Revisión of the genera Nemadiopsis and Falkonemadus (Coleóptera: Leiodidae: Cholevinae)

The genera Nemadiopsis Jeannel, 1936 and Falkonemadus Szymczakowski, 1961 are redescribed and revised. The taxo­ nomic position of Nemadiopsis and Falkonemadus is analy sed using features of male and female genitalia. Two new subgenera, Nemadiopsicus subgen. n. and Nemadiopsella subgen. n. and five new species, Nemadiopsis (Nemadiopsis) grossicornis sp. n., N. (Nemadiopsicus) rotundatus sp. n., N. (Nemadiopsella) curvipes sp. n., Falkonemadus similaris sp. n. and F. avicularis sp. n. are described. Keys to the subgenera and species are given. Nemadiopsis (Nemadiopsis) edwardsi Jeannel, 1936 and N. (N.) irretusus Szymczakowski, 1962 are proposed asjunior subjective synonyms of Nemadiopsis (N.) fastidiosus (Fairmaire et Germain, 1859).


INTRODUCTION
According to Newton (1998), the family Leiodidae con sists of 5 subfamilies, one of which is the subfamily Cholevinae.All taxa in this subfamily exhibit the follow ing: occipital carina or crest (when present) resting against front of pronotum when the head is in repose; cer vical sclerites present; procoxal cavities internally closed behind.Newton (1998) proposed 6 tribes within the sub family Cholevinae, one being the tribe Anemadini Hatch, 1928.This tribe consists of 4 subtribes, including the sub tribe Eunemadina Newton, 1998, characterised by the absence of an epistomal suture and the basal lobes of the parameres of the aedeagus meeting ventrally.This sub tribe includes 19 genera distributed throughout the Nearctic, and particularly the Australian and Neotropical regions.Amongst these genera are Nemadiopsis Jeannel, 1936, found in Chile andArgentina, andFalkonemadus Szymczakowski, 1961, in Chile and the Falkland Islands.Both genera, as indicated in the description, are easily distinguished as Nemadiopsis exhibits the following: apical segment of maxillary palp usually not much shorter than, and at least 2/3 as wide at base as penultimate seg ment; occipital crest strongly elevated throughout; para meres of aedeagus each with two long setae (Figs 3,18 and 37).Whilst the apical segment of the maxillary palp in Falkonemadus is very small, much shorter than and only half as wide at base as penultimate segment (Fig. 40); occipital crest scarcely elevated above neck at mid line; parameres of aedeagus glabrous or with a very small seta (Figs 45,53 and 58).
The genus Nemadiopsis was established by Jeannel (1936), and included two species, N. fastidiosus (Fair maire et Germain, 1859), from Chile, and N. edwardsi Jeannel, 1936, fromArgentina.New descriptions of species were published subse quently by two authors (Jeannel, 1957(Jeannel, , 1962;;Szymc zakowski, 1961Szymc zakowski, , 1962Szymc zakowski, , 1965Szymc zakowski, , 1968Szymc zakowski, , 1971Szymc zakowski, , 1976) ) who provided new data on the geographical distribution of some of the species.Until 1976 the number of species belonging to the genus Nemadiopsis was 8.In this taxo nomic revision in spite of the description of 3 new species, the number of species included in the genus is 7; 3 belong to the subgenus Nemadiopsis s. str., 3 to Nema diopsicus subgen.n. and 1 to Nemadiopsella subgen n.This is because the synonymies of Nemadiopsis edwardsi Jeannel, 1936 and Nemadiopsis irretusus Szymc zakowski, 1962, and the formation of new taxonomic groups.For this reason Nemadiopsis carbonarius Szymc zakowski, 1961, is not mentioned in this paper as it belongs in another genus and neither is Nemadiopsis malleatus Szymczakowski, 1976, which is the type species of a new genus; these changes will be published separately.All the species of the genus Nemadiopsis are known only from Chile and Argentina. Salgado (1991) provides data on three species of the genus Nemadiopsis, and Newton (1998) and Perreau (2000) include all the species in their respective cata logues.
The genus Falkonemadus was established by Szymc zakowski (1961) for F. sphenisci Szymczakowski, 1961.The specimens were found in the corpse and stomach of the penguin, Spheniscus magellanicus Forster, 1781, on the Falkland Islands.No data was published on this genus since 1961.Therefore the two species described in this paper and the data on F. sphenisci are new.Nemadiopsis and Falkonemadus specimens were collected in different types of animal or vegetable material, in this case in humus and fallen leaves in Nothofagus (N.dombeyi, N. pumilio, N. obliqua, N. betuloides and N. alpina), Araucaria araucana and Pinus sp.forests, and beneath Chusquea, Saxegothaea, Podocarpus and Drimys.All collections were made at altitudes of between 25 and 1650 metres.Pitfall traps baited with excre ment or carrion (Newton & Peck, 1975), window (= flight inter cept) traps, Berlese sampling of forest floor leaf litter, and light traps were used.
Prior to the dissection of the genitalia of dried specimens mounted on cards fixed on entomological pins they were first relaxed in boiling water and then softened in 10% KOH for 24 hours.The dissected specimens were then mounted as previ ously described and the genitalia mounted in "Hoyer liquid" on microslides, attached to the same pin.
Trilobed aedeagus with or without setae on lateral mar gins of median lobe (Figs 3,8,18 and 24), internal sac with different structures depending on the species (Figs 9, 14, 29 and 37); parameres with two closely inserted apical setae with or without setae on interior margins (Figs 14,29 and 37).The morphology of female genitalia, which has not been previously examined, is not uniform.In many cases, this will allow the species to be differenti ated from others.The spiculum ventrale, an anterior pro jection of abdominal sternite VIII is very different in some of the species (Figs 5,10,15,31 and 38), as well as the spermatheca, which can be classified as follows (Per reau, 1989), "type 1" with differentiated and well sclerotized lobes (Figs 6,11 and 16) and "type 3" with the spermatheca in the form of an almost completely mem branous elongated globose sac (Figs 21,26 and 32).
5th, 6th, 8th, 9th and 10th antennal segments transverse (Fig. 27).Median lobe of aedeagus wide medially with noticeable small marginal setae (Fig. 29).Internal sac of aedeagus with groups of long sclerotized fibres in the form of an arch (Fig. Diagnosis.Certain characteristics of this subgenus are of considerable taxonomic importance: the transverse 6th, 8th and 10th antennal segments (Figs 2,7 and 12); lateral margins of median lobe lack setae, basal lamina of aedeagus shorter than median lobe (Figs 3,8 and 14); posterior margin of sternite of VII abdominal segment in the female lacks a discernible lobe; and spermatheca with two sclerotized lobes joined by a membrane or hyaline area (Figs 6,11 and 16).
The two main characters defining this subgenus: the lat eral margin of the median lobe lacks setae and the spermatheca has two clearly sclerotized lobes.
Pronotum transverse, 1.65 times as wide as long; coarse, strong, dense, granulate punctures; sides weakly arched, widest at basal third, base wider than anterior region.Elytra together 1.46 times as long as wide and 3.13 times as long as the length of pronotum.Strioles transverse and easily visible, fairly regular and perpen dicular to suture.Median region of mesosternum with feebly-defined carina.Male protarsi expanded, first tarsomere 0.92 times wider than maximum width of tibia: first mesotarsus dilated and somewhat expanded outwards.Genital segment complete (see Salgado, 1991), slightly wider than long (1.08).
Aedeagus with elongate median lobe and blunt apex (Fig. 3).Basal lamina shorter than median lobe, with margin in mid apical region tapering into rounded lobe.Wide, robust parameres, clearly surpassing apex of median lobe; two setae with insertion pores separated.Internal sac with strongly sclerotized structures, in par ticular, a ventral piece on the median base in the shape of a robust tooth expanded at the base; the tooth is normally covered by an elongated oval-shaped lamina; latero-basal spines and fully-formed scaly areas alternating with spines along the median region.
Diagnosis of female.The female is generally more robust than the male, with shorter antennae and more transverse club segments: protarsi and first mesotarsal segment not dilated.
Posterior margin of sternite of VII abdominal segment a wide arch, slightly pointed in the middle (Fig. 4); VIII abdominal segment with spiculum ventrale fully formed towards front, towards back, in the median region of the sternite there is a sclerotized rod shape, which will be referred to as the "sagitta" (Fig. 5)."Type 1" spermatheca (Perreau, 1989) with two sclerotized lobes separated by a membranous region.Apical lobe round, basal lobe slightly oval in shape, wider and larger (Fig. 6).
Median lobe of aedeagus pointed with slightly concave sides (Fig. 8).Base of basal lamina somewhat shorter than median lobe, mid sternal apex in the form of rounded projection.Robust parameres with expanded apices and internal margins with membranous lamina, surpassing the apex of median lobe; fairly long apical setae inserted near external margin with insertion pores very close together.Fully developed and complex sclerotized armature of internal sac (Fig. 9), with numerous spines on the middle, outer ones are symmetrically set lengthways, with imbri cated bases, two large spines differentiated in the ventral median region when the sac is invaginated.Finally, there is a unique arch-shaped structure in the mid basal region.
Diagnosis of female.The female has the same external morphological characteristics as the male, except for the protarsi and mesotarsi, which are not dilated.
Other distinguishing characters are: posterior margin of sternite of VII abdominal segment with slightly pro truding arch in the middle, very similar to N. rufimanus; apex of sternite of VIII abdominal segment slightly indented, spiculum ventrale quite wide and sagitta either very weakly discernible or non-existent (Fig. 10).The spermatheca has two well-defined sclerotized lobes and the interconnecting membranous area is long (Fig. 11).
Remarks.The most outstanding characteristics distin guishing this species are the dense, shallow punctures on the head and the transverse 6th-10th antennal segments; also, the shape of the aedeagus with the complex struc tures of the internal sac, especially the two large spines and the unique arch shape.
Discussion.After examining several specimens from different places, variations were observed in certain char acters, for example, the male protarsi and the expansion of the antennal segments, the width of the apophysis of the genital segment, as in the spiculum ventrale, the more or less extensive membrane of the internal margin of the parameres, the length of the interconnecting area between the spermathecal lobes, and in particular, the concavity of the margins of the median lobe of the aedeagus.
Bearing this in mind, and after examining specimens (2 males and 2 females) belonging to N. irretusus from Rio Negro (Argentina) determined by Szymczakowski and at present in the Natural History Museum, Budapest (Hun gary), as well as numerous specimens from different places in the provinces of Concepción and Chiloé, it was clear that in some the margins of the median lobe are rec tilinear and the spiculum ventrale narrow, whereas in oth ers, the margins are slightly more concave and the spiculum ventrale wide.
According to Szymczakowski (1968), the main differ ence between N. fastidiosus and N. irretusus is that the lateral margins of the median lobe are rectilinear, which is a very variable character.Also, some of the differences used by Szymczakowski (1962Szymczakowski ( , 1968) ) to distinguish between these two species are unreliable.I therefore pro pose that N. irretusus is ajunior subjective synonym of N. fastidiosus.
I must point out that I have not seen specimens of N. brevicollis Jeannel, 1962, considered by Szymczakowski (1968) to be synonym ofN.irretusus.
After examining the "holotype" of Nemadiopsis edwardsi Jeannel, 1936 in the British Museum collection, I must point out it does not differ significantly from the numerous specimens of N. fastidiosus.Neither the shape of the male protarsi, which are clearly wider than the tib iae, nor the antennal segments differ.Similarly, the aedeagus and genital segment of N. edwardsi have the same structure as in N. fastidiosus.In the latter, the inner margins of the parameres are membranous and the pores of the setae are very close together on the external margin.The internal sac has the same structures: two large spines, two rows of imbricated spines and a unique arch-shaped piece.The only specimen described of Nemadiopsis edwardsi in fact belongs to N. fastidiosus, and therefore N. edwardsi should be considered a junior subjective synonym of Nemadiopsisfastidiosus.
Well-defined discrete punctuation on head.Punctures have smaller diameters than distance separating them.Slender maxillary palps, penultimate palpomere not dilated, last one pointed, scarcely longer than previous one.
Pronotum 1.73 times as wide as long, sides uniformly arched, posterior edge wider than anterior edge, widest at middle; small, dense punctures, not granulate.Elytra together 1.69 times as long as wide and 3.42 times as long as the length of pronotum.Carina of mesosternum very fine, slightly raised posteriorly.1st and 2nd male protarsi expanded, 3rd and 4th weakly expanded, with first tarsomere as wide as the maximum width of tibia (1.00).First segment of intermediate tarsi very slightly dilated and not expanded outwards.Genital segment complete (Fig. 13), clearly wider than long, with short, wide apophysis.
Median lobe of aedeagus wide, apex wide and rounded (Fig. 14).Basal lamina slightly shorter than median lobe, ending in an acute angle in mid apex region.Parameres do not extending beyond apex of median lobe, expanding into large apical clubs with two long setae inserted in clearly separated pores; internal margin of parameres with membranous apex and numerous tiny protuberances resembling microdentation laterally (O.M.: 20 x magnifi cation).Inner sac, with four groups of spines and numerous long, narrow symmetrically arranged fibres in the middle; various scaly areas in basal region.
Diagnosis of female.Shape of body similar to male, except for the narrow protarsi and mesotarsi.
The following characters differ: weakly developed pos terior median lobe on sternite of VII abdominal segment: barely marked indention on apex of sternite of VIII seg ment, long spiculum ventrale, somewhat expanded towards apex, sagitta well defined and surpassing median region of sternite (Fig. 15).Spermathecal lobes small, well-sclerotized, the larger of the two being the apical one (Fig. 16).Discussion.This species is easily distinguishable from others belonging to the genus by the shape of the antennal segments, the 7th of which is very robust, and the 4th, 5th and 6th very small segments of the funiculus.The most significant differences are in the overall shape of the aedeagus, which is not pointed at the apex, the parameres with numerous small teeth on the internal margin and the separate insertion pores, as well as the internal sac with 4 groups of elongated fibres and spines.

Type material
Etymology.From the Latin grossus (thick) and cornus (horn), refering to one of the most significant differentiating character istics, the robustness of the 9th and 10th segments, and in particu lar, the 7th antennal segment.

Subgenus Nemadiopsicus subgen. n.
Type species.Nemadiopsis barbarae Szymczakowski, 1968 Diagnosis.The most important characters of the sub genus are: the transverse antennal segments, or at least the 6th and 8th (Figs 17,22 and 27); the genital segment as long as or slightly longer than wide (Figs 23 and 28); the insertion pores of the two setae on the parameres very close together; lateral margins of the median lobe with setae; basal lamina of aedeagus longer than median lobe (Figs 18, 24 and 29); median posterior margin of sternite of VII abdominal segment of females with a differentiated lobe (Figs 19 and 30); membranous sac-shaped spermatheca (Figs 21,26 and 32).
The subgenus can be defined by two basic characters: the lateral margins of median lobe bearing setae and the membranous sac-shaped spermatheca.
Pronotum transverse, 1.56 times as wide as long, sides feebly rounded and not sinuate.Strong, somewhat granu late punctures.Elytra together 1.64 times as long as wide and 3.24 as long as the length of pronotum.Strioles trans verse, strong and almost perpendicular to suture.Mesosternum with fine carina on midline, slightly more pronounced on posterior third.Male protarsi expanded, first segment wider than greatest width of tibia (1.3); intermediate tarsi with first segment weakly dilated and slightly expanded outwards.Genital segment slightly longer than wide (see Szymczakowski, 1968), with short, broad sternal apophysis.
Median lobe of aedeagus lanceolate, narrowing slightly at apex, which has micropunctuation (O.M.: 20 x magni fication) and 3 to 6, normally 3, short setae on lateral margins (Fig. 18).Elongate tubulose basal lamina, longer than median lobe, margin of apex in the form of an arch with wide edge.Robust, wide parameres, completely sclerotized surpassing apex of median lobe; two closely inserted apical setae.Lamina of tegmen well-developed and enclosing basal lamina ventrally and dorsally.Internal sac with small spines on mid region of median lobe and on basal region of lamina base, two long lateral rows of spines on anterior region.Arch-shaped, acute scaly areas along the middle.
Diagnosis of female.The external morphology of the female is very similar to that of the male, but the pro-and mesotarsi are not dilated.
The most important characteristics are the sternite of the VII abdominal segment has a well-developed lobe on the median margin of the apex (Fig. 19); apex of tergite and sternite of VIII abdominal segment rounded or slightly cloven, narrow spiculum ventrale with diffuse sagitta penetrating sternite (Fig. 20).The membranous spermatheca is in the shape of an elongated sac of uni form width (Fig. 21a, b).
Remarks.As indicated by Szymczakowski (1971), though generally smaller and the segments of the antennal club are not elongated, or the male protarsi as expanded, the morphology of this species is very similar to that of Nemadiopsis trichophorus.The most significant differ ences are, undoubtedly, the general structure of the aedeagus, the spermatheca and the VIII abdominal sternite of the female.The paratypes in the Natural History Museum collection, Budapest (Hungary), and specimens from the type locality, El Puelo (Argentina), lack the furrow on the dorsal face of the median lobe mentioned by Szymc zakowski (1968), but do have marginal setae, of great sig nificance but not mentioned by this author.Nemadiopsis trichophorus'.Perreau, 2000: 63.Redescription of male.Length: 3.35-3.65mm.Body elongate and oval-shaped.Colour generally light brown.
Pronotum 1.54 times as wide as long, uniformly arched sides, maximum width near middle.Punctures strong, dense and somewhat granulate.Elytra elongate, 1.60 times as long as wide and 3.00 times as long as the length of pronotum.Strioles transverse, easily visible and quite uniform.Mesosternum with fine carina on midline, some what raised on posterior third.Anterior tarsi with first segment very rounded and much wider than maximum width of tibiae (1.57); 2nd and 3rd segments dilated and cordiform.Intermediate tarsi with first segment dilated and expanded outwards.Genital segment resembling that of other species belonging to the genus, somewhat longer than wide, sternal apophysis short and broad (Fig. 23).
Median lobe of aedeagus pointed apically, with 5 or more short, fine marginal setae, of decreasing length (Fig. 24).Basal lamina much longer than median lobe, ventral region of apex in the shape of a gently arched lobe.Para meres completely sclerotized, very sclerotized internal basal regions, wide, flattened and clearly surpassing apex of median lobe, apex rounded with micropunctuation and two rather short setae very closely inserted near the middle of the apex.Internal sac with two long rows of lat eral spines, with a scaly area in between.
Diagnosis of female.Generally more robust than the male, though the antennal club has shorter segments, and the pro-and mesotarsi are narrower.
The most important distinguishing characters are as fol lows: sternite of VII abdominal segment with welldefined median lobe on posterior margin; sternite of VIII abdominal segment with narrow spiculum ventrale and expanded sagitta reaching mid region (Fig. 25).Membra nous sac-like spermatheca dilated basally and narrow at the apex (Fig. 26).
Remarks.The main distinguishing characters of males are the 7th antennal segment, clearly longer and more robust than the 9th and 10th; the structures of the aedeagus with the narrow, pointed apex of the median lobe, at least 5 marginal setae and parameres much longer than the median lobe.The distinguishing features of females include the well-defined posterior median lobe on the sternite of the 7th abdominal segment, the shape of the spiculum ventrale and the spermatheca.All these charac ters make the species easy to identify.
Pronotum transverse, 1.60 times as wide as long, sides weakly rounded, narrower at front than back; strong, granulate punctures, with dense, transverse striae.Elytra wider than pronotum, together 1.65 times as long as wide and 3.26 times as long as length of pronotum.Mesosternum with well-defined carina along midline, somewhat raised on posterior region.Male protarsi expanded, first protarsomere very dilated, wider than maximum width of tibia (1.57); 2nd to 4th segments dilated and cordiform, becoming progressively more narrow.Mesotarsi with first segment weakly dilated and not expanded outwards.Genital segment complete, as long as wide (Fig. 28), short, broad sternal apophysis.
Aedeagus with median lobe strongly expanded in median region and pointed at apex, bearing 5 to 9 very small marginal setae (O.M.: 20 x magnification), anterior 2 or 3 somewhat longer (Fig. 29).Apex of median lobe and internal apex of parameres with clearly visible micro punctuation (O.M.: 20 x magnification).Basal lamina almost twice as long as median lobe, apical margin gently arched.Robust parameres with internal basal regions strongly sclerotized, surpassing apex of median lobe; broad, rounded club with two fairly long closely inserted setae.Inner sac without teeth or spines, widely arch shaped structure in laminar base formed by long sclerotized fibres, and behind it, numerous spines and scales.
Diagnosis of female.The external morphology of female is very similar to that of the male, in both body shape and structure of the antennal segments, but lacks dilated pro-and mesotarsi.
The most outstanding characters are: the pointed, fullyformed lobe in the posterior median region of the sternite of the VII abdominal segment (Fig. 30); the slight notch in the mid apical region of the sternite of the VIII abdominal segment and the expanded spiculum ventrale in the anterior region, with a slightly marked sagitta in the median region of the sternite (Fig. 31).The sacciform spermatheca is very elongate and membranous (Fig. 32).Discussion.This species is easily distinguished from all other closely related species by the characters of the aedeagus, particularly the wide shape of the median lobe, the small marginal setae (O.M.: 20 x magnification), and the armature of the inner sac, with sclerotized fibres forming an arch.The closest species is, undoubtedly, N. barbarae, because of the head punctures and the shape of the antennae, although the antennal segments are com paratively wider in the new species and the male protarsi are more dilated; the genital segment, VII and VIII abdominal segments of females and general shape of the spermatheca are all very similar.The new species resem bles N. trichophorus, especially in the antennal segments, genital segment, strongly expanded male protarsi and the VII and VIII abdominal segments of females.

Type material
Etymology.The species name come from shape of some of the most important characters of the new species, the rounded shape of the median lobe, the parameres and male protarsi.Subgenus Nemadiopsella subgen.n.Type species.Nemadiopsis (Nemadiopsella) curvipes sp.n.

Diagnosis.
Punctures on head and pronotum fine, numerous and clearly visible.Antennae very long, segments of club clearly separated (Fig. 33).Male metatrochanters strongly pointed and somewhat curved (Fig. 35).Basal lamina shorter than median lobe, very long robust parameres with membranous apex and internal margin bearing microsetae (Fig. 37).Membranous, very irregular spermatheca.
Pronotum markedly narrower than elytra, 1.35 times as wide as long, almost parallel sides, very weakly rounded and weakly narrowing towards base: dense, granulate punctures.Elytra long and convex, 1.65 times as long as wide and 3.30 times as long as the length of pronotum; strioles fine and irregular, perpendicular to sutural stria, which is whole and easily visible.Mesosternal carina scarcely discernible anteriorly, weakly defined posteriorly.Anterior tarsi with first three segments dilated (Fig. 34), first wider than greatest width of tibia (1.25), and slightly longer than wide; intermediate tarsi with first segment weakly widened and slightly expanded outwards.Elongated metatrochanters, very pointed and generally curved towards apex (Fig. 35).Genital segment complete, longer than wide, short, broad sternal apophysis (Fig. 36).
Aedeagus with median lobe subtriangular and a very acuminated apex (Fig. 37).Basal lamina shorter than median lobe, with short protuberance on median margin of apex.Parameres very long and robust, easily sur passing apex of median lobe, two fairly long, clearly separated setae at apex; internal margin with membranous apex and 7-10 microsetae (O.M.: 20 x magnification) on median region.Internal sac bearing two long rows of imbricated, robust teeth and with numerous scaly areas all over the basal, median and apical regions.
Diagnosis of female.The female has the same external morphological characteristics as the male, nevertheless the female is generally more robust, lacks dilated protarsi and mesotarsi, and the metafemurs, have a pointed apex, are shorter and never curved.
Shape of posterior edge of sternite of VII abdominal segment, a wide arch.Apex of sternite of VIII abdominal segment arch-shaped, spiculum ventrale long and narrow, with spatulate apex and median region bearing discerni ble sagitta (Fig. 38).Membranous spermatheca, irregular sac shape, globose, always difficult to define.Discussion.This species is easily distinguished from the others by exterior characters such as the shape of the widely separated antennal club segments, and the unique structure of the male metatrochanters, resembling those of Catops davidsoni Salgado, 1999, and some species belonging to the genera Nargus Thomson, 1867 and Choleva Latreille, 1796.The shape of the aedeagus is striking, with the strongly pointed subtriangular lobe, longer than the basal lamina but much shorter than the parameres bearing microsetae on the internal margins.The sclerotized structures of the internal sac are also very characteristic, with two lateral rows of robust imbricate teeth.The female spiculum ventrale is expanded at the apex.

Type material
Etymology.The species name refers to one of the most striking characters of this species, the long, very pointed some what curved male metatrochanters.
Parameres of aedeagus lacking apical setae or with sin gle, very small one (Figs 45 and 53).Internal sac bearing different sclerotized structures in the basal lamina region, set out in a very characteristic way in each species (Figs 44,51 and 58).Spermatheca membranous or very slightly sclerotized, in the shape of an elongated sac (Figs 48,55 and 61).
The characters distinguishing this genus are: the unique shape of the antennae, with almost moniliform segments; the very short last segment of the maxillary palps; the narrow apophysis of the genital segment; the apex of the parameres with or without 1 seta; the internal sac with strongly sclerotized structures in the basal lamina region; the long, narrow spiculum ventrale, lacking distinct sagitta, and the spermatheca in the shape of a small elongated sac.Falkonemadussphenisci: Perreau, 2000: 59.Redescription of male.Length: 2.40-2.70mm.Gener ally ovoid.Colour dark brown.Punctures on head fine, numerous and fairly close together.Maxillary palps with last segment reduced in size, much smaller than penulti mate segment, which is dilated and slightly longer than wide (Fig. 40).
Aedeagus with elongate, triangular median lobe, apex slightly dilated in lateral view (Fig. 44).Basal lamina shorter than median lobe, with almost straight apical mar gin.Ventral lamina of tegmen short.Parameres easily sur passing apex of median lobe, with large notch on internal margin of apex, where it becomes narrower; on the external margin of the apex of some specimens a pore can be seen and occasionally, a tiny seta (Fig. 45).Internal sac bearing symmetrical sclerotized structures with a unique design in the basal lamina area; towards the middle of the median lobe two fairly large symmetrical areas of numerous, small spines.
Diagnosis of female.The female has the same external morphology as the male, except for the protarsi and first segment of the mesotarsi, which are not dilated.
The most striking characters are: the very narrow sternite of VII abdominal segment, arch on posterior margin facing inwards (Fig. 46), stemite of VIII abdominal seg ment with long, narrow spiculum ventrale, lacking sagitta (Fig. 47).Spermatheca membranous, shaped like an elon gated sac, expanded basally and narrow apically; there is also a long narrow spermathecal duct (Fig. 48).
Remarks.This species is easily identified.The main distinguishing characters are in the genitalia, particularly the shape of the parameres and the armature of the internal sac.It should be pointed out that F. sphenisci has fully-developed membranous flight wings, as Szymczakowski (1961) states that it is apterous.Another out standing character is the varying shape of the basal mar gins of the pronotum in both sexes, which can be narrow or wide; similarly, a more or less marked narrowing can be seen in the basal margins of the genital segment; finally, the pronotal punctures varies in intensity.
Aedeagus with triangular median lobe (Fig. 51), tip of apex blunt and curved in lateral view (Fig. 52).Basal lamina shorter than median lobe, with slightly curved posterior margin.Parameres only just extending beyond apex of median lobe, wide towards apex where there is no pore or seta (Fig. 53).Internal sac in basal region of lamina with a group of sclerotized structures set close together and lacking definite shape, two symmetrical arch shapes in mid region of median lobe, bearing numerous spines.
Diagnosis of female.The shape of the body is similar to the male, except for the protarsi and first segment of mesotarsi, which are not dilated.
The shape of the sternite of the VII abdominal segment closely resembles that in F. sphenisci.The sternite of the VIII abdominal segment has a very long, narrow spiculum ventrale (Fig. 54).The spermatheca is membra nous and quite uniform in width (Fig. 55).
Discussion.The morphology of this species is similar to that of F. sphenisci.The external characters show very little difference, nevertheless F. similaris sp.n. is some what bigger, the club segments more transverse, the male protarsi slightly more dilated and the apophysis of the genital segment shorter and broader.Also, no F. similaris sp.n. specimens show narrowing in the basal region of the pronotum.There is however, a marked narrowing in the basal area of the genital segment.The most out standing differences are in the aedeagus, especially in the blunt apex of the median lobe, the parameres, which extend slightly beyond the apex of the median lobe and widen towards the apex, and the sclerotized structures of the internal sac have a completely different shape.
Etymology.The species name refers to its great similarity to F. sphenisci.
Pronotum small, 1.48 times as wide as long, more narrow than elytra, weakly rounded sides, almost recti linear in posterior third, posterior angles very obtuse.Punctures somewhat granulate, shallow and dense.Elytra elongate, weakly convex with uniformly arched sides, 1.68 times as long as wide and 3.55 times as long as the length of pronotum; striolae transverse, strong, scattered and slightly oblique.Protarsi dilated, with first segment slightly wider than maximum width of tibiae (1.10).Inter mediate tarsomeres with first segment clearly dilated.Genital segment as long as wide, with short, narrow apo physis and latero-basal margins clearly notched.(Fig. 57).
Aedeagus with apex of median lobe very rounded (Fig. 58).Basal lamina longer than median lobe, with ventral margin of apex almost straight.Ventral lamina of tegmen almost as long as basal lamina, weakly defined and scarcely protruding.Parameres robust, not surpassing apex of median lobe, bearing a unique shape resembling a bird's head at apex, and a single very short seta on external margin.Internal sac with well-sclerotized, paired symmetrical structures on laminar base, a small elongate group of spines on median lobe.
Diagnosis of female.The external morphology of the female is the same as that of the male, including body shape, antennae and punctures on head and pronotum, protarsi and first segment of mesotarsi not dilated.
Outstanding characters include: a wide arch on the pos terior margin of the VII sternite of the abdominal segment, with a slight lobe in the middle (Fig. 59); the VIII sternite of the abdominal segment has a very long spiculum ventrale, without a differentiated sagitta (Fig. 60).The spermatheca is an elongate sac shape, with slightly sclerotized walls, wider at the base and narrower at the apex (Fig. 61).It is very likely that this structure is associated with a large spermatic gland, which is very dif ficult to distinguish.Discussion.Falkonemadus avicularis sp.n. is easily differentiated from F. sphenisci and F. similaris sp.n., by the less rounded shape of the pronotum, the shorter apo physis of the genital segment and in particular the unique structure of the apices of the parameres and the sclerotized shapes in the internal sac.These structures make this species unmistakable.

Type material
Other characters such as the weakly defined punctures on the head and pronotum, the basal lamina, which is longer than the median lobe, the very long lamina of the tegmen and the slightly sclerotized spermathecal walls, markedly differentiate this species from the others.
Finally, external morphological characters, including the general shape of the body, pronotum and antennae of F. avicularis sp.n. resemble certain species belonging to the genus Nemadiolus, especially N. oblongus.On the other hand, the internal sac of the aedeagus, the shape of the spermatheca and fully-developed spiculum ventrale, with no sagitta make it impossible to place this species in the genus Nemadiolus.
Etymology.The species name refers to the shape of the apex of the parameres, which in outline is reminiscent of a bird's head.
ACKNOWLEDGEMENTS.I am most grateful to A.F. Newton, Associate Curator of the Field Museum of Natural History, Chi cago, Illinois, for reviewing the manuscript and pointing out the necessary changes, and to both A.F. Newton and P.P. Parrillo, Division of Insects (FMNH), for allowing me access to inter esting entomological material.I would like to thank the National Museum of Natural History (Paris), Mme.N. Berti; the Hun garian Natural History Museum (Budapest), O. Merkl, and the Natural History Museum (London), M. Brendell, for lending me the material, which was of great help in carrying out the comparative studies.