Systematics and phylogeny of the Australian cicada genus Pauropsalta Goding and Froggatt, 1904 and allied genera (Hemiptera: Cicadidae: Cicadettini)

2016·58.9 MB·

by  Christopher L Owen

© The Authors, 2016. Journal compilation © Australian Museum, Sydney, 2016
Records of the Australian Museum (2016) Vol. 68, issue number 4, pp. 117-200.
ISSN 0067-1975 (print), ISSN 2201-4349 (online)
http://dx.doi.Org/10.3853/j.2201-4349.68.2016.1598
Systematics and Phylogeny of the Australian Cicada
Genus Pauropsalta Goding and Froggatt, 1904
and Allied Genera (Hemiptera: Cicadidae: Cicadettini)
Christopher L. Owen and M. S. Moulds
*12 3
1 University of Connecticut, Department of Ecology and Evolutionary Biology,
75 North Eagleville Road, Storrs, CT 06269, United States of America
2 Current address: George Washington University, Computational Biology Institute,
Innovation Hall, 45085 University Drive, Ashbum, VA 20147, United States of America
3 Research Fellow, Australian Museum Research Institute, 1 William Street Sydney NSW 2010, Australia
[email protected] • [email protected]
Abstract. A revision of the Australian cicada genera Pauropsalta and Palapsalta is presented. Twenty-
two new species and five new genera are proposed with supporting evidence from a cladistic analysis of
thirty morphological characters and a maximum likelihood molecular phylogeny that includes five loci.
Most species descriptions include male genitalia drawings, distribution maps, and male calling song to
aid in the identification of species. Furthermore, we include a key to all male and female species within
Pauropsalta and the newly described genera. The following five genera are described as new: Atrapsalta
n.gen., Haemopsalta n.gen., Falcatpsalta n.gen., Relictapsalta n.gen., and Popplepsalta n.gen. The
following 22 species are described as new: Atrapsalta emmotti n.sp., Atrapsaltafurcilla n.sp., Atrapsalta
vinea n.sp., Haemopsalta flammeata n.sp., Haemopsalta georgina n.sp., Popplepsalta aeroides n.sp.,
Palapsaltapalaga n.sp., Palapsalta serpens n.sp., Pauropsalta accola n.sp., Pauropsalta adelphe n.sp.,
Pauropsalta agasta n.sp., Pauropsalta confinis n.sp., Pauropsalta conflua n.sp., Pauropsalta contigua
n.sp., Pauropsalta ewarti n.sp., Pauropsalta herveyensis n.sp., Pauropsalta juncta n.sp., Pauropsalta
katherina n.sp., Pauropsalta kriki n.sp., Pauropsalta similis n.sp., Pauropsalta sinavilla n.sp., and
Uradolichos rotunda n.sp.
Keywords: Pauropsalta; Cicadettini; cicadas; taxonomy; morphology
Owen, Christopher L., and M. S. Moulds. 2016. Systematics and phylogeny of the Australian cicada genus
Pauropsalta Goding and Froggatt, 1904 and allied genera (Hemiptera: Cicadidae: Cicadettini). Records of the
Australian Museum 68(4): 117-200. http://dx.doi.Org/10.3853/j.2201-4349.68.2016.1598
* author for correspondence
118 Records of the Australian Museum (2016) Vol. 68
Pauropsalta Goding & Froggatt, 1904, is currently the largest there. From the original description it would appear that the
genus of Australian cicadas with 26 described species. Those species is allied to Atrapsalta encaustica. The identity of the
species belonging to Pauropsalta have been distinguished type is further clouded because it is a female and it is unlikely
from all other genera in having the following: (a) an that it could be associated with a male to resolve its identity.
infuscation on the hind wing at the distal end of vein 2A; (b) Consequently the name is here treated as a nomen dubium.
complete fusion or abutted forewing veins M and CuA; (c)
The following species have been recently transferred from
head never wider than the mesonotum; (d) exceptionally large
Pauropsalta to other genera, or have been synonymized by
upper pygofer lobes; and (e) ornamented apex of endotheca
Moulds (2012) and consequently are not treated here:
(Moulds, 2012). Those few species currently placed in
Pauropsalta from outside Australia, viz. New Caledonia, basalis Goding & Froggatt, 1904, to Nanopsalta Moulds, 2012
India, and Asia, do not possess the above attributes and will bellatrix Ashton, 1914, to Physeema Moulds, 2012
be treated in future studies. This work forms part of a larger
dubia Goding & Froggatt, 1904, to Platypsalta Moulds, 2012
study of Pauropsalta by the senior author, which will examine
emma Goding & Froggatt, 1904, to Mugadina Moulds, 2012
molecular relationships among genera and species and
explore the tempo and mode of Pauropsalta species as they lineola Ashton, 1914, —Dipsopsalta signata (Distant, 1914)
radiated through Australia. Here we address the taxonomy of nodicosta Goding & Froggatt, 1904, to Clinata Moulds, 2012
Pauropsalta and Palapsalta Moulds, 2012, a closely related signata Distant, 1914, to Dipsopsalta Moulds, 2012
genus, and present a cladistic analysis based upon morphology
Terminology for morphological features and higher
of all described species from which we identify attributes
classification follows those of Moulds (2005). The following
defining new genera and assess intergeneric relationships. In
abbreviations have been used for collections housing
addition, we provide song analyses and molecular evidence
from 5 genes (1 mtDNA, 4 nDNA; Owen et al, 2015) in specimens: AE, collection of A. Ewart; AJE, collection of
support of species and generic differentiation. A. J. Emmott, Noonbah Station; AM, Australian Museum,
The genus Pauropsalta was poorly documented until Sydney; ANIC, Australian National Insect Collection,
Ewart (1989) reviewed the Queensland species of the genus, Canberra; DE, collection of David Emery, Sydney; GAD,
describing 10 new species and providing notes on others. collection of G. A. D. Davis, Hobart; HOPE, Hope
Entomological Collections, Oxford University Museum,
Moulds (1990) provided an overview of 13 species none of
which were those described by Ewart (1989) that had been Oxford; JM, collection of J. Moss, Brisbane; JO, collection
published only a few weeks earlier. Additional notes on of J. Olive, Malanda; LP, collection of Lindsay Popple,
Queensland Pauropsalta species not summarized by Ewart Brisbane; MM, Macleay Museum, University of Sydney;
(1989) or Moulds (1990) (mainly of songs and distribution) MSM, collection of M. S. Moulds, Kuranda; MV, Museum
have been published by Ewart (1990, 1998a,b, 2001 a,b, of Victoria, Melbourne; NHM, Natural History Museum,
2005), Popple (2003), Popple & Ewart (2002), Popple & London; NTM, Museum and Art Gallery of the Northern
Strange (2002), Popple, Walter & Raghu (2007) and Moulds Territory, Darwin; PH, collection of Paul Hutchinson, Perth;
& Owen (2011). QM, Queensland Museum, Brisbane; SAM, South Australian
Despite the above publications, the identities of Museum, Adelaide; UCS, University of Connecticut, Storrs,
11 described species have remained vague, nothing USA; and WAM, Western Australian Museum, Perth.
having been published on them since their brief original
descriptions. The identities of these species are addressed
Methods
below as part of this review of the genus. We describe five
new genera and a further 22 new species. We do not address
those species previously well documented when we have Genitalia preparation and terminology
nothing new to add, although all species are included in our
Genitalia were dissected and prepared following the
cladistic and molecular analyses. A key to all known species
methodology of Moulds (2012). Terminology for genital
of Pauropsalta, Atrapsalta n.gen., Falcatpsalta n.gen.,
structures follows that of Moulds (2005) (Fig. 1). However,
Haemopsalta n.gen., Palapsalta, some Popplepsalta n.gen.,
the homologies of the basal pygofer lobe proved difficult to
Relictapsalta n.gen., and Uradolichos is provided. Song
interpret. Within Pauropsalta and its allied genera there are
analyses are provided for most new species as well as for
two lobed structures on the pygofer wall below the sclerital
those species whose songs were previously undocumented.
suture, the suture of the pygofer wall that indicates the point
The following two names could not be associated with
of fusion between abdominal tergite 9 and stemite 9 that
any species and are treated as nomen dubia in this work.
together form the pygofer. The basal pygofer lobe is that
Pauropsalta stigmatica Distant, 1905. The type is a female situated immediately below the sclerital suture as defined
from Adelaide and because there are no known associated by Moulds (2005).
males its identity cannot be resolved and consequently is In Pauropsalta and allied genera the basal lobe shows
here treated as a nomen dubium. The complete fusion of considerable variation and its identity can be confusing.
fore wing veins M and CuA beyond the basal cell and the In some species such as Pauropsalta ewarti n.sp. the basal
possession of 5 apical cells in the hind wing together with its lobe is large and rounded, clearly positioned immediately
small size, colouring and little to no infuscation on the hind below the sclerital suture and is easily identified (Figs
wing suggest that it may be most closely allied to species of 1A, IB). However, in other species such as Haemopsalta
the Pau. annulata species group. flammeata gen. et n.sp., the basal lobe turns inwards and is
Pauropsalta rubra (Goding & Froggatt, 1904). The type reduced to a small knob-like or spine-like structure (Figs
is a female from Sale, Victoria that according to the original 1C, ID). Simultaneously with reduction of the basal lobe a
description should be in MV. The specimen could not be traced secondary basal lobe, that is undeveloped in most cicadas,
and the Museum has no record of it ever having been housed attains prominence and appears as if it were the true basal
Owen & Moulds: Revision of the cicada genus Pauropsalta 119
Figure 1. Male genitalia: (A) lateral view, Pauropsalta ewarti n.sp.; (B) ventral view, same species; (C) lateral view,
Haemopsaltaflammeata n.gen., n.sp.; (C) ventral view, same species. Terminology after Moulds (2005). (as) anal
style); (cl) clasper; (db) dorsal beak; (end) endotheca; (mdl) median lobe of uncus; (pbl) pygofer basal lobe; (ps)
pseudoparamere; (pyg) pygofer; (sbl) secondary basal lobe; (th) theca; (upl) upper pygofer lobe.
lobe (Figs 1C, ID). A species showing intermediate transition removed from spectrograms to improve song clarity and
in reduction of the basal lobe and increasing development of enhance visualization. Furthermore, most depicted songs
the secondary basal lobe is Pauropsalta confinis (Fig. 16). include an enhanced portion of the waveform to show fine-
scale detail of song structure. Raw songs are available from
Song analyses the senior author upon request.
All song recordings were collected by Simon Lab (University
Type species designation and synonymies
of Connecticut) personnel; primarily, David C. Marshall,
John Cooley, and Kathy B. R. Hill. Songs were recorded in Where possible we have selected holotype specimens
the field with a Marantz PMD-660 compact flash recorder from voucher specimens representing DNA samples and
with a Sennheiser ME-62 omnidirectional microphone sometimes also with associated song recordings. It is our
mounted in a SONYPBR-330 parabolic reflector. Waveforms intention to clearly define holotype specimens as far as
and spectrograms were viewed in Raven Pro 1.4 (Cornell possible using morphological, molecular, and song data
Bioacoustics Laboratory, Ithaca, NY). For most species low in case that a species needs to be separated from potential
frequency noise outside the kilohertz range of the song was cryptic sister species (e.g., Marshall etal., 2011).
120 Records of the Australian Museum (2016) Vol. 68
Uradolichos
Figure 2. Character 1. Postclypeus in dorsal view: (0) substantially protruding, almost semicircular in outline when viewed
from above; (1) tending confluent with anterior margin of head, tending angular in outline when viewed from above.
Listing of synonymies for previously described species taken from those identified as relevant for defining the genus
have been restricted to primary entries only as detailed entries Pauropsalta by Moulds (2012). Four previously described
with all literature references are available elsewhere. In some genera with similar morphological features, and known to be
cases literature not previously listed in earlier published closely allied to Pauropsalta by DNA (Owen et al., 2015),
synonymies is provided. have been included in the analysis to help clarify nodes
representing putative new genera.
Justification for species
Cladistic analysis
The delineation of species is supported by three sources
of data; DNA (Owen et al., 2015), morphology, and song, All described Australian Pauropsalta species together
although DNA and/or song were not available for all species. with those new species described in this monograph, plus
First, 460 specimens were sequenced for c. 800 bp of the exemplars from Graminitigrina, Gudanga, Nanopsalta,
3'COI gene and sequences were run in a Maximum Like¬ Palapsalta, and Uradolichos were scored for the following
lihood analysis (see above) to identify clusters of specimens 30 multistate and binary characters believed to be meaningful
(Owen et al, 2015). The outward appearance of individuals at generic level.
in these clusters was then used to aid in sorting approximately
5,000 collection specimens to putative species. Characters
Once sorted, 450 genitalia preparations were made 1 Postclypeus in dorsal view. (0) substantially
from representatives of all putative species. The genitalia protruding, almost semicircular in outline when
preparations were used as additional evidence in confirming viewed from above (Fig. 2); (1) tending confluent
the species status of the putative species. Described species with anterior margin of head, tending angular in
were then identified from among the putative species
outline when viewed from above (Fig. 2).
recognized by comparison with type material (including male
2 Fore wings: (0) entirely suffused black; (1) entirely
genitalia where practical). Remaining species groupings
hyaline or almost so.
were thus considered new species. Once specimens were
3 Fore wing veins Mand CuA : (0) not touching before
sorted according to DNA and morphology, available song
reaching basal cell; (1) completely fused as one before
recordings were compared. Songs are known to be species
reaching basal cell; (2) abutted for some distance
specific (Marshall et al., 2008) and are thus useful in
before reaching basal cell.
delimiting cryptic species. Thus, based on our sampling we
believe the species we describe in this study represent unique 4 Costa colour. (0) dark brown to black; (1) red.
taxa that do not share identical molecular, morphological, 5 Hind wing apical cells'. (0) 6 cells; (1)5 cells.
and acoustical characteristics with any previously described Note: species with 5 hind wing apical cells usually
species and warrant description. have a small number of individuals with 4 or 6 apical
cells (usually only in one wing). Such specimens are
Justification for genera considered to have 5 apical cells and those minority
individuals with 4 or 6 are regarded as abnormal.
The genera described in this study are based on the results
of a cladistic analysis (detailed below) of a novel dataset of 6 Hind wing apical cell F. (0) of usual size, more than
half the length of apical cell 2; (1) very small, far less
30 binary and multistate characters. This suite of characters
than half the length of apical cell 2.
demonstrates common ancestry among groups of species
examined and provides characters for which we define each 7 Male abdomen'. (0) about as long or shorter than head
new genus. Other characters used for defining genera were plus thorax; (1) longer than head plus thorax.
Owen & Moulds: Revision of the cicada genus Pauropsalta 121
Haemopsalta flammeata n.gen., n.sp. Pauropsalta mneme
Figure 3. Character 13. Upperpygofer lobe shape in lateral view. (0) dorsal margin with basal portion strongly angled to axis of pygofer;
very broad basally and tapering to a broad apex; (1) dorsal margin with basal portion nearly aligned with axis of py gofer; broad basally
and tapering to a broad apex; (2) dorsal margin with basal portion nearly aligned with axis of pygofer; narrow basally so that upper py gofer
lobe tends towards a linear structure with the distal portion slightly tilted downward.
8 Male abdomen. (0) about as wide as thorax; (1) very pygofer; very broad basally and tapering to a broad
wide, much wider than thorax. apex (Fig. 3); (1) dorsal margin with basal portion
9 Male abdomen. (0) ridged along dorsal midline; (1) nearly aligned with axis of pygofer, broad basally
and tapering to a broad apex (Fig. 3); (2) dorsal
broadly rounded across dorsal half.
margin with basal portion nearly aligned with axis of
10 Male abdomen. (0) epipleurites reflexed inwards;
pygofer; narrow basally so that upper pygofer lobe
(1) epipleurites not flexed inwards, abdomen below
tends towards a linear structure with the distal portion
rounded in cross section.
slightly tilted downward (Fig. 3).
11 Pygofer caudal beak. (0) broad, short, not well
14 Pygofer basal lobe: (0) flap-like, sometimes turned
developed; (1) spine-like, well developed.
inwards (Fig. 4); (1) spike-like (Fig. 4); (2) peg with
12 Upper pygofer lobe \ (0) single lobed; (1) bi-lobed. rounded distal end (Fig. 4).
13 Upper pygofer lobe shape in lateral view. (0) dorsal 15 Pygofer secondary basal lobe shape: (0) fold-like, in
margin with basal portion strongly angled to axis of lateral view projecting and broadly rounded, in ventral
Pauropsalta ewarti n.sp. Pauropsalta dolens Haemopsalta flammeata n.gen., n.sp.
Figure 4. Character 14. Pygofer basal lobe: (0) flap-like, sometimes turned inwards; (1) spike-like; (2) peg with rounded distal end.
122 Records of the Australian Museum (2016) Vol. 68
Pauropsalta n r aquila Pauropsalta emmotti n.sp.
Figure 5. Character 15. Pygofer secondary basal lobe shape: (0) fold-like, in lateral view projecting and broadly rounded, in ventral view
either closely aligned with py gofer margin or projecting to varying extent but always totally fused to pygofer margin at rear; (1) plate-like
and well developed, in lateral view clearly projecting and subtriangular with a broad base tapering to a slightly upturned pointed apex, in
ventral view long and spike-like; (2) tending fold-like, in lateral view broadly angled and moderately projecting, in ventral view broad
and rounded; (3) lobe like and domed, in lateral view not or moderately projecting and broadly rounded, in ventral view well developed
with rounded apex that is sometimes two-tiered and fusion with pygofer margin not reaching apex.
view either closely aligned with pygofer margin or dorsal midline; (2) broad, short, distal margin straight;
projecting to varying extent but always totally fused (3) triangular and depressed along dorsal midline; (4)
to pygofer margin at rear (Fig. 5); (1) plate-like and triangular with dorsal surface flat or domed.
well developed, in lateral view clearly projecting and
18 Claspers in lateral view. (0) long, projecting outward
subtriangular with a broad base tapering to a slightly
far beyond margin of pygofer, cavernous below for
upturned pointed apex, in ventral view long and
most of length; (1) extending no more than a little
finger-like (Fig. 5); (2) tending fold-like, in lateral
beyond margin of pygofer, claw-like with minimum
view broadly angled and moderately projecting, in
or no cavity below.
ventral view broad and rounded (Fig. 5); (3) lobe
like and domed, in lateral view not or moderately 19 Claspers'. (0) extreme basal region concave, not
projecting and broadly rounded, in ventral view well developed forward; (1) extreme basal region
developed with rounded apex that is sometimes two- projecting forward as a broad rounded lobe.
tiered and fusion with pygofer margin not reaching
20 Theca: (0) trifid, that is with a pair of dorsal pointed
apex (Fig. 5).
pseudoparameres and a pointed ventral support
16 Secondary basal lobe shape'. (0) outer face rounded; surrounding an exposed endotheca; (1) not trifid,
(1) outer face ridged longitudinally. Note: The with a pair of pseudoparameres but lacking a ventral
secondary basal lobe in Graminitigrina bowensis is support.
small and it is difficult to confirm the presence or
21 Pseudoparameres'. (0) robust, flattened or rounded in
absence of a ridge on the outer face; consequently, for
cross section, not hair-like; (1) hair-like.
this species the character has been scored as “?”.
17 Uncus in dorsal view. (0) linear, dorsal surface flat 22 Pseudoparameres. (0) arising independently at the
or domed; (1) broad, rounded, flat or depressed along base; (1) partly fused beyond the base.
Owen & Moulds: Revision of the cicada genus Pauropsalta 123
Popplepsalta aeroides n.gen, n.sp. Pauropsalta extrema Palapsalta eyrei
Figures 6-8. Characters 25, 29 and 30. (Fig. 6) Character 25, Endotheca shaft. (1) trumpet-shaped in apical region.; (Fig. 7) Character
29, Endotheca apex. (1) with lateral “wing-like” flanges; (Fig. 8) Character 30, Endotheca apex. (1) with a spine-like projection at each
lateroventral corner.
23 Pseudoparameres in lateral view. (0) adjacent to sequence addition (RSA) and 1000 replicate tree searches
or close by theca and not arched high above it; (1) were performed from different starting trees to ensure
arched high above a nearly straight theca. searches did not get stuck in a local optimum. Trees were
24 Pseudoparameres'. (0) distal ends turned outwards; (1) then loaded into CLADOS version 1.2 (Nixon, 1992) to plot
distal ends straight; (2) distal ends turned inwards. character state transformations along branches according
to DELTRAN optimization and from which the final tree
25 Endotheca'. (0) shaft parallel-sided in apical region;
was printed. Branch support was estimated using 1000
(1) shaft trumpet-shaped in apical region (Fig. 6).
bootstrap replications retaining a maximum of 500 trees at
26 Endotheca in lateral view. (0) about as broad as or
each replication.
broader than pseudoparameres; (1) exceedingly thin,
narrower than pseudoparameres.
Molecular phylogeny
27 Endotheca'. (0) dorsal region sclerotized similar to
that of lateral and ventral surfaces; (1) dorsal region Owen et al. (2015) recently estimated the molecular
sclerotized less than lateral and ventral surfaces; phylogeny of Pauropsalta and the new genera described in
(2) much of surface weakly sclerotized and in part this study; therefore, we defer to the molecular phylogeny
translucent. within Owen et al. (2015) to portray the relations among
28 Endotheca apex'. (0) with margin either lacking species and genera addressed in this study. Briefly, the
serrations or partly serrated but not continuously molecular data set included all species in the cladistic
around lateral and ventral margins; (1) with margin analysis except: Pauropsalta rubea, Pauropsalta fuscata,
continuously serrated around lateral and ventral Pauropsalta ayrensis, and Uradolichos longipennis. Fresh
margins. tissue was unavailable for those species. The molecular
sequence data set included 1 mtDNA locus and 4 nDNAloci:
29 Endotheca apex\ (0) without lateral “wing-like”
cytochrome oxidase c subunit 1 (COI), elongation factor
flanges; (1) with lateral “wing-like” flanges (Fig. 7).
1 alpha (EFla), period (Per), acetyltransferase (ARD1),
30 Endotheca apex'. (0) without spine-like projection
and glutaminyl tRNA synthetase (QtRNA). The nucleotide
at each lateroventral corner; (1) with a spine-like
sequences were aligned in Mesquite v.2.75 (Maddison and
projection at each lateroventral corner (Fig. 8).
Maddison 2011) using MUSCLE (Edgar, 2004) and adjusted
The species incorporated in the cladistic analysis are listed by eye. The alignment was partitioned by locus and codon
in Table 1 together with the scoring of their character states. position if a protein-coding gene and partitions were assigned
Uncertainty in a characters state for a species was scored a model of evolution using PartitionFinder v. 1.1.1 (Lanfear
as “?”. Gudanga boulayi was chosen as an outgroup taxon et al., 2012). We estimated a maximum likelihood (ML)
based on the sister relationship of Gudanga to the lineage phylogeny in Garli (Zwickl, 2006) using default parameters
examined in this study (Moulds, 2005; Marshall et al., and 15 independent searches starting from different starting
2016). A parsimony analysis was performed in PAUP* 4bl0 trees. Branch support was estimated using 100 nonparametric
(Swofford, 2003) using the tree bisection-reconstruction bootstrap replicates (BS) with 2 independent tree searches,
(TBR) algorithm with all characters treated as unweighted each from random starting trees, for each replicate. The tree
and unordered. Starting trees were generated with random was rooted with Gudanga boulayi.
124 —Gudanga boulayi Gudanga
8 4-C Uradolichos longipennis
Uradolichos
0>10>10>10>1100>> 1‘— Uradolichos rotunda sp. n.
—Pauropsalta mneme
— Pauropsalta castanea
-Pauropsalta conflua sp. n.
-Pauropsalta juncta sp. n.
51 —Pauropsalta accola sp. n.
-Pauropsalta infuscata
o> 1
—Pauropsalta contigua sp. n.
63
66 —Pauropsalta confinis sp. n.
23
5 20 -Pauropsalta extensa
H-H—
0>10> 10>I 0>1 8 7 -
2£44 2266 1,-hP auropsalta katherina sp. n.
o> io> i ’-FP auropsalta sinavilla sp. n. Pauropsalta
—Pauropsalta elgneri
•Pauropsalta opaca
¥Pauropsalta ewarti sp. n.
—Pauropsalta hen/eyensis sp. n.
50 —Pauropsalta infrasila
73 13 21 29
"I—Pauropsalta walked
41—0— 70 ¥Pauropsalta adelphe sp. n
o>n>o y
I .1—
o> 1 rP auropsalta agasta sp. n.
Pauropsalta extrema
—Pauropsalta similis sp. n.
Pauropsalta kriki sp. n.
—Pauropsalta melanopygia
—Pauropsalta borealis
-|—Nanopsalta basalis Nanopsalta
0>1 15 27
—!>|—21|>—2 PauropsaltaQ aq3u ilus | Falcatpsalta
70 4 l-1-P auropsalta rubristrigata
15 17 83 pf-T
+-h 5 28 0 > 1*-¥lP auropsalta aero/des sp. n.
Popplepsalta
Pauropsalta annulata
0> 1
Pauropsalta ayrensis
18
MHI—
D Palapsalta serpens sp. n.
i >oo> 1
62
-Palapsalta eyeri
17 22 4-C
-Mb O > 1*-¥1P alapsalta palaga sp. n. Palapsalta
—Palapsalta virgulata
70 66
18 Palapsalta circumdata
i- s-f-H
0>1 1 >30> l1--rP alapsalta vitellina
90 -Pauropsalta aktites
414— 19| - -Haemopsalta georgina gen. n. sp. n.
L6j3_ | Pauropsalta rubea Haemopsalta
o> v—¥tH aemopsalta flammeata gen. n. sp. n
-Pauropsalta nigrastriga Relictapsalta
8 21 24 29
-1—ff—|—§— Graminitigrina bowensis I Graminitigrina
5 16 0>10>10>10>1 1
4—I— 22 23 27
o> io>i —^J— |]— ¥[]—Pauropsalta dolens
0> 10> 12 > O
— Atrapsalta emmotti gen. n. sp. n.
Figure 9. One of the three most parsimonious trees (length 56, Cl 67, —- Pauropsalta siccana
RI 92) from an analysis of all Pauropsalta species, both described
— Pauropsalta fuscata
and named as new in this paper, together with exemplars from
four described genera, Graminitigrina, Gudanga, Nanopsalta and —Atrapsalta vinea gen. n. sp. n. Atrapsalta
Uradolichos. Clades representing described genera and new genera are 0> 1
labeled as such. Numbers above branches are bootstrap percentages > — Pauropsalta corticina
50 from 1000 bootstrap replicates. Character transformations generated
Pauropsalta colli na
in Clados shown on branches (include autapomorphies): black bars =
non-homoplasious forward change; grey bars = homoplasious forward — Atrapsalta furcilla gen. n. sp. n.
change; white bars = reversal (whether homoplasious or not).
— Pauropsalta encaustica
Owen & Moulds: Revision of the cicada genus Pauropsalta 125
Molecular and morphological phylogenetic confidence in the generic groupings of the morphological tree.
In the molecular ML phylogeny (Fig. 10) all clades
results and generic groupings
representing described and proposed genera are strongly
Results from the morphological cladistic analysis found 24 supported except Graminitigrina. The ML phylogeny
shortest trees from which a strict consensus tree placed 23 placed Graminitigrina bowensis in a clade (< 70 BS) that
of the Pauropsalta species as unresolved near the base of the includes Pauropsalta corticina, P. siccana, P. dolens, P.
tree that was otherwise reasonably well resolved. However, collina, P. encaustica, and three new species we describe
by de-activating character 21 just three equally parsimonious in this study. The non-monophyly of Graminitigrina may
trees of length of 56, Cl 67 and RI92 were recovered, all with be due to poor taxon sampling for the genus, which is only
the 23 species previously unresolved clustered into a single represented in our molecular study by one of five described
clade and with no other changes in tree morphology. The species (Fig. 10; Ewart & Marques, 2008). Although branch
three trees were identical except for minor differences in the support for Graminitigrina is lacking in the molecular and
placement of Pauropsalta accola n.sp. within Pauropsalta morphological phylogenies, four apomorphies (all be it
and one of these trees is shown in Fig. 9. In this tree clades homoplasious ones) in the cladistic analysis distinguish
representing described genera, and proposed new genera, Graminitigrina from all other species (Fig. 9). Therefore, we
are labelled as such along with bootstrap values. do not dispute the identity of Graminitigrina. Although no
The cladistic analysis recovered all previously described apomorphy could be found to unite the remaining species in
genera as monophyletic except Pauropsalta. Pauropsalta the Graminitigrina clade, they are remarkably similar to each
species were distributed among six clades, five of which we other given the characters we used, and we consider these to
consider to be new genera. Most described genera and clades represent a new genus Atrapsalta, sister to Graminitigrina.
recovered lack branch support, which is most likely due to In conclusion, the generic decisions made here have
a paucity of characters. However, and most importantly, all been based on both the phylogenetic morphological and
the clades representing described genera and proposed new molecular analyses, which together support all proposed new
genera are well supported by a comprehensive molecular genera and elucidate the placement of species in previously
analysis described below and it is on this basis we have described genera.
-Pau. rubristrigata 06.AU.VI.CAW.01
Popplepsalta n. gen.
Po. aeroides n. sp. 06.AU.NS.BES.03
Pau. annulata 05.AU.QL.CAM.02
1001 Pal, eyeri 05.AU.QL.BWC.01
Pal. palaga n. sp. 07.AU.NT.TIL.02
Pal. serpens n. sp. 04.AU.NT.WSM.12 Palapsalta
Pal. vitellina 08.AU.QL.SWN.10
Pal. circumdata 02.AU.QL.LBW.09
Pal. virgulata 05.AU.QL.RAW.01
Pau. corticina 09.AU.NS.OCO.01
A. emmottin. sp. 08.AU.QL.JNW.01
Pau. siccana 08.AU.QL.WBS.01
— Pau. dolens 03.AU.WA.BOY.04 Atrapsalta n. gen.
— Pau. collina 09.AU.QL.GWD.03
Pau. encaustica 04.AU.NS.DHA.02
_A furciila n. sp. 09.AU.VI.NCC.02
A.vinean. sp. 07.AU.SA.CLA.01
Graminitigrina
Gr. bowensis 08.AU.QL.EMK.01 |
Pau. nigrastriga 04.AU.QL.DED. 16 1 R" elictapsalta n. gen.
Haemopsalta n. gen.
Falcatpsalta n. gen.
Nanopsalta
n. basaiis 07.Au.QL.wEN.di |
Pau. kriki n. sp. 08.AU.NT.EDT.il
Pau. melanopygia 06.AU.NTRBM.01
Pau. borealis 04.AU.NT.NKA.10
100Pau. agasta n. sp.10.AU.WA.HCD.01
Pau. adelphe n. sp. 04.AU.NT.KWW.04
Pau. similis n. sp. 06.AU.WA.TSE.03
Pau. extrema 06_AU_WA_SFR_01
Pau. herveyensis n. sp. 09.AU.QL.HER.01
Pau. ewarti n. sp. 04.AU.QL.HMT.13
Pau. opaca 08.AU.QL.DED.01
Pau. infrasila 07.AU.QL.SPL.01 Pauropsalta
Pau. elgneri 07.AU.QL.ARR.01
Pau. walked 04.AU.NT.CCR.10
Pau. confinis n. sp. 06.AU.WA.WSS.06
Pau. contigua n. sp. 07.AU.SA.MRA.01
Pau. accola n. sp. 03.AU.WA.PAU.03
Pau. sinavilla n. sp. 09.AU.WA.HOU.15
Pau. katherina n. sp. 06.AU.WA.MRS.03
Pau.juncta n. sp. 03.AU.WA.SAR.02
Pau. castanea 07.AU.SA.HWK.01
Pau. conflua n. sp. 03.AU.WA.PAU.01
Pau. mneme 06.AU.NS.DHA.01
U. rotunda n. sp. 06.AU.WA.OPS.03 Uradolichos
Gu. boulayi 03.AU.WA.LKD.10 | Gudanga
—oT~
Figure 10. Maximum likelihood tree estimated using Garli 2.0 with 5 loci (1 mtDNA and 4 nDNA). Branch support values are bootstrap
percentages from 100 non parametric bootstrap replicates. Bootstrap support values > 70 are shown. Molecular voucher numbers are
adjacent to species names.
126 Records of the Australian Museum (2016) Vol. 68
Taxonomy
Family Cicadidae Latreille
Subfamily Cicadettinae Buckton
Tribe Cicadettini Buckton
Summary of genera treated in this paper with their included species
Pauropsalta Goding & Froggatt, 1904 Palapsalta Moulds, 2012
accola n.sp. circumdata (Walker, 1852)
adelphe n.sp. eyrei (Distant, 1882)
agasta n.sp. palaga n.sp.
borealis Goding & Froggatt, 1904 serpens n.sp.
castanea (Goding & Froggatt, 1904) virgulata Ewart, 1989
confinis n.sp. vitellina Ewart, 1989
conflua n.sp.
Popplepsalta n.gen.
contigua n.sp.
aeroides n.sp.
elgneri Ashton, 1912
annulata (Goding & Froggatt, 1904), n.comb.
ewarti n.sp.
ayrensis (Ewart, 1989), n.comb.
extensa Goding & Froggatt, 1904
blackdownensis (Popple, 2013), n.comb.
extrema (Distant, 1892)
corymbiae (Popple, 2013), n.comb.
herveyensis n.sp.
decora (Popple, 2013), n.comb.
infrasila Moulds, 1987
granitica (Popple, 2013), n.comb.
infuscata (Goding & Froggatt, 1904)
inversa (Popple, 2013), n.comb.
juncta n.sp.
kobongoides (Popple, 2013), n.comb.
katherina n.sp.
notialis incitata (Popple, 2013), n.comb.
kriki n.sp.
notialis notialis (Popple, 2013), n.comb.
melanopygia (Germar, 1834)
rubristrigata (Goding & Froggatt, 1904), n.comb.
mneme (Walker, 1850)
simplex (Popple, 2013), n.comb.
opaca Ewart, 1989
subtropica (Popple, 2013), n.comb.
similis n.sp.
torrensis (Popple, 2013), n.comb.
sinavilla n.sp.
tremula (Popple, 2013), n.comb.
walkeri Moulds & Owen, 2011
Falcatpsalta n.gen.
Atrapsalta n.gen.
aquilus (Ewart, 1989), n.comb.
collina (Ewart, 1989), n.comb.
corticina (Ewart, 1989), n.comb. Relictapsalta n.gen.
dolens (Walker, 1850), n.comb. nigristriga (Goding & Froggatt, 1904), n.comb.
emmotti n.sp.
Uradolichos Moulds, 2012
encaustica (Germar, 1834), n.comb.
longipennis (Ashton, 1914)
fuscata (Ewart, 1989), n.comb.
rotunda n.sp.
furcilla n.sp.
siccana (Ewart, 1989), n.comb.
vinea n.sp.
Haemopsalta n.gen.
aktites (Ewart, 1989), n.comb.
flammeata n.sp.
georgina n.sp.
rubea (Goding & Froggatt, 1904), n.comb.
Key to species of Pauropsalta and allied genera
Pauropsalta and allied genera share a sclerotized endotheca that lacks a ventral support and all share an
infuscation at the distal end of hind wing vein 2A except most species in the genus Palapsalta.
1 Fore wing veins M and CuA with their stems meeting the basal
cell completely fused as one (view carefully under magnification). 29
- Fore wing veins M and CuA with their stems meeting the basal
cell closely abutted or parted, not fused as one.2