Revision and Analysis of Pseudosaldula Cobben (Insecta: Hemiptera: Saldidae): A Group with a Classic Andean Distribution

2009·78.5 MB·English

by  Randall T. Schuh

REVISION AND ANALYSIS OF PSEUDOSALDULA
COBBEN (INSECTA: HEMIPTERA: SALDIDAE): A
GROUP WITH A CLASSIC
ANDEAN DISTRIBUTION
RANDALL T. SCHUH
Curator
Division of Invertebrate Zoology
American Museum of Natural History
([email protected])
JOHN T. POLHEMUS
Research Associate
Division of Invertebrate Zoology
American Museum of Natural History
([email protected])
BULLETINOFTHEAMERICANMUSEUMOFNATURALHISTORY
Number323, 102 pp., 31figures, 4tables, 8plates
Issued June 30,2009
CopyrightEAmericanMuseumofNaturalHistory2009 ISSN0003-0090
CONTENTS
PART 1: SYSTEMATICS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
RANDALL T. SCHUHANDJOHN T. POLHEMUS
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Materials and Methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Pseudosaldula Cobben. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Key to the species of Pseudosaldula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Pseudosaldula andensis (Distant). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
Pseudosaldula antioquia, new species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Pseudosaldula aurea, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Pseudosaldula bergi (Haglund) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
Pseudosaldula bruesi (Drake) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
Pseudosaldula chilensis (Blanchard). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
Pseudosaldula huamachuco, new species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Pseudosaldula penai, new species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
Pseudosaldula perula (Drake) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
Pseudosaldula pilosa, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
Pseudosaldula salina, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Pseudosaldula saxicola, new species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Pseudosaldula vulgaris, new species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Pseudosaldula yungas, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
PART 2: PHYLOGENETIC AND BIOGEOGRAPHIC ANALYSES. . . . . . . . . . . . . 69
RANDALL T. SCHUH, PAOLA PEDRAZA, ANDJOHN T. POLHEMUS
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
Materials and Methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
Morphological Character Data . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
Discussion of Selected Characters. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
Molecular Character Data. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
Selection of Outgroup Taxa and Rooting of Cladograms. . . . . . . . . . . . . . . . . . . . . 76
Phylogenetic Methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
Phylogenetic Results. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
Ecological Observations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
Historical Biogeographic Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
Plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
PART 1:
SYSTEMATICS
RANDALL T. SCHUH AND JOHN T. POLHEMUS
ABSTRACT
The genus PseudosaldulaCobben, which is restricted to the Andean Subregion of South
America, is revised. Fourteen valid species are recognized, nine of them being described as
new and 10 previously published names are treated as junior synonyms based on the
examination of approximately 3500 specimens from Colombia, Ecuador, Peru, Bolivia,
Argentina, and Chile. All taxa are described or redescribed. A key to the species is
provided. Color habitus illustrations, distributional maps, and detailed measurements are
provided for all species. Scanning electron micrographs of the vestiture, parameres,
parandria, face, and pretarsus are provided for representative species, as are color views of
thefaceandthenymphs.TheconceptofapostclypeusintheSaldidaeisquestionedandthe
term transverse swelling, as coined by Parsons, is applied in discussing distinctive aspects of
facial morphology in Pseudosaldula. A previously unreported, presumably glandular, pore is
documented on the parameres in the Saldinae in the form of a cavernous pit with internal
digitiform processes.
Aphylogeneticanalysisbasedonmorphologicalcharacterdatadocumentsthemonophylyof
Pseudosaldula. Characters treated as synapomorphic for Pseudosaldula are five cells in the
membrane of the forewing, the incomplete connection of the transverse swelling across the
posterior margin of the clypeus, and the straight connection across the posterior margin of
the parandria; nymphal coloration is also distinctive, although treated as ambiguous
because this character was not scored for all species. DNA sequence data from the 16S
rDNA region of the mitochondrion and H3 nuclear region were acquired for 13
Pseudosaldula spp. and five outgroup taxa. The combined analysis of morphological and
sequence data consistently treated Pseudosaldula as paraphyletic. These results are interpreted
astheresultofinadequatesamplingofbothtaxaandgeneregions,inlightofthefactthatthe
patternsofdistributionbecometranspacific,asopposedtoamonophyleticgroupintheAndean
Region. Not unexpectedly, several morphological characters documenting the monophyly of
Pseudosaldula show greater homoplasy in the combined analysis than when analyzing
morphological data alone. Therefore, the results of the morphological cladistic analysis are
further used to examine distributional patterns in the group. Five areas of endemism are
recognized: northern Andes, northern Peru, Puna, central Chile, and subantarctic; the
boundaries of these areas show substantial correspondence with those proposed for other
groups of insects.
INTRODUCTION placed in the family Aepophilidae (King and
Ratcliffe, 1970; Schuh and Polhemus, 1980),
The Saldidae is a worldwide group of and all members of the family Omaniidae
about 335 species that shows its greatest from the tropical Indo-Pacific (Cobben,
diversity in the Northern Hemisphere 1970).
(Schuh et al., 1987). The morphological Thetaxonomichistoryoftheshorebugsis
body plan within the Saldidae shows only complex and the features used to diagnose
limited variation across the group, whereas genera and suprageneric assemblages have
within-species morphological variation can frequently not allowed for the recognition of
be extreme. All species of Saldidae are monophyletic groupings. The details of
predatory and live on damp substrates. morphology in the Saldidae—and Leptopo-
Species in a number of phyletic lineages— domorpha more broadly—were treated in a
including other members of the Leptopo- largely superficial manner until the appear-
domorpha—are capable of withstanding ance of works by the late Rene´ Cobben
prolonged submersion, several of those in beginning in 1957; we make further mention
intertidal environments. These include of Cobben’s relevant works below and
several species of Saldula Van Duzee, such provide citations to them.
as S. laticollis (Reuter) (Bahr and Schulte, The first descriptive work on the Saldidae
1976), all species of Salduncula Brown of South America dates from Blanchard
(Brown, 1959), the truly intertidal Aepophi- (1852). Efforts to describe the fauna were
lus bonnaerei Signoret from western Europe, decidedly piecemeal until Carl J. Drake took
2009 SCHUHANDPOLHEMUS: REVISIONOFPSEUDOSALDULACOBBEN 5
up work on the Saldidae in the late 1940s MATERIALSAND METHODS
(Drake and Carvalho, 1948; Drake, 1949).
ManyoftheavailablenamesfortheSaldidae During the course of this research project,
in South America, and the majority of those matrix code labels were affixed to the more
now placed in Pseudosaldula Cobben, were than 3500 specimens examined as a way to
proposed by Drake and his co-authors. uniquely identify them; these codes are
Drake took at least one field trip to therefore referred to as ‘‘unique specimen
Argentina inthe1920s—on whichhecollect- identifiers’’ (USIs). The USI codes, e.g.,
ed some specimens of Saldidae—and this AMNH_ENT 00021550, are composed of
may have been part of his inspiration to an institution and project code (AMNH_
study the fauna of the region. ENT) and a uniquenumber(00021550).USI
The existing taxonomic literature on the codesareincludedinthelocalitydataandare
SaldidaeofSouthAmericaremainsoflimited used to identify specimens from which
extent and use for taxonomic identification illustrations were made.
andunderstandingofdiversity,distributions, We have listed verbatim label data for the
and relationships. This situation results from holotypes associated with all previously
the fact that most published descriptions available names. Most of these historical
were based on one or a few specimens and specimens are also listed under Specimens
took little or no account of infraspecific Examined as a way of showing the political
variation. The limitations of the literature subdivisions and latitude-longitude coordi-
become particularly evident when one exam- nates we haveassociatedwith them.Ina few
ines the synonymies associated with the cases we were unable to place the type
diagnoses and descriptions provided in the locality with precision and it is therefore
present paper. not georeferenced and not included under
The genus Pseudosaldula, the focus of the Specimens Examined.
present paper, comprises at least 25% of the Alllatitude-longitudedatapresentedinthe
species of Saldidae from South America. SpecimensExaminedsectionofthepaperare
Although much of its distribution lies within in degrees and decimal parts thereof. Most
the tropical latitudes, its restriction to rela- coordinatesweredeterminedthroughtheuse
tively high elevations and high latitudes at of gazetteers, although some were acquired
lower elevations, gives it an effectively directlyinthefieldthroughtheuseofaGPS
temperate-latitude distribution, like so many device. Altitude data are treated as metric.
other members of the family. Please refer to the www.discoverlife.org web-
The current study was undertaken in an site as a way of accessing additional infor-
effort to rectify the shortcomings in the mation on specimens examined and for
existing literature on Pseudosaldula, while at access to interactive mapping tools.
the same time bringing to light information Color digital images of the bugs were
on a large amount of material collected prepared using a Microptics-USA photomi-
mostly by the authors over the last four crographic apparatus using a Nikon SLR
decades. The first fieldwork incorporated digitalcamera,Infinityoptics,andaMicrop-
into this project was conducted by R.T. tics-USA ML 1000 light source, all supplied
Schuh and Janet Crane in Peru during byRoyLarimer.Habitusphotosarepropor-
1971–72, at which time a limited number of tional to the size of the actual specimens so
Saldidaewerecollected.Additionalcollecting thatrelativedimensionscanbededucedfrom
was undertaken by R.T. Schuh in Colombia, comparison of the specimen images. Actual
Ecuador, and Peru during 1976 and in sizes of specimens can be determined by
Bolivia, Colombia, and Peru by J.T. and referringtotable 1. Legs were removed from
D.A. Polhemus during 1989. The southern all imaged specimens through digital editing,
Andes of Chile and Argentinawere surveyed because they were only partially visible,
on field trips by R.T. Schuh during 1981–82, provided no additional information, and
and 1986, by R.T. Schuh and J.T. Polhemus detracted from the informative aspects of
during1993,andbyS.OygurandE.Barrera coloration, vestiture, and texture of the
in 1994. dorsum. Zeiss EVO and Hitachi 4700 scan-
6 BULLETIN AMERICAN MUSEUM OFNATURALHISTORY NO.323
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Body 3.570.220.633.343.97 4.210.481.533.625.15 2.950.150.302.823.12 3.130.170.253.013.26 3.370.110.233.233.46 3.720.260.373.533.91
MeanSDRangeMinMax MeanSDRangeMinMax MeanSDRangeMinMax MeanSDRangeMinMax MeanSDRangeMinMax MeanSDRangeMinMax
a
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Pseudosald P.andensis=M(N9) =F(N8) P.antioqui=M(N3) =F(N2) P.aurea=M(N4) =F(N2)
2009 SCHUHANDPOLHEMUS: REVISIONOFPSEUDOSALDULACOBBEN 7
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MeanSDRangeMinMax MeanSDRangeMinMax MeanSDRangeMinMax MeanSDRangeMinMax MeanSDRangeMinMax MeanSDRangeMinMax
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Pseudosaldul P.bergi=M(N11) =F(N9) P.bruesi=M(N5) =F(N8) P.chilensis=M(N14) =F(N14)
8 BULLETIN AMERICAN MUSEUM OFNATURALHISTORY NO.323
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r
P
Head 0.470.030.070.430.50 0.470.050.120.420.54 0.440.080.180.350.53 0.490.040.100.450.55 0.500.040.080.470.56 0.530.010.010.520.54
Body 2.710.080.212.612.82 2.820.090.202.732.93 3.180.340.792.803.59 3.270.170.493.033.51 2.900.060.142.832.96 3.640.390.703.193.89
MeanSDRangeMinMax MeanSDRangeMinMax MeanSDRangeMinMax MeanSDRangeMinMax MeanSDRangeMinMax MeanSDRangeMinMax
o
a c
Pseudosaldul P.huamachu=M(N6) =F(N6) P.penai=M(N5) =F(N5) P.perula=M(N4) =F(N3)
2009 SCHUHANDPOLHEMUS: REVISIONOFPSEUDOSALDULACOBBEN 9
4
g
e 22404 33199 11202 30033 72549 67404
S 50055 50145 40044 40044 50055 50246
nt 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0.
A
3
g
e 33707 33908 81370 01291 73831 74202
S 40044 40044 30034 40034 50056 50156
nt 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0.
A
2
g
e 64032 05538 93044 05673 55561 85583
S 50156 60156 50156 60056 80179 80179
nt 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0.
A
1
g
e 82460 84033 91380 42335 13267 44291
S 20023 20123 20023 20022 30123 30124
nt 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0.
A
r
a
ulce
Ocan 3702053540 4309333770 3403073239 3202033033 4203103949 4602064349
nter-Dist 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0.
I
m
u
utell 0.720.050.130.670.80 0.760.250.930.060.99 0.810.060.130.750.88 0.790.060.080.750.83 0.860.090.310.741.05 0.940.110.320.761.08
c
S
h m
Widt otu 1906141326 3409281947 2406161834 1611150924 3212391655 4713362966
on 1.0.0.1.1. 1.0.0.1.1. 1.0.0.1.1. 1.0.0.1.1. 1.0.0.1.1. 1.0.0.1.1.
r
P
)
LE1nued Head 0.970.020.070.931.00 1.080.190.680.951.63 0.890.030.080.850.93 0.850.010.020.840.85 1.070.040.171.001.17 1.100.050.111.041.15
Bnti
Ao m
TC u
( utell 0.450.020.060.410.48 0.520.050.160.470.63 0.400.070.170.280.45 0.410.010.010.410.42 0.490.070.290.320.61 0.560.040.130.490.62
c
S
m
u
ut 76976 96222 35278 02292 65987 87167
osc 0.20.00.10.10.3 0.20.00.20.20.4 0.30.00.10.20.3 0.30.00.00.20.3 0.30.00.10.20.4 0.30.00.20.20.4
s
e
h M
gt
n m
e
L u
ot 4503084250 4904154559 4103063844 3804063541 5003084755 5307184361
on 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0.
r
P
Head 0.540.040.120.450.58 0.610.050.150.560.70 0.540.060.150.450.60 0.560.060.080.510.60 0.580.060.210.490.70 0.680.070.220.560.78
Body 3.230.170.463.043.50 3.530.250.723.163.88 3.520.190.523.243.76 3.390.230.333.223.55 3.810.280.923.474.39 4.230.371.023.664.68
MeanSDRangeMinMax MeanSDRangeMinMax MeanSDRangeMinMax MeanSDRangeMinMax MeanSDRangeMinMax MeanSDRangeMinMax
a
Pseudosaldul P.pilosa=M(N7) =F(N11) P.salina=M(N5) =F(N2) P.saxicola=M(N14) =F(N8)
10 BULLETIN AMERICAN MUSEUM OFNATURALHISTORY NO.323
4
g
e 64303 74314 2 22 43617
S 40145 40145 6 66 60066
nt 0.0.0.0.0. 0.0.0.0.0. 0. 0.0. 0.0.0.0.0.
A
3
g
e 93044 04336 7 77 53618
S 30134 40134 6 66 60066
nt 0.0.0.0.0. 0.0.0.0.0. 0. 0.0. 0.0.0.0.0.
A
2
g
e 25651 45673 84088 03774
S 50146 50146 81279 00090
nt 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0. 1.0.0.0.1.
A
1
g
e 52819 62101 22213 02381
S 20022 20123 30033 40034
nt 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0.
A
r
a
ulce
Ocan 3803123446 4002063743 4507104050 5001024951
nter-Dist 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0.
I
m
u
utell 0.780.130.350.610.96 0.810.080.320.660.98 0.940.140.190.841.03 1.100.090.190.981.18
c
S
h m
Widt otu 2615450551 3209341347 4316233254 5911254368
on 1.0.0.1.1. 1.0.0.1.1. 1.0.0.1.1. 1.0.0.1.1.
r
P
)
LE1nued Head 0.930.050.190.841.03 0.950.040.140.861.00 1.180.070.111.131.23 1.220.030.081.181.26
Bnti
Ao m
TC u
( utell 0.450.070.230.350.58 0.470.070.270.340.60 0.510.080.120.450.57 0.650.070.160.560.73
c
S
m
u
ut 66056 96987 60493 77471
osc 0.20.00.20.10.3 0.20.00.20.10.4 0.30.10.10.20.4 0.40.00.10.30.5
s
e
h M
gt
n m
e
L u
ot 4604163854 4803114353 5308114859 6004085664
on 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0. 0.0.0.0.0.
r
P
Head 0.510.080.270.350.62 0.550.060.220.450.67 0.680.080.110.620.73 0.650.100.240.530.78
Body 3.250.361.012.803.81 3.430.260.962.973.93 4.310.430.604.014.61 4.690.400.884.145.02
MeanSDRangeMinMax MeanSDRangeMinMax MeanSDRangeMinMax MeanSDRangeMinMax
a
Pseudosaldul P.vulgaris=M(N15) =F(N13) P.yungas=M(N2) =F(N2)