Elenco delle Memorie della Società Italiana di Scienze Naturali

e del Museo Civico di Storia Naturale di Milano

Volume I

I- CORNALIA E., 1865 - Descrizione di una nuova specie dei

genere Felis: Felis jacobita (Corn.). 9 pp., 1 tav.

Il - MAGNI-GRIFFI F., 1865 - Di una specie d’Hippolais nuova

per l’Italia. 6 pp., / tav.

III - GASTALDI B., 1865 - Sulla riescavazione dei bacini lacustri

per opera degli antichi ghiacciai. 30 pp., 2 figg., 2 tavv.

IV - SEGUENZA G., 1865 - Paleontologia malacologica dei terreni

terziarii del distretto di Messina. 88 pp., 8 tavv.

V - GIBELLI G., 1865 - Sugli organi riproduttori del genere Verru-

caria, 16 pp., 1 tav.

VI - BEGGIATO F. S., 1865 - Antracoterio di Zovencedo e di Mon-

teviale nel Vicentino, /0 pp., 1 tav.

VII - COCCHI I., 1865 - Di alcuni resti umani e degli oggetti di

umana industria dei tempi preistorici raccolti in Toscana. 32

pp., 4 tavv.

VIII - TARGIONI-TOZZETTI A., 1866 - Come sia fatto l’organo

che fa lume nella lucciola volante dell’Italia centrale (Luciola

italica) e come le fibre muscolari in questo ed altri Insetti ed

Artropodi. 28 pp., 2 tavv.

IX - MAGGI L., 1865 - Intorno al genere Aeolosoma. /8 pp., 2

tavv.

X - CORNALIA E., 1865 - Sopra i caratteri microscopici offerti dalle

Cantaridi e da altri Coleotteri facili a confondersi con esse. 40

pp., 4 tavv.

Volume II

I- ISSELA,, 1866 - Dei Molluschi raccolti nella provincia di Pisa,

38 PP.

Il - GENTILLI A., 1866 - Quelques considérations sur l’origine

des bassins lacustres, àpropos des sondages du Lac de Come.

12 pp., 8 tavv.

HI - MOLON F,, 1867 - Sulla flora terziaria delle Prealpi venete.

140 pp.

IV - D'ACHIARDIA., 1866 - Corallarj fossili del terreno nummu-

litico delle Alpi venete. 54 pp., 5 tavv.

V - COCCHI I., 1866 - Sulla geologia dell’alta Valle di Magra. /8

p., 1 tav.

VI - SEGUENZA G,, 1866 - Sulle importanti relazioni paleontolo-

giche di talune rocce cretacee della Calabria con alcuni terreni

di Sicilia e dell’Africa settentrionale. /8 pp., 1! tav.

VII - COCCHI I., 1866 - L'uomo fossile nell’Italia centrale. 82 pp.,

21 figg., 4 tavv.

VII - ‘GAROVAGLIO S., 1866 - Manzonia cantiana, novum Li-

chenum Angiocarporum genus propositum atque descriptum. 8

Pp., 1 tav.

IX - SEGUENZA G.,, 1867 - Paleontologia malacologica dei terreni

terziari del distretto di Messina (Pteropodi ed Eteropodi). 22

Pp., 1 tav.

X- DURER B,, 1867 - Osservazioni meteorologiche fatte alla Villa

Carlotta sul lago di Como, ecc. 48 pp. // tavv.

Volume III

I- EMERY C., 1873 - Studii anatomici sulla Vipera Redii. 16 pp.,

I tav.

Il - GAROVAGLIO S., 1867 - The/opsis, Belonia, Weitenwebera et

Limboria, quatuor Lichenum Angiocarporum genera recognita

iconibusque illustrata. 12 pp., 2 tavv.

III - TARGIONI-TOZZETTI A., 1867 - Studii sulle Cocciniglie. 88

., 7 tavv.

IV - CLAPAREDE E. R. e PANCERI P., 1867 - Nota sopra un Al-

ciopide parassito della Cydippe densa Forsk. 8 pp. 1 tavv.

V - GAROVAGLIO S., 1871 - De Pertusariis Europae mediae

commentatio. 40 pp., 4 tavv.

Volume IV

I- D’ACHIARDIA., 1868 - Corallarj fossili del terreno nummuli-

tico dell’Alpi venete. Parte 11. 32 pp. 8 tavv.

II - GAROVAGLIO S., 1868 - Octona Lichenum genera vel adhuc

controversa, vel sedis prorsus incertae in systemate, novis de-

scriptionibus iconibusque accuratissimis illustrata. 18 pp., 2

tavv.

III - MARINONI C., 1868 - Le abitazioni lacustri e gli avanzi di

umana industria in Lombardia. 66 pp., 5 figg., 7 tavv.

IV - (Non pubblicato).

V - MARINONI C., 1871 - Nuovi avanzi preistorici in Lombardia.

28 pp., 3 figg., 2 tavv.

NUOVA SERIE

Volume V

I- MARTORELLI G., 1895 - Monografia illustrata degli uccelli di

rapina in Italia. 2/6 pp., 46 figg., 4 tavv.

Volume VI

I- DE ALESSANDRI G., 1897 - La pietra da cantoni di Rosigna-

no e di Vignale. Studi stratigrafici e paleontologici. /04 pp., 2

tavv., 1 carta.

II - MARTORELLI G., 1898 - Le forme e le simmetrie delle mac-

chie nel piumaggio. Memoria ornitologica. //2 pp., 63 figg., 1

tavv.

III - PAVESI P., 1901- L’abbate Spallanzani a Pavia. 68 pp., 14

figg., 1 tav.

Volume VII

I- DE ALESSANDRI G., 1910 - Studi sui pesci triasici della

Lombardia. /64 pp., 9 tavv.

Volume VIII

I- REPOSSI E., 1915 - La bassa Valle della Mera. Studi petrogra-

fici e geologici. Parte I. pp. 1-46, 5 figg., 3 tavv.

II - REPOSSI E., 1916 (1917) - La bassa Valle della Mera. Studi

petrografici e geologici. Parte II. pp. 47-186, 5 figg. 9 tavv.

III - AIRAGHI C., 1917 - Sui molari d’elefante delle alluvioni lom-

barde, con osservazioni sulla filogenia e scomparsa di alcuni

Proboscidati. pp. 187-242, 4 figg., 3 tavv.

Volume IX

I- BEZZI M,, 1918 - Studi sulla ditterofauna nivale delle Alpi

italiane. pp. VE 164, 7 figg. 2 tavv. ;

II - SERAG. L., 1920 - Sui rapporti della conformazione della base

del cranio colle forme craniensi e colle strutture della faccia

nelle razze umane. (Saggio di una nuova dottrina craniologica

con particolare riguardo dei principali cranii fossili). pp. 165-

262, 7 figg., 2 tavv.

III - DE BEAUXO. e FESTA E., 1927 - La ricomparsa del Cinghia-

le nell’Italia settentrionale-occidentale. pp. 263-320, 13 figg.,

7 tavv.

Volume X

I - DESIO A., 1929 - Studi geologici sulla regione dell’ Albenza

(Prealpi Bergamasche). pp. /-156, 27 figg., 1 tav., 1 carta.

II - SCORTECCI G., 1937 - Gli organi di senso della pelle degli

Agamidi. pp. 157-208, 39 figg. 2 tavv.

III - SCORTECCI G., 1941- I recettori degli Agamidi. pp. 209-326,

80 figg.

Volume XI

I- GUIGLIA D,, 1944 - Gli Sfecidi italiani del Museo di Milano

(Hymen.). pp. i -44, 4 figg., 5 tavv.

I-II - GIACOMINI V. e PIGNATTI S., 1955 - Flora e Vegetazione

dell’ Alta Valle del Braulio. Con speciale riferimento ai pascoli

di altitudine. pp. 45-238, 31 figg., 1 carta.

Volume XII

I- VIALLI V., 1956 - Sul rinoceronte e l’elefante dei livelli su-

periori della serie lacustre di Leffe (Bergamo). pp. 1-70, 4 figg.

6 tavv.

I- VENZO S., 1957 - Rilevamento geologico dell’anfiteatro more-

nico del Garda. Parte I: Tratto occidentale Gardone-Desenzano.

pp. 71-140, 14 figg., 6 tavv., 1 carta.

III - VIALLI V., 1959 - Ammoniti sinemuriane del Monte Albenza

(Bergamo). pp. 141-188, 2 figg., 5 tavv.

Volume XHI

I- VENZO S., 1961- Rilevamento geologico dell’anfiteatro moreni-

co del Garda. Parte II. Tratto orientale Garda-Adige e anfiteatro

atesino di Rivoli veronese. pp. /-64, 25 figg., 9 tavv., 1 carta.

Il - PINNA G., 1963 - Ammoniti del Lias superiore (Toarciano)

dell’ Alpe Turati (Erba, Como). Generi Mercaticeras, Pseudo-

mercaticeras e Brodieia. pp. 65-98, 2 figg., 4 tavv.

Armando Bilardo & Saverio Rocchi

A revision of the African species of the genus

Neptosternus Sharp, 1882

(Coleoptera Dytiscidae)

MCZ

LIBRARY

APK

1À ì \ ARIDO

HAI

UNIVERSITY

Volume XXXVII - Fascicolo III

Novembre 2012

Memorie della Società Italiana di Scienze Naturali

e del Museo Civico di Storia Naturale di Milano

INDICE

INTRODUCIIONE== === ee "===: Pag. 3

MATERIALAND METHODS .......................-. Pag. 5

ADDIEVIalons' ro en n Pag. 5

THE AFRICAN SPECIES OF THE GENUS

NEPRIOSIERNOSE,O Pag. 6

Group A. Species with elytral pattern arranged

as InscheMeAzgzs, Co IRE Pag. 6

Group B. Species with elytral pattern arranged

asinscheme Been Paolo

Group C. Species with elytral pattern arranged

ds inschemet. Pag. 15

ECGOLONER EE; A n Ana Pag. 38

CHECKLISTOFTHEAFRICANSPECIES...... Pag. 46

Aknowledoements se e Pag. 46

REEPBRENCESSE I, Re ne Pag. 47

Museo Civico di Storia Naturale di Milano

Corso Venezia, 55 - 20121 Milano

In copertina: Neptosternus longitarsis n. sp. Line drawing by C. Pesarini.

Registrato al Tribunale di Milano al n. 6694

Direttore responsabile : Anna Alessandrello

Grafica editoriale: Michela Mura

Stampa: Litografia Solari, Peschiera Borromeo - Novembre 2012

ISSN 0376-2726

Armando Bilardo & Saverio Rocchi

A revision of the African species of the genus

Neptosternus Sharp, 1882

(Coleoptera Dytiscidae)

Abstract - The species of the genus Neptosternus in Africa are revised and the taxonomic characters discussed. The species of

this genus are very difficult to identify; this paper is an attempt to provide a suitable method of identification based above all on a rich

iconography. Four new species are described: bartolozzii, komareki, longitarsis and maliensis. The lectotypes of the following species

are designated: africanus Peschet, 1916, alluaudi Régimbart, 1903, oberthueri Régimbart, 1903, oblongus Régimbart, 1895, ornatus

Sharp, 1882, pumicatus Guignot, 1949 and rotroui (Pic, 1924). The following new synonimies are proposed: meridianus Omer-Cooper,

1970, nigeriensis Omer-Cooper, 1970, rotroui (Pic, 1924) and tropicus Guignot, 1954 = ornatus Sharp, 1882; psephotus Guignot, 1955

= sombuicus Guignot, 1954; si/vester Guignot, 1960 = a/luaudi Régimbart, 1903. N. bilardoi Pederzani & Rocchi, 2008 is downgraded

to subspecies of oberthueri Régimbart, 1903. The neotytpe is designated for N. nuperus Guignot, 1954.

Key words: Dytiscidae, Neptosternus, Africa, review, new species.

Riassunto - Revisione delle specie africane del genere Neptosternus Sharp, 1882 (Coleoptera Dytiscidae).

Viene riesaminata la tassonomia del genere Neptosternus Sharp, 1882 in Africa; si tratta di un genere molto difficile, dove il rico-

noscimento delle specie risulta assai problematico; il lavoro è pertanto corredato da una ricca iconografia, quale supporto fondamentale

per la determinazione dei vari taxa. Vengono descritte 4 nuove specie: bartolozzii, komareki, longitarsis e maliensis. Vengono designati

i lectotipi delle seguenti specie (alcune di queste anche con 1 relativi paralectotipi): africanus Peschet, 1916, alluaudi Régimbart, 1903,

oberthueri Régimbart, 1903, oblongus Régimbart, 1895, ornatus Sharp, 1882, pumicatus Guignot, 1949 e rotroui (Pic, 1924). Vengono

proposte le seguenti nuove sinonimie: meridianus Omer-Cooper, 1970, nigeriensis Omer-Cooper, 1970, rotroui (Pic, 1924) e tropi-

cus Guignot, 1954 = ornatus Sharp, 1882; psephotus Guignot, 1955 = sombuicus Guignot, 1954; silvester Guignot, 1960 = alluaudi

Régimbart, 1903. Viene modificato il rango di bilardoi Pederzani & Rocchi, 2008 da specie a sottospecie di oberthueri Régimbart,

1903. Si designa il neotipo di N. nuperus Guignot, 1954.

Parole chiave: Dytiscidae, Neptosternus, Africa, revisione, nuove specie.

INTRODUCTION

The genus Neptosternus Sharp, 1882 occurs in Africa and

South-Fast Asia. According to the catalogue Nilsson (2001)

the genus comprises 22 African and 70 Oriental species. The

type species of the genus is Neptosternus ornatus Sharp,

1882 described from Madagascar and Zanzibar (Tanzania).

The Oriental species were revised by Hendrich & Bal-

ke (1997) and subsequently updated with descriptions of

new species by Balke et a/. (1997), Balke & Hendrich

(1998 and 2001), Hendrich & Balke (1999, 2000a and

2000b); the Chinese species were recently revised by

Zhao et al. (2012), whereas in recent years no modern

revision was published of the African species. After the

classic revision by Guignot (1959a) only one contribution

to the study of African Neptosternus was published by

Omer-Cooper (1970) with the examination of some types,

but the number of studied specimens was not so high as to

show the range of the intraspecific variability. Neverthe-

less Omer-Cooper (loc. cit.) had the merit of the first at-

tempt to solve identification problems by introducing new

taxonomic characters. Unfortunately Omer-Cooper 's stu-

dy was limited to continental species, with the exclusion

of Madagascan ones, and the study was based on material

from too few localities and low number of specimens, that

caused the establishment of some new species on speci-

mens being within the range of variability of one species.

An example is N. ornatus Sharp, 1882 a very variable spe-

cies widespread in Madagascar, western, central and sou-

thern Africa: its wide area of distribution probably caused

the formation of morpho-clines and locally variable races

that were wrongly described as different species; for that

reason some synonimies proposed here below principally

concern that variable species.

In our foregoing papers (Bilardo & Pederzani, 1978;

Bilardo, 1982; Pederzani & Rocchi, 1982; Bilardo &

Rocchi, 1990 and 1999) the records of Neptosternus we-

re usually reported only in part, due to the difficulty in

reaching reliable identifications; however in spite of ta-

xonomic difficulties, Bilardo & Rocchi (2010) described

four Afrotropical new species clearly distinguishable by

peculiar characters and two possibly new species briefly

described but not denominate, being in need of further

study. Pederzani & Rocchi (2008) described another new

species from Madagascar, whose status will be discussed

here below. The above-mentioned African new taxa were

the first published after Omer-Cooper (1970), that is with

a gap of some 40 years.

4 ARMANDO BILARDO & SAVERIO ROCCHI

For that reasons we felt the need to better investigate

the African species of this difficult genus. Our first tar-

get was the study of type material of all taxa described

up to date, that proved to be an arduous task, but after all

we succeeded in looking over all necessary type material,

except for two species (N. hedychrous Guignot, 1955 and

N. nuperus Guignot, 1954, the latter because the paratype

obtained for study is in fact a N. compsus Guignot, 1953).

The great difficulty in the identification of the species

was always emphasized by authors. Omer-Cooper (1970),

Bilardo & Rocchi (1990), Pederzani & Rocchi (2008) and

Bilardo & Rocchi (2010) agreed upon the unsatisfactory

taxonomic value of the microsculpture; quoting Omer-

Cooper (loc. cit.) the microsculpture is “represented by a

microreticulation and micropunctuation ( ‘“alutacé” Gui-

gnot). This can only be seen under a high magnification,

and with slight changes in the direction of light and in the

focus its appearance may change from one to the other”.

The structure of aedeagus is poorly differentiated in

Neptosternus and somewhat asymmetric, so its median

lobe appears of different shape depending on the orienta-

tion and the angle formed with the optical axis of micro-

scope. Female genitalia are almost alike in all species; in

our opinion they don 't show peculiarities, so we thought it

best not to thoroughly investigate their structures.

Prosternum is obviously keeled as a rule (Guignot,

1959a; Omer-Cooper, 1970); in some species of group B

(see later the subdivision into groups) it is keeled only at base

and transversely convex, not keeled, behind; in N. compsus

Guignot, 1953 it is not keeled in the holotype but shows

traces of a keel in other specimens; therefore its feature is

unreliable as a taxonomic character, and is just mentioned

in a few cases and in the descriptions of the new species.

The apophysis of prosternum has a median spine tapering to

a long, fine point, and lateral spines clearly shorter than the

median (Guignot loc. cit.); the shape of this trident is some-

what variable also in individuals of the same species, so we

cannot rely on it in the identification of species.

The colour of head and its appendages, underside and

legs usually reproduces the colour of elytral marks with

paler or darker gradations of colour depending on the

level of chitinization and also on the age of material (old

specimens are darker as a rule); such colour has no taxo-

nomic value, with some exceptions: N. nuperus Guignot,

1954 represents an exception because its brown underside

is obviously darker than elytral marks. N. batekensis Bi-

lardo & Rocchi, 2010, susinii Bilardo & Rocchi, 2010 and

darker specimens of N. oberthueri bilardoi Pederzani &

Rocchi, 2008 have blackish underside, and are the only

African taxa with completely black head.

Also the testaceous markings on elytra, taken alone,

are often misleading as a taxonomic character, because

the shape and size of the marks show considerable intra-

specific variability. In conclusion, all diagnostic charac-

ters taken one by one are inadequate to provide the usual

diagnosis and descriptions of the species; in fact the sta-

tements and sentences used in the descriptions should be

too repetitive and of little help in the identification. In our

opinion a key to species is unattainable, since it cannot

be based on constant and well definable characters and in

some cases it may be misleading.

Nevertheless, as first noted by Omer-Cooper (loc. cit.),

the basic schemes of arrangement of the elytral marks in

Neptosternus appear to fall into groups that represent a

suitable approach to the study. Therefore the only method

we could work out to identify species is to roughly sub-

divide them into groups, according to the arrangement

of elytral marks, and show for each species, beside basic

information and a general description, a rich set of colou-

red pictures of habitus, penis and parameres. One or mo-

re photos of habitus describe the shape of the beetle and

above all the colour pattern of pronotum and elytra, with

the principal intraspecific variations. The first approach

by photos of habitus and genitalia is integrated by addi-

tional information such as size, shape, and in some cases

also length of mesotarsus (measured without claws): the

latter character (Fig. 1), introduced by Bilardo & Rocchi

(2010), proved to be valid in taxonomy; in fact the con-

spicuous length of mesotarsus is a peculiarity of the genus

Neptosternus and is distinctive of some species, including

one of the new species described below.

The genus Neptosternus in Africa is distinctly splitted in-

to Afrotropical and Madagascan (endemic) species, with the

only exception of N. ornatus Sharp, 1882 that occurs both

in Madagascar and western, central and southern Africa and

represents the most widespread species of the genus.

L mst = L seg.ts 1-5

Fig. 1-N. fasciatus Omer-Cooper, 1970, Gabon, Réserve Lopé Okanda. Right mesotarsus / mesotarso destro. Lmst: = total length of

mesotarsomeres without claws / lunghezza totale dei mesotarsomeri senza unghie.

A REVISION OF THE AFRICAN SPECIES OF THE GENUS NEPTOSTERNUS SHARP, 1882 (COLEOPTERA DYTISCIDAE) 5

MATERIAL AND METHODS

This revision is based on the study of specimens of

the type material of all species, as they were available

to authors, and all material deposited in authors’ col-

lections and received for identification.

Photos were taken by a digital camera CANON

EOS 40 D coupled with a binocular microscope LEI-

CA MZ 12.5, and elaborated at PC by Helicon Focus

and Adobe Photoshop software. Measures were taken

by a micrometric eyepiece of microscope. Genital pie-

ces were dry mounted on the same card as the studied

specimens. The photos of penis were taken from stan-

dard angles so that figures of different species can be

comparable (see explanation of Fig. 2). The penis was

photographed in dorsal aspect, in such a way as to ha-

ve its maximum width perpendicular to the optical axis

of the microscope, and with base and apex on the same

plane. The photo of apex was taken with an inclina-

tion of 45°. Lateral and dorso-lateral aspect of penis is

shown only in case it can help a better understanding

of the shape.

The following information are provided for each

species: literature references (including citations in

faunistic lists), type locality, type (holotype or lectot-

ype) depository, type material examined, additional

material examined, colour pattern of the upper side,

size, body shape, male genitalia (principal features of

penis and parameres), known distribution (according

- DORSAL WIEW

Fig. 2 - N. fasciatus Omer-Cooper, 1970, Gabon, Réserve Lopé Okanda.

Penis right lateral view, direction of the optical axis in the dorsal view

(perpendicular) and in the view of apex (45°). / Pene in vista laterale

destra, direzione dell’asse ottico in vista dorsale (perpendicolare) e in

vista dell’apice (45°).

to material examined and suitable literature data), re-

marks (if necessary). The median lobe of aedeagus

(penis) is briefly described and shown upside-down,

as by classic Authors, that is with basal bulb down, the

right side left and vice-versa.

The parameres were detached from the median lobe

and glued with their inner face up. With a few excep-

tions, no peculiar features were found in either face of

parameres, since they are poorly chitinized as a rule,

and hardly maintain their original shape, so that com-

parison of parameres of different species is without

practical use.

A question mark (?) beside the number of speci-

mens means that the identification is uncertain, in case

of e.g. damaged specimens or females only. Females

with absolutely unrecognizable features were left out

of the list of material, as for instance a series of 7 ex

from Mozambique (Prov. Manica, 45 km NW Chimo-

io, 14-16.XII.2003 leg. A. Kudrna jr., CFP and CAB)

belonging to group A.

The type specimens of the new species as well as

the lectotypes and paralectotypes of the species repre-

sented so far by syntypes (type-series) were provided

with a red label with the printed name of the species

and indication of e.g. “holotypus”, “paratypus”, ‘“ne-

otypus”, ‘“lectotypus” or “paralectotypus”.

Abbreviations

CAB = collection Armando Bilardo, Cassano Magna-

go (VA), Italy

CFP = collection Fernando Pederzani, Ravenna,

Italy

CSR = collection Saverio Rocchi, deposited in the

Natural History Museum of the University of Flo-

rence, Section Zoology “La Specola”, Florence,

Italy

AB = legit A. Bilardo

FP = legit F. Pederzani

ISNB = Institut Royal des Sciences naturelles, Bruxel-

les, Belgium

MNHN = Muséum national d’Histoire Naturelle, Pa-

ris, France

MRAC = Musée Royal de l’Afrique Centrale, Tervu-

ren, Belgium

MSNM = Museo Civico di Storia Naturale, Milan,

Italy

NHML = Natural History Museum, London, UK

NMP = Narodni Muzeum, Praha, Czech Republic

NMW = Naturhistorisches Museum, Wien, Austria

L = total length of beetle (with head)

w = greatest width of beetle

L/w = ratio of total length to greatest width

Lmst/L = ratio of length of the mesotarsus (without

claws) to total length of beetle

Lp = length of penis

ex = specimen/ specimens

m = meters above sea level

mm = millimeters

loc. cit. = the same paper as cited before, (from Latin

“loco citato ”)

6 ARMANDO BILARDO & SAVERIO ROCCHI

THE AFRICAN SPECIES OF THE GENUS NEPTOSTERNUS

This study on African Neptosternus resulted in the de-

scription of 4 new species, the designation of the neot-

ype of N. nuperus, the designation of the lectotypes of 7

species and the proposal of 6 new synonimies. A taxon is

downgraded from species to subspecies rank.

First of all the species were divided into three groups

according to the schematic arrangement of the testaceous

patches on elytra. In that we followed Omer-Cooper

(1970) but we set three species groups instead of two as in

Omer-Cooper (loc. cit.). The resulting schemes are shown

in the figures, and can be described as follows:

Scheme A (the first species group in Omer-Cooper, loc.

cit.) (Figs. 3-53)

a sub-basal band usually divided into two marks: the

inner one close to suture, the outer being an expansion of

the testaceous margin;

two intermediate marks: a juxtasutural inner mark and

an outer squarish or sub-triangular mark connected to the

margin; sometimes the two marks are jointed in a tran-

sverse band, or the inner mark may disappear;

a sub-triangular apical mark.

Scheme B (the second species group in Omer-Cooper,

loc. cit.) (Figs. 54-72)

two sub-basal marks: an inner squarish mark that may

have a backwards prolongement along the suture, and an

outer mark, being an expansion of the marginal border;

these marks may be sometimes jointed;

two intermediate marks, the inner one elongate, pa-

rallel to suture and even connected to the backwards pro-

longement of the inner sub-basal mark; the outer mark

sub-triangular. The two marks may be connected to form

a transverse band;

an additional elongate mark in the middle of elytron

( “extra, short longitudinal testaceus line” in Omer-Coo-

per, loc. cit.);

a sub-triangular apical mark.

Scheme C (derived from Scheme A) (Figs. 73-78)

a sub-basal band, usually broken into up to 4 marks of

various shape and size;

an intermediate band, composed of three marks, the

innermost elongate and parallel to suture, the intermediate

small, the outer sub-triangular; the inner and intermediate

marks sometimes missing;

sometimes a small spot just behind the intermediate

band;

a sub-triangular apical mark.

In each species group the species are treated in alpha-

betical order: known species first, new species next.

GroupA

Species with elytral pattern arranged as in Scheme A

Neptosternus alluaudi Régimbart, 1903 (Figs. 3-5)

Neptosternus Alluaudi Régimbart, 1903: 16; Zimmer-

mann (1920: 28); Guignot (1959a: 605).

Neptosternus alluaudi Régimbart: Bertrand & Legros

(1971: 245); Rocchi (1991: 86); Nilsson (2001: 254).

Neptosternus silvester Guignot, 1960: 99; Rocchi (1991:

86); Nilsson (2001: 256); n. syn.

Type locality: - alluaudi: Madagascar (Fianarantosa);

- silvester: Madagascar (Andobo).

Type depository: MNHN.

Type material examined: - a/luaudi: lectotype 9 (MN-

HN, Fig. 3), designated here: Madagascar, Centre-Sud,

Alluaud 1901 [white label in print], Type [red label in

print], Museum Paris, Coll. Ch. Alluaud [blue label in

print], alluaudi [handwritten white label]; paralectotype

S, 1 ex (MNHN), Mad.[agascar], B. D’Antongil, coll.

Maurice Régimbart, 1908, [median lobe of aedeagus dry

mounted but in bad condition, not suitable for taking pho-

tos]. - silvester: paratypes (1 3, Fig. 5 and 2 99) (MN-

HN), Andobo 190 m, forét Antsingy, det. Antsalova, II-

[19]57, P. Griveaud.

Additional material examined: Madagascar: N. Ma-

dagaskar, Antseranana distr., Sambirana riv., Marovato,

VIII.2001, [leg.] J. Rolcik, 1 &' (Fig. 4) and 2 99 (NMP);

N. Madagaskar, Antseranana distr., Sambirana riv., Ma-

rovato vill., 5-12.12.2001, leg. J. Horak, 1 4 and 1 9

(NMW).

Colour pattern of the upper side: head testaceous, pro-

notum brownish testaceous, anteriorly and posteriorly

dark. Elytra blackish with bright testaceous pattern, ar-

ranged as in Scheme A.

Size: L 3.75-3.96, w 2.09-2.20, L/w 1.75-1.81, Lmst/L

0.16-0.17.

Body shape: regularly oval.

Male genitalia: penis of peculiar shape due to its

clockwise rotation of 90° with respect to the basal bulb;

this peculiar feature is unique in all African Neptoster-

nus. In dorsal view it appears very narrow because of the

twisting, whereas in lateral view it is fairly broad, with the

dorsal side regularly convex and the ventral side sinuate.

Apex pointed; the photo was taken in dorso-lateral view,

to show its greatest width. Right paramere sub-triangular;

left paramere very narrow and elongate.

Distribution: Madagascar.

Remarks: we found no substantial differences between

N. alluaudi and N. silvester, either in body shape, colour

pattern and male genitalia. We therefore consider that N.

silvester is a junior synonym of N. alluaudi.

Neptosternus fasciatus Omer-Cooper, 1970 (Figs. 6-13)

Neptosternus fasciatus Omer-Cooper, 1970: 67; Bilardo

& Pederzani (1978: 120); Bilardo & Rocchi (1990:

178 partim; 2010: 44); Nilsson (2001: 254).

Neptosternus ornatus Sharp: Bilardo & Rocchi (1990:

178; 1999: 232; misidentification).

Type locality: Nigeria (road from Enugo to Makurdi).

Type depository: NHML.

Type material examined: holotype 4 (NHML, Fig. 6),

Nigeria, R. on Enugo rd about n. 7 miles from Makurdi,

24.IV.1963, J.0.C., N. fasciatus n. sp., M.E. Bacchus det.

1978 Holotype, Holotype Brit. Mus. 1978-308.

Additional material examined: Nigeria: Port Harcourt,

new Calabar R., [leg.] O. C. Umeozor, 1 9 ? (NHML).

Gabon: Belinga, 12.3.1963, leg. H.Coiffait, 5 ex (MNW);

A REVISION OF THE AFRICAN SPECIES OF THE GENUS NEPTOSTERNUS SHARP, 1882 (COLEOPTERA DYTISCIDAE) 7

Franceville, villl Mopia, riv. Ikengué, 19.IX.2008,

leg. AB, 2 ex, Fig. 7; 29.VI-1.VIIL2010, leg. AB, 2 ex

(CAB); Franceville, villl Mougouango, Fleuve Ogooué,

23.VIII.1992, leg. AB, 8 ex (CAB), 6 ex (CSR); Lamba-

réné, 7.VIII.1973, leg. AB & FP, 1 ex (CFP); Makokou,

25.5.1963, leg. H. Coiffait, 2 ex (NMW, Fig. 8); Mandji,

route pour Massana, 27.1.1986, leg. AB, 3 ex (CAB, Fig.

9); M. ts Cristal, Kinguélé, 6.VIII.1973, leg. AB & FP, 2 ex

(CAB, Fig. 10), 2 ex (CFP); Ntoum, 1.VIII.1973, leg. AB

& FP, 8 ex (CAB, Fig. 11), 9 ex (CFP), 2 ex (CSR); Réser-

ve Lopé Okanda, 5.VIII.1983, leg. AB, 19 ex (CAB, Figs.

12, 13), 4 ex (CSR); 2.II.1986, leg. AB, 8 ex (CAB).

Colour pattern of the upper side: head and pronotum

reddish testaceous. The arrangement of light marks on

elytra fairly follows the standard of group A; the holot-

ype has a regular inner median spot; specimens from Ga-

bon show some variations: the inner median spot may be

smaller or even disappear and the outer median spot may

be expanded as to reach the apical mark; the two anterior

marks are connected in some specimens. The ground co-

lour of elytra is reddish-brown in the holotype, and the

marks are not very bright, so the contrast of the two co-

lours is rather poor. The specimens from Gabon have a

ground colour brownish-black and the yellow pattern is

more evident.

Size: holotype L 3.30, w 1.87, L/w 1.77, Lmst/L 0.15,

Lp 0.60; specimens from Gabon L 3.17-3.71, w 1.68-2.13,

L/w 1.70-1.84, Lmst/L 0.14-0.17, Lp 0.61-0.70.

Body shape: regularly oval.

Male genitalia: penis distally widened and asymmetri-

cal, apex pointed slightly bent rightwards; right paramere

distally widened, so that it appears sub-rectangular.

Distribution: Nigeria, Gabon.

Neptosternus hedychrous Guignot, 1955

Neptosternus hedychrous Guignot, 1955c: 141; Rocchi

(1991: 86); Nilsson (2001: 254).

Type locality: Madagascar (Sandragato, rivière Saha-

maloto).

Type depository: unknown.

Type material examined: none.

Additional material examined: none.

Colour pattern of the upper side: upper side shown in

the description (Guignot, 1955c: 142, fig. 1). The elytral

pattern consists of a black sutural line and an approxi-

mately median irregular black longitudinal line on a te-

staceus background (Omer-Cooper, 1970: 64); the elytral

marks are arranged as in paler specimens of N. oberthueri

oberthueri Régimbart, 1903.

Size: L3.00.

Body shape: ‘en ovale allongé, assez convexe, attenué

en arrière”.

Distribution: Madagascar.

Remarks: this species was described on a single female

specimen. The type depository is not reported in the origi-

nal description (Guignot, 1955c); we could not trace any

specimen of N. hedychrous in MNHN or elsewhere. There

are no subsequent records of this species in literature. We

just reported the available information on colour and si-

ze as from the original description. Due to the anomalous

elytral pattern with diffuse testaceous colour, Omer-Coo-

per (loc. cit.) did not include this species in her Group I.

However since we find analogies with the variable pattern

of N. oberthueri oberthueri we included hedychrous in our

Group A. This species is closely related to N. oberthueri

oberthueri and possibly a synonym of the latter.

Neptosternus nuperus Guignot, 1954 (Figs. 14-18)

Neptosternus nuperus Guignot, 1954a: 7; Guignot (1954b:

31; 1955b: 182; 1959a: 602); Omer-Cooper (1970:

73); Bilardo & Pederzani (1978: 120); Bilardo & Roc-

chi (1990: 178 partim; 2010: 40; 2011: 195); Nilsson

(2001: 256).

Neptosternus fasciatus Omer-Cooper: Bilardo (1982: 447);

Bilardo & Rocchi (2011: 195; misidentification).

Type locality: (lost type) Democratic Republic of Con-

go (Upemba, Gorges de la Pelenge). Neotype (see remarks

below): Gabon (Franceville, vill. Mopia, riv. Ikengué).

Type depository: (neotype) MSNM.

Type material examined: none. This species was de-

scribed on a single male that should be deposited in ISNB.

In reply to our request of type, they kindly sent a parat-

ype which proved to be N. compsus Guignot, 1953. The

confusion about the wrong paratype(s) of N. nuperus was

already noted by Omer-Cooper (1970: 64); we suppose it

depends either on a mislabelling or on a mistake by Gui-

gnot himself. See below the treatment of N. compsus.

Additional material examined: Cameroun: Kribi Nord,

route pour Edea, 10.1.1979, leg. AB, 9 ex (CAB, Fig. 14),

1 ex (CFP). Gabon: Bakoumba (Franceville), Parc de

Lékédi, 27.VIII 2008, leg. AB, 6 ex (CAB), 1 ex (CFP), 3

ex (CSR); Franceville, vill. Mopia, riv. Ikengué, 29.VI-1.

XWII2010} leg AB+24fcx:(GAB.$Hlgsx15 316: 1)K12 ex

(CSR); Kinguélé, 6.VIII.1973, leg. AB & FP, 1 ex (CFP);

M.ts Cristal, Asseng Assala, 10.VIII.2006, leg. AB, 14 ex

(GABxFig®18),127/ex (CSR);iNtoum;1:VII.19733 leg:

AB & FP, 3 ex (CAB), 1 Sd and 1 £ (CFP).

Colour pattern of the upper side: the ground colour

of elytron is blackish brown; the pattern of yellow marks

slightly differs from the basic arrangement of Group A.

Sub-basal band consisting of two marks: an inner mark

close to suture, and outer mark being the expansion of the

margin; the two marks may be either separate or joined.

The intermediate band is actually post-mediane, that is

displaced a little further back as to the basic arrangement

of Group A; this transverse band is broader at margin and

tapers toward suture, sometimes broken up in elongate

spots or even present only at margin as a sub-triangular

mark. In a few specimens there is also a punctiform spot

at middle of elytron. The apical yellow mark is sub trian-

gular as usual.

Size: L 3.03-3.26, w 1.61-1.76, L/w 1.83-1.89, Lmst/L

0.17-0.19, Lp 0.56-0.58.

Body shape: sub-oval; in some specimens body elon-

gate-oval, as body appears laterally constricted due to less

rounded sides of elytra.

Male genitalia: penis apically widened and asymme-

tric; right side sinuate at apex, showing an inverted ben-

ding, so that apex appears to be bent rightwards; left side

regularly rounded; apex rounded.

Distribution: Cameroun, Gabon, Democratic Republic

of Congo.

Remarks: the above specimens were assigned to this

species on the basis of descriptions and figures by Guignot

8 ARMANDO BILARDO & SAVERIO ROCCHI

(1954a and 1954b) and Omer-Cooper (loc. cit.). Conside-

ring that (1) we were not able to find the holotype of nu-

perus, and (2) also Omer-Cooper (loc. cit.) could not trace

it, we thought it best to designate a neotype of the species,

in order to define this nominal taxon objectively. Since

the lost holotype was described without paratypes and no

material from the formerly type locality is available, we

have chosen as ‘“neotype”’ a male specimen from Gabon,

labelled as follows: Gabon, Franceville, vill. Mopia, riv.

Ikengué, 29.VI-1.VII.2010, leg. AB / &' / (MSNM).

Neptosternus oberthueri oberthueri Régimbart, 1903

(Figs. 19-20)

Neptosternus Oberthiiri Régimbart, 1903: 15; Zimmer-

mann (1920: 28); Guignot (1959a: 600).

Neptosternus oberthuri Régimbart: Bertrand & Legros

(1971: 245); Rocchi (1991: 86).

Neptosternus oberthueri Régimbart: Nilsson (2001: 256)

Type locality: Madagascar (Sainte Marie).

Type depository: MNHN.

Type material examined: lectoype 4 (MNHN, Fig.

19), designated here: S.te Marie de Madagascar, Perrot

Frères, X-XII 1896 [white label in print], Type [red label

in print], Museum Paris, coll. Maurice Régimbart 1908

[blue label in print], Oberthiiri Rég. [handwritten white

label]; paralectotypes 6 ex (MNHN), same data as lec-

totype; paralectotype &' (Fig. 20) and 9 (MNHN), same

data as lectotype, but with red label in print ‘“cotype’”’ and

white label in print “Museum Paris, coll. F. Guignot”.

Additional material examined: none.

Colour pattern of the upper side: head and pronotum

dark-brown. Ground colour of elytra brown to dark-

brown; elytral marks reddish testaceous. The lectotype

has a regular arrangement of the yellow marks; in some

paratypes the reddish-testaceous colour is more extended:

the anterior band is prolonged backwards and even attains

the inner median mark; the outer median mark is wider

and attains the margin of elytron and the apical mark.

Size: L 3.47 (both lectotype Fig. 19 and paralectotype

Fig. 20), w 1.82-1.84, L/w 1.88-1.91, Lmst/L 0.14-0.15,

Lp 0.65-0.67.

Body shape: oblong, narrowed behind.

Male genitalia: penis peculiar for the apical part little

widened, apex progressively pointed, slightly bent righ-

wards.

Distribution: Madagascar.

Neptosternus oberthueri bilardoi Pederzani & Rocchi,

2008 new status (Figs. 21-25)

Neptosternus bilardoi Pederzani & Rocchi, 2008: 91.

Type locality: Madagascar (Andohahela).

Type depository: NMW.

Type material examined: paratype & (CSR, Fig. 21),

Madagascar, Andohahela (Tulear), Isaka, 9.[X.2001, leg.

R. Gerecke & T. Goldschmidt.

Additional material examined: Madagascar: Manam-

patrana Bas. (PO 167), loc. Aff. de la Lantara, Sahanivo-

raky (Andringitra) Riv., 47°00’41”’E - 22°13’33”°S, leg.

Orstom, Antananarivo, 4 £S and 4 99 (NMW, Figs. 22,

23IR0A725).

Colour pattern of the upper side: arrangement of

elytral marks as in standard Group A. Inner interme-

diate mark sometimes missing; in some specimens the

testaceous pattern is reduced: the sub-basal band may

even disappear and the intermediate band is reduced to

only the outer mark, that is an expansion of the testa-

ceous margin of elytron. One specimen has no testa-

ceous pattern at all, and has wholly black elytra: at first

sight it does not look like a Neptosternus and may even

be mistaken for a small Hydrocanthus Say, 1823 (Fam.

Noteridae).

Size: paratype L 4.04, w 2.11, L/w 1.91, Lmst/L 0.15,

Lp 0.79; other specimens L 4.19-4.35, w 2.22-2.26, L/w

1.88-1.96, Lmst/L 0.15-0.16, Lp 0.76-0.82.

Body shape: elongate oval; elytra regularly tapering

from base to apex.

Male genitalia: penis with apical part little widened,

apex progressively pointed, slightly bent rightwards.

Distribution: Madagascar.

Remarks: this taxon differs from the nominotypical

subspecies in much larger size and in minor differences

in body shape and colour pattern, but the median lobe

of aedeagus is alike. This subspecies occurs in southern

Madagascar, while the nominotypical subspecies occurs

in the north.

Neptosternus oblongus Régimbart, 1895 (Figs. 26-28)

Neptosternus oblongus Régimbart, 1895: 150; Régimbart

(1903: 16); Zimmermann (1920: 28); Guignot (1959a:

599); Rocchi (1991: 86); Nilsson (2001: 256).

Type locality: Madagascar (Sikora).

Type depository: MNHN.

Type material examined: lectotype £' (MNHN, Fig.

26), designated here: Annanarivo, F. Sikora [white label

in print], Type [red label in print], Museum Paris, coll.

Maurice Régimbart 1908 [blue label in print], oblon-

gus Rég. [handwritten white label]; paralectotypes 1 d

(Fig. 27) and 3 99 (MNHN), Ankarata, Madag[ascar]

intér[ieur] (Sikora) [handwritten white label], Museum

Paris, coll. Maurice Régimbart 1908 [blue label in print];

3 ex (ISNB), Annanarivo, F. Sikora, det. Régimb.[art]

[18]91, Neptosternus oblongus Rég. [handwritten white

labels], 11195, Coll. R. I. Sc. N. B. [handwritten white

labels].

Additional material examined: Madagascar: Ankara-

tra, Perrier, coll. Maurice Régimbart 1908, 1 £' (MNHN);

Békily, III 1936, coll. F. Guignot, 1 4 (Fig. 28) and 1 ex

of unknown sex (MNHN), 27 ex (ISNB).

Colour pattern of the upper side: head and pronotum

reddish testaceous; pronotum with anterior and posterior

margins brown. Ground colour of elytra brown to dark

brown; elytral marks reddish testaceous; arrangement of

elytral marks as in standard Group A.

Size: L 4.00-4.16, w 2.00-2.20, L/w 1.84-2.00, Lmst/L

0.14-0.15, Lp 0.70-0.73.

Body shape: elongate oval; elytra regularly tapering

from base to apex.

Male genitalia: penis in dorsal view apically asymme-

tric, right side obviously sinuate, apex bent rightwards;

left side from the maximum width onwards, nearly straight

and gently bent to apex; tapering to a long point.

Distribution: Madagascar.

A REVISION OF THE AFRICAN SPECIES OF THE GENUS NEPTOSTERNUS SHARP, 1882 (COLEOPTERA DYTISCIDAE) 9

Neptosternus ornatus Sharp, 1882 (Figs. 29-42)

Neptosternus ornatus Sharp, 1882: 317; Kolbe (1883:

423); Régimbart (1895: 150); Zimmermann (1920:

28); Guignot (1956c: 221; 1959a: 600; 1959b:163);

Omer-Cooper (1957: 22; 1965: 89; 1970: 65); Rocchi

(1991: 86); Nilsson (2001: 256).

Laccophilus rotroui Pic, 1924: 21; n. syn.

Neptosternus Rotroui Pic: Zimmermann (1930: 25); Gui-

gnot (1959a: 606).

Neptosternus rotroui (Pic): Nilsson (2001: 256; 2003:

DeL).

Neptosternus tropicus Guignot, 1954a: 6; Guignot (1954b:

32; 1959a: 603); Omer-Cooper (1970: 71); Nilsson

(2001: 257); n. syn.

Neptosternus meridianus Omer-Cooper, 1970: 66; Nils-

son & Persson (1993: 82); Nilsson (2001: 255); n.

syn.

Neptosternus nigeriensis Omer-Cooper, 1970: 67; Nils-

son (2001: 256); n. syn.

Neptosternus alluaudi Régimbart: Omer-Cooper (1965:

89, misidentification).

Type locality: - ornatus: Madagascar; Zanzibar [Tan-

zania]; - rotroui: Algeria (Sidi-Bel-Abbès); - tropicus:

Democratic Republic of Congo (Riv. Lufira); - meridia-

nus: South Africa (Transvaal); - nigeriensis: Nigeria (road

from Enugo to Makurdi).

Type depository: - ornatus NHML; - rotroui MNHN;

- tropicus MRAC; - meridianus NHML; - nigeriensis

NHML.

Type material examined: - ornatus: lectotype

(NHML, Fig. 29), designated here [this specimen was

chosen because the penis is intact], Madagascar 594 [han-

dwritten white label], Neptosternus ornatus Sharp Cotype

[printed white label], Sharp Coll. 1905-313 [printed white

label]; paralectotypes: 1 ex 4 [with penis a little damaged]

(NHML), Madagascar, Type 594 [handwritten white la-

bel], Sharp Coll. 1905-313 [handwritten white label]; 2 ex

glued on the same card (NHML), Zanzibar, Neptosternus

ornatus, Ind. Typ. D. S. [handwritten on the card], Cotype

[white label in print], Sharp Coll. 1905-313 [printed white

label]; 1 ex S (MNHN, Fig. 30), Zanzibar [handwritten

white label], Sharp det., 1882 Neptosternus ornatus [prin-

ted white label, handwritten in part], Muséum Paris, coll.

F. Guignot [printed white label] 4 ex (ISNB), Zanzibar,

det. Sharp 1882.

- rotroui: lectotype 9 (MNHN, Fig. 31), designated

here: Algeria, Sidi-Bel-Abbès [printed white label], Nep-

tosternus Rotroui Pic, Peyerimhoff det. [printed white la-

bel, handwritten in part], Type [white label printed in red

letters], Museum Paris, coll. Rotrou [blue printed label,

handwritten in part].

- tropicus: holotype & (MRAC, Fig. 32), Congo Belge,

P.N.U., R. Lufira pied Mt Sombwe (700 m), 16.VII.1949,

Mis. G.F. de Witte.

- meridianus: holotype £' (NHML, Fig. 33), S. Africa,

Transvaal, R. Vaal, 23.VIII.1960, M. Chutter, N. meridia-

nus O.C., 4 holotype, M.E. Bacchus det. 1978 Holotype,

Holotype Brit. Mus. 1978-308; paratype 9 (NHML),

Transvaal, Vaal R., 10.XI.1959, M. Chutter, N. meridianus

n. sp. 9 allotype J.0.C., Allotype Brit. Mus. 1978-308.

- nigeriensis: holotype 4 (NHML, Fig. 34), Nigeria,

road to Enugo about 47 miles from Makurdi, 24.IV.1963,

Neptosternus nigeriensis 4 O0.C. holotype; paratype 9

(NHML), Nigeria, stream road Makurdi-Jos, 25.IIT.1963,

J.0.C., Neptosternus nigeriensis O.C. 9 allotype.

Additional material examined: Mali: Mopti, Niger

River, 14°30’N - 4°12’W, leg. Komarek & Mayer, 1 9

(NMW). Sudan: S. Darfur, Cafindibie, stagnant pool ri-

ver bed, V.1984, [leg.] H. Dodd, 17 ex (NHML, Fig. 35);

Shambat, 2.XI.1966, [leg.] P. Stys, 1 4 (NMP). Senegal:

Badi, 15.VIII-25.IX.1955, Mission IFAN au Parc National

du Niokolo Koba, coll. F. Guignot, 7 ex (MNHN, Fig. 36),

1 ex (ISNB), 1 ex (MRAC). Guinea Bissau: 5 km N. de

Madina-Salambande, 11-12.VII.2004, [leg.] A. Kudrna, 1

3 (CFP, Fig. 37). Burkina Faso: Boromo (Balé), VI.2005,

leg. Moretto, 7 ex (CSR). Ivory Coast: Divo, [leg.] J. De-

celler2sXIRl963%eltexX(CAB- Fig.733)Mrex3(MRAO);

Pare Nat. de la Comoé, 150 m, 12.VI.2000, [leg.] Rejn-

ties 1 S' (Fig. 39) e 1 2 (CFP). Nigeria: Kabba Prov.,

Kabba, 19-23.XI.1949, [leg.] B. Malkin, 3 ex (NHML);

Ilorin Prov., Ilorin, 15-18.X1.1949, [leg.] B. Malkin, 7 ex

(NHML, Fig. 40); Sabon Gida, 19.4.1978, [leg.] P. Beron,

16 e2 92 (NMW); Samaru, 13-20.VII.1970, [leg.] P. H.

Ward, light trap 5 ex (NHML); Virgioi (N. E.), Garkida,

18.VIII.1971, [leg.] D. J. Lawis, 1 4° (NHML). Centra-

frican Republic: Lobaye Pr., 50 km S. Boda, Ngoto vill.,

river Lobaye, 500 m, 7-9.XII—.2008, [leg.] A. Kudrna jr.,

1 £ (CFP); Mambere-Kadei, near Banga, 60 Km S. E.

Berberati, river Mambere, 400 m, 7-8.VI.2009, [leg.] A.

Kudrna jr., $ ex ? (CFP); Ombella-Mpoko Pr., Bangui-

Sibut road, 40 Km N Bangui, 380 m, 8-9.V.2009, [leg.]

A. Kudrna j.r, 2 92 ? (CFP); Sangha-Mbaere Prov., 20

Km S.Nola, 600 m, 13-14.XII.2008, [leg.] A. Kudrna

Jr., 1 9 (CFP). Democratic Republic of Congo: Lac Al-

bert, Kasenyi, 11.V-26.VI.1953, J. Verbeke - KEA, coll.

F. Guignot, 1 ex (MNHN); Lac Albert, Kasenyi, U.V.,

26.VI.1953, J. Verbeke - KEA, det. F. Guignot, 1956, 1

ex S' (Fig. 41) e 2 9 9 (ISNB). Tanzania: Ukiriguru, light

trap, 16.1.1960, [leg.] I. A. D. Robertson, 2 44 (NHML,

Fig. 42); Ukiriguru, light trap, 27.11.1960, [leg.] I. A. D.

Robertson, 1 ex (NHML); Zanzibar, N. ornatus, J. Bal-

four-Browne det., 2 ex (NHML). Mozambique: Prov. Ma-

nica, 45 km N. W. Chimoio, 14-16.X1II.2003, [leg.] A. Ku-

drna jr., 2 ex ? (CFP). South Africa: Kwazulu-Natal, S.W.

Magudu 27°34’S - 31°35° W, 4-5.1.2009, [leg.] P. Schiile,

1 4 (NMP); North West 50 km S. Kimberley Ritchie,

22.1.2000, [leg.] J. Halada, 1 9? (NMP); Skukuza, K. N. P,

31.1.1974, [leg.] H.& A. Braack, at light, 1 9 ? (NHML).

Madagascar: Mandrare Bas. (PO345), loc. Berohanga

près de Lotibe, Bezavo Riv., 46°36°07”E - 24°38’57”°S,

Alt. 550 m, 06.06.1994, leg. Orstom, Antananarivo, 1 4

and 1 ? (NMW).

Colour pattern of the upper side: colour and elytral

marks fairly variable. Type materials from Zanzibar and

Madagascar have reddish-brown head and pronotum, and

ground colour of elytra blackish-brown with reddish mar-

ks, so that the contrast of the two colours is rather poor.

Arrangement of elytral marks as in basic Group A. The

specimens from Tanzania, South Africa (as meridianus),

Democratic Republic of Congo, Sudan, Guinea Bissau,

Nigeria (as nigeriensis), Ivory Coast and Senegal have

head and pronotum testaceous to brown testaceous, and

ground colour of elytra brown with obvious testaceous

marks; the inner intermediate mark is elongate as a rule

and nearly attains the sub-basal and apical marks.

Size: L 3.37-3.85, w 1.84-2.04, L/w 1.81-1.89, Lmst/L

0.15-0.17, Lp 0.61-0.74.

10 ARMANDO BILARDO & SAVERIO ROCCHI

Body shape: sub-elliptical; variable: some specimens

are narrower, laterally constricted like nuperus, while

other specimens are very broad and rather abruptly nar-

rowed behind.

Male genitalia: penis in dorsal view somewhat wide-

ned and distally asymmetric; right side sinuate, apex bent

rightwards; left side from the maximum width onwards

nearly straight and gently bent to apex, tapering to a blunt

point. N. tropicus (holotype) shows little differences in

penis shape, i.e. more sinuate outline, right side more

twisted and apex more bent rightwards, but in spite of the

differences we consider it a junior synonym of ornatus.

Right paramere widened at apex as a rule, so that it appe-

ars sub-rectangular.

Distribution: Algeria, Mali, Sudan, Ethiopia, Senegal,

Guinea Bissau, Burkina Faso, Ivory Coast, Nigeria, Cen-

trafrican Republic, Democratic Republic of Congo, Tan-

zania, Mozambique, South Africa, Madagascar.

Remarks: in the description of N. alluaudi, Régimbart

(1903) questioned the occurrence of N. ornatus in Ma-

dagascar as reported by Sharp (1882), deeming that or-

natus only occurs in Zanzibar; the same doubt was put

later also by Zimmermann (1920), Guignot (1959a) and

Omer-Cooper (1970). In our opinion Sharp ’s statement is

trustworthy not only because the type material comprises

specimens from Madagascar, but also because as a result

of our examination of much material, ornatus is the most

widespread Neptosternus in Africa. However we believe

that the occurrence of ornatus in Madagascar and Zanzi-

bar needs further confirmation by new records. In spite of

minor differences in the elytral pattern, we believe that ro-

troui, tropicus, meridianus and nigeriensis belong to the

same variable species.

Neptosternus pederzanii Bilardo & Rocchi, 2010 (Figs.

43-44)

Neptosternus pederzanii Bilardo & Rocchi, 2010: 35.

Neptosternus (?) resartus Guignot: Pederzani & Rocchi

(1982: 82).

Type locality: Republic of Congo (Lague).

Type depository: CAB.

Type material examined: holotype (CAB, Fig. 43), pa-

ratypes 2 ex (CAB, Fig. 44), 13 ex (CFP), 1 9 (CSR):

Popular Republic of Congo, env. Lague, le long du fleuve

Mpama, à la lumière, 17.IV.1980, [leg.] G. Onore.

Additional material examined: Angola: Dundo, alt.

750 m, III.1948, à la lumière électrique, [leg.] A. de Bar-

ros Machado, 2 44 (NHML).

Colour pattern of the upper side: ground colour of

elytra dark-brown with reddish-testaceous markings, ar-

rangement of marks basically as in Group A but sub-basal

outer mark reduced to an expansion of the testaceous mar-

gin; in some specimens all marks are strongly reduced:

the outer and the apical marks look like an expansion of

the testaceous margin and the inner marks are just two

punctiform spots.

Size: L 2.62-3.00, w 1.54-1.75, L/w 1.72-1.88, Lmst/L

0.15-0.17, Lp 0.49-0.51.

Body shape: regularly oval.

Male genitalia: penis of peculiar shape: regularly wi-

dened from base to middle at the maximum width, hence

progressively narrowed to apex. Right side sinuate with

apex slightly bent rightwards; left side reaching apex

with a gentle bending; apex broad, prolonged into a blunt

point.

Distribution: Republic of Congo, Angola.

Neptosternus resartus Guignot, 1954 (Fig. 45)

Neptosternus resartus Guignot, 1954a: 7; Guignot (1954b:

32; 1955b: 182; 1959a: 606); Omer-Cooper (1970: 72);

Bilardo & Rocchi (2010: 44); Nilsson (2001: 256).

Type locality: Democratic Republic of Congo (Upem-

ba, Kaswabilenga).

Type depository: MRAC.

Type material examined: holotype 4 (MRAC, Fig.

45), Congo Belge, P.N.U., Kaswabilenga (700 m), 3-4.

IX.1947, Mis. G.F. de Witte (the holotype was examined

in past times: see Bilardo & Rocchi, 2010).

Additional material examined: none.

Colour pattern of the upper side: elytral marks regular-

ly arranged as in Group A.

Size: holotype L 2.99, w 1.71, L/w 1.75, Lmst/L 0.17,

Lp 0.60.

Body shape: regularly oval.

Male genitalia: penis peculiar for lacking apical wide-

ning, almost sub-parallel at the middle; right side rather

straight to three quarters of its length, hence sinuate and

hollowed at the right; left side shortly widened after base,

hence straight to apex; the latter narrowly rounded in a

blunt point.

Distribution: Democratic Republic of Congo.

Neptosternus sombuicus Guignot, 1954 (Figs. 46-47)

Neptosternus sombuicus Guignot, 1954: 6; Guignot

(1954b: 31; 1955b: 182; 1959a: 602); Omer-Cooper

(1970: 69); Nilsson & Persson (1993: 82); Nilsson

(2001582599?

Neptosternus psephotus Guignot, 1955a: 2; Omer-Cooper

(1970: 68); Nilsson (2001: 256); n. syn.

Neptosternus psephotes (sic !) Guignot: Bertrand & Le-

gros (1975: 681).

Type locality: - sombuicus: Democratic Republic of

Congo (Upemba, Riv. Lufira); - psephotus: Democratic

Republic of Congo (Garamba).

Type depository: - sombuicus MRAC; - psephotus

MRAC.

Type material examined: - sombuicus: holotype &

(MRAC, Fig. 46), Congo Belge, P.N.U., R. Lufira pied

Mt. Sombwe (700 m), 16.VII.1949, Mis. G.F. de Witte;

paratypes 7 ex (MRAC), same data as holotype.

- psephotus: holotype 4 (MRAC, Fig. 47), Congo Bel-

ge, Parc National de la Garamba, Mis. H. De Saeger, I/a/3,

16.XII.1949, Réc. Demoulin, 132; paratype 9 (MRAC),

same data as holotype.

Additional material examined: none.

Colour pattern of the upper side: head and prono-

tum reddish testaceous to light brown (in the holotype).

Ground colour of elytra dark brown; elytral marks reddish

testaceous; elytral marks regularly arranged as in Group

Size: L 3.77-4.04, w 2.13-2.19, L/w 1.76-1.84, Lmst/L

0.15-0.16, Lp 0.78.

A REVISION OF THE AFRICAN SPECIES OF THE GENUS NEPTOSTERNUS SHARP, 1882 (COLEOPTERA DYTISCIDAE) ll

Body shape: elongate oval.

Male genitalia: penis not much narrowed after basal

bulb, so that it appears less widened at the middle; right

side obviously sinuate, with rightwards bent apex; left

side after middle rather straight, hence gently curved to

apex, terminating in a long sharp point.

Distribution: Ethiopia, Democratic Republic of Con-

go.

Remarks: we cannot find any difference between N.

sombuicus and N. psephotus, except for the apex of penis,

that is broken and missing in psephotus, while it is entire

in sombuicus. We therefore consider psephotus a junior

synonym of sombuicus.

Neptosternus subopacus Guignot, 1956 (Fig. 48)

Neptosternus subopacus Guignot, 1956a: 253; Nilsson

(20015257):

Type locality: Democratic Republic of Congo (Tshua-

pa, Bokuma).

Type depository: MRAC.

Type material examined: holotype &' (MRAC, Fig.

48), Equateur: Bokuma, VII.1952, [leg.] R. P. Lootens.

Additional material examined: none.

Colour pattern of the upper side: head and pronotum

testaceous; ground colour of elytra light-brown, elytral

marks testaceous, regularly arranged as in Group A. Up-

perside dull. As far as we know, this is the only African

Neptosternus with obviously microsculptured elytra.

Size: holotype L 3.43, w 1.93, L/w 1.78, Lmst/L 0.17,

Lp 0.70.

Body shape: sides of elytra rather sub-parallel in the

basal two thirds, hence abruptly narrowed to apex.

Male genitalia: penis of peculiar shape: widened from

base to the maximum width after the middle, but not very

broad; right side sinuate with rightwards bent apex; left

side regularly attaining apex in a gentle curve; apex bro-

ad, obviously turned to the right in a sort of hook, with

broadly rounded tip.

Distribution: Democratic Republic of Congo.

Neptosternus tricuspis Guignot, 1954 (Fig. 49)

Neptosternus tricuspis Guignot, 1954a: 6; Guignot

(1954b: 30; 1955b: 182; 1956b: 322; 1959a: 602);

Omer-Cooper (1970: 72); Nilsson (2001: 257).

Type locality: Democratic Republic of Congo (Upem-

ba, Riv. Lufira).

Type depository: ISNB.

Type material examined: paratypes 73 ex (ISNB, Fig.

49), Congo Belge, P.N.U., R. Lufira pied mt. Sombwe

(700 m), 16.VII.1949, Mis. G.F. de Witte. Holotype not

seen.

Additional material examined: Angola: Dun-

do, 17.VIII.1953, [leg.] L. Carvalho, à la lumière, 1 Sd

(CFP).

Colour pattern of the upper side: head and pronotum

testaceous; ground colour of elytra dark-brown; elytral

marks testaceous, regularly arranged as in Group A.

Size: paratype shown in the photo L 3.29, w 1.85, L/w

1.77, Lmst/L 0.17, Lp 0.71; other specimens L 3.21-3.52,

w 1.76-1.92, L/w 1.77-1.83, Lmst/L 0.17.

Body shape: regularly oval.

Male genitalia: penis not much narrowed after ba-

sal bulb, so that it appears less widened at the middle;

right side hardly sinuate, straight at apex; left side almost

straight after the middle, then attaining apex in a gentle

curve; tip broadly rounded.

Distribution: Democratic Republic of Congo, Angola.

Neptosternus bartolozzii n. sp. (Figs. 50-53)

Neptosternus “specie indeterminata A” Bilardo & Rocchi

(2010: 46; 2011: 195).

Neptosternus nuperus Guignot: Bilardo & Rocchi (1999:

232, misidentification).

Type locality: Gabon, M.ts Cristal, Asseng Assala.

Type depository: MSNM.

Type material: holotype 4 (MSNM, Fig. 50), Gabon,

M.ts Cristal, Asseng Assala, 10.VIII.2006, leg. AB; para-

types (same data as holotype) 1 ex 9 (MSNM, Fig. 51), 5

ex (CAB), 20 ex (CSR); Gabon, M.ts Cristal, Andok Fou-

la, 21.VIII.2006, leg. AB, 1 ex (CAB); Gabon, M..ts Cris-

tal, Kinguélé, 6.VIII.1973, leg. AB, 2 ex (CAB); Gabon,

Réserve Lopé Okanda, 5.VIII.19$3, leg. AB, 7 ex (CAB),

2 ex (CSR); Gabon, Franceville, vill. Mopia, riv. Ikengué,

29.VI- 1.VI.2010, leg. AB, 12 ex (CAB, Fig. 52), 21 ex

(CSR), 2 ex (NHML), 2 ex (MNHN), 2 ex (MRAC), 2

ex (ISNB), 2 ex (NMW,), 2 ex (NMP); Gabon, Makokou,

24.VIII.1987, leg. AB, 3 ex (CAB, Fig. 53); Gabon, Fran-

ceville, Bakoumba, Parc de Lékédi, 27.VIII.2008, leg.

AB, 6 ex (CAB), 3 ex (CSR); Gabon, Franceville, vill.

Mougouango, Fleuve Ogooué, 23.VIII.1992, leg. AB, 5

ex (CAB), 2 ex (CSR); Central African Rep., Mambere-

Kadei, near Banga, 60 Km SE Berberati, river Mambere,

400 m, 7-8.V1.2009, [leg.] A. Kudrna jr., 1 ex (CFP).

Additional material examined: 1 ex 9 (CSR), Democra-

tic Republic of Congo, Ituri, Epulu, m 600, 17.VIII.1984,

leg. G. Carpaneto; it cannot be identified with certainty,

so it was not included in paratypes.

Diagnosis: this species belongs to group A; it can be

distinguished by the following combination of characters:

small size, sub-elliptical body outline from the base of

antennae to the extremity of elytra, broadest behind the

sub-basal band of elytron and regularly tapering in front

and behind, pronotum with anterior and posterior borders

dark-brown. Dorsal surface shiny, without detectable mi-

crosculpture.

Head: reddish.

Pronotum: reddish with anterior and posterior borders

of the same brown colour as the ground color of elytra.

Elytra: ground colour light-brown to dark-brown or

piceous, colour pattern typical of the group, but with inner

median mark reduced or missing.

Prosternum: distinctly keeled from base to the apex of

the median spine.

Underside: colour reddish to reddish-brown.

Size: holotype L 2.80, w 1.51, L/w 1.85, Lmst 0.44,

Lmst/L 0.16, Lp 0.60; paratypes.L 2.68-3.03, w 1.44-1.70,

L/w 1.77-1.90, Lmst/L 0.15-0.17, Lp 0.60-0.61.

Male genitalia: penis in dorsal view sinuate, distally

slightly asymmetric; right side sinuate with apex slightly

turned rightwards; left side from the maximum width on-

wards almost straight converging to apex; tip pointed.

Derivatio nominis: this species is named in honour of

12 ARMANDO BILARDO & SAVERIO ROCCHI

Dr. Luca Bartolozzi, chief of the the Zoology Section “La

Specola” of the Natural History Museum of the Universi-

ty of Florence, for the kind co-operation and continuous

help in our entomological work.

Distribution: Gabon, Central African Republic, De-

mocratic Republic of Congo (?).

Group B

Species with .elytral pattern arranged as in Scheme B

Neptosternus africanus Peschet, 1916 (Figs. 54-56)

Neptosternus africanus Peschet, 1916: 316; Legros (1949:

53; 1954: 269; 1958: 211); Guignot (1954b: 29; 1959a:

604); Omer-Cooper (1957: 22; 1958: 52; 1970: 77);

Nilsson (2001: 253).

Type locality: Republic of Congo (Fort Crampel).

Type depository: MNHN.

Type material examined: lectotype £ (MNHN, Fig.

54), designated here: Fort-Crampel, Congo Frangais

[printed white label], Museum Paris 1946, Coll. R. Pe-

schet [printed blue label], Type [printed red label0000],

Neptosternus africanus R. Peschet [handwritten white la-

bel]. In the description, Peschet (1916) mentioned also the

following specimens from the formerly French Congo,

that were not traced in MNHN: 1 ex Quagga (Ovahiné), 1

ex Bahr-Sara, près le confluent de la Tala, 1 ex bassin du

Chari, rivière Gribingui.

Additional material examined: Chad: Logoneufer b.

Lai, Tschadgebiet, AEF., leg. Franz, 1 ex (NMW). Su-

dan: Soudan égyptien, Région de Sennar, 1906, coll. Gui-

gnot, 1 £ (MNHN, Fig. 55), Guinea: Nimba (Guinée),

Yalanzou, M. Lamotte, II-VI 42 (1942), N. africanus, C.

Legros det., 1 ex 9 ? (MNHN). Centrafrican Republic:

Mambere-Kadei, near Banga, 60 Km S. E. Berberati, ri-

ver Mambere, 400 m, 7-8.VI.2009, [leg.] A. Kudrna jr., 2

®© (CFP). Democratic Republic of Congo: Congo Bel-

ge, P.N.U. Kaswabilenga (700 m), 16-X-1947, Mis. G.

F. De Witte, coll. Guignot, 1 4 (MNHN, Fig. 56); Fa-

radje, Aba, 22.1V.1930, [leg.] A. Collart, N. africanus,

det. Gschwendtner,1 ex ? (ISNB). Kenya: Kakamega fo-

rest N.R., 1600 m, 21.IX-10.X.2005, [leg.] J. Holstein &

D. Bartsch, 1 9 ? (NMP). Angola: District de Huilla de

Cutchy au Cuelei, Mission Rohan-Chabot, 1914, janvier,

N. africanus, R. Peschet det. 1924, 1 &° (MNHN). Un-

known country (possibly Democratic Republic of Con-

go): Palanga Dizi, 16.VII.1926, [leg.] A. Collart, N. afri-

canus, R. Peschet det., 1 ex ? (ISNB). Unknown country:

Kai Baku, 8.VII.1926, [leg.] A. Collart, N. africanus, det.

Gschwendtner, 1 ex ? (ISNB).

Colour pattern of the upper side: head and pronotum

reddish testaceous, pronotum with anterior and posterior

borders of the same brown colour as the ground colour of

elytra; ground colour of elytra brown; elytra marks testa-

ceous; Juxtasutural testaceous line broken in some speci-

mens, additional elongate mark in the middle of elytron

(sensu Omer-Cooper, 1970) always detectable.

Prosternum: slightly keeled at base; transversely con-

vex (not keeled) or almost flattened at middle.

Size: L 2.91-3.56, w 1.67-1.91, L/w 1.74-1.90, Lmst/L

0.15-0.19 (this is the only African species showing such a

wide range of variability in size), Lp 0.63-0.64.

Body shape: regularly oval.

Male genitalia: penis symmetrical in the distal part and

apex rounded.

Distribution: Chad, Sudan, Guinea, Cameroun, Cen-

trafrican Republic, Republic of Congo, Democratic Repu-

blic of Congo, Kenya, Angola, Malawi, Zimbabwe, South

Africa.

Neptosternus compsus Guignot, 1953 (Figs. 57-60)

Neptosternus compsus Guignot, 1953: 2; Guignot (1954b:

29; 1959a: 605); Omer-Cooper (1970: 78); Bilardo &

Rocchi (1990: 178; 2010: 42); Nilsson (2001: 254).

Type locality: Democratic Republic of Congo (Upem-

ba, Kaswabilenga).

Type depository: MRAC.

Type material examined: holotype &° (MRAC, Fig.

57), Congo Belge, P.N.U., Kaswabilenga (700 m), 14-

25.X.1947, Mis. G.F. de Witte; paratypes: 4 ex (ISNB),

5 ex (MRAC), Congo Belge, P.N.U., Kaswabilenga (700

m), 14-25.X.1947, Mis. G.F. de Witte; 2 ex (MRAC, Fig.

58), 1 ex (ISNB, Fig. 59), same data but collection da-

te 3-4.XI.1947. [the latter specimen, lodged in ISNB, is

pinned with a printed red label “paratype” and a printed/

handwritten red label “F. Guignot det., 1953 Neptosternus

nuperus n. sp. ”’; it is unboutedly a N. compsus, as it was al-

ready ascertained by Omer-Cooper (1970), and is obviou-

sly mislabelled]; 1 ex (ISNB), Congo Belge, P.N.U., Ka-

swabilenga r. dr. Lufira (700 m), 8.XI.1947, Mis. G.F. de

Witte; 3 ex (ISNB), 4 ex (MRAC), Congo Belge, P.N.U.,

Lukawe affl. r. dr. Lufira (700 m), 6-9.X.1947, Mis. G.F.

de Witte.

Additional material examined: Nigeria: Samaru, 13-

20.V1II.1970, [leg.] P. H. Ward, light trap, 1 ? (NHML).

Gabon: Belinga, 18.3.1963, [leg.] H.Coiffait, 4 ex

(NMW); Makokou, 25.5.1963, [leg.] H.Coiffait, 1 ex

(NMW); Makokou, 1-14. V.1971, [leg.] J. Mateu, 2 ex

(NMP); Makokou, 23.VII.1987, leg. AB, 2 ex (CAB);

Ntoum, 1.VIII.1973, leg. AB & FP, 2 ex (CFP). Centra-

frican Republic: Nana-Mambere Pr., 5 km N.E. Baoro, 60

km S.E. Bouar, 850 m, 31.V-3.VI.2009, [leg.] A. Kudrna

jr., 1 6 (CFP); Mambere-Kadei, near Banga, 60 Km S.

E. Berberati, river Mambere, 400 m, 7-8.VI.2009, [leg.]

A. Kudrna jr., 7 ex (CFP); Nana-Mambere Prov., 20-60

km W Bouar, 650 m, 25-27.III.2010, [leg.] A. Kudrna jr.,

19 ex (CFP, CAB, CSR, Fig. 60). Angola: Dundo, 10-13.

VIII.1953, [leg.] L. Carvalho, à la lumiére, 2 ex (CAB),

16 e 1 9 (CFP). Mozambique: Prov. Tete, ca. 40 km S.

W. Tsangano, 7-8.XII.2005, [leg.] A. Kudrna jr., 14 e 1

2 (CFP). Zimbabwe: South Rhodesia, lower Pungwe Ri-

ver, 20ml. S. Mt. Inyangani, VII-IX.1958, [leg.] R. Wat-

mough, 2 99 (NHML).

Colour pattern of the upper side: head and pronotum

testaceous, pronotum with anterior and posterior borders

of the same brown colour as the ground colour of elytra;

ground colour of elytra brown; elytra marks testaceous;

Juxtasutural testaceous line not attaining the inner sub-ba-

sal mark as a rule; additional elongate mark in the middle

of elytron (sensu Omer-Cooper, 1970) always detectable.

Prosternum: holotype without prosternal keel; in some

paratypes (from the same locality and date) prosternum

distinctly keeled at base, transversely convex (not keeled)

behind, and flattened towards apex of the median spine;

some specimens from Makokou (Gabon) have the proster-

A REVISION OF THE AFRICAN SPECIES OF THE GENUS NEPTOSTERNUS SHARP, 1882 (COLEOPTERA DYTISCIDAE) 13

num distinctly keeled its whole length, while specimens

from Angola lack prosternal keel.

Size: holotype L 2.85, w 1.65, L/w 1.73, Lmst/L 0.22,

Lp 0.55; other ex L 2.60-2.90, w 1.45-1.66, L/w 1.71-1.87

Lmst/L 0.22-0.25.

Body shape: regularly oval.

Male genitalia: penis distally much widened, fai-

rly symmetrical, apex pointed and gently turned

rightwards.

Distribution: Nigeria, Gabon, Centrafrican Republic,

Democratic Republic of Congo, Angola, Mozambique,

Zimbabwe, South Africa.

Remarks: prosternum not keeled as a rule, except for

some specimens with trace of a keel, however all speci-

mens from Centrafrican Republic have more or less kee-

led prosternum. We have chosen to attach little importan-

ce to geographical variations, being not accompanied by

other taxonomic differences. In any case N. compsus is

easy to identify as it differs from other small-sized species

by the peculiar length of mesotarsi and the broad distal

part of penis.

As said before under the treatment of N. nuperus, the

paratype of N. nuperus lent for study by ISNB is not that

species but N. compsus, as already noted by Omer-Cooper

(1970: 64); its prosternum is not keeled and the penis is

the same as in compsus; also size (L 2.82, w 1.60, L/w

1.76, Lmst/L 0.24, Lp 0.53) and habitus agree with the

features of compsus (see Fig. 59).

Neptosternus distinctus Omer-Cooper, 1970 (Fig. 61)

Neptosternus distinctus Omer-Cooper, 1970: 74; Nilsson

(2001: 254).

Type locality: Nigeria (R. Assob about 36 miles from

Jos).

Type depository: NHML.

Type material examined: holotype 4° (NHML, Fig.

61), Nigeria, R. Assob, nr. Jos, 13.1V.1963, J.0.C., Nep-

tosternus &4' distinctus J.0.C., M.E. Bacchus det. 1978,

Holotype, Brit. Mus. 1978-308.

Additional material examined: none.

Colour pattern of the upper side: head and pronotum

testaceous, pronotum with anterior and posterior borders

of the same brown colour as the ground colour of elytra;

ground colour of elytra brown; elytral marks testaceous;

Juxtasutural testaceous line attaining the inner sub-basal

mark; additional elongate mark in the middle of elytron

(sensu Omer-Cooper, 1970) fairly evident.

Diecalt597;iwalaZ7y1L/wal:90}LmstL.0319%Lps1:01

Body shape: regularly oval.

Male genitalia: penis peculiar for its large size; right

side sinuate, deeply hollowed after the middle; left side

progressively tapering to sharply pointed apex.

Distribution: Nigeria.

Neptosternus pumicatus Guignot, 1949 (Figs. 62-64)

Neptosternus pumicatus Guignot, 1949: 14; Guignot

(1959a: 601); Omer-Cooper (1958: 52; 1967: 60;

1970: 78); Franciscolo (1986: 29); Nilsson (2001:

256); Bilardo & Rocchi (2010: 40).

Neptosternus ‘specie indeterminata B ”’ (unidentified spe-

cies B) Bilardo & Rocchi (2010: 47; 2011: 195).

Type locality: Democratic Republic of Congo (Lusen-

gl).

Type depository: ISNB.

Type material examined: lectotype &' (ISNB, Fig. 62),

designated here: Lusengi, 11.IV.1926 [printed white la-

bel], Type 4 [handwritten red label], Dr. F. Guignot det.,

1949 Neptosternus pumicatus Guign. Type [printed white

label, hanwritten in part]; paralectotype 1 ex (ISNB, Fig.

63), same data as lectotype.

Additional material examined: Gabon: M.ts Cristal,

Andok Foula, 21.VIII.2006, leg. AB, 1 ex (CAB, Fig. 64).

Angola: 7 mls. W. Gabela at ligth, 16-18.III.1972 (no col-

lector name), 1 ex (CFP).

Colour pattern of the upper side: head and pronotum

testaceous, pronotum with anterior and posterior bor-

ders of the same brown colour as the ground colour of

elytra; ground colour of elytra brown; elytral markings

testaceous; juxtasutural testaceous line not attaining the

inner sub-basal mark as a rule; the additional mark in the

middle of elytron (sensu Omer-Cooper, 1970) in lectot-

ype is reduced to a short dash on the left elytron whi-

le it is hardly visible as a punctiform spot on the right

elytron (see Omer-Cooper, l. c.: 64) and in paralectotype

it is punctiform on both elytra. The specimen from Ga-

bon has no additional marks at all, but its identification

is based on the feature of aedeagus only, in accordance

with Omer-Cooper (1070: 64) “the presence of this extra

longitudinal testaceus line on the elytra is of taxonomic

importance although its absence, if not accompanied by

an aedeagal difference, would not necessarily invalidate

the species”.

Size: lectotype and paralectotype L 2.81-3.01, w 1.53-

1.61, L/w 1.84-1.87, Lmst/L 0.19-0.20, Lp 0.57.

Body shape: oblong oval, rather compressed at sides.

Male genitalia: penis distally asymmetric, with

rounded apex, like africanus Peschet, 1916.

Distribution: Sierra Leone, Nigeria, Gabon, Democra-

tic Republic of Congo, Angola, Zimbabwe, South Africa.

Neptosternus simulator Omer-Cooper, 1970 (Figs. 65-66)

Neptosternus simulator Omer-Cooper, 1970: 75; Nilsson

(2001257):

Neptosternus africanus Peschet: Omer-Cooper (1957: 22;

1958: 52; misidentification).

Neptosternus resartus Guignot: Omer-Cooper (1965: 90,

misidentification).

Type locality: Rhodesia [Zimbabwe] (River Inganyi,

38 miles from Salisbury).

Type depository: NHML.

Type material examined: paratype 4 (NHML, Fig.

65), Transvaal, Waterberg dist., Poer Zyn R., 19.8.1948,

N. simulator n. sp. 4° type, J.0.C., M.E. Bacchus det.

1978 Holotype, Holotype Brit. Mus. 1978-308 [the label

“Holotype” is wrong: This specimen from Transvaal is a

paratype because the true holotype should be from Rho-

desia, as stated by Omer-Cooper (1970: 75)]; Paratype 9

(NHML), same data, 19.August.[19]48, J.O.C., N. simu-

lator n. sp. 9 allotype, Allotype, Brit. Mus. 1978-308.

Additional material examined: Centrafrican Republic:

Nana-Mambere Prov., 20-60 km W Bouar, 650 m, 25-27.

III.2010, [leg.] A. Kudrna jr., 53 ex (CFP, CAB, CSR, Fig.

66).

14 ARMANDO BILARDO & SAVERIO ROCCHI

Colour pattern of the upper side: head and prono-

tum testaceous, pronotum with anterior and posterior

borders of the same colour as the ground colour of

elytra; ground colour of elytra brown; elytral mar-

kings testaceous; juxtasutural line entire and not

connected with other marks; additional mark in the

middle of elytron (sensu Omer-Cooper, 1970) fairly

evident.

Size: paratype 4 L 3.15, w 1.70, L/w 1.86, Lmst/L

0.19, Lp 0.60; paratype 9 L 3.04, w 1.73, L/w 1.76, other

ex L 2.91-3.21, w 1.67-1.81, L/w 1.73-1.83, Lmst/L 0.21-

0.24.

Body shape: oval, slightly compressed at sides.

Male genitalia: penis in dorsal view distally widened

and asymmetrical: right side sinuate with apex slightly

turned rightwards; left side from the maximum width on-

wards almost straight, progressively attaining apex in a

gentle curve; tip elongate and pointed.

Distribution: Centrafrican Republic, Malawi, Zimbab-

we, South Africa.

Remarks: at first sight this species closely resembles

africanus Peschet, but it differs by longer mesotarsi and

above all by the pointed apex of penis, whereas in africa-

nus the apex of penis is rounded.

Neptosternus komareki n. sp. (Figs. 67-68)

Type locality: Benin, Dep. Platuau, Pobé near Crabé

(village), 7.11.2006, 06°57’24,2°’N - 02°40°28,2”E, 10 m

a.s.1. tributary to Idi river.

Type depository: NMW

Type material: holotype 4 (NMW, Fig. 67), Be-

nin, Dep. Platuau, Pobé near Crabé (village), 7.1I.2006,

06°57°24,2”N - 02°40°28,2” E, 10ma.s.l. tributary to Idi

river, leg. Goergen, Komarek & Houngué; paratypes: 16

(NMW), same data as holotype; Gabon, Parc Nat. M.ts de

Cristal, Kinguélé, riv. Bimvan, à la lumière, 31.V.2012,

leg. AB, 254, 19 (CAB, CSR, Fig. 68); Gabon, M.ts de

Cristal, Vill. Asseng Assala, 1 km route pour la riv. No-

ya, à la lumière, 6-8.VI.2012, leg. AB, 243, 19 (CAB,

CSR).

Diagnosis: this species belongs to group B as for

colour pattern of elytra; it can be distinguished by the

following combination of characters: small size, sub-

elliptical body outline, continuous from the base of an-

tennae to the extremity of elytra, maximum width at the

middle, low ratio L/w, that gives the body a wide and

short outline.

Head: reddish.

Pronotum: reddish, with anterior and posterior borders

of the same colour as the ground colour of elytra.

Elytra: ground colour of elytra dark-brown; elytral

marks-brown-testaceous, arranged as in group B; juxta-

sutural line entire and almost attaining the apical mark;

additional mark in the middle of elytron (sensu Omer-

Cooper, 1970) reduced to a short stripe but fairly evident

on both elytra.

Prosternum: distinctly keeled from base to the apex of

the median spine.

Underside: dark-brown.

Size: L 2.63-2.90, w 1.57-1.67, L/w 1.67-1.75, Lmst/L

0.18-0.19 (holotype L 2.75, w 1.54, L/w 1.79, Lmst/L

0.19, Lp 0.56).

Body shape: sub-elliptical.

Male genitalia: penis distally symmetrical with apex

pointed, very much ressembling that of /angoensis Bilar-

do & Rocchi, 2010.

Derivatio nominis: this species is named in honour

of one of the collectors, Dr. Albrecht Komarek (Vienna),

specialist of Coleoptera Hydrophilidae.

Distribution: Benin, Gabon.

Neptosternus longitarsis n. sp. (Figs. 69-70)

Type locality: Gabon, Franceville, vill. Mopia, petite

riv. alt. 446 m, S 01°54,522”, E 013°38,182°.

Type depository: CAB.

Type material: holotype 4 (CAB, Fig. 69), Ga-

bon, Franceville, vill. Mopia, petite riv. alt. 446 m, S

01°54,522°, E 013°38,182”, 1.VII.2010, leg. AB; para-

types: Gabon, Parc National des Plateaux Batéké, Camp

Ntsa, forét dense, 8-13.IX.2008, à la lumière, leg. AB,

2 92 (CAB, Fig. 70 e CSR); Gabon, Parc Nat. M.ts de

Cristal, Kinguélé, riv Bimvan, à la lumière, 31.V.2012,

leg. AB, 19 (CAB); M.ts de Cristal, Vill. Asseng Assala,

1 km route pour la riv. Noya, à la lumière, 6-8.VI.2012,

leg. AB, 16 (CAB).

Diagnosis: this species belongs to group B as for co-

lour pattern of elytra; it can be distinguished by its small

size and above all by the peculiar length of mesotarsome-

res (mesotarsi without claws).

Head: testaceous.

Pronotum: narrowly testaceous at the middle, with

broad anterior and posterior borders of the same colour as

the ground colour of elytra.

Elytra: ground colour almost black; elytral marks te-

staceous, placed as usual in group B, but with a somewhat

peculiar arrangement (see photo in Figs. 69-70); additio-

nal mark in the middle of elytron (sensu Omer-Cooper,

1970) fairly obvious on both elytra. One of paratypes has

an additional small irregular spot between the juxtasutural

line and. the outer median mark.

Prosternum: distinctly keeled from base to the apex of

the median spine.

Underside: reddish-brown.

Size: L 2.64-3.06, w 1.48-1.62, L/w 1.78-1.88, Lmst/L

0.22-0.24 (holotype L 2.80, w 1.54, L/w 1.82, Lmst/L

0.22, Lp 0.54).

Body shape: regularly oval.

Male genitalia: penis almost symmetrical; in dorsal

view with an obvious distal widening; right and left sides

symmetric in relation to the median axis, and regularly

tapering to the rounded apex.

Derivatio nominis: the species is named in reference to

the anomalous length of mesotarsi.

Distribution: Gabon.

Neptosternus maliensis n. sp. (Figs. 71-72)

Type locality: Mali, 100 km W. Sikasso, Bagoé river,

LIS27ENI 6835 WI

Type depository: NMW.

Type material: holotype &'° (NMW, Fig. 71), Mali,

100 km W. Sikasso, Bagoé river, 11° 27° N, 6° 35° W,

24.2.2000, leg. Komarek & Mayer; paratypes (same data

A REVISION OF THE AFRICAN SPECIES OF THE GENUS NEPTOSTERNUS SHARP, 1882 (COLEOPTERA DYTISCIDAE) 15

as holotype): 6 54 and 11 99 (NMW, Fig. 72), 1 4 and

19 (NMP).

Diagnosis: this species belongs to group B as for the

colour pattern of elytra; it can be distinguished by the

moderate size and the sub-parallel sides of body, abruptly

narrowed behind.

Head: reddish tetaceous.

Pronotum: reddish testaceous with anterior and poste-

rior borders of the same colour as the ground colour of

elytra.

Elytra: ground colour brown; marks brown-testaceous

arranged as in Scheme B, all of them very obvious; sub-

basal marks separate, juxtasutural line continuous and

well defined; additional mark in the middle of elytron

(sensu Omer-Cooper, 1970) fairly evident.

Prosternum: distinetly keeled only at base.

Underside: light-brown.

Size: L 3.06-3.32, w 1.63-1.77, L/w 1.82-1.92, Lmst/L

0.21-0.22, Lp 0.64-0.66 (holotype: L 3.12, w 1.63, L/w

1.91, Lmst/L 0.21, Lp 0.66).

Body shape: elongate, with sub-parallel sides, abrupt-

ly tapering to apex.

Male genitalia: penis almost symmetrical, in dorsal

view slightly narrowed between basal bulb and the distal

part, where it is widened, with right and left sides almost

symmetric in relation to the median axis, and regularly

tapering to the rounded apex.

Derivatio nominis: the species is named in reference to

the country where it was collected.

Distribution: Mali.

Group C

Species with elytral pattern arranged as in Scheme C

Neptosternus batekensis Bilardo & Rocchi, 2010 (Figs.

73-74)

Neptosternus batekensis Bilardo & Rocchi, 2010: 37

Type locality: Gabon (National Park of Plateaux Ba-

téké, Camp Ntsa).

Type depository: MSNM.

Type material examined: holotype 4° (MSNM), Ga-

bon, Parc National des Plateaux Batéké, Camp Ntsa, forét

dense, 8-13.IX.2008, à la lumière, leg. AB; paratypes

(same data as holotype): 4 ex (CAB, Figs. 73-74), 5 ex

(CSR); 1 ex (CAB), Gabon, Franceville, vill. Mopia, riv.

Ikengué, 29.VI-1.VII.2010, leg. AB.

Additional material examined: none.

Colour pattern of the upper side: head, pronotum and

ground colour of elytra black, elytral marks testaceous,

arranged as in Scheme C, with a small spot in the middle

of elytron.

Size: L 2.63-2.71, w 1.48-1.51, L/w 1.75-1.81, Lmst/L

0.20, Lp 0.54.

Body shape: regularly oval.

Male genitalia: penis with distal part symmetrical and

apex rounded.

Distribution: Gabon.

Remarks: N. batekensis and N. susinii Bilardo & Roc-

chi, 2010 are the only African species with head entire-

ly black; besides in batekensis the pronotum is entirely

black, while in susinii it shows a narrow median band

dark-testaceous.

Neptosternus langoensis Bilardo & Rocchi, 2010

(Figs. 75-76)

Neptosternus langoensis Bilardo & Rocchi, 2010:

Type locality: Republic of Congo (National Park

of Odzala, Mboko).

Type depository: CAB.

Type material examined: holotype &' (CAB, Fig.

75), République du Congo, Parc Nat. d’Odzala,

Mboko, riv. Lango, 21.VIII.2002, leg. AB.

Additional material examined: Centrafrican Re-

public: Mambere-Kadei, Carnot-Gadzi road, 15 km

E. Carnot, 600 m, 5-6.VI.2009, A. Kudrna jr., 2 99

UGSB#F19-/63 CEP):

Colour pattern of the upper side: head brown-te-

staceous; pronotum and ground colour of elytra pi-

ceous; elytral marks testaceous arranged as in group

Size: holotype L 2.70, w 1.50, L/w 1.80, Lmst/L

UslOminz0i53%

Body shape: regularly oval.

Male genitalia: penis with distal part symmetrical

and apex rounded, slightly bent rightwards.

Distribution: Republic of Congo, Centrafrican

Republic.

Remarks: the two ex 9 9 designated as paratypes

of this species in Bilardo & Rocchi (2010), possibly

belong to a different species; at the moment we do

not describe that taxon in expectation of a male spe-

cimen.

Neptosternus susinii Bilardo & Rocchi, 2010 (Figs.

77-78)

Neptosternus susinii Bilardo & Rocchi, 2010: 38

Type locality: Gabon (National Park of Plateaux

Batéké, Camp Ntsa).

Type depository: MSNM.

Type material examined: holotype 4° (MSNM),

Gabon, Parc National des Plateaux Batéké, Camp

Ntsa, forét dense, 8-13.IX.2008, à la lumiére, leg.

AB; paratypes (same data as holotype): 15 ex (CAB,

Figs. 77-78), 9 ex (CSR); 1 ex (CAB), Gabon, Parc

National des Plateaux Batéké, Osséré, 4-6.IX.2008,

à la lumiéère, leg. AB.

Colour pattern of the upper side: this species

closely resembles N. batekensis Bilardo & Rocchi,

2010; head, pronotum and ground colour of elytra

pitch-black; in a few specimens pronotum with a

narrow median band dark-testaceous; elytral marks

testaceous, arranged as in group C but somewhat

peculiar (see photo Figs. 77-78); inner and inter-

mediate marks of the median band sometimes mis-

sing.

Size:21002:/9-3:0410w01:51=1:68- L/w=12/6-1.85,

Lmst/L 0.19-0.21, Lp 0.54.

Body shape: regularly oval, but more elongated

than in batekensis and laterally compressed.

Male genitalia: penis with distal part broader that

in batekensis, and with rounded apex.

Distribution: Gabon.

ARMANDO BILARDO & SAVERIO ROCCHI

)

0.5 mm

Figs. 3-5 - 3) N. alluaudi Régimbart, 1903, lectotypus 9 (MNHN), Madagascar Centre-Sud: habitus. 4) N. a/luaudi Régimbart, 1903,

ex £ (NMP), N. Madagascar, Antseranana distr., Sambirana riv., Marovato: a) habitus; b) penis dorsal view and inner face of parameres

/ pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista dorsale; d) penis dorsal-right

lateral view / pene in vista dorsale-destra laterale; e) penis lateral view / pene in vista laterale. 5) N. silvester Guignot, 1960 (new

synonym), ex £' (MNHN), Madagascar, Andobo 190 m, forét Antsingy: a) habitus: b) penis dorsal view and inner face of parameres /

pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista dorsale; d) penis lateral view

/ pene in vista laterale.

0.5 mm

A REVISION OF THE AFRICAN SPECIES OF THE GENUS NEPTOSTERNUS SHARP, 1882 (COLEOPTERA DYTISCIDAE)

0.5 mm

2 mm

ala 0.5 mm

file

2 mm 0.5 mm

8a 8b

Figs. 6-8 - 6) N. fasciatus Omer-Cooper, 1970, holotypus 4 (NHML), Nigeria, stream crossing the road from Enugo to Makurdi: a)

habitus; b) penis dorsal view and inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal

view / apice del pene in vista dorsale. 7) N. fasciatus Omer-Cooper, 1970, ex 4 (CAB), Gabon, Franceville, vill. Mopia, riv. Ikengué:

a) habitus; b) penis dorsal view and inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal

view. 8) N. fasciatus Omer-Cooper, 1970, ex 4 (NMW), Gabon, Makokovu: a) habitus; b) penis dorsal view and inner face of parameres

/ pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista dorsale.

ARMANDO BILARDO & SAVERIO ROCCHI

2 mm

0.5 mm

2 mm

10c

10a 10b

0.5 mm

2 mm

lle

lla llb

Figs. 9-11 - 9) N. fasciatus Omer-Cooper, 1970, ex 4 (CAB), Gabon, Mandiji: a) habitus; b) penis dorsal view and inner face of para-

meres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista dorsale. 10) N. fasciatus

Omer-Cooper, 1970, ex £ (CAB), Gabon, Kinguélé: a) habitus; b) penis dorsal view and inner face of parameres / pene in vista dorsale

e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista dorsale. 11) N. fasciatus Omer-Cooper, 1970, ex £

(CAB), Gabon, Ntoum: a) habitus; b) penis dorsal view and inner face of parameres / pene in vista dorsale e faccia interna dei parameri;

c) apex of penis dorsal view / apice del pene in vista dorsale.

A REVISION OF THE AFRICAN SPECIES OF THE GENUS NEPTOSTERNUS SHARP, 1882 (COLEOPTERA DYTISCIDAE) I 9

0.5mm

2 mm Ù x

ti IDG

2a 12b

0.5 mm

2 mm 130

13a 13b

0.5 mm

2 mm

14c È

14a 14b

Figs. 12-14 - 12-13) N. fasciatus Omer-Cooper, 1970, ex 4 (CAB), Gabon, Réserve Lopé Okanda: a) habitus; b) penis dorsal view

and inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista

dorsale. 14) N. nuperus Guignot, 1954, ex £ (CAB), Cameroun, Kribi: a) habitus; b) penis dorsal view and inner face of parameres /

pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista dorsale.

ARMANDO BILARDO & SAVERIO ROCCHI

0.5 mm

2mm

15c

15b

2 mm il 0.5 mm

16c

16b

2 mm 0.5 mm

L7C

7a 17b

Figs. 15-17 - 15) N. nuperus Guignot, 1954, ex 4 (CAB), Gabon, Franceville, Mopia, riv. Ikengué: a) habitus; b) penis dorsal view

and inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista

dorsale. 16-17) N. nuperus Guignot, 1954, ex 4 (CAB), Gabon, Franceville, Mopia, riv. Ikengué: a) habitus, variations of elytral

pattern and of body shape / variazioni del disegno elitrale e della forma del corpo (in Fig. 17 elytra more narrowed behind giving the

appearance of having been laterally compressed / elitra più stretta posteriormente e che appare compressa lateralmente); b) penis dorsal

view and inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view/ apice del pene in

vista dorsale .

A REVISION OF THE AFRICAN SPECIES OF THE GENUS NEPTOSTERNUS SHARP, 1882 (COLEOPTERA DYTISCIDAE)

4 0.5 mm

2 mm

18c

18b

0.5 mm

19c

19b

0.5 mm

20c

20a 20b 20d

Figs. 18-20 - 18) N. nuperus Guignot, 1954, ex £ (CAB), Gabon, M.ts Cristal, Asseng Assala: a) habitus; b) penis dorsal view and

inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista dor-

sale. 19) N. oberthueri oberthueri Régimbart, 1903, lectotypus (MNHN), Madagascar (S.te Marie): a) habitus; b) penis dorsal view

and inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista

dorsale. 20) N. oberthueri oberthueri Régimbart, 1903, paralectotypus (MNHN), Madagascar (S.te Marie): a) habitus; b) penis dorsal

view and right paramere (inner face) / pene in vista dorsale e paramero destro (faccia interna); c) apex of penis dorsal view / apice del

pene in vista dorsale; d) penis lateral view / pene in vista laterale.

DI ARMANDO BILARDO & SAVERIO ROCCHI

14)

2la 21b 21d

0.5 mm

256

25a 25b 25d

Figs. 21-25 - 21) N. oberthueri bilardoi Pederzani & Rocchi, 2008, paratypus (CSR), Madagascar, Andohahela (Tulear), Isaka: a) habi-

tus; b) penis dorsal view and inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view

/ apice del pene in vista dorsale; d) penis lateral view / pene in vista laterale. 22-25a) N. oberthueri bilardoi Pederzani & Rocchi, 2008,

(NMW), Madagascar, Manampatrana Bas. (PO 167), loc. Aff. de la Lantara, Sahanivoraky (Andringitra): habitus, colour variation of

pattern / variazione del colore del disegno. 25b) penis dorsal view and inner face of parameres / pene in vista dorsale e faccia interna

dei parameri; 25c) apex of penis dorsal view / apice del pene in vista dorsale. 25d) penis lateral view / pene in vista laterale.

A REVISION OF THE AFRICAN SPECIES OF THE GENUS NEPTOSTERNUS SHARP, 1882 (COLEOPTERA DYTISCIDAE)

| 0.5 mm

26c

26a 26b 26d

2 mm

27b

2 mm %

28a 28b

Figs. 26-28 - 26) N. oblongus Régimbart, 1895, lectotypus (MNHN), Madagascar, Annanarivo, F. Sikora: a) habitus; b) penis dorsal

view and inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in

vista dorsale; d) penis lateral view / pene in vista laterale. 27) N. oblongus Régimbart, 1895, paralectotypus (MNHN), Madagascar

intérieur, Ankarata, F. Sikora: a) habitus; b) penis dorsal view and inner face of parameres / pene in vista dorsale e faccia interna dei

parameri; c) apex of penis dorsal view / apice del pene in vista dorsale. 28) N. oblongus Régimbart, 1895, (MNHN), Madagascar,

Békily: a) habitus; b) penis dorsal view and inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of

penis dorsal view / apice del pene in vista dorsale.

24 ARMANDO BILARDO & SAVERIO ROCCHI

2 mm

0.5 dal

290

29a 29b 29d

; 0.5 mm

30c

30a 30b 30d

2 mm

Figs. 29-31 - 29) N. ornatus Sharp, 1882, lectotypus (NHML), Madagascar: a) habitus; b) penis dorsal view and inner face of para-

meres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista dorsale; d) penis lateral

view / pene in vista laterale. 30) N. ornatus Sharp, 1882, paralectotypus (NMHN), Zanzibar: a) habitus; b) penis dorsal view and inner

face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista dorsale; d)

penis lateral view / pene in vista laterale. 31) N. rotroui Pic, 1924 (new synonym), lectotypus 9 (MNHN), Algeria, Sidi-Bel-Abbès:

habitus.

A REVISION OF THE AFRICAN SPECIES OF THE GENUS NEPTOSTERNUS SHARP, 1882 (COLEOPTERA DYTISCIDAE) DS

5 mm

2 mm

2 32c

32b

32d 32€

336

33b 33d 33e

0.5 mm

34c

34a 34b 34d 34e

Figs. 32-34 - 32) N. tropicus Guignot, 1954 (new synonym), holotypus (MRAC), Democratic Republic of Congo, P.N.U., R. Lufira:

a) habitus; b) penis dorsal view and right inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis

dorsal view / apice del pene in vista dorsale; d) penis dorsal-right lateral view / pene in vista dorsale-destra laterale; e) penis lateral view

/ pene in vista laterale. 33) N. meridianus Omer-Cooper, 1970 (new synonym), holotypus (NHML), Transvaal, Vaal River: a) habitus;

b) penis dorsal view and inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view /

apice del pene in vista dorsale; d) penis dorsal-right lateral view / pene in vista dorsale-destra laterale e) penis lateral view / pene in

vista laterale. 34) N. nigeriensis Omer-Cooper, 1970 (new synonym), holotypus (NHML), Nigeria, road to Enugo about 47 miles from

Makurdi: a) habitus; b) penis dorsal view and right paramere (inner face) / pene in vista dorsale e paramero destro (faccia interna); c)

apex of penis dorsal view / apice del pene in vista dorsale; d) penis dorsal-right lateral view / pene in vista dorsale-destra laterale; e)

penis lateral view / pene in vista laterale.

ARMANDO BILARDO & SAVERIO ROCCHI

2 mm

SATO

35a 35b 35d 356

0.5 mm

36c

36a 36b 36d

0.5 mm

37a DID 37d 36

Figs. 35-37 - 35) N. ornatus Sharp, 1882, (NHML), Sudan, S. Darfur, Cafindibie: a) habitus; b) penis dorsal view and inner face of

parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista dorsale; d) penis

dorsal-right lateral view / pene in vista dorsale-destra laterale; e) penis lateral view / pene in vista laterale. 36) N. ornatus Sharp, 1882,

(MNHN), Senegal, Badi, Parc Nat. du Niokolo Koba: a) habitus; b) penis dorsal view and right paramere (inner face) / pene in vista

dorsale e paramero destro (faccia interna); c) apex of penis dorsal view / apice del pene in vista dorsale; d) penis lateral view / pene

in vista laterale. 37) N. ornatus Sharp, 1882, (CFP), Guinea Bissau, 5 km N. de Madina-Salambande: a) habitus; b) penis dorsal view

and inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista

dorsale; d) penis dorsal-right lateral view / pene in vista dorsale-destra laterale; e) penis lateral view / pene in vista laterale.

A REVISION OF THE AFRICAN SPECIES OF THE GENUS NEPTOSTERNUS SHARP, 1882 (COLEOPTERA DYTISCIDAE) Da

0.5 mm

38c

39a

0.5 mm

40a 40b 40d

Figs. 38-40 - 38) N. ornatus Sharp, 1882, (CAB), Ivory Coast, Divo: a) habitus; b) penis dorsal view and inner face of parameres /

pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista dorsale. 39) N. ornatus Sharp,

1882, (CFP), Ivory Coast, Parc Nat. de la Comoé: a) habitus; b) penis dorsal view and inner face of parameres / pene in vista dorsale

e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista dorsale; d) penis dorsal-right lateral view / pene in

vista dorsale-destra laterale; e) penis lateral view / pene in vista laterale. 40) N. ornatus Sharp, 1882, (NHML), Nigeria, Ilorin Prov.,

Ilorin: a) habitus; b) penis dorsal view and inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis

dorsal view / apice del pene in vista dorsale; d) penis lateral view / pene in vista laterale.

28 ARMANDO BILARDO & SAVERIO ROCCHI

0.5 mm

4ld

0.5 mm

42a 42b 42d 42e

0.5 mm

43c

43a 44 43b

Figs. 41-44 - 41) N. ornatus Sharp, 1882, (ISNB), Democratic Republic of Congo, Lac Albert, Kasenyi: a) habitus; b) penis dorsal view

/ pene in vista dorsale; c) apex of penis dorsal view / apice del pene in vista dorsale; d) penis lateral view / pene in vista laterale. 42) N.

ornatus Sharp, 1882, (NMHL), Tanzania, Ukiriguru: a) habitus; b) penis dorsal view and inner face of parameres / pene in vista dorsale

e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista dorsale; d) penis dorsal-right lateral view / pene in

vista dorsale-destra laterale; e) penis lateral view / pene in vista laterale. 43-44) N. pederzanii Bilardo & Rocchi, 2010, (CAB), People

Republic of Congo, env. Lague, fleuve Mpama. 43) holotypus 4, a) habitus; b) penis dorsal view and inner face of parameres / pene

in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista dorsale. 44) paratypus 4, habitus,

colour variation of pattern / variazione del colore del disegno.

A REVISION OF THE AFRICAN SPECIES OF THE GENUS NEPTOSTERNUS SHARP, 1882 (COLEOPTERA DYTISCIDAE) 29

45b 45d

46b 46d

2 mm

0.5 mm

47c

4Ta 47b

Figs. 45-47 - 45) N. resartus Guignot, 1954, holotypus 4 (MRAC), Democratic Republic of Congo, P.N.U., Kaswabilenga: a) habitus;

b) penis dorsal view and inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view /

apice del pene in vista dorsale; d) penis lateral view / pene in vista laterale. 46) N. sombuicus Guignot, 1954, holotypus 4 (MRAC),

Democratic Republic of Congo, P.N.U., R. Lufira pied Mt. Sombwe: a) habitus; b) penis dorsal view and inner face of parameres / pene

in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista dorsale; d) penis lateral view / pene

in vista laterale. 47) N. psephotus Guignot, 1955 (new synonym), holotypus 4 (MRAC), Democratic Republic of Congo, Parc National

de la Garamba: a) habitus; b) penis dorsal view and inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex

of penis dorsal view / apice del pene in vista dorsale.

30 ARMANDO BILARDO & SAVERIO ROCCHI

I JI)

48c

48a 48b 48d

0.5 mm

2 mm /

49c

49a 49b 49d

0.5 mm

50c

5S0a SI 50b

Figs. 48-51 - 48) N. subopacus Guignot, 1956, holotypus £ (MRAC), Democratic Republic of Congo, Equateur, Bokuma: a) habi-

tus; b) penis dorsal view and right paramere (inner face) / pene in vista dorsale e paramero destro (faccia interna); c) apex of penis

dorsal view / apice del pene in vista dorsale; d) penis lateral view / pene in vista laterale. 49) N. tricuspis Guignot, 1954, paratypus &

(MRAC), Democratic Republic of Congo, P.N.U., R. Lufira pied Mt. Sombwe: a) habitus; b) penis dorsal view and inner face of para-

meres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista dorsale; d) penis lateral

view / pene in vista laterale. 50-51) N. bartolozzii n. sp., holotypus £' (MSNM), Gabon, M.ts Cristal, Asseng Assala. 50) a) habitus; b)

penis dorsal view and inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice

del pene in vista dorsal. 51) paratypus £ (MSNM)), habitus.

A REVISION OF THE AFRICAN SPECIES OF THE GENUS NEPTOSTERNUS SHARP, 1882 (COLEOPTERA DYTISCIDAE) 3 l

0.5 mm

2mm

SOC

D29

0.5.mm

IIC

0.5 mm

5S6b 5S6d

Figs 52-56 - 52-53) N. bartolozzii n. sp., paratypes (CAB), Gabon, Franceville, vill. Mopia, riv. Ikengué, 52) 4; a) habitus; b) penis

dorsal view and inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del

pene in vista dorsale. 53) 9 habitus, Gabon, Makokou. 54) N. africanus Peschet, 1916, lectotypus 9 (MNHN), Republic of Congo,

Fort-Crampel: habitus. 55) N. africanus Peschet, 1916, (MNHN), Sudan Région de Sennar: a) habitus; b) penis dorsal view and inner

face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista dorsale. 56)

N. africanus Peschet, 1916, (MNHN), Democratic Republic of Congo, P.N.U. Kaswabilenga: a) habitus; b) penis dorsal view and inner

face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista dorsale;

d) penis lateral view / pene in vista laterale.

209) ARMANDO BILARDO & SAVERIO ROCCHI

0.5 mm

2 mm

Sile

S7a S7b

0.5 mm

2 mm

58c

58a 58b 58d

0.5 mm

2 mm

59a 59b

Figs. 57-59 - 57) N. compsus Guignot, 1953, holotypus (MRAC), Democratic Republic of Congo, P.N.U. Kaswabilenga: a) habitus; b)

penis dorsal view / pene in vista dorsale; c) apex of penis dorsal view / apice del pene in vista dorsale. 58) N. compsus Guignot, 1953,

paratypus (MRAC), Democratic Republic of Congo, P.N.U. Kaswabilenga: a) habitus; b) penis dorsal view and inner face of parameres

/ pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista dorsale; d) penis lateral view

/ pene in vista laterale. 59) N. compsus Guignot, 1953, paratypus (ISNB), Democratic Republic of Congo, P.N.U. Kaswabilenga: a)

habitus; b) penis dorsal view and right paramere (inner face) / pene in vista dorsale e paramero destro (faccia interna).

A REVISION OF THE AFRICAN SPECIES OF THE GENUS NEPTOSTERNUS SHARP, 1882 (COLEOPTERA DYTISCIDAE) 33

0.5 mm

60c

60a 60b

6lc

6la 61b 61d

62b 62d

62a

Figs. 60-63 - 60) N. compsus Guignot, 1953, (CAB), Central African Republic: a) habitus; b) penis dorsal view; c) apex of penis dorsal

view / apice del pene in vista dorsale. 61) N. distinctus Omer-Cooper, 1970, holotypus (NHML), Nigeria, R. Assob, nr. Jos: a) habitus;

b) penis dorsal view and right paramere (inner face) / pene in vista dorsale e paramero destro (faccia interna); c) apex of penis dorsal

view / apice del pene in vista dorsale; d) penis lateral view / pene in vista laterale. 62-63) N. pumicatus Guignot, 1949, (ISNB) Demo-

cratic Republic of Congo, Lusengi. 62) lectotypus d, a) habitus; b) penis dorsal view and right paramere (inner face) / pene in vista

dorsale e paramero destro (faccia interna); c) apex of penis dorsal view / apice del pene in vista dorsale; d) penis lateral view / pene in

vista laterale. 63) paralectotypus, habitus, variation of extra, short longitudinal line / variazione della corta linea longitudinale extra.

34 ARMANDO BILARDO & SAVERIO ROCCHI

0.5 mm

2 mm

64c

64a 64b 64d

0.5 mm

65c

65a 65b 65d

0.5 mm

66c

66a 66b

Figs. 64-66 - 64) N. pumicatus Guignot, 1949, (CAB), Gabon, M.ts Cristal, Andok Foula: a) habitus; b) penis dorsal view and inner

face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista dorsale;

d) penis lateral view / pene in vista laterale. 65) N. simulator Omer-Cooper, 1970, paratypus (NHML), Transvaal, Waterberg dist.:

a) habitus; b) penis dorsal view and inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal

view / apice del pene in vista dorsale; d) penis lateral view / pene in vista laterale. 66) N. simulator Omer-Cooper, 1970, (CAB), Central

African Republic: a) habitus; b) penis dorsal view and inner face of parameres / pene in vista dorsale e faccia interna dei parameri;

c) apex of penis dorsal view / apice del pene in vista dorsale.

A REVISION OF THE AFRICAN SPECIES OF THE GENUS NEPTOSTERNUS SHARP, 1882 (COLEOPTERA DYTISCIDAE) 35

67c

67a 67b 67d

0.5 mm

68c

68a 68b

69c

0.5 mm

69a 70 69b

Figs. 67-70 - 67) N. komareki n. sp. holotypus £ (NMW), Benin, Dep. Platuau, Pobé near Crabé: a) habitus; b) penis dorsal view and

inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista dor-

sale; d) penis lateral view / pene in vista laterale. 68) N. komareki n. sp. (CAB), Gabon, Parc Nat. M.ts de Cristal: a) habitus; b) penis

dorsal view and inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del

pene in vista dorsale. 69) N. /ongitarsis n. sp. holotypus £' (CAB), Gabon, Franceville, vill. Mopia: a) habitus; b) penis dorsal view

and inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in vista

dorsale. 70) N. /ongitarsis n. sp. paratypus 9, (CAB), Gabon, Parc. Nat. Plateaux Batéké, Camp Ntsa: habitus.

36 ARMANDO BILARDO & SAVERIO ROCCHI

7le

0.5 mm

71b

0.5 mm

2 mm

TIC

73a 73b

$ 0.5 mm

2 mm

74c

74a 74b

Figs. 71-74 - 71-72) N. maliensis n. sp. (NMW), Mali, 100 km W. Sikasso, Bagoé river. 71) holotypus 4, a) habitus; b) penis dorsal

view and inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view / apice del pene in

vista dorsale. 72) paratypus 9, habitus. 73-74) N. batekensis Bilardo & Rocchi, 2010, paratypes (CAB), Gabon, Parc. Nat. Plateaux

Batéké, Camp Ntsa: a) habitus; b) penis dorsal view and inner face of parameres / pene in vista dorsale e faccia interna dei parameri;

c) apex of penis dorsal view / apice del pene in vista dorsale.

A REVISION OF THE AFRICAN SPECIES OF THE GENUS NEPTOSTERNUS SHARP, 1882 (COLEOPTERA DYTISCIDAE) 37

0.5 mm

Fiale

75a 76 75b

0.5 mm

IC Va

77a 77b

0.5 mm

2 mm

78c

78a 78b

Figs. 75-78 - 75) N. langoensis Bilardo & Rocchi, 2010, holotypus £ (CAB), Republic of Congo, Parc Nat. Odzala, Mboko, riv. Lango:

a) habitus; b) penis dorsal view and inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal

view / apice del pene in vista dorsale. 76) N. /angoensis Bilardo & Rocchi, 2010, ex 9, (CAB), RCA, Mambere-Kadei, Carnot-Gadzi

road, habitus. 77-78) N. susinii Bilardo & Rocchi, 2010, paratypes (CAB), Gabon, Parc. Nat. Plateaux Batéké, Camp Ntsa: a) habitus;

b) penis dorsal view and inner face of parameres / pene in vista dorsale e faccia interna dei parameri; c) apex of penis dorsal view /

apice del pene in vista dorsale.

38 ARMANDO BILARDO & SAVERIO ROCCHI

ECOLOGY

All species of the genus Neptosternus have peculiarly

long fore and intermediate legs; in particular the interme-

diate legs have long mesotarsomeres and strong claws to

grasp submerged stones and irregularities of the bottom,

in rapid streams. The hind legs bear long natatory setae

that enable the beetle to effectively swim also in running

water. Thanks to such features, Neptosternus are particu-

larly fit for running water environments and usually occur

in all kinds of streams, either slow flowing brooks, even

a few centimetres deep, or in major torrents. According to

our collecting experience in western and central Africa,

Neptosternus prefer streams with gravelly bottom or co-

arse sand, with or without submerged vegetation, either in

habitats exposed to sunlight or in shadowy forests; they

are particularly abundant downstream minor waterfalls,

among semi-submerged stones, even at the middle of the

stream. In forest brooklets with muddy bottom or with de-

caying leaves, they are extremely rare, as well as in the

calm puddles at riversides; such habitats are, on the con-

trary, very rich in other diving beetles, principally Cope-

latus spp.

Researches at the banks of large water streams with sandy

or muddy bottom rarely yielded Neptosternus, but captures

by light-traps near the same water streams sometimes yiel-

ded several specimens, also belonging to different species.

The photos (Figs. 79-93) show some habitats where some

species of Neptosternus have been collected with the net and

localities where hunting at light has been carried out.

The following species of diving beetles were collected

at times associated with Neptosternus in Gabon:

- shallow, slow running brooks: Yola darfurensis Bal-

four-Browne, 1947, Yola gabonica Bistròm, 1983, Lacco-

philus bizonatus Régimbart, 1895, Laccophilus desinte-

gratus Régimbart, 1895;

- sides of streams: Leiodytes hieroglyphicus (Régim-

bart, 1894) and some Hydrophilidae, in one biotope: Afri-

codytes mopiensis Bilardo & Rocchi, 2010.

No diving beetles other than Neptosternus were ever

collected from the middle of rapid streams; the only water

beetles found in association were Elmidae, that also are

well adapted to the rheophilic life, having peculiarly long

legs with strong onychia and grasping claws.

Fig. 79 - Republic of Congo, National Park of Odzala, Mboko, Lango river. Collecting locality of N. /angoensis Bil. & Roc.

A REVISION OF THE AFRICAN SPECIES OF THE GENUS NEPTOSTERNUS SHARP, 1882 (COLEOPTERA DYTISCIDAE)

Fig. 80 - Gabon, Cristal Mounts, Asseng Assala, Sanga river. Disturbed seconde

Guignot and bartolozzii n. sp., occurring in great number.

Fig. 81 - Gabon, National Park of Plateaux Batéké, riv. Mbi. Gallery forest, river bed with sandy ground, collecting locality of N.

batekensis Bil. & Roc.

ARMANDO BILARDO & SAVERIO ROCCHI

A REVISION OF THE AFRICAN SPECIES OF THE GENUS NEPTOSTERNUS SHARP, 1882 (COLEOPTERA DYTISCIDAE) 4]

Fig. 84 - Gabon, National Park of Plateaux Batéké, Mpassa river. East of Ntsa camp, near place of hunting at light.

Ri ia

Fig. 85 - Gabon, National Park of Plateaux Batéké, Mpassa river. Mecistops catafractus on sandbank.

ARMANDO BILARDO & SAVERIO ROCCHI

Fig. 86 - Gabon, Mopia, riv. Ikengué. Gallery forest, rocky/gravelly ground, with dense mats of submerged vegetation, collecting local-

ity of several species, with plenty of water beetles.

Fig. 87 - Gabon, Mopia, riv. Ikengué. Submerged vegetation: detail of Anubias (Araceae) spp.

A REVISION OF THE AFRICAN SPECIES OF THE GENUS NEPTOSTERNUS SHARP, 1882 (COLEOPTERA DYTISCIDAE)

Fig. 89 - Gabon, National Park of Cristal Mounts, small streams near place of hunting at light. Kinguélé.

ARMANDO BILARDO & SAVERIO ROCCHI

01 Tr ati , 5: 3 Ger “I ni di : L

g. 91 - Gabon, National Park of Cristal Mounts, small river near place of hunting at light. Ngonla Ngonla river.

A REVISION OF THE AFRICAN SPECIES OF THE GENUS NEPTOSTERNUS SHARP, 1882 (COLEOPTERA DYTISCIDAE)

Fig. 93 - Gabon, National Park of Cristal Mounts, hunting at light.

46 ARMANDO BILARDO & SAVERIO ROCCHI

CHECKLIST OF THE AFRICAN SPECIES

GroupA

Species with elytral pattern arranged as in Scheme A

Neptosternus alluaudi Régimbart, 1903: Madagascar.

= Neptosternus silvester Guignot, 1960 n. syn.

Neptosternus fasciatus Omer-Cooper, 1970: Nigeria, Ga-

bon.

Neptosternus hedychrous Guignot, 1955: Madagascar.

Neptosternus nuperus Guignot, 1954: Central Africa.

Neptosternus oberthueri oberthueri Régimbart, 1903:

Madagascar.

Neptosternus oberthueri bilardoi Pederzani & Rocchi,

2008 n. stat.: Madagascar.

Neptosternus oblongus Régimbart, 1895: Madagascar.

Neptosternus ornatus Sharp, 1882: Western, central and

southern Africa, Madagascar.

= Laccophilus rotroui Pic, 1924: 21 n. syn.

= Neptosternus tropicus Guignot, 1954 n. syn.

= Neptosternus meridianus Omer-Cooper, 1970 n. syn.

= Neptosternus nigeriensis Omer-Cooper, 1970 n. syn.

Neptosternus pederzanii Bilardo & Rocchi, 2010: Repu-

blic of Congo, Angola.

Neptosternus resartus Guignot, 1954: Democratic Repu-

blic of Congo.

Neptosternus sombuicus Guignot, 1954: Ethiopia, De-

mocratic Republic of Congo.

= Neptosternus psephotus Guignot, 1955 n. syn.

Neptosternus subopacus Guignot, 1956: Democratic Re-

public of Congo.

Neptosternus tricuspis Guignot, 1954: Democratica Re-

public of Congo, Angola.

Neptosternus bartolozzii n. sp.: Gabon.

Group B

Species with elytral pattern arranged as in Scheme B

Neptosternus africanus Peschet, 1916: Western, central

and southern Africa.

Neptosternus compsus Guignot, 1953: Central and sou-

thern Africa.

Neptosternus distinctus Omer-Cooper, 1970: Nigeria.

Neptosternus pumicatus Guignot, 1949: Western and

southern Africa.

Neptosternus simulator Omer-Cooper, 1970: Central and

southern Africa.

Neptosternus komareki n. sp.: Benin, Gabon.

Neptosternus longitarsis n. sp.: Gabon.

Neptosternus maliensis n. sp.: Mali.

Group C

Species with elytral pattern arranged as in Scheme C

Neptosternus batekensis Bilardo & Rocchi, 2010: Ga-

bon.

Neptosternus langoensis Bilardo & Rocchi, 2010: Cen-

tral Africa.

Neptosternus susinii Bilardo & Rocchi, 2010: Gabon.

Acknowledgements

We gratefully acknowledge the Curators of the Muse-

ums who kindly sent types and other materials for study,

namely Dr. Max Barclay (Natural History Museum, Lon-

don, UK), Dr. Marc De Mayer (Musée Royal de 1’ A frique

Centrale, Tervuren, Belgium), Dr. Jiti Hajek (Narodni

Muzeum, Prague, Czech Republic), Dr. Manfred Jàch

(Naturhistorisches Museum, Vienna, Austria), Dr. Pol

Limbourg (Institut Royal des Sciences naturelles, Brux-

elles, Belgium), Dr. Antoine Mantilleri (Muséum national

d’Histoire naturelle, Paris, France), and Miss Christine E.

Taylor (Natural History Museum, London, UK).

We thank for their kindly collaboration Dr. Anna

Alessandrello, Michela Mura and particularly Dr. Car-

lo Pesarini (Museo Civico di Storia Naturale, Milan,

Italy).

We wish to thank also Dr. Luca Bartolozzi, Chief of

the Zoology Section of the Natural History Museum “La

Specola” of the University of Florence (Italy) for his va-

luable suggestions and cooperation in the enforcement of

the International Code of Zoological Nomenclature, and

in obtaining loans from Museums.

Special thanks are due to our friend Dr. Fernando Pe-

derzani (Ravenna, Italy), specialist of Dytiscidae, for the

loan of his entire collection of Neptosternus, the valuable

suggestions and the linguistic help in preparing the En-

glish draft of this paper.

A REVISION OF THE AFRICAN SPECIES OF THE GENUS NEPTOSTERNUS SHARP, 1882 (COLEOPTERA DYTISCIDAE) 47

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Armando Bilardo

Via De Amicis 29, 21012 Cassano Magnago (VA), Italia.

e-mail: armandobilardo@libero.it

Saverio Rocchi

Museo di Storia Naturale dell’Università degli Studi di Firenze, Sezione di Zoologia “La Specola”, Via Romana 17, 50125 Firenze, Italia.

e-mail: rocchisaverio@yahoo.it

A revision of the African species of the genus Neptosternus Sharp, 1882 (Coleoptera Dytiscidae)

Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano

Volume XXXIII — Fascicolo III

III - ZANZUCCHI G., 1963 - Le Ammoniti del Lias superiore (To-

arciano) di Entratico in Val Cavallina (Bergamasco orientale).

pp. 99-146, 2 figg., 8 tavv.

Volume XIV

I- VENZOS.,, 1965 - Rilevamento geologico dell’anfiteatro more-

nico frontale del Garda dal Chiese all’ Adige. pp. 1-82, 1/ figg.,

4 tavv., 1 carta.

Il - PINNA G,, 1966 - Ammoniti del Lias superiore (Toarciano)

dell’Alpe Turati (Erba, Como). Famiglia Dactylioceratidae. pp.

83-136, 4 tavv.

INl - DIENI I., MASSARI F. e MONTANARI L., 1966 - Il Paleo-

gene dei dintorni di Orosei (Sardegna). pp. 13-/84, 5 figg., 8

tavv.

Volume XV

I- CARETTO PG, 1966 - Nuova classificazione di alcuni Brio-

zoi pliocenici, precedentemente determinati quali Idrozoi del

genere Hydractinia Van Beneden. pp. 1-88, 27 figg. 9 tavv.

II - DIENI I. e MASSARI F., 1966 - Il Neogene e il Quaternario dei

dintorni di Orosei (Sardegna). pp. 89-/42, 8 figg., 7 tavv.

INI - BARBIERI F., IACCARINO S., BARBIERI F. & PETRUCCI

F., 1967 - Il Pliocene del Subappennino Piacentino-Parmense-

Reggiano. pp. 143-188, 20 figg., 3 tavv.

Volume XVI

I- CARETTO P. G., 1967 - Studio morfologico con l’ausilio del

metodo statistico e nuova classificazione dei Gasteropodi plio-

cenici attribuibili al Murex brandaris Linneo. pp. 1-60, 1 fig.,

7 tabb., 10 tavv.

II - SACCHI VIALLI G. e CANTALUPPI G., 1967 - I nuovi fossi-

li di Gozzano (Prealpi piemontesi). pp. 61-128, 30 figg., 8 tavv.

III - PIGORINI B., 1967 - Aspetti sedimentologici del Mare Adria-

tico. pp. 129-200, 13 figg., 4 tabb. 7 tavv.

Volume XVII

I- PINNAG,, 1968 - Ammoniti del Lias superiore (Toarciano)

dell’Alpe Turati (Erba, Como). Famiglie Lytoceratidae, Nan-

nolytoceratidae, Hammatoceratidae (excl. Phymatoceratinae)

Hildoceratidae (excl. Hildoceratinae e Bouleiceratinae). pp.

1-70, 2 tavv. n.t., 6 figg., 6 tavv.

II - VENZO S. & PELOSIO G., 1968 - Nuova fauna a Ammonoidi

dell’Anisico superiore di Lenna in Val Brembana (Bergamo).

pp. 71-142, 5 figg., 11 tavv.

III - PELOSIO G., 1968 - Ammoniti del Lias superiore (Toarciano)

dell’ Alpe Turati (Erba, Como). Generi Hi/doceras, Phymatoce-

ras, Paroniceras e Frechiella. Conclusioni generali. pp. /43-

204, 2 figg., 6 tavv.

Volume XVIII

I-_ PINNAG,, 1969 - Revisione delle ammoniti figurate da Giu-

seppe Meneghini nelle Tavv. 1-22 della «Monographie des fos-

siles du calcaire rouge ammonitique» (1867-1881). pp. 5-22, 2

figg., 6 tavv.

II - MONTANARI L., 1969 - Aspetti geologici del Lias di Gozzano

(Lago d’Orta). pp. 23-92, 42 figg., 4 tavv. n.t.

Il - PETRUCCI F., BORTOLAMI G. C. & DAL PIAZ G. V., 1970

- Ricerche sull’anfiteatro morenico di Rivoli-Avigliana (Prov.

Torino) e sul suo substrato cristallino. pp. 93-169, con carta a

colori al 1:40.000, 14 figg., 4 tavv. a colori e 2 b.n.

Volume XIX

I- CANTALUPPI G,, 1970 - Le Hi/doceratidae del Lias medio

delle regioni mediterranee - Loro successione e modificazio-

ni nel tempo. Riflessi biostratigrafici e sistematici. pp. 5-46, 2

tabb. n.t.

II - PINNA G. & LEVI-SETTI F., 1971 - I Dactylioceratidae della

Provincia Mediterranea (Cephalopoda Ammonoidea). pp. 47-

136, 21 figg., 12 tavv.

II - PELOSIO G., 1973 - Le ammoniti del Trias medio di Askle-

pieion (Argolide, Grecia) - I. Fauna del «calcare a Ptychites»

(Anisico sup.). pp. 137-168, 3 figg., 9 tavv.

Volume XX

I- CORNAGGIA CASTIGLIONI O., 1971 - La cultura di Reme-

dello. Problematica ed ergologia di una facies dell’Eneolitico

Padano. pp. 5-80, 2 figg., 20 tavv.

II - PETRUCCI F., 1972 - Il bacino del Torrente Cinghio (Prov.

Parma). Studio sulla stabilità dei versanti e conservazione del

suolo. pp. 81-127, 37 figg., 6 carte tematiche.

III - CERETTI E. & POLUZZI A., 1973 - Briozoi della biocalca-

renite del Fosso di S. Spirito (Chieti, Abruzzi). pp. 129-169, 18

figg., 2 tavv.

Volume XXI

I- PINNAG,, 1974-I crostacei della fauna triassica di Cene in Val

Seriana (Bergamo). pp. 5-34, 16 figg., 16 tavv.

II - POLUZZI A., 1975 - I Briozoi Cheilostomi del Pliocene della

Val d’ Arda (Piacenza, Italia). pp. 35-78, 6 figg., 5 tavv.

III - BRAMBILLA G., 1976 - I Molluschi pliocenici di Villalvernia

(Alessandria). I. Lamellibranchi. pp. 79-128, 4 figg., 10 tavv.

Volume XXII

I - CORNAGGIA CASTIGLIONI O. & CALEGARI G., 1978 -

Corpus delle pintaderas preistoriche italiane. Problematica,

schede, iconografia. pp. 5-30, 6 figg., 13 tavv.

II - PINNA G,, 1979 - Osteologia dello scheletro di Kritosaurus

notabilis (Lambe, 1914) del Museo Civico di Storia Naturale

di Milano (Ornithischia Hadrosauridae). pp. 31-56, 3 figg., 9

tavv.

III - BIANCOTTIA., 1981 - Geomorfologia dell’ Alta Langa (Pie-

monte meridionale). pp. 57-104, 28 figg., 12 tabb., 1 carta ft.

Volume XXIII

I - GIACOBINI G., CALEGARI G. & PINNA G,, 1982 - I resti

umani fossili della zona di Arena Po (Pavia). Descrizione e pro-

blematica di una serie di reperti di probabile età paleolitica. pp.

5-44, 4 figg., 16 tavv.

II - POLUZZI A., 1982 - I Radiolari quaternari di un ambiente idro-

termale del Mar Tirreno. pp. 45-72, 3 figg., 1 tab., 13 tavv.

III - ROSSI F., 1984 - Ammoniti del Kimmeridgiano superiore-

Berriasiano inferiore del Passo del Furlo (Appennino Umbro-

Marchigiano). pp. 73-138, 9 figg., 2 tabb., 8 tavv.

Volume XXIV

I- PINNAG,, 1984 - Osteologia di Drepanosaurus unguicauda-

tus, lepidosauro triassico del sottordine Lacertilia. pp. 5-28, 12

figg., 2 tavv.

II - NOSOTTI S. e PINNA G,, 1989 - Storia delle ricerche e degli

studi sui rettili Placodonti. Parte prima 1830-1902. pp. 29-86,

24 figg., 12 tavv.

Volume XXV

I- CALEGARI G,, 1989 - Le incisioni rupestri di Taouardei

(Gao, Mali). Problematica generale e repertorio iconografico.

pp. 1-14, 9 figg., 24 tavv.

II - PINNA G. & NOSOTTI S., 1989 - Anatomia, morfologia fun-

zionale e paleoecologia del rettile placodonte Psephoderma al-

pinum Meyer, 1858. pp. 15-50, 20 figg., 9 tavv.

III - CALDARA R., 1990 - Revisione Tassonomica delle specie

paleartiche del genere 7ychius Germar (Coleoptera Curculioni-

dae). pp. 51-218, 575 figg.

Volume XXVI

I- PINNAG,, 1992 - Cvamodus hildegardis Peyer, 1931 (Reptilia,

Placodontia). pp. /-21, 23 figg.

IT - CALEGARI G. a cura di, 1993 - L’arte e l’ambiente del Sahara

preistorico: dati e interpretazioni. pp. 25-556, 647 figg.

III - ANDRI E. e ROSSI F., 1993 - Genesi ed evoluzione di fran-

genti, cinture, barriere ed atolli. Dalle stromatoliti alle comunità

di scogliera moderne. pp. 559-610, 49 figg., 1 tav.

Volume XXVII

I - PINNAG. and GHISELIN M. edited by, 1996 - Biology as

History. N. 1. Systematic Biology as an Historical Science. pp.

1-133, 68 figs.

II - LEONARDI C. e SASSI D. a cura di, 1997 - Studi geobotanici

ed entomofaunistici nel Parco Regionale del Monte Barro. pp.

135-266, 122 figg., 23 tabb.

Volume XXVIII

I- BANFI E. & GALASSO G., 1998 - La flora spontanea della cit-

tà di Milano alle soglie del terzo millennio e i suoi cambiamenti

a partire dal 1700. pp. 267-388, 71 figg., 30 tabb.

Volume XXIX

I- CALEGARI G., 1999 - L’arte rupestre dell’Eritrea. Repertorio

ragionato ed esegesi iconografica. pp. 1-174, 268 figg.

Volume XXX

I- PEZZOTTA F. edited by, 2000 - Mineralogy and petrology of

shallow depth pegmatites. Paper from the First International

Workshop. pp. 1-117, 30 figs., 19 tabs.

II - PARISI B., FRANCHINO A. & BERTI A. con la collaborazio-

ne di POTENZA B. & RUBINI D., 2000 - La Società Italiana

di Scienze Naturali 1855 - 2000. Percorsi storici. pp. /-/63,

199 figg.

III - DE ANGELLA. & GARASSINO A., 2002 - Galatheid, chi-

rostylid and porcellanid decapods (Crustacea, Decapoda, Ano-

mura) from the Eocene and Oligocene of Vicenza (N Italy). pp.

1-31, 21 figs., 9 pls.

Volume XXXI

I- NOSOTTI S. & RIEPPELO., 2002 - The braincase of Placodus

Agassiz, 1833 (Reptilia, Placodontia). pp. /-18, 15 figs.

II - MARTORELLI G., 2002 - Monografia illustrata degli uccelli

di rapina in Italia. (1895). Riedizione a cura di Fausto Barbagli.

pp. [XX] 1-216, [14] 46 figg., 4 tavv.

II - NOSOTTI S. & RIEPPEL O., 2003 - Eusaurosphargis dal-

sassoi n. gen. n. sp., a new, unusual diapsid reptile from the

Middle Triassic of Besano (Lombardy, N Italy). pp. /-33, 19

figs., 1 tab., 3 pls.

Volume XXXII

I- ALESSANDRELLO A., BRACCHI G. & RIOU B., 2004 - Po-

lychaete, sipunculan and enteropneust worms from the Lower

Callovian (Middle Jurassic) of La Voulte-sur-Rh6ne (Ardèche,

France). pp. 1-16, 9 figs., 1 pl.

II - RIEPPEL O. & HEAD J. J., 2004 - New specimens of the fossil

snake genus Eupodophis Rage & Escuillié, from Cenomanian

(Late Cretaceous) of Lebanon. pp. 1-26, 13 figs., 1 tab.

III - BRACCHI G. & ALESSANDRELLO A,, 2005 - Paleodiver-

sity of the free-living polychaetes (Annelida, Polychaeta) and

description of new taxa from the Upper Cretaceous Lagerstàt-

ten of Hagel, Hadjula and Al-Namoura (Lebanon). pp. 1-48, 8

figs., I tab., 16 pls.

Volume XXXIII

I- BOESI A. & CARDI F. edited by, 2005 - Wildlife and plants in

traditional and modern Tibet: conception, exploration and con-

servation. pp. 1-88, 30 figs., 9 tabs.

II - BANFI E., BRACCHI G., GALASSO . & ROMANI E,, 2005

- Agrostologia Placentina. pp. 1-80, 7 figs., 1 tab.

IIl - LIVI P. a cura di 2005 - I fondi speciali della Biblioteca del

Museo Civico di Storia Naturale di Milano. La raccolta di stam-

pe antiche del Centro Studi Archeologia Africana. pp. 1-250,

389 figs.

Volume XXXIV

I - GARASSINO A. & SCHWEIGERT G., 2006 - The Upper

Jurassic Solnhofen decapod crustacean fauna: review of the

types from old descriptions. Part I. Infraorders Astacidea, Tha-

lassinidea and Palinura. pp. 1-64, 12 figs., 20 pls. i

Il - FUCHS D., 2006 - Morphology, taxonomy and diversity of

vampyropod Coleoids (Cephalopoda) from the Upper Creta-

ceous of Lebanon. pp. 1-28, 9 figs., 9 pls.

III - CALDWELL M. W., 2006 - A new species of Pontosaurus

(Squamata, Pythonomorpha) from the Upper Cretaceous of Le-

banon and a phylogenetic analysis of Pythonomorpha. pp. 7-42,

18 figs., 1 pl.

Volume XXXV

I- DE ANGELI A. & GARASSINO A,, 2006 - Catalog and bib-

liography of the fossil Stomatopoda and Decapoda from Italy.

pp. 1-95.

Il - GARASSINO A., FELDMANN R.M. & TERUZZI G.,

edited by, 2007 - 3" Symposium on Mesozoic and Ce-

nozoic Decapod Crustaceans. Museo di Storia Natura-

le di Milano May 23-25, 2007. pp. 1-104, 38 figs., 6 tabs.

III - NOSOTTI S., 2007 - Tanystropheus longobardicus (Reptilia,

Protosaura): re-interpretations of the anatomy based on new

specimens from the Middle Triassic of Besano (Lombardy, N

Italy). pp. 1-88, 67 figs., 4 pls., 9 tabs.

Volume XXXVI

I- GALASSO G., CHIOZZI G., AZUMA M. & BANFI E,, a cura

di, 2008 - Le specie alloctone in Italia: censimenti, invasività e

piani di azione. Milano, 27-28 Novembre 2008. pp. 1-96.

Il - MAGANUCO S., STEYER J. S., PASINI G., BOULAY M.,,

LORRAIN S., BENETEAU A. & AUDITORE A., 2009 - An

exquisite specimen of Edingerella madagascariensis (Temno-

spondyli) from the Lower Triassic of NW Madagascar; cranial

anatomy, phylogeny, and restorations. pp. 1-72, 32 figs., 1 tab.,

4 appendix.

II - GARASSINO A.,, 2009 - The thoracic sternum and È

theca in the extant genera of the family Homolidae De Haan,

1839 (Crustacea, Decapoda, Brachyura). pp. 1-80, 2 figs., 18

pIs., 1 tab.

Volume XXXVII

I- DAL SASSO C. & MAGANUCO S., 2011 - Scipionyx samniti-

cus (Theropoda: Compsognathidae) from tha Lower Cretaceous

of Italy. Osteology, ontogenetic assessment, phylogeny, soft

tissue anatomy, taphonomy and palaeobiology. pp. /-287, 186

figs., 10 pls., 3 tabs., 7 appendix.

II - RIGATO F. - Contributions to the taxonomy of West European

and North African Stenamma of the westwoodii species-group.

(Hymenoptera Formicidae). pp. 1-56, 72 figs., 3 tabs., 2 ap-

pendix.

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