially along both sides of median carina and in antennal scrobes. Propodeal spines heavier and longer, approaching 0.7 mm. in L (ca. 0.4-0.5 mm. in mucronata workers), gently divergent; when viewed from the side, more deflnitelj' arched in their basal halves and less strongly elevated than in mucronata. BROWN : ANT TRIBE EOTATOMMINl 255 Shape of petiole shown in Figure 7 ; note the continuous curve formed l\v anterior nodal face and dorsal outline of spine. Gaster markedly depressed and somewhat broadened (Fig. 6), with a deep constriction between postpetiole and second segment, the latter slightly broader and with a conspicuous posterior im- pression extending forward into the main tergital surface from the depressed apical band. Second segment only feebly down- curved, and the exposed sternum correspondingly longer than in 7nvcronata. Apical segments retracted in this specimen, but the slender sting exserted. Gastric sculpture much coarser, denser and more opaque than in mucronaia, consisting of abundant, uneven-sized punctures, slightly larger on the postpetiole than on the second segment, forming a dense irregular rugoreticulum on the sides, less dense in the middle, where narrow, shining interspaces exist on the second segment. Erect hairs abundant, though fewer than in mucronata, but thicker, stift'er and longer. Body color bright ferruginous yellow. Holotype a unique worker [CAS] from Monson Valley, Tingo Maria, Peru, October 26, 1954 (E. S. Ross and E. I. Schlinger leg. ) . [2] ACANTHOPONERA CRASSA Sp. nOV. (Figure 10) Holotype worker: TL 6.2, HL 1.24, HW excluding eyes 1.07 (CI 86), greatest diameter of eye 0.30, scape L 0.85, WL 1.82 mm. Similar to A. minor, but a little larger. Head narrowing from a point in front of eyes toward occiput, narrowest, and sides sloping inward, behind eyes near the much-rounded occipital corners ; occipital margin short, median section straight, or even feebly convex. Median pronotal eminence low, but better de- veloped than in other species of the genus. Metanotal groove distinct, broad and rather deep. Propodeal spines (L ca. 0.35 mm.) straight and obliquely elevated as seen from the side; diverging, with tips feebly curved mesad as seen from above. Petiolar node (Fig. 10) thicker than in minor (Fig. 9), its posterapical tooth deflected more or less dorsad. Gaster very much like that of minor (Fig. 4) in general form, but slightly larger. 256 bulijEtin : museum of comparative zoology Sculpture in general a little coarser on head, alitrunk and node than in minor, the interspaces larger and shining. Pronotal rugules forming a V around the median eminence. Gaster smooth, shining, with abundant fine, separated punctulae, giving rise to dense, reclinate golden-brown pubescence. Longer fine erect hairs abundant. Body color orange-brown, legs more yellowish. Holotype [CAS] one of a series of 4 workers taken 6 miles west of Santo Domingo de los Colorados, Pichincha, Ecuador, February 23, 1955 (E. S. Ross and E. I. Schlinger leg.). The remaining three workers are paratypes [CAS, MCZ]. HL 1.22-1.25 mm., CI 87-89. Considered as belonging to A. crassa is a short series of workers [USNM, MCZ] from Hamburg Farm, Santa Clara Prov., Costa Rica, February 26, 1925 (F. Nevermann leg.). The MCZ specimen measures HL 1.19, IIAV excluding eyes 1.03 mm. (CI 87). The color is much lighter than in the type series, being bright ferruginous yellow, and the occipital angles are some- what more abruptly rounded ; transverse part of occipital border broader than in type, concave in middle. Eyes a little larger, propodeal teeth shorter and feebly bent in side view, and the posterapical petiolar teeth flattened and only slightly deflected dorsad. In color and head shape, the Costa Rican samples are some- what intermediate, tending toward minor, but in the characters of total size, metanotal groove and petiolar thickness and form, they are clearly related to crassa. The differences from minor could possibl}^ represent geographical variation, but the known distribution of minor and crassa does not support this very well. [3] The taxonomy of the New World Heteropo7iera, like that of Acanthoponera, remains in a very unsatisfactory state, due pri- marily to the lack of sufficient material and to the inadequacy of some of the descriptions of forms undoubtedly belonging here. There are a fow firm points, however. There is not much doubt about what dolo is; Roger's description fits reasonably well this large, yellowish species, the commonest one in collections, and distributed widely from Rio de Janeiro south to Uruguay and Misiones (the westward extent of its range is unknown). I have BROWN : ANT TRIBE ECTATOMMINI 257 seen quite a few series and strays from within this area, and the species is reasonably constant: the worker head length (HL) runs from about 1.20 to about 1.50 mm., and the cephalic index from 85 to 91. The occiput seen full-face is slightly concave in outline. Judging from the original description, Forel's var. a urea is apparentlj^ a semicallow worker of dolo. H. carinifrons is a distinct species, easily recognized by its small size, blackish color and unarmed node ; it is now known from various localities in Chile, and the variation from one series to the next is slight. H. ni {crops Borgmeier is very small, has minute eyes and an unarmed, rather scale-like petiolar node, and is light ferruginous in color. In addition to the types, from southeastern Brazil, I have seen specimens of microps taken at Venecia, near Medellin, Colombia (S. Flanders leg.), a rather surprising extension of the range. The Colombian sample was taken near a sample of Typhlomyrmex pusillus Emery in the soil ; T. pusilUis is also an ant usually thought of as having a much more southerly distribution in South America. With the deniinodis complex, we come to the serious confusion in Heteroponera. Up to now, there have been named in this group dentinodis itself, plus three varieties : panamensis, inermis and schweheli. Mayr described dentinodis from southern Brazil- ian examples with a short tooth on the posterior apex of the node in larger workers, but said that this tooth was obsolete on the nodes of smaller workers. However, there is no definite statement that Mayr's large and small workers come from the same nest series, so we cannot be sure that they represent the same species. The few specimens I have seen from the southern part of Brazil are strays, and so we still have no assurance that the two types of workers represented in collections are conspecific, though this does seem likely. Emery complicated the situation by describing var. inermis from a female without developed posterapical nodal tooth ; this female indicates either that the tooth character does not vary in correlation with size (i.e., is not markedly allometric, as Mayr implied), or, more likely, that another species exists with poorly developed tooth in all sizes of workers and females as well. We now know, in fact, that such a species exists in Liiderwaldt 's var. schweheli, but schweheli seems too large to be the same as inermis. It may be that Emery's measurement was too low, but we shall not be sure of the identity of inermis until the type can be reviewed. 258 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Referred to dentinodis provisionally are the following : a larger worker from Parecy Novo, Rio Grande do Sul; HL 1.02, HW without eyes 0.90, WL 1.35 mm., CI 88. One smaller worker each from Nova Teutonia, Santa Catarina (F. Plaumann leg.), and Agudos, S. Paulo (W. W. Kempf leg.), measure HL 0.78- 0.79, HW 0.70-0.71, WL in both examples 1.04 mm. ; a winged female from Petropolis, Rio de Janeiro (Coll. Borgmeier) : HL 0.97, HW 0.85, WL 1.32, forewing L about 3.5 mm. The cephalic index for these three large and small workers and one female ranges from 88 to 90. The female has a well developed and acute posterapical nodal tooth. An anteroposteriorly compressed node is characteristic for all of these specimens. The holotype of var. schweheli, from Estacao Alto da Serra, S. Paulo, has HL 1.12, HW 0.92, AVL 1.51 mm. ; CI 82. A similar, but slightly smaller specimen from Jaguara, Parana, in Coll. Borgmeier," has HL 0.98, HW 0.78, WL 1.21 mm. ; CI 78. Both of these, as well as a dealate female from S. Teresa, Espiritu Santo (0. Conde leg.), have dense, irregularly punctate sculp- ture, opaque for the most part ; interpunctural spaces of post- petiole narrow and finely reticulate or coriaceous. A similar specimen from Agudos, S. Paulo (R. Miiller leg.). Coll. Kempf, measures HL 1.07, HW 0.87, WL 1.43 mm.; CI 81. This last example is more shining, especially on the gaster, where the inter- punctural spaces are largely smooth and shining on postpetiole as well as on the succeeding segment. So far as I can see, schwe- heli is distinct from the sympatric dentinodis in shape of petiolar node (see key) as well as in the narrower head. The female node, however, is more anteroposteriorly compressed than in the worker. Total size averages larger than in the few dentinodis I have seen, but w^orker size even here overlaps slightly. For the time being, it seems best to recognize schweheli as a species in its own right, though it may later prove to be the same as inermis, as mentioned above. Two workers in the Borgmeier Collection (Nr. JBV 47) from Campinas, Goias, Brazil, resemble large dentinodis in size and general appearance, but are light yellowish-ferruginous in color and have thicker nodes (not as thick from front to rear as in worker schweheli, however) with the posterior apex continued backward as a stout subconical projection or tooth. Whether these BROWN : ANT TRIBE ECTATOMMINI 259 specimens represent a different species, or mere geographical vari- ation in dentinodis, cannot be decided without material from intervening areas in Brazil. Concerning var. panamensis, I have asked some questions about the type in the British Museum, kindly answered by Mr. G. E. J. Nixon, who made comparison for me with a paratype worker of the species I describe below as H. inca sp. nov. Mr. Nixon 's remarks are adapted in part and follow in the next three paragraphs. H. panamensis type is slightly smaller than inca, and more slender. Outstretched length (the small apical gastric segments are hidden and not included in length) of panamensis type about 3.8 mm. Color virtually the same as in inca. Head distinctly longer than broad and decidedly narrower than in inca, less convex dorsally and without the occipital lobes of inca; occipital margin straight. Propodeal teeth very short, forming angular projections not longer than wide at base. The second gastric segment is much less strongly differentiated from the first than in inca, and is nearly the same width as the first segment (postpetiole). Left mandible with 7 teeth, the seventh tooth forming the basal angle ; the three teeth next to the seventh tooth very weak. Sculpture much finer than in inca; surface of head between the frontal area and occiput and between the scrobes quite dull, very finely longitudinally striate and with the median carina in full relief. Between scrobes and eyes the surface finely rugose, not easily defined at 40 X ; toward the occipital angles the sculpture thickens slightly. From this characterization, it appears that panamensis is much closer to dentinodis than to inca, and it is not impossible that panamensis and dentinodis are synonymous. However, the distributional facts and the darker color of panamensis are, it seems to me, sufficient cause to hold up synonymy until a direct comparison of the type with other material can be made. For the time being, panamensis is raised to nominal species status. Heteroponera inca sp. nov. (Figures 12, 13) Holotype worker : TL to apex of second gastric segment 4.8, HL 1.14, HW including eyes 1.10, HW excluding eyes 1.06 (CI 260 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 93), exposed scape L 0.76, greatest diameter of compound eye 0.24, WL 1.44 mm. General habitus that of H. dentinodis, but more robust in build, darker in color, with a differently shaped head, longer propodeal teeth, thick petiolar node with calcarate ventral process, and with a proportionately larger postpetiole. Head in full-face view as in Figure 12 ; note especially the somewhat depressed and lobiform occipital angles. Seen from the side, head strongly convex across vertex and central occiput ; occipital lobes auriculate, broad, obliquely subtruncate, with narrowly rounded corners above and below, much as in Gnampto- genys cosiata and G. menadensis. Antenual scrobes weakly de- fined curving above the eyes and terminating indefinitely some distance behind toward the occipital angles. Eyes strongly con- vex, each with a circummarginal sulcus. Mandibles triangular, basal angles blunt ; apical tooth and a smaller preapieal tooth strong, acute, remainder of margin with coarse, shallow crenula- tion. Clypeus convex in the middle and with narrow, depressed anterior apron, the margin of which is feebly convex seen in outline from dorsal view. Scapes short, incrassate toward apices ; when laid straight back or nearly so from their iiLsertions, falling slightly short of the occipital margin. Funiculi gradually thick- ened toward their apices ; the last three segments could be con- sidered as forming an indistinct club ; penultimate and ante- penultimate segments about as long as thick, segments II-VIIl thicker than long. Alitrunk robust, pronotum rounded above, separated from the narrower mesonotum by a deep-cut (possibly mobile) suture; inferior pronotal margin obtusely angled on each side. Meso- notum forming a single gentle convexity with propodeal dorsum ; metanotal groove marked by a very indistinct transverse line or sulcus. Propodeal teeth well developed and acute, larger than in any other Hetcroponera so far described; seen from above divergent, slighth' longer than the distance between the inner sides of their bases. Petiole as in Figure 13 ; seen from above, the node is slightly broader than long, excluding posterapical tooth, which in this view is broad at the base, tapering to a nar- rowly rounded point. BROWN : AXT TRIBE ECTATOMMINI 2(31 Postpetiole large, semiglobose, broader than long, slightly nar- rowed around its apical border. Succeeding segment as long as postpetiole, but narrower (about as broad as long seen from above), weakly downcurved and tapering toward apex. Apical .segments tapered rapidly apicad, at least partially retractile. Legs robust ; each middle and posterior tibia with a single large pectinate spur; all tarsal claws slender and simple (without submedian teeth or denticles). Mandibles smooth and shining (when clean), with scattered punctures. Antennae, gula, gastric apex and legs with very fine indistinct punctulation or reticulation, subopaijue to opaque. Head, alitrunk, node and gaster irregularly reticulo-rugulose over an extremely fine microsculpture, opaque, with inconspicu- ous coarse punctures crowded on head and alitrunk, but becom- ing fewer and still less conspicuous posteriorly. Upper scrobes with fine transverse rugulae. Normally exposed surfaces of body covered with an abundant and mostly decumbent, fine, light-colored pubescence, sparse only on the mandibles, gula and retractile segments of gaster. Body and appendages also with an abundant pilosity of fine, tapered erect hairs, uneven in length, the longest about twice the length of the propodeal teeth. Body color deep brownish- red, legs more reddish. Holotype. one of a long series of workers taken about 6 miles west of Cali. Valle, Colombia, at an altitude of 1630 m., March 20, 1955 (E. I. Schlinger and E. S. Ross leg.), deposited in California Academy of Sciences. The paratypes have the same data as the type. TL 4.7-5.0, HL 1.11-1.20,' HW 1.00-1.11, CT 1)1-93. Propodeal teeth vary slightly in length and thickness, but in general are longer than broad at base. Posterapical tooth of petiolar node varies considerably in length, sometimes being markedly shorter than as show^n in Figure 13, and sometimes more slender and acute, but not mucli longer. Color ranging from that of the holotype to very dark reddish-brown. Paratypes in CAS, Coll. Borgmeier, USXM, MCZ. [4] Hetej'oponera imhellis (Emery). This little species is wide- spread in eastern, southeastern and southwestern Australia, where it makes .small nests under stones or woody fragments, or 262 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY directly in the soil without cover. It forages at least partly dur- ing the daytime, and solitary workers can be seen leaving the tiny, inconspicuous nest entrance to move slowly about in the open and among the leaves and debris of the soil cover. Although Wheeler (1934, p. 140) thought it hypogaeic in habits, I have not found it so. One worker was seen carrying unidentifiable remains of a very small insect into the nest. H. imbellis does not seem to be very conspicuous anywhere, but it is not rare. I have found as many as four nests in a single day's collecting at one locality. One nest contained over 70 workers and several ergatoid females. The ecological tolerance of this ant is surprisingly wide ; it occurs in a great variety of vegetation zones, from the wet mountain forest of the Australian Alps (at least to 1300 m. altitude) and southeastern Queensland to dry savannah woodland in the southern Flinders Ranges of South Australia (Mt. Remarkable) and the treeless savannah west of Melbourne. Considerable variation is seen in size, color, sculpture, length and abundance of pilosity, angularity of propodeum, and antero- posterior thickness of the petiolar node. A large part of the color variation is due merely to the long period of adult tenerality, which may cause whole broods of workers to appear very dif- ferent from one nest to another. The color also affects the visi- bility of sculptural patterns. I have examined 21 nest samples, mostly in good series, from nearly as many scattered localities, including 6 cotypes of occidentalis and a type of var. scabra, and I believe that the forms listed in the synonymy can be defended at best only as local or nest variants of imbellis. All of the characters mentioned appear in varying combinations and de- grees, and I find considerable intergradation connecting all series. The present material shows no obvious "racial" pattern on a broad geographical scale, even for single characters. [5] Cotypes of H. hroiini from the Canterbury Museum at Christchurch, N. Z., received through the kindness of Dr. R. R. Forster, were compared directly with the type series of subsp. kirki and found to be as closely identical as series from two different nests can be. Wheeler's diagnosis of kirki is not as representative as Forel's. I have followed Wheeler in recognizing BROWN : ANT TRIBE ECTATOMMINI 263 the obvious lapsus or typographical error in Forel's original spelling, "hrownii," and in emending the name to hrouni. The species is widely distributed in the North Island of New Zealand, l)ut I have seen no specimens from the South Island. [6] Justification of new synonymy and other systematic changes in Riiyfkloponcra attributed to Dr. E. 0. Wilson are included in the manuscript he is preparing on the Ponerinae of Melanesia, amounting to a revision of the species of this region. Wilson col- lected extensively in Melanesia during 1954-1955, and on his return to the United States, he was able to make critical com- parisons of MCZ material, including specimens forwarded to him out of his own Melanesian samples, with types in several of the principal European collections. This type-comparison work was important in establishing many of the new synonymies put forward here, both in his name and in my own. [7j The imprcssa group, R. aspera and B. croesiis were revised by Brown (1954b :1-11) and a number of names put into syn- onymy under these species. I have omitted these junior syn- om^ms from the list. [8] In his revision of the larger Australian Rhytidoponera, Clark (1936) placed a number of names proposed by earlier authors in the synonymy. Unfortunately, he failed to offer evi- dence for most of the synonymies, and it is possible that some of them are incorrect. In any case, they should all be reviewed. However, a spot check convinces me that most of his assignments in synonymy are probably correct. None of the synonyms from the 1936 work will be listed here. [9] A previous guess (Brown 1952b :137) that laevior Stitz was based on part of the type series of aciculata (Fred. Smith) is strongly backed by an obscure reference of Roger (1860:307) that I had not noticed before. Roger here cites aciculata ma- terial in the Berlin Museum that was sent by Smith. There can be little doubt that this is part of the material used by Stitz to describe laevior. In order to clear up any doubt about this 264 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY synonymy, I here select Hunter River, New South Wales, as the type locality for laevior, since this is also the type locality cited by Smith for aciculata. [10] R. scaherrima and R. malandensis, described from northern Queensland localities very close together, cannot be distinguished. Although malandensis was described as a subspecies of R. lacimi- osa Viehmeyer, from New Guinea, laciniosa clearly is not closely related to the Queensland form. [11] I found this little species {horealis) fairly common every- where in the open tall eucalypt woodland southeast of Darwin, at Berrimah, at Howard Springs, and along the edges of the Howard River gallery forest, in the Northern Territory of Australia. Foraging workers are often found abroad in bright sunlight. Nests are made directly in the soil, or under logs, at least during the dry season. I also have a worker from the Fin- niss R., N. T. (R. Parkinson leg.). Clark's description of hrnnnea, while failing to mention some details of sculpture, nevertheless fits this species passably well. It is the common representative of the metallica group in the Darwin area. [12] R. anceps has been poorly known, due chiefly to its rarity and local distribution. AVilson has compared specimens from near Brisbane (W. M. Wheeler leg.) with Emery's type in Genoa, and he finds that they match well. In the Museum of Comparative Zoology are other samples from Tamborine Mt. (A. M. Lea) and National Park (H. Hacker), both localities also in southeastern Queensland near the New South Wales border. Very likely the species occurs in northeastern New South AVales. The most obvious character, and one Emery does not properly emphasize, is the very low, blocky petiolar node of the worker and female. Seen from the side, the worker node averages just about as long as high without the ventral process, the proportions varying slightly either way in different specimens. As seen from above, the node is as long as broad, or very nearly so. The dorsal surface is distinct, even subtruncately set off", but is always slightly convex and usually rises slightly from front to rear. The posterior face is vertical or slightly receding ventrad. The BROWN : ANT TRIBE ECTATOMMINI 265 head is not (juite so wide behind as in related mefo/Zica-complex forms, and there is a weak impression in the middle occipital region, best seen when the head is tilted slightly back from full- face view. The scapes are rather slender and, when laid straight back, they surpass the occipital border by distinctly more than their greatest width. Funicular segments all more or less dis- tinctly longer than broad. Color brown or reddish-brown, the gaster often darker brown, without metallescence. Second gastric segment finely striate in a transverse arch, which passes pos- teriorly into a longitudinal, concentric ellipse ; this posterior region somewhat more shining. In addition to the Queensland samples, I have two workers and a headless male that I provisionally call anceps, collected at the Thomas River Station, east of Esperance, Western Australia, by E. 0. Wilson and C. P. Haskins. The locality is a wooded de- pression in the heath-covered sandplain, the trees being mainly paperbark {Melaleuca cidieularis), yate (Eucalyptus cornuta) and wattles (phyllodineous Acacia spp.), the last-named con- tributing in places to the shrubby undergrowth. Thin leaf litter exists in some parts of the depression, but in the summer this is very dry. The two workers apparently represent a rather dense popula- tion, for Wilson's notes mention, in addition to an abundance of R. mcfaUica nesting in the ground, a smaller, "more slender" species, also nesting in the earth and foraging on trees and shrubs, especially at night, at the Thomas River locality. The specimens resemble the Queensland anceps very closely in all respects, except that the petiolar node is not quite so long. In dorsal view, the node is distinctly wider than long, but it is still much longer and lower than the nodes of inornata and metallica. In general habitus without magnification, the Thomas River an- ceps looks much like inornata, and probably acts like that species in the way it accompanies metallica in a restricted, relatively favorable habitat. If this Thomas River population is actually conspecific with anceps from Queensland, or even if the two populations represent very closely related but distinct siblings, as their external morphology suggests, their present distribution is a remarkable example of a relict peripheral persistence. 266 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY [13] R. convexa and E. violacea are very closely related, the best difference so far seen being the metallic green-violet surface reflections of violacea, lacking in its eastern twin. The ranges of the two forms appear to be separated by a broad gap in central Australia ; neither has been found in the Northern Terri- tory, despite intensive collection by G. F. Hill and others, in- cluding myself. R. violacea, widespread in Western Australia, has been taken along the Transcontinental Railway in that state as far east as Kalgoorlie (Wilson leg.) and even 1072 Mile Siding (Brown leg.), but it apparently does not cross the Nullar- bor Plain. R. convexa ranges widely in eastern Queensland, and has isolated populations in the moister parts of the Flinders Ranges of South Australia [19], but its western limits are yet to be determined. The species separation is maintained here arbi- trarily in the absence of decisive information concerning their status relative to each other. The form described as opacior by Crawley is only a variant of violacea in which the metallescence is more obscure than usual, the green component especially being suppressed. Such indi- viduals have been taken from the same nest series as some with the more usual violet-green coloration, e.g., in a nest series from Pioneer Siding, north of Norseman, W. A. (Brown leg.). R. rufescens, heretofore placed as a variety or race of convexa, is really a good species, distinct in its more widely-spaced puncta- tion, as well as in the shape of its node, which averages wider at the base. -The color is yellowish-ferruginous in rufescens speci- mens from Townsville (type locality by present selection), but a sample from Bowen, Queensland (F. H. Taylor leg.) is reddish- brown, approaching the hue of lighter C07ivexa workers. I have true convexa workers also labelled as from Townsville (W. M. Wheeler leg.), but there is no sign of intergradation between the two forms in this region. [14] R. inornata, described by Crawley as a variety of metallica, is clearly a species apart, restricted to the extreme southwestern strip of Western Australia and certain of the islands off Fre- mantle. On the eastern side, it reaches the Darling Range in back of Perth, where it overlaps R. metallica without producing intergrades so far as known. R. metallica appears to be scarce BROWN : ANT TRIBE ECTATOMMINI 267 ill the main part of tlie iiioi-nata distribution, so that the two species approximately replace each other. Workers of inornata vary in color from light tan to blackish-piceons, the alitrunk usually beino- a trifle lighter than head and gaster ; metallescence is lacking. The striation of the postpetiole is coarser than in Western Australian metalUca, with broader interspaces, and is usually strongly arched over a small median area of more or less longitudinal rugulosity situated near the slightly emarginate posterior border. Punctures on the gaster vary from strong and numerous to virtually completely obsolete. The sculpture of the second postpetiolar (second gastric) segment is arched rather strongly, and the raised costulae are separated by shining interspaces, whereas the sculpture of this segment in fiietallica is very fine, dense and more or less sericeous. A normal dealate female and winged males were taken No- vember 3, 1931, at Margaret River, W. A. (W. M. Wheeler). This female differs from the worker in the ways usual for poner- ine ants, and has well-developed ocelli, compound eyes, thoracic flight sclerites and wing stumps. Wheeler's name carhonaria applies to common variants well within the normal spread of inornata. R. inornata combines several features of the two related species tasmaniensis and victoriae, from eastern Australia, where they replace metalUca in damp forests. This situation is interpreted as "character displacement" by Brown and Wilson, 1956. [15] Rhytidoponera kurandensis sp. nov. Holotype worker : A small species, to the naked eye resembling E. tenuis, or to a lesser extent B. victoriae. TL 4.3, HL 1.02, HW 0.84 mm. (CI 82), WL 1.38 mm. Head broad-oval, almost sub- circular in outline seen full-face ; greatest width at or immedi- ately behind compound eyes (excluding eyes themselves), slightly narrowed in front of eyes. Occipital angles gently rounded, occipital border convex, with a very slight flat or concave tendency in the \evy middle. Clypeus convex, rather strongly projecting and forming an obtusely rounded angle in the middle. Eyes large (ca. 0.21 mm. greatest -^ameter) and strongly convex, occupying roughly Va the sides of the head. Antennal scrobes 268 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY short and shallow, not extending backward past the eyes. Anten- nae slender; scape 1.04 mm. long from basal collar; when laid directly back, surpassing the occipital border by ^ its length. Mandibles large, finely denticulate. Alitrnnk rather slender (more slender than in tenuis), convex in profile, with a feeble impression in the region of the metano- tum. Propodeal dorsum and declivity forming one rather flat curve, sloping downward caudad. Petiole distinctly smaller than in victoriae, and slightly smaller than in tenuis, with a short but distinct pedicel in front ; free part of node nearly as high as long, with steep anterior and posterior faces rounding into a convex dorsal face ; ventral process forming an acute triangle. Node seen from above broader than long, semicircular in outline, with the curved surface anterior. Gaster a little more slender than in victoriae or tenuis, but the lengths of the postpetiole and succeeding segment are in about the same proportion (3:4) to each other. Sculpture of head, alitrunk and petiolar node consisting chiefly of coarse, contiguous foveolae or punctures with shining bottoms. Ridges separating these foveolae form rugulae, longitudinal on the head, mixed in direction on pronotum, and finer, closer and transversely oriented on the posterior half of the alitruncal dorsum. In general, this sculpture is coarser and more shining than in tenuis, and it lacks the interspersed dense punctulation found over the alitrunk and node of tenuis. Postpetiole with loose, irregularly-spaced costulae arching over a median posterior area of indistinct, mostly obsolete costulae forming a concentric longitudinal fusiform pattern, with a few coarse, shallow elon- gate punctures ; costular interpaces and much of the postero- median area nearly smooth and shining. Second gastric segment costulate on the same pattern as the postpetiole, but much more weakly and with wider, smoother interspaces ; a large postero- median area completely smooth and shining, with scattered small punctures. Erect hairs fairly abundant over most of body, mandibles, scapes and legs ; similar in abundance and distribution to those of victoriae, but averaging a little shorter. Appressed hairs of gastric dorsum few and very small, inconspicuous, more scattered BROWN : ANT TRIBE ECTATOMMINI 269 than in victoriac, and about the same as or fewer than in tenuis. Tolor deep, more or less reddish brown; mandibles, antennae and leg's lighter, more yellowish. Ilolotype worker [MCZ] collected at Kuranda, near Cairns, northern Queensland (W. L. Brown leg.) . Paratypes : 14 workers representing- six or more nest series from Kuranda and immediate vicinity (W. M. ^Yheeler leg., October 19, 1915; Brown leg. October 29-30, November 1, 1950). Also, four workers labeled "Cairns dist." (F. P. Dodd leg.) which probably came originally from Kuranda, near Dodd's home. My collections were made in rotting logs in rain forest, and consist of strays rather than colonies, probably because I did not at the time distinguish be- tween knrandensis and victoriae, one or both of which were com- mon at the same site. The only full colony of victoriae I took here, however, was in a large, rotting polypore fungus growing from the earth of the forest floor. The knrandensis workers may have belonged to diffuse colonies living in the rotten wood, but my notes do not mention the discovery of any brood with them. \"ariation within the paratype series is slight. All workers are close to the holotype in size and proportions ; in some of them, the head is a little more nearly quadrate, owing to less convex sides and occiput. There is slight variation in the height and thickness, as well as the angularity, of the petiolar node as seen from the side, and different specimens differ slightly in the density and distinctness of the gastric sculpture. Color Agarics from light brown (tenerals?) to piceous. The appressed pubescence of the second gastric segment is at best very sparse and inconspicuous, and in some specimens can scarcely be made out even in the best lights and magnifications. Paratypes to be deposited in USNM, ^ICZ and one or more Australian collections. [16] Rhytidoponera chnoopyx sp. nov. Ilolotype v\-orker and two paratype workers : Like knrandensis, but with the entire exposed tergital surface of the second gastric segment densely and uniformly covered with fine, short appressed pubescence, directed posteriorly and mesad. This pubescence forms a distinct brownish-yellow cast over the segment, but does not quite conceal the sculpture beneath. The individual setae of 270 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY the pubescence arise from myriad fine, separated punctulae, but otherwise the sculpture is like that of Ixurandensis. R. pulchella Emery, of New Caledonia, is very different in other ways, but has somewhat similar dense pubescence on the gastric dorsum ; however, in pulchella the pubescence is less fine and is more whitish in color. Holotype and two paratype workers taken (apparently to- gether) at Millaa Millaa, Atherton Tableland, northern Queens- land, at an altitude of about 2500 feet, undoubtedly in rain forest (P. J. Darlington leg., April, 1932). Deposited in MCZ. Except for the gastric pubescence, kurandensis and chnoopyx are so similar that they could represent the same species. I prefer to separate them specifically for the time being, since I feel that sibling species may be rather common in this limited Ather- ton Tableland-Cairns region. The similar, but longer-headed species tenuis and peninsularis [27] form a more or less parallel Arfenkreis, and it is worth noting that no two of the four forms mentioned in this paragraph have ever been taken at exactly the same locality. Bhytidoponera of this general group are well worth further investigation in northern Queensland. [17] The group of species related to E. punctata includes robust, medium-sized entities of black or brown color, very finely and evenly reticulo-punctulate over most of the body, so that the integumental surface is largely opaque. In addition, the head and alitrunk (especially) bear larger, conspicuous punctures or foveolae. These forms are distributed widely in the drier parts of South, central and southwestern Australia, but up to now, they have been only very sporadically collected, which is unfor- tunate for the taxonomy, seeing that there is a great deal of variation in sculptural and minor structural details from nest to nest and especially from place to place. As might be expected, there seem to be more names than there are species, in large part due to splitting by Clark and Crawley. I have not seen the types, or even topotypical material, of R. punctata, so I shall suggest no formal synonymy here. However, it does seem to me that R. flmdersi is very likely to prove a synonym of punctata. And material referable to punctata from Forrest, Western Australia [C. Barrett leg., MCZ] and near Balladonia, W. A. BROWN: ANT TRIBE ECTATOMMINI 271 [E. 0. Wilson and A. Douglas leg., MCZ] shows that punctata may be distributed widely, if sporadically, across the wastes at the head of the Great Australian Bight, or at least that the dis- tance separating eastern and western populations of the complex is not so great as has generally been assumed. In Western Australia, the two forms douglasi (^ levior Crawley) and ru- fonigra are yevy like punctata; in douglasi the conspicuous punc- tures are much smaller and farther apart ; however, some speci- mens (e.g., from Dongara, AV. A.) are difficult to assign, and maj^ be intergradient. A close study should be made of these forms in the Perth area ; if they are found to intergrade there, syn- onymy with each other and with punctata would seem fairly certain. However, it is also possible that character displacement is operating here. The punctata-gYow^ representative in central Australia is B. incisa, of the MacDonuell Ranges, where it is fairly common in my experience. This form is the most distinct of the group, and seems to be well isolated from the other species ; however, we must expect further members of this complex to turn up from ranges to the south of the MacDonnells, in South Australia, and these may well link incisa to punctata if they do occur. [18] Wilson (MS. notes) has compared an MCZ specimen from Broken Hill, N. S. W. (F. W. Shepherd leg.) with types of maniac and spatiata in the Forel Collection. He finds that, "maniac is the same as spatiata — the types are nearh^ identical ; spatiata is somewhat larger and with a proportionately broader head, but the Broken Hill specimen is intermediate in both characters." I have also found that the Broken Hill specimens (determined as spatiata by Clark) are very similar to some collected by Zietz, Tepper and myself in the Lofty Ranges east of Adelaide. The types of both species probably came also from this neighborhood, more precisely the dry eastern scarp of the Lofties, such as the locality, at Barren Falls, near Mannum, South Australia, where I found maniac avoiding the heat of the day in December, and foraging at the same time as mayri after late afternoon shadows fell on the canyon floor. R. maniac is essen- tially a species of the Murray-Darling drainage area. To the 272 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY north and west, in the Flinders Ranges, it is apparently replaced l)y another, similar species that I have not yet satisfactorily determined [19]. Wilson has called attention to the close similarit}' between maniac and R. hiJIi Crawley, from the north of the Northern Territory. The two are indeed similar, but hiUi averages lighter in color. The hilli types (Stapleton, N. T.) are brown with reddish gaster, while a series I toolc at the Mataranka Hot Spring, on the Roper River, is a rather uniform rich orange-brown. In these two series, the second gastric segment is usually transversely striolate, but occasional examples have the posterior part with longitudinally or obliquely arched striolation approaching the condition in maniae. R. hilli also tends to have a slightly more opaque and finer sculpture on head and alitrunk. Whether the two populations represent separate species remains to be decided. [19] In the Flinders Ranges of South Australia, where I searched intensively for Rhyfidoponera, I found only three of the larger species {R. mctalUca was also present). Of these, the largest and most tolerant of xeric conditions, and hence the most continuously widespread species, is R. mayri. The smallest clearly agrees with R. convcxa, which is usually thought of as a tropical Queensland species. The middle-sized species of the three is similar to maniac [18], but it is a little larger and is more heavily and rugosely sculptured over head and alitrunk. With convexa, this middle-sized species occupies the more moderate localities within these largely very arid ranges, especially such moister localities, often with some woodland cover, as Mt. Re- markable, AVonoka Creek and Wilpena Pound. I have not been able to match this species satisfactorily with any of Clark's (1936) descriptions. The name 7iigra could possibly apply to it, but it seems more likely that nigra is Clark's designation for the Flinders Ranges populations of convexa; his description and figures are not good enough to make the identification certain. (A belated look at specimens of R. nigra, determined by Clark and belonging to the original series from which he described nigra [Owieandana, South Australia, Hale and Tindale leg.], establishes, as far as I am concerned, the synonymy of nigra with convexa. I have not changed the list to indicate this synonymy.) BROWN : ANT TRIBE ECTATOMMINI 273 [20] R. mayri was described from material erroneously labeled as liavinp- come from New Zealand, but which probably came instead from the hinterland of Adelaide, South Australia. Dr. Wilson has been able to compare material from various localities with the )naijfi type, and he has been able to make a very close match with specimens from Mildura, Victoria (F. H. Taylor leg.), these latter matching in turn Clark's concept of the "mallee species," R. dixoni. The samples of the mayri complex come from many arid inland localities in the southeastern quarter of Australia, ranging from southwestern Queensland through western New South Wales and Victoria into South Australia. The complex is not known from south-central Australia, and it is not yet known to extend continuously, or even as a chain of isolated populations, across the arid inland of South and Western Australia. R. mayri complex is absent from, or at least very uncommon in, the extreme south of Western Australia, but is met again in the dry country to the north and northeast of Perth. Specimens from Geraldton and the Mullewa area, more or less near to the coast of Western xiustralia, are very like eastern R. mayri, and show a large part of the same variation in size, head shape, color, sculpture, pilosity and especially node shape, that marks the eastern series. Going farther inland, at Meeka- tharra, Wiluna, Lake Violet and Yandil, W. A., (leg. W. M. Wheeler, P. J. Darlington), one encounters populations with a larger proportion of reddish individuals, and in which the rugos- ity of the body, especially the pronotum, tends to be reduced and looser, with wider, very finely reticulate sculpture in the inter- spaces. Both occidentalis and glabrior (as represented by speci- mens in the ^ICZ determined l)y Clark) appear to represent the more southerly, more coastal type with heavier sculpture, which I do not feel from present evidence should be nomencla- torially separated from mayri. Clark relied heavily on the shape of the petiolar node in separating what he thought were different species, but it seems that he failed to realize the amount of variation in this segment. In 1)oth the eastern and western populations, and also in single nest series, one can find nodal variation completely exceeding the narrow limits Clark apparently allowed. Variation is strong in other characters, also, particularly those already mentioned. 274 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Some series have numerous short, erect hairs over the body and appendages, others are bare or nearly so. The striolation of the gaster, particularly of the second segment, varies from transverse, through strongly arched, to longitudinal. In the population called quadriceps by Clark, from Tennant Creek and the Mae- Donnell Ranges (Mt. Frances, leg. G. F. Hill) of Central Australia, the short erect hairs are abundant evervwhere on the body, limbs and scapes, and the second gastric tergite is longi- tudinally striolate. The sample described as stridulator, from inland New South AA^ales, is intermediate between quadriceps and South Australian populations in having the average second gastric segment with longitudinally arched striolation ; the short hairs are mostly erect, with a sparse growth of appressed, pubescence-like hairs also, especially on the gastric dorsum. The extreme quadriceps variant may represent a case of character displacement in the mayri population of the MacDonnell Ranges, because a very similar species is known from there in B. mirahiUs, a form notable for its virtual lack of erect hairs and for the substitution of a rather dense pubescence-like pilosity of short, appressed to decumbent, fine hairs; it is also worth noting that the dorsigastric striolation of mirahilis is weakly arched-trans- verse in contrast to the longitudinal orientation of the quadriceps sculpture. At present, this interpretation must be considered a very tentative one, awaiting more material from the MacDonnell Ranges and from areas between these mountains and South Australian districts in the known range of mayri. In my own collecting at Tennant Creek, and around Alice Springs in the heart of the MacDonnells, I found no /Jtat/n-group species (begin- ning of cold season), but took numerous samples of taurus, incisa and one or two other large species. Apparently, the mayri group is near the limit of its range here, in an area largely dominated by taurus (a large species that is similar in many ways). R. petiolata refers to specimens from northeastern South Australia with "dome-shaped" nodes (type loc. : L. Killal- paninna, S. A.). Specimens in the MCZ from this general area (Cooper R., leg. Reuther; L. Callabonna, leg. Zietz) have rounded nodes much as indicated by Clark for petiolata, but nodes similar to these can also be found in series from the Flinders Ranges in South Australia (Brown leg.). The petiolata kind of petiolar BROWN : ANT TRIBE ECTATOMMINI 275 node seems, therefore, to be only a part of the normal variation of mayri in South Australia. The variation already cited for mayri is of the kind that would undoubtedly be described by ornithological colleagues in terms of the ''polytypic" species, with the various species of Clark mostly ranking as races. It seems preferable, however, that future students of this very interesting group be encouraged to examine it from the vicAvpoint of independent character variation studies. Another point worth study in the future is the relationship of mayri to aciculata [9]. [21] The mefaUica complex deserves a full-scale study which, when it comes, will undoubtedly furnish wonderful data on geographical variation. The species metallica is one of the most abundant, easily collected and variable of Australian insects. I have examined the specimen in the British Museum labeled "type" and here designate it as lectotype; the specimen with others, is from Adelaide. These specimens are, of course, old ones, and their color may have changed, or they may have come from some distance away from Adelaide proper. However this may be, they can be matched rather closely by certain specimens taken by myself in the Adelaide district in 1950 and 1951. Here the color varies from metallic green to purple with green over- tones. As one travels northward in the dry Flinders Ranges of South Australia, the color of metallica series gradually and rather irregularly shifts more and more toward a uniform dark purple, and dark purple clearty predominates in the series I took around Alice Springs, in central Australia. It is to this northern desert variant that Wheeler gave the name purpiirascens. Clark's pulchra, from farther west, does not seem to differ from this dark form in any significant way, at least judging from his descrip- tion. B. caeciliae doubtless belongs here also; I have had a hurried look at a cotype in USNM. In country with somewhat better rainfall, such as the south Victorian savannahs and the New South Wales tablelands, metal- lica workers are more often predominantly green, though with purplish tones usually present on the sides of the alitrunk. Northward in New South Wales and Queensland, one meets a form in which the alitrunk is predominantly reddish-violet. 276 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY shading into golden on the lower pleura, and with the green restricted to a mid-dorsal strip, or even absent. The gaster of most such s])ecimens, unlike that of the central desert populations, is solid, bright green. In some places, especially at the University of Queensland Farm at Moggill, near Brisbane, I found what seemed two slight color forms living side by side under stones in sparsely-treed savannah ; one of these was the form jiLst men- tioned, while the other was noticeably more completely green. The difference is still visible in the cabinet series obtained, but it was more obvious as seen in the many living colonies opened in the field. Besides color, I can find no constant differences between these series. Whether this variation marks two sibling species cannot be determined from such casual and incomplete evidence, but the possilnlity cannot be overlooked. These same two color forms are connected by all degrees of intermediate coloration in other localities, especially farther south, but it is possible that this is only another case of ''character displace- ment" [14]. However, there was no obvious sign of ecological seg- regation between the two forms at Moggill. In the far north of Queensland, in the Mareeba-Koah district, the open monsoonal sclerophyll woodland and savannah wood- land supports an abundant population of a smallish, rather uni- formly dull green tiietallica-like form, quite distinct from the more southern types. However, we do not know how these populations hook uj) west and south of the Atherton Tableland, so it is impossible to say whether the Mareeba district form is an independent species (maleclictaf) or not [31]. In addition to the extensive and striking variation in color, metallica shows a range of minor differences in size, shape of head and of petiole, length of appendages, sculptural details, and so on. Crawley's var. va)iau.s is a form in which the superficial punctures of the gastric dorsum are coarser and more numerous tliaii usual. But such variants are found not only in the Darling IJange, Western Australia, the varions type source, but also in various parts of southeastern Australia. Intergrades to forms with weak or obsolescent punctures are found both east and west, so that varians cannot be considered as a separate population. Ir is interesting to note that sculptural variation shows no BROWN : ANT TRIBE ECTATOMMINI 277 geographical concordance with color variation over broad areas, iit least in tlie fairly abundant material present in the MCZ. Part of the varians type material was teneral, it should be noted. The metallica complex is represented in the extreme south- west by inornata [14], in the far north by horealis and trachypyx [11, 28], and in moister parts of the southeast, by tasmaniensis. All of these species are very nearly or completely non-metal- lescent, and in their more brownish or reddish color phases, they resemble one another, to the naked eye. R. tasmaniensis meets and intermingles with metallica at various points in the savannah countrj" of western Victoria, in the environs of Melbourne, and on the New South Wales tablelands, but the two remain distinct and easily separal)le at a glance at the living colony, even when their galleries are only a few feet apart. However, teneral speci- mens of iDciaUica isolated in the cabinet may he mistaken for ias)na}iiensis, since apart from color, the two species are closely similar. The var. cristulata Forel is the same as tasmaniensis (Emery. 1912:79) ; types have been compared by Wilson. Tas- maniciisis is brown or reddish broAvn in color, the gaster darker brown and sometimes faintly bronzy in tone. [22] I took a few workers of R. reflexa from a small nest on the Koolpinyah Track, near the western entrance of Koolpinyali Station, Darwin district. Northern Territory, Australia. Kool- pinyah, one of the original localities, is here selected as type locality. My nest was taken in the middle of a typical, broad tioodplain savannah, floored by heavy soil covered with hard, lumpy earthen hillocks up to 15 or 20 cm. high and about the same in diameter, formed by an unknown agency. This savannah, shallowly flooded in the wet season (my visit was made in the early dry season), is characterized by widely scattered, spindly, low Melaleuca and Bajilcsia trees, and bj' a poor ground cover consisting of sedges and certain special flowering herbs. The ants taken here, while not al)undant, are special forms not seen else- where during my stay in the Darwin area ; they include R. reflexa, a species of Polyrhachis, and a small, bright yellow Monomorium, the flrst two nesting in the hillocks, and the last in rotten sticks on the ground. It is possible that the ants escape part or all of the monsoon floods l)y iiieans of the hillock nest sites, although 278 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY I have never seen this area myself when it was in flood, so cannot verify the surmise. Clark's description and figure of this species are rather crude and sketchy, but it is so aberrant a form that there can be little question of identifying it. Clark records it also from Bathurst Island, and it is probably widespread in suitable localities in the Darwin area. It will be interesting to see what happens to B. reflexa when and if the projected rice-culture areas are begun there. [23] I took R. reticulata in open savannah woodland at Kath- erine. Northern Territory, on July 4-5, 1951. Workers were for- aging from a single small craterless nest entrance in the hard soil. Males were present ; like the lighter-colored workers, they are yellowish-brown. Foraging is nocturnal. [24] R. rufiventris was originally described as a variety of R. convexa, but the two are quite distinct in size, shape of head and petiole and in their distribution, although they are sympatric over very large parts of Queensland. R. rufiventris is common and conspicuous in the open forest country of north Queensland ; in Western Australia, it appears to be widespread but erratic and local in occurrence. The MCZ has specimens from this state labeled as from the Fortescue River (L. Glauert leg.), from Subiaco (Hamburg S. AY. Australia Expedition, ex Forel Coll., det. Forel as R. convexa var. violacea), and from Derby (J. Gr. Campbell leg.) ; the last are probably part of the type series of Crawley's synonym B. castanea. Many of the specimens from the region west of Kuranda, north Queensland, are rather uni- formly dark brown, instead of having the gaster red. I did not find this species in the neighborhood of Darwin or elsewhere along the Darwin- Alice Springs highway in the Northern Terri- tory. R. rufiveritris needs to be considered carefully in relation to the very close forms maniae and hilli [18] and also rufescens [13]. From evidence present in publications and collections I have seen, it is not impossible that some or all of these forms are geographic variants of the same species. A dark brown form with red gaster, appearing superficially very like rufiventris BROWN : ANT TRIBE ECTATOMMINI 279 to the naked eye (when alive) occurs at Alice Springs, in the central Australian MacDonnell Ran<>'es. However, this form is smaller and has the head shape of small convcxa; it may repre- sent an independent species. [25] I have examined a specimen labeled "Typus" of B. spoliata from the Genova Museum, courtesy of Dott. Delfa Guiglia. This worker is further labeled as from "Kamerunga" in Queensland, ]>rol)ably wrongly, since the type locality cited with the descrip- tion is Mt. Bellenden Ker, northern Queensland. Podenzana collected at both localities, but Bellenden Ker is the more likely locality for spoliata ; I have other specimens taken in the Cairns- Kuranda area hy T. Greaves, C. P. Haskins and others, nesting in epiphytic Platyccriiim in rain forest. Emery's brief descrip- tion fits this specimen well enough, but does not mention the somewhat sunken eyes, surrounded by circular rugules. In his collection, Wheeler confused this species with laticeps, but laficeps has tlie gaster almost completely smooth and shining, and the rugules near the eyes do not pass circle-like around them ; instead, those rugules anterior to the eye pass out radially from it (Wilson notes from type). The species described and figured by Clark in 1936 as B. spoliata, from Mutchilba, northern Queensland (A. D. Selby leg.), according to a specimen sent by Clark to the MCZ, and evi- dently from this same series, is really B. scahra Mayr. [26] I have a worker taken on the south shore of Lake Eyre North (G. F. Gross leg.) that agrees fairly well with Clark's ilescription of viridis, except that in my worker, the posterior border of the head is feebly convex, not concave. The head is narrower behind than in nietallica, the scapes are longer, the striation of the second gastric segment is finer, and the size is a little larger. [27] There is a worker type of B. tenuis in the MCZ which, although it is headless, otherwise matches very closely three specimens from Cairns, Queensland (W. M. Wheeler leg., 191-1). Cairns and MackaA- (the type locality) are about 400 miles apart, so it is possible that tenuis ranges along most of the 280 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY northern Qneensland coastal strip. This small brown species is easily recognized by its moderately elongate head (HL 1.10-1.18 mm.; CI 72-75) with transverse, slightly convex occipital border (a slightly concave median portion is seen in some views), long- scapes, and large, strongly convex eyes. Spaces between foveolae of alitrunk, sides of alitriink, node and a broad posterior strip on the postpetiole, densely and finely retieulo-punctiilate and opaque. Rhytidoponera peninsularis sp. nov. Holotype worker : Very similar to B. tenuis, but differing in sculpture. TL 4.9, HL 1.18, HW 0.87 mm. (CI 74), WL 1.61 mm. The foveolae or coarse punctures are larger and more closely contiguous, with strongly shining bottoms, and such interspaces as exist are mostly more or less smooth and notably shining, not densely punctulate as in tenuis. On the sides of tlie posterior alitrunk, the sculpture tends to form indefinite oblique costulae that run over the top of the propodeum and become very distinct where a few of them cross the propodeal de- clivity, where the interspaces are very smooth and shining. As in tenuis, the postpetiole is finely and regularly arched-striate, but the striation continues back to the posterior tergital border (re- placing the punctulate band of tenuis there) and surrounds con- eentrically a posteromedian area where the striae form a longi- tudinal fusiform pattern. The posterior reticulo-punctulate band of tenuis is not present in peninsularis. Second gastric segment shining, superficially arched-striate in front, becoming smooth behind, w' ith scattered inconspicuous punctures ; no fine reticulo- striate pattern as in tenuis. Ill addition to the sculptural differences, peninsularis has a slightly thicker and more squarely-cut petiolar node, as seen from the side. The anterior and dorsal slopes are separated by a more abrupt angle, and the posterodorsal angle slightly overhangs the posterior face. Color medium brown. Holotype a single worker [MCZ] from the Rocky Scrub (rain forest) in the Mcllwraith Range, northeast of Coen, Cape York Peninsula, Queensland (P. J. Darlington leg., June 19, 1932). BROWN : ANT TRIBE ECTATOMMINI 281 R. tenuis and E. peyiinsula)is are distingaiishable from all species of similar size and color found in northern Queensland by means of their elongate head (CI distinctly less than 80, and perhaps normally under 76) and generally convex occiput. The related species R. kiirandensis and R. chnoopxjx have shorter lieads (CI normally 80 or more). The common R. victoriae is distinct from all of these species in having a short, broad head with broadly concave occipital l)order and salient occipital angles; also, victoriae has very short scapes that reach or barely surpass the occipital l)order, in contrast to very much longer scapes for the other four species. There are, of course, other differences as well, but the features mentioned should serve to distinguish the small brown species at present known from northern Queensland. [15, 16, 31] [28] Khytidoponera trachypyx sp. nov. Holotype worker: With the general form of R. mctaUica and relatives, especially R. horealis. Size a little larger than that of boreal is, but smaller than the average for metaUica. TL 5.1, HL (including clypeus and occipital lobes) 1.22, HW 1.06, WL 1.66. petiole L 0.45 mm. Head narrower than in the average mctaUica worker ; CI 87 ; occipital angles very prominent ; posterior border seen in full-face view rather deeply concave. Antenna! scape slender; when laid straight back from insertion, overreaching the occipital border (at the point of intersection) by a distance greater than the length of the first funicular segment. Petiolar node as in horealis, i.e., similar to that of metallica, but more rounded above. The be.st characters are in the gastric sculpture. Large punc- tures of petiole and postpetiole coarse, deep and close together, with shining bottoms. Between these deep punctures the surface is finely and densely reticulo-punctulate and obscurely rugulose, the orientation predominantly longitudinal. The succeeding seg- ment is longitudinally costulate, the costulae being numerous and distinct, although becoming more irregular and tending slightly to anastomose anteriorly. Intercostal spaces finely reticulate, and with scattered large punctures. By comparison, R. horealis has each of the two segments finely and densely reticulate-striate 28^ BULLETIN : MUSEUM OF COMPAKATIVE ZOOLOGY in a pattern arching over median posterior areas of fine reticulo- punctulation ; scattered over both segments are large punctures which, however, are shallow and sculptured like the surrounding surface. Color brownish-red, legs moi'e yellowish ; petiole and gaster pieeous, appearing nearly black to the naked eye. This color is much as in horealis and some samples of tasmaniensis, although horealis frequently is darker over head and alitrunk. Other characters much as in horealis. Holotype a single worker taken foraging in the late afternoon on the sandy river liank at Katherine, Northern Territory, Australia (W. L. Brown leg.) ; deposited in the Museum of Com- parative Zoology. Paratype a single worker in the MCZ, labeled, "Darwin, N. T." in W. M. Wheeler's hand. I consider it likely that this is not the precise locality, but represents instead a shipping point or base locality for the collector (possibly Wesselmann). Other ants, such as R. taurus, in the MCZ collections bear similar labels, but it is virtually certain these were collected much farther south, near the center of the Northern Territory. The upper parts of the Northern Territory, nearest Darwin, are occupied by the closely related common species U. horealis [11], and present indi- cations are that B. trachypyx may replace the latter in the drier savannah woodland zone of the central Territory. [29] Rhytidoponera tyloxys sp. nov., Brown and Douglas (Figures 36, 37) Holotype worker : TL, adjusting downward for expansion of gaster 7.8, HL 1.57, HW 1.19 (CI 75), L head with closed mandibles 0.93, scape L 1.32, greatest diameter of eye 0.50, WL 2.37, petiole L 0.62, gaster L, greatly extended, 3.46 mm. Details of form of head, mandibles, antennae and petiolar node with adjacent segments as shown in Figures 36 and 37. Ali- trunk convex in profile, highest in mesonotal area, with a very shallowly concave portion along the propodeal dorsum. Pro- mesonotal suture fine but distinct ; other sutures obliterated on BROWN : ANT TRIBE ECTATOMMINI 283 dorsum of alitrunk. Pronotum evenly rounded, its lower left- liand border not forming the acute tooth (this tooth is an im- ]K)rtant generic character), but instead produced only as a sub- rectangular (actually slightly obtuse) angle. The border on the right side forms a more conventional toothlike projection, but even this is not so conspicuous or acute as is usual for the genus. 37 Figures 36 and 37. Ehytidoponera iyloxys Brown and Douglas, sp. nov. Fig. 36, full-face view of head. Fig. 37, side view of petiole and adjoining structures. Sculpture characteristic. The body, with mandibles and ap- pendages, is very finely and densely punctulate and opaque, except that the posterior half of the gaster has the sculpture shallower and weakly shining. In addition, there are abundant coarse, shallow umbilicate punctures or small foveolae (not inter- rupting the punctulation), contiguous to subeontiguous on head and pronotum, becoming smaller, spaced and indistinct on mesonotum, sides of alitrunk and petiolar node ; nearly or quite obsolete on propodeum and postpetiole. Between the foveolae on the clypeus and dorsal surface of the head are longitudinal rugulae that fan outward over the vertex and become lost in a reticulum behind the eyes. The punctulation of the mandibles passes into fine striation apicad. 284 BULLETIN : MUSEUM OF COMPARATI\^ ZOOLOGY Body largely devoid of conspicuous erect pilosity, those hairs remaining being mostly short and stiff : about 10 on anterior elypeus ; a few on masticatory and ventrolateral mandibular borders ; a few on oral Iwrder of gula and on smaller mouthparts ; about 4 along the extensor surface of each scape, and a group at each tip of scapes ; a circlet on each funicular segment ; a widely spaced pair on the pronotal dorsum ; a few on the legs, including the coxae ; a row across the posterior part of the postpetiolar sternum ; a spaced short pair on the middle of the second gastric segment ; a circlet around the posterior borders of the second and succeeding gastric segments, the hairs becoming longer toward the apex. In addition to the erect hairs, there is a sparse growth of fine, small, inconspicuous, appressed or decum- bent hairs, one each in most cephalic foveolae and in a few of the foveolae elsewhere, also forming a very dilute pubescence on scape, mandibles, gula and on legs and gaster. Color deep orange- brown, the liead, alitrunk and petiole a trifle darker than the appendages and gaster. Holotype [Western Australian Museum, Perth] one of a series of seven workers taken by Mr. K. C. Buller at Woodstock Station, about 900 miles north of Perth, Western Australia, in June, 1952. Paratypes: the remaining six workers in the type nest series differ from the holotype only very slightly in dimen- sions and proportions ; the holotype is about average for the series. Some workers have the convex median portion of the occipital outline slightly emarginate at its summit ; there is some variation in the distinctness of the inferior pronotal teeth and in the length, thickness and acuteness of the posterodorsal process or tooth of the petiolar node, some specimens having this point slightly less sharp and salient than as shown in Figure 37. There is also some difference in number and position of erect hairs, but this is at least partly due to abrasion of the integument. From the few other Rhytkloponera with petiolar node produced to an apical point, R. fyloxys can be distinguished immediately by means of the structure of its head, particularly the large eyes, and by the peculiar sculpture. [30] Wilson manuscript notes on comparison of types in the Forel Collection: "R. haeckeli and R. tunieri are closely related BROWN : ANT TRIBE ECTATOMMINI 285 but distinct species. In haicl-di, sculpture of posterior half of head consists of fairly regular rugae which originate in the space between the frontal lobes and radiate outward toward the occi- pital corners. In itirneri. sculpture in the same area consists of a solid rugo-reticulum with no orientation." B. lamellmodis Santschi also belongs to this group of species, which have high, angulatp propodeum and thin, scale-like, dorsally emargiuate petiolar node. Differences, if any, separating haeckeli and lamel- linodis remain to be clarified. [31] R. vicforiae is a common and widespread eastern Australian species, ranging from Mt. Spurgeon, northern Queensland (P. J. Darlington leg.) around the moister eastern and southeastern fringes of the continent to the grasslands of Western District in Victoria. It is the smallest of the common Australian members of its genus, and is a familiar inhabitant of home gardens in the suburbs, and even in the large central parks of a city like Melbourne. The variants called rnodesfa, scrobiculaia and cedarensis were never satisfactorily distinguished from victoriae, and it seems that they were described only because Emery and Forel lacked sufficient material of vicioriac to appreciate the extent of its variation. Wilson has now been able to compare the types of all these forms in the Emery, Forel and Andre collections, either directly or Avith samples furnished from MCZ. B. modesta is a variant showing very feeble bluish metallescence over a blackish or piceous ground pigmentation ; this form is most prevalent in moist forests and upland grassy clearings in southeastern Queensland. In northern Queensland, the ground color varies from dark to very light brown in fully adult workers; and metallescence is not present. In northern Queensland also, especially near Kuranda, the gastric sculpture shows ex- tremes of variation in the fineness and coarseness of the post- petiolar and gastrodorsal striation. It is not possible from the present material to rule out entirely the occurrence of more than one sibling species in northern Queensland, but now it seems better to take the conservative course in recognizing only the single species victoriae as representing the complex in north- ern Queensland. 286 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY R. maledicta, described as a race of victoriae, seems distinct in lacking a well-defined posterior serobal area ; the corresponding- area is coarsely rngo-reticulate instead of longitudinally rugo- striate. The relationship of maledicta to the dnll-green North Queensland metallica-Vike population needs study [21]. [32] The Chalcoponera hilli described by Clark in 1941 becomes a homonym of R. hilli Crawley through the merger of the two genera, a fact duly noted by Donisthorpe when he proposed the new name clarki to replace Clark's hilli. The MCZ has cotypes or nidotypes of this large, robust, mc tallica -like brown species ; these specimens lack metaliescence entirely. It seems likely that Forel's ohscurmn, an earlier but also preoccupied name, applied in part to the same form. Forel mentioned that some specimens had greenish heads, but it seems possible that he may have had mixed samples, including some large examples of mciallica with faded or otherwise obscured metaliescence on head and especially on alitrunk, petiole and gaster. The available name is clarki Donisthorpe. This species apparently is not common, and is found only along the central Queensland Coast and on certain Barrier Reef islands. [33] The Rhytidoponera of new Caledonia So far as our present information goes, there are seven species of Rhytidoponera on New Caledonia and the islands on its coasts. This figure includes two species described here as new, and leaves out two now placed as synonyms. Undoubtedly there yet remain species to be described, particularly from the poorly-known highlands of the north and from the coastal islands, but a key should help to make further study easier. Up to now, most of the work on the Rhytidoponera has been done by Emery (1883, 1914), although Andre and Viehmeyer each described a single species. Unfortunately, in his 1914 paper, Emery momentarily confused the two common species R. fidgens and R. numeensis, so that most of his remarks on nunieensis, at least on the workers, actually apply to fidgens. In this work he described the real niimeerisis over again as acupuncta. The suspicion that some such mixup applied here has been fully con- firmed, and the confusion itself finally cleared, by Wilson's BROWN : ANT TRreE ECTATOMMINI 287 examination of the types and other specimens in the collections of Emery (Genoa) and Andre (Paris), with fresh specimens of the species concerned at hand for comparison. Wilson has also noted that the type of var. socrula differs from ' ' typical ' ' fulgens chiefly in the more definitely oriented rugation of the head (longitudinal) and alitrunk (transverse, especially on prono- tum ) ; in Wilson 's extensive series from the southern half of New Caledonia, one can find all degrees of intergradation linking ex- tremes of the two kinds of sculpture, without particular terri- torial attachment of the variation. Along with the two new species he collected in the countrj', Wilson's most helpful contribution is the detailed information contained in his notes on the distribution, ecology and behavior of the New Caledonia species. This information can be sum- marized as follows. Collections were made in several localities, both disturbed and relatively undisturbed, in the southern half of New Caledonia. In all of these localities, at altitudes below about 800 m., there occurred the large metallic green species, R. fulgens, and usually also the smaller, black or brown species, R. nnmeensis and R. pulchella. At three of the forested upland localities, there was found a new small species close to numeensis, but with a thinner node. All of these species were taken foraging during broad daylight on low foliage in the forest understory or at the forest border, and all of them were taken in Berlese funnel samples from the forest floor leaf litter. Mostly, their nests were found under rocks in the soil, but occasionally one or more of these forms nests in rotting wood. A second new species of the small-sized group was taken twice at a single locality {R. versicolor', Montague des Sources), at altitudes of about 800 and 1000 m. ; significantly, no other mem- bers of the genus were seen at these altitudes at this locality, nor, except for R. acanfhoponeroides, were any other species of Rliyiidoponera taken above 800 m. anywhere by Wilson. The food of these species, so far as observed, consisted of small arthropods, some of which were certainly taken alive. To what extent the New Caledonia Rhytidoponera feed on plant sugars and other foods is not vet known. 288 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY The brief review of the species offered below is based on Wil- son's ample collections from New Caledonia, made during Decem- ber, 1954, and January, 1955 ; smaller samples, all from the vicinity of Noumea, were sent me by N. L. H. Krauss. Key to the Rhytidoponera species of New Caledonia — workers 1. Larger forms, full length of alitrunk (WL) exceeding 2.0 mm.; body color of mature specimens with rich metallic green or purple tones .2. Smaller forms, full length of alitrunk (WL) less than 2.0 mm.; body color black, brown or combinations of these with red or yellowish; no metallic colors 4. 2. Petiole produced backwards as an acute apical tooth ; gastric dorsum smooth, with only shallow traces of coarse punctures, the surface cloudy Avith a bluish opalescence ac-anthnporieroides Vieh. Petiole bluntly subtruncate or rounded above, not produced as a tooth from the apex; gastric dorsum largely or wholly striate or reticulate . .3. 38 39 40 Figures 38-40. Rhi/tidoponera spp. from southern New Caledonia. Workers, side view of petiole and adjoining structures. Fig. 38, E. wilsoni sp. nov., paratype. Fig. 39, B. numeensis Andre. Fig. 40, B. versicolor sp. nov., paratype. 3. Coarse sculpture of body overlain by finer sculpture, rendering the surface largely opaque ; postpetiole and succeeding segment very finely reticulate atropurpurea Emery Coarse punctures of body shining, without overlying fine sculpture of much account; postpetiole and succeeding segment transversely- to arched-striate fulgens Emery BROWN : ANT TRIBE ECTATOMMINI 289 4. Ventral process of petiole forming a long, straight, very slender spine; the small oval fenestra is situated behind the spine near its base (Fig. 40) 5. Ventral process of petiole not wholly slender spiniform, but more sub- triangular, irregularly tapered, and much broader, wholly containing the oval fenestra (Figs. 38, 39) 6. 5. Gastric dorsum, especially of the second segment, covered densely with fine whitish pubescence which nearly obscures the finely reticulate- striolate sculpture of much of its surface; color black, with brown legs, mandibles and antennae, becoming yellowish toward the apices pulchella Emery Gastric dorsum very smooth and shining, except for some feeble striation across the postpetiole anteriorly ; pubescence very sparse and inconspicu- ous; color mahogany or piceous, the gaster orange, brown or piceous; legs, antennae and mandibles yellow throughout (Fig. 40) versicolor sp. nov. 6. Petiolar node strongly compressed anteroveutrally, its ventral process narrowed to a digitiform apical portion; spaces between pronotal punc- tures not densely and finely striate (Fig. 3S) wilsoni sp. nov. Petiolar node not strongly compressed anteroventrally, its ventral process without a digitiform apical portion; spaces between pronotal punctures densely and finely striate (Fig. 39) niimcensis E. Andre RpiYTiDOPONERA ACANTPioPONEROiDES Viehmever This species was described from a single worker from New Caledonia, locality unspecified. It has not been reported since the description, but now Wilson has taken a colony on Mt. Mou, north of Paita, under a stone in an opening in cloud forest dominated by Araucaria and Podocarpus on the summit ridge below the summit, altitude probably somewhere near 1000 m. This sample fits the original description well except for the color, stated to be "dark brown, with more yellowish brown appendages and mandibles; body steely blue." Viehmeyer describes the smooth gastric dorsum well, with its cloudy blue-gray opales- cence ; Wilson notes that his samples were predominantly metallic green while alive, and these were the colors seen in the specimens as they came out of alcohol two months after capture. After two years in the dry state, they have changed to a predominantly bluish-purple over head and alitrunk; the legs are a contrasting 290 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY light reddish, as are also the mandibles and antennae. The worker of this species is unmistakable because of the postero- dorsally mucronate petiolar node, found in no other New Cale- donian species, but occurring in similar form in several Austra- lian Ehytidoponera, as well as in other ectatommine genera in both hemispheres. The male (previously undescribed) measures TL 7.1, HW 1.30 (across and including compound eyes), WL 2.41, scape L (ex- cluding basal neck) 0.27 mm. Scape only about twice as long as broad, twice as long as the first flagellar segment, and half as long as the second flagellar segment. Mesothorax robust ; notauli distinct, impressed, forming a complete "Y. " Wing venation of the complete type ; fine traces are present of first radial crossveiu in one of two male specimens examined. Hindwing (of measured specimen and one additional specimen) with 6-8 subapical and 0-3 submedian hamuli. Petiole nearly twice as long as high, feebly arched, profile rising slowly from front to gently rounded summit near pos- terior end ; ventral process lacking (ventral process of worker is of the versicolor and pulchella type, Fig. 40). Genital capsule a little more elongate than in some other species of the genus; parameres and hypopygium (IX sternite) rather more slender than average, the latter rounded apieally. Penis valves obliquely truncate at apex, the corners rounded oif. Integument predominantly smooth, moderately to strongly shining, on the body overlain by weak bluish opalescence ; certain areas slightly roughened or vaguely punctulate, such as the dorsum of head mesad of compound eyes. Propodeum and petiole more definitely roughened, the latter opaque, and with a few superimposed, separated rugae. Pilosity of abundant fine light brown hairs, more tapered and mostl}" more curved than in the numeensis male. Pubescence dense, appressed to decumbent, yellowish-gray and conspicuous over most of body and appendages. General body color dark brown (with faint opalescence), head darkest; mandibles, legs, and antennae ligliter; genital capsule yellowish. BROWN : ANT TRIBE ECTATOMMINI 291 Rhytidoponera pulchella Emery Emery (1914) noted the variation in size and gastric sculptnre shown by this species, evident in the extensive series before me. The largest of these are as large as large numecnsis, while the smallest are slightly smaller than the smallest versicolor, so that pulchella just about matches the total size variation of the other three small New Caledonian species taken together. This species is readily distinguished by the shape of its ventral petiolar process, which is like that of versicolor (Fig. 40), by the conspicuous whitish pubescence of the second gastric seg- ment, by the fine, centrally obsolescent sculpture of this same segment, which is consequently rather strongly shining under its vestiture, and by the more or less yellowish apices of the antennae and tarsi, and edges of the mandibles, which contrast with the black of the body. Both Wilson and Krauss found this form rather common in relatively moist forested localities all over southern New Caledonia. It nests commonly under stones in the earth. Rhytidoponera numeensis E. Andre An average worker of this species measures 5.5 mm. TL ; the IIL is 1.28, nW 1.15, WL 1.82 mm.; CI 90, ex Mt. Mou, No. 147. This dark brown species is the commonest and ecologically most tolerant of the smaller Rhytidoponera of the island. It is easily recognized by its light reddish legs and antennae, contrasting with the darker body, by the robust build, by the striation mixed with the coarse punctures of the alitrunk, and by the shape of the ventral process of the petiolar node (Fig. 39). Wilson found numeensis foraging on the ground and on foliage, together Avith pulchella, and, in forested areas in the hills, with wilsoni. It was not taken in southern New Caledonia above 800 m., and the majority of collections came from below 500 m. This species nests in rotten wood where it is available ; Wilson's nests were taken in rotten branches lying on the forest floor, as well as in larger, moss-covered logs. In shaded pastures and other disturbed areas, more nests of this species seem to occur under rocks in the ground. Colonies were usually small, with 100 workers or so, but one or two larger nests were seen. In one nest, the larvae were seen feeding upon a small talitrid amphipod. 292 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Rhytidoponera atropurpurea Emery I have not seen this species, and Wilson 's notes do not mention the type material. Emery's description compares it with "7?/;. numeensis" chiefly ; as mentioned already, he was really referring to fulgens, bnt cnriously enough he also mentions fulgcns by name in the description. At any rate, the picture emerges of a large form similar to fulgens, but with a superficial, finely reticu- late sculpture overlying the coarser sculpture, and replacing the striation of the gastric dorsum so characteristic of fulgens. The color is cited as "violaceous black; mandibles, funiculi, legs and apex of gaster deep brown." The male is stated to be like that of fulgens, except that the gaster is subopaque, finely and superficially reticulate. Known only from Ouedjo I., near Hienghiene, in the northern part of the main island. Rhytidoponera versicolor sp. nov. (Figure 40) Holotype worker: TL i.o, IIL 1.04, IIAV (excluding eyes) 0.!)2 (CI 89), AVL 1.38, scape L 0.90, max. diameter eye 0.22 mm. Similar to numeensis, but smaller and less robust. Occipital border of head feebly and broadly concave in full-face view, but occipital angles gently rounded ; as seen from the side, the posteroventral corners a bit better developed and more nearly rectangular. Clypeal apron distinct, translucent, forming a bluntly rounded angle in the middle. Alitrunk lower and more slender, in profile forming a single convexity, with only the pro- podeal declivity breaking the outline slightly as it falls off steeply from the dorsum. Petiolar node (Fig. 40) much lower than in numeensis (Fig. 39), and with the ventral process very long and slender, like that of pulchella. Seen from above, the node is only slightly broader than long (much broader than long in numeensis) . Head, alitrunk and petiole coarsely reticulopunctate through- out; a few of the dorsomedian rugules on the head forming longitudinal costulae (including a median carinula), but the sculpture otherwise without definite orientation, and without the interspersed striation so widespread in numeensis. Gaster smooth and shining, witli scattered small piligerous punctures; BROWN : ANT TRIBE ECTATOMMINI 293 postpetiole with a few indistinct striae arching from the sides across the anterior face. Erect pilosity much as in numeensis, the hairs fine, whitish, abundant on all parts, uneven in length. Appressed pubescence extremely sparse except on extremities of antennae and legs. Head, alitrunk and petiole deep mahogany (nearly black to naked eye). Gaster bright orange-yellow. Legs, antennae and mandibles clear j^ellow. liolotype [MCZ] from near the Dumbea Road, just beloAv Montague des Sources, New Caledonia. This locality is mixed forest, mainly angiosperm evergreens, but with large Arancaria and Agathis prominent in the upper story, surrounding the head- waters of the Dumbea River; altitude ca. 800 m. (E. 0. AVilson leg., XII-17-1954, No. 185). The holotype and several paratype workers were collected from the vegetation at the forest border during daylight hours. Another collection from Montague des Sources (Wilson, No. 169) was made on tlie same day higher up (ca. 1000 m.) in second-growth Arancarki forest, where the workers were taken foraging on a small angiosperm shrub in the understory during midafternoon. Variation: The workers from the upper site (No. 169) are much darker in color ; the head and alitrunk are dead black or very nearly so, while the petiole and gaster are dark brown above (blackish to naked eye), medium brown below. This darker coloration is approached by certain specimens from the lower site (No. 185) with brownish-orange or brown gaster. All speci- mens maintain the clear yellow mandibles, antennae and legs. h\ several specimens, particularly in the upper-site series, the petiolar node is a little more rounded above, and the sculpture of its anterior face varies more or less toward a smooth, shining surface crossed by weak transverse rugae or costulae, becoming concentric or longitudinal on the summit. These same specimens often tend to have the obsolescent striation of the postpetiole a little better developed and extending back over as much as % of the surface, which is consequently often not so strongly shining as in the holotype. A few individuals have the declivity of the propodeum meeting the dorsum through a distinct but obtuse angle, and in these, the declivity may l)e weakly concave as seen in side view. One specimen has a feeble impression or step in 294 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY the region of the metanotum. Variation in size and in the chief proportions is very slight ; the darker specimens from the upper site average a trifle smaller. The paratypes are to be deposited in MCZ, USNM, and one or more Australian and New Cale- donian collections. Rhytidoponera wilsoni sp. nov. (Figure 38) Holotype worker: TL 5.2, HL 1.25, HW (excluding eyes) 1.06 (Ci 87) WL 1.73, scape L 1.12, max. diameter eye 0.26 mm. Very similar to numeensis, from which it differs principally in having a notably more strongly anteroposteriorly compressed node, with a different ventral petiolar process (Fig. 38) ; the posterior face of the node is concave as seen from the side. The head of wilsoni has a more broadly concave occipital margin (straighter in some views) and slightly more sharply rounded occipital corners than in numeensis. The sculpture differs in sev- eral details ; generally speaking, the punctures are coarser and more closely contiguous over head, alitrunk and gaster, and the ridges between tend to form coarse, rather indefinite rugules (longitudinal on head, diagonal on sides of alitrunk, transverse on propodeum and anterior face of petiolar node) in place of the fine, dense striation of these same areas in numeensis. Postpetiole rather finely arched-striate, passing into a posteromedian area of finer, concentric striae that are interrupted by and interspersed with fine, indistinct punctulation. Coarse piligerous punctures of this segment very shallow, indistinct, and not so clearly elon- gate as the corresponding ones in numeensis. Second gastric segment very finely and densely striate, the pattern arched over a posteromedian weakly shining area of very superficial elliptical- concentric striation (longitudinal axis) that is very finely inter- rupted or coriaceous (clearcut, uniform, and predominantly longitudinal striation in nuuiee^isis) . Coarse piligerous punctures of this segment small and inconspicuous, much smaller than the corresponding ones of numeensis. Erect pilosity and appressed pubescence fairly abundant but not dense, uneven in length, the pubescence conspicuous at all only on the gaster and appendages. Color piceous, the head, BROWN : ANT TRIBE ECTATOMMINI 295 petiole and gaster lighter and a bit more reddish than the ali- trunk. To the naked eye, the bodj^ appears very dark, nearly or quite black. Legs, mandibles and antennae orange-brown. Coloration in general much like that of numeensis. Ilulotype [MCZ] a stray diurnal forager taken on foliage at 180-400 m. on j\It. Mou, north of Pa'ita, New Caledonia (E. 0. Wilson leg., XII-10-1954, No. 110). Paratype workers were taken with the holotype and at Le Chapeau Gendarme, east of Yahoue, and at Ciu, near Mt. Cauala (Wilson Nos. 73, 74, 80, 84, 96 and without numbers). These were taken foraging during the daylight hours on foliage and over rocks on the forest floor. One nest was found beneath a stone ; other workers came from leaf litter berlesates. The Ciu sample was very dark, and had the legs dark brown. The occipital border varied in degree of con- cavity. Paratypes in MCZ, USNM and elsewhere. [34] Among Ectatomma species, the closest to aztecuni (judging from the holotype, kindly sent by Dott. Delfa Guigiia) is ruidum. The color is dark, as in ruidum, but the node of aztecum is slightly thicker as seen from the side. The sculpture of gastric segment I (postpetiole) is slightly different in aztecum; the costulae are arched transA^ersely across the anterior descending face of the segment, but otherwise are longitudinal, close and at most feebly undulant in a few places. The outstanding distinction, however, is the extreme development of the erect pilosity; this is rather short, but present in great abundance on most body and appendage surfaces. The species has never been taken a second time ; its type locality is ' ' Michoacan. ' ' Since this form is very close to the highly variable ruidum, it would be desirable to have additional material from southwestern Mexico in order to see exactly what the relationship is between these two species. [35] The type locality of E. coiifine apparently lies somewhere in the present Colombia or Panama. Mayr's description is very sketchy, and the only specimen I have seen that fits it fairly well is a unique worker (Fig. 2) from Tela, Honduras (D. M. Bates leg.). This worker was found mixed with ruidum and morgani in the Wheeler Collection, and at first glance it appears to be intermediate between these two species. However, it differs from 296 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY both ill the disproportionately strong development of the median pronotai eminence; this is high, somewhat compressed laterally, and forms a blunt angle as seen from the side. The lateral pro- notai eminences are rather small, but their size is probably some- what variable in this as in other species. The greatest eye diam- eter and the apical antennal segment are very nearly precisely the same in length, and the petiolar node is thicker than that of ruidum as seen from the side. The sculpture of the second gastric segment is composed of fine, sericeous striolation, as in ruidum. Whether this identification is correct can only be determined after the review of more material in comparison with the type in the Mayr Collection in the Vienna Museum. It is not impos- sible that confine will finalh- prove to be a geographical variant of the species represented in the Amazon Basin by morgani [38]. [36] E. edcnfatum has been confused with muticum several times (Forel, l!jl2b :31), but the two are easily separated by eye size (see key) and sculpture of gastric segment II. In edentatmn, the tergum of the second segment is very regularly and very liiiely striolate in a pattern of varying direction, and in general is sericeous-opaque. Jn muticum, the same segment has a much more indefinite and looser transverse striation that is virtually eifaced over a large area disead ; the surface in general is defi- nitely shining, even glassy in the center and between striae. Forel 's arrangement of 1912 depends on characters, such as antennal proportions and metallic luster of integument, that seem too variable (and often allometric in expression) to have more than local or individual significance ; neither Forel nor Santschi seems to have expended much effort in trying to define limits of variation for these forms. E. macdonaghi is poorly de- scribed as a race of edentatum, of which species it may be an ergatoid or pathological intercaste. E. morgani [38] maj^ be a northern variant of edentatum. [37] The names strigosum, permagnum, confusa and aerea seem to me to represent slight local variants of one species, widespread ill southern Brazil and Bolivia. The name strigosum unfortu- nately must fall as a primary homonym of Rhytidoponera stri- gosa (Emery), originally described in Ectatomma, and Forel's BROWN : ANT TRIBE ECTATOMMINI 297 name permagna (emended to the proper form permagnum), being next available, is raised from varietal to species rank to take its place. Since permagnum and lugens are separated only by minor and variable characters and are allopatric as at present known, it is possible that these two names represent extreme {geographical populations of the same species. Collections from the region just south of the Amazon are needed to decide whether intergradation is completed in this area. In addition to the sculptural differences (see key, couplet 1), which may only reflect character displacement against the closely related species opaci- ventre in the south, the samples of lugens I have seen have nar- rower heads and are generally more slender OA'crall than are the permagnum examined from Bolivia and southeastern Brazil. [38] E. morgani is uncommon in collections, and has remained enigmatic due to its almost universal confusion with the similar )-uidi(m. MCZ has stray workers taken near Port-of-Spain, Trini- dad (R. Thaxter leg., beating foliage) and in British Cluiana : source of R. Essequibo (Ogilvie leg.) ; Kartabo vie. (W. M. Wheeler, H. 0. Lang leg.). Constant characters distinguishing this species from ruichnn (see key) are the proportionately longer and narrower head, somewhat more convex behind as seen in full-face view ; eyes smaller and placed a little farther from the posterior border ; scapes and legs a trifle longer ; petiole thicker from front to rear, and not biconcave in side view, as in ruidum; fine striolation of second gastric segment less distinct basad, where it merges with some fine, dense, irregular reticulo-punctu- lation or broken striation (but this last character is very vari- able). There is no doubt that uiorgani is really very closely related to cdentatum, and the two forms may well be geographical seg- ments of the same species. A worker sent b}^ Father Borgmeier from the Mato Grosso is intermediate in some respects between my Guiana-Trinidad morgani samples and the few specimens of edentatum available from the State of Sao Paulo (W. AV. Kempf leg.). The material available to me is not sufficient as a basis for formal action on this probable synonymy at the present time. 298 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY [39] E. opaciventre is a large species of the La Plata-Parana drainage. It apparently does best in districts that are arid, or at least that have prolonged annual dry spells. The color is very variable, ranging from reddish through a series of bicolored forms to piceous ; the variety concolor is a part of this variation. Shape of head and complete lack of transverse rugules on the postpetiolar dorsum distinguish this species from permagnum. [40] E. planidenii is a puzzling species very close to quadridens (see key, couplet 4), but both of these are present at some locali- ties in Brazil, without known intergradation (Agudos, S. Paulo State, leg. W. W. Kempf). A small, light female from central Brazil owned by Father Borgmeier, however, appears transitional to quadridens, and it is not beyond possibility that planidens is merely a morph of the more familiar species. Wilson (MS notes) has located the probable type of E. quadridens in the Museum d'Histoire Naturelle in Paris; it is a specimen from the Bosc Collection, now in the general collection. [41] E. muticum has been confused with E. edentatum; the characters are discussed under the latter species [36]. The rec- ords and specimens of muticum I have seen indicate that this species is restricted to the dry eastern bulge of Brazil, in Ceara and neighboring states. There is only one exception to this pat- tern, and this exception is so far off that it is difficult to credit it. I refer to a single worker in the MCZ bearing the label "Acapulco, Mex./Fredk. Knab." and which is identical, so far as I can tell, to specimens from northeastern Brazil collected by W. M. Mann, now in the same museum. Acapulco is close to Michoacan, the type locality of aztecum, but this circumstance is probably only a coincidence. Until such time as the Mexican record for muticum is confirmed by further collections, the oc- currence of the species in that country should be regarded as doubtful. [42] E. tuhercidatum varies widely in color and sculpture, as pointed out by Weber (1946). The greatest variation seems to focus in northern South America and the Amazon Basin, where hardly two nest series can be found that are alike. Especially BROWN : ANT TRIBE ECTATOMMINI 299 striking is the brown variant, contrasting with the more familiar yellowish types from farther north and south, as well as from this same area in the middle of the range. E. acrista differs modestly from tuberculatum in having the occipital margin of the head more ronnded as seen in full-face view ; I have specimens of this kind from the Parana River, Paraguay (Fiebrig leg.). Other samples from central South America may represent transi- tions to acrista from tuberculatum, but at present the evidence is not as complete as it should be to dismiss acrista as a southern peripheral variant of tuberculatum. Figures 41 and 42. G-namptogenys spp., workers. Fig. 41, G. grammodes sp. nov., paratype, full-face view of head. Fig. 42, G. mecotyle sp. nov., paratype, dorsal view of petiolar node. [43] G. acuminata samples in the MCZ : Campinas, Goias, Brazil (Schatzmaier leg.) ; Santa Helena, Bolivia (W. M. Mann leg.) ; Bartica District, British Guiana (A. E. Emerson leg.). Varies from ferruginous to black in color; very similar to G. sulcata, but differing in the more striking posterodorsal production of the node. [44] G. annulata. Samples in the MCZ indicate a very wide range for this species in the Amazon-Orinoco drainage, west as far as Ecuador, Peru and Bolivia, and north to Costa Rica and Honduras. I have seen specimens from Rio de Janeiro State, and the species is reported from Santa Caterina. 300 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY The male is a little shorter than workers from the same nest, piceous with brown legs and antennae, smooth and shining except for rugose propodeum ; petiole finely rugulose, shining ; front of head finely punctulate, shining. Notauli complete, well- marked. Forewing with "complete" venation, i.e., Rsf2 and 3 and Mf2 and all present. [45] Samples of G. aculeaticoxae were examined from Kama- kusa, British Guiana (H. 0. Lang leg.), Barro Colorado Island, Panama Canal Zone (J. Zetek leg.), and from Bolivia: lower Rio Madidi and Blancaflor and Huachi on the Rio Beni (W. M. Mann leg.). [46] In a recent paper (Brown 1957), several forms were dealt with under Ilolcoponera. Var. antillana Santschi was placed as a synonym of striatula Mayr [79], and the nominal species or subspecies satzgeri Forel, spuriuni Forel and foreli Santschi were synonymized under simplex Emery. The new species mina and acuta were described, and Rhopalopone relicta Mann was trans- ferred to Holcoponera. There are some new records of acuta from Peru : Tingo Maria, Monson Valley ; and Colonia Perene, 18 miles northeast of La Merced, Junin (E. S. Ross and E. I. Schlinger leg.). [47] In a review of the Stictoponera coxalis group (Brown, 1954c) all of the species here indicated were discussed, and synonymy proposed for some of them. S. hiroi Emery (Fig. 17) has been taken by E. 0. Wilson at Bisianumu, near Sogeri, Papua, at about 500 m. A single dealate female (No. 654) and a small colony with callow winged males and females (No. 675) were taken in moist rotting logs in rain forest. [48] G. panda is very close to G. taivanensis, but the adult color is reddish-brown (blackish in taivanensis), and the postpetiole of panda is much more closely, finely and opaquely sculptured, with the interfoveolar spaces forming distinct oblique costulae, so that the surface appears in some lights to be punctate-striate. In panda, the sculpture of the second segment is nearly closed in across the middle, whereas in taivanensis nearly the whole BROWN : ANT TRIBE ECTATOMMINI 301 dorsal-diseal area is smooth and shining, with only the coarse piligerous punctures. I have seen additional specimens of panda from Muping- (2000 m.), on the Ya An to Mou Kung track, Sikang Province, about 90-100 miles west of Chengtu, western China (D. C. Graham leg.) ; this locality is not far airline from the type locality near Kuanhsien, Szechuan. The petiole of panda is shown in side view in Figure 18. [49] 1 have examined specimens of G. hispinosa taken by G. C. Wheeler at Changuinola district, Bocas del Toro, Panama, and by W. M. ]\Iann in Costa Rica at Zent, Colombiana Farm and Hamburg Farm, Santa Clara Province. The Hamburg Farm series includes a male : TL 8.2, HL without cervical rim of occiput 1.28, L head with closed mandibles 1.73, HAV with compound eyes 1.33, greatest diameter of compound eye 0.52, WL 2.61, petiole L 0.80 mm. Antenna : scape L 0.35, L funicular segment 1 0.19, fun. II 0.55, fun. Ill 0.54 mm. L forewing ca. 5.5 mm. Mandibles triangular, distinctly dentate. Ocelli large and clear, set close together on a darkened callus ; distance between anterior and posterolaterals less than an ocellar diameter, distance be- tween posterolaterals slightly greater than an ocellar diameter. Xotauli and parapsidal furrows very distinct, complete. Maxil- lary palpi with 5, labials with 3 segments. Petiolar node long and low, nearly twice as long as high, with a subrectangular ventral process in front below. AVings as in other larger Gnamp- togenys, Mf2 short or completely contracted, Rsf2*3 present. Body and appendages shining, completely smooth except for mandibles and clypeus, Avhich are feebly and obscurely longitudinally striate, and propodeum, which bears a bold pattern of sharp rugae. Color ferruginous yellow; gaster yellowish-tan. The worker of this species has a palpal formula of 3, 2. The "Wheelers have described and figured the larvae (1952a, pp. 132-133, pi. 5, figs. 1-5). [50] G. alfaroi. I have examined a short series (USNM, MCZ) from Turrialba, Costa Rica (0. L. Cartwright leg.) of this very rare, local and interesting species. It is a rather aberrant mem- ber of the mordax group, large in size, black in color, long-headed, with short, thick mandibles. In many respects, it resembles 302 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY species of the genus Cylindromurmex (Subfamily Cerapachyi- iiae?), and the relationships between Cylindromyrmex and Gnamptogenys are worth investigation on this account. [51] I was able to compare the female type of G. haenschi with workers from the lower Rio Madidi, Bolivia (W. M. Mann leg.) ; the workers are very similar in their robust build and very fine costulation, but they are decidedly smaller than the female. Like the type, their color is brownish red. In the worker, the petiolar node is compressed anteroposteriorlj^ but is not quite twice as broad as long. A female from Tingo Maria, Peru (E. S. Ross and E. I. Schlinger leg.) is similar to the type, but is much darker, nearly black in color. [52] G. Jiartmani remains known only from the unique type worker, from Iluntsville, Texas, far outside the known range of other species of Gnaniptogeriys. Creighton (1950, p. 36) gave it as his opinion that the tarsal claws of Jxnrimani were simple, a condition he "regularly encountered in . . . subgenus Gnampto- genys." However, careful examination of the hartmani holotype, as well as all available species of Gnamptogenys and Parecta- tonima in the old sense, reveals that all have an extra tooth on each of the six pairs of tarsal claws. Among these groups, simple tarsal claws must be exceptional, if they occur at all. [53] The dealate female type of epinotalis has been examined and compared with workers of the same species (Fig. 20) taken by Wilson at Ebabaang, ca. 1400 m. altitude, on the Mongi Watershed, Huon Peninsula, New Guinea, in irregular galleries in the earth under a stone. The Ebabaang locality is in mountain rain forest. The ants resembled Ponera in life. G. epinotalis is very close to G. luzonensis Wheeler, differing mainly in details of sculpture. The extra teeth on the posterior tarsal claws of epinotalis and luzonensis are very minute and reclinate, and are situated close to the base of each claw. They can be seen only at magnifications over 120 X, and then only under the most favorable conditions. Dr. Chapman has turned over to me some workers of luzonensis taken by D. Empeso in the vicinity of BROWN : ANT TRIBE ECTATOMMINI 303 Dumaguete, Negros Oriental, Philippiue Islands. These two species are among the smallest Gnamptogenys ; and the eyes are much reduced in the workers. Among the other small species of the Indo-Melanesian area, formerly placed in Rhopaloponc, I have seen types of malaensis (Mann) and reliable specimens of dammermani (Wheeler) de- termined by Wheeler himself from Buitenzorg, now Bogor, Java (Dammerman leg.), as well as a series from the Cuernos Mts., near Dumaguete, Negros Oriental, Philippines, at about 1800 feet altitude (J. W. Chapman leg.). Wilson has compared these with the types of crihrata (Emery) and diehli (Forel), as well as with the very different major (Emery), on deposit at Genoa and Geneva. Wilson took crihrata twice at the lower Busu River, near Lae, New Guinea, in lowland rain forest leaf litter. The worker does have vestiges of compound eyes, but these are minute. [54] Dr. Wilson has compared worker specimens from the MCZ Collection, taken at Kartabo, British Guiana (A. E. Emerson leg.), Avith the female type of exarata, and he feels that they are conspecific. He notes that the type female has a head length of about 0.93 mm., or less than females associated with the Kartabo workers, but a little larger than these workers themselves. A Kartabo female has HL 1.12 mm., compound eye L 0.27 mm. The compound eye L of the type female was about 0.23 mm. Wilson's measurements were, however, not made under ideal conditions. The type has about 30 costulae between the compound eyes, with a possible error of about ±1. In the females of the Kartabo series, the costulation of the petiolar dorsum varies from con- centric to longitudinal ; in the type, it is longitudinal. G. exarata is a member of the mordax group, related to both continua and interrupta, but it does not compare with the types of these two species [55, 56]. In exarata, the anterior clypeal border is concave in the middle. [55] The worker type of G. interrupta borrowed from the Vienna ^luseum has HL 1.12, HW 0.95 mm. The eyes are very small, 304 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY with about 12 pigmented facets, siirroimded by a ring of un- pigmented facets, colored like the surrounding integument. The pigmented part of the eye is slightly smaller than the greatest scape thickness, but if the unpigmented part is included, the eye diameter would more or less equal the scape thickness. The sculpture is fine, 50 ±3 costulae between the compound eyes. Second gastric segment smooth and shining, with a few coarse punctures. Color rich ferruginous red ; legs and antennae more yellowish. ' ' Patr. ? M. Dresd./Sudamerika. ' ' Like continua, but larger and with finer sculpture. (Lectotype.) Specimens from Lombardia, Honduras (W. M. Mann leg.) agree well with the type, but have coarser costulation. The size variation in this series is marked. [56] G. continua is the smallest known species of the mordax group ; lectotype HL 0.74, HW 0.58 mm. ; about 35 costulae between the compound eyes; eyes themselves small, only about 5-6 facets, their diameter less than the greatest scape thickness. Sides of head subparallel ; greatest width at about the anterior third. Clypeus as in mo^rlax. Metanotal groove shallow, but dis- tinct. Propodeal angles blunt, sublaminate, feebly overhanging the concave borders of the declivity. Petiole approximately as broad as long, broadest behind, longitudinally costulate ; anterior face of node distinct, its sculpture effaced. Second gastric seg- ment rather finely longitudinally costulate above, but smooth and shining on the sides behind. I have seen samples referred to continua from the following localities : Las Hamacas, near Santiago Tuxtla, Veracruz, Mexico (E. 0. Wilson leg.) ; nest in a piece of rotten wood in leaf litter, tropical evergreen forest. Boquete, Chiriqui Mts., Panama (F. M. Gaige). Barro Colorado Island in the Panama Canal Zone, April, (A. E. Emerson, No. 110). Mandeville, Jamaica (A. Wight). Tingo Maria, Monson Valley, Peru (E. S. Ross and E. I. Schlin- ger leg.). Variation among samples extends to density of costula- tion, size, extent of shining area on second gastric segment, and depth of pigmentation; some series are deep reddish, others are almost black. The Las Hamacas series was most like the lectotype ("Brasilieu") among my samples, even though it was on the opposite extreme of the range. BROWN : ANT TRIBE ECTATOMMINI 305 The Barro Colorado series includes a male, which is a little shorter than the accompanying workers, is dark piceous in color with yellowish antennae, legs and genitalia, and which has deep and complete notauli. The integument is smooth and shining except for the cephalic dorsum, propodeum and petiole, which are finely and densely punctulate and more or less opaque. The second and third abscissae of Rs are lacking. Some of the Panamanian specimens have additional suturation on the alitruncal dorsum and otherwise betray themselves as ergatoid females or other worker-female intermediates. Sant- schi's Panama variety appears to be about the same as the Barro Colorado sample from the brief characterization ; there seems no need to maintain his name panamensis. The Jamaican record of coniinua indicates an introduction, very probably made within historical times. This island has received many introduced ants from South and Central America. [57] The three varieties of concinna were described chiefly on sculptural differences now recognized as individual variations that can be expected even in single nest series. This is confirmed for the type of semicircularis, matched with a worker from one variable nest series in the MCZ. The var. romani had as an additional character the color of the gaster, "entirely black." Whether this blackness is caused by decomposed gastric contents or some secondary influence, I do not know, but it does not seem likely that it is a character of specific value. [58] Gnamptogenys chapmani sp. nov. (Figures 19, 44) Ilolotype worker: TL 3.4, HL 0.71, HW 0.56 (CI 79), L head with closed mandibles 0.93, scape L 0.47, greatest diameter of eye 0.17, WL 1.04, petiole L 0.38 mm. Head as shown in Figure 44. Extremities of occipital angles rather sharply rounded and laterally compressed, but in side view not forming distinct "ears" as in costata and menadensis. Masticatory borders of mandibles feebly crenulate, almost .straight. Alitrunk subcylindrical, only slightly narrower than head, tapering only slightly from front to rear. Humeri bluntly 306 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY subangulate seen from above ; dorsal sutures of alitrunk oblit- erated. In lateral view, dorsal profile gently convex, almost straight in central portion, falling off through a sharper curve to the almost vertical upper part of the propodeal declivity (Fig. 19), which is marginate above and on the sides, but Figures 43 and 44. G^iamptogenys spp., Avorkers, full-face view of head. Fig. 43, G. Icalahit sp. uov., pai'atyi:ie. Fig. 44, G. cliapmani sp. nov., liolo- type. there are no propodeal teeth. The node is also shown in the figure ; seen from above, it is widest behind, and slightly wider than long. Gaster slender, the first segment (postpetiole) trun- cate in front, slightly wider than long. Second segment longer than the first, but slightly narrower, strongly downcurved. Apical segments retractile. Integument smooth and shining, with abundant round foveolae spaced on dorsum of head, becoming more crowded and finally contiguous on sides and gula ; numerous but separated on alitrunk dorsum, very few on sides; few on sides of petiolar node and succeeding segment, becoming fewer and obsolescent on dorsal surfaces of these segments. Second and apical gastric segments, undersurface of first, propodeal declivity, sides of alitrunk, scapes, legs and mandibles predominantly smooth and shining, with occasional coarse punctures, especially on mandibles. BROWN : ANT TRIBE ECTATOMMINI 307 Fairly abundant erect fine hairs of uneven length over most surfaces of body, becoming more frequently reclinate on ap- pendages and mandibles. Color brownish red ; mandibles, legs and antennae yellowish. Ilolotype [MCZ] taken by Dr. J. W. Chapman near his vaca- tion home. Camp Lookout (altitude about 600 m.), in the Cuernos Mts., near Dumaguete, Negros Oriental, Philippine Islands, on March 19, 1924. Paratype workers : two workers taken with the holotype ; one worker taken with the female paratype (see below) at or near tlie type locality, on April 27, 1924 (J. W. Chapman leg.) ; two workers from Romblon Island, Philippines, April 5, 1924 (L. Morato leg.). Dr. Chapman thinks that the Cuernos Mts. collec- tions came from nests in rotting wood in rain forest ravines. The paratype workers range downward in size from the holotype to one of the two specimens from Romblon : TL 3.0, HL 0.66, HW 0.48 (CI 74), WL 0.90, petiole L 0.35 mm. In smaller speci- mens, the head is disproportionately narrower, and even more noticeably so is the petiole ; in the smallest specimens, the node is distinctly longer than broad ; in these same specimens also, the foveolae of the sculpture tend to be smaller and farther apart. The shape of the occipital concavity varies slightly by indi- viduals. In some workers and the female, the margins of the I)ropodeal declivity form a distinct angle with the dorsum, in some the angle being almost like a blunt tooth or tubercle. Female paratype, dealate : Size and proportions much as in larger workers, but alitrunk a little more robust and with flight sclerites differentiated in normal (but minimal) (jueen fashion; wing stumps present. Compound eyes only very slightly larger than in worker of same size ; ocelli modest in size, but distinct. G. chapniani is close to G. laevior Forel, but is much smaller and much lighter in (full adult) color than any of the laevior specimens I have seen; also, laevior has the node slightly longer than broad, a circumstance that is reached only in the smaller workers of cJiapmani. 308 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY [59] Gnamptogenys kalabit sp. nov. (Figure 43) Holotype worker: TL 5.0, HL 1.06, HW 0.84 (CI 79), L head with closed mandibles 1.37, scape L 0.76, greatest diameter of eye 0.24, WL 1.47, petiole L 0.62 mm. Head as shown in Figure 43; note the almost perfectly straight and parallel sides and the rather sharp occipital angles. Seen from the side, the occipital corner is rather sharply rounded and margined by a narrow translucent border or carina, but no distinct "ear" is formed. Antennal scapes do not quite reach occipital angles, but reach the occipital border when laid straight back from their insertions. Clypeus not or very indistinctly sulcate in the middle ; mandibles denticulate. Antennal funiculus stout, median segments very short, only about half as long as broad ; last three segments form- ing an indistinct club ; only the first and apical funicular seg- ments longer than broad. Alitrunk seen from above with obtusely angled (vertically margined) humeri ; inferior border of pronotum forming a salient right angle in front of each fore coxa. Seen from the side, the alitrunk is deeper and a little more convex dorsally than in cliapmani, but the conformation of the propodeum is similar. Coxal tooth acute but subconical, with broad base. Petiolar node paniform, of the general form of costata, meyiadensis and chap- mani, slightly longer than high, with rounded dorsal surface; seen from above, node about as broad as long, narrowed an- teriorly. Ventral process with posterior border sloping back gradually from an obtuse posterior angle. Base of postpetiole subtruncate (more rounded than in cliapmani) ; postpetiole slightly broader than long; succeeding segment considerably longer and slightly narrower, strongly downcurved. Middle and hind tibiae each with a single slender, very minutely denticu- late or subpectinate spur ; all tarsal claws with a single extra strong subbasal tooth. Head covered with deep, circular, umbilicate shining-bottomed f oveolae, the ridges separating these forming a coarse reticulum ; cervical surface of occiput smooth and shining, with a few fine striae in the middle ; clypeus coarsely longitudinally striate, f oveolate along sides of triangular median lobe ; mandibles BROWN: ANT TRIBE ECTATOMMINI 309 smooth, shining, coarsely punctate. Alitrunk, petiole and sides of postpetiole with conspicuous, shining, mostly separated foveolae, still more widely separated over the postpetiolar dorsum (disc). The following areas smooth and shining : a median mesonotal- propodeal strip, metathoracic portion of sides of alitrunk, pro- podeal declivity, a small area on the nodal summit, the broad interfoveolar spaces on the postpetiolar dorsum, tergum of the second segment (except for posterior and lateral margins), and the apical segments of the gaster. Scapes and legs smooth and shining, with scattered punctures. There is also a superficial fine sculpture, characteristic of this species, found on the alitrunk, node and sides of the first two gastric seginents in those interfove- olar spaces that are not smooth and shining. This fine sculpture ranges from a fine striation to a broken striolation or even to a finely coriaceous or reticulo-punctulate condition, the extremes grading through all intermediate substriate stages. This sculp- ture is more arched-striate and weakly shining on the disc of the pronotum and on the petiolar node ; obliquely substriate or striate on the sides of the alitrunk and on the lateral and latero- ventral surfaces of the gaster, which are subopaque, especially the sides of the postpetiole ; transversely striolate along the posterior marginal band of the postpetiole. Fore coxae trans- versely striate, middle and hind coxae densely and finely punc- tulate, opaque, pubescent. Sternum of postpetiole more or less smooth and shining in the middle. Pilosity, etc. much as in chapmani. Color medium brownish-red ; antennae and legs more yellowish. Holotype [j\ICZ] one of a nest series labeled, "Kalabit Coun- try/N. Borneo/E. Mjoberg/ 3000 ft." Paratype workers : 13 workers from type nest series : TL 4.6-5.3, IIL 1.00-1.12, HW 0.79-0.90 (CI 79-80), WL 1.37-1.56 mm. Variation in form and sculpture is very slight. In some specimens, the ventral process of the petiole is a rounded lobe ; in some, the occipital concavity is a little more or less broad and rounded than in the holotype. Paratype female, dealate (from type nest series) : TL 5.7, IIL 1.08, HW 0.88 (CI 82), L head with mandibles 1.40, scape L 0.78, greatest diameter of compound eye 0.29, WL 1.73 mm. Alitrunk bulky, normal flight sclerites and wing sclerites present. 310 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Mesonotum longitudinally rugose-foveolate ; propodeal dorsum smooth and shining in the middle. Ocelli small but distinct, each with an adjacent blackened callus. Males, 2 specimens from holotype nest series, of which one was measured : TL without genital capsule 4.9, forewing length ca. 3.8, HL 0.76, HW including compound eyes 0.87, WL 1.54, petiole L 0.52 mm. Antennal segment lengths : scape 0.38, funicu- lus I 0.15, fun. II 0.22, fun. Ill 0.21 mm. Occipital margin broadly and shallowly concave in the middle as seen full-face. Mandibles finely dentate, with the clypeus finely and indistinctly striolate, the mandibles and anterior clypeal apron longitudinal- ly, remainder of clypeus transversely striolate. A strong median carina connects clypeus to anterior ocellus. Ocelli large, clear, separated by their own diameters or a little more. Compound eye large and rounded, greatest diameter 0.31 mm. Dorsum of head with distinct, circular, contiguous to subcontiguous foveo- lae ; interfoveolar ridges forming a reticulum. Pronotum sculp- ture like head, but smooth and shining in tlie middle ; scutum smooth, shining, with scattered foveolae ; notauli very distinct, complete, coarsely foveolate. Scutellum rugose-foveolate. Pro- podeum smooth, shining along anterior margin and in center of declivity; elsewhere rugulose, weakly shining. Sides of meso- thorax smooth to obscurely striolate, shining. Node smooth and shining above, foveolate laterally. Gaster and legs smooth, shin- ing, with scattered punctures. Wings with venation as in macretes male [61]; microtrichiae yellow-tan. Color piceous to black ; gaster, legs and antennae more reddish or brownish, geni- talia yellowish. Larva described and figured by G. C. and .1. Wheeler (1952a: 122-123, pi. 3, figs. 1-8) as Sfictoponera sp. This species is easily recognized by means of its rectangular head and substriate inter- foveolar sculpture, especially of the sides of the postpetiole. [60] Gnamptogenys grammodes sp. nov. (Figure 41) Holotype worker: TL 4.9, HL 1.09, HW without eyes 0.93 (CI 85), L head with closed mandibles 1.44, scape L 0.97, greatest diameter of eye 0.28, WL 1.56, petiole L 0.52 mm. BROWN: ANT TRIBE ECTATOMMINI 311 Head shape as shown in Figure 41, drawn from a paratype, except that in the holotype the occipital border is a trifle flatter and longer, and the occipital angles not quite so gently rounded to the compound eyes. Seen in side view, occipital lobe rounded, but with a small, thin, subrectangularly rounded lamina or "ear" on the curve. Alitrunk much as in the other medium- and large-sized Old World Gnamptogcnys, convex, but the humeral angles rounded, with scarcely a trace of the corners prominent in menadensis and relatives. Promesosuture marked by a faint line ; mesonotum with a narrow l^ut deep and well-defined median longitudinal sulcus; metanotal groove fairly well marked, and the alitruncal dorsum slightly impressed in this region. Propodeum convex above and rounded evenlj^ into the declivity, the latter short, feebly concave, bounded above and laterally by indistinct mar- gins, and guarded on each side hy a very small, rectangular to subacute vestige of a tooth. Coxal tooth slender, tapered to a blunt point. All tarsal claws with Avell-developed subl)asal teeth. Petiolar node paniform, shaped much as in the menadensis group, but with the ventral process reduced to a very small anterior tooth, directed obliquely downward and forward ; node seen from the side a little longer than high ; seen from above just about as long as it is broad; greatest breadth at about the posterior third. Gaster much as in menadensis ; second segment perhaps a little shorter relative to first, but still long and strongly downcurvecl. Apical borders of mandibles finely denticulate ; shining, striate and with scattered punctures. Clypeus shining, with large punc- tures behind and short longitudinal rugulae in front ; bisected by a narrow, deep longitudinal sulcus with shining bottom. Head with circular, shining-bottomed foveolae numerous, smaller in the median dorsal region, where they are mixed with longi- tudinal striation ; larger and contiguous on the sides and beneath. Alitrunk covered with large, circular, shining foveolae, mostly contiguous above, becoming smaller and separated on the sides ; in the metanotal area, the foveolae are mixed with some indistinct transverse striation. Interfoveal spaces here, especially a postero- median triangular area on the pronotum, smooth and shining, as is also the upper part of the propodeal declivity. Metapleura 312 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY and lower declivity with a few coarse horizontal striae. Node of petiole smooth and shining, with a few separated, shallow foveo- lae scattered over its surface ; a few indistinct transverse rugulae or coarse striae across the lower anterior face of node. First gastric segment rather irregularly costulate or coarsely striate, with coarse, elongate punctures interspersed, transversely, across the basal third, longitudinally over the remainder of the dorsum, becoming a little smoother along the lateral margins. Sternum of first segment smoother, shining, loosely rugulose in a transverse direction and with large shallow foveolae. Second segment mostly smooth and shining discad, with small scattered punctures, al- though this smooth area is bordered along both side margins and in the posterior corners by short costal remnants with large punctures; along the anterior border there is a narrow band of transverse striation, followed by a brief zone of longitudinal costulation with coarse punctures or foveolae, extending about ^ to 1/4 the length of the segment. The sternum of the second segment is rugoso-punctate. Apical segments retracted in holo- type, but these are smooth and shining in a paratype worker. Coxae of middle and hind legs finely punctulate, subopaque, as are also the antennal funiculi and the tarsi. Anterior coxae finely and closely striate horizontally, shining. Femora, tibiae and scapes smooth and shining, the scapes with one or two fine longi- tudinal rugae on inner and outer margins. Body generally with abundant long, tapered, erect hairs, mostly 0.15-0.20 mm. long, becoming shorter, more nearly decum- bent and denser on the extremities, including a dense brush on the gastric apex. Color reddish piceous, almost black to the naked eye ; antennae and legs light red. Holotype [MCZ] a worker from Bisianumu, near Sogeri, Papua, at about 500 m. altitude, from a small colony or colony fragment nesting in a piece of rotten wood on the forest floor (E. 0. Wilson leg.. No. 671, March 15-20, 1955). Paratypes consist of 8 additional workers from collection No. 671 with the holotype, plus a stray worker, No. 655, taken from a rotten log in which lived a dealate female of G. macretes as well as colonies of Phei- dole and Cardiocondyla, at the same locality as the holotype nest series. Size variation was relatively slight: HL 1.04-1.12, TL BROWN : ANT TRIBE ECTATOMMINI 313 4.7-5.1 mm. In some specimens, the occipital border is more evenly rounded (Fig. 41) than in the holotype, the clypeus is more sharply striate, even within the sulcus, and the propodeal teeth may be either more acute and more dorsally directed, or smaller and more obtuse, scarcely distinguishable from the sculp- ture. The transverse costulation of the first gastric segment may extend back over nearly half of the tergum of the segment, and in some specimens there are rather wide, shining spaces between some costulae. The distinctness and extent of the bordering remnant sculpture on the second gastric segment is also very variable, in some examples being almost completely obsolete, so that the entire tergum is smooth and shining, with only the scat- tered piligerous punctures to mar the surface. The color of the callows is dull yellowy and all intergradations of pigmentation occur in nest series No. 671, from yellow through brownish-red to nearly black. The legs are usually conspicuously lighter and more reddish or yellowish than the rest of the body. The female and male are unknown. [61] Gnamptogenys macretes sp. nov. Holotype worker: TL 5.3, HL 1.25, HW 1.07 (CI 86), L head with closed mandibles 1.67, scape L 1.20, greatest diameter of eye 0.31, WL 1.90, petiole L 0.62 mm. Very similar to grammodes, but larger overall and with a finer, more regular and more extensive striate or costulate com- ponent of the sculpture. Head shape as in grammodes, but the occipital border a little more broadly transverse (more truncate) and with the concave section in the middle broader and shallow. Humeri more distinctly angulate as seen from above, rectangular. Ventral process of petiole broader and longer, averaging larger than the coxal tooth of the same specimen. Sculptural differences: Foveolae of head averaging a trifle smaller, the spaces between longitudinally striate right out to the sides, curving above and behind the eyes and then running transversely across the gula. Dorsum of pronotum coarsely striate longitudinally between foveolae on its posterior half, especially in the middle, where a narrow sulcus is all that repre- sents the broader smooth triangular space of grammodes. Meso- notum with interfoveolar rugae converging posteriad, transverse 314 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY along posterior margin, so as to form a U. Median sulcus here obscure, partitioned into a row of coarse foveolae. Sides of alitrunk sculptured much as in grammodes; propodeal declivity very short, smooth and shining. Petiolar node shining, coarsely transversely striate, with a few scattered shallow foA^eolae. First gastric segment (postpetiole) evenly and distinctly striate or costulate (more finely and densely than in grammodes), trans- verse across the anterior quarter of the tergum, arching over the longitudinally-oriented remainder, with coarse interspersed punctures. The sides of this segment are smooth and shining, with sparse coarse foveolae. Second segment somewhat more finely striate longitudinally, with interspersed punctures, the striae reaching nearly half the length of the segment in the middle. A small anteromedian sliver, separated from the re- mainder of the surface by a curved line resembling the mark left by pressing a fingernail into something soft, is delicately transversely striate. Sides and posterior half of this tergal sur- face smooth and shining, with scattered piligerous punctures. Sternum of first segment predominantly smooth, with a few indistinct transverse wrinkles and grooves, shining. Sternum of second segment shining, indistinctly obliquely striate over the triangular side pieces. Sculpture otherwise as in grammodes, except that tarsi are only sparsely punctulate and are shining, and mandibles and clypeus more distinctly and more completely striate, including median clypeal sulcus. Color brownish-red ; mandibles, antennae and legs yellowish. The darker-colored parts of the body, especially alitrunk and gaster, overlain with a feeble pinkish-violet to bluish metal- lescence as viewed in certain lights. Holotype [MCZ] a worker from colony series of collection No. 649, taken at Bisianumu, Papua, the same locality from which came the types of grammodes (see above), by E. 0. Wilson. Paratypes: 6 workers from the holotype nest series (No. 649) : TL 5.2-5.4, HL 1.21-1.28, HW 1.04-1.11 mm. (CI 86-87). These specimens vary slightly in density of sculpture (especially on the petiolar node), in size of ventral petiolar process, and in stria- tion of second gastric segment, which in some examples reaches, and even surpasses, the midlength of the segment. Some speci- mens have the occipital border straight or nearly so, seen full- face. BROWN : ANT TRIBE ECTATOMMINI 315 Paratypes : 3 dealate females — 2 from the holotype nest series, and a unique (Collection No. 655) taken from a rotten log in which a G. grammodes worker was also taken (see under grammodes paratypes, above) at Bisianumu l\y Wilson. TL 6.7-6.8, HL 1.27-1.29, HW 1.11-1.12 (CI 87 in all 3 specimens), scape L 1.22-1.23, greatest diameter of eye ca. 0.35, WL 2.2, petiole L 0.65-0.66 mm. Alitrunk bullcy; mesonotum foveolate, with longitudinal rugae between. Pronotum coarsely foveolate, the interspaces smooth and shining, especially at midline ; sides of alitrunk behind pronotum predominantly coarsely longitud- inally striate. Color darker than worker, mahogany, nearly black to naked eye ; legs, scapes reddish-yellow. Males, 3 specimens for holotype nest series, of which only one was measured : TL Avithout genital capsule 6.0, forewing L ca. 4.2, HL 0.86, HW including compound eyes 0.98, AVL 1.76, petiole L 0.52 mm. Antennal segment lengths : scape 0.33, funiculus I 0.14, fun. II 0.29, fun. Ill 0.29 mm. Mandibles tri- angular, finely dentate, with larger apical tooth. Mandibles and convex clypeus finely longitudinally striate. A strong median carina connects clypeus and anterior ocellus. Ocelli large, clear, separated from one another by a distance a little greater than the diameter of each. Compound eye large and strongly rounded, greatest diameter ca. 0.29 mm. Dorsum of head and alitrunk shallowlj' foveolate, with predominantly longitudinal rugae inter- spersed. Epimera convex, mostly smooth and shining, with a few foveolae. Propodeum steep, gently rounded, declivity not distinct from dorsum, unarmed, vermiculate-rugose. Petiolar node de- pressed-paniform, longer than broad and about half again as long as high, mainly smooth and shining, with a few scattered shallow foveolae ; ventral process represented by a small oblique tooth or angle. Caster slender, smooth and shining, with piligerous punctures ; an irregular transverse wrinkle or fold line across the tergum of the first segment in all of my specimens. Veins Mf2 and Rsf2-3 missing from forewing. Abundant long, fine, tapered erect hairs on most surfaces of body, becoming shorter and more decumbent on legs. Wings densely covered with brown microtrichiae. Color piceous to black, legs brown, trochanters and genitalia yellowish. The holotype nest contained 30-40 workers and several males and dealate females; the colony was in a large rotting log in rainforest. Paratypes will be deposited in MCZ and elsewhere. 316 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY [62] G. horni is clearly a good species apart from regularis; honii is much smaller, and differs in other ways; besides, it is sympatric with regularis in the Guianas and probably elsewhere in South and Central America, but no intergrades between the two forms have yet been reported. The MCZ has numerous examples from each of several localities in British Guiana and the Panama Canal Zone, plus a sample each from Huachi, Rio Beni, Bolivia (W. M. Mann leg.) and from San Francique, Brasso, Trinidad (A. E. Emerson leg.). [63] The status of Spaniopone IwyUana has been discussed in the main section of the generic revision as one of the Holcoponera Group of genera. See p. 218. [64] G. menozzii is represented in the MCZ by type workers from Eio Grande do Sul, and by a worker from Volta, Parana : color blackish-brown, HL 1.16-1.21 mm., CI 109, 21-24 costulae between eyes. The ventral process of the petiole has a salient posterior corner, either digitiform or acutely dentiform. Two types of G. schubarti (Monte Alegre, Sao Paulo, 900 m.) are also blackish-brown, HL reaches 1.40 mm., but CI is still 109 ; 25-26 costulae between eyes. Ventral process of petiole with posterior corner reduced, obtusely rounded, shorter than anterior corner. A specimen similar to both foregoing species is from Corcovaclo, Rio de Janeiro (T. Barbour leg.) : color bright ferruginous (prob- ably teneral), HL 1.28 mm., CI 106; 22 or 23 costulae between the eyes. Ventral process of petiole with posterior corner obsolete, the process consisting only of the short, digitiform remnant of the anterior angle. It looks as though menozzii, schuharti and the Corcovado specimen may belong to one geographically variable species. More collections are needed. [65] In 1896, when Emery first set up Alfaria, he included two species then new: a larger, very finely striolate one (simulans) and a smaller, opaquely and extremely finely subgranulose one {miniita). Since then, other authors have added six species to Alfaria. In addition, Mann's OpistJioscyphus scahrosus is clearly a member of the same group, and thus makes seven additions since the original description of the first two species. BROWN: ANT TRIBE ECTATOMMINI 317 Bufonis is close to simulans, and in fact resembles in nearly every particular moderate-sized dark brown simulans specimens from Costa Rica ; the simulans types I have seen are somewhat larger and are dull yellowish in color, but they may be teneral. The only clear difference shown by hufonis is the well-developed metanotal groove in Mann 's unique type, which conceivably could be an ergatoid female or some other departure from the true worker caste. Until we know more about the normal variation in simulans, it seems best to recognize the Mexican hufonis as a species apart. Five of the six other post-Emery species all appear to merge into a single variable species indistinguishable from minuta. This has been determined from careful comparison of types (except mus) one against another and against what little other material could be assembled. To begin with the final and most important check : Wilson has compared the tjT)es of A. minuta and A. emeryi found in the Forel Collection, and in turn matched these with a topotypic worker of panamensis that I had previously checked directly against a panamensis cotype, carinata cotypes, and the holotype of 0. scabrosus, as well as additional samples from southeastern Brazil and Costa Rica sent by Father Borg- meier. Although there is some variation in size and sculpture, particularly of the gaster, these samples all appear to represent one species, minuta, which ranges from Central America to Bolivia and southern Brazil. The integument of this species is frequently fouled with a whitish incrustation, apparently a dried secretion like that found on many dacetines ; this deposit very deceptively alters the appearance of the surface features, and can make even nest mates look very different. Examinations of the 0. scabrosus holotype [USNM] reveals that this genus and species was described on the basis of the artist's unrepresentative depiction of the mandibles, rather than from the specimen itself. Actually, the mandibles are in no way unusual for minuta. Most of the differences cited in the other species descriptions are based on small errors of obserA^ation or differences in viewing planes. The type review showed these distinctions to be non-existent or completely trivial. Though the mus type was not seen, the description tells enough to make this synonymy reasonably safe. The frontal groove mentioned by Santschi is undoubtedly the weak frontal carina as viewed in haste. 318 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY The most recently described species, Borgmeier's striolata, pro- vides a link between the more "typical" Alfaria and the other Gnamptogenys species, and is itself a distinctive form. [66] Gnamptogenys mecotyle sp. nov. (Figure 42) Holotype worker.- TL 6.0, HL without cervical flange of occiput 1.16, HW 1.10 (CI 95), L head with closed mandibles 1.70, scape L 1.16, greatest diameter of eye 0.30, WL 1.85, petiole L 0.69 mm. General habitus of the larger species in the G. rastrata group, particularly aculeaticoxae, menozzii, etc., but diifering in the following respects. (1) Scapes long, when laid straight back from insertions, sur- passing occipital margin by more than 14 their total exposed length; their dorsal surfaces coarsely longitudinally striate throughout. (2) Costulation of head much less even, the costulae nearest the middle mostly much coarser than those near the eyes, mixed in gauge. Although a count is difficult and uncertain, it appears that there are about 28 costulae between the compound eyes, and about 11 between the frontal carinae. The longitudinal costula- tion of the clypeus is finer and regular. (3) The longitudinal costulation of the alitrunk does not con- tinue all the way forward to the anterior pronotal margin, but instead runs into an arched pattern across the anterior pronotum, where 5 or 6 costulae are transverse. (4) Propodeal teeth (L. ca. 0.13 mm.) distinctly longer and broader at base than in aculeaticoxae, and about ^^3 as long as the distance between the centers of their bases. Propodeal declivity narrowed above, margined, smooth and shining. (5) Node (Fig. 42) longer than deep and slightly longer than broad, gently arched, crossed by about 13 transverse cos- tulae. Ventral process shallow, in the form of two short, blunt subequal teeth joined by a low concave lamina. (6) Extreme base of postpetiole truncate in both dorsal and lateral views, leaving a small vertical anterior face that is nearly or completely smooth and shining. At the center of the border limiting this face above, the border is produced forward slightly as a blunt tubercle. Coarse costulae of postpetiole slightly wavy, becoming more wavy and erratic on sides of this segment near base. BROWN : ANT TRIBE ECTATOMMINI 319 (7) As minor characters, may be mentioned the cervical flange of the occiput, which is quite plainly visible in full-face view, and is convex in the middle ; and the gula and postpetiolar ster- num, both predominantly transversely costulate. Mandibles coarsely striate (smooth, punctate at apices) and weakly dentate along the masticatory borders. Color deep brownish-red ; man- dibles, antennae and legs more yellowish. Holotype [USNM] taken by W. M. Mann on the lower Rio Madidi, Bolivia, in February, 1922. Four worker paratypes [USNM, MCZ, Coll. Borgmeier] taken with the holotype are similar, differing slightly in size, depth of color and sculptural details. The size ranges from that of the holotype up to that of a worker: TL 6.3, HL without cervical flange 1.22, HW 1.14 (CI 94), L head with closed mandibles 1.77, scape L 1.23, greatest diameter of eye 0.33, WL 1.96, petiole L 0.73 mm. This distinct species is intermediate between hispinosa and aculeaticoxae, but is closer to the latter. It is apparently sym- patric with aculeaticoxae at or near the mecotyle type locality. [67] The variety ericae was based on a small Colombian speci- men of tornata having the costulae on the propodeum, especially the declivity, longitudinal. Such specimens are commoner south of Guatemala, but Costa Rican and other series are found in which the patterns of this region are longitudinal, transverse, and intermediately U- or V-shaped, even in single nest series. [68] The name fielirigi applies to specimens from south of the Amazon (type locality is San Bernardino, Paraguay). I have seen material from Goias, Minas Gerais and Sao Paulo states with the shallower and more shining costulation and the less acutely bidentate ventral petiolar process described by Forel. The more northern of these series tend toward the "typical" regularis from Ecuador, British Guiana and Venezuela (in the ]\ICZ). Var. splendida apparently applies to somewhat callow ^Mexican specimens. Samples in the MCZ come also from Costa Rica. [69] G. iiiordax is a very variable species. Size and gastric sculpture are particularly unstable characters, but extremes of 320 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY variation are connected by intergrades both inter- and intra- nidally. From its description, it seems fairly certain that La- treille's Formica nodosa is the same as mordax. However, the name nodosa has not been used except in catalogs, and I believe that it should not replace mordax, which has been used for this common ant for nearly a century. Forel's description of var. purensis applies well to one common variant of mordax, and I have examined a type of sehastiani, which is close to mordax types received from the British Museum. [70] Specimens in the MCZ from Mexico (Guadalajara, Guer- rero, Chiapas), as well as from Guatemala and Costa Rica, show that Santschi's differentiation of curtula and stolli as races is based on individual, partly allometric variation. Larger workers have transverse sutures on the posterodorsal alitrunk indicated as impressed lines. Apparently this species can live in somewhat drier places than the habitat of G. sirigata, the related species having much the same range. [71] In a nest series of porcata from Hamburg Farm, Costa Rica (F. Nevermann leg.), some specimens have the costulation of the posterodorsal alitrunk departing from a strictly longitudinal pattern ; in one specimen, the propodeal area is covered by a well- developed whorl, approaching the condition as described by Santschi for the magnifica types. The distinction between porcata and magnifica is thus seriously weakened, and it may be that these two names represent a single relict species now in the process of being replaced by the centrifugally spreading relative, G. pleurodon. [72] G. pleurodon is closely allied to porcata, but is smaller. It is widely distributed in the Amazon drainage. After examin- ing the nidotype series from Bolivia of var. recta (USNM, MCZ), assigned by Santschi to emery i, as well as a series from Yurac, 67 miles east of Tingo Maria, Peru (E. S. Ross and E. I. Schliuger leg.), I believe that the characters upon which Santschi differentiated pleurodon, emery i and recta are individual or allo- metric variates that may be expected in almost any nest. Emery's BROWN : ANT TRIBE ECTATOMMINI 321 female pleurodon was separated as vidua hy Santschi, but the differences are only those one might expect between worker and female of one species. [73] Gnampfogenys rastrata group. This group is essentially the old subgenus Parectatomma. There exist in this group many uncertainties at the species level, due to the present scarcity of material, to the fact that several names were based on single specimens (and even these hardly adequately described), and to the circumstance that females are mostly not yet properly associ- ated with workers. Some species must remain in doubt for the time being, but it seems wise here to review the material available and outline the problems involved. G. rastrata. The type (NM Vienna) is a winged female from "Brasil/F. Sahib, [erg]." HL 0.90, HL plus closed mandibles 1.40, HW including compound eyes 0.92, HW without eyes 0.83 mm. About 24 or 25 costulae between compound eyes : 9-10 between frontal carinae. Mandibles as in mcnozzii and schuharti, etc. (Fig. 16), masticatory border almost straight, feebly concave in basal third, faintly convex in mid third, then feebly concave again in apical third, showing a tendency toward the condition in mediatrix; no defined denticulation. Dorsal costulation of mandibles coarse, but not as bold as in scliuharti, shading off to a shining, nearly smooth surface near the inner basal angle ; costu- lation replaced by a smooth surface with shallow punctures on the apical third of the blade. Teeth of propodeum minute blunt tubercles ; coxal tooth short, triangular, as in schuharti w'orker ; shape of petiolar node and its ventral process also very much as in schuharti types, and like those of acideaticoxae. Costulae of petiole longitudinal, diverging behind as seen from above ; trans- verse across anterior face of node. Color brownish red. The worker from Alajuela, Costa Rica, determined by Emery as "rostratum Mayr" can scarcely be the same as the rastrata type from Emerj' 's account, and considering the remoteness of the locality. For the present, I am unable to guess what Emery had from Costa Rica (Emery, 1890a :41). Wheeler (1925:5) determined some specimens collected by Holmgren at Llinguipata, Peru, as rastrata, but I have examined four of these specimens and found them to be the form described by myself as Holcoponera mina, now placed in Gnaynptogenys [46]. 322 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY A worker in the collection of Father Thomas Borgmeier, from Nova Teutonia in southeastern Brazil, 300-500 m. altitude (F. Plaumann leg.) is close to rastrata, and may actually be its worker. HL 0.82, HW with compound eyes 0.86, HW without eyes 0.83 mm., 27-28 costulae between compound eyes. Eyes large. General habitus, mandibles, etc. about as expected for worker of rastrata, as compared with the female type. Petiolar node slightly but distinctly broader than long; costulae trans- verse across the anterior face ; as seen from above, costulae longitudinal in the center, but around this concentrically circular, with only 2-3 transverse at the posterodorsal border. The ventral process is as in the type worker of G. menozzii, i.e., with concave ventral border, rounded anterior corner, and acute posterior corner. Color red-brown, head slightly darker than rest of body. The variation in nodal sculpture and form of ventral process of petiole may be normal for a single species ; only further material will tell. G. trigona. Emery described this form from a unique female from Novo Friburgo, Santa Caterina, Brazil. The description is in the form of a very brief comparison with triangularis Mayr; the size is "a little smaller . . . L. 4.5 mm." Sculpture much coarser than in triangularis ; antennae thicker and shorter, the scape scarcely surpassing the occipital border. On the petiole, the costulation is disposed in concentric, but transverse, arches. Except for the transversely concentric petiolar costulation, this characterization would match fairly well the type of rastrata; however, the petiolar sculpture may possibly be vari- able enough to include the rastrata and trigona patterns within one species. Other specimens I have seen, determined as trigona in various collections, are as large as or larger than triangularis, and have coarse costulation, but this is concentric on the petiole without being in the least transverse in orientation. I believe that these large specimens, such as the worker from Corcovado, discussed above [64], are probably not the true trigona. I conclude for the jDresent that trigona may be the female of the small species next described below, or else it is a synonym of rastrata. BROWN : ANT TRIBE ECTATOMMINI 323 Borgmeier Coll., Nr. 5756. Angra dos Reis, Rio de Janeiro State, Brazil (Lopes et Lent leg.). Four workers: HL 0.74-0.76, HW without compound eyes 0.69-0.73 mm., CI 93-96. Eyes large, but only feebly convex. 24-28 costulae between eyes. Mandibles as in rastrata tj^pe and the possible rastrata worker (Nova Teu- tonia, see above), but costulation coarse and extending more completely into basal corners. Propodeal teeth reduced to small tubercles, inconspicuous alongside the larger, protruding, obliquely-facing spiracles, just as in menozzii. Petiolar node about as broad as long ; costulae concentric, forming a trans- versely elliptical pattern, variable in its center from specimen to specimen. Ventral process of petiole of the menozzii type. Teeth of posterior coxae shorter than in menozzii, but still digiti- form or spiniform. Color red-brown. The total length of these specimens varies from about 3.1 mm. to 3.5 mm., outstretched with mandibles. This size is considerably less than the 4.5 mm. measurement Emery gave for the trigona female, and Emery's measurements usually fall under the ones I make on the same specimens of other ant species. Nevertheless, the correspondence of other characters, particularly the sculpture, with Emery's de- scription, leaves the possibility open that Sample 5756 represents the worker caste of trigona. G. triangularis. Four workers from: Ipiranga, S. Paulo State, Brazil (Luederwaldt leg.) ; Tafi Vie jo (F. Silvestri leg.) and Tafecillo (N. Kusnezov leg.) in Tucuman, Argentina, measure HL 1.14-1.21, HW 1.12-1.16, eyes excluded. The head is variable in proportions, some specimens having the length slightly greater, others the width (CI 94-102). Only slight variation in gauge and density of costulation, from about 31 to about 35 costulae between the compound eyes. Propodeal teeth small but distinct and acute. Petiolar node varying from about as broad as long to distinctly broader than long, as seen from above more or less transversely costulate ; sometimes only one or two transverse costulae in the middle, surrounded by concentric, ellipsoidal pattern on a trans- verse axis; sometimes the costulae transverse, meeting far down the sides of the node in a V or U pattern. Ventral process of petiole concave beneath, anterior corner rounded, posterior corner subrectangular. Color red-brown to blackish. Forel's race richteri reads like a common variant of triangularis, and it seems likely that Forel misidentified typical triangularis. 324 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY [74] Santschi's reichenspergeri is not a Heteroponera, to judge from a type worker, several males and a dealate female received from the Basel Museum and from Father Borgmeier. The frontal carina is lacking, and habitus and sculpture clearly place the species in Gnaynptogenys, among the species of which reichen- spergeri seems nearest to relicta. However, reichenspergeri has a well developed air of propodeal angles, and the striate sculpture of the alitruncal dorsum is effaced medially in the worker ; the gastric sculpture is also largely effaced, broken up into short, bilaterally oblique sections of striation or fine rugulation, with large areas virtually smooth and definitely shining. The males are blackish in color, and generally resemble the species formerly placed in Ilolcoponera, but vein Rsf2*3 is present and situated as in G. mordax (Brown and Nutting, 1950, PI. 8, fig. 5). The eyes of the worker are minute but distinct, and the promesonotal suture is cut through on the dorsum ; the position of the metanotal groove is indicated by a faint impressed line. The petiolar node is anteroposteriorly compressed, and resembles those of relicta and the former Spaniopone and Rhopaloponc species. EA^ery thing considered, this aberrant Gyiamptogenys appears to be close to a perfect intermediate species linking the old genera Gnampto- genys, Spaniopone, Ilolcoponera and Rhopalopone. [75] (/. rimulosa is known only from the vicinity of Rio de Janeiro, where it is infrequently collected. Specimens sent by Father Borgmeier under this name agree well enough with the description. [76] From the brief original characterization, qiiitcnsis is ap- [jurently only a minor variant of tortuolosa, which is widespread in Ecuador. [77] Gnamptogenys semiferox sp. nov. (Figure 14) Holotype worker: TL 7.0, HL 1.43, HW (across eyes) 1.44, HW (at anterior corners) 1.39, closed mandibles extend about 0.81 mm., WL 1.99, petiole L 0.74, greatest eye diameter 0.29, scape L 1.11, absolute exposed length of left mandible 1.32; CI BROWN : ANT TRIBE ECTATOMMINI 325 97. Differs from Gnampiogenys {-=Emeryella) schmitti workers of about the same size in its narrower head; shorter, broader, differently formed mandibles (Fig. 14) ; and in the longer petiolar node, which is a trifle longer than broad seen from above. The median clypeal lobe is more prominent, and its an- terior border is entire and evenly convex in outline. The sculp- ture differs in a number of ways. The costulation throughout, but particularly on the head and pronotum, is finer, less uniform and even, less shining, and a little less distinct. The interspersed punctation, very sparse and insignificant in schmitti, is more abundant, coarser and con- siderably more distinct in semiferox, especially on the head, but it still does not interfere seriously wdth the regularity of the costulation except in the areas just mesad of the compound eyes and on the dorsolateral surfaces of the propodeum. The punc- tures are mostly indistinctly bounded, but contain piligerous tubercles. The dorsal face of the propodeum is longitudinally and subvermiculately costulate (evenly and transversely in schmitti). Color deep piceous, as in fully pigmented schmitti, appearing black to the naked eye. The appendages are more reddish. The holotype [MCZ] and two very similar paratypes [Coll. Borgmeier, MCZ] all bear the same data : Mt. Diego de Ocampo, 3000-4000 feet altitude, Dominican Eepublic, July, 1938 (P. J. Darlington leg.). Dr. Darlington tells me that the type locality was chiefly rain forest in which small palms were prominent, but that the land was being cleared for agriculture even at the time of his visit. In the development of the mandibles, semiferox is intermediate between schmitti and hanksi on the one hand and mediatrix sp. nov. on the other. In sculpture, semiferox is closest to schmitti, but show's tendencies toward the irregular, coarse, partly reticu- late or punctate sculpture even better developed in hanksi. In the lack of a dorsal tooth on the posterior coxa, and in the smooth postpetiolar sternum, schmitti and semiferox are alike and dis- tinctive, but other Gnamptogenys lack the coxal armament, and hanksi has a partly smooth postpetiolar sternum. 326 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY [78] Gnamptogenys mediatrix sp. nov. (Figure 15) Holotypye worker : TL 5.4, HL 1.04, HW (across eyes) 1.13, HW (near anterior corners) 1.08, closed mandibles extend about 0.53 mm., WL 1.64, petiole L 0.59, scape L 1.02, absolute exposed L of left mandible 0.88 ; CI 104. Very similar except in size and subfalcate form of mandible (Fig. 15) to G. acid eat icoxae and to me7iozzii and schubarti. Propodeal teeth short but sub- acute ; coxal teeth very slender, with feebly clavate apices. Petio- lar node broader than long and very nearly as long as high, its dorsal profile only feebly convex and only slightly higher pos- teriorly than in front (similar in shape to that of aculeaticoxae) ; anterior face steep, posterior face distinct and abruptly vertical (not distinct from dorsal convexity in aculeaticoxae) . Subpeti- olar process with both anterior and posterior free corners rounded, the latter obtuse, oblique ventral margin very feebly concave in outline, nearly straight (form of process as in schu- barti types). Petiolar costulation predominantly transverse, as in aculeaticoxae. Eyes large, convex, their anterior margins about at midlength of head. Costulation almost precisely as in aculeaticoxae samples from British Guiana so far as density and direction are con- cerned ; 26-28 costulae between the eyes. Mandibles costulate on basal %, except near basal angles, where costulation becomes finer and then obsolescent ; apical third of blade smooth, with coarse piligerous punctures. Color piceous, with reddish tinge ; appendages lighter, more yellowish. Holotype and one very similar paratype [MCZ and Coll. Borgmeier respectively] collected together at the present Belem do Para, Brazil (Dr. Fred Baker leg.), presumably in the out- skirts of the suburb of Souza (Mann, 1916:399). Other paratypes are all from the environs of Rio de Janeiro [Coll. Borgmeier, MCZ] : 3 workers from "Rio de Janeiro" and one from the Jardim Botanico (H. Souza Lopes leg.) ; one alate female from Engenlio Novo (C. R. Goncalves). The workers vary somewhat in size, the largest being about the same size as the female in dimensions and proportions of the head. There is slight variation in the density of costulation and in the shape of the BROWN : ANT TRIBE ECTATOMMINT 327 subpetiolar process. The female is scarcely larger than the largest worker, and the eyes are only a little greater in diameter. HL 1.14, HW (near anterior corners) 1.21 mm. Forewing L 4.5 mm. Venation of typical Gnamptogenys pattern ; Mf2 present but very short. Petiolar node a trifle shorter and broader than in the worker. CI 106. [79] The material available to me, in conjunction with the dis- cussions and figures of Santschi's 1929 revision, leads me to consider the names provisionally synonymized as representing parts of the continuous variation of one species. I have already dis.iussed in general terms (Brown, 1957, pp. 489-490) the difiiculties in Santschi's treatment of the variation of characters like gauge and density of costulation, body color, development of mesepisternal lobe or flange, and shape of ventral petiolar process; these very difficulties are most exaggerated in his han- dli)ig of striatula, hrasiUcnsis and relatives. From Santschi's own data, it seems unlikely that the types he saw of striatula and hrasiliensis can be different species. But it seems possible that not all the infraspecific forms he described under hrasiliensis also belong to the same species. Just wdiat constitute species sep- arations between striatula, the form with nominal priority, and other forms in or near this complex, we cannot yet say because of the lack of sufficient material and the inadequacy of detail in our knowledge of distributions of the various jDopulations. In general, one might guess that most striatula-like specimens with more than 30 costulae between the compound eyes are striatula itself, while most with less than 30 are other, closely related species. An indication that this is so may be furnished on sympatric series from Agudos, S. Paulo State, Brazil (W. W. Kempf leg.), one of which is finely costulate (ca. 40 costulae between eyes) and has large eyes, while the other has smaller eyes and about 26-28 costulae between them. This latter type with coarser sculpture perhaps corresponds to the forms named by Santschi as regularis, arcuata and rustica, from southern Brazil and Paraguay, and may also be the same as the variant simplicoides of hrasiliensis. At first view, there seems to be intergradation between the coarse and fine sculpture types, but this may be due in part to Santschi's inaccuracy in counting the 328 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY costulae (his counts are sometimes grossly contradictory as be- tween key and text). Other intergradation in costular density may indicate that a character displacement situation exists be- tween coarse and fine sculpture types when they occur sympatri- cally. The value of the eye diameter as a character has not been determined. G. striatula is widespread in tropical South America, where the subpetiolar process usually or always has two rather distinct, subequal corners or lobes in its free portion. Specimens from Venezuela and Colombia look like striatula, but have the poste- rior corner or lobe much reduced, so that the process is more or less triangular or forms a single lobe, the condition character- istic of G. curtula [70]. The possibility should be entertained for the time being that curtula is the geographical variant of striatula in Central America and Mexico. In Costa Rica, there is a counterpart of the rustica group of South America in wheeleri, w^hich maintains itself distinct from the curtula of Costa Rica in the form of the ventral process. The relationship of striatula to wasma7ini cannot be decided without more material ; its node is rather thick and its ventral process subtriaugular, so wasmanni seems to me to be closer to 2)leurodon. The synonymy suggested and the opinions offered here con- cerning species-level relationships are strictly to be considered as "shooting from the hip," although it is felt that such treat- ment is justified at this time as a small beginning at making the necessary revision of the striatula group. It will perhaps be worthwhile reemphasizing for the benefit of future revisers the unreliability of one particular character given much weight by previous authors dealing with "Holcopo7iera" species. I refer to the mesepisterual lobe or flange. Not only is this structure highly variable in single nest series, but it has been very loosely dealt with by Santschi. Reviewing a substantial part of the same material, including unique types, that Santschi used in writing his revision of 1929, I have found that in several cases his drawings or descriptions are unrepresentative. For instance, his figure 19 shows the mesepisterual lobe of ivheeleri with a small posterior portion that does not exist in the types ; actually, it seems that Santschi was drawing a small ridge that traverses BROWN : ANT TRIBE ECTATOMMINI 329 the sternum at this level, and which can be seen by focusing down only slightly from the plane of the lobe. In other cases, the "tooth" at the posterior end of the lobe proved to be a stray bit of glue or dirt. In some series, the lobe was visible on only one side of one specimen; in the other specimens it was covered by glue or otherwise hidden. This means that Santschi 's idea of the form and constancy of the lobe was usually drawn from a small fraction of the actual specimens he saw. [80] G. sirigaia is a small, coarsely sculptured forest species. Wilson took it in rotten wood at Las Ilamacas, near Santiago Tuxtla, Veracruz (tropical evergreen forest), and at Las Perlas, near Orizaba, at about 2000 m. in Liquidamljar-'h.ovvibQwai forest. C. and M. Goodnight took a specimen at about 1600 m. altitude in a pine grove at Union Juarez, Chiapas. [81] G. teffensis is a large species for which we now have quite a few localities in the Amazon-Orinoco drainage, from the Guianas to Bolivia. The color is variable, but is most often some shade of ferruginous brown. The holotype of teffensis has been compared directly with the type of var. concinna, and in my opinion the two specimens fall within the normal variation of a single species; the posterior tooth of the teffensis type's mese- pisternal lobe proved to be only a piece of dirt adhering to the lobe, and once cleaned, the lobe looks like that of concinna. The very closely related nioelleri, with its color variant splen- dens, has been recorded by specific locality only from the south- eastern extremity of Brazil, particularly from Santa Caterina. Mann's Madeira-Mamore record applies to teffensis. Actually, teffensis and moelleri may be geographic strains of one species, although the more slender funicular segments of moelleri still seem to separate this form satisfactorily from teff'ensis, even considering that the funicular segments of teffensis are variable in thickness to some extent. [82] G. sulcata is highly variable in color, and less so in density and gauge of its costulate sculpture. The principal color of worker and female ranges from uniform ferruginous (perhaps usually in callows) to black. The head is always dark in adults 330 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY near or past full coloration, and is piceous or black ; the mandibles are always pale yellowish. The MCZ now has a number of series representing the stages of intergradation in both color and sculpture, and I have matched the types of ypirangensis, hufonum and var. nitens directly with these. Var. lineata has long been considered only a color variant of sulcata, and this follows the description well. Var. cearensis is described as a fine-sculpture form, and the details fit MCZ specimens well enough. A male from Barro Colorado in the Panama Canal Zone (N. Banks leg.) is blackish in color, but has the pronotum a bright contrasting ferruginous yellow in hue. It is about the right size to fit workers of sulcata from the same locality, so I believe that it may well be the corresponding sex of this species. [83] Just as Wheeler puts it, the only important difference (size) between the minor syntype series and the schmiiti workers taken by Mann at Diquini, Haiti, is an "average" one. Differences between Forel's type and later collections are not apparent from Forel's description. Normal size variation is clearly great even within single limited series. Internidal size and sculpture varia- tion is stronger in this than in some other Gnamptogenys species, but not as strong as, for instance, in mordax. [84] Kennedy and Talbot (1939) claimed for the worker-female of P. silaceum one-segmented maxillary palpi, but their figures 9, 10 and 12 show that these palpi have two segments — a flat- tened apical one and a short basal one. The labial palpi were depicted with two elongate segments. This pattern has been found also in dissections of croceum and relictnm; the maxillary palp of the former is shown in Figure 26. The arrangement of the first and second segments, with the second segment attached to the first by a slender lateral neck, holds through all of the species for which palpi have been examined, and is apparently characteristic of the genus. Kratochvil (1944) shows the palpi of worker melinum (under the name fialai) in his figures 2a, 2b, and these are like the sample of perganclei I dissected, with a formula of 4, 3, the apical segments being fairly elongate in form. The holotype of stictum, viewed without dissection, shows three exposed segments BROWN : ANT TRIBE ECTATOMMINI 331 for the maxillary palpus, the basal one evidently corresponding well to the second segment of the foregoing species ; from this, I assume that sticfum has four-segmented maxillary palpi, although the labial palpi of this species cannot be made out. A single female of micrommatum dissected had two segments in the labial palps, while the maxillary palps appeared to have three segments on the right side, and two on the left. The two- segmented palpus has resulted from what seems to be a fusion of the basal segment with the slender basal neck of segment II. This condition may be an anomaly for the species; only further material will settle this. According to Kennedy and Talbot {loc. cit.), the male of silaceum has a formula of 4,2, but here again their figures show that they did not count the small basal segment of the maxillary palpus. The formula is actually 5,2 as interpreted from their figures, and the second segment of the maxillary palpus is formed in much the same way as in the worker and female. We should expect that the formula for other species of Proceratium males is ordinarily 5,3 or 5,2. [85] The distinctions drawn between the southern European mayri and the North African algiricum do not appear to be very striking. I have not seen any North African specimens referable to either species, but judging from the characters cited in the literature, it would seem that algiricutn merely represents geographical variation of the European species. Final decision in this case must wait for critical comparison of the available material. [86] Dr. George Arnold has kindly sketched for me the type worker of arnohli, and 1 have presented these drawings here (Figs. 1, 34, 35) with the realization that they are perhaps not strictly comparable with the remainder of the figures of the genus given here. Undoubtedly Dr. Arnold and I have empha- sized different points in a slightly different fashion, and, of course, the scale is not the same. 332 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY [87] P. californicum was described by Cook on the basis of a male stated to have been taken by himself at Glenwood in the Santa Cruz Mountains of coastal California. The description and figure fit well enough a Proceratiiim, but of course so few males are known in the genus that this stray specimen cannot easily be compared with any other species. This is the only record of a Proceratium from west of the Great Plains in North America north of Mexico. The type locality is not an impossible one by any means; undoubtedly the mountains of California still hold some surprises for ant collectors. Nevertheless, it would be well to confirm the presence of Proceratium in California through further collections before this record is unreservedly accepted, especially since other data in Cook's book is open to some ques- tion (see reviews: Brown and Wilson, 1953; G. C. Wheeler, 1954; E. S. Ross, 1954). [88] A series from the Cuernos Mts., near Dumaguete, Negros Oriental, Philippines (J. W. Chapman leg.) fits the diagnosis of carinifrons well enough. The relationship of this species to longigaster needs investigation. [89] This species, closely related to micrommatum, lacks the median clypeal projection supposed to be characteristic of Sys- phincta. [90] 1 have examined a specimen of P. itoi through the courtesy of Dr. K. Yasumatsu ; this example is from Setagaya, Tokyo (A. Haga leg.). [91] Wilson has compared the type of lomhokense with Philip- pine specimens of carinifrons; the latter are much smaller than lomhokense — according to Wilson's rough guess, only about 1/4 the bulk of the lomhokense type. [92] In the MCZ there is a dealate female of mancum from Quirigua, Guatemala (W. M. Wheeler leg.) and a worker labeled simply, ' ' Guatemala. ) ) BROWN : ANT TRIBE ECTATOMMINI 333 [93] Ixoger's description of micrommatuni was good for its time, and from certain aberrant features mentioned, especially the contiguous frontal carinae, its identity seems assured with speci- mens from tlie following localities : Barro Colorado I., Canal Zone (N. A. Weber leg.) ; the cavernicola type, from Cliilibrillo Caves, Panama; from Soledad, Las Villas Prov., Cuba (H. B. Mills leg.) ; Chichicastle, Tabasco (F. Bonet leg.) ; and Pueblo Nuevo, near Tetzonapa, Veracruz, Mexico (E. 0. Wilson leg.). S. micrommatum is apparently widespread but rare in circum- Caribbean lands. With the exception of the cavernicola type, the above specimens were all taken in soil-cover berlesates. Borg- meier (1957 :118) has recently dealt fully with P. micrommatum ; he concurs in synonymizing cavernicola. [94] When Koger first described melinuni from specimens in the Berlin Museum, he recorded them as having come from "Caro- lina," mentioning that they were "with" the specimens he made the types of P. croceum. In view of other evidence, it seems possible that the association of melinum and croceum specimens in the Berlin Museum may have been the reason why both of these species have been considered to be North American since Roger's time. The record might have come under serious doubt had not Wheeler (1905) reported P. melinum (as Sysphincta melina) from the collections of P. J. Schmitt, made in southwestern Pennsylvania; this Pennsylvania record has been repeated since (e.g., Creighton, 1950 :42), but without anyone's having reviewed the specimens upon which the report was based. I have tried to locate the relevant specimens in all the major collections in the United States and Europe, as well as in Schmitt 's collection, now at St. Vincent Archabbey, near Latrobe, Pennsylvania, but the only species I could find collected by Schmitt were silaceum and pergandei (among Pennsylvania-collected material). In view of the fact that Wlieeler did not mention directly any specimens that he had examined and determined as melinum, and since no specimens can be found corresponding to this record, the record is open to question. To my knowledge, no one has taken melinum in North America for about a century. Of course, such negative evidence must not be over-emphasized ; after all, Discothyrea testacea Roger, described from the same general area, has now 334 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY been rediscovered in the southeastern U. S. [103]. However, there is other evidence to be considered. Melinum belongs to a group not otherwise known from North America, but which is widespread in temperate Eurasia. In 1895, Emery pub- lished reasonably good figures of all castes of melinum from the types, and careful examination of these figures reveals no significant difference between the types and the species described by Forel in 1886, and later by Emery himself, as europaea. In my opinion, melinum and europaea represent one and the same species, regardless of where their types came from. Of course, I consider it most likely that the melinum types really were collected in Europe, and I do not believe the species occurs naturally in the United States. In view of the greater familiarity of the name europaea, it might seem best to attempt to conserve it despite the priority of melinum. However, the best characteri- zation of the species up to recent times was that of Emery under the name melinum (1895a), and the name melinum has been used in numerous publications since. In this case, I favor the prior name. The other melinum group members are itoi of Japan, fialai of central Europe, and rossica of southern Russia. The last two seem to me to be slight geographical variants of melinum that are not worth formal names. The characterizations are based on scanty material, and little allowance is made for normal variation within series of "typical" melinum, which includes specimens with more convex alitruncal profiles approaching the condition as described and figured for fialai and rossica. Kratochvil shows clearly that in fialai the palpal formula is 4,3, probably the same for all melinum. I have examined a fialai eutopotype or paratype through the kindness of Dr. K. Samsinak. [95] From Santschi's characterizations of numidicum and nor- mandi, it is not clear that the differences between them are worth species separation. I have not seen any North African material myself, but Wilson has kindly compared for me a specimen in the MCZ Collections (ex Finzi Coll.) from Tirana, Albania (Ravasini and Lona leg.), with a numidicum type in the Emery Collection at Genoa. Wilson states that the type has more abundant erect hairs on head and alitrunk than does the BROWN: ANT TRIBE ECTATOMMINI 335 Albanian specimen, a point which apparently also is supposed to distinguish numidicum from normandi. Probably future study will show that normandi represents only slight individual or internidal variation within one species, numidicum. The Albanian record is, I believe, the first from Europe for this species, and represents a very considerable range extension ; older records are all North African. [96] A specimen taken by "Wilson in a mixture of rotten wood chips and soil between the buttresses of a rain forest tree at Tumnang, Mongi Watershed, 1500 m., Huon Peninsula, New Guinea, agrees well with Emery's description and with a type of Donisthorpe's Ponera caeca (nee P oner a coeca Santschi) that was compared directly, courtesy of Dr. E. S. Koss of the California Academy of Sciences. The type of Donisthorpe's species came from Maffin Bay, New Guinea. A single specimen taken in rotten wood in rain forest at Malanda, 800 m., northern Queensland (W. L. Brown leg.), is similar to the Tumnang specimen in size and in its weak, more or less shining sculpture, but has the propodevim gently rounded into the declivity as seen from the side, with only the most obtuse trace of angulation ; the color is also lighter, but color means little in this genus. Another Proceratium worker was taken by myself in a small rotten log on the floor of second-growth. Eucalyptus gigas-dom- inated rain forest on the summit of Tamborine Mt., southeastern Queensland, at about 600 m. altitude. The Tamborine worker may well belong to a species apart from papuanum, for it is slightly larger than the Tumnang and Malanda specimens, and a little more robust in body build ; the petiolar node is slightly lower and thicker, and the sculpture is definitely rougher and opaque over the head and alitrunk, subopaque over the post- petiole. I do not find it expedient to describe a new species from a unique worker in such a tightly-related and little-known group of the genus at this time. However, wdth the description of P. stictum sp. u. below [99] added to the above records, it is clear for the first time that Proceratium is represented rather well in eastern Australia, nearly or quite south to New South Wales, and that at least two, and possibly even three, species are included in the fauna. 336 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY [97] Several workers of pergandei were found by myself under a rook in rich mesophytic forest on Great Blue Hill, Milton, Massachusetts, the most northeasterly record for the species yet known. These workers were confined in a plaster nest for tw'o weeks, but could not be induced to feed on various dissected fresh insects offered them ; they finally died one at a time. Dr. Wilson tells me that Mr. D. W. Pfitzer of the University of Ten- nessee once told him that he had observed workers of pergandei clustered around a spider egg-case, into which they appeared to be trying to bite their way. [98] P. silaceum is the common species of the genus in North America. Wilson and I have found it feeding on spider and other arthropod eggs in nature, and have maintained it on spider eggs almost exclusively for weeks in the artificial nest. When w^ell supplied with spider eggs, it will have nothing to do with other insect remains, including eggs, larvae or pupae of other ant species, upon which the related species crocemn was fed by Haskins (1930). In both natural and artificial nests so far ex- amined, the silaceum workers tend to store more eggs than they will immediately need. P. silaceum ranges northward as far as Lexington and Woburn, Massachusetts, where I have found it extremely rare, nesting under stones. It nests under stones also in moist, wooded hilly sections of eastern Pennsylvania, but in the adjacent lowlands and pine barrens along the Delaware River and in New Jersey, the nests are almost invariably in rotten logs or stumps. The synonymy of silaceum, long a source of confusion, was settled by Creighton in his penetrating study of 1950 (pp. 36-40) ; his conclusions are followed here. [99] Proceratium stictum sp. nov. (Figures 45, 46) Holotype worker: TL (to posterior curve of second gastric segment') 4.7, HL 1.08, HW 0.97, WL (with cervix) 1.48 mm.; CI 90. The general form of the head and body is shown in Figures 45 and 46. Notable is the broad clypeus, separated from the remainder of the cranium by a reasonably distinct suture, BROWN : ANT TRIBE ECTATOMMINI 337 and not markedly tilted from the horizontal position. The broad median lobe is bicarinate, the space between the carinae concave and emarginate at the anterior border; this broad lobe corre- sponds to the much more strongly reduced median process in the pergayidci and melinum group workers and females, and it may well be the homologue of the various modifications of the bicari- nate or bidentate clypeus as found in Monomorium and many other myrmicine genera. Maxillary palpi with at least three segments (probably four) ; labial palpi not seen. Antennae 12-segmented, gently incrassate toward apex, not forming a distinct club, but the apical segment much longer and thicker than the rest. Figures 45 and 46. Proceratiiim stictum sp. nov., holotypc worker. Fig. 45, lateral view of head and body. Fig. 46, full-face view of head. Drawings by Dr. F. Y. Cheng. Mandibles triangular, with oblique apical (masticatory) mar- gins set with four coarse teeth counting the blunt basal angle. Eyes distinct but small,' each with a single clear, convex facet, placed a little anterior to the cephalic midlength. 338 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY Alitrunk stout, compact, convex above, without traces of dorsal transverse sutures, seen from above broadest across the humeri. Propodeal teeth short but prominently projecting, dorso- ventrally flattened and with upturned apices. Propodeal de- clivity short and broad, not margined above or on sides between upper and lower (metapleural) teeth; the latter teeth broad, laterally compressed, subtriangular. Petiolar node low and paniform; seen from above slightly but distinctly longer than broad and very slightly narrowed in the anterior half, suggesting incipient formation of a peduncle ; node anteriorly squarely truncate, with prominent anterodorsal angles opposing the mesial surfaces of the lower propodeal teeth ; upper nodal surface evenly convex in both directions ; ventral appendage divided into three small, acute teeth. Basic sculpture of head, alitrunk, petiole and postpetiole densely and irregularly rugulose and with numerous well-spaced, narrow, but deepset punctures with central tubercles at their bottoms, this sculpture boldest on the petiole. Cephalic dorsum inside the eye with punctures indistinct, but the rugules more linear and running obliquely posterolaterad from the region of the antennal insertions; vertex with transverse rugulation. Cervical dorsum and lower propodeal declivity smooth and shin- ing; first gastric segment also smooth and shining, but abund- antly sown with small, distinct punctulae. Clypeus very finely rugulose. Mandibles very strongly striate up to apices, which are smooth, their bases with elongate punctures between striae. Legs and antennae very finely and densely punctulate, sub- opaque. (Legs stout, middle and posterior tibiae each with a single broadly pectinate spur ; tarsal claws moderately well developed, simple, as in other Proceratium species.) Pilosity fine and erect, moderately long and abundant, dis- tributed evenly over the entire body, including mandibles, an- tennae, legs and gula. Pubescence fine, dense, suberect to sub- reclinate, but not hiding sculpture. Color rich ferruginous red; appendages, mandibles and apical segments of gaster yellowish- ferruginous. Holotype [MCZ] a unique worker taken at Kuranda, northern Queensland, Australia, in a rotten log in heavy rain forest at an altitude of about 370 m., November 1, 1950 (W. L. Brown). BROWN : ANT TRIBE ECTATOMMINI 339 This species is abundantly distinct from the members of the pergandei and micrommatiim groups, to which it is most similar. The broad median clypeal lobe, the form of the propodeum and its teeth, and the low, elongate petiolar node are key characters, all of which seem to be near the generalized type of the genus. The single-facetted eyes and solidly fused alitrunk are, however, as specialized as these structures ever become within the genas, except that the single facet of the eye, while hyaline, is some- what larger than the eyes in some other species. It would be most interesting to see the winged sexes and the larva of this appar- ently very primitive Proceratium. [100] The holotype of P. ioscliii has been examined through the kindness of Prof. Athos Goidanich. I have drawn the middle front section of the head (Fig. 25) and the petiolar node and adjacent segments (Fig. 30) from the holotype; the difference between my figure of the node and Consani 's figure 1, no. 2, may be explained by the fact that Consani included in his drawing a large piece of dirt that adhered to the apex of the node and was matted in the hair, appearing to be a part of the node itself. [101] The four species described from southern Australia are bidens, crassicornis, leae and turtoni. I have collected workers that compare well with the types of turtoni in the National Museum at Melbourne ; these specimens come from the Dande- nong and Warburtou Ranges to the east of Melbourne, where the species is fairly common in the fern gullies as well as under stones in grassy-floored sclerophyll forest at 600-700 m. on the Dande- nong summits. I am unable to separate this species, which varies somewhat in color of integument in the queens, from the type of leae, and I think that further study may prove them synony- mous. The type of bidens in the MCZ is a large, dark-colored worker with fairly prominent propodeal angles or teeth that are directed obliquely upward, in this differing from the workers of turtoni I have seen. The bidens type, however, has a very faint trace left of the promesonotal suture across the alitruncal dorsum, and therefore it could be an ergatoid female. Thus, its specific identity with turtoni or leae, or with both, is not ruled out. The species crassicornis, retaken by W. M. Wheeler at Margaret 340 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY River in southwestern Australia, is smaller than turtoni, and, at least in the series before me, has eight-segmented antennae. Clark states that the antennae of his types had nine segments, but his figure 4 shows ten segments, indicating that he really did not have a clear count of the segments. The propodeal angles of crassicornis workers are also more prominent than are those of turtoni. Clark told me personally in 1950 that he thought turto7ii and leae were probably synonyms. The original collector of hidens, Mr. F. E. Wilson, informed me in a conversation that the real type locality of this species was Cement Creek, at about 700 m. altitude, and some miles up into the Warburton Range, away from the town of Warburton. In addition to the species already discussed, I have found what appears to be another distinct species at One Tree Hill in the Dandenong Range, east of Melbourne. This species is smaller than turtoni, has less noticeable pubescence and more distinct propodeal angles, and has seven-segmented antennae (two worker specimens, antennae on both sides counted). I have considered that these differences might be due to allometry within a single species, and continuing to take allometry into account, it seems best to wait until all of the available Australian Discotliyrea ma- terial can be examined in one place before attempting to decide the status of the seven-segmented variant. Most of this material, including some belonging to the MCZ, is still with the Clark Collection at Canberra. [102] The African species of Discotliyrea fall into two groups: those with the clypeo-frontal fusion process flat-topped and broad (Fig. 48) — ociilata, sculptior and mixta; and those in which the process forms a simple convex or angular vertical plate — traegaordhi, hcwitti, poiveri and patrizii. In the first group, mixta is distinct by the characters mentioned in the description [105]. Santschi's scidptior, described as a variety of oculata, may in fact be no more than a variant of that species. I have not seen the types of either oculata or sculptior, although I have seen a single worker collected by N. A. Weber in Kenya that in most respects answers to the ocidata description, except that the eyes have less than 30 facets. BROWN : ANT TRIBE ECTATOMMINI 341 In the second group, the species are apparently all very close, and they may, in fact, represent only a single species. D. poweri. with its twelve-segmented antennae, seems the most distinctive form, and possibly it is really distinct. Antennal segmentation is particularly dangerous to use as a species character in this group, since Santschi, in his original description of tracgaordhi, mentioned that segmentation varied between eight and nine in this species, and he later admitted that he had seen balsam prep- arations of the same species with only six segments, "the others more or less fused and indefinable," according to Bruch (1919: 400). The statement of Weber (1949 :3) in differentiating patrizii is accordingly of little value ; he states, ' ' The eight-segmented antennae separate this species from nine-segmented hewitti Arnold, ocidafa Emery, and traegaordhi Emery [sic], as does also the smaller size. The lobate fused frontal carinae of hewiffi are produced as a triangular plate. ' ' Santschi gave as the original measurement for traegaordhi worker, "Long. 1,5 mill.," while Weber giA^es 1.6 mm. as the length of his patrizii worker, so Weber's "smaller size" does not hold, from this information. D. hewitti differs from traegaordhi only in characters that usually differ between female and worker castes of the same species, so far as their respective descriptions go, so we cannot tell whether they are really distinct or not. The fronto-clypeal plate of these species is probably not so different as the characterizations indi- cate ; the posterodorsal margin is probably always nearly straight and steep, while the anterior margin is broadly rounded. The plate could therefore be variously described as convex, rounded, or even "triangular," according to what feature of the outline is stressed. I have two workers that belong to this complex, which I shall call D. traegaordhi at least provisionally, from Ebolowe, Cameroons (A. I. Good). [i03] The New World species so far described number eight, seven of which are keyed by Borgmeier (1949:205) ; the eighth species was added by Borgmeier (1954:191) ; see also Borgmeier 1957:122. Smith and Wing (1954) have redescribed and dis- cussed the type of the genus, D. testacea, from southeastern North America. Some of the species are described only from females, others only from workers, and the total material so far placed 342 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY under study is so small that the present arrangement cannot be considered close to final. In separating the species, too much reliance has been placed on antennal segmentation counts, which may be unstable in single species here as elsewhere ; the compara- tive statements on total length are also suspect, because measure- ments of ants this small have been notoriously inaccurate, due to lack of standardization as well as to poor optical aids. Perhaps four to six good species are represented. A definitive study must await more material and re-examination of the types. One addi- tional word of warning should be heeded in studying Discothyrea species, particularly the New World forms : it is entirely possible that some forms now established in the Americas are tramps originating in Africa or Asia. This possibility makes it impera- tive that the Old World forms all be duly considered in any New World study of the genus. [104] D. remmgtoni was described from a single worker from near La Foa, New Caledonia. Wilson has since collected two unique workers of this species or near it; one of these is larger than the type, the other is smaller; the larger specimen has the head as well as the alitrunk dark brown, while in the smaller one, the head is yellow and the alitrunk ferruginous. These specimens also show variation in the convexity of the alitruncal profile, and in the thickness and height of the petiolar node. They may or may not belong to the same species as remingtoni; with such limited material, it is impossible to decide. The larger specimen comes from Ciu, near Mt. Canala, 300 m., rain forest soil cover berlesate ; the smaller worker is from Mt. Mou, at about 180 m., both localities in New Caledonia. Another, much larger Discothyrea was taken by N. L. H. Krauss at an unspecified locality in southeastern New Caledonia (southeast of Noumea) ; this individual is 2-3 times the bulk of the workers, and has a faint sutural line behind the pronotum, indicating that it may be an ergatoid female. The eyes are larger, and the sculpture of the head is coarsely but shallowly punctate, with coriaceous intervals, the whole surface virtually opaque. This may be the ergatogyne of remingtoni, but if so, the caste differences are unusually strong in this species. BROWN : ANT TRIBE ECTATOMMINI 343 [105] DiSCOTHYREA MIXTA sp. nOV. (Figures 47, 48) Ilolotype worker: TL 2.4, HL (without mandibles) 0.66, HW 0.59, WL 0.61, scape L 0.43, funiculus L 0.52 mm. Form of head and body shown in Figures 47 and 48. Similar to the species D. velutina, D. hryanti and D. oculata, particularly the last, but smaller in size and with smaller compound eyes. Funiculus Figures 47 and 48. Discothyrea mixta sp. nov., paratype worker. Fig. 47, lateral view of head and body. Fig. 48, full-face view of head. 10-segmented, as determined from examination of two funiculi under compound microscope ; median segments extremely short, not distinct on the intact specimen as viewed with reflected light. 344 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Frontoclypeal platform narrower than in oculata, with slightly raised borders. Depressed scrobal areas distinct. Palpal seg- mentation undetermined. Propodeum concave, with a sharp concave margin above joining the propodeal angles, which are subrectangular as seen from above, though acute and dorsally inclined as seen from the side. Petiole thick disciform, as seen from above, truncate anteriorly and posterodorsal border without a distinct tooth or process. Postpetiole longer than broad, ovoid as seen from above, much larger than the succeeding segment. Head and body densely and rather coarsely punctulate, the punctulae nearly or quite contiguous, but becoming a bit more spaced on the gaster. Scrobal depressions indistinctly and finely transversely striolate, becoming smooth and shining mesad. Mandibles and appendages very finely and superficially punctu- late. Integument generally opaque, except for shining bottoms of scrobes and parts of the gastric dorsum, which are feebly shin- ing in some lights. Body, including mandibles and appendages, covered with dense, fine, very short, whitish decumbent pubescence, becoming more nearly erect on alitrunk, and obliterated from the shining parts of the scrobes. Color medium ferruginous, appendages more yellowish. Holotype [MCZ] one of five workers taken together at Bolahun, Liberia (J. C. Bequaert leg.). Paratypes [MCZ, USNM] are the four workers taken with the holotype, one of which has now been destroyed during the course of dissection. Varying only slightly in size, and scarcely at all in form, from the holotype. In the smallest workers (HL 0.63 mm.) the head tends to be a little narrower in proportion to its length, and the sides of the head may be a trifle more nearly straight. In some speci- mens, the occipital margin is a little straighter than as shown in the figure. Color varies from yellowish ferruginous to medium ferruginous. The lighter specimens show a more or less distinct pair of brownish-tinged longitudinal bands on the gastric dor- sum ; these may be internal structures showing through the integument. BROWN: ANT TRIBE ECTATOMMINI 34o ACKNOWLEDGEMENTS In preparing this paper, I received indispensable aid from a number of colleagues whose names are mentioned in the text in connection with their particular contributions. To all, I am deeply grateful. To Father Thomas Borgmeier of Jacarepagua, Brazil, and Dr. E. 0. Wilson, of Harvard University, I am par- ticularly indebted for long hours of effort on my behalf and for the many specimens they have loaned or given me. Dr. M. R. Smith of the U. S. National Museum, Dr. E. S. Ross of the Cali- fornia Academy of Sciences, and Dr. W. W. Kempf of Sao Paulo are also thanked for the loan of specimens under their care. Pub- lished bv a grant from the Wetmore Colles Fund. LIST OF REFEREXCES CITED AxDR^, Ern. 1889. 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Formiciden der australischeu Faunenregion. Ent. Mitt., 13:219- 229. Weber, N. A. 1938. New ants from stomachs of Bufo inarinus L. and Typhlops reticulatus (L.). Ann. Ent. Soc. Amer., 31:207-210. 1939. New ants of rare genera and a new genus of ponerine ants. Ann. Ent. Soc. Amer., 32:91-104. 1940. Eare ponerine genera in Panama and British Guiana. Psyche, 47:75-84. 1946. Two common ponerine ants of possible economic significance, Ectatomma tuberculatum (Olivier) and E. ruidum Eoger. Proc. ,Ent. Soc. Washington, 48:1-16. 1949. New ponerine ants from Equatorial Africa. Amer. Mus. Novit., 1398:1-9. Wesson, L. G., and E. G. Wesson 1940. A collection of ants from southeentral Ohio. Amer. Midi. Nat., 24:89-103, (cf. pp. 90-91). Wheeler, G. C. 1954. (Eeview.) The ants of California, by Thomas Wrentmore Cook. Ann. Ent. Soc, Amer., 46:618-619. BEOWN : ANT TRIBE ECTATOMMINI 355 Wheeler, G. C, and J. Wheelee 1952a. The ant larvae of the subfamily Ponerinae — ■ Part I. Amer. Midi. Nat., 48:111-144, 5 pis. l(tr/2b. The ant larvae of the subfamily Ponerinae — Part II. Amer. Midi. Nat., 48:604-672, 6 pis. Wheeler, W. M. 1905. An annotated list of the ants of New Jersey. Bull. Amer. Mus. Nat. Hist., 21:371-403. 1906. The ants of Japan. Bull. Amer. Mus. Nat. Hist., 22:301-328, pi. 41. 1914. The ants of the Baltic Amber. Schrift. phys.-okon. Ges. Konigs- berg, 55:1-142. 1915a. Some additions to the North American ant-fauna. Bull. Amer. Mus. Nat. Hist., 34:389-421. 1915b. Paranomopone, a new genus of ponerine ants from Queensland. Psyche, 22:117-120. 1915l'. Hymenoptera. [Family Formicidae of White Expedition to northern South Australia.] Trans. Eoy. Soc. S. Australia, 39: 805-823, pis. 64-66. 1916. Prodiscotliyrea, a new genus of ponerine ants from Queensland. Trans. Eoy. Soc. S. Australia, 60:33-37, pi. 4. 1917. A new Malayan ant of the genus Prodiscotliyrea. Psyche, 24:29- 30. 1922. Ants of the American Museum Congo Expedition. VII. Keys to the genera and subgenera of ants. VIII. A synonymic list of the ants of the Ethiopian Region. Bull. Amer. Mus. Nat. Hist., 45:631-1004. 1923a. Wissensehaftliehe Ergebnisse der schwedischen entomologischen Eeise des Herrn. Dr. A. Eoman in Amazonas 1914-1915. Ark. Zool., 15(7): 1-6. 1923b. Ants of the genera Myopias and Acanthoponera. Psyche, 30:175- 192. 1924. Ants of Krakatau and other islands in the Sunda Strait. Treubia, 5:1-20. 356 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 1925. Neotropical ants in the collections of the Royal Museum of Stockholm. Parti. Ark. Zool., 17A(8) :l-55. 1929. Ants collected by Professor F. Silvestri in Formosa, the Malay Peninsula and the Philippines. Boll. Lab. Zool. Portici, 24:2")-6-i. 1930. A new Emeryella from Panama. Proc. New England Zool. Club, 12:9-13. 1934. Contributions to the fauna of Itottnest Island, Western Australia. No. IX. — The ants. Jour. Roy. Soc. W. Australia, 20:137-103. 1936. Ants from Hispaniola and Mona Island. Bull. Mus. Comp. Zool., 80:193-211. Wheeler, W. M., and W. M. Mann 1914. The ants of Haiti. Bull. Amer. Mus. Nat. Hist.. 33:1-61. Wilson, E. O., and W. L. Brown, Jr. 19.')3. The subspecies concept and its taxonomic application. Syst. Zool., 2:97-111. INDEX included here are names of ant species, genera and higher categories mentioned in the body of the paper. Names mentioned in the Appendix and captions to figures are excluded where direct reference is made to them through bracketed numbers at the primary reference. The pages of the primary references are given in bold face below. Abbreviations for generic names are as follows : Ac. = Acanthoponera, D. = Diseothyrea, E. = Ecta- tomma, G. = Gnamptogenys, H. = Heteroponera, Pr. = Proce- ratium, E. = Rhytidoponera. alMloininalis, E., 202 Acanthoponera, 177-182, 188-194, 196, 217, 230, 244 acanthoponeroides, E., 202, 288-290 aeieulata, E., 202 acrista, E., 208, 211 aeuleaticoxae, G., 214, 21.1, 223, 227, 230, 237 acuminata, G., 227, 236 acupuncta, E., 204 acuta, G., 217, 227, 230, 232 aeneseens, E., 202 aerea, E., 209 Agioecomyrniex, 179 albiclava, G., 212, 221, 227, 230, 239 Alfaria, 177, 211, 213, 221-223, 230 alfaroi, G., 214, 223, 227, 234 algiricum, Pr. 247 Amblyoponini, 183, 230 Anacanthoponera, 177, 194, 196, 230 anceps, E., 200, 202 angustiloba, G., 229, 230 angustipleura, G., 229, 230 annulata, G., 223, 227, 234 antarctica, D., 253 araneoides, E., 197, 199, 202 Archiponera, 181 arcuata, G., 227, 230, 233 arcuata, E., 202 arnoldi, Pr., 243, 246, 247 aruana, E., 205 aspera, E., 200, 202 aterrima, G., 221, 227, 230, 240 atropurpurea, E., 202, 288, 292 Aulacopone, 177-182, 187, 206 aurata, E., 198, 202 aurea, H., 195 aztecum, E., 208, 211 banksi, G., 212, 215, 227, 230, 233 Barbourella, 178, 212, 215, 230 barnardi, E., 202 barretti, E., 202 bicolor, G., 227, 230, 240 bidens, D., 253 liinghami, G., 227, 230, 239 biroi, G., 227, 230, 238 bispinosa, G., 211, 215, 223, 227, 230 borealis, E., 202 Bradoponera, 180 lirasiliensis, G., 229, 230 brouni, H., 195, 196 brownii, H., 195, 263 bruchi, G., 212, 214, 227, 230, 236 brunnea, E., 202 bryanti, D., 253 bufonis, G., 222, 227, 230, 231 bufonum, G., 229 caeca, Ponera (Pr. ), 248 caeciliae, E., 204 358 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY calcarata, G., 229, 230 californicum, Pr. 247 Camponotus, 201, 219 carbonaria, R., 203 Cardiocondyla sp., 312 carinata, G., 228, 230 carinata, R., 202 carinif rons, H., 195, 197 carinifrons, Pr., 247 cavernicola, Pr., 248 cearensis, G., 229 cedarensis, R., 205 celtinodis, R., 202 ceramensis, R., 202 Cerapachyinae, 302 cerastes, R., 202 Chalcoponera, 177, 198, 201 chalybaea, R., 202 ehapmani, G., 221, 223, 227, 240 clinoopyx, R., 202 clarki, R., 203 clavata, Paraponera, 205 clavicornis, D., 249, 253 eoeca, Ponera, 335 Commateta, 178, 206, 212, 213, 230 concimia (Pr. Smith), G., 212, 223, 225, 227, 229, 230, 235 concimia (Santschi), G., 229, 230 concolor, E., 208 confine, E., 208, 210 confusa, E., 209 conica, G., 227 continua, G., 227, 235 convexa, R., 203 convexiceps, Pr., 247 cornuta, R., 203 costata, G., 227, 230, 238 coxalis, G., 211, 221, 223, 228, 230, 238 crassa, Ac, 191, 194 crassicorne, Pr., 248 crassicornis, D., 253 crassieornis, G., 228, 230, 239 crassinodis, R., 203 Crematogaster, 183 crenaticeps, G., 221, 228, 230, 240 cribrata, G., 228, 230, 241 cristata, R., 203 cristulata, R., 205 croeeum, Pr., 246, 247 croeaus, R., 203 curtula, G., 228, 230, 233 curvata, R., 204 Cylindromyrmex, 302 Dacetini, 180 dammermani, G., 228, 230, 241 densestriata, E., 208 denticulata, D., 253 dentinodis, H., 195, 197 diehli, G., 212, 228, 230, 241 Discothyrea, 178, 180, 182, 183, 186, 248-253 dixoni, R., 204 dolo, H., 194, 195, 197 douglasi, R., 203 dubia, R., 203 Ectatomma, 178, 180-182, 187-189, 197, 199, 205, 206.211, 212, 230 key to spp., 209 Ectatommini, 175-179, 183, 185 key to genera, 185 edentatum, E., 208, 210, 211 Electroponera, 181 Emeryella, 177, 181, 206, 211, 212, 215, 230, 325 emeryi (Forel), G., 228, 230 emeryi (Santschi), G., 229, 230 epinotalis, G., 211, 224, 228, 230, 241 eremita, R., 203 ericae, G., 229 Escherichia, 178 europaea, G., 181 europaea, Pr., 248 exarata, G., 228, 235 BEOWN : ANT TRIBE ECTATOMMINI 359 f erruginea, E., 203 fialai, Pr., 246, 248 fiebrigi, G., 229 fla\-icornis, E., 203 flavipes, E., 203 flindersi, E., 203 foreli, E., 203 foveolata, E., 203 froggatti, E., 202 fulgens, E., 203, 288, 292 fuliginosa, E., 203 gagates, E., 204 glabrior, E., 203 globus, D., 253 Gnamptogenys, 177-182, 187, 196, 206, 207, 211-241, 324, 325, 327 key to New World spp., 230 key to Old World spp., 237 goeldii, Ac, 193, 194 goyana, Ac, 192, 194 gracilis, G., 228, 230, 232 gramniodes, G., 221, 228, 239 greavesi, E., 203 gregoryi, E., 203 haeckeli, E., 203 haenschi, G., 214, 228, 234 hanieli, E., 203 haytiana, G., 212, 218, 221, 228, 230, 231 hartmani, G., 214, 228, 230, 234 Heteroponera, 176-183, 188, 189, 194- 197, 199, 206, 226, 244, 324 keys to spp., 196 hewitti, D., 253 liilaris, H., 196 hilli (Clark), E., 203 hilli Crawley, E., 203 Holcoponera, 177, 211-213, 216-219, 223, 225, 230, 324 horni, D., 253 horni, G., 228, 235 humilis, D., 253 hybrida, G., 229, 230 icta, D., 253 imbellis, H., 195, 196 impressa, E., 198, 199, 203 impressinodis, E., 202 inca, H., 196, 197 incisa, E., 198, 203 inermis, H., 196, 197 iuops, E., 203 inornata, E., 203 interrupta, G., 228, 235 intricata, E., 204 inversa, E., 208 iris, E., 208 irregularis, E., 209 isthmica, D., 253 isthmica, G., 230 itoi, Pr., 247 japonicum, Pr., 247 kalabit, G., 228, 240 kirki, H., 195 kuraudensis, E., 203 laciniosa, E., 200, 203 laevior, G., 228, 230, 240 laevior, E., 202 lamellinodis, E., 203 laticeps, E., 203 leae, D., 253 leae, H., 194, 196 levior, E., 203 lineata, G., 229 lobulifera, E., 208 lombokense, Pr., 247, 248 longigaster, Pr., 247, 248 lucida, G., 221, 228, 230, 240 lugens, E., 208, 209, 211 luzonensis, G., 228, 230, 241 macdonaghi, E., 208 maeretes, G., 228, 238 magnifica, G., 228, 230, 233 major, G., 228, 230, 233 major, E., 204 360 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY malaensis, G., 228, 230, 240 nialandensis, R., 204 maledieta, R., 203 manciim, Pr., 247, 248 maniae, R., 203 mayri, G., 230 mayri, Pr., 247, 248 mayii, R., 199, 200, 203 mecotyle, G., 215, 228, 237 mediatrix, G., 215, 228, 234 meieri, Bradoponera, 180 melinum, Pr., 243, 246, 247, 248 menadensis, G., 223, 228, 230, 239 menozzii, G., 228, 230, 237 metalliea, R., 198-200, 204 micans, R., 204 micrommatum, Pr., 241, 247, 248 mierops, H., 196 Mictoponera, 212 mina, G., 217, 218, 228, 230, 232 minor, Ac, 188, 191, 193, 194 minor, G., 229, 230 minuta, G., 222, 223, 228, 230, 231 nniuita, H., 196 mirabilis, R., 204 mixta, D., 249, 253 modesta, R., 205 moelleri, G., 228, 230,233 Monomorium sp., 277 mordax, G., 214, 215, 223, 225, 228, 235 morgani, E., 208, 210, 211 mucronata, Ac, 188, 192, 193, 194 mus, G., 228, 230 muticum, E., 208, 210 Myrmecina, 251 Myrmica, 199, 200, 225, 245 Myrmicinae, 183 neotropica, D., 253 nexa, R., 204 nigra, H., 196 nigra, R., 204 nigrifrons, G., 214, 228, 236 nitens, G., 229 nitens, R., 205 nitida, R., 204 nodifera, R., 204 nodosa, G., 228 normandi, Pr., 247, 248 nudata, R., 204 numeensis, R., 204, 289-295 numidicum, Pr., 247, 248 obscura, R., 203 oecidentalis, H., 196 oceidentalis, R., 204 oculata, D., 249, 253 opacior, R., 205 opaciventre, E., 207, 208, 209 Opisthoscyphus, 178, 212, 213, 222, 230 panamensis Santschi, G., 227 pananiensis (Weber), G., 228, 230 panamensis, H., 196, 197 panda, G., 221, 228, 230, 239 papuanum, Pr., 247, 248 Paranomopone, 177, 194, 196 Paraponera, 176, 178, 180-182, 185, 199, 205, 207 Paraponerini, 175, 176 Parectatomma, 177, 206, 212, 213, 222, 230 patrizii, D., 253 paulina, G., 229, 230 peninsularis, R., 204 pergandei, Pr., 243, 246, 247, 248 permagnum, E., 208, 211 pernambucana, G., 229, 230 peruviana, Ac, 193, 194 petiolata Viehmeyer, R., 203 petiolata Clark, R., 204 Pheidole, 183, 312 pilosula, Myrmecia, 202 pilosula, R., 204 BROWN : ANT TRIBE ECTATOMMINI 361 planidens, E., 209, 'JIO Platythyreini, 178, 179 plaumanni, Ac, 192, 193, 194 pleurodon, G., 229, 230, 233 Polyrhachis sp., 277 Poneraeantha, 178, 206, 211, 212, 213, 215, 230 Ponerinae, 175, 176, 178 Ponerini, 179 porcata, G., 223, 229, 230, 233 posteropsis, G., 229, 230, 238 poweri, D., 248, 250, 253 Prionopelta, 177, 230 Probolomyrmex, 178 Proeeratiini, 175, 176, 178 Proeeratium, 176-178, 180, 182, 183, 187, 231, 241-248, 250, 252 Prodiscothyrea, 178, 248, 249, 253 Pseudosphincta, 248 Pseudosysphineta, 178, 248, 250, 253 pulchella, E., 204, 289, 290, 291 pulchra, E., 204 punctata, E., 200, 204 punctigera, E., 204 punctigerum, E., 209 punctiventris, E., 204 purensis, G., 228 purpurascens, E., 204 purpurea, E., 204 quadriceps, E., 203 quadridens, E., 207, 209 quitensis, G., 230 rastrata, G., 215, 229, 230, 237 recta, G., 229, 230 reflexa, E., 204 regularis Mayr, G., 229, 235 regularis (Santschi), G., 227, 230 reichenspergeri, G. or Ac, 196, 217, 218, 229-231 relicta, Aulacopone, 206 relicta, G., 217-221, 223, 229-231 relicta, H., 180, 195, 196 relictum, Pr., 243, 245, 247, 248 remingtoni, D., 253 reticulata, E., 204 Ehopalopone, 177, 211, 212, 217-222, 226, 230, 324 Ehytidoponera, 177, 178, 180, 182, 187, 197-205, 207 key to New Caledonia spp., 288 richteri, G., 230 rimulosa, G., 229, 235 romani, G., 227 rossica, Pr., 248 rotundiceps, E., 204 rufescens, E., 204 rufithorax, E., 204 rufiventris, E., 204 rufonigra, E., 204 ruginoda, Poncra or E., 202 rugosa, E., 202 rugulosum, Pr., 248 ruidum, E., 207, 208, 209, 210 rustica, G., 229, 230, 233 sauteri, D., 253 scaberrima, E., 204 scabra, H., 196 seabra, E., 204 scabrosus, G., 212, 228, 230 schlaginhaufeni, E., 204 schmitti, G., 181, 211, 215, 223, 227, 229, 230, 233 schubarti, G., 229, 230, 237 schwarzi, Ac, 190, 194 schwebeli, H., 195, 198 scrobiculata, E., 205 sculptior, D., 253 sebastiani, G., 228 semicircularis, G., 227 semiferox, G., 215, 229, 234 sexarticulata, D., 250, 253 silaceum, Pr., 241, 245, 246, 247, 248 simillima, E. (Prionopelta), 230 simplex, G., 229, 230, 232 362 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY simplicoidea, G., 229, 230, 233 simulans, G., 211, 222, 229, 230, 231 socrula, R., 203 socrus, R., 204 Spaniopone, 178, 212, 217, 218, 221, 230, 324 spatiata, R., 203 spininodis, Ac, 190, 191, 194 spiralis, G., 229, 230, 239 splendens, G., 228, 230 splendida, G., 229 spoliata, R., 198, 204 Stictoponera, 177, 206, 211, 212, 213, 217-222, 230 stictum, Pr., 243, 244, 247, 248 stolli, G., 228, 230 striata, R., 203 striatula, G., 211, 223, 229, 230, 233 stridulator, R., 203 strigata, G., 217, 229, 230, 232 strigosa, R., 200, 204 strigosum, E., 209 striolata, G., 221, 222, 229, 230, 235 subcyanea, R., 205 sulcata, G., 229, 236 Sysphincta, 178, 241, 243, 244, 247 Sysphingta, 178, 241 taivanensis, G., 221, 229, 230, 239 Tammoteca, 178, 206, 212, 213 tasmaniensis, R., 205 taurus, R., 198, 205 tefPensis, G., 229, 230, 233 tenuis, R., 205 testacea, D., 248, 253 tornata, G., 211, 214, 229, 236 tortuolosa, G., 230, 236 toschii. Pp., 243, 246, 247, 248 trachypyx, R., 205 traegaordhi, D., 253 transversiruga, R., 205 triangularis, G., 212, 215, 225, 230, 237 trigona, G., 230, 237 tuberculatum, E., 207, 208, 209, 211 turneri, R., 205 turtoni, D., 253 tyloxys, R., 205 Typhlomyrmex, 177 Typhlomyrmicini, 177, 219 varians, R., 204 velutina, D., 248, 250, 253 versicolor, R., 205, 289-293 vestitum, Pr., 248 vietoriae, R., 200, 205 vidua, G., 229, 230 violacea, R., 205 viridis, R., 205 wagneri, Ac, 192, 194 waigeuensis, R., 204 wallacei, R., 205 wasmanni, G., 230, 233 watasei, Pr., 244, 248 wheeleri, Archiponera, 181 wheeleri, G., 230, 233 Wheeleripone, 178, 212, 219-221, 230 wilsoni, R., 205, 289, 291, 294, 295 yorkensis, R., 205 ypirangensis, G., 229 Bulletin of the Museum of Comparative Zoology AT HARVAED COLLEGE Vol. 118, No. 6 THE SPIDER SUBFAMILY CLUBIONINAE OF THE UNITED STATES, CANADA AND ALASKA (ARANEAE: CLUBIONIDAE) Bt Robert J. Edwards With Twenty-Three Plates CAMBRIDGE, MASS., U.S.A. PRINTED FOR THE MUSEUM June, 1958 Publications Issued by or in Connection WITH THE MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD COLLEGE Bulletin (octavo) 1863 — The current volume is Vol. 118. Breviora (octavo) 1952 — No. 86 is current. Memoirs (quarto) 1864-1938 — Publication was terminated with Vol. 55. JoHNSONiA (quarto) 1941 — -A publication of the Department of Mollusks. Vol. 3, no. 35 is current. Occasional Papers of the Department op Mollusks (octavo) 1945 — Vol. 2, no. 21 is current. Proceedings of the New England Zoological Club (octavo) 1899-1948 — Published in connection with the Museum. Publication terminated with Vol. 24. The continuing publications are issued at irregular intervals in num- bers which may be purchased separately. Prices and lists may be obtained on application to the Director of the Museum of Comparative Zoology, Cambridge 38, Massachusetts. Of the Peters "Check List of Birds of the World," volumes 1-3 are out of print ; volumes 4 and 6 may be obtained from the Harvard Uni- versity Press; volumes 5 and 7 are sold by the Museum, and future volumes will be published under Museum auspices. Bulletin of the Museum oi Compaiative Zoology AT HAKVAED COLLEGE Vol. 118, No. 6 THE SPIDER SUBFAMILY CLUBIONINAE OF THE UNITED STATES, CANADA AND ALASKA (ARANEAE: CLUBIONIDAE) By Robert J. Edwards With Taventy-Three Plates CAMBEIDGE, MASS., U.S.A. FEINTED rOE THE MUSEUM June, 1958 No. 6 — The Spider Subfamily Cluhioninae of the United States, Canada and Alaska (Araneae: CluhionidaeY By Robert J. Edwards Introduction. The subfamily Clubioninae has been in a state of confusion for some time, and little has been done to rectify this condition except for the fine paper by Gertsch (1941, Amer. Mus. Novitates, no. 1148) on the ahhotii group of the genus Clul)iona. Many of tlie original descriptions and figures are of no diagnostic value. During this study it became evident that common and widely distributed species have subspecies. Lack of sufficient material prevented a careful study of these. Acknowledgments. This study was conducted in the zoologi- cal laboratories of the University of Rochester under the super- vision of the late Dr. S. C. Bishop, to M'hom I am greatly indebted for his constant interest, helpful advice and criticism. I am especially grateful to Dr. W. J. Gertsch of the American Museum of Natural History, and to the late Miss E. B. Bryant of the Museum of Comparative Zoology, for the use of the very extensive collections at those institutions and for very valuable suggestions and information. Large collections w'ere also made available for study through the courtesy of Dr. H. K. Wallace, University of Florida; Mr. T. Kurata, Royal Ontario Museum of Zoology ; and Dr. H. Dietrich, Cornell University. All figures were drawn by Miss Carolyn Fallon, former staff artist for the Department of Zoology at the University of Rochester. 1 This thesis was submitted, in partial fulfillment for the degree of Doctor of Philosophy, to the faculty of the University of Rochester in May 19.51. Since it remained unpublished, I received permission from the author to prepare it for publication. It was rewritten, shortened, and keys to the females of the genus Cluhiona, and plates of the cleared epigyna were added. I want to express my thanlis for the help of Prof. K. Cooper ; to the edit