The genus Anthelephila ( Coleóptera : Anthicidae )

The genus Anthelephila Hope, 1833 and its type species, Anthelephila cyanea Hope, 1833, are redescribed. Based on examination of the type material, the following new synonymy is proposed, Anthelephila Hope, 1833 (= Formicoma Motschoulsky, 1845 syn. n.) and A. cyanea Hope, 1833 (= Notoxus caeruleus Thunberg, 1787 syn. n.). Five genus-group names are regarded as unjustified emendations and are placed as synonyms: Anthelephila Hope, 1833 (=Anthelephilus LaFerté-Sénectére, 1849; Formicosoma Motschoulsky, 1845; Myrmecosoma Mannerheim, 1846; Formicomus LaFerté-Sénectére, 1849; Orthauchen KrekichStrassoldo, 1925 syn. n.). A lectotype is designated for Anthelephila cyanea Hope, 1833.


INTRODUCTION
There are two closely related genera currently placed in the Formicomini: Anthelephilus LaFerte-Senectere, 1849, comprising about 20 species, and Formicomus LaFerte-Senectere, 1849 comprising more than 350 species.Unfortunately, the present nomenclatorical status of both these names is in serious contradiction with the ICZN (1999) rules, since they were knowingly and unjustifiedly proposed by Hope (1840) and LaFerte-Senectere (1849a, b) to replace the older names Anthelephila Hope, 1833 and Formicoma Motschoulsky, 1845, with new spellings for these two genera accepted by nearly all subsequent authors.
The present paper is aimed to solve the above men tioned problem, and to provide a redescription of the genus Anthelephila Hope and its type species.Further more, brief remarks are made upon the general topics of biology, geographic distribution, and position of Anthele phila within the subfamily Anthicinae.

M ATERIAL AND M ETHODS
Specimens were examined with a stereoscopic microscope using diffused lighting, and illustrations were made using an ocular microgrid.Morphological measurements were taken using the ocular micrometer.Male and female genitalia were illustrated after having been cleared by boiling in 10% KOH solution.
Author's comments on the type material are found in square brackets.Exact label data are quoted for the type specimens only.Separate labels are indicated by slashes (\).The publica tion date o f the papers by LaFerte-Senectere follows Chandler (2000) .

Taxonomic history of the treated generic names
Anthelephila, Anthelephilus.The genus Anthelephila was originally established by F. W. Hope as a subgenus of Anthicus Paykull, 1798.The description was initially read at the meeting of the Zoological Society in London on May 28th 1833, and then published as a report of this meeting in an abbreviated format (comprising only more important characters of the genera and species described) in July of the same year in the Proceedings o f Zoological Society o f London (Hope, 1833).The full description was published the next year (Hope, 1834) in the Transactions o f the Zoological Society o f London.Since the description in the Proceedings is clearly attributed to Hope and lists several characters for recognition of the genus Anthele phila and its type species, Anthicus cyaneus Hope, it is regarded as the original description (cf.Sherborn, 1928;Neave, 1939).Saunders (1834) added another two species and was the first to treat Anthelephila as a genus, without providing any comments on his rationale for this eleva tion.A change in gender to Anthelephilus was first used, without comments, by Hope (1840) to replace Anthele phila.Although he used Anthelephilus in combination with Hope's authorship, LaFerté-Sénectere (1849a, c) was apparently quite unaware of the preceding paper by Hope, and proposed more formally the same emendation, along with a generic description.
According to the ICZN (1999), both Formicosoma and Formicomus are unjustified emendations of Formicoma.On the other hand, Formicoma was used later as a valid name only by Kolenati (1846) and Motschoulsky (1849), and the youngest name Formicomus is undoubtedly in prevailing usage and could thus be conserved (ICZN, 1999: Article 23.9.1.1-2).This action, however, would be unnecessary for the reasons described below.
Myrmecosoma.The name Myrmecosoma was proposed by Mannerheim (1846) to replace Formicosoma, since he believed the latter name to be incorrectly composed.It was used later only by Truqui (1855) and usually has been treated as synonym of Formicomus.According to the ICZN (1999), it is regarded as unjustified emendation of Formicosoma.
Orthauchen.The subgenus Orthauchen was estab lished by Krekich-Strassoldo (1925) to accomodate the species of Formicomus with long neck smoothly merging with head.It was recently synonymized with the nominotypical subgenus by Kejval (2000).
Carteromerus.Pic (1911) listed Carteromerus as synonym of Formicomus with the following data: "Car teromerus Laf.(Monogr. 1848, p. 71, nota 1)."According to the respective note (pages 2, 71) in LaFerté-Sénectère (1849b, c), Carteromerus is the original name LaFerté-Sénectère meant to use before he decided to emend Motschoulsky's Formicoma.It occurs on the original labels in the LaFerté-Sénectère's collection, however it was never officially established since there were no explicitly included species, and he never used it as a valid name.Consequently, it is herein regarded as an unavail able name.

Comments on the newly proposed synonymy
The only character traditionally used to separate the genera Formicomus and Anthelephilus is the absence of hind membranous wings in the latter genus, which is related to the obsolete elytral humeri, and sometimes also to the truncate elytral apices.This difference was first noted by LaFerte-Senectere; although he had not seen the type specimen of Anthelephila, he assumed its aptery solely from a copy (Fig. 8) of the original habitus illustra tion (Fig. 9) provided by Chevrolat (LaFerte-Senectere, 1849a: 1).However, as was already stressed by some early workers (Lacordaire, 1859;King, 1869), there is no supporting evidence for this division.The reduction of hind wings is well known in numerous genera of Anthicidae, and their tendency toward reduction has been observed in a group of closely related species of Formi comus (Kejval, 2000).The most widely known species treated under the name Anthelephilus (A. ruficollis Saun ders, 1834) appears to vary in this character as f have seen fully winged, brachypterous, and apterous specimens.Finally, examination of the type material has shown that the type species of the genus Anthelephila is not apterous (see the redescription of A. cyanea below).
Motschoulsky (1845a) designated "Anth.pedestris F." as the type species of Formicoma.Why Motschoulsky and Dejean (1837) attributed this species to Fabricius is not clear.ft is evident from the data given by Fabricius (1801), that hejust proposed one new combination for the species described as Carabus pedestris by Rossi (1790), and accordingly Anthicus pedestris is not listed as a Fabrician species by Zimsen (1964).The major part of the collection of P. Rossi is now deposited in MNHB as a part of the so called "Historical collection," and present there is the series labelled "Pedestris R.* [R.= Rossi, * = type material], Anthicus.pedestris Fab., N. thoracicus Pz.," comprising specimens of various origin, including the type(s) of Carabus pedestris Rossi, 1790.Having examined the whole series, f found it homogeneous and belonging to Formicomus pedestris (Rossi, 1790) in its present sense, which is a well known species clearly con generic with Anthelephila cyanea.Considering these facts, Formicoma, Formicosoma, Myrmecosoma and For micomus are all regarded as junior synonyms of Anthelephila.
Sexual dimorphism.Adults of most species of Anthelephila show rather distinct sexual dimorphism.Males are usually of smaller size and exhibit a more slender bodyform than females, sometimes have slightly longer anten nae, and display a number of conspicuous secondary sexual characters: front legs almost exclusively modified, have been found running together with ants (Hemp & Dettner, 1997;Hingston, 1925;Wasmann, 1898), even near their nests, but only a few species may be potentially regardedasmyrmecophilous (Wasmann, 1898).Relationships.Based on male characters, Bonadona (1974) established the tribe Formicomini for the genera Formicomus, Anthelephilus, Stenidius LaFerte-Senectere, 1847 and Andrahomanus Pic, 1903, which all have sternite VIII of the males strongly modified.This tribe was both omitted (Uhmann, 1976) and accepted (Bucciarelli, 1980;Hemp, 1994) by subsequent authors.Although I am not quite convinced of the status of the tribe Formicomini, the following three characters may repre sent synapomorphies suggesting (in combination) the placement of Anthelephila with Stenidius, Andrahomanus and Chileanthicus: i) mesepisterna simply connected with mesepimera, their margins not raised and bare (Fig. 11); ii) the intercoxal process of abdominal sternum III widely rounded to subtruncate and incompletely bordered (Fig. 4); iii) male sternite VIII modified, with distinct posteri orly projecting prongs.
In the other genera of Anthicinae that I have examined the postero-lateral margins of mesepisterna were found to be completely (as in Fig. 20) or at least partly raised in the posterior half above the more or less recessed anterior portion of mesepimera.Similarly, I failed to find a spe cies from any of the other genera sharing both of the fol lowing two characters.The examples where one of these characters occurs represent, in my opinion, convergent evolution rather than evidence of relationship.As far as I know, only males of Acanthinus LaFerte-Senectere, 1849 and some Vacusus Casey, 1895 possess forms of sternite VIII that may resemble the structures developed as in the above mentioned genera (see for example Fig. 43 by Werner 1970).However, the intercoxal process of abdominal sternum III in these two genera is always acutely angulate apically and with the margins completely bordered; representatives of all major groups of the large genus Acanthinus were examined, including the Austra lian species (D.S. Chandler, pers. comm.).On the other hand, the comparatively wide, apically rounded intercoxal process of abdominal sternum III occurs in various genera of Anthicinae and it can be even extremely finely, indis tinctly bordered apically (although very rarely).However, sternite VIII in males of the respective species was always found to be formed by a simple flattened sclerite, with the posterior margin at most emarginate to incised medially.The genera Phalantias Heberdey, 1936 from South India and Sri Lanka, and Falsoformicomus Pic, 1948 from Madagascar, were tentatively included in the Formicomini by Hemp (1994), but share neither of the mentioned synapomorphies and exhibit other striking dif ferences (see Bonadona, 1958Bonadona, , 1982)).
Within the group of related genera, Anthelephila differs by the following major characters: anterior angle of mesosternum distant from anterior margin of mesothorax due to the medially broadly joined mesepisterna (angle situated at least very near this margin in the remaining three genera); all femora clavate (meso-and metafemora narrow in Stenidius); ovipositor with well developed styli (ovipositor without apparent styli in Chileanthicus and Andrahomanus).
Head.Head 1.2 times as long as wide, moderately widely rounded posteriorly in dorsal view; tempora arcu ately narrowing towards base; hind temporal angles rounded; base distinctly differentiated from short neck; dorsal face of head evenly vaulted; frontal region slightly, longitudinally impressed on each side along slightly raised lateral margins.Dorsal surface distinctly, largely simply punctured; interspaces among punctures much wider than their diameter, without traces of corrugation, mostly smooth, finely punctured postero-laterally; pubes cence rather short, composed of decumbent setae and few, sparsely scattered, suberect to erect setae.
Antennae moderately long, distinctly exceeding elytral humeri, distinctly enlarged and flattened distally; antennomere I thick, 1.7 times as long as wide; antennomere II shortest, 1.2 times as long as wide; antennomeres III-VII each about twice as long as wide; antennomere X 1.2 times as long as wide; antennomere XI twice as long as wide, slightly narrowed at about midlength and bluntly pointed apically.Antennomeres mostly sparsely setose, with several longer erect setae towards apices; distal antennomeres VIII-XI much more densely and shortly setose, with erect setae less conspicuous.
Thorax.Pronotum 1.5 times as long as wide, as long as and distinctly narrower than head including eyes, widest before midlength, distinctly constricted (impressed later ally) in posterior half; anterior portion somewhat angulately shaped (Figs 5,6), with rounded edge and distinct incision laterally to collar, surface behind collar flattened to slightly concave dorsally and dorso-laterally.Dorsal punctation of pronotum similar to that of head, slightly denser mid-dorsally; antero-lateral sides with only few, very fine punctures dorso-laterally, largely smooth and glossy, bottom of latero-basal impressions shortly, finely wrinkled and small vaulted area situated posterior to impressions finely punctured.
Mesostemum (Figs 1, 2) modified, equal in both sexes; distinctly impressed along antero-lateral margins and with rather distinct, rounded postero-median bulge situated at level of anterior margins of mesocoxae; paired antero lateral impressions shallowly interconnected medially and becoming gradually more prominent laterad; surface largely smooth and glossy, only the intercoxal process more distinctly punctured and setose.Metasternum simple in both sexes, only shallowly impressed postero medially.
Elytra 1.5 times as long as wide, less elongate, rather convex, moderately transversely impressed dorsally in post-scutellar area in basal third, somewhat subtruncate apically; humeri less protruding but distinct.Surface rather glossy; punctation double, setigerous punctures sparser (especially in basal third) than on dorsal side of head and pronotum, and finer, except for few coarse punctures scattered in post-scutellar area and bearing longer tactile setae, interspaces among setigerous punc tures very finely sparsely punctured, asetose; pubescence rather short, at most slightly longer than on head and pro notum, becoming denser and shorter towards elytral api ces, composed of subdecumbent to nearly appressed short setae and slightly longer erect setae, with some distinctly longer suberect to erect setae in post-scutellar area.Mem branous wings well developed (Fig. 16) Legs.Penultimate tarsomere of all tarsi flattened dis tally, with apical margin slightly emarginate and with ter minal tarsomere articulated on its dorsal side, distinctly before apex.Male front legs modified (Fig. 22 femora moderately dilated on inner side proximally; pro tibiae with small, blunt, dent-like protrusion in distal half.Femora finely and comparatively sparsely setose, setae appressed to subdecumbent; tibiae (especially pro-and mesotibiae) distinctly more densely setose than femora; terminal margins of tibiae (especially metatibiae) with fringe of stiff, pointed, black setae laterally; tarsi with numerous longer, black bristle-like setae.
Male abdomen.Sternum VII (Fig. 23) with posterior margin shallowly emarginate medially and with short and wide, apically rounded process projecting from dorsal side of sternum closely before posterior margin; margins of median process bearing numerous long and stiff setae.Tergum VII (Fig. 24  clearly differentiated into five sclerites: paired prongs slim, moderately divergent distally, with three longitudi nal, rather densely setose edges on medial and ventral side, each prong armed apically with three more sclero-tized and somewhat pointed projections and with two membranous lobes; median sclerite rather wide, trans verse, with posterior margin straight, slightly emarginate medially; paired latero-basal sclerites medium-sized, moderately dilated on medial side.Tergite VIII (Fig. 28) nected medially, simply rounded and long setose postericomposed of a pair of flattened sclerites narrowly con-orly.
Aedeagus (Fig. 29); tegmen with apical portion 0.7 times as long as basal-piece, rather distinctly trilobed api cally, lateral lobes divergent, rounded and slightly enlarged apically, middle lobe bluntly pointed and moder ately shorter than lateral lobes; median lobe of aedeagus terminating in a pair of small, membranous projections.
Female abdomen.Sternum VII (Fig. 30) simply shaped, with posterior margin at most slightly produced medially; tergum VII similar to that in male, but more narrowed posteriorly and with posterior margin more densely and longer setose medially; both sternite and tergite VIII (Figs 17,18) conspicuously long and densely setose along posterior margin.Ovipositor (Fig. 19).Differential diagnosis.Anthelephila cyanea may be easily distinguished by the conspicuous metallic blue reflection of elytra, the angulately shaped anterior portion of the pronotum, double elytral punctation, modified mesosternum, and by the male characters.Remarks.Anthelephila cyanea was described from Australia and never subsequently recorded until now.Having examined the type material, I found this species quite dissimilar to all its Australian congeners, placed in the here synonymized genus Formicomus, and identical in all characters with F. caeruleus (Thunberg, 1787) from southern Africa.Although there are examples of close relationships between the faunas of Australia and southern Africa (see Endrodi-Younga, 1978), Anthele phila does not seem to present this situation.Based on my studies, it appears to be rather derived genus, whose members colonized Australia from the Oriental region (see Distribution of the genus above).For these reasons, I believe that either the type locality of A. cyanea is errone ous, or the respective specimens were introduced to Aus tralia.
Originally, Formicomus caeruleus was described by Thunberg (1787) in the genus Notoxus Geoffroy, 1762.Later, the species was listed under Anthicus by Dejean (1837), and then transferred to Formicomus by LaFerté-Sénectere (1849b, c).The examined syntypes are iden tical in all characters, including morphology of the prongs of male sternite VIII (their apical portion is exposed in the male specimen) with the types and the specimens additionally examined of Anthelephila cyanea.Conse quently, F. caeruleus is regarded as junior synonym of the latter species.
genus is extremely diverse in southeast Asia, but I know o f only a few species that are east o f W allace's line and occuring in Australia.All Australian species belonging to Anthelephila (about half o f known Australian "Formicomus") show a clear relationship to some o f their Oriental congeners, and they are most frequent in the humid northern and north western regions o f Australia.The species with dentate metafemora, which predominantly inhabit the southern arid regions o f Australia, belong in my opinion to the genus Chileanthicus Werner, 1966 (Kejval, in prep.).The only species known from N ew Caledonia, F. austrocaledonicus Montrouzier, 1854, belongs to a different, unrelated genus.
) simply and somewhat widely rounded posteriorly, with a pair of transverse, densely spinulose patches near base.Sternite VIII