On phyletic closeness between South American and New Caledonian spilopyrines (Chrysomelidae, Eumolpinae, tribe Spilopyrini)

2006·3.5 MB·

by  Krishna K Verma

Bonnerzoologische Beiträge Band 54(2005) Heft4 Seiten 297-303 Bonn, Oktober2006
On Phyletic Closeness Between South American and New Caledonian
Spilopyrines (Chrysomelidae, Eumolpinae, Tribe Spilopyrini)1
Krishna K. Verma" & Pierre JOLlVET2'
"ßorsi, Dure, India
" Pans, France, and Florida State Museum ofEntomology, Gainesville, FL, USA.
Abstract. Among members ofthe eumolpine tribe Spilopyrini (Reid 1995), the South American genera Hornius and
Stenomela and the New Caledonian Bohumiljania are especially similar, while the Australian and the New Guiñean
forms seem to stand a little apart. In this communication, by tracing taxonomic history ofspilopyrines, and by taking
intoaccounttheirfood plants,palaeogeological historyofthe Southern Hemisphere,anatomical featuresofspilopyrines,
and their developmental stages, a possible phyletic closeness between the South American and the New Caledonian
generahasbeen inferred.
Keywords.Taxonomy,ecology,zoogeography
1. INTRODUCTION These bits oftaxonomic history clearly support the no-
tion ofphyletic closeness between the S. American and
REID (2000) separated some members of the chry- theNew Caledonian forms.
somelid subfamily Eumolpinae, and put them in a new
subfamily, Spilopyrinae. Earlier these genera had been
regarded by REID (1995) as a tribe under Eumolpinae, 3. FOOD PLANTS
the Tribe Spilopyrini. These primitive Eumolpinae are
Hornius of Chile, as larva and adult, is a specialised
being referred to in this communication as spilopyrines,
following Verma & Jolivet (2004). The spilopyrin feeder on leaves and bark ofNothofagus trees (Fagaceae)
(JEREZ 1996). Stenomela ofthe same region, on the other
genera include: Spilopyra Baly, 1860 (Australia and
New Guinea), Macrolema Baly, 1861 (Australia and hand feeds on different species of Myrtaceae (Blepharo-
New Guinea), Richmondia Jacoby, 1898 (Australia), jcaanliyaxccarluekdsohnainckasiiinNthieeddeisntzauntaNnedwLCuamlaedosnppi.a),.aBloshoufmeield-s
Cheiloxena Baly, 1860 (Australia), Stenomela Erichson,
on Myrtaceae (Sizygium spp.). This is a pointofsimilarity
1847 (Chile), Hornius Fairmaire, 1848 (Chile and Ar-
gentina), and Bohumiljania Monrós, 1958 (New Cale- between Stenomela and Bohumiljania. But in spilopyri-
donia). As pointed out by Verma & Jolivet (2004), the nes, Cheiloxena in Australia also feeds on Myrtaceae
Chilean and the New Caledonian spilopyrines are espe- {Eucalyptus spp.) and Spilopyra on Sapindaceae (Cu-
paniopsis and Guioa). Host plants of Macrolema and
cially similar. This communication aims to emphasize a
Richmondiaarenotonrecord(Jolivetetal. 2003).
possible phyletic closeness between the South American
and the New Caledonian spilopyrines. In this context
PALAEOGEOLOGICAL HISTORY
the following are specifically notable. 4.
In the context ofdistribution ofspilopyrines, the Gond-
TAXONOMIC HISTORY
2. wana Hypothesis has been invoked byJerez (1996) and
JOLIVET (1954), describing the wing venation in the Reid (2000). Holloway (1979) and Verma & Jolivet
New Caledonian spilopyrin, referred to it as Stenomela (2004) have pointed out that in the late Cretaceous (75
caledonica. MONRÓS (1958) created a new generic mya) continental shelves ofS. America and ofthe Aus-
name for this spilopyrin, Bohumiljania. SEENO & WIL- tralian Plate were still connected with those ofthe Ant-
COX (1982), following MONRÓS (1958), placed arctica, while Africa and India, that also resulted from
Stenomela and the New Caledonian Bohumiljania to- the break, had moved away northward considerably re-
moved from Gondwanaland.
gether under the Tribe Stenomelini, a tribe under Eu-
molpinae. The Tribe Hornibiini, with a single genus In a recent paper SANMARTIN & RONQUIST (2004) gave
Hornius, has been placed close to the Tribe Stenomelini. a detailed and interesting account of Gondwanian
history, supported by explicit maps of the southern
Paperpresentedtothe6lhInternational SymposiumontheChry- hemisphere in different geological periods. The fol-
somelidae, Bonn,Germany,May7,2004. lowing parts ofthe history, as mentioned in that paper
298 Bonnerzoologische Beiträge 54(2005)
121 my
Africa
Madagascar
•
India
Gondwanaland 84my 30 my
NewZealand
125 my NewCaledonia
South America
•
80 my
Australia^Floral/faunalexchanges
V continuedintoPleistocene
probablythrough
60 my NewGuineiantermediateislands.
30 my
Fig. 1. Cladograms showing sequence ofseparation ofland masses in the Southern Hemisphere (based on data in Sanmartín&
Ronquist2004, and in a summaryofherstudiesprovided by I. Sanmartín, Uppsala, pers. comm. 2004).
and in a summary of their work, provided by Isabel (v) Opening ofthe Drake passage between Antarctica
Sanmartín (Uppsala, pers. comm. 2004), are of special and South America at the boundary between Eo-
interest in the present context. cene and Oligocene (30 to 28 mya led to the onset
ofthe Circum-Antarctic Current, and this resulted
(i) "Consistent with other studies, the animal data are
in cooling ofthe Antarctica.
congruent with the geological sequence of Gond-
wana breakup Trans-Antarctic dispersal (Aus- Jerez (1996) has pointed out that Nothofagus arose in
tralia southern South America) is also sig- the Australian part ofGondwana, and from there it dis-
nificantly more frequent than any other dispersal persed to New Zealand and S. America.
event in animals, which may be explained by the
long period ofgeological contact between Austra- As mentioned above, Hornius is a specialized feeder on
tliíana&ndRSoonuqtuhisAtme2r0i0c4a).via Antarctica." (Sanmar- Npoetrsheodfagtuos.AusIttraalpipae,arNsetwhatCaplreidmiotniivae asnpdiloSp.yrAimneesridcias,-
along with temperate flora, like Nothofagus, and subse-
(ii) In Palaeocene (60 mya) New Zealand + New quently, due to cooling and development ofsubantarctic
Caledonia separated from Australia. conditions in the southern part ofS. America, and due to
separation ofNew Caledonia as an island from the Aus-
(iii) In mid to late Tertiary (mid Tertiary 40 to 30 mya)
New Caledonia separated from New Zealand. tralian Plate, the primitive spilopyrines in these parts
became shielded from competition with more modern
(iv) During the period 60 to 30 mya Australia and forms, and have survived as Hornius and Stenomela in
South America remained in contact across Antarc- southern S. America, and as Bohumiljania in New Cale-
tica (Fig. 5), and this continued up to Eocene- donia. One may ask: New Guinea is also an island, and
early Miocene. At this time the Antarctica had a it shares Spilopyra and Macrolema with Australia, then
warm temperate climate, and had a rich flora, how island isolation of New Caledonia could help the
dominated by Nothofagus, as evidenced by fossils survival ofBohumiljania as a primitive spilopyrin? One
ofthis plant found in coastal parts ofthe Antarc- answer to this question: New Zealand + New Caledonia
tica. Ready dispersal of Nothofagus to Australia separated from Australia in Palaeocene (60 mya),
and South America is probable. I. Sanmartín whereas floral and faunal exchanges between Australia
(Uppsala, pers. comm. 2004) points out, "This and New Guinea could continue even into the Pleisto-
(= a warm temperate climate in the Antarctica) cene (I. Sanmartín, Uppsala,pers. comm., 2004)(Fig. 5).
presumably established a significant dispersal
route for the exchange ofsouthern temperate biota
between South America and Australia"
KrishnaK. Verma & PierreJolivet: SouthAmerican andNew Caledonian Spilopyrini (Chrysomelidae) 299
5. ANATOMICAL FEATURES and robust body form like typical Eumolpinae. All the
New Caledonian and S. American spilopyrines have an tinhrceoelSo.urAmoenrtihceahno-stNepwlanCta.leOdnontihaenogtheenrerhaanardetlhieghAtugsrterean-
elongated and cerambycid like body shape (Fig. 2). The lian genera show brightand diverse metallic colours.
Australo-Papuan genera in contrast have a more compact
mm
1
Fig. 2. New Caledonian and South American spilopyrines. (i)Bohumiljcinia (from Jolivetetal. 2003); (ii) Hornius (trom Jerez
1996); (iii)Stenomela(fromJerez 1996).
300 Bonnerzoologische Beiträge 54(2005)
(iii) The ventrally directed basal orifice of the
aedeagus is relatively restricted anteroposteriorly.
In contrast: (i) In Spilopyra, Macrolema, Cheiloxena
and Richmondia the differentiation of the basal hood
and the aedeagus proper is somewhat bettermarked than
in the S. American and New Caledonian forms; (ii) In
Richmondia, Cheiloxena and to some extent in Spi-
lopyra the ventral curvature ofthe aedeagal tube is more
marked than in S. American and New Caledonian spi-
lopyrines, and in Richmondia and Cheiloxena this cur-
vature is almost as strong as in higher Eumolpinae; and
(iii) the anteroposterior extent of the basal orifice is
large, specially in Macrolema, Cheiloxena and Rich-
mondia (For differences between typical eumolpine and
spilopyrin aedeagi see Verma & Jolivet2004).
Thus the aedeagal structure in Hornius, Stenomela and
Bohumiljania is more primitive than in the remaining
spilopyrines, and almost of the chrysomeline type (For
features of the chrysomeline type of aedeagus see
Verma 1996).
SPD
i
\
3
Fig. 3. Aedeagi, lateral view, (i) ofBohumiljania; (ii) ofHor-
niiis; (iii) oiStenomela(from Verma& Jolivet2004).
The aedeagal structure in spilopyrines in general is
primitive, as discussed by Verma & Jolivet (2004).
But in Hornius, Stenomela and Bohumuljania it is even Fig. 4. Spermathecal complex ofBohumiljania (from Verma
more primitive; the primitive features of their aedeagi & Jolivet2004).
include:
(i) Poor differentiation of the basal hood and the Spermatheca in all spilopyrines shows a characteristic
eumolpine feature, namely the proximal part of the
aedeagus proper(Fig. 3).
spermathecal capsule is differentiated into a swollen
(ii) Only moderate ventral curvature ofthe aedeagus, or bulbous section, which receives the duct of the
spread throughout the length ofthe organ, and not spermathecal gland and from which the spermathecal
confined to the basal part ofthe aedeagus proper. duct starts. In Bohumiljania this proximal part is some-
In Stenomela, however the curvature is more what elongated and presents some irregular coiling
marked in the basal part ofthe aedeagus proper. (Fig. 4). Such acoiling is seen also in the spermathecaof
Krishna K. Verma & PierreJOLIVET: SouthAmerican and New Caledonian Spilopyrini (Chrysomelidae) 301
ii iii
Fig. 5. Southern hemisphere, (i) Gondwana (-140 my). End ofJurassic; (ii) Polar view ofthe Southern hemisphere (-80 my).
Mid-Cretaceous; (iii) Polarview ofthe Southern hemisphere (- 60 my). Early Eocene. The Drake Passage isjust forming. New
Zealand, New Caledonia, Norfolk ridge and Lord Howe ridge are also figured (from Isabel Sanmartín summary (2002), pers.
comm. 2004, withpermission).
302 Bonnerzoologische Beiträge 54(2005)
Megascelis does not have a mediocubital patch in the
hind wing like spilopyrines. Richmondia, however has a
dark spot in this position, and not a well defined me-
diocubital patch (Verma & Jolivet 2004).
DEVELOPMENTAL STAGES
6.
Eggs ofHornius are laid under an oothecal cover, made
of excrement material, and a single cover includes 7-8
eggs (JEREZ & CERDA 1988). In Stenomela, eggs are
also provided with excretory matter cover, each cover
protecting two eggs. In Bohumiljania the ootheca is
made up ofexcretory material mixed with vegetal fibres
and a glandular secretion, and there are four eggs per
ootheca (Jolivet et al. 2003). Perhaps it will not be out
place to mention that in Megascelis there is also a mem-
branous protective cover for a batch of about 10 eggs
(Cox 1988). In remaining spilopyrines laid eggs have
not yet been observed and reported.
The neonate larva ofBohumiljania bores onto the stem
ofthe host plant, but later instars are exposed feeders on
leaf lamina (Jolivet et al. 2003). There is a similar
habit with the first instar larvae of Hornius and
Stenomela; they pierce into leaf buds and leaf lamina,
but later instars are well exposed, feeding on leaves.
Larval habits have not been recorded so far for the Aus-
tralian spilopyrines.
Pupae ofHornius are formed in soil in a specially con-
structed cell (JEREZ 1996). Pupae ofStenomela and Bo-
humiljania have not been observed yet, but they too are
believed to be formed in soil (Jerez 1995; Jolivet et
al. 2003).
CONCLUDING REMARKS
7.
Phyletic closeness between spilopyrines of New Cale-
donia and S. America has been hypothesized on the ba-
2
sis ofthe above discussion. This hypothesis will hope-
Fig. 6. Bohumiljania caledonica (Jolivet). (1) starting to fly; fully gain additional support through further studies on
(2) on a Syzygium cumini L. ( Myrtaceae) leaf, in New Cale- the biology ofthe Australian spilopyrines, and through
donia DNA
analysis studies.
Megascelinae (Suzuki 1988). Megascelinae are essen-
tially Central and South American in distribution, and Acknowledgements. We must thank here Dr. Michael
are closely related to Eumolpinae (JOLIVET 1957-1959; Schmitt, from Zoologisches Forschungsmuseum Alexander
Reid 1995; Cox 1998). Should this situation be re- Koenig, Bonn, who welcomed the junior author at the
g&arAdSedHLaOsCcKove(r1e9d91b)yiwnhtahtehfaosllboeweinngdepsacirliboefdtbheyirMtAeYxtR? CsfihrtroyymsCtoohnmefeelFriledonrcisedy,ampMionussMieauuymmh2eo0lf0d4E,dnutaroinmndoglDtorhg.ey,AMfiGrcaihicanaeenlsvBiTilholdoeim,vaeFsrL,-
"A serious problem is posed by propensity ofgenotypes who has provided laboratory access to the junior author
to produce a certain phenotype, such as stalked eyes in while in the USA. Our best thanks also to Dr. Isabel San-
certain acalyptrate dipterans, which is manifested in martin (Uppsala University, Sweden) who provided decu-
only some ofthe possessors ofa genotype. Parallelism mentation and illustrations.
in this case may be defined as homologous similarity,
since the common ancestor evidently had the genetic
propensity even ifit was not expressed phenotypically."
In this context it would be relevant to point out that
.
Krishna K. Verma& PierreJolivet: South American andNewCaledonian Spilopyrini (Chrysomelidae) 303
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