Cladistic systematics of the genus Amphimallon ( Coleoptera : Scarabaeidae : Melolonthinae )

A phylogeny of fifty-eight cockchafer species belonging to the genus Amphimallon Berthold, 1827 is proposed, based on sixty-five morphological characters. The cladistic analysis provides seventy-two equally parsimonious trees. The genus Amphi­ mallon is redefined and species-groups are introduced and defined: A. pini-group (seven species), A. vernale-group (five species), A. solstitiale-group (seven species), A. arianae-group (two species), A. peropacum-group (one species), A. fuscum-group (eleven spe­ cies), A. naceyroi-group (seven species), A. majale group (five species), A. lusitanicum-group (six species). Other species previously placed in Amphimallon are considered species incertae sedis in this paper: amphibolum Peyerimhoff, 1949, and a monophyletic group composed of six North African species: altifrons Baraud, 1971, julieni Baraud, 1972, melillanum Baraud, 1972, scutellare Lucas, 1846, subcristatum Fairmaire, 1879, subparallelum Escalera, 1913. Four new Amphimallon species are described: A. adanense sp. n. from Turkey, A. maniense sp. n. from Greece, A. jeannae sp. n. and A. safiense sp. n. from Morocco. The following taxonomic conclusions are proposed: A. seidlitzi Brenske, 1891 = 1971) comb, n.; Miltotrogus caucasiens Gyllenhal, 1817 comb, i 1817) stat. n.; A.javeti Stierlin, 1864 stat. n.


INTRODUCTION
The cockchafers of the genus Amphimallon Berthold, 1827 form one of the most important groups in western Palaearctic Rhizotrogini by the number of described taxa.This genus, considered in its traditional definition, is composed of more than sixty species (see Appendix 1) according to the most important recent works dealing with the western Palaearctic Melolonthidae (Medvedev, 1951; Baraud, 1985Baraud, , 1992)).The present paper adds spe cies not studied bythe above authors and describes new species.These species are distributed from Morocco throughout Europe and central Asia to Siberia.In the Mediterranean Region they are absent from the south eastern part.In spring and summer, adults, and especially the males searching for females, are particularly active for a short time in the morning or in the evening twilight, flying above ground level of grassland or around the tops of the trees.Larvae feed upon plant roots.Some species, like A. solstitiale (L.), and especially A. majale (Razoumowsky), are known for causing severe damage, mostly to grass, in Europe (Régnier, 1939(Régnier, , 1940)), but also in the United States (Schwardt & Withcomb, 1943;Gambrell, 1946;Shorey & Gyrisco, 1960), where A. majale has been introduced (Gambrell et al., 1942).
Amphimallon and Rhizotrogus are traditionally distin guished by the number of antennal segments: 10segmented antennae in Rhizotrogus (Fig. 1), 9-segmented antennae in Amphimallon (Figs 2-3).However, some aberrant Rhizotrogus specimens with 9-segmented antennae have been found, and have wrongly been con sidered as Amphimallon (Baraud, 1977a;Montreuil, 1997), introducing a doubt in using this character to dis criminate Amphimallon from Rhizotrogus.In her works, Coca Abia has proposed a new diagnostic character to separate both genera: the development of the sclerotized lateral apophyses in the endophallus.These apophyses are long and strong in Rhizotrogus (Fig. 4) and clearly reduced in Amphimallon (Fig. 5).
Amphimallon is a rather well-known group, but it has not been the subject of a complete recent revision, or of a phylogenetic study.The main purpose of this paper is to establish the phylogenetic relationships of Amphimallon species using a morphological approach, in order to revise systematics of the genus.The main groups of Amphimallon that can be defined by reference to this phy logeny are introduced.Taxonomic conclusions are also offered, and the description of four new species belonging to this genus are given in the last part of this paper: Amphimallon adanensis sp.n. from Adana, Turkey, separated from A. nigripenne Reitter, 1902; A. jeannae sp.n. from Morocco, until now confused with A. theryi Peyerimhoff, 1949; A. safiense sp.n. from Safi, Morocco, separated from A. litigiosum Fairmaire; and A.  5. Amphimallon and Rhizotrogus.1-3: Head (dorsal view; setae and punctation omitted).1 -Rhizotrogus marginipes Mulsant; 2 -Amphimallon vitalei Luigioni; 3 -A.atrum (Herbst).4, 5: Aedeagus and endophallus (lateral view; p -parameres; phphallobasis; sa -sclerotized apophyses; b -body of endophallus; las -lateral apical saccules; dbs -dorso-basal area of spicules; vs -ventral area of spicules).4 -Rhizotrogus aestivus (Olivier); 5 -Amphimallon pygiale Mulsant.maniense sp.n., recently discovered in the Peloponnese, Greece.

MATERIAL AND METHODS
According to the recent general works which deal with Amphimallon (Medvedev, 1951;Baraud, 1985Baraud, , 1992)), and including described taxa not treated in these works, new taxo nomic conclusions based on comparison of types and new spe cies described in this paper, the genus Amphimallon Berthold, 1827 sensu auctorum is composed of sixty-seven species (Appendix 1).Fifty-eight Amphimallon species are considered in the phylogenetic analysis.The missing species which could not be studied in this work will be treated later in a general taxo nomic revision in preparation.
In order to solve the phylogenetic relationships between the Amphimallon species considered, the cladistic method has been used, together with the principle of parsimony (Crisci, 1982;Forey et al., 1992;Darlu & Tassy, 1993).The sixty-five mor phological characters used in this phylogenetic analysis are listed in Appendix 2. They were examined on dry mounted adult specimens, mostly conserved in the Muséum National d'His toire Naturelle, Paris, France (MNHN).Specimens of each spe cies were observed directly, and specimens of common species were dissected and observed in lactophenol.Particular structures such as endophallus, digestive duct or bursa copulatrix were extracted from the abdomen and stained (Carayon, 1951(Carayon, , 1969)).Only three characters have been found in structures because the females of many species are still unknown or were not available during this study.
The different states of each character were coded for each species in a data matrix with missing data and polymorphic characters scored "?" (Table 1).For each character, observed states at ingroup root are scored with "0" in the matrix after the analysis, in order to standardize the data presentation.In order to limit ad hoc hypotheses, the characters are unweighted, and multistate characters have been considered as unordered.The character states were polarized using outgroup comparison (Wiley, 1981;Nixon & Carpenter, 1993).Three species belonging to two closely related genera (Coca Abia, 1995) were used as outgroups: Geotrogus inflatus (Buquet, 1840), Rhizotrogus aestivus (Olivier, 1789) and Rhizotrogus marginipes Mulsant, 1842.The data were analysed using a heuristic search routine with PAUP Version 3.1 (Swofford, 1992).The resulting trees were analysed with McCLADE (Maddison & Maddison, 1993).
Taxonomic and nomenclatural conclusions proposed in this paper are based on the study of typical material deposited in the MNHN, or borrowed from the Hungarian Natural History Museum, Budapest, Hungary (HNHM, O. Merkl), from the Deutsches Entomologisches Institut, Berlin, Germany (DEIC, L. Zerche) and from the Naturhistoriska Riksmuseet, Stockholm, Sweden (NHRS, B. Gustafsson and J. Ferrer).

RESULTS AND DISCUSSION
Cladistic analysis generates seventy-two equally parsi monious trees, with a total length (L) = 223 steps, consis tency index (CI) = 0.4 and retention index (RI) = 0.8.The strict consensus of seventy-two trees is given in Fig. 6.
This consensus shows a main monophyletic resolved group which comprises most species of Amphimallon.The type-species of Amphimallon Berthold, 1827, Scarabaeus solstitialis L., 1758, belongs to this group.This group can be therefore considered as Amphimallon s. str.Unresolved phylogenetic relationships between this group, Rhizotrogus species, and seven species previously classified within Amphimallon (A. amphibolum and six species forming a monophyletic group called A. scutellare-group) are observed.This basal polytomy is the consensus of trees which show two main patterns.In the first pattern (Fig. 7), Rhizotrogus-species + A. scutellare-group + A. amphibolum form a monophyletic group presenting sister-group relationship with Amphi mallon.In the second pattern (Fig. 8), Rhizotrogusspecies + A. scutellare-group + A. amphibolum form a paraphyletic group.In both patterns, the A. scutellaregroup and A. amphibolum never occur within the Amphi mallon s. str.These species have 9-segmented antennae like Amphimallon, but also have long and well developed sclerotized lateral apophyses in endophallus, like Rhizo trogus.The species belonging to the A. scutellare-group were sometimes placed in the particular subgenus Amadotrogus Reitter, 1902.The present analysis does not help to place these taxa, and they are here removed from Amphimallon and are temporarily considered species incertae sedis.

Redescription of the genus Amphimallon Berthold, 1827
Antennae 9-, exceptionally 8-segmented (character 8).The club, longer in the male than in the female, composed of three segments.Basal margin of pronotum enlarged or thin, but never lacking.Meso-and metasternum densely clothed with long pale hair-like setae.Inner edge of male protibiae tridentate, exceptionally with only one or two teeth; protibiae of females tridentate.Claws equal, outer side simple, ventral edges generally with a basal tooth.Sclerotized apophyses of endophallus feebly developed, thin and shorter than one third of the length of endophal lus, sometimes absent (character 61).Basis of endo phallus dorsally with a dense area of spicules which may or may not expand basally along the sides (character 63).Species of moderate size: 10-18 mm.Basic colour yellow-brown, redd-brown or totally brown or black, pro notum often darker than elytra, rarely paler.

Groups of Amphimallon
Three lineages, based on the main clades in Fig. 6, can be recognized.Each lineage is composed of groups named from the species first described in the group.The species composition of each group is given and the main characters defining each group are presented.Unambi guous character changes are presented in Appendix 3. New taxonomic and nomenclatural considerations are added to this presentation when necessary.the "pini-group" introduced by Reitter (1902) and rede fined by Baraud (1967Baraud ( , 1992)).A seventh species from Greece, A. maniense sp.n., is added, which branches off in basal position.
The monophyly of this group is supported by six synapomorphies: odd elytral interstriae strongly elevated (character 35), but the fifth not so strongly in A. maniense; odd elytral interstriae with fine transversal ridges (character 36) which generally conceal the punctation (in A. maniense, these ridges are sparsely distributed and do not conceal the punctation); frons of male without clear carina (character 6); disc of abdominal sterna, except the last one, darkened (character 39); characteristic colour pattern of head (character 5) (except in A. nigrum which is a totally black species), with the clypeus yellowbrown and the cephalic capsule brown or black (in A. maniense, this colouration is not so contrasted due to the cephalic capsule feebly darkened); pronotum with a median furrow present at least in the posterior part (char acter 23).
The clade A. gianfranceschii-nigrum-pini-pygiale--vitalei-vivesi is supported by four synapomorphies: pro notum with a very fine punctation without setae (char acter 26) in addition to the fine, dense and regular main punctation (character 25); second antennal club segment with only its dorsal margin smooth, the ventral margin with area of sensilla (character 11); disc of pronotum black or dark brown, presenting an obvious contrast with the yellowish sides, except in A. nigrum (character 31) (in A. maniense, the pronotal disc has only confused dark ened flecks).
The clade A. gianfranceschii-nigrum-pygiale-vitalei--vivesi is supported by the shape of the antennal club which is strongly elongated and curved before apex (char acter 9).The sensillar area of inner side of first antennal club segment is prolonged and developed on the outer side basis (10).
Amphimallon vernale (Brulle, 1832) is usually consid ered a synonym of A. caucasicum (Gyllenhal, 1817) (Reitter, 1902;Medvedev, 1951;Baraud, 1992).The comparison of the types of Melolontha caucasica Gyllen hal, 1817, preserved in Stockholm (NHRS), and of Rhizo trogus vernalis Brulle, 1832, preserved in Paris (MNHN), shows that the synonymy between both names is false.The type of Melolontha caucasica does not correspond to the species named A. caucasicum by the authors.In addi tion, this taxon does not belong to Amphimallon, but to Miltotrogus Reitter, 1902 by the lack of the basal pronotal margin and by the shape of the aedeagus.
Amphimallon leuthneri Brenske, 1902 was originally described as a variety of A. caucasicum (Gyllenhal, 1817).It was raised to the rank of species by Medvedev (1951).This conclusion is confirmed by sister-group rela tionship between this taxon and A. adanense + A. nigripenne.
This group is defined by four synapomorphies: clypeus strongly raised in the corners and apparently bilobate (character 3), but this character is not constant in A. ver nale; lateral edges of pronotum clearly raised in anterior part (character 20); tridentate external edge of female pro tibiae strongly enlarged (character 45); apex of parameres, in lateral view, with a curved tooth, salient towards ventral side of parameres (character 55), except in A. fissiceps which presents a blunted apex.
The monophyly of the clade A. fissiceps-adanenseleuthneri-nigripenne is supported by two synapo morphies: apex of parameres in lateral view regularly curved (character 54); two paramedian lines of short and dense setae on dorsal side of labium lacking (character 13).
The monophyly of the clade A. adanense-leuthnerinigripenne is strongly supported by five synapomorphies: lateral edges of pronotum not crenellated in front part (character 21), due to the lack of punctation which usu ally bears hair-like setae on the sides of pronotum; disc of pronotum with short and inclined setae (character 28); punctation of pronotum regular, not very dense, fine (character 25) but quite stronger in A. adanense; disc of pronotum with a median furrow in posterior part (char acter 23); endophallus with extremely reduced ventral median area of spicules (character 63).
Sister-group relationship between A. pini-group and A. vernale-group is supported by three synapomorphies: a wide ventro-median sub-apical non-sclerotized area of paramere (character 58); basal margin of pronotum enlarged (character 22, this character is not constant in all the species of this group and can be occasionally present in species which do not belong to it); external edge of protibiae tridentate, the median tooth close to the basal (character 44).A reversal of this character is observed in the clades A. adanense-nigripenne and A. nigrumpygiale-vivesi.In A. pini, the male protibia is unidentate.
Amphimallon javeti Stierlin is usually considered the Sicilian subspecies of A. solstitiale (L.).Its colour pattern and the lack of sub-apical spicules on the dorsal side of parameres, which are present in A. solstitiale, A. ochraceum and A. alatavicum, allow us to raise it to the species rank.
The monophyly of this group is supported by two synapomorphies: apex of parameres in lateral view straight and elongated (character 54); apex of parameres in dorsal view with an internal concavity (character 56).
altaicum, ochraceum and solstitiale form a monophyletic group which is supported by three synapomorphies: sides of pronotal disc with irregular inclined short setae which often form a conspicuous white fleck (character 29); elytral disc with long sparse setae (character 37), shorter in volgense, lacking in posterior part in A. ochraceum; pygidium bearing very long and raised setae (character 42) which are generally absent in A. ochraceum.
The clade A. altaicum-ochraceum-solstitiale is sup ported by a single synapomorphy: dorsal side of parameres with a subapical area of spicules (character 57).
Sister-group relationship between this group and A. pini-group + A. vernale-group is supported by a single synapomorphy: punctation of the outer side of mandible fine and superficial, hardly perceptible (character 15), but a reversal is observed in A. fissiceps.
The monophyly of this group is supported by a single synapomorphy: posterior margin of first visible sternum medially V-shaped (character 38).
Sister-group relationship between this group and A. pini-group + A. vernale-group + A. solstitale-group is supported by three synapomorphies: metasternum with robust spinose setae in addition to the hair-like setae (character 33); second antennal club segment with ventral and dorsal margin smooth, both without sensillar area (character 11); frons of male with a strong transverse carina which reaches the sides of the head (character 6).However, reversals for this last two characters are observed in the A. pini-group.

The A. peropacum-group
This group includes a single species from Portugal, A. peropacum Reitter, 1911, which is one of the most recog nizable species of Amphimallon.In addition to the char acters treated in this phylogenetic analysis, this species is characterized by many other autapomorphies, and in par ticular the shape of its parameres, the raspy punctation of its pronotum and the rough surface of its tegument.
Sister-group relationship between this species and other groups composing the A. solstitiale-lineage is supported by two synapomorphies: inner spur of protibiae clearly inserted between median and apical teeth (character 47); ventro-median sub-apical non sclerotized area of paramere narrow (character 58), but a reversal is observed in the A. pini-group +A.vernale-group.

The A. fuscum-lineage
This lineage is composed of two monophyletic groups and presents sister-group relationships with the A. solstitiale-lineage.
Amphimallon furvum (Germar, 1817) was considered a simple variety, and recently (Miksic, 1970) a subspecies of Amphimallon fuscum (Scopoli, 1786).Its particular elytral colour pattern and the presence of spicules on the dorsal side of parameres allow us to raise this taxon to the species rank.
Amphimallon jeannei (Baraud, 1971) comb.n. was originally described in Monotropus Erichson, 1848 on the basis of its 8-segmented antennae.In fact, this species finds a place in the A. fuscum-group of Amphimallon.
This group is defined by a single synapomorphy: pro notum and elytra have a distinctive tint (character 32).The pronotum is dark reddish brown and elytra reddish brown, but the monophyletic clade A. jeannae-theryi pre sents pale reddish brown elytra (convergence with maevae) and the clade A. brucki-furvum-jeannei presents yellowish-brown elytra (convergence with A. ruficorne, A. evorense and A. arianae).
In addition to its particular colour pattern, the mono phyletic group A. jeannae-theryi is defined by two syna pomorphies: edge of clypeus strongly lifted all around, the clypeus deeply concave (character 3); second antennal club segment with ventral and dorsal margin smooth, both without sensillar area (character 11).
The monophyly of the clade A. brucki-furvum-jeannei is supported by a single synapomorphy in addition to the colour pattern: dorsal side of parameres with a subapical area of spicules (character 57) (convergence with a part of A. solstitiale-group), however, only traces of spicules are observed in A. jeannei.
Type comparison and the study of many specimens allows to establish the new synonymy between A. litigiosum Fairmaire, 1860 and A. galleti Baraud, 1970.This group is supported by a single synapomorphy: clypeus enlarged (character 2).
The clade A. safiense-litigiosum-insculptum is defined by the superior spur of metatibiae enlarged before apex (character 48) and by the elytra clearly laterally enlarged in the front part (character 34).
The clade A. menori-naceyroi is supported by the frons of the male with a strong transverse carina (character 6).
Sister-group relationship between A. fuscum-group and A. naceyroi-group is supported by two synapomorphies: absence of the two paramedian lines of short and dense setae on the dorsal side of labium (character 13), with a reversal in a part of A. fuscum-group; endosymbiotic crypts in mesenteron absent in the male (character 51), also with a reversal in a part o f A. fuscum-group.Sister-group relationship between the A. fuscum-lineage and the A. solstitiale-lineage is supported by two synapomorphies: frons of the female with a strong transverse carina (character 7); tegument of pronotum microreticu lated (character 24), with a reversal in A. circumligatusvulpecula (A.naceyroi-group) and A. maniense (A.pini-group).

The A. ruficorne-lineage
This lineage is composed of two monophyletic groups and presents sister-group relationships with the two pre vious lineages.
The monophyly of this group is supported by three synapomorphies: pygidium with short but clearly percep tible regularly distributed raised setae (character 42), shorter in A. ruficorne; elytral disc with short regularly distributed raised setae (character 37); pronotal disc with short setae from fine punctuation (29).
The monophyly of the clade A. assimile-burmeisterimajale-pseudomajale is supported by four synapomorphies: pronotum with a very fine and dense punctation, bearing short inclined setae (character 25) which give the pronotum a silky aspect; pronotum transverse (character 17); narrow prosternal basisternum with a transversal furrow (character 30); ventral edge of second tarsomere of protarsus clearly and strongly dentate (character 49).The monophyly of the clade A. majale-pseudomajale is supported by the absence of main punctation of pronotal disc (character 27).The pronotal disc presents only a very fine and dense punctation bearing short inclined setae.Hair-like setae are absent from the disc because of the lack of main punctation.
The description of Amphimallon trisinuatum Reitter, 1902 was based on a single female from Portugal, Sierra da Estrella.This species has never been recaptured since its description.The authors who have studied the Iberian fauna (Baguena, 1959(Baguena, , 1967;;Baraud, 1977b;Martin Piera, 1985) did not know this species, which was even called "enigmatic" by Baraud (1992).I have studied the type of this name which is preserved in Budapest (HNHM).The lack of setae at the basis of pronotum, con sidered by Reitter as a specific character, is due to the poor condition of the specimen.In fact, this specimen is barely different from Amphimallon seidlitzi Brenske, 1891, a species common in Sierra da Estrella, by the shape of the clypeus, which is trisinuate.I have also met this particular clypeal shape in a specimen of Amphi mallon majale (Razoumowsky, 1789) from Montpellier, France (MNHN, A. Mackanga leg).Other specimens, like the type-specimen of Amphimallon litigiosum Fairmaire, 1860 (MNHN), or a specimen of Amphimallon solstitiale pictum Kraatz, 1902 from Greece (MNHN), present only a single lateral sinuosity which gives the clypeus an asymmetrical shape.These specimens may be teratologic cases (e.g., due to defective maturation).
The monophyly of this group is supported by four synapomorphies: pronotum with a very fine superficial punc tation without setae between main punctation (character 26) (convergence with a part of A. pini-group); narrow ventro-median sub-apical non-sclerotized area of parameres (character 58); dorsal basis of endophallus with dense area of spicules which does not expand basally on the sides (character 63); ventro-median spiculate area of endophallus clearly delimited, its anterior part M-shaped (character 64).
The monophyly of the clade A. cantabricum-sainziiseidlitzi-lusitanicum-roris is supported by two synapomorphies: posterior part of lateral edge of pronotum sinuate (character 19) (convergence with a part of A. rujicorne-group and A. solstitiale-group), reversed in roris; parameres, in lateral view, thick (character 53) (convergence with a part of A. solstitiale-group).
The clade A. sainzi-seidlitzi-lusitanicum-roris is def ined by two synapomorphies: hair-like setae of pronotal disc lacking at least from the front part and on the sides (character 27); parameres in dorsal view clearly inflated (character 52).
The clade A. seidlitzi-lusitanicum-roris is supported by three synapomorphies: apical segment of labial palpas strongly inflated (character 16); narrow prosternal basisternum, with a transversal furrow (character 30) (conver gence with a part of the A. rujicorne-group); ventral edge of second tarsomere of protarsus clearly and strongly den tate (character 49) (convergence with a part of A. rujicorne-group).
The clade A. lusitanicum-roris is defined by the strongly enlarged head (character 1).
Sister-group relationship between A. lusitanicum-group and A. rujicorne-group is supported by a single synapomorphy: pygidium with a regular, dense and strong punc tation (character 41).

DESCRIPTIONS OF NEW SPECIES
Amphimallon adanense sp.n.
Description.Length: 13-15 mm.Reddish brown head, antennae, mouth-parts and legs yellow-brown.Disc of pronotum pale reddish brown, with a median line and symmetrical flecks dark reddish brown, sides yellowbrown.Dark reddish brown elytra.Last sternum yellowbrown, disc of other sterna pale reddish brown.Yellowbrown pygidium, with a reddish brown median fleck.
Head (Fig. 15).Frons hairy, armed with a strongly ele vated transverse carina, interrupted in the middle, stronger in the female.Edge of clypeus strongly lifted and sinuated in the middle.Antennae 9-segmented, club shorter than the other segments together.
Pronotum.Sides rounded, in front part, with a thin, uncrenellate and lifted edge.Posterior part of disc with a feeble, but clearly perceptible, median furrow.Surface of tegument microreticulated.Punctation dense, punctures spaced by 1-1.5 times their diameter.Clothed with short, inclined setae, hardly perceptible.Basal margin enlarged, but thinner scutellum.
Aedeagus (Figs 9-10).Parameres regularly curved before apex in lateral view.Apex with a small curved tooth toward ventral side.Endophallus as in Fig. 11.
Etymology.Species named after its occurrence in Adana.
Remarks.Amphimallon adanense sp.n. belongs to a natural group which includes also two species described from the same locality in Turkey, Adana: Amphimallon nigripenne Reitter, 1902 and Amphimallon leuthneri Reit ter, 1902.Amphimallon adanense sp.n. differs from both species by the stronger punctation of pronotum and elytra,  which seems rough, by its colour pattern, and by its smaller size.Amphimallon safiense sp.n.
Head (Fig. 16).Clypeus enlarged, its edge weakly raised.Clypeus and frons with fine and very dense punctation which bears long hair-like setae.Frons without carina.Antennae 9-segmented, club shorter than the other segments together.
Pronotum.Surface of tegument smooth or scarcely microreticulated.Punctation fine and dense, irregular with very long hair-like setae.Basal margin thin.Edge regularly curved.
Surface clothed with very short hardly visible setae on disc, clearly longer around scutellum.
Pygidium.Punctuation superficial, regularly spaced.Surface clothed with hardly perceptible short and inclined setae.
Etymology.Species named after its occurrence in Safi.
Remarks.Amphimallon safiense sp.n. is close to Amphimallon litigiosum Fairmaire, 1860, with which it was until now confused.Both species can be separated by the punctation, by the pilosity of the head and pronotum, and by the shape of endophallus which is very particular in A. safiense sp.n. (Fig. 14).
Head.Clypeus wide, its edge raised, straight or feebly sinuate in middle.Frons without transverse carina.Antennae 9-segmented, club shorter than the other seg ments together.
Pronotum.Posterior part with a feeble median furrow.Surface of tegument smooth or scarcely microreticulated.Punctation double, large points mixed with smaller points, regularly distributed and dense, spaced by less than one diameter.Hair-like setae are short but regularly distributed and clearly perceptible.Basal margin enlarged, but thinner in front of scutellum.
Etymology.Species named after its occurrence in Máni.
Remarks.The specimens were collected during the evening flight, above the ground level of a grassland.Amphimallonjeannae sp.n.
Head (Fig. 21).Frons hairy, armed with two parame dian transverse elevations on a central gibbosity which do not reach.Semicircular clypeus, with regularly lifted bor ders.Antennae 9-segmented, club shorter than the other segments together.
Pronotum.Convex and transverse, with barely angular, curved sides.Basal margin thin.Surface of tegument microreticulated.Punctation dense, double: size of punc tures is irregular.Punctures spaced by one diameter.Sur face clothed with long golden hair-like setae.
Elytra.1st and 3rd interstriae feebly elevated.Puncta tion fine and dense, odd interstriae more sparsely punc tured.Surface clothed with very short setae on disc, clearly longer around scutellum.
Pygidium.With a superficial punctation, punctures regularly spaced.Clothed with very short and inclined setae, hardly perceptible.
Etymology.Species named after Jeanne Charbonnel for her important work in the Melolonthidae collections in MNHN.
Remarks.This species is close to Amphimallon theryi Peyerimhoff, 1949 with which it was until now confused.Both species could be separated by the characters in Table 2. CONCLUSION Phylogenetic analysis of the genus Amphimallon Berthold based on a study of morphological characters enabled systematic conclusions, in particular the rede scription of the Amphimallon s. str.and the definition of the main groups of this genus.Position of seven species previously included in Amphimallon is questioned.Phylo genetic hypothesis for a group allows also to propose evolutionary scenarios in biogeography and biology, and this is intended for Amphimallon in forthcoming papers.Appendix 1. Synonymical list of taxa belonging to Amphi mallon Berthold, 1827 sensu auctorum.Names of species not considered in the phylogenetic analysis are marked with *.Spe cies excluded from Amphimallon s.str.are in square brackets.Amphimallon adanense sp.n.Amphimallon alatavicum Medvedev, 1951 [Amphimallon altifrons Baraud, 1971

Table 1 .
Data matrix for sixty-one species and sixty-five characters used in the cladistic analysis.See character coding explana tions in text.