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Chaetocnema Stephens and related genera (Coleoptera, Chrysomelidae,Alticinae): A systematic revision, with description of a new genus from theIndian Ocean (Seychelles)Maurizio Biondi a

a Dipartimento di Scienze Ambientali, University of L'Aquila, Coppito, L'Aquila, Italy

To cite this Article Biondi, Maurizio(2002) 'Chaetocnema Stephens and related genera (Coleoptera, Chrysomelidae,Alticinae): A systematic revision, with description of a new genus from the Indian Ocean (Seychelles)', Italian Journal ofZoology, 69: 4, 355 — 366To link to this Article: DOI: 10.1080/11250000209356481URL: http://dx.doi.org/10.1080/11250000209356481

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Ital. J. Zool., 69. 355-366 (2002)

Chaetocnema Stephens and relatedgenera (Coleoptera, Chrysomelidae,Alticinae): a systematic revision, withdescription of a new genus from theIndian Ocean (Seychelles)

MAURIZIO BIONDIDipartimento di Scienze Ambientali, University of L'Aquila,I-67010 Coppito, L'Aquila (Italy)E-mail: bicmdi@univaq.it

ABSTRACT

Chaetocnema Stephens and its related flea beetle genera Terpno-chlorus Fairmaire, Carcharodis Weise, Pepila Weise, CollartalticaBechyné, Biodontocnema Biondi, and Seychellaltica n. gen. areconsidered for a taxonomic analysis. Scanning electron micro-graphs of particular morphological aspects of these genera are in-cluded. The new genus Seychellaltica from Seychelles is de-scribed. The following species are attributed to it: Seychellalticamahensis (Maulik) (type species), S. krishna Maulik, and two newspecies S. gardineri sp. n. and S. praslinensis sp. n. Line drawingsof male and female genitalia of the Seychellaltica species are illus-trated. A cladistic analysis of this taxonomic group using a parsi-mony method is presented. Chaetocnema shows most affinitywith the Afrotropical genera Carcharodis, Collartaltica andBiodontocnema, while Pepila, the new genus Seychellaltica, andabove all Terpnochlorus, appear to have originated in earliertimes.

KEY WORDS: New genus - New species - Chrysomelidae Altici-nae - Chaetocnema - Terpnochlorus - Seychellaltica n. gen. -Biodontocnema - Carcharodis - Pepila - Collartaltica - Cladisticanalysis.

ACKNOWLEDGEMENTS

I am very grateful to colleagues who enabled me to study thevaluable material preserved in their respective institutions: C. Bel-lamy (TMSA), N. Berti (MNHN), M. Cludts (ISNB), MargieCochrane (SAMC), M. De Meyer (MRAC), E. Matthews (SAM), G.A. Samuelson (BPBM); S. L. Shute (BMNH) and B. Viklund(NHRS). This research was supported by a grant from the "Mini-stero dell'Universita e della Ricerca Tecnologica e Scientifica"(M.U.R.S.T. ex 40%).

INTRODUCTION

The flea beetle genus Chaetocnema Stephens, 1831,with over 300 species described to date from all thezoogeographical regions, is one of the most representa-tive genera of the whole chrysomelid subfamily Altici-nae. Other genera, closely related to Chaetocnema andoften confused with it, have been described mainlyfrom the Afrotropical and Australian Regions.

The aim of the present paper is to clarify the taxono-my of this group of genera including: ChaetocnemaStephens, 1831, Terpnochlorus Fairmaire, 1904, Car-charodis Weise, 1910, Pepila Weise, 1923, CollartalticaBechyne, 1959, Biodontocnema Biondi, 2000 and Sey-chellaltica n. gen. From the morphological point ofview, this taxonomic group is mainly characterized byhaving: (1) middle and hind tibiae both with ciliate den-tate dorsal emargination; (2) 1st and 2nd abdominal ster-nites joined. The taxa here considered also share thefollowing character-states: (1) closed procoxal cavities;(2) simple apical spur of the hind tibiae; (3) simpletarsal claws; (4) elytral punctation at least partiallyarranged in regular rows.

Other genera, considered by previous Authors as close-ly related to Chaetocnema, have been excluded from thisanalysis on the basis of the following considerations:

- Crepicnema Scherer, 1969: for having middle tibiaewithout ciliate dentate emargination; 1st and 2nd ab-dominal sternites separate, not joined; pronotum withevident antebasal transverse impression; pronotalbase strongly sinuous;

- Xenidea Baly, 1862: for having middle tibiae without ci-liate dentate emargination; hind tibiae only with a weaklateral emargination; 1st and 2nd abdominal sternitesseparate, not joined [differently from the opinion of Cha-puis (1875: 47), who reported "Abdomen a 1 et 2 seg-ments soudes"]. On the basis of the metafemoral springmorphology this Australian genus is included in thePhyllotreta morpho-group by Furth & Suzuki (1998: 97).

Dorsal ciliate dentate emarginations on the hind tibiaeare more or less evident also in other flea beetle generabelonging to other groups, such as: (1) Podontia Dal-man, 1824, Blepharida Chevrolat, 1837, Polydada Che-vrolat, 1842 etc., easily distinguishable by the presenceof bifid tarsal claws; (2) Euplectroscelis Crotch, 1873,with entirely confused elytral punctation and an evidentantebasal transverse pronotal impression [this genus,occurring in Mexico, is considered by Furth (1998: 5) asrelated to Blepharida]; (3) Heikertingerella Csiki, 1940(cf. Heikertinger & Csiki, 1940), with open anterior cox-al cavities and almost no punctation on dorsal integu-ment; (4) Oedyonychus Berthold, 1827, PhysonychisClark, I860, Physodactyla Chapuis, 1875, PhiloponaWeise, 1903, with open anterior coxal cavities, prono-tum subdepressed laterally widely channeled, and en-tirely confused elytral punctation.

(Received 24 January 2002 - Accepted 2 May 2002)

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356 M. BIONDI

MATERIALS AND METHODS

Studied material

The studied material consists of preserved dry insects providedby courtesy of the following institutions: The Natural History Mu-seum, London, United Kingdom (BMNH); Bernice P. Bishop Mu-seum, Honolulu, Hawaii, U.S.A. (BPBM); Institut Royal des Sci-ences Naturelles de Belgique, Brussels, Belgium (ISNB); MuseumNational d'Histoire Naturelle, Paris, France (MNHN); Musee Royalde l'Afrique Centrale, Tervuren, Belgium (MRAC); NaturhistoriskaRiksmuseet, Stockholm, Sweden (NHRS); South Australian Muse-um, Adelaide, Australia (SAM); South African Museum, CapeTown, Republic of South Africa (SAMC); Transvaal Museum, Pre-toria, Gauteng, Republic of South Africa (TMSA). The specimenswere examined using WILD MZ8 and MZ12.5 binocular micro-scopes. The scanning electron micrographs were taken by aPHILIPS SEM XL30 CP.

Phylogenetic analysis

Cladistic analysis was performed using the licensed packagePAUP 4' (Ver. 4.0b8; Swofford, 2000). The character states andcoding used in this analysis are:

1. Shape of the distal margin of the frons: (0) not medially in-cised (Fig. 10); (1) medially deeply incised (Figs 7-8).

2. Frontal sulci: (0) long, present around the eyes (Figs 7, 10);(1) very short, absent around the eyes (Figs 13-15).

3- Width of the interantennal space: (0) at most two and a halftimes as wide as the distance between the antennal socketand the relative inner ocular margin (Figs 9, 13, 15); (1) atleast four times as wide as the distance between the antennalsocket and the relative inner ocular margin (Fig. 7).

4. Shape of the interocular space: (0) without transverse carinae(Figs 13, 14); (1) at least with an evident transverse carina(Fig. 7).

5. Position of the mouthparts: (0) not sunken (Fig. 15); (1) sunken,with exception of the labrum and the mandibles (Fig. 9).

6. Shape of the labrum: (0) without longitudinal median carina(Figs 9, 15); (1) with an evident longitudinal median carina(Fig. 8).

7. Shape of the antennae: (0) filiform; (1) subclavate, with anten-nomeres 7-11 enlarged.

8. Shape of the prothorax: (0) subcylindrical; (1) not subcylindri-cal, subdepressed.

9. Punctation of the elytra: (0) arranged in 11 (+ 1 long sutural)regular striae; (1) arranged in 9 (+ 1 short sutural) striae atleast partially regular.

10. Shape of the prosternum: (0) anteriorly moderately convex,not able to receive mouthparts in resting position; (1) anteri-orly strongly convex, able partially to receive mouthparts inresting position (Figs 10-12).

11. Ratio of metastemal width/metastemal length: (0) <2.5; (1)>2.5

12. Median small hollow near the posterior metasternal margin:(0) present (Fig. 16); (1) absent.

13. Shape of the hind apical tibial socket: (0) unmodified (Figs17, 19); (1) very wide, spoon-shaped (Fig. 18).

14. Shape of the 1st tarsomere of the pro- and mesotarsi: (0) sym-metric; (1) asymmetric (Fig. 20).

15. Length of the 1SI hind tarsomere (a): (0) shorter than 2nd; (1)longer than 2nd.

16. Length of the 1st hind tarsomere (b): (0) longer than 2nd butshorter than twice the length of the 2nd and 3 rd together (Figs17, 18, 21); (1) longer than twice the length of the 2nd and 3rd

together (Fig. 19).17. Shape of the 1st hind tarsomere: (0) subrectangular; (1) sub-

triangular.18. Shape of the apical tarsal segment of the hind legs: (0) not

globose (Figs 17, 18); (1) more or less globose (Fig. 21).

19. Apical margin of the hind tibiae: (0) with short setae; (1) den-ticulate.

20. Hind tibiae: (0) dorsally not channelled in the proximal half,with inner margin not visible (Fig. 17); (1) dorsally channelledin the proximal half, with evident inner margin (Fig. 19).

21. Lateral emargination of the hind tibiae: (0) unidentate (Fig.17); (1) bidentate, dorsally with both inner and outer marginciliate [type Seychellaltica] (Fig. 19); (2) bidentate, dorsallywith only outer margin ciliate [type Biodontocnema] (Fig. 18).

Abbreviations used: LAED, length of median lobe of aedea-gus; LAN, length of antennae; LB, total length of body; LE, lengthof elytra; LP, length of pronotum; LSP, length of spermatheca; WE,width of elytra; WP, width of pronotum.

TAXONOMIC ACCOUNTS

Genus Biodontocnema Biondi(Figs 2, 18)

Biodontocnema Biondi, 2000: 347

Type species - Biodontocnema brunnea Biondi, 2000by monotypy (type in SAMC examined).

Distribution - South-West Africa.

Remarks - Only one species is known for this genusfrom the Republic of South Africa (B. brunnea Biondi,2000). No information about host plants is available forthis genus.

Genus Carcharodis Weise(Figs 1, 7, 8)

Chaetocnema subg. Carcharodis Weise, 1910: 434Carcharodis Weise (upgraded to genus by Bechyne, 1954: 683)

Type species - Chaetocnema rugiceps Baly, 1877 bysubsequent designation (Bechyne, 1954: 683) (type inBMNH examined).

Distribution - Sub-Saharian Africa, Madagascar.

Remarks — Carcharodis was considered by some Au-thors as a synonym of Chaetocnema (cf. Seeno & Wilcox,1982: 139). On the contrary, this African genus showsmany characteristics (see the key below) that allow Car-charodis to be considered as a separate genus. At themoment, Carcharodis includes the following four species:C. congoana (Weise, 1916); C. durbanensis Bechyne,1954; C. malvernensis Bechyne, 1954, from ContinentalAfrica and C. rugiceps (Baly, 1877) from Madagascar. Noinformation about host plants is available for this genus.

Genus Chaetocnema Stephens(Fig. 17)

Chaetocnema Stephens, 1831: 325

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CHAETOCNEMA AND RELATED GENERA 357

1 mm

\

Figs 1-6 - Habitus. 1 - Carcharodis rugiceps (Baly). 2 - Biodontocnema brunnea Biondi. 3 - Collartaltica tenebrosa (Laboissiere). 4Pepila megalopoides (Baly). 5 - Seychellaltica mabensis (Maulik). 6 - Terpnochlorusperrieri Fairmaire.

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358 M. BIONDI

Odontocnema Stephens, 1831: 285 (nomen nudum)Plectroscelis Chevrolat, 1837: 417 (synonymized by Weise, 1886: 750)Udorpes Motschulsky, 1845: 107 (synonymized by Heikertinger &Csiki, 1940: 375)Tlanoma Motschulsky, 1845: 108 (synonymized by White, 1996: 20)Ydorpes Motschulsky, 1845: 107 (synonymized by White, 1996: 20)Hydropus Motschulsky, I860: 235 (synonymized by Heikertinger& Csiki, 1940: 375)Exorhina Weise, 1886: 750 (synonymized by Heikertinger & Csiki,1940: 376)Brinckaltica Bechyne, 1959a: 237 (synonymized by Scherer, 1961:259)

Type species - Chrysomela concinna Marsham, 1802by subsequent designation (Westwood, 1838: 42) (loca-tion of the type unknown).

Distribution - All the zoogeographical regions.

Remarks - Within this genera-group here discussed,Chaetocnema is the most widespread genus. It consistsof over 300 described species, of which at least 83 oc-cur in the Palaearctic Region (Gruev & Doberl, 1997),59 in the Nearctic Region (White, 1996), 80 in theNeotropical Region (Scherer, 1983), 99 in the Afrotropi-cal Region (Biondi, in press), and an undeterminednumber in the Oriental and Australian Regions. The di-agnostic characters used to separate the genera andsubgenera Exorhina, Tlanoma and Brinckaltica are notconstant and/or taxonomically significant. Many botan-ical families include host plants for this genus, particu-larly Chenopodiaceae, Polygonaceae, Cyperaceae, Gra-minae (Jolivet & Hawkeswood, 1995).

Genus Collartaltica Bechyne(Figs 3, 9-12)

Collartaltica Bechyne, 1959b: 27

Type species — Collartaltica cryptostoma Bechyne,1959 (type in ISNB not traced).

Distribution - Central and southern Africa.

Remarks — Three species belong, at present, to thisgenus (Biondi, 2002): C. cryptostoma Bechyne, 1959; C.nigrolucens (Bechyne, I960) (type in MRAC examined);C. tenebrosa (Laboissiere, 1942) (type in MRAC exam-ined) (Biondi, in press). No information about hostplants is available for this genus.

Genus Pepila Weise(Figs 4, 21)

Pepila Weise, 1923: 127

Type species - Chaetocnema megalopoides Baly, 1877by original designation (type in BMNH examined).

Distribution - Australia.

Remarks - At the moment, this Australian genus in-cludes the only species P. megalopoides (Baly). Howev-er, other species originally attributed to the genusChaetocnema Stephens, such as C. fuscomaculata Baly,1877, C. submetallescens Baly, 1877, C. crassipennis(Blackburn, 1896), C. pygmaea (Waterhouse, 1838) etc.and other new taxa are to attribute to Pepila Weise(Biondi & D'Alessandro, unpubl. data). Some species ofthis flea beetle genus are associated with plants of Euca-lyptus and Melaleuca (fam. Myrtaceae) (pers. data).

Genus Seychettaltica n. gen.(Figs 5, 14-16, 19-20, 22-30)

Type species- Chaetocnema mahensis Maulik, 1931.

Diagnosis — This new genus shows some similaritieswith Pepila Weise, with which it shares the channelledhind tibiae and the very short frontal sulci (this lastcharacter-state is present also in Terpnochlorus Fair-maire). However, Seycahellaltica gen. n. is distinguish-able from Pepila and the other genera here considered,mainly by having: hind tibiae with a ciliate dentateemargination both on inner and outer margin (Fig. 19);metasternum very elongate (Fig. 16); 1st tarsomere ofthe pro- and mesotarsi distinctly asymmetric (Fig. 20).

Description - Dorsal integument from pale to darkbrown, often with yellowish areas on the elytra. Headwith wide frons; vertex and interocular space uniformly,but weakly punctate. Frontal sulci very short, absentaround the eyes; internantennal space convex, about aswide as length of 1st antennal segment; eyes large, subel-liptical; antennae little longer than half body length.

Pronotum transverse (Fig. 14), laterally straight orweakly rounded, convergent towards base; anterior an-gles distinctly slanted; lateral and basal margin clearlybordered; punctation sparse, formed by punctures ofdifferent size, impressed on clearly microreticulate orsmooth surface.

Elytra laterally subparallel or slightly rounded, com-pletely covering pygidium; punctation arranged in 9(+1 sutural) regular striae; interstriae almost flat, sub-convex or subcarinate; humeral callus distinct; wingsmacropterous; scutellum semicircular or subtriangular,with finely shagreened surface.

Prosternum impunctate; mesosternum wide, distinct;metasternum very elongate, with an evident median de-pression near posterior margin (Fig. 16); abdominal sternalsegments impunctate, with sparse yellowish pubescence.

Middle tibiae with simple dentate ciliate emargination;hind tibiae dorsally channelled, with ciliate dentateemargination both on inner and outer margin (Fig. 19);1st tarsomere of the pro- and mesotarsi distinctly asym-metric; 1st posterior tarsomere longer than twice thelength of 2nd and 3 rd together; 3rd anterior and middletarsomere strongly dilated, medially not incised; hind

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CHAETOCNEMA AND RELATED GENERA 359

Figs 7-14 - Scanning electron micrographs. 7 - Head of Carcharodis malvernensis Bechyne (X 86). 8 - labrum of C. malvernensis (x 200).9 - Head and mouthparts of Collartaltica tenebrosa (Laboissiere) (X 148). 10 - Head and prosternum of C. tenebrosa (x 122). 11 -Prostemum of C. tenebrosa (x97). 12 - Prosternum of C. tenebrosa (X118). 13 - Head of Pepila megalopoides (Baly) (x99). 14 - Headof Seychellaltica krishna (Maulik) (x 100).

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360 M. BIONDI

femora moderately enlarged; metafemoral spring (Fig.22) attributable to Chaetocnema morpho-group (Furth& Suzuki, 1998: 97), with extended arm of dorsal lobeshort and depressed, ventral lobe with no recurvedflange, dorsal edge straight and strongly angled downapically, basal edge flat.

Etymology - The name of this new genus is from theSeychelles, islands where this genus lives, and Altica,nominate genus of the subfamily Alticinae.

Distribution — Indian Ocean: Seychelles.

Remarks — Four species from the Seychelles belong tothis new genus: Seychellaltica mahensis (Maulik, 193D,S. krishna (Maulik, 1931), Seychellaltica gardineri sp.n., Seychellaltica praslinensis sp. n.

Seychellaltica mahensis (Maulik) n. comb.(Figs 5, 15-16, 23, 27)

Chaetocnema mahensis Maulik, 1931: 250 (partim)

Material examined - Lectotype 9 (M. Biondi des.;specimen figured by Maulik, 1931: 250, fig. 7) and fiveparalectotypes (two &($ and three 99), SEYCHELLES:Mahe, 1908-9, Seychelles Exp., Percy Sladen Trust Expe-dition, B.M. 1927-444 (BMNH). Lectotype designatedbecause the description of this species was based onseveral syntypes and the type series was mixed (5. ma-hensis and S. gardineri n. sp.).

Distribution — Indian Ocean: Seychelles (Mahe).

Morphometrics (mean ± SD) - Male: LB = 2.16 ± 0.23mm; LAN = 1.61 ± 0.06 mm; LP = 0.41 ± 0.04 mm; WP =0.62 ± 0.07 mm; LE = 1.54 ± 0.16 mm; WE = 0.86 ± 0.10mm; LAED = 0.74 ± 0.01 mm; LAN/LB = 0.74 ± 0.05;WP/LP = 1.51 ± 0.01; LE/LP = 3.76 ± 0.01; LE/LAED =2.24 + 0.08.

Female: LB = 2.29 + 0.10 mm; LAN = 1.56 ± 0.09 mm;LP = 0.42 ± 0.01 mm; WP = 0.68 ± 0.03 mm; LE = 1.63 ±0.06 mm; WE = 0.91 ± 0.04 mm; LSP = 0.22 ± 0.02 mm;LAN/LB = 0.69 ± 0.07; WP/LP = 1.58 ± 0.03; LE/LP =3.81 ± 0.05; LE/LSP = 7.73 ± 0.12.

Morphological remarks— Median lobe of aedeagus (Fig.23) slender with smooth surface; laterally wider at mid-dle; apical part sub-rounded with a median small tooth;ventral sulcus wide, distinct, interrupted about middle;dorsal sulcus wide, but indistinct; median lobe in lateralview medially strongly curved; apical part ventrallycurved. Spermatheca (Fig. 27) comparatively large, withbasal and distal parts elongate; ductus arcuate, uncoiled.

Ecological notes - Concerning the ecology of thisspecies, Maulik (1931: 251) reported: "evidently a

species characteristic of the endemic forests" (altitude500-2000 feet).

Seychellaltica krishna (Maulik) n. comb.(Figs 14, 19-20, 24, 28)

Chaetocnema krishna Maulik, 1931: 251 (partim)

Material examined - Lectotype C? (M. Biondi des.)and six paralectotypes (two <S(S and four 99), SEY-CHELLES: Mahe, 1908-9, Seychelles Exp., Percy SladenTrust Expedition, B.M. 1927-444 (BMNH). Lectotypedesignated because the description of this species wasbased on several syntypes and the type series wasmixed (5. krishna, S. gardineri n. sp. and S. praslinensisn. sp.).

Distribution - Indian Ocean: Seychelles (Mahe).

Morphometrics - Male: LB = 2.33 ± 0.07 mm; LAN =1.70 ± 0.12 mm; LP = 0.48 ± 0.03 mm; WP = 0.74 ± 0.04mm; LE =1.64 + 0.09 mm; WE = 1.05 ± 0.06 mm; LAED= 0.96 ± 0.02 mm; LAN/LB = 0.73 + 0.04; WP/LP = 1.55± 0.09; LE/LP = 3-42 ± 0.17; LE/LAED = 1.78 ± 0.04.

Female: LB = 2.48 ± 0.21 mm; LAN = 1.49 ± 0.03 mm;LP = 0.48 ± 0.06 mm; WP = 0.80 ± 0.07 mm; LE = 1.77 +0.09 mm; WE = 1.17 + 0.09 mm; LSP = 0.22 + 0.01 mm;LAN/LB = 0.63 ± 0.08; WP/LP = 1.67 + 0.06; LE/LP =3.70 + 0.26; LE/LSP = 8.63 ± 0.06.

Morphological remarks — Median lobe of aedeagus(Fig. 24) slender, comparatively large, with smooth sur-face; laterally narrower in basal half; apically lanceolatewith a median small tooth; ventral sulcus narrow, indis-tinct; dorsal sulcus present in distal third; median lobein lateral view straight in basal half and strongly curvedin distal half; apical part slightly sinuous. Spermatheca(Fig. 28) comparatively large, with basal part subglo-bose; ductus straight, uncoiled, comparatively short.

Ecological notes - Information about this species wasreported by Maulik (1931: 252): "found generally in thehigh forests from about 1000 feet to over 2000 feet".

Seychellaltica gardineri n. sp.(Figs 26, 30)

Chaetocnema mahensis Maulik, 1931: 250 (partim)Chaetocnema krishna Maulik, 1931: 251 (partim)

Type material - Holotype, <?, SEYCHELLES: Silhouette,1908, Seychelles Exp., Percy Sladen Trust Expedition, B.M.1927-444 (BMNH: ex syntype of Chaetocnema mahensisMaulik). Paratypes, one 9, same data of the holotype(BMNH: ex syntype of Chaetocnema krishna Maulik).

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CHAETOCNEMA AND RELATED GENERA 361

Figs 15-22 - Scanning electron micrographs. 15 - Head and mouthparts of Seychellaltica mahensis (Maulik) (x 182). 16 - Pro-, meso- andmetasternum of S. mahensis (x 99). 17 - Hind tibia of Chaetocnema hortensis (Geoffroy in Fourcroy, 1785) (x 157). 18 - Hind tibia ofBiodontocnema brunnea Biondi (x 161). 19 - Hind tibia of Seychellaltica krishna (Maulik) (x 131 and x 400). 20 - Tarsal segments ofanterior leg of S. krishna (x 185). 21 - Tarsal segments of hind leg of Pepila megalopoides (Baly) (x 200). 22 - Metafemoral spring in an-terior and posterior view of Seychellaltica praslinensis n. sp.

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362 M. BIONDI

26

30

Figs 23-30 - Median lobe of aedeagus in lateral, ventral and dorsalview (Figs 23-26). 23 - Seychellaltica mahensis (Maulik). 24 - S.krishna (Maulik). 25 - S. praslinensis n. sp. 26 - S. gardineri sp.n.. Spermatheca (Figs 27-30). 27 - S. mahensis (Maulik). 28 - 5. kri-shna (Maulik). 29 - S. praslinensis n. sp. 30 - S. gardineri n. sp.

Diagnosis - On the basis of external morphology, thisnew species shows some similarities with S. krishna(Maulik), while it shares with S. praslinensis n. sp. thesimilar shape of spermatheca and of median lobe ofaedeagus. Diagnostic characters for identification of thisnew species are given in the key.

Description of the holotype (cT) - LB = 2.30 mm; LAN =1.70 mm; LP = 0.48 mm; WP = 0.77 mm; LE = 1.74 mm;WE = 1.08 mm; LAED = 0.82 mm; LAN/LB = 0.74;WP/LP = 1.59; 1E/LP = 3-59; LE/LAED = 2.13.

Dorsal integument bright pale brown. Body shape elon-gate-elliptical, weakly convex. Maximum pronotal widthnear anterior angles; maximum elytral width at middle.

Head with frons and vertex densely punctate; punc-tures weakly impressed on almost smooth surface; frontalsulci very short, not visible around eyes; frontal tuberclesabsent; interantennal space as wide as length of 1st anten-

nomere; frontal carina wide and rather flat; labrum andpalpi reddish brown; antennae slightly longer than halfbody length; antennomeres 1-2 and 7-8 partially dark-ened, 3-6 yellowish; 9-11 clearly blackened; length ofeach antennal segment proportional to numerical se-quence 59:49:44:57:62:54:52:55:46:46:65 (1 = 0.003 mm).

Pronotum moderately transverse, anteriorly wider,laterally straight and convergent from anterior angles tobase; lateral and basal margin clearly bordered; anteriorangles distinctly slanted and posterior angles slightlyobtuse, both with setigerous pore bearing a seta; punc-tation sparse with large punctures weakly impressed onfinely microreticulate surface.

Elytra elongate, entirely covering pygidium, laterallyvery weakly rounded; punctation arranged in 9 regularrows (+1 sutural interrupted at basal third); punctureswell impressed on disc but more weakly on apical de-clivity; interstriae subcarinate, with sparse fine puncta-tion; humeral callus evident; macropterous. Scutellumsubtriangular, with finely shagreened surface.

Legs entirely pale; anterior tibiae clearly enlarged,laterally compressed; hind tibiae with apical spur short,reddish brown; 1st anterior and middle tarsomere clear-ly dilated, cordiform.

Ventrally bright reddish brown; thorax glabrous; ab-domen with yellowish pubescence. Last sternite sparse-ly punctate on almost smooth surface; particular preapi-cal impressions absent.

Median lobe of aedeagus slender (Fig. 26), withsmooth surface; in ventral view apically lanceolate, lat-erally convergent from apical fifth to base; apex strong-ly acute; ventral sulcus wide but indistinct; dorsal sulcuswide, moderately impressed; median lobe in lateralview medially weakly arcuate, apically straight.

Paratype (9) - LB = 2.18 mm; LAN = 1.40 mm; LP =0.44 mm; WP = 0.73 mm; LE = 1.60 mm; WE = 1.03mm; LSP = 0.16 mm; LAN/LB = 0.64; WP/LP = 1.68;LE/LP = 3.68; LE/LSP = 10.04.

Paratype examined darker than holotype (becausemore mature). Body shape in female similar to male;antennae comparatively little shorter; 1st tarsomere ofthe pro- and mesotarsi narrower; spermatheca (Fig. 30)very small, with basal part subglobose and ductusstraight, uncoiled, comparatively long.

Etymology - This species is named after John StanleyGardiner, the organizer of the Percy Sladen Trust Expe-dition to the Seychelles in 1908-1909.

Distribution - Indian Ocean: Seychelles (Silhouette).Maulik (1931: 251-252) reported Chaetocnema mahensisand C. krishna from Silhouette, from the following local-ities: "near Mont Pot-a-eau, circa 1500 feet, viii.1908, 1example; highest forests, over 2000 feet, ix.1908, 2 spec-imens." (C. mahensis); "near Mont Pot-a-eau, circa 1500feet, viii.1908, and Mare aux Cochons, circa 1000 feet,ix.1908.". These data are attributed to S. gardineri n. sp.

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CHAETOCNEMA AND RELATED GENERA 363

Ecological notes — The specimens attributed to thisnew species were collected in August and September inthe high endemic forest from about 1000 feet to over2000 feet (Maulik, 1931: 251-252). Maulik (1931: 251--252) reported five specimens from Sihouette, but it wasnot possible to trace the three missing specimens in thecollection of the BMNH.

Seychellaltica praslinensis n. sp.

(Figs 22, 25, 29)

Chaetocnema krishna Maulik, 1931: 251 (partim)

Type material-YloXotype, <$, SEYCHELLES, Praslin, 1908,Seychelles Exp., Percy Sladen Trust Expedition, B.M. 1927-444" (BMNH: ex syntype of Chaetocnema krishna Maulik).Paratypes, one C?, one 9, same data as the holotype(BMNH: ex syntypes of Chaetocnema krishna Maulik).

Diagnosis — This new species is the smallest one ofthe genus. The median lobe of aedeagus and particular-ly the spermatheca are very similar in shape to S. gar-dineri n. sp. However, S. praslinensis n. sp. shares withS. krishna (Maulik) a similar body shape, coloration ofthe dorsal integument, and elytral interstriae weaklyconvex. Diagnostic characters for identification of thisnew species are given in the key.

Description of the holotype (C?) - LB = 1.80 mm; LAN =1.37 mm; LP = 0.39 mm; WP = 0.59 mm; LE = 1.27 mm;WE = 0.82 mm; LAED = 0.66 mm; LAN/LB = 0.76;WP/LP = 1.52; LE/LP = 3.25; LE/LAED = 1.93.

Dorsal integument bright reddish brown. Body shapeelongate-elliptical, weakly convex. Maximum pronotalwidth in distal part; maximum elytral width at apical third.

Head with frons and vertex sparsely and weakly punc-tate; surface almost smooth; frontal sulci very short, notvisible around the eyes; frontal tubercles absent; interan-tennal space about as wide as length of 1st antennomere;frontal carina wide and flat; labrum and palpi yellowish;antennae slightly longer than half body; antennomeres 1--8 entirely pale, 9-11 clearly blackened; length of eachantennal segment proportional to numerical sequence53:40:52:43:62:56:54:55:49:42:66 (1 = 0.003 mm).

Pronotum transverse, anteriorly wider, laterally slightlywider than posteriorly; laterally weakly rounded; lateraland basal margin clearly bordered; anterior angles distinct-ly slanted and posterior angles slightly obtuse, both withsetigerous pore bearing a seta; punctation sparse withcoarse punctures weakly impressed on smooth surface.

Elytra elongate, entirely covering pygidium, laterallymoderately rounded; punctation arranged in 9 regularrows (+1 sutural interrupted at basal third); punctureswell impressed on disc but more weakly on apical de-clivity; interstriae subconvex, rarely punctate; humeralcallus evident; macropterous. Scutellum semi-circular,dark brown, with finely shagreened surface.

Legs entirely yellowish; anterior tibiae unmodified;hind tibiae with apical spur short, reddish; 1st anteriorand middle tarsomere moderately dilated.

Ventrally bright reddish brown; thorax glabrous; ab-domen with sparse yellowish pubescence. Last sternitenot punctate, with very finely shagreened surface; par-ticular preapical impressions absent.

Median lobe of aedeagus (Fig. 25) slender, withsmooth surface; in ventral view wider at basal half, api-cally lanceolate; apex acute; ventral sulcus very narrow,visible from apical part to base; dorsal sulcus wide,weakly impressed; median lobe in lateral view almoststraight but apically slightly sinuous.

Paratypes - Male: LB = 1.74 mm; LAN = 1.13 mm; LP= 0.35 mm; WP = 0.60 mm; LE = 1.24 mm; WE = 0.78mm; LAED = 0.65 m; LAN/LB = 0.70 + 0.08; WP/LP =1.72; LE/LP = 3.55; LE/LAED = 1.91. Female: LB = 1.99mm; LAN = 1.21 mm; LP = 0.40 mm; WP = 0.69 mm; LE= 1.54 mm; WE = 1.03 mm; LSP = 0.13 mm; LAN/LB =0.61; WP/LP = 1.73; LE/LP = 3.87; LE/LSP = 11.85.

Paratypes examined similar in colour and sculpture toholotype. Body in female not much longer than inmale; antennae relatively shorter; 1st tarsomere of thepro- and mesotarsi narrower; spermatheca (Fig. 29) verysmall, with basal part subglobose and ductus short,straight, uncoiled.

Etymology — The specific name refers to the island ofthe Seychelles where the type locality is located,Praslin.

Distribution - Indian Ocean: Seychelles (Praslin).Maulik (1931: 252) gave the following locality forChaetocnema krishna from Praslin, "Cote d'Or Estate,forest of Coco-de-Mer Palms (Lodoiced) in the Vallee deMai, xi.1908.", which must refer to this new species.

Remarks - The specimens attributed to this newspecies were collected in November in the endemic for-est of Lodoicea (Maulik 1931: 252). Maulik (1931: 252) re-ported 14 specimens from Praslin, but the location of the11 missing specimens has not been possible to establish.

Key to species q/'Seychellaltica n. gen.

1. Dorsal integument yellowish, sometimes partially darker on ely-tra and pronotum. Elytra narrow, laterally subparallel. Medianlobe of aedeagus (Fig. 23) in ventral view wider at middle; inlateral view medially strongly bent. Spermatheca (Fig. 27) large,with elongate basal part and strongly arcuate ductus.

S. mahensis (Maulik)

Dorsal integument uniformly brown. Elytra wider, laterallymoderately rounded. Median lobe of aedeagus in ventral viewnot wider at middle. Spermatheca with subglobose basal part.

2.

2. 8th antennomere mostly darkened. Elytral interstriae subcari-nate. Anterior tibiae in male enlarged and laterally compressed.Median lobe of aedeagus (Fig. 26) in ventral view apicallylanceolate, with very acute apex; in lateral view, medially mod-

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364 M. BIONDI

erately arcuate, apically almost straight. Spermatheca (Fig. 30)small, with elongate and straight ductus.

S. gardineri n. sp.

8th antennomere entirely yellowish. Elytral interstriae flat orsubconvex. Anterior tibiae in male unmodified.

3.

3. Size smaller (LB < = 2.10 mm). 1st anterior tarsomere in malerelatively narrower. Median lobe of aedeagus (Fig. 25) in ven-tral view with wide ventral sulcus; apex acute, subtriangular; inlateral view almost straight but apically weakly sinuous. Sper-matheca (Fig. 29) small.

S. praslinensis n. sp.

Size larger (LB > 2.10 mm). 1st anterior tarsomere in male rela-tively wider. Median lobe of aedeagus (Fig. 24) in ventral viewwith narrow ventral sulcus; apex with a small rounded mediantooth; in lateral view, strongly curved in apical third. Sperma-theca (Fig. 28) large.

S. krishna (Maulik)

Genus Terpnochlorus Fairmaire(Fig. 6)

Terpnochlorus Fairmaire, 1904: 269

Type species - Terpnochlorus perrieri Faimaire, 1904by original designation (type in MNHN examined).

Distribution — South America (Venezuela), central andsouthern Africa, Madagascar.

Remarks — Originally, this genus was attributed to thesubfamily Galerucinae, later it was included by Labois-siere (1932: 575) among the Alticinae. Terpnochlorusshows a possible distribution of the Gondwanian type.In fact, this genus is known from the Afrotropical Re-gion (including Madagascar) with the species T. perrieriFairmaire, 1904 and T. janssensi Bechyne, I960, andfrom the Neotropical Region with the species T ameri-canus]. Bechyne & B. Bechyne, 1967. These specieslive in moist environments and are associated with Jun-caceae (Bechyne & Bechyne, 1967; personal data). Onthe basis of the examination of spermatheca, medianlobe of aedeagus, metafemoral spring and wing vena-tion, Furth & Suzuki (1994) assert that the systematicposition of this genus is unclear. My opinion is that allthese morphological characters can be included in thenormal variability of the genera-group here discussed.

Key to genera1. Prothorax subcylindrical. Elytral punctation at least partially reg-

ular and strongly impressed, arranged in 11 rows (+1 long su-tural). Elytra often with two or three lateral longitudinal carinae.1st tarsomere of hind legs subtriangular and shorter than 2nd.Apex of hind tibiae with short bristles. Ventral margin of hindfemora rounded, not carinate. Eyes roundish, protuberant.

Terpnochlorus Fairmaire (Fig. 6)

Prothorax not subcylindrical, more or less dorsally depressed.Elytral punctation generally more weakly impressed, arrangedin 9 rows (+1 short sutural) at least partially regular. Elytra with-out lateral longitudinal carinae. 1st tarsomere of hind legs sub-

rectangular and longer than 2nd. Apex of hind tibiae finely den-ticulate. Ventral margin of hind femora carinate. Eyes generallysubelliptical and weakly protuberant.

2.

2. Hind tibiae with dorsal margin bidentate (Figs 18-19).3.

Hind tibiae with dorsal margin unidentate (Fig. 17). [SometimesPepila has an indication of a dentate emargination in the innerdorsal margin of the hind tibiae, but this genus is easily distin-guishable for other characters].

4.

3. Frontal sulci very short, not visible around eyes (Fig. 14). Bodyshape elongate, with pronotum weakly transverse. Ratiometasternal width/metasternal length equal or less than 2.5. 1st

hind tarsomere at least twice as long as 2nd and 3rd together(Fig. 19). 1st tarsomeres of the pro- and mesotarsi distinctlyasymmetric (Fig. 20). Hind tibial socket narrow (Fig. 19).

Seychellaltica n. gen. (Fig. 5)

Frontal sulci long, clearly visible around eyes. Body shape oval,with pronotum clearly transverse. Ratio metasternal width//metasternal length greater than 2.5. 1st hind tarsomere shorterthan 2nd and 3rd together (Fig. 18). 1st anterior and middle tar-somere symmetric. Hind tibial socket wide (Fig. 18).

Biodontocnema Biondi (Fig. 2)

4. Interocular space at least with evident transverse carina (Fig. 7).Distal margin of the frons medially distinctly incised (Figs 7-8).Interantennal space at least twice as wide as length of 1st an-tennomere (Fig. 7).

Carcharodis Weise (Fig. 1)

Interocular space without transverse carinae. Distal margin offrons medially not incised. Interantennal space less than twiceas wide as length of 1st antennomere.

5.

5. Hind tibiae dorsally channelled in proximal half. Hind apicaltarsal segment normally more or less globose (Fig. 21). Thirdhind tarsomere small, not or very weakly medially incised.Frontal sulci short or finely impressed, not visible around eyes(Fig. 13). Pronotum generally smaller (LE/LP>3.10).

Pepila Weise (Fig. 4)

Hind tibiae not dorsally channelled. Hind apical tarsal segmentnot globose (Fig. 17). Third hind tarsomere larger, often medial-ly clearly incised. Frontal sulci long, deeply impressed, clearlyvisible around eyes. Pronotum generally larger (LE/LP ? 3.10).

6.

6. Prosternum anteriorly strongly convex, shaped to receivemouthparts in resting position, except labrum and mandibleswhich act as 'cover' (Figs 10-12). Maxillae and labium sunken(Fig. 9). Pronotal punctation with a mixture of punctures of dif-ferent sizes. Vertex of head in lateral view protruding.

Collartaltica Bechyne (Fig. 3)

Prosternum anteriorly moderately convex, not able to receivemouthparts in resting position. Maxillae and labium notsunken. Pronotal punctation uniform. Vertex of head in lateralview moderately convex.

Chaetocnema Stephens

Phylogenetic analysis

As reported above, this natural group of flea beetlegenera is characterized by the following synapomorphies:(1) middle and hind tibiae both with ciliate dentate dorsalemargination; (2) 1st and 2ncl abdominal sternites joined.The taxa considered also share the following character--states: (1) procoxal cavities closed; (2) apical spur of

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CHAETOCNEMA AND RELATED GENERA 365

hind tibiae simple; (3) tarsal claws simple; (4) elytralpunctation at least partially arranged in regular rows.

A cladistic analysis of all the genera here consideredbased on morphological characters was undertaken.Parsimony method was used with PAUP 4.0b8 (Swof-ford, 2000). Multistate characters were treated as un-ordered characters. The genus Terpnochlorus was con-sidered in the analysis as an ancestral group. The polar-ity of the character-states (Table I) was determined withreference to the ancestor.

The strict consensus tree (tree length = 23; consisten-cy index = 0.9565; homoplasy index = 0.0435; retentionindex = 0.8571), obtained from the parsimony analysisbased on 21 morphological characters (see below), isshown in Figure 31. In this tree, the following mono-phyletic groups can be recognized:

- (Seychellaltica (Pepila (Chaetocnema, Collartaltica(Biodontocnema, CarcharodisJ))), characterized byfive synapomorphies (characters 8, 9, 15, 17, 19) (seetext and Table I) and distinctly separate from thegenus Terpnochlorus,

- (Pepila (Chaetocnema, Collartaltica (Biodontocnema,Carcharodis))), characterized by two synapomorphies(characters 11, 12) (see text and Table I);

- (Chaetocnema, Collartaltica (Biodontocnema, Car-charodis)), characterized by one synapomorphy(character 2) (see text and Table I), including all thegenera, but not Chaetocnema, endemic of theAfrotropical Region; however, the node Collartaltica--Chaetocnema is not resolved;

- (Biodontocnema, Carcharodis), characterized by twosynapomorphies (characters 3, 13) (see text and Table I).

It can also be pointed out that the endemic genera ofthe Afrotropical Region, Collartaltica, Biodontocnemaand Carcharodis, show the most affinity with Chaeto-cnema, while Seychellaltica, from the Indian Ocean,and Pepila, from the Australian Region, apparently

8 9 1 5 1 7 1 90000 0

141620 21a

eea e— SEYCHELLALTICA

11 12

-e-e-

• CHAETDCNEMA

51000 CdLLARTALTICA

7 2 1 b

313-©-©-

BiaoQNTOCNEMA

C A R G H A R O D I S

18 200 D P E P I L A

• TERPNaCHLDRUS

Fig. 31 - The strict consensus tree from the parsimony analysisbased on 21 morphological characters (see text).

evolved in earlier times. In this analysis, only character20 (see text and Table I) exhibits homoplasy.

For this genera group, a Gondwanian origin followedby events of differentiation to the genus level mainly inthe present Afrotropical Region (Biodontocnema, Car-charodis, Collartaltica and Seychellaltica) and Australia(Pepila) can be hypothesized. Regarding Chaetocnema,to localize its area of origin is not easy. Anyway thisgenus, unlike others of this group, radiated widely inthe Nearctic and Palaearctic Regions.

TABLE I - Matrix of the character-states considered in the cladistic analysis (see text).

10 11 12 13 14 15 16 17 18 19 20 21

Terpnochlorus

Pepila

Carcharodis

Seychellaltica

Biodontocnema

Collartaltica

Chaetocnema

0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0

1 1 1 1 0 1 0

0 0 0 0 0 0 0

1 0 0 0 1 1

1

1

1

1

1

1

1

1

0

0

0

0

1

1

0

1

1

1

0

1

0

1

0

10

0 1 0 0 1 0 0 1 1 1 1

0 1 0 0 0 0 0 1 1 0 1

0 0 0 0 0 0 0 0 0 0

0 0

0 0 1 0

0 0

0

0

1

0

0

0

1

1

1

1

1

1

1

0

0

0

0

0

1

1

1

1

1

1

1

0

1

0

0

0

0

0

1

2

0

0

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366 M. BIONDI

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