Mediterranean-southern African disjunct distribution and flea beetles: the example of the “black species” of Longitarsus associated with Boraginaceae (Coleoptera, Chrysomelidae, Alticinae).

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Key words. Alticinae, Boraginaceae, Afrotropical Region, Longitarsus, Mediterranean area, disjunct distribution.
SUMMARY
In this paper the disjunct distribution of two Longitarsus species-groups associated with Boraginaceae: capensis-group,
restricted to the southern Africa and presently including 15 species, and anchusae-group (sensu Biondi, 1995), widespread
in the western Palaearctic area with 4 species, is discussed. Some hypotheses explaining the separate distributions of
Mediterranean and South African groups are proposed. Finally, the host-plant shift from native Lobostemon spp. to the
introduced plant Echium plantagineum L. shown by some species of the capensis group is also discussed.
INTRODUCTION
Longitarsus Latreille, as described in Berthold (1827), is presently the largest
genus within Chrysomelidae, Alticinae with about 600 known species occur-
ring throughout all zoogeographical regions (Biondi & D’Alessandro, 2008).
This flea beetle genus is well represented in the Mediterranean area (about
160 species), where is better known from an autecological standpoint (Biondi,
1996). Longitarsus species occurring in the Mediterranean area are known to
feed, as adults, on plants of different botanical families, the preferred being
Boraginaceae (26.4%), Lamiaceae (24.9%), Asteraceae (13.8%), Plan taginaceae
(9.5%), and Scrophulariaceae (9.5%) (Fig. 1). Particularly, species-groups, such
as aeneus, anchusae, exsoletus and echii feeding on Boraginaceae, show an eleva-
ted degree of trophic specialization (Jolivet & Hawkeswood, 1995; Biondi, 1996)
and include mainly oligotopic species, generally termophilous (Figs 2-3). In the
Biogeographia vol. XXIX - 2008
(Pubblicato il 30 dicembre 2008)
The Mediterranean-southern African disjunct
distribution pattern
Mediterranean-southern African disjunct
distribution and flea beetles: the example of the
“black species” of Longitarsus associated with
Boraginaceae (Coleoptera, Chrysomelidae,
Alticinae)
MAURIZIO BIONDI, PAOLA D’ALESSANDRO
Dipartimento di Scienze Ambientali, Università de L’Aquila,
67010 Coppito, L’Aquila, Italy
e-mail: biondi@univaq.it, paola.dalessandro@univaq.it.

Mediterranean area, in fact, 34% of the Longitarsus species associated with
Boraginaceae are monophagous (feed on one or two closely related botanical
genera), 56% oligophagous (feed on more botanical genera of one or two clo-
sely related families) and only 10% polyphagous (feed on many botanical spe-
cies not closely related). The trophic categories are those proposed by Biondi
(1996) and Fernandez & Hilker (2007). Therefore, in the Mediterranean area
90% of the Longitarsus species associated with Boraginaceae (34% monopha-
gous and 56% oligophagous) feed exclusively on them (Biondi, 1996) (Fig. 3).
In the Afrotropical Region, the genus Longitarsus presently includes about 120
described species, but many other yet undescribed taxa exist (Biondi, unpubl.
data). Auto-ecological data available for the Afrotropical species of this flea bee-
tle genus are extremely scarce or completely lacking if compared to the
Mediterranean species. In the literature there are only a few reports of Longitarsus
species associated with Boraginaceae in the sub-Saharan Africa, namely
Longitarsus punctifrons Weise, 1895 (= gossypii Bryant, 1941) collected on
Heliotropium sp. (Furth, 1985), and species of the capensis group collected on
Lobostemon spp., Anchusa capensis Thumb., Echium plantagineum L. and
Heliotropium sp. (Biondi, 1999; Biondi & D’Alessandro, 2008).
This paper includes some observations about the distribution of two
Longitarsus species-groups associated with Boraginaceae: capensis-group, restric-
ted to the southern Africa and presently including 15 species, and anchusae-
group (sensu Biondi, 1995), widespread in the western Palaearctic area with
4 species (see list below) (Fig. 6).
100
Fig. 1 - Percentages of the Mediterranean Longitarsus species for host botanical families in the Mediterranean area
(redraw from Biondi, 1996).

The most part of zoogeographical and autecological data on the species of
the Longitarsus capensis-group reported in this contribution were collected
during zoological collecting trips that were part of an Italian research project
(PRIN 2004057217) aimed at interpreting the disjunct distribution of diffe-
rent plant and animal groups in the Mediterranean-South African regions (cf.
Balinsky, 1962; La Greca, 1970, 1990; Axerold & Raven, 1978; Jürgens, 1997;
Coleman et al., 2003).
101
Fig. 2 - Percentages of ecological categories, based on environmental preferences, in the Mediterranean Longitarsus
species associated with Boraginaceae. ERY: eurytopic; OLM: oligotopic-mesophilous; OLT: oligotopic-termophilous;
STT: stenotopic-thermophilous.
Fig. 3 - Percentages of trophic categories in the Mediterranean Longitarsus species associated with Boraginaceae. POL:
polyphagous; OLI: oligophagous; MON: monophagous.

COMMENTED LIST OF THE SPECIES
capensis species-group:
Longitarsus afromeridionalis Biondi & D’Alessandro, 2008
Distribution. Southern-Western Afrotropical element (cf. Biondi &
D’Alessandro, 2006), distributed in the southern-western coastal area of the
Western Cape Province (Republic of South Africa) (Fig. 7).
Host plants. Boraginaceae: Lobostemon cf. lucidus (Lehm.) H. Bueck.
Trophic range. Monophagous.
Phenology. September (Tab. I).
Longitarsus capensis Baly, 1877
Longitarsus capensis: Biondi, 1999; Biondi & D’Alessandro, 2008
102
Fig. 4 - Habitus, median lobe of aedeagus, and spermatheca of Longitarsus afromeridionalis Biondi & D’Alessandro,
2008, a member of the South African capensis species-group. LE: length of the elytra; LAED: length of the median
lobe of aedeagus; LSP: length of the spermatheca.

Distribution. Southern-Western Afrotropical element (cf. Biondi &
D’Alessandro, 2006), distributed in the western part of the Western Cape
Province (Republic of South Africa) (Fig. 7).
Host plants. Boraginaceae: Lobostemon fruticosus (L.) H. Buek, L. cf. lucidus,
Anchusa capensis Thumb. and Echium plantagineum L.
Trophic range. Oligophagous.
Phenology. July, September, October (Tab. I).
Longitarsus cedarbergensis Biondi, 1999
Longitarsus cedarbergensis: Biondi & D’Alessandro, 2008
Distribution. Southern-Western Afrotropical element (cf. Biondi &
D’Alessandro, 2006), distributed in the Cedarberg area (Republic of South
Africa: Western Cape Province) (Fig. 7).
103
Fig. 5 - Habitus, median lobe of aedeagus, and spermatheca of Longitarsus anchusae (Paykull, 1799), a member of
the West Palaearctic anchusae species-group. LE: length of the elytra; LAED: length of the median lobe of aedeagus;
LSP: length of the spermatheca.

Host plants. Boraginaceae: Lobostemon cf. dorotheae M. H. Buys.
Trophic range. Monophagous.
Phenology. September (Tab. I).
104
Fig. 6 - General distribution of the Longitarsus species of the South African capensis group and the West Palaearctic
anchusae group.

Longitarsus debiasei Biondi & D’Alessandro, 2008
Distribution. Southern-Western Afrotropical element (cf. Biondi &
D’Alessandro, 2006), distributed in the Langeberg area (Republic of South
Africa: Western Cape Province) (Fig. 8).
Host plants. Boraginaceae: Echium plantagineum.
Trophic range. Monophagous (?).
Phenology. October (Tab. I).
Longitarsus grobbelaariae Biondi & D’Alessandro, 2008
Distribution. Southern-Eastern Afrotropical element (cf. Biondi &
D’Alessandro, 2006) distributed in Kwazulu-Natal (Republic of South Africa)
(Fig. 7).
105
Fig. 7 - Distribution of: Longitarsus afromeridionalis Biondi & D’Alessandro, L. capensis Baly, L. cedarbergensis Biondi,
L. grobbelaariae Biondi & D’Alessandro, L. melanicus Biondi and L. transvaalensis Biondi.

Host plants. Boraginaceae: Heliotropium sp.
Trophic range. Monophagous.
Phenology. January, February (Tab. I).
Longitarsus hexrivierbergensis Biondi & D’Alessandro, 2008
Distribution. Southern-Western Afrotropical element (cf. Biondi &
D’Alessandro, 2006), distributed in the Hexrivierberg area (Republic of South
Africa: Western Cape Province) (Fig. 8).
Host plants. Boraginaceae: Echium plantagineum.
Trophic range. Monophagous (?).
Phenology. October (Tab. I).
106
Fig. 8 - Distribution of: Longitarsus debiasei Biondi & D’Alessandro, L. hexrivierbergensis Biondi & D’Alessandro, L.
luctuosus Biondi, L. lugubris Biondi, L. neseri Biondi, L. piketbergensis Biondi & D’Alessandro and L. rouxi Biondi
& D’Alessandro.

Longitarsus luctuosus Biondi, 1999
Longitarsus luctuosus: Biondi & D’Alessandro, 2008
Distribution. Southern-Western Afrotropical element (cf. Biondi &
D’Alessandro, 2006), distributed in the southern-western part of the Western
Cape Province (Republic of South Africa) (Fig. 8).
Host plants. Boraginaceae: Echium plantagineum.
Trophic range. Monophagous (?).
Phenology. July, September (Tab. I).
Longitarsus lugubris Biondi, 1999
Longitarsus lugubris: Biondi & D’Alessandro, 2008
Distribution. Southern-Western Afrotropical element (cf. Biondi &
107
Fig. 9. Distribution of: Longitarsus malherbei Biondi & D’Alessandro and L. sudafricanus Biondi & D’Alessandro.

D’Alessandro, 2006), distributed in the Bokkeveldeberg and Cedarberg areas
(Republic of South Africa: Northern and Western Cape Province) (Fig. 8).
Host plants. Boraginaceae: Lobostemon cf. dorotheae and L. cf. trichotomus
(Thumb.) DC.
Trophic range. Monophagous.
Phenology. September (Tab. I).
Longitarsus malherbei Biondi & D’Alessandro, 2008
Distribution. Southern-Western Afrotropical element (cf. Biondi &
D’Alessandro, 2006), distributed in the Cedarberg area (Republic of South
Africa: Western Cape Province) (Fig. 9).
Host plants. Boraginaceae: Lobostemon cf. dorotheae and L. cf. trichotomus.
Trophic range. Monophagous.
Phenology. September (Tab. I).
108
Fig. 10. Host-plants of the South African Longitarsus capensis species-group.

Longitarsus melanicus Biondi, 1999
Longitarsus melanicus: Biondi & D’Alessandro, 2008
Distribution. Southern-Western Afrotropical element (cf. Biondi &
D’Alessandro, 2006), distributed in the southern coastal area of the Western
Cape Province (Republic of South Africa) (Fig. 7).
109
Tab. I - Phenology of the Longitarsus species of the South African capensis group and the West Palaearctic anchusae
group. Light grey rectangles indicate negative samplings in places where the species was previously collected.
JAN FEB MAR APR MAY JUN JUL AGO SEP OCT NOV DEC
capensis species-group
afromeridionalis
capensis
cedarbergensis
debiasei
grobbelaariae
hexrivierbergensis
luctuosus
lugubris
malherbei
melanicus
neseri
piketbergensis
rouxi
sudafricanus
transvaalensis
anchusae species-group
anatolicus
anchusae
hittita
saulicus

Host plants. Boraginaceae: Lobostemon cf. marlothii Levyns.
Trophic range. Monophagous.
Phenology. September (Tab. I).
Longitarsus neseri Biondi, 1999
Longitarsus neseri: Biondi & D’Alessandro, 2008
Distribution. Southern-Western Afrotropical element (cf. Biondi &
D’Alessandro, 2006), distributed in the Piketberg area (Republic of South
Africa: Western Cape Province) (Fig. 8).
Host plants. Boraginaceae: Echium plantagineum.
Trophic range. Monophagous (?).
Phenology. July (Tab. I).
Longitarsus piketbergensis Biondi & D’Alessandro, 2008
Distribution. Southern-Western Afrotropical element (cf. Biondi &
D’Alessandro, 2006), distributed in the Piketberg area (Republic of South
Africa: Western Cape Province) (Fig. 8).
Host plants. Boraginaceae: Lobostemon fruticosus.
Trophic range. Monophagous.
Phenology. September (Tab. I).
Longitarsus rouxi Biondi & D’Alessandro, 2008
Distribution. Southern-Western Afrotropical element (cf. Biondi &
D’Alessandro, 2006), distributed in the Cedarberg area (Republic of South
Africa: Western Cape Province) (Fig. 8).
Host plants. Boraginaceae: Lobostemon sp., probably L. cf. dorotheae or L.
cf. trichotomus.
Trophic range. Monophagous.
Phenology. April, September (Tab. I).
Longitarsus sudafricanus Biondi & D’Alessandro, 2008
Distribution. Southern-Western Afrotropical element (cf. Biondi &
D’Alessandro, 2006), distributed in the western mountain areas of the Western
Cape Province (Republic of South Africa) (Fig. 9).
Host plants. Boraginaceae: Lobostemon cf. dorotheae, L. fruticosus, L. cf. tri-
chotomus and Echium plantagineum.
Trophic range. Oligophagous.
Phenology. September (Tab. I).
110

Longitarsus transvaalensis Biondi, 1999
Longitarsus transvaalensis: Biondi & D’Alessandro, 2008
Distribution. Southern-Eastern Afrotropical element (cf. Biondi &
D’Alessandro, 2006), distributed in Mpumalanga Province (Republic of South
Africa) (Fig. 7).
Host plants. This species was collected at forest edge on leaves of an uni-
dentified plant belonging to the Borage family.
Trophic range. Monophagous (?).
Phenology. December (Tab. I).
anchusae species-group:
Longitarsus anatolicus Weise, 1900
Longitarsus anatolicus: Biondi, 1995, 1996; Gruev & Döberl, 1997, 2005.
Distribution. Central Asiatic element (cf. Vigna Taglianti et al., 1999),
extended to the Anatolian region and Bulgaria.
Host plants. Boraginaceae: Anchusa spp.
Trophic range. Monophagous.
Phenology. May, June (Tab. I).
Longitarsus anchusae (Paykull, 1799)
Longitarsus anchusae: Biondi, 1995, 1996; Gruev & Döberl, 1997, 2005.
Distribution. European element (Vigna Taglianti et al., 1999), extended
to western Asia.
Host plants. Boraginaceae: Anchusa spp., Asperugo sp., Buglossoides arven-
se (L.) Johnst., Cerinthe sp., Cynoglossum spp., Echium plantagineum, Myosotis
sp., Nonnea sp., Pulmonaria sp., Solenanthus (?) sp., Symphytum spp.
Trophic range. Oligophagous.
Phenology. March, April, May, June, July (Tab. I).
Longitarsus hittita Biondi, 1995
Longitarsus hittita: Biondi, 1996; Gruev & Döberl, 1997, 2005.
Distribution. Anatolian endemic element (cf. Vigna Taglianti et al.,
1999), known from Turkey.
Host plants. Boraginaceae: Solenanthus stamineus (Desf.) Wettst.
Trophic range. Monophagous.
Phenology. May, June (Tab. I).
Longitarsus saulicus Gruev & Döberl, 2005
= Longitarsus morio Sahlberg, 1913: Biondi, 1995, 1996; Gruev & Döberl,
1997.
111

Distribution. E-Mediterranean element (cf. Vigna Taglianti et al., 1999),
distributed in Israel and Jordan.
Host plants. Boraginaceae: Alkanna strigosa Boiss. & Hohen., Anchusa
spp., Cynoglossum creticum Miller, Echium glomeratum Poiret, Symphytum pale-
stinum Boiss.
Trophic range. Oligophagous.
Phenology. February, March, April (Tab. I).
DISCUSSION
Both these Longitarsus species-groups here considered, containing exclusi-
vely species with a black integument, are strictly associated with Boraginaceae
generally in thermophilous environments, and share the following updated
morphological characteristics that differentiate the anchusae and capensis grou-
ps within Longitarsus (Biondi, 1999; Biondi & D’Alessandro, 2008): i) head
with impunctate vertex and frons with some large punctures impressed near
the frontal grooves; ii) elytra apically subtruncate or widely and independen-
tly rounded; iii) humeral callus always completely absent in capensis-group,
rarely developed in anchusae-group; iv) hind wings strongly reduced (species
always subapterous in capensis-group; brachypterous, sub-brachypterous or
very rarely macropterous in anchusae-group); v) large spermatheca (generally
longer than 0.30 mm) with ducts often widely arcuate and usually uncoiled
or with one coil, very rarely with two coils; vi) median lobe of aedeagus stron-
gly sclerotized, mostly with a distinct apical median small tooth and a ventral
sulcus invariably with clear impressions; vii) impressed elytral punctation gene-
rally dense and without signs of striae even in sutural area. The latter charac-
ter is very important for distinguishing the species of the anchusae and capen-
sis groups from the black species of Longitarsus associated with Lamiaceae, com-
mon in both the Mediterranean area and southern Africa.
Species of the capensis and anchusae groups are remarkably similar (Figs 4-
5). In fact there is only one morphological trait, namely the degree of metatho-
racic wing reduction, that distinguishes the two groups; only subapterous spe-
cies are known in the capensis species-group whereas brachypterous, sub-bra-
chypterous or macropterous species can be found in the anchusae species-
group. The similarities, if shown to be synapomorphic by a future phyloge-
netic analysis, might suggest that the two groups comprise a monophyletic
unit whose Mediterranean and South African subgroups became geographi-
cally separated relatively recently.
Alternative hypotheses could explain the separate distributions of
Mediterranean and South African anchusae and capensis species-groups. This
particular type of distribution is widely documented for many plant and ani-
112

mal groups (cf. Balinsky, 1962; La Greca, 1970, 1990; Axerold & Raven,
1978; Jürgens, 1997; Coleman et al., 2003). Excluding unlikely recent long-
distance dispersal events, the most probable hypothesis to explain this type of
geographical distribution is to accept there were ecological connections
between the Mediterranean and South African areas in the past. Species in the
capensis-group are significantly associated with “fynbos”, a Mediterranean-type
of vegetation that is widespread in the south-western part of South Africa and
probably formed no earlier than the Late Miocene (cf. Axelrod & Raven, 1978;
Richardson et al., 2001). Based on this observation, the migratory flow of one
or more Mediterranean capensis ancestors from the north towards South Africa
may have occurred during the Quaternary period via “arid corridors” (sensu
Balinski, 1962) which “appeared” in eastern Africa due to climatic changes
that took place in the Northern Hemisphere during the glacial periods (cf.
Jürgens, 1997).
In the Cape region, the capensis species-group radiated into different species,
often confined to rather limited areas (Figs 7-9). In the Mediterranean, during
the Quaternary period, species diversification was very limited in the anchusae-
group, as well as in other closely related species-groups (cf. Biondi, 1995). This
occurrence is frequent in many plant and animal groups in the Cape region, which
have a comparatively high biodiversities compared with ecologically similar areas.
Different factors have been suggested to explain this phenomenon: (1) topo-
graphical complexity; (2) edaphic complexity; (3) pollinator specialization; (4)
fire; and (5) short dispersal distance (Barraclough, 2006).
Mediterranean-type ecosystems are included among the main biodiversity
hot spots around the world (Myers et al., 2000; Goldblatt & Manning, 2002)
and fire is considered one of the main drivers of diversification in these eco-
systems. Recurrent fires over millions of years, with an average frequency of
about 15-50 per year, have been suggested to trigger dramatic diversification
by selecting taxa with short generation times and/or to instigate the isolation
of populations (Verdù et al., 2007). In our opinion, the high number of spe-
cies in the capensis-group in such a restricted geographic area can be explained
by the important role fire played in leading to the evolution of different life
history strategies and driving the process of speciation or local extinction in
fynbos plants and in animals associated with them.
As mentioned above, species of the capensis-group are primarily associated
with fynbos and plants of the genus Lobostemon (Boraginaceae). Lobostemon
includes perennial shrubs with alternating leaves and flowers, mostly bell-sha-
ped and usually pink or blue. This genus includes twenty-eight species ende-
mic to South Africa and is largely confined to the winter rainfall area from
Springbok to Mossel Bay and further eastward along the coast to near
Grahamstown, where it rains throughout the year (Buys, 2006).
113

It is important to emphasize that 6 species of the 14 attributed to the capen-
sis-group (Fig. 10) were collected from Echium plantagineum, an alien plant
introduced into South Africa from Europe after 1825 and now present in
Western and Eastern Cape and in the temperate mountain areas of Free State,
Lesotho and Kwazulu-Natal, with some reports from southern Mpumalanga
(cf. Retief & Van Wyk, 1998). From a systematic point of view, the genus
Echium is a sister-group of Lobostemon (Böhle et al., 1996), so the host-plant
shift from Lobostemon to Echium observed in some species of the capensis-group
could have easily occurred. However, in our opinion, this ecological event
could dramatically alter the autecology and evolutionary mechanisms of the
species involved (Biondi & D’Alessandro, 2008). The presence of E. planta-
gineum in South Africa represents, in fact, a new “shared” trophic resource for
the sympatric species of the capensis-group and, simultaneously, an unexpec-
ted connection among allopatric species as consequence of the easy spread of
this invasive plant. On the basis of other analogous adaptive processes obser-
ved in native species in response to the introduction of invasive plants (Mooney
& Cleland, 2001; Strauss et al., 2006), it is possible that unexpected evolu-
tionary events could result in several different scenarios: (1) catalyze the local
breakdown of reproductive isolation between native sympatric parent species;
(2) bring into contact parent allopatric species and contribute to the reinfor-
cement (or deletion) of reproductive isolating mechanisms; and (3) promote
speciation by hybridization as an adaptive response of two or more native spe-
cies to the new host-plant, as observed in species of the genus Rhagoletis Loew
(Diptera Tephritidae) in testing invasive plants of Lonicera sp. (Caprifoliaceae)
in North America (Schwarz et al., 2005, 2007).
With the knowledge presently available, it is impossible to evaluate which sce-
nario among those listed above may play the most significant role in determin-
ing the high rate of species diversification within the capensis-group. Only fur-
ther analyses focused on phylogeography by mean biomolecular analyses (e.g.
coalescent simulation, biomolecular clock) may help clarify the paleobiogeogra-
phy of the capensis species-group.
ACKNOWLEDGEMENTS
We are grateful to the following colleagues who allowed us to study valu-
able material preserved in their respective institutions: S. L. Shute (The Natural
History Museum, London, United Kingdom), A. Vigna Taglianti (Zoological
Museum, University of Rome “La Sapienza”, Italy), E. Grobbelaar (South
African National Collection, Plant Protection Research Institute, Pretoria,
Gauteng, Republic of South Africa), R. Müller (Transvaal Museum, Pretoria,
Gauteng, Republic of South Africa) and M. Uhlig and H. Wendt (Museum
114

für Naturkunde der Humboldt-Universität, Berlin, Germany). We would also
like to offer special thanks to our friends, P. A. Audisio, M. A. Bologna and
A. De Biase (Italy, Rome), fellow travellers on many collecting trips in South
Africa. This study was funded by grant PRIN 2004057217, “Zoogeography
of Mediterranean-southern African disjunct distributions by a multi-method
approach” from the Ministero dell’Università e della Ricerca.
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