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A description of the larva of Heliaeschna idae Brauer from Borneo, with a supplementary note on the larva of H. univervulata Martin (Odonata: Aeshnidae)

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The final instar larva of Heliaeschna idae Brauer is described and figured for the first time based on the exuvia from an advanced female larva collected in Sarawak, Borneo (East Malaysia). It is compared with the known larvae of the genus and is concluded to be most closely allied to Heliaeschna simplicia Karsch, with which it shares a unique structure on the anterior margin of the prementum of the labium, along with several other distinctive characters. The dorsal structure of the head of H. uninervulata is re-examined and illustrated. It is shown to bear a prominent convexity and tuft of long setae on the vertex which is similar to a structure previously recorded only in H. simplicia, and which is only weakly developed in H. idae. The taxonomic implications of these partly conflicting observations are discussed and it is concluded the Oriental members of the genus Heliaeschna might be separated into two or three separate genera, which are as yet unnamed.
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A description of the larva of
Heliaeschna idae Brauer from Borneo,
with a supplementary note on the larva
of H. univervulata Martin (Odonata:
Aeshnidae)
Albert G. Orr a , Robin W.J. Ngiam b & Rory A. Dow c
a Griffith, School of the Environment , Griffith University ,
Nathan , Q 4111 , Australia
b National Biodiversity Centre , National Parks Board , Singapore
Botanic Gardens, 1 Cluny Road. S(259569), Singapore
c Naturalis Biodiversity Center , PO Box 9517, 2300 RA , Leiden ,
the Netherlands
Published online: 16 Jul 2013.
To cite this article: International Journal of Odonatology (2013): A description of the
larva of Heliaeschna idae Brauer from Borneo, with a supplementary note on the larva of
H. univervulata Martin (Odonata: Aeshnidae), International Journal of Odonatology, DOI:
10.1080/13887890.2013.813827
To link to this article: http://dx.doi.org/10.1080/13887890.2013.813827
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Downloaded by [Albert G. Orr] at 15:14 16 July 2013
International Journal of Odonatology, 2013
http://dx.doi.org/10.1080/13887890.2013.813827
A description of the larva of Heliaeschna idae Brauer from
Borneo, with a supplementary note on the larva of
H. univervulata Martin (Odonata: Aeshnidae)
Albert G. Orra*, Robin W.J. Ngiamband Rory A. Dowc
aGriffith, School of the Environment, Griffith University, Nathan, Q 4111, Australia; bNational
Biodiversity Centre, National Parks Board, Singapore Botanic Gardens, 1 Cluny Road. S(259569),
Singapore; cNaturalis Biodiversity Center, PO Box 9517, 2300 RA, Leiden, the Netherlands
(Received 28 February 2013; final version received 14 April 2013 )
The final instar larva of Heliaeschna idae Brauer is described and figured for the first time based on the
exuvia from an advanced female larva collected in Sarawak, Borneo (East Malaysia). It is compared with
the known larvae of the genus and is concluded to be most closely allied to Heliaeschna simplicia Karsch,
with which it shares a unique structure on the anterior margin of the prementum of the labium, along with
several other distinctive characters. The dorsal structure of the head of H. uninervulata is re-examined and
illustrated. It is shown to bear a prominent convexity and tuft of long setae on the vertex which is similar
to a structure previously recorded only in H. simplicia, and which is only weakly developed in H. idae.
The taxonomic implications of these partly conflicting observations are discussed and it is concluded the
Oriental members of the genus Heliaeschna might be separated into two or three separate genera, which
are as yet unnamed.
Keywords: Aeshnidae; dragonfly; Heliaeschna idae; larva; Sarawak
Introduction
The genus Heliaeschna Selys currently includes 11 recognised species occurring in tropical Africa
and south-east Asia (Schorr & Paulson, 2012). Present thinking holds that the five African species
and six Asian species are not congeneric (Dijkstra, 2005), and it has been suggested on the basis
of larval characters that the south-east Asian species may represent at least two, possibly three
genera (Kawashima & Sasamoto, 2007; Orr & Ngiam, 2011; Butler & Orr, 2013). To date there
is no available description of the larva of any African species but the larvae of three Oriental
species are known and described. These are H. filostyla Martin, a species endemic to Sulawesi,
described by Kawashima and Sasamoto (2007), H. uninervulata Martin from Singapore (Orr &
Ngiam, 2011) and H. simplicia Karsch from Borneo (Butler & Orr, 2013). The adults of these
three species are all very distinct. Another distinctive grouping is formed by the two very closely
allied species H. idae (Brauer) and H. crassa Krüger, the affinities of which might be clarified by
larval characters. The position within the genus of the sixth Oriental species, H.bartelsi Lieftinck,
is also uncertain.
*Corresponding author, at: 26 Currimundi Rd, Caloundra, Q4551, Australia. Email: agorr@bigpond.com
© 2013 Worldwide Dragonfly Association
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2A.G. Orr et al.
Recently, one of us (RWJN) collected an advanced female larva from Sarawak, Borneo. The
adult emerged soon after and has been identified as H. idae on the basis of comparisons with
voucher specimens of female H. idae and H. crassa taken in copula. The separation of females of
these two species has long proved virtually impossible (Orr, 2003), and therefore the determination
as H. idae should probably still be considered provisional. Here we describe the final stadium of
female larva of H. idae based on the exuvia left after emergence.
We compare the larval characters of this species with those of the three Heliaeschna species for
which larvae are already known and discuss their taxonomic implications. In addition we amend
the description of H. uninervulata (Orr & Ngiam, 2011) following further examination of exuviae
of both sexes of this species.
The larva of Heliaeschna idae
Material examined
One exuvia ex larva, collected 3 May 2012, Malaysia, Sarawak, Usun Apau National Park,
03.02666N, 114.64799E, alt. 969 m, muddy forest pool, leg. Robin Ngiam. One adult female,
emerged from larva, same locality data.
Description
Diagnosis. Habitus typically aeshnid (Figure 1); head large with prominent eyes and strongly
developed mouthparts elongating the face, outer margins of mandibles angulated and prominent
in dorsal view, legs moderately long, abdomen fusiform with moderately long anal appendages
and lateral spines on S5–S10. Coloration of exuvia pale greyish ochre with a fairly well-defined
pattern and prominent dark speckles associated with small tubercles at the base of larger setae
(Figure 1); in life reddish brown with heavy dark markings, eyes yellowish. Body surface generally
finely pilose with patches of heavier setae in places.
Head. Large, robust, roughly rounded pentagonal, with very prominent eyes (Figure 2a). Pos-
tocular lobes well developed and evenly rounded. Glabrous suboval areas immediately behind
bases of eyes reaching almost to hind margin, each ringed by a line of small tubercles surmounted
by setae. Remainder of postocular area weakly pilose, with strong marginal and submarginal setae
in the posterolateral regions, the bases of these setae mounted on small dark tubercles presenting
a strong speckled appearance. Vertex very slightly swollen behind ocelli with a small compact tuft
of dark setae. Anterior part of head well developed. Dorsally frons and clypeus broad. Labrum
with row of coarse setae along its distal margin. In dorsal view labrum not completely covering
labium, with movable hooks and part of palps clearly visible. Mandibles very prominent in dorsal
view, their outer margins meeting at an angle of c.90; outer angle somewhat bulbous and bearing
dense setae. Antennae 7 segmented, mainly dark with pale marking basally on S1 and distally on
S2; overall moderately long. Length ratio of segments 1–7 as follows: 1.0 : 1.1 : 1.7 : 1.1 : 1.3 :
1.0 : 1.0. Head with rather low profile (Figure 3a); vertex scarcely rising above level of eyes; frons
slightly convex anteriorly but with median transverse groove; ante- and postclypeus broad and
well defined but almost in same plane, labrum slanted sharply downward. Labium (Figure 2b, c,
d, e) with prementum long and robust, the distal third expanded laterally with a rounded margin
bearing a row of 30+strong, dark, recurved spines. This row continues basad onto the dorsal
surface of the prementum with up to 10 more smaller spines, gradually diminishing in size. Most
spines are associated with a single long pale seta, inserted anteroventrally at the base of the spine
(Figure 2c). Lateral margins in basal two thirds of prementum taper evenly to a narrow base.
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A description of the larva of Heliaeschna idae 3
Figure 1. Heliaeschna idae exuvia, habitus.
Anterior margin of the prementum with two prominent lobes each occupying about one fifth of
its total width, separated centrally by a deep V-shaped incision, and also defined laterally by two
further incisions on each side, before the base of the labial palps which in dorsal view overlap a
second, smaller, outer pair of fleshy lobes, clearly seen in ventral view (Figure 2d). The inner lobes
have a thin, naked, blade-like rim centrally, extending slightly beyond the fleshy part of the lobe;
this rim ends in a well-defined step at approximately one third of the lobe width measured from
the central ‘V’, after which the margin continues smoothly bearing a row of 30+short stout setae
inserted ventrally and reaching the outer shoulder of the lobe (Figure 2d, e). The labial palps bear
long, fairly robust hooks, strongly curved inward apically and forming a blunt point; inner margin
with row of coarse serrations. Movable hooks long, slender and curved. Mandibles in ventral view
(Figure 2f) with protruding, outer corners forming an angle of c.90and moderately long, with
complex well-defined dentition arising from both incisor and molar surfaces, the former bifid on
the left, weakly trifid on the right, the latter strongly bifid on both sides.
Thorax. Relatively short; narrower than head. Pronotum shield-shaped, terminating in a blunt
point on each side which overlaps the supracoxal processes, just visible in dorsal view (Figure 1).
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4A.G. Orr et al.
Figure 2. Heliaeschna idae exuvia: (a) dorsal view of head, labium removed; (b) dorsal view of anterior part of
labium, with detail of lateral spines shown to left; (c) ventral detail of spines, showing associated setae; (d) ventral detail
of anterior part of prementum showing insertion of rows of setae along anterior lobes; (e) dorsal detail of anterior margin of
prementum; (f) ventral view of mandibles; (g) detail of prothoracic supracoxal processes; (h) dorsal view anal appendages;
(i) ventral view S9–S10 with gonapophyses.
Supracoxal processes robust, the anterior one finger-like, the posterior one broad and subquadrate,
the two lying close together and separated by a narrow ‘V’shape. Both processes covered in small
tubercles and bearing coarse, sparse setae basally and apically (Figure 2g). In the posterior part
of the thorax are slight protuberances above meso- and metacoxae, the intervening area being
saddle-shaped (Figure 1). Legs moderately long, distinctly banded. Wing sheaths reaching anterior
margin of S4 or a little beyond; distinctly banded.
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A description of the larva of Heliaeschna idae 5
Figure 3. Profile of head, labium and maxillae removed, showing tuft on vertex: (a) H. idae; (b) H. uninervulata; (c)
H. simplicia (redrawn from Butler & Orr, 2013).
Abdomen. Moderately high in profile, in section a somewhat flattened inverted catenary shape.
Dorsal spines absent. In dorsal view fusiform. S5–10 with ventrolateral spines, longest on S7–8
(Figure 1).
Appendages (Figure 2g) long; epiproct 1.6 times length of S10, terminating bluntly; row of small
spines along dorsum. Paraprocts slightly longer, turned slightly inwards apically and terminating
in a sharp point; outer surface with scattering of small blunt spines especially along outer margin;
overall with a few long fine setae and a sparse covering of fine short setae. Cerci relatively very
long, almost reaching end of epiproct, slightly hooked inward toward the tip and terminating in
a sharp point. Gonapophyses (Figure 2i) extend for full length of sternum 9 and overlap base of
S10; outer gonapophyses extending beyond tips of inner gonapophyses, their inner margin with
a subapical step.
Measurements (mm). Total length 39.0; head width 9.2; pronotum dorsal width 5.7; length of
paraprocts 3.0.
Habitat
The larva was found in a pool formed by a fallen log in an upland forest stream. The pool was
about 0.5 m deep, muddy and filled with leaf litter. The main stream from which the pool was
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6A.G. Orr et al.
formed had a sandy, gravely substrate. The surrounding vegetation was pristine, dense, tall and
mossy dipterocarp forest.
Heliaeschna uninervulata
Following the description of the larva of this species (Orr & Ngiam, 2011) the larva of H.
simplicia was discovered and described (Butler & Orr, 2013). H. simplicia presented an unusual
character, a swelling on the top of the vertex incorporating the ocellar mound bearing a dense
tuft of long, thick setae. The character is easily overlooked, and in one specimen the tuft was
poorly developed or damaged, but this prompted us to re-examine material of H. uninervulata,
specifically to check for this character.
Material examined
1exuvia (reared), collected as F-4 larva, 14 July 2010, Singapore, forest pool at Chestnut streams
within Central Catchment Nature Reserve, RWJN leg., emerged 12 September 2010. 1 exuvia
(reared), collected as F-3 larva, 23 December 2010, Singapore, forest pool at Chestnut streams
within Central Catchment Nature Reserve, RWJN leg., emerged 9 February 2011.
Head. Seen in profile upper side of head strongly raised, culminating in a slightly swollen
summit immediately behind ocelli (Figure 3b). This area bears a tuft of long thick setae.
Discussion
In almost all characters the larva of H. idae bears a strong resemblance to that of H. simplicia
and many of these characters are not shared with other Heliaeschna or Gynacantha species, with
which they share a general similarity in habitus. Both species have lateral spines on abdominal
S10 (lacking in H. filostyla and H. uninervulata); in both the anterior process of the supracoxal
armature is distinctly smaller than the posterior process and separated by a V-shape (a condition
approximately found also in H. filostyla but subequal and separated by a broad ‘U’in H. uninervu-
lata); in both the mandibles are exceptionally developed laterally forming an acute or nearly acute
outer angle (the mandibles are much weaker and distally rounded in H. uninervulata). However
the principal character which unites these two species is the form of the anterior margin of the
prementum. The blade-like inner bilobate structure flanked on either side by a well-defined row
of setae is of identical structure in both species, differing only in the relative widths of blade and
row of setae with respect to the lobe which bears them. Such a structure is, to the best of our
knowledge, unknown elsewhere in the Aeshnidae. It strongly suggests, along with other consid-
erations, that H. idae and H. simplicia, and presumably H. crassa, are congeneric. Conversely
these features suggest they are not congeneric with either H. filostyla described by Kawashima
and Sasamoto (2007) or H. uninervulata described by Orr and Ngiam (2011).
The main differences between H. idae and H. simplicia are: the cerci are relatively much longer
in H. idae; the shape of the supracoxal processes differs, especially the posterior process which
is subquadrate in H. idae rather than triangular as in H. simplicia; seen dorsally the head is more
robust in H. idae with the postocular lobes better developed; the mask is generally slightly more
sturdy, with the fixed hooks on the labial palps blunter and heavier, with very much coarser
serrations on their inner margin, suggestive of a different diet; the mandibles are quite similar in
their outer profile, the outer angle being only a little more acute in H. simplicia, but the internal
dentition is quite different, with H. idae having shorter incisor and molar processes, the latter being
bifid rather than a single long spine as in H. simplicia. The most obvious difference between the
two species is the presence in H. simplicia of a strongly elevated area of the vertex surmounted by
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A description of the larva of Heliaeschna idae 7
a tuft of long setae (Figure 3c). In H. idae this area is reduced to a low, slightly swollen convexity
and bears only a modest tuft of short dark setae.
The presence of a tuft on the vertex of H. uninervulata, outwardly similar to that found in H.
simplicia, raises a conundrum. With the possible exception of H. filostyla, all Heliaeschna larvae so
far known have a general form similar to that commonly found in Gynacantha, which is consistent
with the views of von Ellenrieder (2002), based on adult characters. In the case of H. uninervulata
the form of the mask falls easily with the range of variation known in Gynacantha (Matsuki,
1986a, 1986b; Theischinger, 2002, 2007; C.Y. Choong, pers. comm.), as do other characters. On
the other hand no Gynacantha has been recorded with a tuft of setae on the vertex. It might be
argued that the character is easily overlooked, and indeed this is the case, but we are informed
that a range of Gynacantha species in the collection of S.G. Butler bore no tuft (S.G. Butler,
pers. comm.). On this evidence, we suggest that the tuft may represent a synapomorphy, confined
to the Oriental Heliaeschna excepting H. filostyla (A. Sasamoto, pers. comm.), which is now
believed to lie well outside the GynacanthaHeliaeschna group (Kawashima & Sasamoto, 2007).
Its very modest development in H. idae may indicate it is quite a labile character, and while it
is strikingly similar in form in H. simplicia and H. uninervulata, the overall form of the head
and mouthparts in these species is very different. In particular in H. uninervulata the labial palp
bears five long, strong setae, a character shared, with variation in the number of setae, with all
known Gynacantha and Triacanthagyna species as presented by Heckman (2006), as well as
Austrogynacantha heterogena Tillyard (Theischinger, 2002). The securiform processes on the
labial palps of H. uninervulata are also typical of Gynacantha.
Therefore on present knowledge we should consider two taxonomic scenarios. Conservatively,
all African Heliaeschna plus H. uninervulata might be moved to Gynacantha. The remaining
Oriental species would include two new genera (because the type species of Heliaeschna is the
African H fuliginosa Selys), one including H. filostyla and another with the remaining species, H.
bartelsi,H. crassa,H. idae and H. simplicia. For this group the genus group name Malayaeschna
Foerster is available (Foerster 1909) with generotype H. simplicia. A less conservative approach,
and one which would reconcile the presence of a tuft of setae on its vertex, would be to erect a
new genus for H. uninervulata, as suggested by Orr and Ngiam (2011). Obviously further light
could be shed on this problem by DNA analysis, but as yet too little data is available to resolve
the problem raised here by considerations of larval morphology.
Acknowledgements
We thank in particular Stephen Butler, who donated a specimen of H. simplicia larva for comparison and provided much
helpful information and discussion, and Akihito Sasamoto and Gunther Theischinger for their critical comments and
helpful suggestions on the manuscript. RJWN and RAD are grateful to the Sarawak Forest Department and Sarawak
Forestry Corporation for granting permission to collect Odonata in Sarawak and to enter Usun Apau National Park.
RJWN was funded by the National Parks Board, Singapore. Fieldwork in Usun Apau National Park was also supported by
grants from The Mohamed bin Zayed Species Conservation Fund and the International Dragonfly Fund. RJWN and RAD
owe special thanks to Luke Southwell andYusof Tegong for their assistance with all aspects of the Usun Apau expedition,
and to Graham Reels for his contribution.
References
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(Odonata: Aeshnidae). International Journal of Odonatology,8, 1–32. doi: 10.1080/13887890.2005.9748240
Foerster,F. (1909). Beiträge zu den Gattungen undArten der Libellen. Jahrbücher des Nassauischen Vereins für Naturkunde
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Heckman, C.W. (2006). Encyclopedia of South American Aquatic Insects: Odonata – Anisopteras. Dordrecht: Springer.
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Kawashima, I., & Sasamoto, A. (2007). Descriptions of the last two instar larvae of Heliaeschna filostyla Martin, 1906
(Anisoptera, Aeshnidae, Aeshninae) from Sulawesi Island, Indonesia. Tombo, Matsumoto,49, 9–14.
Matsuki, K. (1986a). Description of the possible larva of Gynacantha hyalina Selys from Taiwan (Odonata, Aeshnidae).
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landesmuseum.at/biophp/de/arti_det.php?litnr=25388&artinr=59805
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... There is an additional complication in that H. idae was described from a female but H. crassa from a male, so it is at least possible that H. crassa is a junior synonym of H. idae and that the species referred to as H. idae here is actually unnamed. I can find no reliable means to separate females of these taxa at present, and for this reason the female Heliaeschna reared from a larva described as that of H. idae in Orr, Ngiam & Dow (2013) is regarded as of uncertain status here. Note 53. ...
... Note 53. Orr, Ngiam & Dow (2013), on the basis of larval characteristics, questioned the inclusion of this species in Heliaeschna. This is an issue that is only likely to be resolved through molecular analysis. ...
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The larva of Heliaeschna uninervulata is described and figured for the first time. Its characters mostly fall within the limits of variation of Gynacantha spp. Comparison of the larval characters of H. filostyla, the only other member of the genus for which the larva is known, suggests that it is not congeneric with H. uninervulata.
Article
Abstract A cladistic analysis of the world Aeshnidae is presented, based on fifty-eight characters of adult and larval anatomy. The ingroup taxa include all the extant genera of Aeshnidae, and the austropetaliid genera Phyllopetalia and Hypopetalia were chosen as the outgroup. The strict consensus tree obtained after successive weighting shows that the subgroups defined traditionally for Aeshnidae are paraphyletic or polyphyletic. The previous reclassification derived from analyses based on wing venation is supported in terms of the monophyly of Aeshnidae, Gomphaeschninae and its sister group comprising the remaining Aeshnidae. Gomphaeschninae is confirmed as sister group of the remaining Aeshnidae (= Aeshnodea Bechly). The sister-group relationships between Gomphaeschna+Sarasaeschna and Linaeschna+Oligoaeschna are corroborated. Within Aeshnodea, three monophyletic groups emerged: Boyeria + (Petaliaeschna+ (Limnetron+Gynacanthaeschna+Periaeschna)) + ((Cephalaeschna+Caliaeschna) + (Allopetalia (Notoaeschna+Spinaeschna))); Dendroaeschna+ (Epiaeschna+ (Aeschnophlebia+ (Nasiaeschna+ (Tetracanthagyna+Brachytron)))); and Polycanthagyna+ (Basiaeschna+ (Amphiaeschna+ (Indaeschna+ (Oplonaeschna+ (Racenaeschna+Plattycantha+Agyrtacantha+Triacanthagyna+ (Subaeschna+Austrogynacantha+Gynacantha) + (Heliaeschna+ (Neuraeschna+Staurophlebia))) + ((Castoraeschna+Coryphaeschna+Remartinia) + (Oreaeschna+ (Aeshna+ (Anaciaeschna+ ((‘A.’isosceles+Andaeshna) + (Anax+Hemianax)))))). Additional informative characters are required to test the relationships suggested here between the main groups of Aeshnodea and some enigmatic basal taxa (Antipodophlebia, Austroaeschna, Acanthaeschna, Telephlebia, Austrophlebia and Planaeschna).
Description of the possible larva of Gynacantha hyalina Selys from Taiwan (Odonata, Aeshnidae)
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The final instar larvae of Gynacantha rosenbergi Kaup and Antipodogomphus proselythus (Mar-tin) (Odonata, Aeshnidae & Gomphidae) Linzer biologische Beiträge Available fromA phylogenetic analysis of the extantAeshnidae (Odonata:Anisoptera)
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World Odonata List Retrieved from http://www.pugetsound Preliminary keys for the identification of larvae of the Australian Petaluridae
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Schorr, M., & Paulson, D. (2012). World Odonata List [Internet]. University of Puget Sound, WA. Retrieved from http://www.pugetsound.edu/academics/academic-resources/slater-museum/biodiversity-resources/dragonflies/ world-odonata-list/ Theischinger, G. (2002). Preliminary keys for the identification of larvae of the Australian Petaluridae, Archipetaliidae, Austropetaliidae, Telephlebiidae and Aeshnidae (Odonata). Thurgoona, NSW: Cooperative Research Centre for Freshwater Ecology.
Descriptions of the last two instar larvae of Heliaeschna filostyla Martin
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Dordrecht: Springer. Downloaded by [Princeton University] at 03:59 26 September 2013 238 A.G. Orr et al. Kawashima, I., & Sasamoto, A. (2007). Descriptions of the last two instar larvae of Heliaeschna filostyla Martin, 1906 (Anisoptera, Aeshnidae, Aeshninae) from Sulawesi Island, Indonesia. Tombo, Matsumoto, 49, 9–14.