Molecular phylogeny of the pond skaters (Gerrinae), discussion of the fossil record and a checklist...

© Koninklijke Brill NV, Leiden, 2013 DOI 10.1163/1876312X-44042105

Insect Systematics & Evolution 45 (2014) 251–281 brill.com/ise

Molecular phylogeny of the pond skaters (Gerrinae), discussion of the fossil record and a checklist of species assigned to the

subfamily (Hemiptera: Heteroptera: Gerridae)

Jakob Damgaarda,*, Felipe Ferraz Figueiredo Moreirab, Tom A. Weirc and Herbert Zetteld

aLaboratory of Molecular Systematics, Botanical Garden and Museum Sølvgade 83 Opg. S, 1307 Copenhagen K, Denmark

bLaboratório de Entomologia, Departamento de Zoologia, Instituto de Biologia Universidade Federal do Rio de Janeiro, Avenida Carlos Chagas Filho, 373, CCS

Cidade Universitária, CEP 21941–971, C.P. 68044, Rio de Janeiro, RJ, Brasil cAustralian National Insect Collection, CSIRO Entomology, GPO Box 1700

Canberra, ACT 2601, Australia dNatural History Museum Vienna, Entomological Department,

Burgring 7, 1010 Vienna, Austria *Corresponding author, e-mail: [email protected]

Published online 12 May 2014; published in print 10 August 2014

AbstractThe phylogenetic relationships among selected species and genera of Gerrinae (Heteroptera: Gerridae) were investigated in a parsimony analysis of 2268 bp of DNA sequence data from the genes encoding COI + II, 16S rRNA and 28S rRNA. The taxa represented 12 of 15 recognized genera of Gerrinae and with outgroup taxa from all other subfamilies of Gerridae, including three of five recognized genera of Eotrechinae, which is considered the sister-group of Gerrinae. The resulting phylogeny shows that Gerrinae is not monophyletic, since a clade comprising Gerris, Aquarius, Limnoporus, Tachygerris, Eurygerris and Gigantometra is more closely related to representatives of the subfamily Eotrechinae than to a clade comprising Limnogonus, Neogerris, Limnometra, Tenagogerris and Tenagogonus. The two currently recognized gerrine tribes, Tachygerrini and Gerrini, were also paraphyletic, since Eurygerris was sister-group to Gigantometra, while Tachygerris was sister-group to Limnoporus + Aquarius + Gerris. Limnogonus and Neogerris were found to be strongly supported sister-taxa, and their sister-group was a clade comprising Limnometra, Tenagogerris and Tenagogonus, none of which were monophyletic. Finally, Aquarius chilensis (Berg, 1881) was sister-group to a clade comprising Aquarius amplus (Drake & Harris, 1938), A. remigis (Say, 1832) and A. remigoides (Gallant & Fairbairn, 1993), recognized as the A. remigis species group, and the entire clade was sister-group to Gerris. Based on the phylogenetic reconstruction, we outline possible diagnostic character combinations for a future revision of the Gerrinae and discuss the fossil record. While some of the relationships reinstate earlier ideas, e.g., Limnogonus and Neogerris being sister groups, many others are poorly supported and poorly diagnosed, and therefore, we retain from drawing taxonomic conclusions until data is available from the remaining genera, which can support a future generic revision of the Gerrinae. The updated checklist of species assigned to the subfamily is there-fore based on the established taxonomy.

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252 J. Damgaard et al. / Insect Systematics & Evolution 45 (2014) 251–281

KeywordsWater striders; systematics; simultaneous analyses; parsimony

Introduction

Pond skaters of the subfamily Gerrinae are among the best known and most frequently encountered semi-aquatic bugs (Gerromorpha) on all types of stagnant and slow flow-ing freshwater habitats throughout the world (Andersen 1982). In northern temperate regions, most species of water striders (Gerridae) can be assigned to just three genera of Gerrinae: Gerris Fabricius, 1794, Aquarius Schellenberg, 1800 and Limnoporus Stål, 1868. Of these, only Limnoporus is truly Holarctic, while Gerris extends into the Oriental and Afrotropical Regions, and Aquarius, as currently defined, is represented in all zooge-ographical regions (Andersen 1982, 1990, 1993, 1995a; but see Damgaard & Cognato 2005). Limnogonus Stål, 1868 and Neogerris Miyamoto, 1913 are found in tropical and subtropical areas of the world; Eurygerris Drake, 1958 and Tachygerris Hungerford & Matsuda, 1957 are restricted to the Neotropical Region; Tenagogerris Hungerford & Matsuda, 1958 is confined to Australia; Gerrisella Poisson, 1940, Tenagometra Poisson, 1949 and Tenagometrella Poisson, 1958 to the Afrotropical Region; Gigantometra Hungerford & Matsuda, 1958 to eastern Asia; and members of Tenagogonus Stål, 1853 and Limnometra Mayr, 1865 to the Old World tropics (Andersen 1995a).

As the name pond skaters implies, most species of Gerrinae are found in small, lentic freshwater habitats, such as ponds, pools, and sometimes lakesides and backwaters of streams; a few species enter brackish habitats, and others are confined to lotic habitats, where they may occur in large schools (Andersen 1982). Pond skaters are easy to collect and study, and have a broad outreach in studies of ecology, behavior and biogeography (Spence & Andersen 1994; Arnqvist 1997; Arnqvist & Rowe 2002; Han & Jablonski 2009, 2010; Rowe & Arnqvist 2012).

Matsuda (1960) was the first to revise the Gerridae, when he established five presumably monophyletic subfamilies: Gerrinae, Halobatinae, Ptilomerinae, Rhagadotarsinae and Trepobatinae, and within Gerrinae he included four tribes: Charmatometrini, Cylindrostethini, Eotrechini and Gerrini. In a later revision, Andersen (1975) erected the four tribes to subfamiliar rank, and subdivided his Gerrinae in Tachygerrini, containing Eurygerris and Tachygerris, and Gerrini contain-ing the remaining genera. Finally, Andersen (1995a) performed a phylogenetic analysis of 67 morphological characters scored for all gerrine genera and found two major clades in Gerrini, one comprising Gigantometra, Gerris, Aquarius and Limnoporus, and another comprising Limnogonus, Tenagometrella, Tenagogonus, Limnometra and Tenagometra, and that Tenagogerris and Neogerris + Gerrisella were successive sister-groups to the entire clade.

A series of papers have addressed the phylogenetic relationships among species of Gerris, Aquarius and Limnoporus (Andersen 1993, 1995a; Andersen & Spence 1992; Sperling et al. 1997; Damgaard et al. 2000; Damgaard & Sperling 2001; Damgaard & Cognato 2003, 2005), and were summarized by Damgaard (2006) in an analysis of 66 morphological characters and almost 3 kb of DNA sequence data from COI + II,

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J. Damgaard et al. / Insect Systematics & Evolution 45 (2014) 251–281 253

16S rRNA and 28S rRNA (D3–D5) and using Gigantometra gigas (China, 1925) as outgroup. The study confirmed most of the species-groups established by Andersen (1990, 1993) and Andersen & Spence (1992), but revealed that their relationships dif-fered considerably from what was previously recognized, mainly in Aquarius not being a monophyletic taxon, since A. remigis (Say 1832) and its close relatives A. amplus (Drake & Harris, 1938) and A. remigoides (Gallant & Fairbairn, 1993), together rec-ognized as the A. remigis-group, were more closely related to Gerris than to congeners, and that the Neotropical A. chilensis (Berg 1888) was placed unresolved at the root of the tree. No other gerrine genera have been studied in similar detail, but Damgaard et al. (2010) studied the phylogeny of Limnogonus using DNA sequence data from the same compartment of genes as Damgaard (2006), and suggested the subgeneric clas-sification in Limnogonus (s.str.) and Limnogonus (Limnogonoides) be replaced with a number of monophyletic species-groups.

The relationships within and among families and subfamilies of Gerromorpha have been studied repeatedly using molecular (Muraji & Tachikawa 2000), morphological (Andersen & Weir 2004) and combined analyses (Damgaard et al. 2005), and were sum-marized by Damgaard (2008a) in an analysis of the same compartment of DNA sequence data as Damgaard (2006), and with addition of 66 morphological characters obtained from the literature. The study revealed poor support and lack of diagnostic character com-binations for the Gerrinae represented by Gerris lacustris (Linnaeus, 1758) and Eurygerris fuscinervis (Berg, 1898) thus questioning the monophyly of the entire subfamily. The close relationship between Gerrinae and Eotrechinae already suggested by Andersen (1982) has never been seriously questioned, and the two subfamilies were also found to be well diag-nosed, although poorly supported, sister-groups by Damgaard (2008a).

The present study investigates the molecular phylogenetic relationships of Gerrinae by combining the studies by Damgaard (2006, 2008a) and Damgaard et al. (2010) with a number of genera and species from the Gerrinae and Eotrechinae not previously included in phylogenetic analyses. Figure 1 shows the phylogenetic relationship within and between these two subfamilies from Damgaard (2012) based on a compilation of two studies, each including a complete sampling of genera (J.T. Polhemus & Andersen 1984; Andersen 1995a).

In addition to the phylogenetic study we present an updated checklist of Gerrinae as compared to Andersen (1995a,b) and with additions from Andersen’s unpublished cata-logue, from recently published literature, and with new records based on author’s studies.The two species of Tenagogonus described by Gupta & Chaturvedi (2010) and Gupta et al. (2011) are not valid, because depositions of type-specimens were not indicated. Therefore, they are not included in the checklist.

Materials and methods

Taxonomic sampling

From Damgaard (2006) we obtained a complete taxon sample of Limnoporus and Aquarius and a dense sample of Gerris with inclusion of representatives from all

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Fig. 1. Phylogeny of the Gerrinae and Eotrechinae (after Damgaard 2012 and based on J.T. Polhemus & Andersen 1984 and Andersen 1995a). The relationships of Gerrinae are based on one of two trees (each 185 steps long; CI = 0.44; RI = 0.59) obtained from a parsimony analysis of 67 morphological characters and selected by both implied weighting and successive weighting, while the relationships of the Eotrechinae are based on J. T. Polhelmus & Andersen (1984), Andersen (1995a) and Tran & J. T. Polhelmus (2009).

Outgroup

Chimarrhometra

Eotrechus

Amemboa

Amemboides

Onychotrechus

Tarsotrechus

Tachygerris

Eurygerris

Neogerris

Gerrisella

Tenagogerris

Tenagogonus

Limnometra

Tenagometra

Tenagometrella

Limnogonus (s.str.)

Limnogonus(Limnogonoides)Gigantometra

Aquarius

Limnoporus

Gerriselloides

Macrogerris

Gerris s.str.

EOTREC

HIN

AE

GERRIN

AE

GERRINI

TACHYGERRINI

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subgenera and species-groups according to Andersen (1993), and with Gigantometra gigas as outgroup. From Damgaard et al. (2010) we obtained 17 species of Limnogonus, as well as Tenagogerris euphrosyne (Kirkaldy, 1902) and Onychotrechus esakii Andersen, 1980 (Eotrechinae). Finally, from Damgaard (2008a) we obtained Eurygerris fuscinervis, Metrocoris strictus Chen & Nieser, 1993 (Halobatinae), Ptilomera kirkaldyi Hungerford & Matsuda, 1965 (Ptilomerinae), Brachymetra unca Shaw, 1933 (Charmat ometrinae), Trepobates subnitidus Esaki, 1926 (Trepobatinae), Rhagadotarsus kraepelini Breddin, 1905 (Rhagadotarsinae), Cylindrostethus costalis Schmidt, 1915 (Cylin drostethinae) and Velia caprai Tamanini, 1947 from the family Veliidae. The lat-ter species is used as outgroup in the present study. To this assemblage we add three species of Tachygerris, one species of Eurygerris, four species of Neogerris, two species of Tenagogerris, 11 species of Limnometra and six species of Tenagogonus, most of which have not been used in molecular phylogenetic studies before. Table 1 lists the material used in this study with collecting data, references, and GenBank accession numbers.

Molecular data

Total genomic DNA was extracted from ethanol preserved specimens following the QIAGEN DNEasy Purification Kit Protocol (Qiagen, Santa Clara, CA, USA), except for the final elution, which was done twice in 200 µl EB buffer using a single centrifu-gation step. All PCR amplifications took place in a cocktail of 2 µl DNA template, 20 µl GATC mix (0.02 mM of each nucleotide), 14 µl ddH2O, 5 µl Amplicon III 10 × key buffer (15 mM MgCl2) (VWR International, West Chester, PA, USA), 5 µl of each of the sense and anti-sense primers and 0.2 µl Amplicon III Taq polymerase (VWR International). In case of no amplification, 5 µl of ddH2O was replaced by 5 µl Amplicon III 10 × standard buffer (15 mM MgCl2) (VWR International). The PCR amplifications were performed under the following conditions: 10 min 94°C (hot start), followed by 25–30 cycles of 45 s at 94°C (denaturing), 45 s at 45–48°C (anneal-ing) and 60 s at 72°C (extension). PCR-amplicon electrophoresis was done on a 2% low melting agarose gel and stained with SYBR® (Invitrogen, Eugene, OR, USA) under UV-light. Positive amplicons were purified using QIAQUICK PCR Purification Kit (Qiagen) and sequenced using the ABI Sequencing Reaction Kit (Applied Biosystems, Foster City, CA, USA). Sequencing products were purified using QUIAGEN DyeEx 2.0 Spin Kit (Qiagen) and run on an ABI 3130xl Analyzer (Applied Biosystems). All vouchers and DNA extractions are stored at the Natural History Museum of Denmark. Table 2 lists the primers used in this study. A number of internal primers were designed in order to amplify and sequence overlapping fragments of COI + II for the many deviant taxa, and the sequences of these can either be deduced from the sequences or obtained from the senior author.

Phylogenetic analyses

Sequence editing and contig construction for each segment was performed in Sequencher 4.10 (Gene Codes, Ann Arbor, MI, USA). PAUP* b4.08 (Swofford 1998)

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sas,

John

son

Cre

ek20

02J.

Dam

gaar

d47

9AY

9513

14AY

9513

29D

Q69

3330

Ger

ris (G

erris

) bab

ai M

iyam

oto,

195

8C

hina

, Tia

njin

(lab

. cul

t.)19

93J.R

. Spe

nce

534

AF25

1108

AY42

5225

DQ

6833

25G

erris

(Ger

ris) b

rasil

i Poi

sson

, 194

1Sp

ain,

Mur

cia,

Sie

rra

Segu

ra,

R

io M

oro

1999

G. A

rnqv

ist10

8AY

4252

38AY

4252

29D

Q68

3343

Ger

ris (G

erris

) bue

noi K

irkal

dy, 1

911

Can

ada,

Ont

ario

, Otta

wa

????

J.R. S

penc

e42

7AY

9513

12AY

4251

91D

Q68

3327

Ger

ris (G

erris

) cau

casic

us K

anyu

kova

, 198

2Ir

an, G

uila

n Pr

ov.

????

R. H

osse

ini

545

AY95

1313

AY95

1328

DQ

6833

28G

erris

(Ger

ris) c

osta

e cos

tae (

Her

rich-

Schä

ffer,

18

50)

Austr

ia, S

chei

bbs,

Dür

rens

tein

????

F. S

eyfe

rt35

8AY

4252

36AY

4252

26D

Q68

3334

Ger

ris (G

erris

) com

atus

Dra

ke &

Hot

tes,

1925

Can

ada,

Ont

ario

, Otta

wa

????

J.R. S

penc

e42

8AY

9513

15AY

4252

22D

Q68

3337

Ger

ris (G

erris

) gib

bifer

Sch

umm

el, 1

832

Den

mar

k, B

ornh

olm

,

Bølsh

avn

1997

J. D

amga

ard

807

AF25

1117

AY42

5195

DQ

6833

44

Ger

ris (G

erris

) gill

ette

i Let

hier

ry &

Sev

erin

, 189

6U

SA, N

ew M

exic

o,

Je

mez

Mts.

, Fen

ton

L.19

96G

. Arn

qvist

29AF

2511

05AY

4252

18D

Q68

3323

Ger

ris (G

erris

) inc

ogni

tus D

rake

& H

otte

s, 19

25U

SA, N

ew M

exic

o, B

osqu

e

de

Apa

che

1996

G. A

rnqv

ist25

0AY

4252

47AY

4252

07D

Q68

3322

Ger

ris (G

erris

) inc

urva

tus D

rake

& H

otte

s, 19

25C

anad

a, B

ritish

Col

umni

a:

Ab

botsb

ed??

??L.

Row

e10

7AY

4252

40AY

4252

20D

Q68

3339

Ger

ris (G

erris

) ins

pera

tus D

rake

& H

otte

s, 19

25U

SA, A

laba

ma:

Cah

aba

R. H

ist. P

ark

2000

J. D

amga

ard

491

AY95

1316

AY95

1331

DQ

6833

40

Ger

ris (G

erris

) lac

ustri

s (Li

nnae

us, 1

758)

Den

mar

k, G

l. H

olte

(lab.

cul

t.)19

93J.R

. Spe

nce

577

AY42

5239

AY42

5227

DQ

6833

41

Ger

ris (G

erris

) lat

iabd

omin

is M

iyam

oto,

195

8Ja

pan,

Sak

urai

1992

T. H

arad

a38

AF25

1110

AY42

5215

DQ

6833

31G

erris

(Ger

ris) m

acul

atus

Tam

anin

i, 19

46Tu

nesia

, Tam

ara

1986

ZM

UC

-exp

.21

AF25

1116

AY42

5224

DQ

6833

42G

erris

(Ger

ris) m

argi

natu

s Say

, 183

2U

SA, N

. Car

olin

a,

W

eave

rvill

e, L

. Lou

ise19

97G

. Arn

qvist

105

AY42

5241

AY42

5211

DQ

6833

38

Ger

ris (G

erris

) nep

alen

sis D

istan

t, 19

10Ja

pan,

Hon

shu,

Koc

hi19

91T.

Har

ada

22AF

2511

06AY

4251

92AB

0347

91

Tabl

e 1.

(C

ont.)

<UN> <UN>


Ger

ris (G

erris

) odo

ntog

aste

r (Ze

tters

tedt

, 182

8)D

enm

ark,

Gl.

Hol

te

(la

b. c

ult.)

1993

J.R. S

penc

e53

6AF

2511

09AY

4252

19D

Q68

3329

Ger

ris (G

erris

) pin

gree

nsis

Dra

ke &

Hot

tes,

1925

Can

ada,

B.C

., Al

bert

a D

unsta

ble

(lab

cult.

)19

93J.R

. Spe

nce

537

AY95

1311

AY42

5214

Ger

ris (G

erris

) sph

agne

toru

m G

auni

tz, 1

947

Swed

en, S

mål

and,

Bjä

djes

jö19

96J.

Dam

gaar

d1j

d1AF

2511

03AY

4252

06D

Q68

3321

Ger

ris (G

erris

) sw

akop

ensis

Stå

l, 18

58K

enya

, Mas

ai M

ara

1994

J.R. S

penc

e79

5AF

2511

12AY

4252

23D

Q68

3332

Ger

ris (G

erris

) tho

racic

us S

chum

mel

, 183

2D

enm

ark,

Tås

trup

1996

J. D

amga

ard

26AF

2511

14AY

4251

87G

igan

tom

etra

gig

as (C

hina

, 192

5)V

ietn

am, C

he L

inh

NE

Han

oi??

??L.

Row

e34

AY42

5235

AY42

5183

DQ

6832

92

Lim

nogo

nus c

apen

sis (C

hina

, 192

5)Ta

nzan

ia, M

t. Ro

ngw

e19

98P.

Gra

vlun

d13

8G

U64

5151

GU

6450

45G

U64

5100

Lim

nogo

nus h

ypol

eucu

s(Ger

staec

ker,

1873

)Ta

nzan

ia, M

inzir

o Fo

r. Re

s.19

98P.

Gra

vlun

d13

9G

U64

5157

GU

6450

51–

Lim

nogo

nus i

nter

med

ius P

oiss

on, 1

941

Uga

nda,

Kib

ale

For.

Res.,

Nya

bike

re19

95M

. Num

mel

in14

0G

U64

5159

GU

6450

53G

U64

5107

Lim

nogo

nus n

igre

scens

Poi

sson

, 194

1Bo

tswan

a, O

kava

ngo

Del

ta19

98L.

Gra

u N

ersti

ng98

GU

6451

60G

U64

5054

GU

6451

08Li

mno

gonu

s pec

tora

lis (M

ayr,

1865

)Th

aila

nd, P

atta

ni B

ay19

97C

. Sw

enne

n21

1G

U64

5179

GU

6450

73G

U64

5126

Lim

nogo

nus p

oisso

ni A

nder

sen,

197

5U

gand

a, K

ibal

e Fo

r. Re

s.,

N

yabi

kere

1995

M. N

umm

elin

32b

GU

6451

55G

U64

5049

GU

6451

04

Lim

nogo

nus c

ereiv

entri

s (Si

gnor

et, 1

862)

Rodr

igue

s, Sa

int G

abrie

l20

09M

. Mad

l23

90G

U64

5170

GU

6450

64–

Lim

nogo

nus f

ossa

rum

fossa

rum

(F. 1

775)

Th

aila

nd, B

angk

ok20

08J.

Dam

gaar

d22

90G

U64

5183

GU

6450

77G

U64

5130

Lim

nogo

nus f

ossa

rum

gilg

uy A

nder

sen

& w

eir,

19

97Au

stral

ia, N

.T, N

hulu

nbuy

2007

T. W

eir

2397

GU

6451

88G

U64

5082

GU

6451

35

Lim

nogo

nus w

indi

Hun

gerfo

rd &

Mat

suda

, 196

1Au

stral

ia, W

.A.,

Prin

ce

Re

gent

Riv

er19

98C

ALM

Sur

vey

2394

GU

6452

03G

U64

5097

GU

6451

48

Lim

nogo

nus f

ranc

iscan

us (S

tål,

1859

)Ar

uba,

Boc

a Pr

ins

2009

N.F

. M

unch

-Pet

erse

n24

06G

U64

5175

GU

6450

69G

U64

5122

Lim

nogo

nus r

ecen

s Dra

ke &

Har

ris, 1

934

Hon

dura

s, C

orte

s, Yo

joa

2002

A.I.

Cog

nato

1437

GU

6451

69G

U64

5063

GU

6451

17Li

mno

gonu

s adu

ncus

Dra

ke &

Har

ris, 1

933

Arge

ntin

a, Ju

juy,

Agua

s Cal

ient

es20

04S.

Maz

zucc

oni

2344

GU

6451

64G

U64

5058

Lim

nogo

nus i

gnot

us D

rake

& H

arris

, 193

4Ar

gent

ina,

Cha

co,

R

.V.S

. El C

acha

pé20

08S.

Maz

zucc

oni

2365

GU

6451

67G

U64

5061

Lim

nogo

nus p

rofu

gus D

rake

& H

arris

, 193

0Ar

gent

ina,

Misi

ones

,

P. P.

Salto

Enc

anta

do20

04S.

Maz

zucc

oni

2342

GU

6451

68G

U64

5062

( Con

tinue

d )

<UN> <UN>


Taxo

nC

olle

ctin

g sit

eYe

arC

olle

ctor

Ext.

CO

I + C

OII

16S

rRN

A28

S rR

NA

Lim

nogo

nus h

unge

rford

i And

erse

n, 1

975

Mal

aysia

, Sar

awak

, Lip

ad R

. 20

00??

??10

14G

U64

5192

GU

6450

86G

U64

5139

Lim

nogo

nus l

uctu

osus

(Mon

trous

ier,

1865

) IN

ew C

aled

onia

, Man

dgel

ia20

08J.

Dam

gaar

d22

92G

U64

5196

GU

6450

90G

U64

5143

Lim

nogo

nus l

uctu

osus

(Mon

trous

ier,

1865

) II

Austr

alia

, N.T

., D

arw

in??

??T.

Wei

r &

V. Th

ink

149

GU

6452

00G

U64

5094

GU

6451

45

Lim

nogo

nus n

itidu

s (M

ayr,

1865

)La

os, V

ient

iane

Pro

vinc

e20

08J.

Pede

rsen

&

A. S

olod

ovni

kov

2190

GU

6451

74G

U64

5068

GU

6451

21

Lim

nom

etra

bor

neen

sis H

unge

rford

& M

atsu

da,

1958

Mal

aysia

, Sar

awak

, Lip

ad R

.20

00??

??10

15K

C88

0946

KC

8808

95K

C88

0926

Lim

nom

etra

cursi

tans

(F. 1

775)

Austr

alia

, QLD

, Woo

badd

a R

.20

05G

. Arn

qvist

1477

KC

8809

47K

C88

0896

Lim

nom

etra

fem

orat

a M

ayr,

1865

Phili

ppin

es, L

eyte

Is.,

Bayb

ay20

05H

. Zet

tel &

C

.V. P

anga

ntih

on10

54K

C88

0948

KC

8808

97

Lim

nom

etra

fluv

ioru

m (F

. 179

8)Sr

i Lan

ka, K

andy

, Blu

e H

eave

n19

95??

??22

7K

C88

0945

KC

8808

94K

C88

0925

Lim

nom

etra

lipo

vsky

i Hun

gerfo

rd &

Mat

suda

, 19

58Au

stral

ia, Q

LD, I

ron

Ran

ge

N

P19

98A.

Cal

der

1071

KC

8809

49K

C88

0898

Lim

nom

etra

mat

suda

i (M

iyam

oto,

196

7)C

hina

, Yun

nan,

Men

glun

????

L. C

heng

256

KC

8809

51K

C88

0900

Lim

nom

etra

nig

ripen

nis M

ayr,

1865

Phili

ppin

es, M

asba

te Is

l.,

Tu

gbo

2005

H. Z

ette

l10

63K

C88

0952

KC

8809

01K

C88

0932

Lim

nom

etra

pal

auan

a Es

aki,

1925

Chu

uk, W

eno

Is.

1999

J.T. P

olhe

mus

230

KC

8809

53K

C88

0902

KC

8809

33Li

mno

met

ra p

seudo

insu

laris

Nie

ser &

Che

n, 1

992

Brun

ei, S

unga

i Bel

ait A

rea

2007

H. Z

ette

l19

09K

C88

0954

KC

8809

03K

C88

0934

Lim

nom

etra

pul

chra

May

r, 18

65Pa

lau

1999

J.T. P

olhe

mus

231

KC

8809

55K

C88

0904

KC

8809

35Li

mno

met

ra sp

.PN

G, W

este

rn H

ighl

ands

,

Kur

umul

2006

M. B

alke

1562

KC

8809

50K

C88

0899

KC

8809

30

Lim

nopo

rus e

saki

i (M

iyam

oto,

195

8)Ja

pan,

Hon

shu

????

J.R. S

penc

e13

4AY

9513

05AY

4252

16D

Q68

3293

Lim

nopo

rus c

anal

icula

tus (

Say,

1832

)C

anad

a, O

ntar

io??

??J.R

. Spe

nce

136

AY42

5251

AY42

5208

DQ

6832

94Li

mno

poru

s diss

ortis

(Dra

ke &

Har

ris, 1

930)

Can

ada,

Ont

ario

, Otta

wa

????

J.R. S

penc

e42

5AY

9513

06AY

4252

28D

Q68

3295

Lim

nopo

rus n

otab

ilis (

Dra

ke &

Hot

tes,

1925

)C

anad

a, B

ritish

Col

umbi

a,

Va

ncou

ver

????

J.R. S

penc

e40

6AY

9513

07AY

4252

10D

Q68

3296

Lim

nopo

rus g

enita

lis (M

iyam

oto,

195

8)Ja

pan,

Hok

kaid

o??

??J.R

. Spe

nce

135

AY95

1308

AY42

5217

DQ

6832

97Li

mno

poru

s ruf

oscu

tella

tus (

Latre

ille,

180

7)Fi

nlan

d (la

b cu

lt.)

????

J.R. S

penc

e13

1AY

9513

09AY

4252

12D

Q68

3298

Tabl

e 1.

(C

ont.)

<UN> <UN>


Neo

gerr

is he

sione

(Kirk

aldy

, 190

2)U

SA, N

. Car

olin

a, C

huns

Cav

e, A

shvi

lle??

??G

. Arn

qvist

134

KC

8809

56K

C88

0881

KC

8809

12

Neo

gerr

is lu

bricu

s (W

hite

, 187

9)G

uyan

a, T

imiti

ro A

irpor

t??

??M

. Num

mel

in13

3K

C88

0961

KC

8808

82K

C88

0913

Neo

gerr

is pa

rvul

us (S

tål,

1859

)La

os, C

ham

pasa

k Pr

ovin

ce20

08J.

Pede

rsen

&

A. S

olod

ovni

kov

2123

KC

8809

57K

C88

0883

KC

8809

14

Neo

gerr

is vi

sendu

s (D

rake

& H

arris

, 193

4)Ve

nezu

ela,

“T

VM

0200

”??

????

?18

64K

C88

0939

KC

8808

80K

C88

0911

Tach

yger

ris a

dam

soni

(Dra

ke, 1

942)

Trin

idad

, Par

ia R

iver

1991

J.G. N

ielse

n24

4K

C88

0962

KC

8808

77K

C88

0908

Tach

yger

ris o

pacu

s (C

ham

pion

, 189

8)Pa

nam

a, C

erro

Cam

pana

Park

1993

J.T. P

olhe

mus

&

A. G

illog

ly21

7K

C88

0964

KC

8808

79K

C88

0910

Tach

yger

ris q

uadr

illin

iatu

s (C

ham

pion

, 189

8)Be

lize,

Cay

o D

istric

t,

Rio

Frio

????

P.W. K

ovar

ik28

5K

C88

0963

KC

8808

78K

C88

0909

Tena

goge

rris

euph

rosy

ne I

Kirk

aldy

, 190

2Au

stral

ia, N

SW,

M

urra

mar

ang

????

N.M

. And

erse

n20

4G

U64

5150

GU

6450

44G

U64

5099

Tena

goge

rris

euph

rosy

ne II

Kirk

aldy

, 190

2Au

stral

ia, Q

LD, B

roke

n R

.20

05G

. Arn

qvist

1481

KC

8809

42K

C88

0891

KC

8809

22Te

nago

gerr

is fem

orat

us A

nder

sen

& W

eir,

1997

Austr

alia

, N.T

., K

akad

u N

. P.

1992

R. H

ause

r13

6K

C88

0943

KC

8808

92K

C88

0923

Tena

goge

rris

palli

dus A

nder

sen

& W

eir,

1997

Austr

alia

, N.T

, Gre

gory

N. P

.20

01T.

Wei

r &

P. Bo

ucha

rd24

73K

C88

0944

KC

8808

93

Tena

gogo

nus a

ustra

liens

is An

ders

en &

W

eir,

1997

IAu

stral

ia, Q

LD, W

ooba

dda

Riv

er20

05G

. Arn

qvist

1479

KC

8809

59K

C88

0887

KC

8809

18

Tena

gogo

nus a

ustra

liens

is An

ders

en &

W

eir,

1997

IIAu

stral

ia, Q

LD, H

enrie

tta

C

reek

2005

G. A

rnqv

ist14

82K

C88

0941

KC

8808

88K

C88

0919

Tena

gogo

nus s

p.G

abon

, Ipa

ssa

2006

E. G

uilb

ert

1566

KC

8809

58K

C88

0884

KC

8809

15Te

nago

gonu

s mad

agas

carie

nsis

Hob

erla

ndt,

1947

Mad

agas

car

2004

M. M

adl

1240

KC

8809

40K

C88

0885

Tena

gogo

nus z

ambe

zinu

s (Po

isson

, 193

4)Ta

nzan

ia, E

ast U

sam

bara

,

Aman

i19

97P.

Gra

vlun

d24

2K

C88

0960

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was used for assembling and aligning the combined sequences manually and in com-bination with ClustalX (Thompson et al. 1997) using default parameters. Combined parsimony analyses were performed in PAUP* using 100 random taxon addition replicates, and with gaps scored as either a fifth character state or as missing data. Branch support measurements were evaluated by bootstrap analyses (Felsenstein 1985) using 500 simple taxon addition pseudoreplicates and by Bremer support (BS) (Bremer 1994) using constrained files generated in TreeRot v.2a (Sorensen 1999), and run in PAUP* with all other parameters set as default.

Results

Data sets

The 16S rRNA and 28S rRNA primers worked well for most samples, and only a few specimens failed to amplify and sequence these genes. The protein coding genes COI + II, on the other hand, were more problematic due to numerous base substitu-tions, especially in third codon positions, and a number of specimens failed to amplify and sequence the entire segment. The assembled and aligned data set consisted of 2268 characters of which 1–1817 (1818 characters) came from COI + II, 1818–2270 (453 characters) from 16S rRNA and 2271–2858 (588 characters) from 28S rRNA, and is available from Treebase (www.treebase.org). All sequences have been deposited in GenBank (see Table 1 for accession numbers).

A phylogenetic analysis of all data and with gaps scored as a fifth character state (altogether 828 parsimony informative characters) gave four equally parsimonious trees, each 9082 steps long (CI = 0.1895; RI = 0.5124). A strict consensus of these trees is shown in Fig. 2 and reveals that Gerrinae is not monophyletic since Eotrechinae (represented by members of Amemboa Esaki, 1925; Eotrechus, Kirkaldy 1902 and

Table 2. Primers used in the present study.

Primer Sequence (5′→3′) Reference

28S rRNA 28SL CCCGTCTTGAAACACGGACCAA Muraji & Tachikawa 2000 28SH CCACAGCGCCAGTTCTGCTTAC Muraji & Tachikawa 2000

16S rRNA 16sA CGCCTGTTTAACAAAAACAT Cognato & Vogler 2001 16sB2 TTTAATCCAACATCGAGG) Cognato & Vogler 2001COI + II C1–J-1751 GGATCACCTGATATAGCATTCCC Simon et al. 1994 C1–N-2191 CCCGGTAAAATTAAAATATAAACTTC Simon et al. 1994 C1–J-2183 CAACATTTATTTTGATTTTTTGG Simon et al. 1994 TL-2–3014 TCCAATGCACTAATCTGCCATATTA Simon et al. 1994 C1–J-2798 CCWCGWCGWTAYTCWGAYTATCC Damgaard & Cognato 2005 C1–N-3389 CCACAAATTTCTGAACATTGACCA Simon et al. 1994 C2–N-3554 GTTCATGARTGWARCACATC Damgaard & Cognato 2005

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http://www.treebase.org


Fig. 2. Strict consensus of four equally parsimonious trees resulting from a phylogenetic analysis of all data and with gaps scored as a fifth character state. Numbers above branches refer to bootstrap support values and numbers below branches refer to Bremer support values.

Gerris comatusGerris marginatusGerris incurvatusGerris insperatusGerris alacrisGerris latiabdominisGerris thoracicusGerris brasiliGerris gibbiferGerris lacustrisGerris maculatusGerris swakopensisGerris costae costaeGerris caucasicusGerris odontogasterGerris babaiGerris buenoiGerris argentatusGerris nepalensisGerris argenticollisGerris gilletteiGerris pingreensisGerris incognitusGerris sphagnetorumGerris gracilicornisGerris yezoensisGerris insularisGerris asperGerris lateralisAquarius amplusAquarius remigisAquarius remigoidesAquarius chilensisAquarius adelaidisAquarius liliAquarius philipinensisAquarius distantiAquarius antigoneAquarius fabriciiAquarius paludum amamiensisAquarius paludum paludumAquarius ventralisAquarius conformisAquarius nebularisAquarius elongatusAquarius cinereusAquarius najasLimnoporus dissortisLimnoporus notabilisLimnoporus genitalisLimnoporus rufoscutellatusLimnoporus canaliculatusLimnoporus esakiiTachygerris adamsoniTachygerris quadrilineatusTachygerris opacusEurygerris fuscinervisEurygerris �avolineatusGigantometra gigasEotrechus petraeusAmemboa sp.Onychotrechus esakiiLimnogonus cereiventrisLimnogonus franciscanusLimnogonus fossarum fossarumLimnogonus fossarum gilguyLimnogonus hungerfordiLimnogonus luctuosus Limnogonus luctuosus IILimnogonus aduncusLimnogonus recensLimnogonus profugusLimnogonus ignotusLimnogonus nitidusLimnogonus intermediusLimnogonus pectoralisLimnogonus poissoniLimnogonus hypoleucusLimnogonus nigrescensLimnogonus capensisLimnogonus windiNeogerris hesioneNeogerris lubricusNeogerris visendusNeogerris parvulusTenagogerris euphrosyne ITenagogerris euphrosyne IITenagogerris pallidusTenagogonus australiensis ITenagogonus australiensis IITenagogerris femoratusTenagogonus �jiensisLimnometra lipovskyiTenagogonusTenagogonus zambezinusTenagogonus madagascariensisLimnometra cursitansLimnometra sp.

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Onychotrechus Kirkaldy, 1903) is nested as sister-group to a clade comprising Gerris, Aquarius, Limnoporus, Tachygerris, Eurygerris and Gigantometra, and another clade, comprising Limnogonus, Neogerris, Limnometra, Tenagogerris and Tenagogonus, is placed as sister-group to the entire clade. Within Gerris s.str. we find G. latiabdominis Miyamoto, 1958 and G. thoracicus Schummel, 1832 as successive sister-taxa to a strongly supported clade comprising G. comatus Drake & Hottes, 1925 + G. marginatus Say, 1832 + G. incurvatus Drake & Hottes, 1925 + G. insperatus Drake & Hottes, 1925 and G. alacris Hussey, 1921. The sister-group to this clade comprises G. costae (Herrich-Schäffer, 1950) + G. swakopensis (Stål, 1858) and G. brasili Poisson, 1941 + G. gibbifer Schummel, 1832 + G. lacustris + G. maculatus Tamanini, 1946. The succes-sive sister-group to the entire clade includes a clade comprising G. caucasicus Kanyukova, 1982 + G. odontogaster (Zetterstedt, 1828) + G. babai Miyamoto, 1958 + G. buenoi Kirkaldy, 1911 + G. argentatus Schummel, 1832, G. nepalensis Distant, 1910, G. argenticollis Parshley, 1916 and a clade comprising G. gillettei Lethierry & Severin, 1896 + G. pingreensis Drake & Hottes, 1925 + G. incognitus Drake & Hottes, 1925 + G. sphagnetorum Drake & Hottes, 1925. The sister-group to Gerris s.str. is a clade comprising representatives of subgenera Gerriselloides (G. asper (Fieber, 1860) + G. lateralis Schummel, 1832) and Macrogerris (G. gracilicornis (Horváth, 1879) + G. insularis (Motschulsky, 1866) + G. yezoensis Miyamoto, 1958).

The sister-group to Gerris is a strongly supported clade comprising Aquarius amplus, A. remigis and A. remigoides, together the A. remigis group, and A. chilensis (Berg, 1881) as sister-group to this clade. The remaining species of Aquarius form a clade in which A. ventralis (Fieber, 1861) is sister-group to a clade comprising A. paludum paludum (Fabricius, 1794) + A. p. amamiensis (Miyamoto, 1958) + A. antigone (Kirkaldy, 1899) + A. fabricii Andersen, 1990 + A. distanti (Horváth, 1899) + A. philippinensis Zettel & Ruiz, 2003 + A. adelaidis (Dohrn, 1860) + A. lili D.A. Polhemus & J.T. Polhemus, 1994. Aquarius cinereus (Puton, 1869) + A. najas (De Geer, 1773) is sister-group to a clade comprising A. elongatus (Uhler, 1896) + A. conformis (Uhler, 1878) + A. nebularis (Drake & Hottes, 1925).

Limnoporus is sister-group to the larger Aquarius clade and includes L. canaliculatus (Say, 1832) and L. esakii (Miyamoto, 1958) as successive sister-taxa to a clade comprising L. rufoscutellatus (Latreille, 1807) + L. genitalis Nieser & Chen, 1992 + L. dissortis (Drake & Harris, 1930) + L. notabilis (Drake & Hottes, 1925). Tachygerris and Eurygerris + Gigantometra gigas are successive sister-groups to the Limnoporus + Aquarius + Gerris clade.

The remaining species of Gerrinae form a relatively well-supported clade. In this, Limnogonus is monophyletic and relatively well supported, and includes a few well-supported clades and sister-taxa, such as L. aduncus Drake & Harris, 1933 + L. recens Drake & Harris, 1934 + L. profugus Drake & Harris, 1930 + L. ignotus Drake & Harris, 1934 and L. cereiventris (Signoret, 1862) + L. franciscanus (Stål, 1859) and L. hungerfordi Andersen, 1975 + L. luctuosus (Montrouzier, 1865), even though the latter itself is not monophyletic. Neogerris is strongly supported, and so is the sister-group relationship with Limnogonus, while relationships among species of Neogerris have little support. The sister-group to Limnogonus + Neogerris is a clade of Limnometra,

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Tenagogerris and Tenagogonus, and none of the three genera are monophyletic, but comprise two clades.

In the first clade, the two specimens of Tenagogonus australiensis Andersen & Weir, 1997 are nested within Tenagogerris as a well-supported sister-group to T. euphrosyne + T. pallidus Andersen & Weir, 1997, while T. femoratus (Mayr, 1865) is strongly supported sister-group to the entire clade, and Tenagogonus fijiensis Hungerford & Matsuda, 1958, Limnometra lipovskyi Hungerford & Matsuda, 1958 and a clade of African Tenagogonus (T. madagascariensis Hoberlandt, 1947 + T. zambezinus (Poisson, 1934) + an unidentified nymph from Gabon) are successive sister-taxa to this clade. In the second clade, Limnometra fluviorum (Fabricius, 1798) is strongly supported sister-group to a well-supported clade comprising Tenagogonus bergrothi Hungerford & Matsuda, 1958 and Limnometra matsudai (Miyamoto, 1967), while the remaining species of Limnometra form a strongly supported sister-group to this clade. A poorly supported clade comprising representatives of Rhagadotarsinae + Trepobatinae and Cylindrostethinae is sister-group to the poorly supported Gerrinae + Eotrechinae clade, and a representative of Charmatometrinae is relatively well-supported sister-group to the entire clade.

Treating gaps as missing data (804 parsimony informative characters) also gave four equally parsimonious trees, each 8902 steps long (CI = 0.1837; RI = 0.5071). A strict consensus tree (not shown) was very similar to the tree with gaps treated as a new character state except for a shift in relationships among species of Limnometra, where L. nigripennis nigripennis Mayr, 1865 and L. pulchra Mayr, 1865 form a sister clade to L. cursitans (Fabricius, 1775) + an unidentified specimen, L. sp. #1562, + L. palauana Esaki, 1925 + L. pseudoinsularis Nieser & Chen, 1992.

Discussion

Paraphyly of Gerrinae, Gerrini and Tachygerrini

Andersen (1982) listed the sclerotized male dorsal vesical plate as a single synapomor-phy for the subfamily Gerrinae, but later (Andersen 1995a) found it to be a weak character (absent in several genera), and instead listed the reduced male parameres; the tightly coiled distal section of the fecundation gynatrial complex; the modified metasternal area surrounding the scent orifice; the absence of evaporative scent grooves; and the distance between pro- and mesoacetabula less than 3.5× the distance between meso- and metacetabula.

Damgaard (2008a) found the Gerrinae to be poorly supported, and without convincing diagnostic characters, while the sister-group relationship with Eotrechinae was supported by the sclerotized inner lobes of the first female gonapophyses and the glandular areas of female gynatrial sac being divided in two lateral glandular areas. The present study includes representatives of two more eotrechine genera, Eotrechus and Amemboa, and the subfamily includes three additional genera, i.e., Amemboides J. T. Polhemus & Andersen, 1984, formerly a subgenus of Amemboa, but later raised to genus rank by Tran & J. T. Polhemus (2009), and two monotypic genera represented

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by Chimarrhometra orientalis (Distant 1879) and Tarsotrechus polhemi Andersen, 1980. While J. T. Polhemus & Andersen (1984) found Amemboa and Onychotrechus to be sister groups, we find a sister group relationship between Amemboa and Eotrechus, which calls for future investigations of the relationships among genera of Eotrechinae.

According to Damgaard (2008a) the sole diagnostic character for the Eotrechinae was lack of sclerotization of the accessory scent gland, a reversal also found in Ptilomerinae, and the subfamily is not strongly supported by the present study. Andersen (1995a) diagnosed the Tachygerrini on the scent orifice being situated on a longitudinal ridge or carina and protected by a tuft of hairs; pear shaped scent reser-voir; and second bifurcation of M + Cu in fore wing far removed from the anterior cross-vein, and he diagnosed Gerrini on the absence of a distinct lateral intersegmental suture between meso- and metanotum, and loss of pretarsal arolia in the adult stage. Andersen (1990) found that Aquarius chilensis had retained the pretarsal arolia, but nevertheless considered other characters sufficiently convincing to retain it in Aquarius, but stated (p. 45) that “if it is hypothesized that the pretarsal arolia have only been lost once within the subfamily Gerrinae, chilensis might well be placed as the sister-group of all other species belonging to the tribe Gerrini”. Our study shows that A. chilensis is placed as sister-group to the A. remigis group, and that Eurygerris and Tachygerris are not each others closest relatives, which makes it likely that the pretarsal arolia have been lost and/or regained several times in Gerrinae, and therefore have little diagnostic value. According to Andersen (1995a), Tachygerris can foremost be diagnosed on the reduced apical lobes of second gonapophyses, while Eurygerris is diagnosed on a num-ber of character reversals.

The paraphyly of Gerrinae, Gerrini and Tachygerrini makes it reasonable to question the current classification of the entire clade containing Gerrinae and Eotrechinae. The Eotrechinae is diagnosed by the genital characters mentioned above, but has little branch support. According to Andersen (1995a: Table 1: character 19), members of Aquarius, Gerris (Macrogerris), Gigantometra, Limnogonus, Limnometra, Tenagogerris, Tenagogonus, Tenagometra and Tenagometrella all share the modified “omphalium-type” of metasternal scent orifice sensu Matsuda (1960). If Tenagometrella is sister-group to Limnogonus and if Tenagometra is related to Limnometra and Tenagogonus as shown in Fig. 1, the “ompha-lium” type of metasternal scent orifice could be a synapomorphy for the clade comprising these taxa, even though it has been lost in Neogerris, and has been independently derived in Gigantometra, Aquarius and Gerris (Macrogerris), which belong to another clade.

Matsuda (1960) considered Eotrechinae a tribe of Gerrinae along with Gerrini, Cylindrostethini and Charmatometrini, all of which were instated as subfamilies by Andersen (1975). Monophyly of the entire clade Eotrechinae + Gerrinae + Charmatometrinae + Cylindrostethinae was confirmed by Damgaard (2008a), but the clade could not be diagnosed on any morphological character combinations. Based on the present study, one could consider downgrading Eotrechinae to a tribal rank, which consequently would reinstate Gerrinae as a monophyletic group.

The relatively well-supported clade comprising genera with the “omphalium-type” metasternal scent orifice could qualify as another tribe, but as mentioned above the structure of the scent orifice is neither universal in - nor confined to - this clade.

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Also the clade comprising the remaining gerrine genera lack support and diagnostic character combinations. This clade includes reprensentatives of the “Tachygerrini” as well as of the “Gerrini”, incl. the enigmatic Gigantometra gigas, which, uniquely in Gerrinae, has evaporative scent grooves, otherwise found in Charmatometrinae and Cylin drostethinae (Andersen 1995a). Matsuda (1960) assigned Gigantometra gigas as the closest relative of Gerris, Aquarius and Limnoporus on basis of the placement of the metathoracic spiracle, and it has been used as such ever since, but from our study it is evident that the species has no close relationship with this clade.

Speciesgroups of Gerris, Aquarius and Limnoporus

Andersen (1990, 1993) and Andersen & Spence (1992) divided Aquarius, Limnoporus and Gerris into a number of presumably monophyletic species-groups, each represent-ing one or more species, and these hypotheses were tested in a series of studies (Sperling et al. 1997; Damgaard et al. 2000; Damgaard & Sperling 2001; Damgaard & Cognato 2003, 2005; Damgaard & Zettel 2004; Damgaard 2005, 2006, 2008b). Based on our results we can confirm the Gerris marginatusgroup (G. alacris, G. comatus, G. insperatus, G. incurvatus and G. marginatus). The group is diagnosed on having a short lateral process of proctiger, and absence of pale, marginal stripes on anterior pronotum. According to Andersen (1993), G. comatus, G. marginatus and G. incurvatus are diag-nosed on the ventrally keeled male segment 8, and G. marginatus and G. incurvatus on basis of the laterally modified male proctiger. The very close relationship between G. comatus, G. incurvatus and G. marginatus found in our study calls for additional studies of the taxonomic validity of these taxa.

Andersen (1993) placed the Mexican G. firmus Drake & Harris, 1934 from Mexico as “seeming closest to the G. marginatus species-group” even though the species is smaller than other New World species of Gerris, and its relatively long antennae (more than one half of body length) is not congruent with the diagnosis of Gerris. Finally, G. firmus is predominantly apterous, which is unique in the predominantly macropter-ous G. marginatusgroup. G. latiabdominis was tentatively assigned to the G. marginatusgroup by Andersen (1993), but Damgaard (2006) found that G. latiabdominis was sister-group to G. argenticollis although the support for this constellation was weak. In the present study the relationship between G. latiabdominis and the G. marginatusgroup is reinstated, and may be diagnosed on the presence of silvery pubescence on pronotum, even though this is not present on G. alacris. The support for the constella-tion is weak, and we prefer to maintain G. latiabdominis in its own species-group until more evidence is provided. The sister-group to the G. latiabdominis + G. marginatus group clade is G. thoracius although the clade is poorly supported. Andersen (1993) placed G. thoracicus with G. costae and G. sahlbergi in a G. thoracicus-group, but Damgaard (2006) found that G. thoracicus and G. costae were not closely related and down ranked G. sahlbergi to a subspecies of G. costae (along with G. c. costae, G. c. fieberi and G. c. poissoni). In our study, G. costae is sister-group to G. swakopensis, which is widely distributed in the Afrotropical Region. The G. swakopensisgroup includes a few additional species from the Afrotopical Region, such as G. gobanus

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Poisson, 1940 and G. zuqualanus zuqualanus Poisson, 1940 from Ethiopia and G. z. milloti Poisson, 1949 from Madagascar. The sister-group of G. costae + G. swakopensis is the G. lacustrisgroup (G. brasili, G. gibbifer, G. lacustris, G. maculatus). The G. lacustrisgroup includes a few more species from the eastern Palaearctic Region (G. angulatus Lundblad, 1934 and G. lobatus Andersen & Chen, 1993). The sister-group to all abovementioned taxa is the G. odontogastergroup, comprising G. odontogaster, G. argentatus; G. babai, G. caucasicus, and G. kabaishanus Linnavuori, 1998 (this species not studied here) from the Palaearctic Region, and G. buenoi from the Nearctic Region. Damgaard & Sperling (2001) named the entire clade “the derived Gerris”, and G. nepalensis and G. argenticollis were placed as successive sister-taxa to this clade, and the entire assemblage is Gerris s.str. The recently described G. curvus Tran & J.T. Polhemus, 2012 from Viet Nam was placed as sister species to G. nepalensis by its authors, and will be important to include for investigating the relationships among species of Gerris s.str. Finally, we found the subgenera Gerriselloides and Macrogerris to be sister-groups. Both subgenera have a number of species awaiting inclusion, and which will help to shed light on the basal relationships of Gerris s.l.

Within Limnoporus we find the same constellation of species as Damgaard (2006), i.e., an L. esakiigroup containing L. esakii from East Asia, a L. canaliculatusgroup comprising L. canaliculatus from eastern North America, and a L. rufoscutellatusgroup containing L. genitalis from East Asia, L. rufoscutellatus from the Palaearctic Region and Alaska in the Nearctic Region and L. notabilis and L. dissortis from eastern and western North America respectively. Aquarius (minus the A. remigis-group and A. chilensis) was found to be sister-group to Limnoporus, and we can confirm the A. conformis-group from Andersen (1990) comprising A. elongatus from East Asia and A. conformis and A. nebularis from eastern North America and the A. paludum-group sensu Damgaard & Zettel (2004) comprising A. paludum from the Palaearctic Region (incl. A. p. amamiensis from Amami-Oshima and the Ryukyu Islands south of Japan); A. antigone and A. fabricii from Australia; A. distanti from the Afrotropical Region; A. adelaidis from the Oriental Region, A. philippinensis from the Philippines; and A. lili from Timor. Andersen (1990) established an A. najas-group comprising A. najas, A. cinereus and A. ventralis from the Palaearctic Region, and tentatively placed A. chilensis in this group due to resemblance in several aspects, such as size difference between the smaller male and bigger female, the depressed meso- and metasternum of the male, and the incrassate and arched male fore femora. Andersen was concerned that A. chilensis had several characters distinguishing it from most congeners, such as second and especially third antennal segments relatively long; anterior pronotum only slightly elevated behind eyes; metathoracic spiracles removed a little more than their own length from wing bases; middle tibia subequal in length to femur; connexival spines extremely short in both sexes; and gynatrial sac of female with long and narrow basal part, and bristle-like ventral arolia on pretarsi of all legs. In our study we find a poorly supported sister-group relationship between A. chilensis and the A. remigis-group comprising A. amplus, A. remigis and A. remigoides, and the clade is sister-group to Gerris, even though the relationship is poorly supported. The lack of strong branch support and diagnostic characters for the entire clade makes it likely that A. chilensis

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and the A. remigis-group should be transferred to two new genera. A monotypic genus containing A. chilensis could be diagnosed on the characters mentioned above, while a new genus for A. remigis and relatives could be diagnosed first of all on the modified male genitalia and the relative length of the antennal segments.

Sistergroup relationship between Neogerris and Limnogonus

Earlier authors treated Neogerris as a subgenus of Limnogonus, but Andersen (1975) reinstated its generic rank, and stated that the two genera were quite distantly related. This was also found by Andersen (1995a), who placed Neogerris as sister-group to Gerrisella on basis of the relative short and broad head, the bottle shaped accessory gland sclerite, and absence of claws on the middle tarsus, while Limnogonus was placed in a clade comprising Tenagogonus, Limnometra, Tenagometra and Tenagometrella fore-most on basis of the female gynatrial sac with narrow base and widened distal part. The strongly supported sister-group relationship between Limnogonus and Neogerris revealed in the present study suggest that the earlier authors were right in assuming a close relationship between the two. Both Neogerris and Limnogonus are relatively well supported in the present analysis, but none of the genera are particularly well-supported in terms of morphological characters. Andersen (1995a) diagnosed Neogerris on basis of the relative length of pronotal lobe in apterous form (if present) covering anterior part of mesonotum and absence of inner lobe of first gonapophysis, as well as two reversals: posterior margin of sternum 7 of female more or less produced in mid-dle, concealing gonocoxae (also found in Eotrechinae and in Eurygerris + Tachygerris), and phallotheca ventrally sclerotized, dorsally membranous (also found in some Gerris). Andersen (1995a) diagnosed Limnogonus on basis of one reversal (lateral sclerites of vesica slender, spoon shaped), and two homoplasies: fore femur of male thickened, more than twice as thick as middle femur, and middle femur distinctly shorter than body length. Traditionally, Limnogonus has been divided in two subgenera, Limnogonus (s.str.) and L. (Limnogonoides) Poisson, 1965, which are foremost recognized on the presence respective absence of two light elongated spots on the anterior part of the pronotum, but as shown by Andersen (1995a) only Limnogonus (s.str.) could be diag-nosed on convincing synapomorphies, and Damgaard et al. (2010) found that the subgenera were not reciprocally monophyletic. While Limnogonus was not strongly diagnosed by Andersen (1995a), he diagnosed the clade comprising Limnogonus and Tenagometrella on the unique structure of the metasternal tubercle and scent orifice as well as several character reversals. Inclusion of representatives of Gerrisella and Tenagometrella will be important for understanding the basal diversification within the entire clade.

Within Limnogonus we recognize the following clades from Damgaard et al. (2010): Clade I (L. aduncus, L. recens, L. profugus and L. ignotus, all from the Neotropical Region), Clade II (comprising L. nitidus Mayr 1865, from the Oriental Region), Clade III (comprising L. franciscanus from the New World and L. cereiventris from the Afrotropical Region), Clade IV (comprising L. hungerfordi and L. luctuosus from the Oriental and Australasian Regions) and V (comprising L. fossarum F., 1775 from

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the Oriental and Australasian Region); and Clade VI (comprising L. intermedius Poisson, 1941 from the Afrotropical Region and L. pectoralis (Mayr, 1865) from the Oriental Region). Damgaard et al. (2010) also had a Clade VII (comprising L. hypoleucus (Gerstaecker, 1873), L. nigrescens Poisson, 1941, L. poissoni Andersen, 1975 and L. capensis China, 1925 from the Afrotropical Region), but in our study we have L. windi Hungerford & Matsuda, 1961 from Australia nested within this clade. The preliminary clade names from Damgaard et al. (2010) should be replaced by proper species group names when the remaining species of Limnogonus have been included in a phylogeny.

Taxonomic status of Limnometra, Tenagogerris and Tenagogonus

As mentioned above, the relationship between Limnometra, Tenagogerris and Tenagogonus is relatively well supported, even though none of the genera are themselves monophyletic. Andersen (1995a) recognized a close relationship between Limnometra and Tenagogonus diagnosed on the relatively long fourth antennal segment, the pre-dominantly pale pronotum, and the convex plate-shaped accessory scent gland sclerite, and he considered Tenagogerris as sister-group to a clade not only including Limnometra and Tenagogonus, but also Tenagometra, Tenagometrella and Limnogonus and another clade comprising Gigantometra, Limnoporus, Aquarius and Gerris.

When describing Tenagogonus australiensis, Andersen & Weir (1997) considered it to be closest related to T. fijiensis and T. valentinei Hungerford & Matsuda, 1961, both Fijian, but stated, “In both these species, however, the posterior corners of the female connexivium are more distinctly prolonged and may overlap each others. Besides, the pronotum of apterous individuals of T. fijiensis has a very characteristic, narrow projec-tion [Matusuda, 1960: Fig. 375]. The male abdominal end of both Fijian and the Australian species is relatively simple, not modified as in seveal other Tenagogonus species, including T. kampaspe (Kirkaldy, 1901) from New Guinea. An undescribed species, closely related to T. australiensis sp. nov. occurs in Papua New Guinea [J.T. Polhemus, in litt.]”. In our study, we find the two Australasian species, T. australiensis and T. fijiensis, in a clade along with the three species of Tenagogerris and Limnometra lipovskyi, all from Australia. The strong support for the clade containing the three species of Tenagogerris and Tenagogonus australiensis calls for par-ticular attention, and suggests the latter should actually be moved to Tenagogerris, but such a step will rise the question what to do about Tenagogonus fjiensis and Limnometra lipovskyi both of which are members of the same larger clade, while the likewise Australian L. cursitans is deeply nested within the clade containing the remaining species of Limnometra plus Tenagogonus bergrothi. The sister-group to the Australasian clade is comprised by three African members of Tenagogonus: T. madagascariensis, T. zambezinus and an unidentified nymph from Gabon (T. sp. #1566). This nymph may represent T. albovittatus Stål, 1855, which is a widespread species in Africa, but this should be confirmed with inclusion of identified material of this species.

A number of additional species are found in the Australasian region, such as Limnometra faracii Zettel, 2007 and Tenagogonus valentinei from Fiji, L. grallator

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D.A. Polhemus & J.T. Polhemus, 1998, L. monochroma Nieser & Chen, 1992 and Tenagogonus kampaspe from Papua New Guinea, Limnometra hysterema Nieser & Chen, 1992 from the Solomon Islands, L. kallisto (Kirkaldy, 1899) from Indonesia, Papua New Guinea and the Solomon Islands, and L. ciliata Mayr, 1865 and L. cilioides Andersen & Weir, 1997 both of which are found in the Australasian and the Oriental Region. Inclusion of these species will be important in order to investigate whether they belong to the Australasian clade containing L. lipovskyi, Tenagogonus australiensis and T. fijiensis or the Oriental clade containing the remaining included species of Limnometra and Tenagogonus bergrothi. A number of species of Limnometra and Tenagogonus are found in the Oriental Region (see checklist), and should be investi-gated in order to see if any of these have close relatives in the Australasian Region. The outcome of our study is that three zoogeographically restricted clades exist within the Limnometra, Tenagogerris and Tenagogonus clade, but that none of the presently accepted genera are monophyletic.

Hungerford & Matsuda (1958) listed Limnometra pulchra from Tanzania, but this record is most likely erroneous. The only species of “Limnometra” entering the Afrotropical Region is L. fluviorum, which is found in India and Sri Lanka and on Réunion Island, but according to our analysis it is not a typical Limnometra.

Fossil history of Gerrinae

Despite being undoubtedly monophyletic and strongly supported, the family Gerridae as currently defined can hardly be diagnosed on any convincing morphological synapomorphies (Damgaard 2008a). On the other hand, most members of the Gerridae have long and slender legs, and cannot be confused with other extant groups of organisms, and only the extinct orthopteroid family Cresmodidae may have inhabited a similar environment (Grimaldi & Engel 2005). Along with members of the family Hydrometridae (water measurers), the Gerridae are the only fossil taxa that can be reliably identified from compression fossils, since other gerromorphan families have much more generalized appearances (Damgaard 2008c). While the represented subfamilies, Ptilomerinae, Halobatinae, Rhagadotarsinae and Trepo batinae, can be identified on reliable, external, macroscopic characters, members of the Charmato metrinae, Cylindrostethinae, Eotrechinae and Gerrinae can hardly be separated on basis of characters available from fossils. Andersen & Poinar (1992) established a new subfamily Electrobatinae for an inclusion in Mid-Miocene (20–17 ma) Dominican amber, and the presence of just three pairs of cephalic trichobothria in this subfamily places it among the basalmost lineages of Gerridae.

Nevertheless, a number of fossil taxa have been assigned to Gerrinae, and several of these have convincingly been placed in extant genera, such as Aquarius lunpolaensis (Lin, 1981) from Miocene (25–5 Ma) Tibet, which was placed in the A. najas-group by Andersen (1998), and Limnoporus wilsoni Andersen, 1998 from Lower-Middle Eocene (52–47 Ma) shales in British Columbia and placed in the L. rufoscutellatus-group.

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Damgaard (2008c: 263) suggested that members of Telmatrechus Scudder, 1890, with three described species from Lower-Middle Eocene deposits in North America, were congeners to the extant A. remigis-group sharing the same relative lengths of the antennal segments, and we agree that they should be re-examined in a future revision of Aquarius. According to Andersen (1998), the three species of Paleogerris Andersen, 1998 from Paleocene-Eocene moclay deposits in Denmark may be closely related to Gigantometra gigas sharing the distinct evaporative scent channels on the metathorax and the metathoracic spiracle close to the wing basis as in this species. Palaeogerris is also of considerable size (27–31.5 mm body length) and thereby matches Gigantometra gigas (32–36 mm), but males of Paleogerris do not have the extraoridinarily long middle and hind legs characteristic of this species. Based on Andersen (1998) and the present study, it is very likely that Paleogerris is sister-group to Gigantometra, and belong to a clade that used to be much more widely distributed in the Palaearctic Region, and with a single species left today having a peculiar disjunct distribution in Viet Nam and southern China (Hainan) (Andersen, 1995b). The well-preserved Electrogerris kotashevichi Andersen, 2000 and Succineogerris larssoni Andersen, 2000 are both from Eocene-Oligocene (54–38 Ma) Baltic amber, and were placed as relatives of Eurygerris and Neogerris respectively by Andersen (2000) and Zettel & Heiss (2011).

Acknowledgements

We wish to thank everybody who has contributed material, information or ideas to the present study and to two anonymous reviewers for suggestions that have greatly improved our paper. The study was supported by grants from the Villum Kann Rasmussen Foundation to J.D.

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Molecular phylogeny of the pond skaters (Gerrinae), discussion of the fossil record and a checklist...

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