THE MYRMECOPHILOUS PTINIDAE (COLEOPTERA), WITH A KEY TO AUSTRALIAN SPECIES

JOHN F. LAWRENCE' AND HANS REICHARDT"

CONTENTS INTRODUCTION

Alistract 1 The family Ptinidae includes about 50 Introduction _ 1 genera and 600 species, which are dis- Notes on Terminology , 2 tributed the of Classification of Myrmecophiles 3 throughout major regions The Australian Ectrephines 5 the world but are particularly abundant Key to the Genera and Species of Myrme- in the drier parts of the subtropical and cophilous Ptinidae Occurring in Australia __ 7 temperate zones. Of the described species, Notes on the Genera 8 almost 50 have been recorded from the Origin and Evolution of Myrmecophily in the Ptinidae 10 nests of ants and at least 42 appear to be Haliitats and Food Habits 10 true inquilines. The purposes of this paper Bi()lo,g\- of the Myrmecophilous Species .. 11 are 1 to certain in ( ) propose changes the Origin of Myrmecophily 13 classification of the myrmecophilous species E\olutionary Trends 15 on the basis of discovered Catalogue of the Myrmecophilous Ptinidae 16 newly characters, List of Ant Host Species and Associated In- (2) to provide a key to the species oc- 21 in quilines curring Australia, ( 3 ) to summarize what Acknowledgments 22 is presently known about the biology and Literature Cited 23 distribution of mynnecophilous ptinids in the form of a and to ABSTRACT catalogue, (4) spec- ulate on the possible origin and evolution A review is presented of the current of myrmecophily in the family. knowledge on ptinid beetles known to Those species of Ptinidae which are here occur with ants. Inckided are: 1) a cata- considered to be true ant guests fall into of the 2 a five logue myrmecophilous Ptinidae, ) main groups: the two tropical Amer- summar)' of biological and distributional ican genera Gnostus and Fabrasia, the data, 3) a revision of the genera and key South African Diplocotidus, the southeast to the species occurring in Australia, and Asian genus Myrmecoptimis, and the sev- a 4) discussion of the origin and evolution eral Australian genera which have usually of mynnecophily in the family. Eight gen- been placed in the subfamily Ectrephinae. era and 43 species are treated, and 8 The first four of these will be discussed previously recognized genera are placed briefly below and the last will be treated into synonymy. A Hst of ant host species in detail in a later section. is also given. In addition to the above, several ptinids have been recorded from the nests of ants. ^ Museum of Harvard Comparatixe Zoology, Tlie Ptinus Sturm and University, Cambridge, Mass. European suhpilostis - P. Panzer have been taken Departamento de Zoologia, Secretaria da Agri- scxpunctatiis cultura, Sao Paulo, Brazil. with the ants Acanthomyops fuliginosus

Bull. Mus. Comp. Zool., 138(1): 1-28, April, 1969 1 Vol. No. 1 2 BuUetin Museum of Comparative Zoology, 138,

scribed from the same area. The (Latreille) and A. hnmneus (Latreille); species both beetles occur in other habitats as well species differs considerably from other and (Linsley, 1944; Howe, 1959). The species mvrmecophilous ptinids probably rep- an line. The construc- Diplocotidus moscii Braiins was described resents independent the sub- from specimens collected under a rock with tion of the prothorax (Fig. 29), the ant Mcmnopliis peringucyi Emery in quadrate and concave clypeus (Fig. 3), South Africa (Wasmann and Brauns, 1925). and the aedeagus with reduced parameres the are all The Andreae (in litt.) has examined type (Figs. 26, 27) unique. grooves to and considers it to be an old and rubbed on the pronotum are somewhat similar Fair- those of P. and some of the South specimen of Pscudomczium coqucrcli ( coqucrcli but the does not maire). This record with ants is probably African gibbiines, species to related to of these. accidental. Finally, the species Leioptinus appear be closely any — hutteli infhtus \\'asmann (1928) was described yiyrmccoptinus. Myrmecoptinus 1916 was described from from a nest of Mijnnicoria amchnoides ^^'asmann ( ) Smith on Sumatra. This species may be Sumatra and associated with the ant Crc- hutteli Forel. Was- myrmecophilous, but from the description mafogastcr Although several and illustration it does not appear to us to mann's type has not been examined, from belong in the family Ptinidae. specimens have been seen Geylon, Gnostus and Fohrosia.—These two gen- Borneo, China, North Vietnam, and Burma that well with the era, which contain all of the New ^^'orld conform fairl>- descrip- in the same mvrmecophiles, have been treated else- tion and probably belong where (Lawrence and Reichardt, 1966a) group. The elytral humeri are well devel- and will not be discussed in detail here. oped, the wings are present, the clypeus is that in Ptinus Since the publication of the above revision, flattened and triangular like 2 antennal modifications are however, two new records have come to ( Fig. ) , and our attention. A specimen of Fohrasia completely absent. The pronotum is rela- horfimcicri Lawrence and Reichardt, in the tively simple but has a well-defined, nar- collection of the Departamento de Zoologia, row, transverse groove at the base, which was collected at P090S de Caldas, Minas connects lateral pairs of processes resem- with the Gerais, Brazil, in February, 1933, bling the trichomes found in other inqui- scnc.x ant Camponotus (Mijnnohrachys) lines. The color is black and the vestiture This a southern (F. Smith). represents is sparse, consisting of fine hairs as in other extension and a new of ant range species m\rmecophilous forms. One of the speci- host. A of F. wheeleri Lawrence specimen mens examined was collected by Mjoberg and Reichardt, in the collection of the in an ant nest, but the locality is indecipher- CaHfornia Insect Survey, was collected 6.3 able. These beetles are probably myrme- miles west of Tehuantepec, Oaxaca, Mexico, cophiles, but they do not appear to be on July 21, 1952, by E. E. Gilbert and C. D. closely related to the other groups. MacXeill (ant host not recorded). This is —The majorit\' of myrme- the first record of Fabrosia from Mexico Ectrephinae. cophilous Ptinidae occur in Australia and and represents a considerable northern are placed in the subfamily Ec- range extension for F. wheeleri, kno\\n generally These will be discussed further previously from Colombia. trephinae. below. Diplocotidus.—\\'ith the removal of D. moseri (see above), this genus contains a Notes on Terminology single species, D. formicola Peringuey, which occurs in the interior of South Africa. There are a few terms used in the key

litt. to true and discussions that need clarification, Andreae ( in ) considers these be may the same myrmecophiles and has seen two unde- although the usage is essentially Myrmecophilous Ptinidae • Lawrence and Reichardt

as that found in Lawrence and Reichardt two processes arising on different sides of (1966a). a cleft or fovea. Perhaps this is a device This is to Chjpcus. equixalent the area for collecting a drop of liquid. In the fol- referred to Stickne\- 1923 as the all by ( ) post- lowing discussions of these prothoracic ch'peus. In most Ptinidae it consists of a structures are called trichomes. Patches of well-defined, broad, triangular plate lying short hairs or pile are usually found on the just in front of and between the antennal last antennal segment (Figs. 17-19), and in fossae, the anatomical base of the plate some species they are concealed within a the of the forming apex triangle (Fig. 2). concavity at the tip of the segment (Figs. The word "apex" in the text refers to the 15, 16). Occasionally the hairs may be anatomical apex to which the labrum is absent (or rubbed off), but the extremely attached. Many of the myiTnecophilous fine punctation may indicate the presence

ha\e a more and tumid of a area 22 . species elongate glandular ( Fig. ) clypeus (Figs. 1, 4, 7, 8), while in several Measurements. The total length is the others it consists of a flattened, cleft plate maximum length as measured from above (Figs. 5, 6, 9, 10). (tips of elytra to the most anterior point

TricJwmes. In the to Torre- of the . E\\^ supplement clypeus ) EL and refer to great- Bueno's Glossary of Entomology (Tulloch, est elytral length and greatest elytral width, I960), trichomes are defined as "modified respectively. hairs present on certain mynnecophilous insects which give off secretions which ants CLASSIFICATION OF MYRMECOPHILES imbibe." the word be used Although may The placement of myrmecophiles in a to describe any group of hairs which are system of classification has always pre- to be it is thought glandular, commonly sented a problem to taxonomists because of restricted to distinct setiferous processes the development in these fonns of special usually found on the prothorax, antennae, adaptations associated with inquilinous or abdomen of elytra, myrmecophilous habits. In some groups normally consena- beetles. statements about the Actually, tive characters, such as the number or function of these structures are seldom, if shape of antennal segments, may be highly ever, accompanied by histological or behav- variable, while in others entirely new struc- ioral studies (see p. 12). tures, such as trichomes, may be present. In the Ptinidae there are four main kinds It is not surprising that a number of these of structures which be 1 glandular: ( ) may species have been placed in separate fam- femoral elytral pores, (2) trichomes, (3) ilies until more careful studies have clarified pronotal trichomes, and (4) antennal pilos- their relationships. A few of the many ity. The first two structures are found only beetle families originally proposed for in the genus Fabrasia; the elytral pores the inclusion of inquilines but no longer secrete a white waxHke substance. In some recognized are the Pseudomorphidae ptinids, such as Ectrephes fonnicanim Carabidae ( ) , Clavigeridae ( Pselaphidae ) , Pascoe 28 and Enasiha tristis Olliff ( Fig. ) , Rhysopaussidae (Tenebrionidae), and Cos- the pronotal trichomes bear distinct pubes- syphodidae ( Tenebrionidae ) . cence and resemble those in the occurring During the last hundred years there has Paussidae (Darhngton, 1950). Many spe- been considerable difference of opinion as cies, however, have lateral clefts and to the placement of the myrmecophilous processes 30-32, that to (Figs. 34) appear ptinids within the higher classification of be to the trichomes of E. homologous for- the Coleoptera. Although Westwood micannn but that do not bear patches of (1855) considered Gnostits to be related to hairs. One feature that all of these struc- the Xylophaga of Latreille, later workers tures share is the proximity of the tips of placed it near the Paussidae, Pselaphidae, 4 Bulletin Museum of Comparative Zoology, Vol. 138, No. 1

the or Scydmaenidae; the taxonomic history of species in the family has led to proposal The the genus is discussed by Lawrence and of several more subfamilies. genus Reichardt (1966a). Pascoe (1866) com- Ectrephes Pascoe, along with Westwood's and pared his genus EctrcpJies with Gnostus genera Diplocotcs and Polyplocotcs and the Paussidae. Gemminger and Harold Olliffs Enasiba and Diphobia, was placed while Gnostus was (1869) proposed the family Gnostidae to in the Ectrephinae, include both Gnostus and Ectrephes, while placed in the Gnostinae. Mjoberg (1916) the di\ided the into three sub- W'asmann ( 1894 ) proposed family ectrephines most Ectrephidae for Ectrephes and \\'estwood's families: Polyplocotinae, containing those two genera Diplocotcs and Polyplocotcs. of the species; Paussoptininae, for Sharp and Muir (1912) noted the similar- species with flattened antennal segments; ities between Pohjplocotes and Ptinus with and Ectrephinae, including those species respect to the basal connection of the lateral originally placed in Ectrephes. He placed and median lobes of the aedeagus. In 1916, Diphobia and the South African genus both ^^'asmann and Mjoberg treated the Diplocotidus Peringuey in the subfamily ectrephines as members of the Ptinidae. Ptininae, because of the lack of antennal 1926 out the modifications. Martinez and Viana Forbes ( ) pointed relationship (1965) for of Gnosius to the ptinids on the basis of proposed the name Fabrasiinae the wing venation, but the complete absence of genus Fabrasia, and Costa Lima (1962) wings in the ectrephines made a similar placed his genus Plaumanniola in the comparison impossible. Our studies defi- Plaumanniolinae. The latter genus was the nitely confirm the inclusion of this group found to belong to family Scydmae- within the Ptinidae. Diplocotcs familiaris nidae (Lawrence and Reichardt, 1966b). are of the that there are at (Olhff ), one of the least speciaUzed of the We opinion Australian species, differs little from other least five independent stocks of myrme- wingless ptinids, and a gradation can be cophilous ptinids: Gnostus, Fabrasia, Dip- seen from D. familiaris to the highly mod- locotidus, Myrmecoptinus, and the ectre- ified forms discussed below. The aedeagi phines. The last group may be divided into of D. familiaris (Fig. 2.3), D. laticornis two subgroups (see below), but their 24 and derivation from a ancestor is indi- ( Lea )( Fig. ), Ectrephes formicarum single Pascoe (Fig. 25) all have the same basic cated by the numerous parallelisms dis- structure found in Gnostus and Fabrasia played by the two. Myrmecoptinus differs (Lawrence and Reichardt, 1966a) and in only slightly from the ptinines and could various other ptinids (Hinton, 1941). hardly be placed in a separate subfamily. \\'ithin the Ptinidae, the classification of Gnostus and Fabrasia, on the other hand, these myrmecophilous species presents a are (juite distinct, although they are deriv- problem, owing to the present need for a able from some of the more elongate and complete revision of the family. Two sub- parallel-sided winged Ptininae. Diploco- groups, the Gibbiinae and the Ptininae, tidus is also a very distinct form and might are usually recognized. In the former, the well be placed in a separate subfamily, for which a new elytra are laterally compressed and partly name would be required. The enclose the reduced abdomen; the group Ectrephinae are linked with the Ptininae includes Gibbium, Mcziuni, Damarus, and by forms like D. familiaris, and the only a few other genera, the species of which character which could be used to distin- are most abundant in South Africa. The guish the two groups is the form of the remainder of the ptinids are usually in- clypeus, which is elongate and tumid or cluded in the Ptininae, which contains a cleft in Diplocotcs and its relatives. If sub- diverse assemblage of fonns. families or tribes are recognized, then, four The inclusion of \arious m)rmecophilous would be required: one for each of the Myrmecophilous Ptixidae • Lawrence and Reichardt

Neotropical groups, one for the South In Ectrc))Jics Pascoe the antennae are 3- African Diplocotidtis, and one for the Aus- segmented and the last segment forms a trahan ectrephines. solid, cylindrical club (Fig. 15), while in The second problem is whether or not Bitrcphcs Oke they are 2-segmented and these myrmecophilous forms are different the terminal segment is somewhat wedge- other enough from ptinids to be placed in shaped (Fig. 35). Finally, in the genus separate groups and what rank to assign to Encisiha Olliff the antennal segments are these groups. \\'ithin the large subfamily peculiarly shaped, as in Figure 16, and the Ptininae, there are several other clusters of pronotum is elongate, with four trichomes. genera which are just as distinct, or more \\'e feel that the above classification is so, if one disregards the special adaptations unnatural for several reasons. First, any of myrmecophiles. These include: Dieij^oiis classification which relies so heavily on a and Xylodes from Mauritius; Casopus and single character complex is suspect. Fur- its relatives from the Canary Islands; CijJin- thermore, the particular characters used are d I opt inns and its relatixes from Southeast highly variable vvithin the group and are Asia; iSiptus and its allies; SpJwericus; and associated with a narrow specialization; Tri(ionop,cniiis. Until a thorough study is thus they would have a low taxonomic of made the generic classification in the weight according to the tenets originally family Ptinidae, we think it serves no useful proposed by Darwin. The adaptive nature purpose to place the incjuilinous species in of the antennal modifications becomes separate subfamilies or tribes. obvious if one considers the numerous cases of convergence with respect to these charac- THE AUSTRALIAN ECTREPHINES ters in unrelated myrmecophilous groups. The antenna of The 35 described species of myrmeco- short, stump-like Ecirephcs for is philous ptinids inhabiting Australia are formicarum (Fig. 15), instance, ver\' similar to that of and the same generally placed within 14 genera, which Gnosius, are characterized mainh- on the basis of type of antenna mav be found in several unrelated such as Euclarkia antennal segmentation. Diphobio Olliff m}rmecophiles,

, Trochoideus contains species with relatively simple, 11- ( Colydiidae ) ( Endomychi- and Poussus antennae 21 . dae), Fustiger (Pselaphidae), segmented ( Fig. ) Those Another is the lack species in which the penultimate antennal (Paussidae). objection of correlation of antennal characters with segment is enlarged (Figs. 17, 18) are other Paus- placed in three genera according to the features; Diplocotcs armicoUis, and total number of antennal segments: Dip- soptinus laticomis, Mesectrephes kingi, for have different antennal locotes Westwood (11), Decemplocotcs instance, very 10 tvpes and yet are similar with respect to Mjoberg ( ) , and Poh/pJocotes Westwood the and A further criti- (9). In the monotypic Hexoplocotes Lea, pronotum clypeus. cism be made of the use of meristic the antennae are 6-segmented and the last may

characters number of antennal , two segments are enlarged and flattened ( segments ) which should be low for (Fig. 22). In several species the antennal given weight obvious reasons. The loss or segments are strongly compressed and may independent fusion of antennal has been be partially or completely fused to form a segments demonstrated in a number of beetle club (Figs. 19, 20); these fonns have been groups 1965, for the Coccinellidae; Law- placed in Fau.ssoptinus Lea (10 segments; (Chapin, rence, 1967, for the and extreme see p. 9). Lcaptinus Mjoberg (10 seg- Ciidae), reductions in ments), Paussoceros Mjoberg (9 segments), parallel myrmecophilous beetles have Pseudectrephes Pic (7 segments), Monec- been discussed by Darlington for the Paussidae Park trephes Mjoberg (3 segments; see p. 10), (1950) and (1942) and Mescctrephes Mjoberg (3 segments). for the Pselaphidae. 6 Bulletin Museum of Comparative Zoologij, Vol. 138, No. 1

In an attempt to find other characters to easily derivable from that occurring in most of use in classifying these AustraHan species, Ptinidae and since the simplest type a study \\as made of the clypeal region. antenna also occurs in the first group, it is This structure had been ignored in the past reasonable to assume that this clypeal type because most of the specimens were glued is primitive. The joint possession of a to cards. In most Ptinidae the clypeus is relatively complex and derived clypeus by broadly triangular and somewhat flattened, members of group II and the occurrence and the labrum is small and transverse within each group of characteristic antennal evidence (Fig. 2). \\'ith the exception of Myrmc- and pronotal types present strong is one. The coptinus hiittcli, all of the myrmecophilous that the division a natural of certain antennal species have the clypeus somewhat mod- parallel development ified. The Australian forms fall into two modifications in the two groups is not distinct groups on the basis of clypeal surprising for myrmecophiles and would of structure. In the first group (Figs. 1, 4, 7, tend to support the ultimate derivation a ancestor. S), the clypeus is slightly to strongly tumid both stocks from common and subtriangular, with the median portion, Within group I, it is difficult to segregate too at least, distinctly punctate. In the second the species further without using many names. If the antennae are group (Figs. 5, 6, 9, 10), the clypeus is generic used, elevated, flattened, and deeply cleft at the five genera might be recognized, when in the apex, forming a U-shaped clypeal plate, meristic differences are ignored an which is impunctate and shiny. The species group characterized by having enlarged included in each group are given below penultimate antennal segment. In the struc- D. P. kiti- according to their present generic assign- ture of the prothorax, armieollis, ments: eornis, P. eorinaticeps, M. kingi, and B. are similar 30 and Group I: Diphohia familiari.s, D. fovcafa, cunciformis (Figs. 35), D. infricota, D. Ion

28 . is the oldest name for sufficient to place the species with a reason- ( Fig. ) EetrepJies do not able degree of certainty in one group or the this genus. The remaining species other. \'ary considerably \\'ith respect to pronotal Since the first type of clypeus is the most and antennal characters and are all in- Myrmecophilous Ptinidae • Lawrence ami Reichardt 7

eluded in the genus Pohjplocotes West- at tlu' apex 4 brushes of yellow hairs wood. D. intricatus (Lea) Antennal segments 6-10 as long as or the classification Although generic pre- longer than broad, not compressed, and sented here represents an improvement over without brushes of yellow hairs 8 the older classification, further modification ^ Elytral punctures very small and shallow, scarcely visible under lower magnifica- may still be needed. When more species tion; pronotum without basomesal fovea or character complexes are studied, it may D. familiaris (Olliff) be to further sulodivide the necessary gen- Elytral punctures larger and more deeply era. A key to the genera and species of impressed; pronotum with basomesal fovea D. ectrephines is included below, followed by foveatus (Lea) Pronotum expanded apically forming 2 a brief discussion of each genus as it is lateral projections _.. 10 here delimited. A resume of the proposed Pronotum without apicolateral projections 11 classification, with generic and specific 10. Pronotum about as long as wide at base, of synonymies, is given in a later section. apicolateral projection consisting a sharp tooth; cKtral punctures bearing very short, decumbent hairs to the Genera and Key Species — D. armicollis Lea of Myrmecophilous Ptinidae Pronotum more than 1.25 X as long as wide at base, apicolateral projection con- Occurring in Australia sisting of a rounded process bearing several small 1. Clypeus usually subtriangular (Figs. 1, tubercles; elytral punctures 4), slightly to strongly tumid, occasionally bearing relatively long, suberect hairs elexatfd, forming a narrow, median ridge - D. foveicoUis Olliff H- Total 1.50 (Figs. 7, 8) (genus Diplocotes West- length more than mm; basal of not in wood ) 2 impression pronotum expanded middle D. howittanus — Chpeus elevated, flattened, and deeply Westwood Total less than 1.25 basal cleft at apex, forming a U-shaped clypeal length mm; of plate (Figs. 5, 6, 9, 10) _.. 20 impression pronotum slightly expanded in middle a shallow fovea 2. Antennae 11-segmented 3 forming D. minutus Oke — Antennae with less than 11 segments 12 12. Antennae 10-segmented 13 3. Antennal segment 10 not enlarged, shorter Antennae with less than 10 segments 15 than segment 11 (Fig. 21) 4 13. Antennal segments 2^10 strongly com- — Antennal segment 10 enlarged, longer pressed (Fig. 19) D. laticornis (Lea) than segment 11 and usually as long as Antennal segments 2-10 not compressed ^ 14 segments 8 and 9 combined ( Fig. 17 ) 9 14. Pronotum with basolateral trichomes 4. Antennal segment 11 very long, as long D. hrevipennis (Pic) as segments 8 to 10 combined Pronotum without basolateral trichomes .. D. longicornis ( Lea ) D. strigicollis Lea — Antennal 11 not as segment long as seg- 15. Antennae 9-segmented 16 ments 8 to 10 combined 5 Antennae with less than 9 segments 19 5. Pronotum without basolateral trichomes, 16. Clypeus elevated in middle, forming a occasionally with weak lateral foveae 6 narrow ridge ( Fig. 7 ) ; pronotum shorter — Pronotum with distinct basolateral tri- than wide at base, with apicolateral teeth chomes and deep lateral foveae 7 D. carinaticeps ( Oke ) 6. Elytra shorter and broader, EL/EW less — Clypeus somewhat tumid, but not fonu- than 1.25, dark metallic blue in color, ing a narrow ridge (Fig. 8); pronotum the punctation not distinctly seriate, each more than 1.20 X as long as wide at puncture rounded and bearing a relati\'ely base, without apicolateral teeth 17 long, subercct hair D. mciallicus (Lea) 17. Antennal segment 9 longer than 8 — and Elytra longer narrower, EL/EW D. apicalis ( Oke ) more than 1.2.5, reddish brown in color, — Antennal segment 8 longer than 9 18 the punctation distinctly seriate, each 18. Total length less than 2.50 mm; elytral puncture elongate and bearing a very punctures bearing very short, decumbent short, decumbent hair __. hairs D. similis (Oke) D. Lea — myrmecophilus ( ) Total length more than 2.50 mm; elytral 7. Antennal segments 6-10 shorter than punctures bearing relatively long, sub- broad, compressed, each segment bearing erect hairs D. pilosus (Mjoberg) 8 Bulletin Museum of Comparative Zoology, Vol. 138, No. 1

19. Antennae with 3 segments, the last seg- 28. Pronotum w ithout basal impression; mesal ment strongly compressed portion of pronotal disc not longitudinally P. nitidus Westwood D. kingi ( Westvvood ) strigose mesal — Antennae with 2 segments, the last seg- — Pronotum with basal impression; disc ment wedge-shaped (Fig. 35) portion of pronotal longitudinally at least 34) 29 D. cuneifarmis ( Oke ) strigose, basally (Fig. 29. Antennal 3 almost 3 as 20. Pronotum with a narrow, median, longi- segment X long as wide P. Lea tudinal groo\e, and with 2 lateral and 2 longipes than 1.50 mesal trichomes w ithin a trans\ erse basal — Antennal segment 3 not more as as wide 30 impression; antennal segments 2 and 3 X long 30. of disc very broadly joined, segments .3-5 somewhat Highest portion pronotal and and triangular or wedge-shaped with the base finely sparsely punctate shiny; anterior of with a trans- broader than the apex, and segments edge pronotum verse row of ( .34 ) .— 6-10 somewhat compressed (Fig. 16) deep punctures Fig. P. \\'estwood (genus Enasiha Olliff) 21 longicolU.s- — of disc more — Pronotum without a median, longitudinal Highest portion pronotal coarsely and densely punctate, granulate, groove, without trichomes or with only or not anterior of 2 lateral ones 22 strigose, shiny; edge pronotum without row of deep punctures .. 31 21. Head, pronotum, and bases of elytra 31. Ehtral punctation dual, with series of bearing short, stout bristles, which are smaller and larger pimctures less than 3 X as long as wide; remainder P. castaneus Lea of elytra bearing ver>' short and fine — Elytral punctation not dual 32 hairs, which are barely visible under 32. Antennal segments 4-7 slightly com- lower magnification; antennal segments pressed, segment 8 distinctly compressed; 3-5 only slightly broader at base than at total length 2.25 mm; Northern Territory apex £. microcera Clark P. scabricollis Lea — Head, pronotum, and elytra bearing — Antennal segments 4-7 monilifomi, seg- longer and finer bristles, which are about ment 8 only slightly compressed; total 8 X as long as wide; antennal segments length 3 mm; South Australia 3-5 much broader at base than at apex .... P. ovipennis Lea £. tristis Olliff .33. Pronotum without median fovea near the 22. Pronotum without lateral margins (Figs. base (Fig. 28); antennae 3-segmented, ._ 23 33, 34 ) ( genus Pohjplocotes W'estwood ) the last segment forming a solid, cylin- — Pronotum with lateral margins sharp drical club without any trace of sutures 33 (Fig. 28) (genus Ectrephes Pascoe) E. Pascoe ( Fig. 15 ) formicarum 23. Antennae 10-segmented, segments .3-10 — Pronotum with median fovea near the P. (Lea) compressed dolichognathus base; antennae 7-segmented, or 3-seg- less than 10 24 — Antennae with segments mented with the last segment flattened with the last 2 24. Antennae 6-segmented, (Fig. 20) 34 and segments enlarged compressed ( Fig. 34. Antennae 7-segmented, the last 5 seg- P. 22) sulcijrom (Lea) ments not fused togetlier _. E. chtatus Mjoberg — Antennae 9-segmented 25 — Antennae 3-se,gmented with the club flat- 25. Antennal segments 3-9 compressed 26 tened and apparently consisting of 5 fused — Antennal segments 8 and 9 or none of segments (Fig. 20) E. pascoei Westwood them compressed 27 26. Total length more than 2.50 mm; 9th Notes on the Genera antennal segment buried within the apex

P. anicnnalis ) the of the 8th ( Mjoberg Diphcotcs.—In most of the species, — Total length less than 2.50 mm; 9th an- clypeus is simple and only slightK' convex tennal segment not buried within the but in others it is more tumid and apex of 8th P. crenui.sioga.siri (Lea) (Fig. 1),

a short carina above 4 . In 27. Pronotum without basolateral trichomes may have ( Fig. ) of or a basal impression; mesal portion 8 it is somewhat constricted in kingi ( Fig. ) pronotal disc strigose (Fig. .33); chpeal the middle, and in carinaticeps (Fig. 7) a plate with a lateral perforation beneath it is P. perforatu.s- Lea narrow, median ridge formed. — Pronotum with basolateral trichomes; if The antennae may have from two to basal impression absent, then mesal por- eleven segments. The simplest or least tion of pronotal disc not strigose; clypeus of antenna occurs in mil- without perforation 28 modified type fa Myrmecophilous Ptinidae • Lawrence and Reichardt 9

iaris (Fig. 21); there are 11 segments, most fovea and trichomes are quite distinct. In of which are longer than wide, and the last several species {brevipennis, armicoUis, lati- segment is not modified at the apex. In corni.s,kingi, cuncifonni.s,iind carinaticeps), foveatus, longicomis, mctaUicus, and imjr- die pronotum is short and broad, \\ith sharp mecophilus, the antennae are also 11-seg- anterior teeth, a median fovea, and distinct mented, but the segments tend to be shorter trichomes that are comiected w ith longi- and the last is truncate and tudinal the disc 30 segment pubes- grooves on ( Figs. and cent at the apex. In longicomis the last 35). D. foveicoUis has a similar pronotum, segment is \er\' long, and in intiicatus most but it is not as short and broad, and the of tlie segments are transverse and several anterior teeth are replaced by tuberculate have apical pubescence. In several other processes. species, the penultimate segment is en- The elytra in almost all of the species are larged and longer than the last segment bulbous and strongly convex, but in D. (Fig. 17); these forms may have 11-seg- myrmccophilus they are somewhat flat- mented (armicoUis, fovcicollis, howittanus, tened. In some species there may be several and minutus), 10-segmented [sirigicoUis distinct pits at the bases of the elytra, but and brevipennis), or 9-segmented (similis, these are absent in foveicollis, armicoUis, apicaUs, pUosiis, and carinaticeps) anten- laticornis, kingi, ciineiformis, and carinati- nae. In kingi the antennae are 3-segmented, ceps. and the last segment is enlarged and Folijplocotes.—The clypeal structure in strongly flattened, \\'hile ciineiformis has this genus does not vary considerably, but 2-segmented antennae with the second seg- the shape of the clypeal plate and the ment wedge-shaped (Fig. 35). D. laticornis extent of the apical cleft differ among the presents a problem, since there has been species. In P. poforatiis there is a lateral some confusion concerning the number of perforation beneath the clypeal plate, segments in the antenna. There are ap- The antennae are less xariable than in parently 10 segments, and the last nine are Diplocotes, and in most of the species they strongly flattened (Fig. 19). Mjoberg are 9-segmented with the penultimate seg- (1916) states that there are 11 segments ment enlarged (Fig. 18). In P. longipes and that segment 2 is greatly reduced, as most of the antennal segments are elongate, in some species of Paussidae (see Darling- but in longicoUis, nitidus and several others ton, 1950). After examining carefully the they are more robust and the last two are antenna of this species, we cannot agree somewhat flattened. The antennae of sid- with on this the 22 are but there Mjoberg matter; apparent cifrons ( Fig. ) very similar, second segment is actually a condyle at- are only six segments. InP. dolichognatJius, tached to the real segment 2. It is more P. antennalis, and P. cremastogastri most probable that reduction of segments has of the antennal segments are flattened, as taken place at the apex of the antenna. in Diplocotes laticornis (Fig. 19); in the The simplest t\pe of pronotum in this first species the antennae are 10-segmented, genus also occurs in familiaris (Fig. 32); while antennalis and cremastogastri have a transverse, basal impression is present, 9-segmented antennae. anterolateral teeth and a median fovea are Most of the species have a simple type absent, and trichomes are small and indis- of pronotum with a weak basal impression tinct. Other species may have various com- and small trichomes usually not visible binations of the above characters; D. 5fn'g/- from above (Fig. 34). In P. perforatiis collis, for instance, has sharp anterolateral (Fig. 33) the basal impression is absent teetli on the pronotum but lacks a median and the pronotum is narrowed anteriorly; fovea and trichomes, while in D. longicomis basolateral trichomes are absent, but there (Fig. 31) the teeth are lacking, but the are tufts of yellow hairs along the anterior 10 Bulletin Museum of Compamfive Zoology, Vol. 138, No. 1

edge of the pronotiim that may represent think that Mjoberg had not misidentified lateral trichomes that haxe been displaced. the species bvit had merely transposed the Ectrcphcs.—The species in this genus names. If this hypothesis is acceptable, fall into two groups: E. formicanim with a then Ectrephes and Monectrephes are simple pronotal disc (Fig. 28) and the based on different species and are not terminal segment of the 3-segmented an- objective synonyms, although they are tenna forming a solid, cylindrical club here considered to be subjective synonyms.

15 , the other ( Fig. ) and two species ( pas- ORIGIN AND EVOLUTION OF coei and clavatiis) with a foveate pronotum MYRMECOPHILY IN THE PTINIDAE and the distal antennal segments strongly flattened and variously fused so that the Habitats and Food Habits total number of is three 20 segments ( Fig. ) Most ptinid beetles are scavengers, and or seven. It is that these interesting species they are known to feed on a wide variety and also D. and D. have kin^i cuneifonnis of animal and plant substances. The major the tibiae and enlarged angulate proxi- kinds of habitats from which ptinids have mal ly. been recorded are: (1) under bark and in In the there has been some con- past decomposing wood, (2) in caves, (3) in the fusion the names and concerning Ectrephes nests of birds and mammals, (4) in the Mone ctre divided plies. Mjoberg (1916) nests of other insects, and (5) in association into three the Ectreplies genera: original with various stored products of man. Be- with its E. genus Ectrephes type fonni- cause of the economic importance of these carinn for E. Pascoe, Mesectrephes kin^ii beetles, there have been several attempts and for E. Westwood, Monectrephes pas- to summarize the infonuation pertaining to coei Westwood. He also described another their biology; most of the following discus- E. \\'hich he referred to species, clavatiis, sion is based on papers of Hinton (1941), Then in discussions of the Monectrephes. Linsley (1944), and Howe (1959), all of various he genera continually transposed which contain extensive bibliographies. the names. He mentions that is Ectrephes In Europe, Ptiniis siihpilosiis Sturm, P. the least differentiated, the last having palliatiis Ferris, and P. licheniim Marsham of the antennae more or less segments are commonly collected under bark and in and the joined together penultimate seg- decaying wood. According to Crowson ment excavated at the apex and receiving 1962 P. occurs on ( ) , siibpilosus chiefly old, the last is a of E. (this description pascoei, mossy, partly dead oaks in Scotland, and not E. . He then states that in formicanim) the larvae have been found in decaying oak Monectrephes there are only three joints, wood. The larva of P. Uchenum has been the last one being very long and fonning to bore into reported dry fig wood ( Howe, a solid club refers to E. . (this formicanim) 1959), but no ptinid is known to utilize This type of transposition occurs in several P. wood as food. Reitter ( 1911 ) records other the to the places, including key suhpilosiis from dry moss, P. pilosus Miiller Because of later authors genera. this, (Lea, from old deciduous trees, and P. sexpunc- 1917; Pic, 1929) synonymized Monectrephes tatiis Panzer from beneath pine and maple with Ectrephes, assuming that both were 1944 lists four other bark, and Linsley ( ) based on the same E. species, formicarum. records from European literature. In North In first discussion of the Mjoberg's genera, America, two species of Ptinus were col- £. was referred to as however, formicarum lected by beating live oaks in southern the of E. type Ectrephes and pascoei was California (Fall, 1901), a specimen of P. definitely placed in Monectrephes. The bimaciilatiis Melsheimer was taken in Span- latter species is designated as type of ish moss in Louisiana, and specimens of Monectrephes in the catalogue below. We P. fur Linnaeus have been found in associa- Myrmecophilous Ptinidae • Lawrence and Reichardt 11

tion with Pinus in New England. Althougli products has been adequately covered by the Ixx'tles be on may feeding rotten wood, Howe and others. Hinton (1941) lists 21 it is also that possible they ntilize insect economically important species, and Howe remains or other organic debris. (1959) has made detailed studies of 14 of Several ptinids haxe been recorded from these. Some of the many items on which where ca\'es, they usnally feed on bat guano ptinids have been reported to feed are: or other mammal excrement. Brown 1959 ( ) wheat, flour, rye, cereals, bread, hay, corn, described Pscudciirostus kellcri and Niptus ginger, nutmeg, cacao, opium, fish meal, ahditus from taken in caves in specimens sugar, pepper, tallow, wool, tobacco, paste, Utah, and four other species have been dried mushrooms, dried fruits, rubber, collected in caves in Europe and northern leather, plant and insect collections, and Africa (Howe, 1959). An undescribed books. Although some of these records are was found on raccoon 'Niptus feeding scat suspect, the total range of food items is still in a Texas cave 1966 . ( Reddell, ) Although very great. mammal dung appears to be an important food source in all the more sheltered hab- Biology of the Myrmecophilous Species itats mentioned here, ptinid beetles have Very little is known about the habits of never been found in cattle droppings and inquilinous ptinids, and the present discus- are rarely collected in other kinds of dung sion is based partly on knowledge of the in open places. One notable exception is food habits of non-myrmecophilous species, given by Andrews (1967), who collected facts concerning the distributions and nest- two Central American Ptimis in coati ing habits of host ants, interpretation of Nostio narica scat ( ) on exposed surfaces in structural adaptations in the myrmecophi- a dry forest. tropical lous Ptinidae, and information on the The most important natural habitats of behavior of other myrmecophiles which the Ptinidae are the nests of mammals, have evolved analogous structural features. birds, bees, and ants. wasps, Linsley if Most, not all, of the myrmecophilous (1944) and Howe (1959) Hst 11 of species ])tinids would be considered or from symphiles ptinids mammal nests, 17 species from true guests, according to the ecological clas- bird nests, and 12 species from the nests of sification proposed by Wasmann (1898). hymenopterous insects. The actual of tvpe Most of the species have the characteristic food utilized varies with the kind of nest, "symphilie features" mentioned by Donis- but here again excrement is an probably thorpe (1927), Wheeler (1928), and Park important food item, with dead in- along 1964 : ( ) smooth and shiny, black or reddish sects, hair or feathers, and pollen, possiblv integument, reduced mouthparts, trichomes, living larvae. to and According Linsley and antennal modifications. Behavioral MacSwain (1941), Ptinus califamicus Pic, observations indicate that some species, at which lives in the nests of Osniia, does not least, are tolerated and even protected by attack the bee larva but cause its death may the ant colony. Clark (1923) noted that indirectly through competition for pollen. specimens of Enosiba tristis and E. conifer P. scxpunctatus, on the other feeds hand, in an observation nest an active on containing healthy larvae of P. fur (Howe, 1959). colony of Iridomyrmcx conifer "seemed to A number of eeonomicallv important spe- be on friendly terms with the ants, and, cies, hololcucus (Falder- including Niptus during the three months' confinement were mann), feed on rat and mouse commonly never interfered with, but were allowed to feces, and the infestation of warehouses move freely through the various chambers, and mills these beetles be associated by may including the nurseries which always con- with the of rodents. presence tained a large number of larvae and pupae The of stored biology ptinids attacking of the ants." He further observed that "one, 12 Bulletin Museum of Comparative Zoology, Vol. 138, No. 1

and sometimes two ants were seen to attach lant that they become temporarily dis- themselves to the antennae, and appeared oriented and less sure of their footing." to be getting great satisfaction by nibbHng The predaceous reduviid bug Ptiloccrus and licking the apical joints, stroking the ochraccus Montandon produces a secretion beetle nlcan^^'hile with their antennae. which not onlv attracts ants but narcotizes None of the ants were seen to attach them- them as well, enabling the bug to penetrate selves to the fascicles on the prothorax." and feed on the paralvzed prey (Wheeler, The follo\\ing comments were made abont 1928; Donisthorpe, 1927). to \\' heeler 1928 the modi- Diplocotidiis fonnicola by W'asmann and According ( )

: is not fied antennae of some are Brauns ( 1925 ) "The species only mynnecophiles tolerated by the ants but seems to be adapted to stroking the ant host, or function treated as a true guest like most of the as handles for the ants to carry the beetles Paussidae. When opening the ants' nest about in the nests. According to Mou I have often observed that the Flagiolcpis (1938) and Janssens (1949), however, the [AnopJolcpis spp.] grasped the beetles and antennae of paussids are definitely glan- carried them away into security. . . . The dular and probably serve the same function behavior of the beetle is unconcerned. It as the trichomes on the prothorax. The runs carelessly in the thickest throng of the observations on Enasiba mentioned above otherwise vicious and carnivorous Plagio- indicate that this is also true in some in Ptinidae. 1950 stresses the lepis and can be obser\'ed sitting copula Darlington ( ) in the crowd of ants." Although there is no apparent loss of the tactile function in other direct evidence pertaining to the paussid antennae, as suggested by the re- biology of myrmecophilous ptinids, studies duction or loss of the pedicel and the of other inquilinous beetles may shed some associated Johnston's organ. light on certain features, such as the func- Since most other ptinids are scavengers, tions of trichomes and antennal modifica- it is unlikely that the inquilinous forms feed tions, feeding and means of dispersal. on ant larvae as do staphylinids and paus- The tufts of hairs or trichomes on the sids. The most probable sources of food prothorax, elytra, or abdomen of mynne- are excrement, dead ants, and the remains cophiles secrete some substance which is of prey. In some of the Ptinidae, however, imbibed by ants. The licking of trichomes the mouthparts are reduced, indicating that by ants has been observed for a number they may be fed regurgitated food by the of beetle groups, including the "lomechu- ants. Free-li\"ing ptinids, such as Ptinus fur sine" Staphylinidae (\\^asmann, 1920; (Figs. 11 and 14) have normal chewing Wheeler, 1928), the clavigerine Pselaphi- mandibles and well-developed maxillae dae (Donisthoipe, 1927; Park, 1964), and with two distinct lobes. In Diplocotes re- the Paussidae . In both structures are somewhat ( Reichensperger, 1948 ) farnilioris a detailed anatomical and histological duced, and in Ect replies formicarum the study, Mou (1938) demonstrated that the mandible is narrow and scoop-like (Fig. 13 is trichomes of the Paussidae are definitelv ) and the maxilla highly reduced ( Fig. glandular, producing a secretion which 12). Reduction of mouthparts and feeding evaporates slowly, leaving a greasy residue by ants is a common feature in other sym- which is difficult to dissolve in alcohol but philes. According to Donisthorpe (1927), soluble in acetate. Artemeles elicits food from easily ethyl Park ( 1964 ) (Staphylinidae) made the following comment on the behav- worker ants by tapping with its antennae, ior of ants which had been feeding on but can survive on ant larvae and dead

". . . an clavigerine trichomes: on several insects as well. Park ( 1964 ) described occasions, I haxe obser\'ed the workers to individual of Adranes lecontei Brendel be so overwhelmed bv this trichomc stimu- being fed regurgitated food by one worker Myrmecophilous Ptinidae • Lawrence and Reichardt 13

while two more ants were its tri- licking with the loss of the wood-boring habit ehomes. being the most important factor involved in Since of many species Ptinidae, inclnding the divergence of the group. He considers almost all of the inquilines, are wingless the ptinid genus XyJodes to be intermediate in both sexes, the means of in dispersal between Hedobia and a primitive Ptiniis, these insects is difficnlt to A cine explain. such as P. paUiatiis. Assuming that there be fonnd in observations made on may was a shift from xylophagy to general certain which are cla\'igerine Pselaphidae, scavenging, the ancestral forms may have also wingless. Adrancs lecontei Brendel is first fed on rotten wood mixed with excre- to known ride the gaster of worker ants ment and insect remains. Some extant and has been observed to be carried in this species have retained this habit, as noted manner for as as minntes long ninety ( Park, above. Two independent trends must then

1964 ) . testaceus Claviger Preyssler may have taken place: (1) the movement away attach itself to females and males winged from the arboreal or woody habitat, and of its host, and 1927 noted the Donisthorpe ( ) (2) increased utilization of animal by- that two of specimens Claviger longicornis products, including stored food, feces, and Miiller sat for hours on the frequently carrion. It is not surprising that many of an gaster Aconthovu/ops ftdiginosiis species came to inhabit the nests of birds, queen. It is then, that a quite possible, mammals, and insects, since the greatest number of m>'rmecophilous insects mav be accumulations of suitable food items would to new nests transported by queen ants. be found in these habitats. Also, the Ptini- Tlie small number of available host re- dae have become particularly successful in cords makes it impossible at present to arid regions where nests of various kinds determine the extent to which particular provide one of the few habitats in which are restricted to one ptinids species or one sufficient moisture is available for the com- of ants. In the New W'orld, Gnostus group pletion of the life cycle. Howe (1959) is known from nests only Crematogaster stresses the fact that ptinids require "drink- and Fahrasia is associated with Cam- only ing" water, which would be present in nests but in Australia the ponotus, situation is in the form of urine, moist excrement, more complex. Many of the Australian honey, or other fluids. As mentioned above, forms are associated with species of Irido- the species which invaded warehouses and but several have been myrmcx, found in mills were probably first associated with Crematogaster and nests. Rlujtidoponcra rats and mice. From these general nest Seven different have ptinids been taken in scavengers evolved the depredators, such the nests of and Jridomyrmcx conifer Forel, as Ptinus californicus and P. sexpunctatiis, some have been associated with as many and the myrmecophiles. as three ant species. Iridomyrmex conifer The invasion of the nests of social insects, occurs in the southwestern dry part of such as ants, presents a problem to the Australia, and it builds large, conical potential inquiline, who must overcome the mounds of and leaves. Most of the twigs strong defenses of the colony. Unlike the Australian ant hosts are ground nesting, myrmecophilous Staphylinidae, the slow- while and Camponofus Crematogaster in moving ptinids certainlv have never been the New World build off usually nests the active predators, and the first species to ground in rotten wood, and vines. twigs, penetrate ant colonies were probably indif- ferently tolerated forms or synoeketes, in Origin of the Myrmecophily terminology of W'asmann (1898). According to Crowson (1955), the Ptini- Ptinus sid)piIosus is a living species which dae have probably evolved from an anobiid- may be a synoekete, since it lacks trichomes like ancestor resembling the genus Hedobia, or other symphilie features but appears to 14 Bulletin Museum of Comparative Zoology, Vol. 138, No. 1

have invaded successfully the nests of Acan- outside the nest, nor do they supply a thomyops in Europe. These secretive forms significant amount of nutriment to the may be able to survive in the nest by colony. Symphiles are known to occur in occupying abandoned chambers or by feign- small numbers throughout the active por-

disturbed letisimulation . tions of the nest and to ing death when ( ) they appear be The latter type of behavior, which has been accepted as part of the colony. It may be 1964 for the that discussed by Park ( ) Pselaph- argued any "foreign" insect producing idae, may have been very important in the secretions which stimulate the ants to feed initial adjustment to the ant society. It is would be devoured rather than protected, likely that beetles exhibiting letisimulation but several factors may prevent this. First, would be ignored by worker ants or pos- it has been assumed that the initial invaders sibly carried off with the debris. It is also have penetrated the nest in the manner possible that certain structural features of discussed above and have thus taken on the Ptinidae have preadapted them for the colony odor. Second, most of these myrmecophilous existence. All ptinids are symphiles resemble ants and some of them, relatively small and compact beetles with a at least, exhibit antlike behaxior. Third, the heavy cuticle. The head is declined and aggressive behavior of a foraging or defend- the prosternum shortened, so that the ing worker is unlike that of an ant within mouthparts and antennae may be hidden the nest, where a common activitv is the beneath the body. If a beetle were to be exchange of regurgitated food or secretions attacked, the com]:)act fonn and thick cuti- (trophallaxis). If a myrmecophile elicits cle might provide sufficient protection, but no defensive response from the ants, and this would not be necessary for survival if if trophallactic exchange is a common pat- enough of the population avoided encoun- tern among individuals within the nest, it ters with the ants. Once this initial adjust- is conceivable that the development of ment was made, the inquiline would be attractive secretions by an inquiline might provided \\'ith a comjoatible enxironment lead directly to a situation in which the (especiallv in arid regions), an abundant insect is not only protected but also fed food supplv, and the protection of the by the ants. colony. Selection would favor the sub- In the preceding discussion it is assumed sequent evolution of features either that that trichomes developed in beetles which would improve the defenses of the inquiline already had invaded successfullv the nests or allow it to become integrated into the of ants. It is also possible that these glands ant society as a protected symphile. were a necessary prerequisite to the in- There is little doubt that in several vasion of nests. Cazier and Mortenson such as the 1965 that in the groups, Ptinidae, Paussidae, ( ) suggest scarabaeid and Pselaphidae, the evolution of true beetle Crcmastochcilu.s trichomes mav svmphily has been associated with the serve to attract ants to the beetles which development of glandular structures, such are then carried into the nest as items of as trichomes, but the functions of these food. A third alternative is that trichomes glands and the steps leading to their evolved independently and serve no neces- development are far from being clearly sary function in myrmecophily. We think understood. It is known that ants may that the occiuTence of these structures in imbibe the secretions and that insects pos- so man\' unrelated myrmecophiles makes sessing trichomes are often protected and this last suggestion improbable. fed by ants, but the relationships between As mentioned above, myrmecophily in symphiles and their hosts have never been the Ptinidae probably originated several studied in detail. Unlike aphids and coc- times. Mijrmecoptinus butteli, a winged cids, these insects are not tended in groups species associated \\'ith Crcmatogaster in Myrmecophilois Ptimdae • Laurence and Reichardt 15

the forests of Southeast Asia, is the least the Australian inquilines are all wingless modified of the inquihnes and may have (which may be related to water conserva- evolved recently from non-m\rmecophilous tion in dry areas), while the New World forms, ^^'e ha\-e suggested (Lawrence and species are winged. The lack of wings will Reichardt. 1966a that the two New \\'orld ) obvious!)' reduce tlie rate of gene exchange Gnostus genera, and Fahrasia. e\olved between populations. If, as suggested direct!)- and independent!)' from ear!)- above, the wingless myrmecophile must Ptinus-MVQ ancestors. Tl-ie species in botli attach itself to a new queen, then the groups are high!)' modified sympliiles, but probabilit)' of outbreeding will be very low. they ha\-e retained their wings and are Finally, the Australian ant fauna contains associated witli arl)orea! or wood-nesting a large numl^er of ground-nesting species, ants in the American tropics. The South such as Iridomyrmex conifer, which build African Diplocotidus and the Australian mounds and have relatively large and per- ectrephines are all wingless, and most of manent colonies. The South American ants, them occur with ground-nesting ants in on the whole, have smaller and less per- open, arid or semi-arid regions. Tliese two manent colonies. These Iridomyrmex nests groups may have evolved from ancestors provide a suital^le en\ironment for the which had already left the ancestral forest Ijuild-up of inquiline populations, which, habitat and associated were with ground because of difficulties of dispersal, ma)' debris or mammal burrows in dr\' areas. be isolated from other such populations for Diplocotidus fomiicolo is a true symphile, long periods of time. Such conditions judging from the observations of ^^'asmann would fa\or the evolution of a dixerse and Brauns (1925), but it has not evolved myiTnecophilous fauna. some of the more obvious antenna! and pronota! modifications found in the Aus- Evolutionary Trends tralian forms. The are the most ectrephines In examining the various modifications di\erse and numerous of the m)-rmecopl-ii- of myrmecophilous ptinids, we may dis- lous ptinids, but have almost certain!\' the)' tinguish between those unique to inquilines e\olved or from one possibly two related and those found in other Ptinidae as well. forms; all are but the probably s)'mphiles, The former group would include: elonga- of structural modification varies degree tion of clypeus and labrum, reduction of considerably. mouthparts, enlargement, flattening, and six of Only species mynnecophilous fusion of antenna! segments, formation of occur in the ptinids Neotropical region, glandular structures on antennae, prono- while 35 species are known from Australia. tum, elytra, and hind legs, changes in shape the World fauna consists Furthermore, New of tibiae, and loss or extreme reduction of of three related in each of closely species hairs and scales over most of the body the two while the Australian fauna genera, surface. The following would fall into the is so diverse that the were formerlv species latter group: reduction of eyes, declination in 14 placed genera. Certainly the habitat of head, fonnation of deep antenna! fossae, of South America exceeds that of diversity loss of antenna! segments, formation of Australia, and the ant faunas of the two grooves and pits on pronotum, loss of hind- areas are The comparable. following fac- wing, reduction or loss of scutellum and tors are to have been !il

in to that idae, including Meziiim, Gihhium, Niptus, Ptinidae is similar many ways in the Paussidae, and a number and Trigonogeniws, while some of the myr- occurring of can be seen in the two mecophiles ha\e fully developed wings convergences the has antennae (Gnostiis and Fahrasia). Since most of groups. Protopaiissws simple areas or in like those of is, while wingless ptinids occur in drier Diplocotes familiar the loss and their rela- stored products, it is possible that Homopteru-s, Cerapterus, tives have the flattened of antennae of wings is associated with \\'ater conserxa- type in laticornis. Lebioderus and tion. found D. have flattened antennae Changes in the shape of the clypeus and FcntapIatartJinis with a reduced number of and labrum are unique to the myrmecophilous segments, in Ceratoderus and fur- species, and mav sene to protect the mouth- PJatyrhopalopsis ther fusion and reduction can be seen; this parts. In some of the highly modified finds a in forms, such as Ectrcphes, the concave inner series parallel Diplocotes kingi, surface of the labrum forms with the hol- Ectrcphes clavatiis, and £. pascoci among the The club-like antenna found lowed and elongate mandibles a tube which ptinids. in E. and the Gnostus is may be used for feeding on liquids supplied formicarum genus Paussus. bv the ants. In these same forms, the similar to that of some chewing surface of the mandible is absent and the maxillae are reduced. CATALOGUE OF THE Modifications of the legs of these inqui- MYRMECOPHILOUS PTINIDAE lines probably represent three different kinds of adaptations. In Gnostus, the tibiae Myrmecopfinus Wasmann are broad and and serve compressed may Mijrmecoptinus Wasmann, 1916: 206-207. Type structures buttcli as protective shields; very similar species, by monot\py, Myrmecoptinus occur in the paussid genera Homoptcrm Wasmann. The tibiae of and Ceraptcms. Ectrcphes Included species: 1. formicarum and certain other ectrephines are somewhat bent and angulate; these Myrmecopfinus buffeli Wasmann ants. hind might be used to cling to The Miirmcco]^finus Jnitteli Wasmann, 1916: 207, pi. 4, tibiae and femora of Fahrasia are peculiarly fig. 20. enlarged and bear tufts of yellow hairs: Distribution: SUMATRA (Soengei, Bam- these are structures glandular . probabl\- ban ) serving the same function as trichomes. Host: Crcmatogaster buttcli Forel. The types and possible functions of pro- notal and antennal trichomes have been Diplocofidus Peringuey of discussed abo\e. Several different types Diplocotidus Peringuey, 1899: 245. Type species, glandular systems have evohed in this l^y monotypy, Diplocotidus jormicola Peringuey. but the is not as group, di\ersity nearly Included species: 1.

great as in the Paussidae ( see Darlington. 1950). Perhaps the first structures were Diplocotidus formicola Peringuey small isolated scattered on the bod\ glands Diplocotidus jormicola Peringuey, 1899: 245-246, 1925: surface. A consolidation of these small pi. 7, figs. 13, 13a; Wasmann and Braims, 112-113. glands gave rise to more complicated organs, such as trichomes, and to various Distribution: UNION OF SOUTH AF- accessory grooves and pits for the collection RICA (Prince Albert; W'illowmore; Lady- of fluid. The fusion of antennal segments smith; Kimberley). would serve to consolidate glandular mate- Hosts: Acantholcpis capensis Mayr; rial and form a storage vesicle as well. Anoplolcpis custodiens (F. Smith); Anop- The evolution of antennal types in the lolcpis steingroeveri (Forel). Myrmecophilous Ptinidae • Laurence and Reichardt 17

Gnostus Westwood Distribution: BRAZIL: RIO GRANDE Gno.ftus Westwood, 1855: 90-92; Lawrence and DO NORTE (iNatal). Reichardt, 1966a: 32-34. Type species, by Host: Unkno\\n. nionot>p\-, Gnostus formicicola Westwood. Fabrasia borgmeieri Lawrence Included 3. species: and Reichardt Gnosfus formicicola Westwood Fabrasia borgmeieri Lawrence and Reichardt, 1966a: 41-42, figs. 6-7. Gnostus formicicola Westwood, 1855: 92, pi. 8, Distribution: BRAZIL: lies. 1-21; Lawrence and Reichardt, 1966a: 34, GOIAS (Campi- fis:. 4. nas); MINAS GERAIS (P090S de Caldas). Host: Distribution: BOLIVIA: LAPAZ(Map- Camponotus (Myrmothrix) reng- iri); BRAZIL: PARA (Santarem; Villa geri Emery; Camponotus (Myrmobrachys) sencx Nova); AMAZONAS (Sao Paulo de Oli- (F. Smith). ven9a); \L\TO GROSSO (Utiariti). Fabrasia wheeleri Lawrence Hosts: Crcrnatogaster victima F. Smith; and Reichardt hrasilicnsis Crcmatogaster sp. (near Mayr). Fabrasia uheeleri Lawrence and Reichardt, 1966a: 42-44, figs. 8-12. Gnosfus floridanus Blatchley Distribution: COLOMBIA: MAGDA- Gnostus floridanus 1930: 111-112, fig. Blatchley, LENA (Sevilla); MEXICO: OAXACA 1; Lawrence and Reichardt, 1966a: 37, figs. 1, mi. \\' . (6.3 ) 2. 5, 13, 14. Tehiiantepec Ho.^: Camponotus (Myrmothrix) bug- Distribution: Point, BAHAMAS (Clifton nioni Forel. New Providence); FLORIDA (Dunedin; Plantation Key). Diplocotes Westwood Hosts: Crcrnatogaster ashmcadi Mayr; Diploeotes Westwood, 1869: 317. Type species, by nionot\p\-, hotcittanus Westwood. Crcrnatogaster sanguinca lucatjana \Mieeler. Diplocotes Diphohia Oilif'f, 1886: 837-838. Type species, by monot\p\-, Diphohia familiaris Olliff. NEW Gnosfus meinerti Wasmann SYNONYMY. Gnostus nicincrti Wasmann, 1894: 121, 216; Law- Paussoptinus Lea, 1905: 381. Type species, by rence and Reichardt, 1966a: 35, fig. 3. monotvp\-, Paussoptinus laticornis Lea. NEW SYNONYMY. Distribution: PANAMA Barro Colorado ( Decemplocotes Mjoberg, 1916: 2; Lea, 1917: 146 Is.; Erwin Is.; Gatun Lake); VENEZU- (proposed synon\iny with Diplocotes) . Type ELA: CARABOBO (Trincheras, near species, by present designation, Diplocotes strig- icollis Lea. Valencia ) . Mesectreplies Mjoberg, 1916: 3. Type species, by Host: litnata dextella Crcrnatogaster nionot\p\-, Ectrephes kingi Westwood. NEW Santschi. SYNONYMY. Bitrephes Oke, 1926: 291; Oke, 1928: 27. Type Fabrasia Martinez and Viana species, bv monotvpv, Bitrephes cuneiformis Oke. NEW SYNONYMY. Fahia Martinez and Viana, 1964: 8, not Dana, Fahoectrephes Pic, 1929: 35. Type species, by 1851. Type species, by original designation, nionotvp\', Ectrephes kingi Westwood. NEW Fahia alvarengai Martinez and Viana. SYNONYMY. Fabrasia Martinez and Viana, 1965: 18; Lawrence Included 19. and Reichardt, 1966a: 37-40. species:

Included species: 3. Diplocotes apicalis (Oke), NEW COMBINATION Fabrasia alvarengai (Martinez and Viana) Poltjplocotes apicalis Oke, 1928: 26. Fabia Martinez and 1964; alvarengai Viana, 9, Distribution: AUSTRALIA: VICTORIA figs. 1-3. Fabrasia alvarengai (Martinez and \'iana), 1965: (Hattah). 18; Lawrence and Reichardt, 1966a: 41. Host: Iridomyrmex sp. 18 Bulletin Museum of Comparative Zoology, Vol. 138, No. 1

Diplocofes armicoll'is Lea Diplocofes foveafus (Lea), Diplocotes armicoUis Lea, 1910: 219-220, figs. NEW COMBINATION 25-26. Diphobia fovcata Lea, 1917: 154-155. Distribution: AUSTRALIA: SOUTH Distribution: AUSTRALIA: QUEENS- AUSTRALIA (Adelaide). LAND (Townsville). Host: Unknown. Collected under bark Host: Unknown. with ants. Diplocofes foveicollis Olliff Diplocotes brevipennis (Pic) Diplocotes foveicollis Olliff, 1886: 839-840; Lea, 1910: 219; 1917: 149; 1919: Paussoptinu.i brevipennis Pic, 1909: 215; Lea, Lea, Lea, 257; 1910: 223. Clark, 1921: 103. - Decemplocotes brevipennis, Mjoberg, 1916: 6, Distribution: AUSTRALIA: NEW fig. 6. - SOUTH WALES; SOUTH AUSTRALIA Diplocotes brevipennis, Lea, 1917: 146. Diplocotes decemarficulatus Lea, 1910: 220-221, (Lucindale); WESTERN AUSTRALIA 1917: 146 fig. 57: Lea, (syn.). (Serpentine River; Perth; Swan River). Distribution: AUSTRALIA: \\ESTERN Host: Iridomijrmex conifer Forel. AUSTRALIA (Geraldton; Moora). howiffanus Westwood Host: Unknown. Diplocofes Diplocotes houittantis \\'est\voocl, 1869: 317; Westwood, 1874: 103, pi. 3, fig. 6; Lea, 1910: Diplocotes carinaficeps (Lea), NEW COMBINATION 219; Oke, 1928: 26. Distribution: AUSTRALIA: SOUTH Polyplocotes carinaticeps Lea, 1919: 254-255, pi. AUSTRALIA VIC- 25, fig. 39; Zeck, 1920: 246, pi. 19, fig. 3: Oke, (Gawler, Adelaide);

1928: 27. . TORIA ( Grampians ) Distribution: AUSTRALIA: VICTORIA Ho.st: Unknown. Collected under bark Swan with ants. Hattah ; WESTERN AUSTRALIA ( ) (

River ) . Diplocofes infrlcafus (Lea), Hosts: Crematogasfer laeviceps F. Smith; NEW COMBINATION Iridomijrmex conifer Forel. Diphobia intricata Lea, 1917: 155-156. Diplocotes cuneiformis (Oke), Distribution: AUSTRALIA: QUEENS- NEW COMBINATION LAND (Stewart River). Bitrephes cuneiformis Oke, 1926: 291; Oke, 1928: Hosts: Crematogaster sp.; Iridomijrmex 27-28. sp. Distribution: AUSTRALIA: VICTORIA kingi (Westwood), Lake Hattah . Diplocofes ( ) NEW COMBINATION Host: Iridomijrmex rufoniger (Lowne). Ectrcphes kingi Westwood, 1869: 316; Westwood, 1874: 104, 3, 2; Lea, 1910: 222; Lea, Diplocofes familiaris (Olliff), pi. fig. 1917: 147. NEW COMBINATION - Mesectrephes kingi, Mjoberg, 1916: 3. - Diphobia familiaris Olliff, 1886: 838; Lea, 1905: Falsoectrephes kingi, Pic, 1929: 35. 381; Lea, 1910: 223. Distribution: AUSTRALIA: WESTERN Diplocotes niger Poll, 1886: 238; Blackburn, 1892: 300 (syn.). AUSTRALIA (Swan River; Mullewa). Distribution: AUSTRALIA: SOUTH Ho.st: Iridomijrmex sp. AUSTRALIA (Adelaide); VICTORIA Diplocofes laficornis (Lea), (Kiata). NEW COMBINATION Hosis: Iridomijrmex nitidus Mayr; Irido- Paussoptimis laficornis Lea, 1905: 382-383, pi. 28, Andre; mijrmex nitidiceps Rlujtidoponera fig. 7; Lea, 1917: 147; Tillyard, 1926: 215, pi. metallica (F. Smith). 16, fig. 14. Myrmecophilous Ptinidae • Lawrence and Reichardt 19

Distribution: AUSTRALIA: SOUTH Diplocotes similis (Oke), AUSTRALIA (Mindarie; Murat Ray; Mur- NEW COMBINATION Port Lincoln); VICTORIA ray Rridge; Polyplocotes si77}ilis Oke, 1928: 27. (Rirchip; Rendigo; Sea Lake). Disiribution: AUSTRALIA: VICTORIA Hosts: CrcmatOfiasier kieviceps F. Smith; (Rendigo; Inglewood). Iridomijrmcx nitidus Mayr; Iridomyrmex Host: Crematogaster laeviceps F. Smith. spp.

Diplocotes strigicollis Lea (Lea), Diplocotes longicornis Diplocotes .stri<:icollis Lea, 1910: 221; Lea, 1917: NEW COMBINATION 150. - Dcceinplocotes strigicollis, Mjoberg, 1916: 2. Dipho})ia loiif^icorni.s Lea, 1919: 256, pi. 25, fig. 40; ZLck, 1920: 245, pi. 19, fig. 4; Clark, 1921: Distribution: AUSTRALIA: SOUTH 103. AUSTRALIA (Adelaide; Port Lincoln). Distribution: AUSTRALIA: WESTERN Host: Iridomyrmex sp. AUSTRALIA (Victoria Park; Swan River). Westwood Hosts: Iridoynyrrucx conifer Forel; Irido- Polyplocotes Polyplocotes 1869: 316. myrmex sp. Westwood, Type species, by present designation, Polyplocotes longicollis Westwood. Diplocotes mefallicus (Lea), Hexaplocotes Lea, 1906: 224. Type species, b\- NEW COMBINATION nionoKpv, Hexaplocotes sulcifrons Lea. NEW SYNONYMY. metallica Lea, 1917: 155. DipJiolna Leaptinus Mjoberg, 1916: 3; Lea, 1917: 146 Distribution: AUSTRALIA: QUEENS- (proposed synonymy with Paussoptinus). Type LAND species, by monotvpv, Paussoptinus dolicho- (Stewart River). gnathus Lea. NEW SYNONYMY. Host: Iridomyrmex sp. Paussoceros Mjoberg, 1916: 13. T\pe species, by monotvp\-, Paussoceros anteunalis Mjoberg. NEW SYNONYMY. Diplocotes minufus Oke Included species: 11. Diplocotes minuta Oke, 1928: 26.

Distribution: AUSTRALIA: VICTORIA Polyplocotes antennalis (Mjoberg),

( Rendigo ) . NEW COMBINATION Host: Rhytidoponera sp. Paussoceros atitenualis Mjoberg, 1916: 13-14, fig. 4.

Diplocotes myrmecophilus (Lea), Disiribution: AUSTRALIA: WESTERN NEW COMBINATION AUSTRALIA (Derby, Kimberley District). Dii)hohia myrmecophila Lea, 1917: 153-154. Host: Unknown. Collected mider bark. Distribution: AUSTRALIA: NORTH- Polyplocotes castaneus Lea ERN TERRITORY (Darwin). Polyplocotes castaneus Lea, 1912: 75. Hosts: OecophyJIa smara^dina (Fabri- Distribution: AUSTRALIA: WESTERN cius), Rhytidoponera hilli Crawley. AUSTRALIA (Northwestern Australia). Host: Unknown. Diplocotes pilosus (Mjoberg), NEW COMBINATION Polyplocotes cremastogastri (Lea),

Polyplocotes pilosus Mjoberg, 1916: 11-12, fig. 3. NEW COMBINATION Disiribution: AUSTRALIA: WESTERN Paussoceros cremastogastri Lea, 1917: 153.

AUSTRALIA ( Noonkanbah, Kimberley Distribution: AUSTRALIA: \\'ESTERN District). AUSTRALIA (Derby). Host: l^nknown. Host: Crematogaster sp. Vol. 138, No. 1 20 Bulletin Museum of Comparative Zoology,

scabricoilis Lea Polyplocotes dolichognathus (Lea), Polyplocofes NEW COMBINATION Polyplocotes scabricollis Lea, 1917: 151-152. NORTH- 1912: Lea, Distribution: AUSTRALIA: Paussoptinus dolichognathus Lea, 76; 1917: 146. ERN TERRITORY (Stapleton). - 1916: 3. Leaptimis dolichognathus, Mjoberg, Ho.st: Unknown. Collected in ant nest. Distribution: AUSTRALIA: SOUTH sulcifrons (Lea), Central Aus- Po/yp/ocofes AUSTRALIA ( Killalpanima, NEW COMBINATION

tralia ) . Hexaplocotes .sulcifrons Lea, 1906: 224-225, pi. Host: Unknown. 1921: 18, fig. 8; Lea, 1917: 150; Clark, 104; Tillvard, 1926: 215, pi. 16, fig. 13. Westwood WESTERN Polyplocofes longicoHis Distribution: AUSTRALIA: 1869: 316; AUSTRALIA Newcastle; Perth). Polyplocotes longicotlis Westwood, (Beverley; 1910: Forel. Westwood, 1874: 103, pi. 3, fig. 4; Lea, Host: Iridomyrmex conifer 219; Lea, 1917: 147. 1903: 183. PPolijplocotes nigropiceus Pic, Ecfrephes Pascoe 1869: 315. Distribution: AUSTRALIA: WESTERN Ectrephe.s Pascoe, 1866: xvi; Westwood, monotypy, Ectrephes formi- Swan River; Sorrento Type species, by AUSTRALIA (Cue; carum Pascoe. Beach, Perth District). Anapestus King, 1866: 317; Westwood, 1869: 315 Host: Iridomyrmex sp. (proposed synonymy with Ectrephes). Type kreusleri species, by monotypy, Anapestus King formicarum Pascoe). Lea {= Ectrephes Polyplocofes longipes Monectrephes Mjoberg, 1916: 3; Lea, 1917: 145 with Ectrephes). Type Polyplocotes longipes Lea, 1917: 152-153. (proposed synonymy WESTERN species, by present designation, Ectrephes pas- Distribution: AUSTRALIA: coei Westwood. sub- AUSTRALIA (Derby). Pseudectrephes Pic, 1929: 35 (proposed as a of 1935: 4 (raised Host: Cremotogaster sp. genus Ectrephes); Schenkling, to rank of genus). Type species, by monotypy, Ectrephes clavatus Mjoberg. nitidus Westwood Po/yp/ocofes Included species: 3. West- Polyplocotes nitidus Westwood, 1869: 316; 1910: 219. wood, 1874: 103, pi. 3, fig. 5; Lea, Ecfrephes clavafus Mjoberg 1916: 5. Distribution: AUSTRALIA: WESTERN Ectrephes clavatus Mjoberg, 14, fig. Ectrephes (Pseudectrephes) clavatus, -Pic, 1929: AUSTRALIA (Swan River). 35. Hosi: Unknown. Collected with ants. Distribution: AUSTRALIA: WESTERN AUSTRALIA (Swan River). Lea Polyplocotes ovipennis Host: Iridomyrmex nitidus Mayr. Polyplocotes ovipennis Lea, 1914: 256-257. Ecfrephes formicarum Pascoe Distribution: AUSTRALIA: SOUTH formicarum Pascoe, 1866: xvi-xvii; Central Ectrephes AUSTRALIA (Lake Callabonna, Westwood, 1869: 315; Westwood, 1874: 104, 1919: . 1910: 256, Australia ) pi. 3, fig. 1; Lea, 222; Lea, 1920: Ho.st: Unknown. pi 25 figs. 41-42; Zeck, 246, pi. 19, fig. 1; Clark, 1921: 103; Tillyard, 1926: 215, pi. 16, fig. 15. Po/yp/ocofes perforafus Lea Anapestus kreusleri King, 1866: 317, pi. 16; West- wood, 1869: 315 (syn.). Polyplocotes pcrforatus Lea, 1917: 150-151, pi. 12, figs. 7-8. Distribution: AUSTRALIA: SOUTH Distribution: AUSTRALIA: WESTERN AUSTRALIA; WESTERN AUSTRALIA Swan AUSTRALIA (Beverley). (Jandakot; Perth; River). Host: Unknown. Collected in nests of Hosts: Iridomyrmex conifer Forel; Rhy- ants. tidoponera metallica (F. Smith). Myrmecophilous Ptinidae • Lawrence and Reichaidt 21

Ecfrephes pascoei Westwood Distribution: AUSTRALIA: WESTERN Ectrcphe.s pascoei Westwood, 1869: 315-316; AUSTRALIA ( Bunbury; Busselton). Westwood, 1874: 104, pi. 3, fig. 3. - Host: Iridonujrmex conifer Forel. Mouc'ctrcphe.s pascoei, Mjoberg, 1916: 3. Distribution: AUSTRALIA: WESTERN Enasiba frisfis OH iff AUSTRALIA (Swan River). Host: Unknown. Enasiba tri.sti.s Olliff, 1886: 839; Lea, 1919: 254, pi. 25, figs. 37-38; Zeck, 1920: 245, pi. 19, fig! Enasiba Oil iff 2; Clark, 1921: 103; Clark, 1923: 46; Tillyard, 1926: Enasiha Ollitf, 1886: 838-839. Type species, by 215, pi. 16, fig. 12.

m(mot>p\ , Enasiba tri.sti.s Olliff. Distribution: AUSTRALIA: WESTERN Included species: 2. AUSTRALIA (King George's Sound; Perth; Enasiba microcera Clark Swan River). Euasihd luicroccra Clark, 1923: 44-46. Host: Iridomynnex conifer Forel.

LIST OF ANT HOST SPECIES AND ASSOCIATED INQUIUNES

Fonnicidae Ptinidae Ponerinae Ectatommini Rhytidoponera hilli Crawley D. myrmecophihis (Lea) R. metallica (F. Smith) D. familiaris (Olhff) E. formicantm Pascoe Rhytidoponera sp. D. minutus Oke

Myrmicinae Crematogastrini Crematogaster ashmeadi iMayr G. floridanus Bkitchley C. butteli Forel M. butteli Wasmann C. F. Smith laeuiceps D. carinaticeps (Lea) D. Jaticornis (Lea) D. similis (Oke) C. limata dextella Santschi G. meinerti Wasmann C. sanguined Iucaya7ia Wheeler G. floridanus Blatchley C. victimo F. Smith G. formicicola Westwood Crematogaster sp. G. formicicola West\vood D. intricatus (Lea) P. cremastogastri (Lea) P. longipes Lea Dolichoderinae Tapinomini Forel Iridomyrmex conifer D. carinaticeps (Lea) D. foveicoUis Olliff D. longicornis (Lea) P. sulcifrons (Lea) E. formicarum Pascoe E. microcera Clark E. tri^tis Olliff 7. Andre nitidiceps D. familiaris (Olliff) Vol. \o. 1 22 BuIIelin Museum of Comparative Zooh^ij. 138,

D. I. nitidus Mayr fomiJiaris (Olliff) D. laticomis (Lea) E. clacatus Mjoberg D. 7. nifonioer (Lowne) cimeiformis (Oke) D. Iridomyrmex sp. opicalis (Oke) D. intricatus (Lea) D. kingi (Westwood) D. laticomis (Lea) D. Jongicornis (Lea) D. mctoUicus (Lea) D. stri^icoUis Lea P. longicoUis \\'estwood Formicinae Plagiolepidini D. Acantholepis capensis Mayr formicoh Peringuey F. Smith) D. Peringuey Anoplolepis custodiens ( formicoh D. A. steingweveri (Forel) formicoh Peringuey Oecophvllini OecophyUo smoragdina (Fabricius) D. mijrmecophihis (hea.) Camponotini Camponotus {Mijrmobrachys) senex and Reichardt (F. Smith) F. horgmeieri Lawrence F. uhcclcri Lawrence and Reichardt C. (Myrmothrix) bugnioni Forel F. Lawrence and Reichardt C. (Myrmothrix) renggeri Emery horgmeieri

ACKNOWLEDGMENTS Melbourne, Australia (A. Xeboiss); Ohio State University, Columbus, Ohio (R. W. the mdivid- We are indebted to following Hamilton and'B. D. Valentine); South for available uals and institutions making _^^,straHan Museum, Adelaide, Australia to us collections in their care: M. Alva- (Q. F. Gross and A. X. McFarland); United Rio de Brazil; T. renga, Janeiro, Borgmeier, g^^^^^ National Museum, \\'ashington, D. C. Rio de Janeiro, Brazil; British Museum and T. ( j. m. Kingsolver J. Spilman). London, (J. (Xatural History), England ^y^ ^j^^ ^^.-^j^ ^^ ^Yiank W. L. Brown, Jr, Balfour-Browne and R. D. Cahfor- Pope); ^^- ^^^ Kempf, and R. \\'. Taylor for ant San Francisco, nia Academy of Sciences, determinations and information on the California B. Leech); California Insect H. (H. i^i^logy and distribution of ant species, Survey, University of California, Berkeley, Andreae for information concerning Dip- California A. H. E. (J. Chemsak); Departamento Jocotidiis, and P. J. Darlington, Jr., de Zoologia, Secretaria da Agricultura. Sao Evans, H. F. Howden, E. Mayr, E. Munroe, K. for their Paulo, Brazil (W. C. A. Bockerman, W. G. Eberhard. and M. J. West Lenko, A. Martinez, and M. A. Vulcano); criticisms and suggestions. Thanks are given K. Field Museum of Xatural History, Chicago, to NL Catron for her illustrations and to the Illinois (H. S. Dybas); Hope Department Pearson for the t\'ping of manuscript, was conducted with of Entomology, Oxford University, Oxford, Part of the research a Xational Science Foundation England (E.' Taylor); Museum National the aid of from d'Histoire Xaturelle, Paris, France (A. M. Grant (GB-4743) and a Grant-in-aid of the Xi. Villicr); Xational Museum of Victoria, the Society Sigma Myrmecophilous Ptinidae • Lawrence and Rcichardt 23

LITERATURE CITED Gemmixger, M., .axt) E. von Harold. 1968. Catalogus Coleopterorum hujusque descrip- R. 1968. Notes on the of two Andrews, biology' torum s\nonymicus et systematicus. Tom. Central American Ptiiuis Ptin- (Coleoptera: . II. Dytiscidae .. Scaphidiidae. Monaco, with a of a new idae) description species. Gumm, pp. 425-752. 74 191-202. Psyche (3): HixTox, II. E. 1941. The Ptinidae of economic Blackburx, T. 1892. Notes on Anstralian Cole- importance. Bull. Ent. Res. 31: 331-381. optera with descriptions of new species. Proc. Howe, R. W. 1959. Studies on beetles of the Linn. Soc. New South Wales 7(2): 283-300. family Ptinidae. X\TI.—Conclusions and Blatchley, W. S. 1930. On a family of Cole- additional remarks. Bull. Ent. Res. 50: 287- optera new to the fauna of North America 326. of with description one new species ( Cnos- Jaxssexs, E. 1949. Sur la massue antennaire de tidae). Ent. News 41: 108-112. Paussus Linne et genres \'oisins. Inst. Rov. Browx, W. J. 1959. Niptiis Boield. and allied Sci. Nat. Belg. 25 (22): 1-19. genera in North America (Coleoptera: Ptin- Klxg, R. L. 1866. Description of Anapesttis idae). Canad. Ent. 91: 627-633. kreusleri: a species of coleopterous insect Cazier, M. a., and M. a. Mortensox. 1965. inhabiting ants' nests in South Australia. Bionomical obserxations on m\rmecophilous Trans. Ent. Soc. New South Wales 1 (5): 16. beetles of the genus Cremastocheilus ( Cole- 316-318, pi. optera: Scarabaeidae). Jour. Kansas Ent. Soc. L.a%\'REXCE, J. F. 1967. Delimitation of the 38 (1): 19-44. genus Ceracis (Coleoptera: Ciidae) with a Ch-APIN, E. a. 1965. The genera of the Chilo- revision of North American species. Bull. corini (Coleoptera, Coccinellidae). Bull. Mus. Mus. Comp. Zool. 136 (6): 91-144. AND H. Reichardt. 1966a. Revision of Comp. Zool. 133: 227-271. , the Gnostiis and Fahrasia Clark, J. 1921. Notes on Western Australian genera (Coleoptera: 73: 30-45. ant-nest beetles. Jour. Proc. Rov. Soc. West- Ptinidae). Psyche em Australia 6 (2): 97-104. 1966b. The systematic position

of Plaumanniola Costa Lima ( Coleoptera: . 1923. A new mynuecophilous beetle. Bull. 20: 39-42. Jour. Proc. Rov. Soc. Western Australia 9 Scydmaenidae). Coleopt. (1): 44-46. Lea, a. M. 1905. On Nepharis and other ant's nest beetles taken Mr. C. Gondie at Costa Lima, A. da. 1962. Micro-coleoptero by J. Proc. Rov. Soc. Victoria 17 representante da nova subfamilia Plauman- Birchip. (n.s.) 371-385. niolinae (Col., Ptinidae). Rev. Bras. Biol. (2):

. 1906. of new of 22 (4): 413-418. Descriptions species Australian Part \TII. Proc. Linn. Crowsox, R. a. 1955. The Natural Classifica- Coleoptera. Soc. New South Wales 31 (2): 19.5-227, tion of the Families of Coleoptera. London, 18. Lloyd, 187 pp. pi. . 1910. Australian and Tasmanian Cole- . 1962. Observations on Coleoptera in optera inhabiting or resorting to the nests of Scottish oak woods. Clasgow Nat. 18: 177- ants, bees, and termites. Proc. Rov. Soc. 195. Victoria (n.s.) 23 (1): 116-230, pis. 25-27. Daxa, J. D. 1851. Conspectus Crustaceorum . 1912. Australian and Tasmanian Cole- in orbis terraruni quae circumnavigatione, optera inhabiting or resorting to the nests of Carolo Wilkes e classe Reipublicae Foederatae ants, bees, and termites. Supplement. Proc. duce, lexit et descripsit. [Part.] Proc. Acad. Rov. Soc. Victoria (n.s.) 25 (1): 31-78, Nat. Sci. Philadelphia 5: 247-254, 267-272. pi. 2.

Dahlixgtox, p. J., Jr. 1950. Paussid beetles. . 1914. Notes on some miscellaneous Trans. Amer. Ent. Soc. 76: 47-142. Coleoptera, with descriptions of new species. DoxisTHORPE, H. 1927. The Guests of British Trans. Proc. Rov. Soc. South Australia 38: Ants. Their Habits and Life Histories. Lon- 249-344, pi. 16.'

. 1917. Notes on some miscellaneous don, Routledge, 244 pp. Coleoptera, with descriptions of new species. Fall, H. C. 1901. List of the Coleoptera of Part III. Trans. Proc. Rov. Soc. South Aus- Southern California. ... San Francisco, Calif. tralia 41: 121-322, pis. 12-15. Acad. Sci., 282 pp. . 1919. Notes on some miscellaneous

. 1905. Revision of the Ptinidae of boreal Coleoptera, with descriptions of new species. America. Trans. Amer. Ent. Soc. 31: 97- Part V. Trans. Proc. Roy. Soc. South Aus- 7. 296, pi. traha 43: 166-261. pis. 25-27. T. \l. 1926. F(jRBES, W. The wing folding LixsLEY, E. G. 1944. Natural sources, habitats, patterns of the Coleoptera. Jour. New York and reservoirs of insects associated with stored Ent. Soc. 34: 42-68, 91-1.39, pis. 7-18. food products. Hilgardia 16: 187-224. Vol. No. 1 24 Bulletin Museum of Comparative Zoology, 138,

AND W. MacSwain. 1941. The bio- Reitter, E. 1911. Fauna Germanica. Die Kafer , J. III. nomics of Ptiniis calijornicus, a depredator in des deutschen Reiches. Band Stuttgart, the nests of bees. Bull. So. Calif. Acad. Sci. Lutz, 436 pp. S. In: W. 40 (3): 126-137, pis. 11-14. Schenkling, 1935. Ectrephidae. Junk and S. cata- Martinez, A., and M. J. Viana. 1964. Una Schenkling, eds., Coleopterorum nueva subfamilia de Coleopteros (Ptinidae; logus 10 (141): 3-4. Fabiinae). Neotropica 10: 7-14. Sharp, D., and F. Muir. 1912. The compara- tive of the male tube in . 1965. Un caso de homoniniia genital , anatomy Trans. Ent. Soc. London 1912: . 11: en Ptinidae ( Coleoptera ) Neotropica Coleoptera. 18. 477-642, pis. 42-78. MjoBERG, E. 1916. Results of Dr. E. Mjoberg's Stickney, F. S. 1923. The head-capsule of Swedish scientific expeditions to Australia Coleoptera. Univ. Illinois Biol. Monogr. 8 1910-1913. 8. Ptinidae. Ark. Zool. 10 (6): (1): 1-104. 1-15. Tillyard, R. J. 1926. The Insects of Australia Mou, Y. C. 1938. Morphologische und histo- and New Zealand. Sydney, Angus and Robert- logische Studien iiber Paussidendrusen. Zool. son, 560 pp. G. S. 1960. Torrc-Bueno's Jahrb. (Abt. Anat.) 64: 287-346. TuLLOCH, — Glossary Oke, C. 1926. Two entomologists in the Mallee. of Entomology' Supplement A. Merrick, Vict. Nat. 42: 279-294. New York, Brooklyn Ent. Soc, 36 pp. E. 1894. Kritisches \'erzeichniss der . 1928. Notes on Australian Coleoptera, Wasmann, Arthro- with descriptions of new species. Part i. Proc. myrmekophilen und termitophilen Linn. Soc. New South Wales 53 (2): 1-30. poden. Berlin, Dames, 231 pp. und Ter- . der Ameisen Olliff, S. a. 1886. Remarks on Australian 1898. Die Gaste 3: 161- Ptinidae and descriptions of new genera and miten. Illustr. Zeit. Ent. 145-149, species. Proc. Linn. Soc. New South Wales 164, 179-181, 19.5-197, 209-211, 225-227, 10: 833-840. 243-246. einer Park, O. 1942. A study in Neotropical Psel- . 1916. Wissenschaftliche Ergebnisse Ostindien. ... V. Ter- aphidae. Northwestern Univ. Stud. Biol. Sci. Forschungsreise nach Med. 1: 403 pp. mitophile und myrmecophile Coleopteren. . . . the Zool. 39: 169-210, 4-5. . 1964. Observations upon behavior Jahrb. pis. ihre of mvrmecophilous pselaphid beetles. Pedo- . 1920. Die Gastpflege der Ameisen, Probleme. biologia 4: 129-137. biologischen imd philosophischen zur theoretischen Pascoe, F. P. 1866. [Note.] Trans. Ent. Soc. J. Schaxel's Abhandlungen xvii 176 London, ser. 3, vol. 5, Jour. Proc, 1866: Biologic, Heft 4, + pp. und xv-xvii. . 1928. Ptinidae aus Sumatra Java zur Kenntnis der Perixguey, L. 1899. Fifth contribution to the (Coleopt. ). (275 Beitrag Ent. Mitt. 17: 242-244. South African coleopterous fauna. Ann. South Myrmecophilen. ) 1925. and African Mus. 1(2): 240-330, pis. 6-7. -, and H. Brauns. New genera of South African and Pic, M. 1903. Diagnoses generiques et spe- species myrmecophilous beetles. So. Afr. Nat. Hist. cifiques de divers Coleopteres exotiques. termitophilous L'Echange 19: 182-183. 5: 101-118, pis. 9-10. O. 1855. of a new . 1909. Ptinidae. In: W. Michaelsen and Westwood, J. Description insects the R. Hartmeyer, eds.. Die Fauna Sudwest- genus of coleopterous inhabiting Trans. Ent. Australiens 2 (13): 215. interior of ants' nests, in Brazil. S. Soc. Ser. 2, 3: 90-94, pi. 8. . 1912. Ptinidae. In: W. Junk and London,

. on the Schenkling, eds., Coleopterorum catalogus 10 1869. Remarks genus Ectrephes, exotic (41): 1-46. and descriptions of new Coleoptera. 1929. liber Wasmann's Ectrephidae. Trans. Ent. Soc. London 1869: 315-320. Oxon- . 1874. Thesaurus Entomologicus Wien. Ent. Zeit. 46 ( 1 ) : 34-35.

ensis. . .. O.xford, Clarendon, 205 pp. Poll, J. van de. 1886. Description of a new Social Insects. gnostid. Notes Leyden Mus. 8: 238. Wheeler, W. M. 1928. The Har- Reddell, J. R. 1966. A checklist of the cave Their Origin and Evolution. New York, fauna of Texas. II. Insecta. Texas Jour. Sci. court. Brace, 378 pp. 18 (1): 25-56. Zeck, E. H. 1920. Myrmecophilous Coleoptera. Af- Reichensperger, a. 1948. Die Paussiden Austral. Zool. 1 (8): 245-247, pi. 19. rikas. Abhandl. Senckenberg. Naturforsch. Ces. No. 479, 32 pp. (Received 19 October 1967.) Myrmecophilous Ptinidae • Lawrence and ReicJiaidl 25

0.13 mm

Plate 1. 1-10. Facial Figs. region (clypeus, labrum, and mandibles) of various Ptinidae. Fig. 1. Diplocotes familiaris (Ol- liff). Fig. 2. Ptinus californicus Pic. 3. formicola Fig. Diplocotidus Peringuey. Fig. 4. Diplocotes laticornis (Lea). Fig. 5. Enosifaa tristis Olliff. 6. Westwood. 7. Fig. Polyplocotes longicotl'n Fig. Diplocotes carinaticeps (Oke). Fig. 8. Diplocotes kingi (Westwood). Fig. 9. Ectrephes pascoei Westwood. Fig. 10. Ectrepbes formicarum Pascoe. Vol. No. 1 26 Bulletin Museum of Comparative Zoology. 138.

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111 25

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14. Man- 11. Plinus fur Linnaeus. 12. formicarum Poscoe. Figs. 13, Plate 2. Figs. 11, 12. Maxilla. Fig. Fig. Ectrephes 15. for- Pascoe. 14. Pf/nus fur Linnaeus. Figs. 15-22. Antenna. Fig. Ectrephes dible. Fig. 13. Ectrephes iormicarum Fig. 17. foveicollis Olliff. Fig. 18. Polyphcotes /ong/co/lis micarum Pascoe. Fig. 16. Enasibo tristis Olliff. Fig. Diplocotes 20. Westwood. Fig. 21. Diplocotes familiarls Westwood. Fig. 19. Diplocotes laticornis (Lea). Fig. Ectrephes poscoei 23-28. (with basal piece removed). Fig. 23. Diplocotes (Olliff). Fig. 22. Po/yplocotes sulcifrons (Lea). Figs. Aedeagus 25. Iormicarum Pascoe. Fig. 26. D/p/ocot/dus formi- familiaris (Olliff). Fig. 24. Diplocotes hticorriis (Lea). Fig. Ectrephes cola Peringuey, ventral. Fig. 27. Same, lateral. Myrmecophilous Ptinidae • Lawrence and Reichardt 27

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Pla^e 3. Figs. 28-34. Pronofum of various myrmecophilous Ptinidae. Fig. 28. Ectrephes formicarum Poscoe. Fig. 29. Dip-

locotidus formicola Peringuey. Fig. 30. Diplocotes laticornis (Lea). Fig. 31. Dip/ocofes longicornis (Lea). Fig. 32. Diplo- cofes famillarls (Olliff). Fig. 33. Po/yp/ocofes perforatus Lea. Fig. 34. Polyp/oco/es longicol/is Westwood. Fig. 35. Dip- locotes cuneiformis (Oke).