C H I A S O G N A T .

H I N A E
TA X O N O M Y ETHOLOGY
OF THE ANDES
BIOGEOGRAPHY
For the naturalist and the traveller, the American Andes are one of the richest parts
of the world f rom a faunistic, geological and botanical point of view – not forgetting
the multiplicity of habitats and the history of the peoples who have colonized them.
Following in the footsteps of Aimé Bonpland and Alexandre von Humboldt, to mention
but them, many naturalists have prospected the Andes and numerous works have been
devoted to them, extolling their richness and beauty. In their turn, F. Chalumeau, by profession
a writer, and B. Brochier, a pharmacist and biologist, both with a passion for entomology,
have devoted some thirteen years to studying a difficult and predominantly Andean
subfamily of Coleoptera, consisting of thirty-three representatives in South America, two
in Australia, and a fossil one in Europe.

The purpose of this book is ambitious. First of all it is to provide a practical tool for
the determination of all the surveyed species within an accurate taxonomical f ramework.
In order to do this, the authors visited South America many times, with the aim of collecting
specimens, and making ecological and faunistic observations. Apart f rom having had the
assistance of well-known specialists, they have consulted the collections of the most
important national museums (with their precious types) and those of many private individuals.
Finally, with the faunistic f ramework created, one of the most important aspects of this
work was tackled: the biogeography of the group f rom a worldwide perspective.

The reader will no doubt appreciate the complexity of the subject-matter, given
the scale of the studied area! Establishing distribution maps may be a relatively easy task,
but it is far more difficult to answer questions such as the following: how does one trace
the populations across the Andes, and explain the causes of subsequent vicariance
in the Chiasognathinae? What conclusions can be drawn? In other words: what are the factors
linked to the presence (today or in the past) of the taxa – including the “exotic” species,
i.e. those in Australia, and the one found in Europe? Where did all these fauna originate,
and how did they survive and multiply? So, apart f rom examining tectonic factors
and the geography of the areas covered, and using a floristic approach (and one concerned
with other biota), the authors have been obliged to look at the main stages in the history
of the earth’s crust, continental drift, and subsequent developments. The result is a work
whose language is sometimes complex, but always clear and concise, which is enhanced
by plentiful illustrations created by talented professional artists.

This book, a classic taxonomic study and an ecological approach to biogeography,

is intended for professional and amateur entomologists alike, as well as bio-geographers,
but also for all those who are passionate about Nature and wish to discover more about
this exceptional region known as the American Andes. They will find here a rich source
of information and escapist pleasure.
 Summary (Fig. 12 A - B, pages 34 - 35).

The Sphaenognathus form a homogenous group within the two subgroups: mandibular subgroup A, with series 4 of
the genus; mandibular subgroup B, with the series 1 to 3 of the genus. Nowadays and f rom a morphological perspective,
considering the structure of the male mandibles, it would seem that the genus Sphaenognathus represents the most evolved
stage of the subfamily. However, species with two mandibular carinae (genera Protognathinus, Chiasognathus, Bomansodus,
Chiasognathinus and its subgenera), forming a heterogeneous group, would seem to be less evolved.

In the females, the shape of the mandibles is more homogeneous; each of them showing, f rom a dorsal view,
the appearance of a half-circle. With the jaws closed, the mouthparts look like an almost circular plateau. One observes
a lower carina which is coarsely toothed, an upper carina which is raised and forming the half-circle of the mandible, along
with a smaller inner carina and an explanate outer one. As in the males, individual variations are considerable.

? spielbergi Protognathinus

queenslandicus Australognathus

Inf.
1A. grantii

beneshi
Chiasognathus
latreillei

jousselinii

2 carènes
Groupe 1 Sup. impubis Bomansodus

peruvianus
Chiasognathinus
gaujoni
Int.
1B.
Chiasognathinus
monguilloni
Sphaenognathinus

? kolbei

mandibularis Chiasornithodus

Fig. 12A: Mandibular groups and subgroups (1A and 1B).

34
 curviceps

lindenii

metallescens

inf. à peine metallifer

ébauchée série 4
2A. oberon

pubescens

rotundatus

taschenbergi

feisthamelii

giganteus
3 carènes série 3
Groupe 2
garleppi

higginsi
Sphaenognathus

bordoni

bellicosus
série 2
inf. nette armatus
2B.
nobilis

prionoides

spinifer

furumii série 1

alticollis
Fig. 12B: Mandibular groups and
subgroups (2A, 2B),
signatus and series of Sphaenognathus of
the determination keys (1, 2, 3, 4).

35
 Map 13: Colombia - climatic areas.

74

Barranquilla
0 200
km

Monteria

ca
u
Ca
Cucutá
uca
Ara
Medellín

alena
Atrato
Puerto Careño

Magd

ueoq
Orén
Bogotá

Buenaventura
are
Guavi

Tumaco
Patia Va
upé
Pasto s
Mitú
Ya R.
ri Ne
gro

Caqu
etá

Putumayo

one
Amaz
Napo
Steppe climate, xerophilous vegetation and rainfalls at midday.

Moist climate in the temperate and cold areas. Temperature depends of the altitude (from 18 °C to 20 °C).

Moist climate in the cold areas, low páramo (2,500-3,000m) and cloudy forests.

High tropical mountain climate. Páramos. Altitude above 3,100 m. Temperature under 10 °C. No vegetation.

Very moist climate of equatorial forest, with rainfalls along the year.

Moist climate along the year but with short rainfalls times.

Pasture climate periodically moist with rainfalls at midday.

Desert climate, xerophilous vegetation and rainfalls at midday.

86
 Map 14: Colombia - rain flow rates.

74

0 200
Barranquilla
km

Monteria

a
Cucutá

uc
Ca
uca
Ara
Atrato

Medellín

alena
Puerto Careño

Magd

ue
oq
Bogotá

Orén
4
Buenaventura
are
Guavi

Patia
Va
Tumaco upé
Pasto s
Mitú
Ya
ri

R. N
egr
o
Caqu
etá

Putumayo

one
Napo Amaz

less than 500 mm

from 500 to 1,000 mm

from 1,000 to 2,000 mm

from 2,000 to 3,000 mm

from 3,000 to 4,000 mm

more than 5,000 mm

« alto Páramo » area

87
 Fig. 40 - 42: Chile, Chiloë Island. Northern valdivian forest (Chilote).
Species living there: grantii.

110
Fig. 43 - 46: Chile, Cautín region. Villarica volcano, 1,450 m.
Moist valdivian forest. Species living there: grantii.

111
 2.3.2.5. The N. dombeyi and N. betuloides evergreen forests.

They spread from the Valdivian rain-forest to Tierra del Fuego, occupying the western slopes of the Cordillera and
the southern half of Chiloé island. In the northern continental part and in the southern half of Chiloé, the coigüe forest is
dominant. These forests reach as far as the 48th parallel, that is to say, far beyond the limits of the Chiasognathus * population.
The climate is cool, with an average annual temperature of 8.2 °C and a reduced thermal amplitude. Precipitation decreases
from north to south; it ranges from more than 4,000 mm to 1,000-2,000 mm in Tierra del Fuego. First mixed with species from
the Valdivian forest which are still present, the coigüe forest quickly becomes predominant, giving way, at around 1,000-1,200
metres altitude, to the lenga forest. Beyond the 48th parallel, the guindo forest (N. betuloides) then appears.

2.3.3. Geonemy of the Chiasognathinae.

The data that we have gathered show that the Chiasognathus have only been observed hitherto between 36° south
(Longaví) and 45.3° south (Coyhaique), both in Chile and Argentina. With the Andean cordillera separating southern America
into two unequal parts, with a distinct geonemy of species, we shall study their distribution in Chile, then in Argentina.

A - Chile.

Fig. 52: Chile, Cautín region. Villarica volcano. Moist valdivian forest.
Species living there: grantii, impubis. Species likely present: jousselinii, latreillei.

*The extreme distribution limit of grantii is situated near 45.3° (Coyhaique).

116
 Chile has been divided into three structural units of southern orientation: coastal range in the west; the Central Valley,
and the Andes range, in the east. Natural regions occur in regular succession according to the latitudes and longitudinal zoning.
Chiasognathus and Bomansodus are present only in the Valdivian zone.

About 1,000 km long (from north to south, between 36° and 48° south), this zone includes the following thirteen admi-
nistrative regions: Linares, Ñuble, Concepción, Arauco, Bío Bío, Malleco, Cautín, Valdivia, Osorno, Lanquihue, Chiloé, Palena
and Aysén. The town of Longaví (36° S., Linares) is probably the northern limit of the range of the Chiasognathinae, with only
the jousselinii (topotype of schoenemanni), while Coyhaique (45° S., Aysén) seems to be the southern limit of the expansion of
the subfamily, with grantii. Between those two extremes, areas inhabited by chiasognaths are much more numerous between
Bío Bío and Llanquihue. Chiasognathus latreillei, jousselinii, grantii, beneshi and Bomansodus impubis are present in the same
localities – grantii is the only species on Chiloé.

The listed sites are as follows: Linares (Estero de Leiva, Longaví), Ñuble (Chillán, Las Trancas), Concepción (Concepción),
Arauco (Contulmo, Lago Lanalhue, Caramavida, Renaico), Bío Bío (Termas Pemehue), Malleco (Angol, Curacautín, Malalcahuello,
Lago Jesús María, Las Raíces, Lonquimay, Nahuelbuta, La Fusta, Victoria), Cautín (Lago Caburga, Volcan Villarica, Cautín,
Cunco, Cherquenco), Valdivia (Valdivia, Antilhue, Río Bueno, Las Trancas, Río Correntoso), Osorno (Puerto Octay, Puyehue,
Antillaca), Llanquihue (Frutillar, Llanquihue, Lago Llanquihue, Lago Chapo, Correntoso, Hornohuinco, Río Blanco), Chiloé
(Chiloé island), Palena (Chaitén, Río Michimahuida, Mocopullí, Puerto Cárdenas, Lago Yelcho), Aysén (Aysén, Coyhaique,
Puente Zaranda, Río Simpson, Río Cisne Medio, Manihuales).

Fig. 53: Chile, eastern Temuco region, 1,600 m, with view on the Lonquimay volcano.
Species likely present: grantii, latreillei, jousselinii, impubis.

117
Sphaenognathus oberon Kriesche, 1922,

Sphaenognathus oberon Kriesche, 1922, 
 C H I A S O G N AT H I N U S G A U J O N I ( O b e r t h ü r, 1 8 8 5 )
(Fig. N° 92 - 98)

Chiasognathus gaujoni Oberthür, 1885. Bull. Soc. Ent. France: 198.

Chiasognathinus (Chiasognathinus) (Oberthür) Chalumeau & Brochier, 1995.

Derivatio nominis. – As a tribute to the Abbé Gaujon, the first collector of this species.

Bibliography: Nonf ried, 1891: 278; Felsche, 1898: 6; Heyne & Taschenberg, 1908: 48, pl. 6, fig. 8; Van Roon, 1910: 4;
Kriesche, 1919: 94; Campos, 1921: 50; Didier, 1937: 11; Blackwelder, 1944: 195; Didier & Séguy, 1952: Série A, 28, pl 2, fig 1-8;
Didier & Séguy, 1953: Série A, 27: 63 & 71; Benesh, 1960: 150; De Lisle, 1967: 522; Numhauser, 1981: 17; Lacroix, 1984: 7:
Chalumeau & Brochier, 1995: 20; Krajcik, 2001: 82.

Synonymies: [= leoni Lacroix, 1983 (Chalumeau & Brochier, 1995)].

Typical series: Many male specimens.

Male lectotype. With the labels: Équateur Loja Abbé Gaujon (black bordered); Chiasognathus Gaujoni R. Oberthür, type Bull. soc.
Ent. France 1885. p. CXCVIII (black bordered ); TYPE (red ); MUSÉUM PARIS 200.D.; SYNTYPUS (red ) det. L. Bartolozzi 1988;
LECTOTYPE (red ) Chiasognathinus (Oberthür) Dés. F. Chalumeau & B. Brochier ‘92; Sbg CHIASOGNATHINUS Didier &
Séguy. – At the MNHN, Paris. Species described from Ecuador.

Surveyed localities: Ecuador, region of Loja.

Number of specimens examined: 60 M + 3 F.

Collections: ANSP (2 M); BMNH (8 M, 1 F); CARN (7 M, 1 F); CAS (3 M); CBOM (6 M, 1 F); CBRO (2 M); FMNH
(8 M); IREC (2 M); IRSN (1 M ); MNHUB (6 M); MZ (2 M); SMTD (9 M); UZM (4 M).

Diagnosis: Lenght: M: 32-41 mm - F: 28-33 mm.

Male: Mandibles black, plunging, imbricate at their apex with, on the upper side, a strong tooth directed towards inside
at their base and directed towards the rear end. Under side of the mandible with an acuminate manubrium. Elytra greenish
to mauve. Pronotum with violet metallic glints, covered with a yellow ochre pubescence. Legs bicoloure.

Female: Same general aspect than the male. Mandibles semi-circular, the upper carina not reaching the base; it finishes at the level of
the median tooth of the lower carina.

Synonymies

• Sphenognathus leoni Lacroix, 1983. Bull. Sci. Nat, 40, p. 7, pl. 2.

• Chiasognathinus (Chiasognathinus) leoni (Lacroix) (Chalumeau & Brochier).

Derivatio nominis. – As a tribute to Mr Carrère.

Bibliography: Chalumeau & Brochier, 1995: 20; Krajcik, 2001: 82.

170
Fig. 92 - Chiasognathinus gaujoni Fig. 93 - Chiasognathinus gaujoni
Ecuador, Loja Ecuador, Loja
Fig. 94 - Chiasognathinus gaujoni Fig. 95 - Chiasognathinus gaujoni

171
 Typical series: one specimen.
Male holotype. Body length: 30 mm; elytra width: 12 mm; Mandibles length: 5,5 mm – coll. Lacroix. Described from: Perú, Tingo
María. X.1969. Allotype not designated. Female unknown.

Note: Described by Lacroix from a single specimen, we thought this species was synonym of gaujoni. Indeed, the differences (based
on the slender aspect and the coloration of this specimen) are not significant. The examination of a very large series shows, as in
all Chiasognathinae, important differences as much in the size as in the colour. In gaujoni, we have observed specimens with elytra
either bronze green or reddish brown, and the pronotum with green or violet iridescence.

Frequency: Pretty common, but localized.

Fig. 96 - Chiasognathinus gaujoni
Fig. 97 - Chiasognathinus gaujoni  Fig. 98 - Chiasognathinus gaujoni

Ecuador, Loja Ecuador, Loja Ecuador, Loja

172
Chiasognathinus gaujoni
- x2.8,  - x3.1
 C H I A S O G N AT H I N U S M O N G U I L L O N I ( L a c r o i x & B o m a n s , 1 9 7 2 )
(Fig. N° 114 - 122)

Sphenognathus monguilloni Lacroix & Bomans, 1972. Bull. Ann. Soc. R. Belg. Ent., 108, page 47, fig. 6.
Chiasognathinus (Sphaenognathinus) monguilloni (Lacroix & Bomans) (Chalumeau & Brochier, 1995).

Derivatio nominis. – As a tribute to Mr Monguillon.

Bibliography: Chalumeau & Brochier, 1995: 20; Krajcik, 2001: 82.

Typical series: seven specimens

Male holotype. Body length: 30 mm, elytra width: 14,4 mm, mandibles length: 7 mm. With the labels: Sphenognathus
monguillonii HOLOTYPE Lacroix J.P. LACROIX det.1971; legue de l ’auteur à MNHN Paris J.P. LACROIX det. 1973 (orange
colour bordered ); PERou Mt Carpich 2800 m Tingo Maria Monguillon 1970.; TYPE (red ); Figuré Jla + (stamped signature) – At
the MNHN, Paris. With 6 male paratypes: coll. Monguillon, Bomans, Lacroix. Allotype not designated. Species described
from Peru, surroundings of Tingo María, Mt Carpish, 2.800 m.

Surveyed localities: Peru: regions of Huánuco (Mts Carpish, Tingo María), San Martín (Uchiza Huállagas), Cajamarca
(Cajamarca).

Number of specimens examined: 93 M + 28 F.

Collections: BMNH (1 M, 1 F); CALA (4 M); CARN (4 M); CBOM (4 M, 3 F); CBOR (4 M); CBRO (38 M, 14 F); CM
(1 M, 1 F); FMNH (7 M, 2 F); IREC (26 M, 7 F); IRSN (1 M); IZA (3 M);

Diagnosis: Lenght: M: 33-58 mm - F: 32-43 mm.

Male: Mandibles elongate, slender and plunging; with, at the base of the upper carina, a strong horizontal tooth directed
towards inside; on the under side, presence of an acuminate manubrium. Elytra greenish. Lacinia strongly fleshy at its apex.

Female: Black. Base of the nasus sinuous. Mandibles broadened and flat on the upper side, with an arc-shaped half-carina more
or less raised.

Frequency: Common.

184
Fig. 114 - Chiasognathinus monguilloni Fig. 115 - Chiasognathinus monguilloni
Peru, Carpish Huánaco Peru, Carpish Huánaco
Fig. 116 - Chiasognathinus monguilloni Fig. 117 - Chiasognathinus monguilloni

185
 Fig. 118 - Chiasognathinus monguilloni
Fig. 119 - Chiasognathinus monguilloni
Fig. 120 - Chiasognathinus monguilloni

Peru, Carpish Huánaco Peru, Carpish Huánaco Peru, Carpish Huánaco

Fig. 121 - Chiasognathinus monguilloni
Fig. 122 - Chiasognathinus monguilloni 

Peru, Carpish Huánaco Peru, Carpish Huánaco

186
Sphaenognathus monguilloni
- x2.25,  - x2.9