Scholarly article on 2-Ethylhexanol 104-76-7 from Journal of the American Chemical Society p. 813,819

DOI: 10.1163/187631202X00154

Source and publish data:

Journal of the American Chemical Society p. 813,819 (1938)

Update date:2022-08-24

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Authors:

Magnani Magnani

McElvain

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Article abstract of DOI:10.1163/187631202X00154

Full text of DOI:10.1163/187631202X00154

The

surrounding

Part 6.

bibliography

JOHN T. POLHEMUS and DAN A. POLHEMUS

of New

Guinea

and

Trepobatinae

(Gerridae)

with

review

of the world

fauna.

regions,

Phylogeny,

and

world

addenda

biogeography,

final

taxonomic

checklist,

The small waterstridersof the

have radiated

on New

extensively

subfamilyTrepobatinae

Guinea and

The

contributionis the sixth in a series of

surroundingarchipelagos.

present

reportsdealingwith this endemicfauna, and presentsa cladisticanalysisat the genericlevel

for the

a world

appendix

checklist, completebibliography,

and

subfamily, biogeographicanalysis,

final taxonomic addenda.The taxonomic

contains new distributionalrecords for

mubiPolhemus& Polhemus,and

of the

descriptions

newtaxa:thenew

following

Stygiobates

genus Talaudiato hold the previouslydescribedAndersenellanilsi Chen & Nieserfrom the

Talaud

Metrobatoides

n. from northern central Irian

bifurcatus sp.

NewGuinea.Thecurrentcladistic

and

Archipelago;

Jaya;

Iobates ivimka n. fromsouthern

sp.

Papua

analysissupports

the

tribalclassification,withthe

previouslyproposed

Metrobatini,Trepobatini,Naboandelini,

and Stenobatiniall resolvedas discrete

Our cladisticand

monophyleticgroups.

biogeograph-

that the currentdistributionof

in theNew Guinea

trepobatinegenera

also

suggest

analyses

region has been strongly influencedby past tectonic events, most notably the accretionof

islandarcterranesderivedfromthe SolomonsandHalmaheraarc

Thishasledto dis-

systems.

tinct areasof freshwaterandmarineendemism

areas

the Metrobatiniand Stenobatiniin the

among

Halmahera,the

Peninsula,and northeasternNew Guinea

surrounding

Vogelkop

plus

region sug-

islands.

contrast,the distributionof Naboandeliniin the New Guinea

nearby

gestsa vicariancepatternwithAustraliaacrosstheTorresStrait.

J. T. Polhemus,

USA.

of Colorado

3115 S. York

CO.

80110,

University

Museum,

St., Englewood,

D. A. Polhemus,

MRC 105, Smithsonian

Department SystematicBiology,

Institution,

WashingtonD.C., 20560,

USA.

Introduction

onomic details,

one additional new

species.

including

and two additional new

This

the sixth and final

of our

paper represents

part

of New Guinea

genus

Given the

New Guinea

of the

in the

monograph

Trepobatinae

diversity

Trepobatinae

and

In the

five

region,

fully expect

further new

surrounding regions.

parts

of this series (Polhemus & Polhemus 1993, 1994,

and

even new

to be dis-

genera

species

possibly

995, 1996, 2000) we proposed four new tribes,

covered by future collecting,

on the

particularly

to the east and west of

taxonomic revisions for each,

more remote island

providing

descriptions

In this final section we

including

species.

groups

Even

of 10 new

and 47 new

New Guinea

so, any

additions to the

genera

proper.

known biota can be treated in future

provide

at the

phylogenetic

currently

to the current

plements

sup-

which has

for the

level, fol-

analysis

lowed

subfamily

generic

monograph,

sufficient for

discussion, a world bib- reached a

by biogeographic

stage

completion

synthetic phylogenetic

below.

graphic analyses provided

pres-

and a world checklist. We also attach an entation of the

and

liography,

biogeo-

with a few small

tax-

appendix dealing

remaining

eredto be

Elaborationsare consideredto

plesiomorphic.

Phylogeny

Character

but an

apomorphic,

of these is

apomorphicsequence

Extensive character

have been

previously provid-

unclear.

.) Bodysize (character2).Anessentiallyequalbody

sizein malesandfemalesis consideredto be

analysis. -

analy-

ses of

gerrid genera

plesiomor-

sizebetweensexes

Matsuda (1960) and Andersen (1982).

whilea

differencein

body

phic,

great

Efforts to use these for the

is considered

present study proved

gerrid gen-

apomorphic.

Matsuda's work covers all

3.) Headshape(character4). Theplesiomorphichead

shape is consideredto be that of Rhagadotarsu.r.Four

frustrating.

era known at that time, but for the

ed here is

inadequate

amount of material available for certain

intend-

purpose

for several reasons: 1.) the

other

apomorphicmorphologiesprovide unambiguous

charactersfor the four tribes of

the of

however

Trepobatinae,

key gen-

in Stenobatini and

anageneticsequence development

is not clear.

era was

4.) Omphaliumposition(character11).The omphali-

um, or metastemal scent gland opening, occurs at the

inadequate, especially

Naboandelini; 2.) almost half of the genera we

of the metastemum, or is

vestigial

posterior edge

have before us were unknown at that time; and 3.)

absent, in all trepobatinegeneraexcept those now held

the illustrations of the male

for

in the tribe Stenobatini;in these latter five

genera,

the

aedeagus, provided

for which males were available to him,

is medial in three, and locatednear

in the other two. The more anterior

all

omphaliumposition

genera

are inaccurate,

the anterior

margin

is unusual,

details of some

omitting important

in the Gerridae:

position

Andersen

perhaps unique,

sclerites while at the same time

incorrectly depict-

states that 'when

the

(

1982: 201)

present,

as sclerites darkened

of the

metastemalscentorifice

of theGerridaeis

ing

cuticle and internal structures. Andersen's work

covers the of the with

regions

aedeagal

(Omphalium)

half of the

located in the

but this

posterior

segment,'

clearlydoes not hold

true for all Stenobatini.

entirety

Gerromorpha,

of all features and

.) Wings (character 13).Wings,or vestigesthereof,

and

analyses

descriptions

major

are found in all

of Gerridae

the

except

genera

truly

of these insects, and is thus

understanding

many

because of the

use-

marine

Thusthe invariantlack of

charac-

aspects

extremely

homologies

genera.

wings

Trepobatinae.

Halobatinaealso

terizes the tribe Stenobatini within the

ful in

the

and variabili-

Other marine

in the

subfamily

genera

knownin the

structures and features.

Conversely,

of this

are

form.

wingless

only

broad

extremely

coverage

6.) Hind tarsal claws

character 18).

The hind tarsal

(

work, there is not enough included detail concern-

claws of trepobatinegenera vary widely,from short or

hair-like, long

slenderand blade-like.The latter well

the

to establish

single subfamily Trepobatinae

ing

claws are found in the marine tribe

developed

a character set for

In addition, as with

genera.

Stenobatini.The three

of the tribe Naboandelini

genera

Matsuda's work, almost half of the

here were unknown to Andersen. The character

treated

genera

all have shortor hair-likeclaws.The

remaininggenera,

in the tribes

and Metrobatini, have

Trepobatini

long

slender claws reachingalmost to the apex of the tarsi,

that follows is therefore based

analysis

primarily

the

Metrobates in which the claws are

genus

except

upon original investigations.

Characters have been

shorter,not nearlyreachingthe tarsal apex.

with

analyzed

regard

7.) Male anterior tibia, distal comb (character 19).

The distal comb of the male fore tibia has different

forms:the more elaborateare consideredto be apomor-

phic, and help define clades, but the functionalsignifi-

canceof the variousformsis unclear.

trends and

For some char-

stability.

developmental

acters, the apomorphic state can be inferred by rel-

tests or observations, i. e.: charac-

atively simple

ters with

occurrence

(characters

8, 30,

exceptional

5); characters that have arisen de novo (charac-

ters 25, 26, 31, 32); total or loss of a struc-

8.) Posterior femur

The

length (character 21).

Trepobatinaetendto haveshortermiddleandhindfemo-

ra than

in other subfamilies,with some in

gerrids

genera

shortmiddlefemora.In mosttre-

partial

particularhavingvery

ture found in

or ancestors

closely

character 10); characters that exhibit elaboration

of structure (characters 3, 5, 6, 7, 9, 12, 16, 17, 20,

3, 29, 34). Conversely, the plesiomorphic charac-

ter state can be inferred if a structure is homolo-

groups

the hind femur is shorterthan or

to the

pobatines

equal

(

but in a few

it is

genera

than the

body length,

longer

Whilethis characterthus

to define

body.

helps

genera,

forthe

doesnot

provide unambiguoussynapomorphy

subfamily.

.) Posteriortarsal segmentation(character23). The

with that in more

taxa or ancestors

gous

characters 14, 15).

Particularly important,

character

primitive

tarsi of

exhibit various tarsal

Trepobatinae

posterior

(

lengthratios,andmaybecompletelyfusedin somegen-

era. In the

the two tarsal

areeither

controversial,

systems (ie.,

those for which

Trepobatini

segments

I is the longest.In the

I is about 1/2to 2/3 the

of II,

length

of about

equallength, segment

ambiguous

cannot be

Stenobatini

segment

inferred a

are

polarity

discussed in

directly

priori)

in some

allarefused.The

although

in the Naboandelini is similar to that of Stenobatini

for the

where the

species

segmentation

detail below.

greater

except

genusCalyptobates,

segments

.) Predominant color (character 1). Unicolorous in all species are always fused. The segmentsare uni-

withoutextensiveadditional

is consid-

fused in the

genera

of Metrobatini, in the

except

grey,

markings,

formly

Metrobates where

I is about 1/4 the the external and internal structure of the

proctiger

genus

segment

two.Thefunctional

of the

significance

(and

related on other

length segment

Metrohatopsis

genera closely

differentthan of the latter

any genera,

isunclear,

definetribalclades.

segmentation except help

grounds) quite

0.)Scleritesof the vesica (chracter26). The vesical suggestingthat their originsand functionsare different.

scleritesandthe versus states the small size of these waterstridersand

apomorphic

plesiomorphic Unfortunately,

of various

arrangements

have been treated Andersen the

in them in the field

consequentdifficulty observing

(

1982, 1991).Of particularinterestis the proposal that has limited our understandingof the function of these

the

state is

plesiomorphic

the four

lateral

likely,

Involvement in

seems most

representedby

major

basal (bs),

ds),

spines.

however.

copulation

medial sclerites [anterior (as), dorsal

(

ventral(vs)]

and

beingseparated

as individual

12.) Male

parameres, shape

(character 28). Small

appearing

sclerites, whereas the continuously coiled vesica that simpleparameres,or their absence,are most likely the

continues

as seenin the

apomorphic.

state in Gerridae. The

or more

larger

anteriorly

genusRhagadotarsus plesiomorphic

is consideredto be

This

in our

are consideredto be

and

analysis,

complex shapes

apomorphic,

view,has thepolaritiesreversed.Areviewof thevesical these may exhibit several forms whose significanceis

architecture of a wide

and unclear,

that

delineate the

genus

range

Homoptera

except

they help

Trepobatini

analysis possesslarge complexparameres.

as the basal (character29). In contrastto

13.) Female

that the continuous vesica is

Metrobate.s ,a nd the tribe

whereinall

Heteroptera argues

ple-

genera

siomorphic.Furthermore,

Andersen's cladistic

Rhagadotarsinae

ovipositor

poorly developedovipositor

of the Gerridae

the

though

places

even

subfamily,

he holds that the male

the

of most

mem-

a well

genitalia

gerrids,

subfamilyRhagadotarsinaepossess

the most derivedcharacterstate for the vesical bers of the

possess

sclerites, thus even a computer generated cladogram developed,longovipositorwithdistinctserrationsonthe

on the basis of

that the continuous first

to be assocatedwithembed-

argues

parsimony

plesiomorphic.

that the continuous

gonapophyses ,t hought

vesicais

in tissues.Andersen(1982:224)believes

dingeggs plant

vesical that this is

To our

propose

sclerites represent the

uninterrupted

state

probably secondarydevelopment.

we discoveredthat an with serrate

plesiomorphic

tarsus), withthe followingtrends increasinglyapomor- second gonapophysesis also present in all genera of

in the in a firm sub-

(Rhagado- surprise,

ovipositor

phic, although perhaps not exactly

following

figures

Stenobatini,

suggesting egg deposition

sequence(abbreviationsgiven

are those used in

strate,

the

are ovalandshownomodifica-

although eggs

series): tionsto indicatea siteor substrate

particularoviposition

of this

published previous parts

monograph

.) medial sclerites (as, ds, bs, vs) increasingly frag- (Polhemus& Polhemus 1996: 287; Andersen & Weir

mentedand/orreduced;2.) lateral sclerites(Is)

1998:

531, fig.38).

14.) Mesothorax

present,

distally,

simple, rod-like; 3.)

lateral sclerites modified

(character32). The mesotho-

shape

bifurcate,expanded,knob-like,etc.; 4.) lateral sclerites rax of Trepobatinaeis usuallytumid ventrally,however

plate-like(e. g. Metrobates,many Trepobatini);5.) ap-

of the medial sclerite(ms) as a new structure

in two

dorsoventrally

genera

flattened.The

of Stenobatini the mesothorax is

pearance

of this feature

significance

(

e. g. Andersenella);6.) appearanceof the apicolateral is unknown,but may aid in phoresy.

sclerite(als)as a new structure;7.)baseof anteriorscle-

15.)

Bodyshape(character

34).The usual

bodyshape

rite

(

bifurcate, even

waterstridersis

trepobatine relatively

narrowandnot

expanded, broadly

lyre-shaped

Stenobatini);8.) loss of all median sclerites; usually flattened.The generaMetrobatesandMetrobatoidesare

accompaniedby 9.) sclerotization

of the distal

heavy

broad,

flat-

portions unique possessing very

dorsoventrally

onthemidstreamsectionsof

skating

of the

as well as

basal, dorsal and tened associatedwiththeir mode of liv-

phallotheca,

body,probably

whichinvolves

ventralsclerotization(character24).

1.)Maleproctiger;ie., abdominalsegment10(char- largestreamsand rivers.

acters 27, 29). The is rather liable to

ing,

proctiger

modification,and its use as an indicatorof phylogenetic

must be

with caution.Various Characters

plastic,

relationships

approached

modificationsand elaborationssuch as lobes,

protuber-

ances,tuftsof spines,andlateraloutgrowthsin the form Morphological data were analyzed to obtain a set

of or are seen in various

digitateprotuberances spines

of characters suitable for

the

phyloge-

determining

gerridlineages g.

e. Halobatinae(someHalobates),Ptil-

In all, 39 characters were

ny trepobatine genera.

omerinae(somePtilomera)and

(allNabo-

Trepobatinae

identified that offered useful

infor-

phylogenetic

andelini, some Metrobatini, all Stenobatini). In some

the lateral

are

outgrowths very large

and accom- mation. These characters and their states are

pygophore described below. The character states are coded

groups

modificationsto the

paniedbycorresponding

and/or eighth segment. In these groups (e. g. Nabo-

with 0;

of these

using integers beginning

characters were coded as multistate, with the num-

many

andelini,Stenobatini,someMetrobatini)the morpholo-

seems

more stablethan in those

groups

exhibiting

Lateral spines or

considerably

modificationsto the

alone.

appear

ber of discrete states for a

character

proctiger

given

The character

varying

that

homolo-

protuberances

from 1-8

Tab.

(see

1).

priori assumptions

numbering

as to character

gous in their form, functionor origin may not be. The

genus Metrobatopsis has lateral protuberances that

indicates no a

state

to be

with those of the Naboandelini

analogous

polarity

weighting.

appear

(

Naboandelus, Hynesionella and Calypobates) and

Stenobatini (Rheumatometroides,Stenobates, Steno- Descriptions of

howevercarefulexaminationrevealsthatboth

characters and character states

Characters followed

for genera of Trepobatinae. -

batopsis),

had no

of polarity and were

0 = absent

= present

by '(U)'

hypothesis

run unordered in all iterations of the phylogenetic

. Mesonotumwithmediansutureor sulcusin female

0 = absent

analysis.

= present

10.Metastemal

scent

(omphalium)

glandopening

Body

in both sexes,largein males

prominent

. Predominantcolor pattern (groundcolor mentioned

= reducedor vestigial

first) (U)

= secondarilyabsent

grey*

1.Omphaliumposition(U)

= black,

or blackish grey with yellowish or

grey

leucinemarkings

= black with

locatedat posteriormarginor absent

= locatedmedially

grey markings(exceptlight spot

light

locatednear anteriormargin

pronotum)

2.Mesothoracicspiracleregion

= light brown, leucine,yellow or orangewith dark

= undifferentiated

markings

=differentiated,cuticle surroundingspiracle defi-

= blackish brown with yellowishor leucine mark-

differentthanthat of adjacentmesonotum

nitely

ings

= black to blackishbrown with light brown to red

Wings

= sometimespresent

brown

size

markings

never

present,

alwayswingless

Body

= essentiallysamein malesand females

Abdomen

dimorphic,malesmuch smaller

sexually

4.Malebasal abdominalstemites

= wide

Head

= narrow,ring-like

. Antennaewith stiffspine-likesetaedistally

5.FemaleventriteVII

= absent,or scatteredwidelyon segmentsI and II

verticalor angledposteriorly;may

edge

ondistal 1/4on at least segmentsI andII,

posterior

present

few in number

on distal extreme of at least segmentsI

or may not exposegonocoxae

angled anteriorly or excavated;

posterior edge

broadlyexposinggonocoxae

present

and II

. Head shape(U)

= Head long, extending anteriorly; gular region Legs

antennalsocketremovedfromeyeby width 16.Anteriortarsi,cross section

long;

round,ovate, flattened,

narrow

of socket;base of rostrumat mid eye level

= Same as 0 except base of rostrum at ventraleye

1 =blade-like,wide

of first

level

17.Anteriortarsi, setiferation

segment

without

setae

= Head short, gular region short, usually tumid,

long

belowventraleye level;antennalsock-

with

long

setae

ventrally

extending

et closeto eye,not removedfromeye bywidthof 18.Hindtarsalclaws

socket;baseof rostrumenclosedin ventrallypro-

jecting

eye level,

= short,oftenhair like

differentiatedrostralcavity,belowventral

present,longslender,tapering,reaching

apexof tarsi

almostto

directed

slightlyposteriorly

= Sameas 2, exceptgularregionnot tumid;base of

rostrum not enclosed, rostral cavity planar with

2 = present, same as 1,

ing apexof tarsi

but shorter,not nearly reach-

at about ventral

level

3 = present,long slender,blade-like

4 = falcate

gularregion,usually

eye

gularregionnot tumid;base of

= Sameas 2,

except

rostrum arising anteriorly,not enclosed; rostral

at or above gular region, arising at about

5 =hairlike, extending far beyond tip

absent

of tarsus, or

cavity

ventraleye level.

19.Maleanteriortibia,distalcomb (U)

. Malefourthantennalsegment,cross section

withnormalbrushof moderate

with a well defined

lengthstout setae

length

brush of moderate

elongate

setae

= roundor oval

widenedand compressed

slightly

= widenedand stronglyflattened

2 = with an elongate brush of short to

moderate

longerspines

. Rostrumshape,segmentIII

lengthsetaemixedproximally

or spine-likesetae

with

= linear or tapering, posterior margin straight to

withdifferentiated

shortor

curved

pad closelypacked

slightly

margintumid, stronglycurved

mediumlengthsetae

distal male

spur,

posterior

of rostralsegmentIII

20.Anteriortibial

absent

1 =present

21. Posteriorfemurlength

= shorterthan or equal to body includinghead

than

head

Length

= normal,lengthapproximately3x segmentI

= long,length4x segmentI or more

lengthapproximately6x segmentI

verylong,

Thorax

. Mesonotum with median membranous region in 22. Middletarsi,modificationsfor rowing

female

0 = unmodified,withoutswimingplume

longer

body including

= with swimmingplumeof long silkysetaepresent

1= as, ds, bs present;vs well formed;Is long,paired

or bifurcatedistally

3. Posteriortarsi. segmentation(U)

= withtwo distinct

of aboutequal length,

same as 1

large, broad, expanded

except:

betweenls, lobe-likein lateralview

bs ms

segments

segment longest

= with two distinct segments,segmentI about 1/2

as, ds, present;

long, large (may poorly

to 2/3 the

fused

of II, sometimes secondarily

sclerotized);ls long,rod-likeor slender

length

4 = as, ds, bs present; ms short, narrow; Is large,

plate-like;as basallylarge,expandedbetweenIs

= with two distinct segments,segmentI about 1/4

sameas 4

except: absent;

two

(1+1)auxiliiary

the

of II

length

alwaysfused

basal scleritespresent.

segments

= as, ds, bs present,and sometimesvs;ms absentor

poorly formed; Is narrow, basal, reaching only

Genitalia, terminalia

4. Phallotheca(U)

middleof

phallotheca

= Normal, more or less

membranousdistally

basally,

7 = as, ds, bs short or reduced,

lower dis-

cylindrical, open

occupying

tal

of phallotheca;Is long, slender;all other

part

scleritesabsent

all

medial sclerites lost

(as, ds, bs,

= Sameas 0,

distallypartlyclosedand scle-

except

rotized,forminga U-shapedwall [Ciliometra]

= Sclerotizedat least dorsally and distally,acumi-

major

absent);ls commencingbasally,long, lobed dis-

nate

[Metrobatopsis ,S tygiobate s] ]

als

tally ; present

dorso-distally

5. Male

withbroad internalphragma

27. Maleabdominal segment 10 (proctiger), with long

lateral anteroventrallydirected digitate or spine-like

proctiger

= absent

present,

simple

processes(U)

6. Vesicalsclerites(U)

0 = absent,or modifiedin differentform

= all four major sclerites (as, ds, bs, vs) present,

Is narrow,medial

1= present

28. Male

parameres,shape

(U)

forming ring;

Table1.Morphologica lc haracterdatafor25generaof Trepobatina ea nd4 outgrouptaxa(Aquarius,Charmatometra,

andRhagdotarsus).For definitionsof charactersand characterstates see text.

Cylindrostethus

.Strictconsensustree of 16MPTsfor

basedon 39

generaof Trepobatinae

characterswith a

morphological

Figure

charactersrun unordered(for discussionsee text).

= 102,CI = 0.676, RI = 0.851. Numbersabovebranche

Length

indicatebranch

lengths.

= small,

long.

complex

or absent

the

Charmatometra

simple,vestigial

blade-like, falcate or corkscrew

genera. Subsequently,

genera

(Kirkaldy) coded], Aquariums

and

I =

shaped;

bakeri

Kirkaldy [C.

Schellenberg [A. remigis (Say) coded],

Cylin-

9. Female

ovipositor

without

serrationsor teeth drostethus Fieber [C. erythropu.s (Herrich-Schaef-

plate-like,

significant

serrate

= first

gonapophyses

fer) coded]

were also added in order to

provide

taxa

exemplar outgroup

= second

serrate

gonapophyses

broad

from other

representation

subfamilies.

gerrid

Additional characters

0.Mesothorax,

shape

(U)

= tumid

ventrally

flattened,not tumid

shape

Parsimony analysis

dorsoventrally

ventrally

adapted

1.Male fore femur,

The 39 characters described above were coded in

for a matrix and analyzed using the computer program

PAUP 4.Ob8 (Swofford 2001 ).Parsimony analyses

= not

curvedor modified

strongly

and

strongly

smoothly

curved,

phoresy

modified, thickenedand sometimesbent,

highly

were

the heuristic

intially implemented using

with basal or medial

and often

large

sculpturing

with TBR branch

option

and

ventral

swapping,

protuberances

2.Body,

parsimony-uniformative characters excluded. The

same data were then

shape

relatively

very

narrow,not flattened

a Heuristic

analyzed

using

broad,

dorsoventrally

length

flattened

search with 100

of random

repetitions

stepwise

3.Middlefemur,

addition

which discovers 'islands' of

than middletibia

sequences,

shortest trees that may not be detected by other

addition All charac-

longer

= shorterthanmiddletibia

4.Fore wings,cells

sequences (Maddison 1991).

ters were run unordered, and no

= withmore than two closedcells

= withtwo closedcellsin basalhalf

were

assumptions

made as to character

These

5.Fore

weighting.

analyses

trees (MPTs)

wings

= withouttransverseline of weakness

both

16 most

parsimonious

100, CI 0.690, and RI 0.860.

= =

produced

I = withtransverseline of weakness

6. Ocularsetae

length

A strict consensus tree

from this ini-

generated

1) was examined with

than four

facets

longer

eye

= shorterthan four

tial set of MPTs

(Fig.

7. Lateral

suture between meso- and MacClade 3.01 (Maddison & Maddison 1992), to

assess the distribution of individual character

intersegmental

metathorax

= distinct

states on the tree. A second

parsimony analysis

was then run with all characters for which a rea-

= indistinctor absent

8. Base of rostrum

= not enclosed

= enclosed

sonable

the basis of

could be made on

or other evidence (see

hypothesis

polarity

morphoclines

9. Gularregion

Character

section above) run as ordered;

Analysis

this included all characters in the matrix except 1,

, 11, 19, 23, 26, 27, 28, and 30. This analysis pro-

= not tumid

= tumid

duced 18 trees of

110, CI = 0.627, RI =

was then run

length

Outgroups

.837. This same

analysis

again

Branch and Bound

rigorous algo-

Matsuda (1960)

inae and

esis

stead

subfamilies constitute the

the

the basal

the

terminal to the

the tree, a

these considerations, the

Breddin (R.

Rhagadotarsinae,

our initial

that the

the more

proposed

are sister

Andersen (1982: 237), who in-

that the

Rhagadotars-

using

rithm, resulting in the same 18 MPTs. Although

Trepobatinae

rejected by

hypothesized

groups, hypoth-

than the trees

from the

of ordered char-

slightly longer

unordered

generated

seven

(as is

remaining

sister

gerrid

analysis typical

of acter sets), the ordered-character trees were

probable

which his

clade. In Andersen's

group

signif-

more resolved. A strict consensus tree

icantly

gen-

erated from these latter trees

Rhagadotarsinae,

gerrid

Trepobatinae

analysis placed

2) contained

cladogram

position

(Fig.

form a sister clade one

well resolved clades

near the base of Stenobatini, Naboandelini,

the tribes

representing

and

Rhagadotarsinae

with which we

Trepobatini,

Given

Metrobatini, and improved internal resolution

within the Metrobatini, a

topology

agree.

genus Rhagadotarsus

Breddin coded),

zoo-

group

particular

interest in relation to the current

kraepelini

was chosen as the

subfamily

for

geographic

Branch

study.

for these clades was assessed

under

(Felsenstein

parsimony

outgroup

support

phylogenetic analysis

trepobatine

Bootstrap analysis

. Strict consensustree of 18MPTsfor

based on 39

generaof Trepobatinae

characterswith

morphological

Figure

certaincharactersrun as ordered(fordiscussionsee text).

102,CI = 0.676,RI = 0.851.Plaintext numbers

Length =

above branches indicate branch

pseudoreplicates

bold face numbers below branches indicate

from 500

lengths,

bootstrap support

in PAUP.

per pseudoreplicate

with 10random-additionheuristicsearches

Figure3. Taxon-areacladogramfor generaof Trepobatinae,basedon tree shownin Fig. 2.

with 10random-

per pseudoreplicate and parsi-

characters excluded; these

em Indo-Pacific, with their greatest diversity in

Melanesia and no representation in the Western

985) using 500 pseudoreplicates

replicates

mony-uninformative

addition

Hemisphere.

The Naboandeliniare also an

entirely

Hemisphererepresenta-

known

Paleotro-

values are shown in

2, and indicate

clades. The

forms the basis for the subse-

with no Western

pical group,

Bootstrap

Fig.

for all

major

tion. This tribe

includes the

represented

only Trepobatinae

in NewGuineaby a single

branch

strong

support

fromAfrica.It is

tree thus

generated

that also occurs in Australia and

genus, Calyptobates,

discussion of

the New Guinea region.

quent

trepobatine biogeography

extendswestward

Ceylon.

Borneoand the Andamansto

through

Western

.) The Trepobatini are a predominantly

with

two

genera, Cryptobates

Hemispheregroup,

only

Biogeography

and Gnomobates,present in the Eastern Hemisphere,

where

are confinedto SoutheastAsia,fromBorneo

they

to India.

The Metrobatini are

centered in New

primarily

The

of New Guinea is

biogeographic history

interwoven with the

tectonic his-

closely

tory

complex

4.)

Guinea and surrounding islands, with a single large

basalclade

Metrobates,in the Neotropicalregion

of the island's formation. As outlined by

genus,

and another

Polhemus (1996), and further discussed by

Polhemus & Polhemus (1998), New Guinea con-

Rheumatometra,in Australia.

genus,

sists of several

were

of island arc terranes that

the

to the fauna of New Guinea, which is

focus of this work, the island's assem-

primary

layers

laminated onto the northern

regard

successively

of the Australian craton between the

was derived

margin

blage

Trepobatinae

independently

Eocene and the Pliocene, with each episode of from three

tribes, the Metrobatini,

separate

Naboandelini and

accretion

a distinctive faunal

that are centered in

and do not form a

producing

Davies (1987) identified 32

signature.

ter- different

Stenobatini,

geographic regions

cluster of sister

Pigram

ranes

separate

north of the former Australian cratonic

on the clado-

lying

monophyletic

gram.

biota,

summary cladogram

using component analysis.

have also had

groups

and these

those

be divided into two

The island therefore has a multicentric

margin,

groups,

may

major

from an Eocene to

resulting

and will

appear

as a

area on

composite

any

derived from this data

of arc collision,

the

area

Oligocene episode

involving

follows Polhemus

Arc

these tribes

histories in

Papuan

Polhemus 1998), and those

(terminology

addition,

differing biogeographic

due to their

from a subse-

deriving

Miocene to Pliocene collison event, involv-

the

with

quent

region

divergent ecologies,

the Solomons Arc (Fig. 14-16). Polhemus & the Metrobatini and Naboandelini

ing

Polhemus (1998) provided evidence from aquatic

that the Eocene to

terranes harbored faunas whose affini-

inhabiting

freshwater, while the Stenobatini are marine. In

light

Heteroptera indicating

Oligocene

ties extended to the west, toward the Malay

while the Miocene to Pliocene ter-

of these

considerations,

of the three tribes are discussed

the

biogeographic

separate-

patterns

in the

sections.

following

Archipelago,

ranes had faunas with affinities extending to the

Metrobatini

The

east, toward the Solomons. Recently,

researchers

and animals

the terrane accre-

with other

dealing

have also

tion

groups

incorporate

plants

of Metrobatini shows a basal

cladogram

split

begun

in this tribe between the

occurs in the

Metrobates, which

and a clade con-

all of which are

genus

Neotropical region,

remaining genera,

into their analyses of regional bio-

hypthesis

geography (Welzen 1997;

In the context of the terrane accretion hypothe-

Heads 1999).

the

taining

in Australia, New Guinea,

Paleotropical, occurring

sis, the distributions and relationships

and

This

nearby archipelagos.

pattern sugges-

on New Guinea and

Trepobatinae

archipelagos

surrounding

are instructive. A taxon-area clado-

tive of an initial Gondwana distribution, which

could have been

vicariance follow-

of Australia from

that was initiated near 95 Ma

fragmented by

the eventual

separation

for the

was created

gram

by replacing

with the

genera

Trepobatinae

ing

Antarctica, a

the terminal taxa on our

tree

preferred

process

Australia contains

areas

(Fig. 3).

geographic

they occupy

(Kroenke 1996).

only single

with two

& Weir 1998), while New Guinea and

This taxon-area

reveals some obvious

cladogram

Rheumatometra,

genus,

species

the worldwide distribution of

patterns regarding

tribes:

(

Andersen

its associated

a localized

archipelagos support

.)The Stenobatiniare entirelyconfinedto the east- radiation of six genera,

two of which,

Stygiobates

. Distributionsof four

of Metrobatiniin the NewGuinearegion.

genera

Figure

and

extend westward to the north-

islands and accreted terranes of Miocene to

Pliocene but is not known from drainages on

Metrobatopsis,

ern Moluccas and eastward to the Solomons

age,

of the taxa of the Australian craton south of the island's central

respectively (Fig. 4). Many

Metrobatini

that conform to the

the island as

New Guinea have

divide

4, 5).

the

has many

occupying

ranges

(Fig.

Although

on the islands near New Guinea,

genus

of endemic

in Polhemus & Polhemus

major geological provinces

species

a single widely distributed species, M.

depicted

only

flavonotatus

or to individual terranes within these

Esaki, occurs within the northern

(

1998),

provinces

Fig. 4-7).

Three

as outlined

& Davies (1987)

zone of Miocene terranes on New Guinea

by Pigram

proper

(Fig. 5).

This

that the

pattern strongly suggests

(

of Metrobatini are confined to overall distribution of the

has been influ-

genus

genera

of New Guinea and

islands that

enced

the tectonics of the Solomons Arc (com-

portions

were derived from

adjacent

of the Miocene -

4, 16), and the concept ad-

supports

portions

Pliocene Solomons Arc as defined

pare Fig.

vanced

ranes

tion even after such arc

into a larger land mass.

2.) Andersenella is

Kroenke

Polhemus (1996) that species on arc ter-

may

retain their insular

of distribu-

are

intergrated

(

1984, 1996):

.) Metrobatopsis

Solomons, through the Bismarcks, and along the

northern coastal of New Guinea,

patterns

fragments

is distributed from the

confined to the

ranges

occupying

apparently

. Distributionof

in the New Guinea

region.

Figure

Peninsula

Metrobatopsisspecies

accreted ter- basin

(Fig. 4, 20).

The terranes of the

Papuan

(Fig. 4), complex

rane that was sutured to the remainder of New

Guinea sometime between the Eocene and the

Miocene. The exact tectonic

Peninsula is still a matter of debate; the circum-

scribed distribution of Andersenella in modem

deeper

mountains in the latter area are of Eocene but

age,

north of

volcanics

sequence

Tertiary

lying

and linked to a later

of the

them

be of Miocene

of arc accretion

history

Papuan

may

episode

field

age

(P.T. Freeport

Indonesia

case, the

entirely

geologists, pers. comm.).

current distribution of the

any

falls

New Guinea

mass was

island until

the

that this land

supports

separate

relatively

hypothesis

genus

from the remainder of the within the zone of accreted terranes in the north of

recent

time,

the island, and

mediated.

to have been

geological

suturing hypothesis

appears

tectonically

the Miocene

thereby favoring

(

see further discussion in Polhemus & Polhemus

998).

.) Metrobatoides is known from the Torricelli

None of the above three genera are known from

the portion of New Guinea formed from sections

of the Australian continental craton, lying south of

Mountains of northern coastal New Guinea, an the island's central mountain chain. Instead, three

area

to the Torricelli terrane of other

of Metrobatini have

corresponding

genera

into this

ranges extending

Davies (1987) which was accreted to the

island in the Miocene, and from the northern face

Pigram

region:

.) Stygiobates occupies

from Morotai in the Northern

tion

distribu-

elongate

of the central

in the Mamberamo River

ranges

extending

. Distributionof

in the New Guinea

Stygiobatesspecies

region.

Figure

Moluccas, south through Waigeo and the

therein), followed by dispersal of the genus east-

the ward

cratonic New Guinea south of the

Vogelkop Peninsula,

southern New Guinea

as the Lakekamu River basin of

and then eastward

along

foreland as far mountains

through

this fusion event. Con-

premontane

following

possible

New

it is

that evolved on

Papua

to be com-

versely,

Stygiobates

Guinea

4, 6). This

the Halmahera Arc, which

north of New

the Miocene

(

Fig.

absent from the accreted terranes of

New Guinea's northern coast,

genus appears

lay

Guinea from the

pletely

Miocene

Oligocene through

14-16), since one

Morotai, a current element in this arc (Baker &

Malaihollo 1996), while another is endemic to the

is endemic to

species

age along

that the

(Fig.

and

areas

indicating

were not influenced

accretion of terranes

Arc, a notable conclusion, since the western termi-

nus of this arc has not been well defined

Vogelkop

Raja Ampat

the Miocene to Pliocene

belonging

to the Solomons

island

which is also

Waigeo/Gam

complex,

hypothesized to contain terranes derived from this

arc (Charlton et al. 1991);

this

hypothe-

geologi-

following

on the

sis,

the arrival of the

genus

Vogelkop

ter-

cally.

The current distribution of

rane would be a more recent event. This latter sce-

Stygiobates suggests

that it

have instead evolved on the

nario is also

the fact that a clear mor-

may

microcontinent, which was fused to the main body

of New Guinea in the Miocene or Pliocene

Vogelkop

suggested by

phocline exists in terms of male abdominal modi-

fications, with the least

early

Polhemus 1998 and references

complex morphologies

on Morotai and while

(

Polhemus

occurring

species

Waigeo,

. Distributionof Iobates speciesin New Guinea.

Figure

(and therefore

and

2), both

Stygiobates (Fig.

strongly

progressively

sumably

occur in

complex

derived)

pre-

morphologies

Peninsula

ing Metrobatopsis

of which show

arc-linked distributions,

also have evolved on arc

recently

found in the

does

that it

species

and southern New Guinea

Vogelkop

It is also

suggest

may

north of New Guinea and then been sub-

proper.

and its sister

signif-

on the

systems

icant that

into the biota of the modem

Stygiobates

cladogram, Metrobatopsis,

exclusive

in the northern coastal terranes of

genus

sequently integrated

island.

occupy mutually

3.) Ciliometra occurs

New Guinea,

throughout

lowland river basins both north and

ranges

New Guinea

(

Fig.

occupied allopatric

cent arc that formed

4),

indicating

that

they may

adja-

the southern

occupying

have

distributions within

south of the New Guinea central

drainage

divide,

on the island of to the

Waigeo

and is also

present

Sea northwest of the

systems

along

boundaries of the Caroline and

Peninsula. Unlike

Philippine

Vogelkop

and Iobates, this

plates (Fig. 14-16).

Stygiobates

the

genus

Peninsula, with different endemic

also occurs on

.) Iobates displays quadripartite

distribution,

confined to the north-

of New Guinea, another, I.

Papuan

species occurring

some endemic Ciliometra

species

with one

on the north and south sides.

species, affinis,

ern coastal

ranges

Although

show congruence with arc terranes on Waigeo and

in the

Peninsula, the broad dis-

salawati, in the Vogelkop area, a third, I. somare,

the southern fore-

widespread through

premontane

Papuan

tribution of

generally

on lowland rivers

land of New Guinea as far east as the Kikori River

basin, and a fourth, I. ivimka, in the southern

in this

species

in mainland New Guinea, which flow through

7). basins between mountain formed

genus

Peninsula east of the Aure

Papuan

The

Scarp (Fig.

into

separate species

lying

accreted terranes, makes it difficult to determine

how tectonic events have influenced their dis-

ranges

of this

segregation

genus

in the northern, southern, and

and

Vogelkop

may

In the

Peninsula sections of the island is a

tributional

Papuan

pattern

typical

aquatic

history.

2) this

phylogenetic analysis

sister to

seen in

other

many

groups

(Fig.

genus

consistently

fishes (Allen 1991), and con- Metrobatoides, which is confined

to the

organisms, including

entirely

limited information about the role of tecton-

northern zone of accreted terranes, and this

veys

coup-

ics, if any, in structuring the distribution of this led with localized speciation of Ciliometra in the

However the

of Iobates in the

to the clade contain-

Peninsula

that the latter

genus

genus.

cladogram

position

group

Papuan

have had an arc-mediated distribution as well.

may

suggests

as the sister

It is notable that two of the three

of tive of an intial arc-based

with

subsequent

genera

south of the central

pattern

southward onto the Australian craton

Metrobatini

occurring

lobates and Ciliometra, are also

ranges,

in the

dispersal

several

arc collision.

present

northern section of New Guinea, while none of the

genera following

the current distributional and

summary,

evidence is

(Fig. 8)

that the Metrobatini diversified into a

phy-

hypoth-

found in the north of the island

in the south. In addition, the one

with the

remaining genera

are also

logenetic

esis

compatable

present

that occurs in southern New Guinea but is

of local

on one or more

genera

early

genus

not found in the north,

complex

Tertiary

and that these

the biota of New Guinea following a series of tec-

has

arc

north of New

Guinea,

Stygiobates,

Peninsula, the

represen-

systems lying

tatives in the

were later

into

Vogelkop

Raja Ampat

genera

incorporated

Islands, and the Northern Moluccas. This strongly

that the

Metrobatini occurred on arc

diversification

of tonic collisions that occurred between the Eocene

suggests

major

that formed

of the Caroline and

and the Pliocene.

systems

the southern

along

Philippine

margins

and

Sea

collided with

eventually

plates

Naboandelini

northern New Guinea

followed

(

Fig. 14-16),

into the

of certain

subsequent dispersal

genera

Only

single genus

Naboandelini, Calypto-

is also

southern section of the island after the

occurs on New Guinea. This

episodes

bates,

genus

the

arc terrane accretion. This is view is further

known to occur on

Australia,

the Andaman

Borneo, Islands,

and Sri

sup-

repre-

Moluccas,

but

the fact that the

Australian

ported by

only

Lanka,

of the island

sentative of the Metrobatini, Rheumatometra, is has so far not been collected in

any

east or southeast of New Guinea.

not a basal clade sister

to the South

arcs

group

American Metrobates, but is instead in a more ter-

minal subclade

that it was

lying

it is

that the distribution of this

Although

is still

incompletely documented,

likely

(

Fig. 2), indicating

genus

due to the

which are eas-

it is still clear

derived from ancestors on New Guinea or small size of its constituent

species

antecedent land masses. This weakens the case for

an initial vicariance of Gondwana metrobatine lin-

overlooked

most

collectors,

ily

that its

extends

range

ward from New

primarily

in contrast to the

south and west-

in the Mesozoic, and

the

eages

suggests

since a tectonic mech-

possibili-

Guinea,

which occur from New Guinea

north and eastward. In

ty sweepstakes dispersal,

anism which metrobatines could have colo-

Metrobatini,

pri-

all three

nized the arcs that formed

marily

addition,

so far known from New

mountains,

of the island. These lines of

the southern mar-

along

of the Caroline and Pacific

species

Calyptobates

Guinea occur south of the central

in the

gins

plates

early

while

evident.

Australia is not immedi-

the cratonic

Tertiary

bypassing

portions

evidence all

ately

suggest

genus

and that it was not

that

Calyptobates

of the area

discussed

Gondwanan

Australia,

derived from the subtree for north of New Guinea that later contributed accret-

that reached New Guinea from

summary

relationships

8, which

above is

on the arcs

present

provided

Fig.

represents

taxon-area

cladogram

the Metrobatini, in which the tectonic units dis-

cussed herein have been substituted for terminal

ed terranes to the island.

level taxa. This

illustrates once

in the tree is

generic

again

analysis

taxon

Stenobatini

that

distributed

only single

deep

on the Australian craton,

exclusively

The Stenobatini are a

predominantly

genera

the coasts of New

Melanesian

in this tribe

Guinea,

and

Halmahera,

while all

confined to arc

taxa have distributions either

Four of the five known

remaining

group.

and their accreted rem-

are

the

systems

present along

remaining genus

nants, or to such systems plus the New Guinea

occurs in

nearby

of the Australian craton, exclusive of with

as one moves

any

direction. The con-

widely varying patterns

portion

diversity dropping rapidly

Australia

this

of area

from Melanesia in

proper. Although

pattern

the

away

stituent

does not

that

show

relationships

preclude

Paleotropical

possibility

Metrobatini were

genera

basal

their overall distributions:

Thetibates is found

lineages

on the extreme northern

of the

in the

Solomons,

only

the coast of the

present

margin

1.)

the

Australian craton which later became New Guinea

and

Bismarcks,

to Adelbert-Finisterre terrane in northeastern New

along

and

onto the arcs

dispersed subsequently

the north and northeast, we feel it is more

lying

sugges-

Guinea

(Fig. 9).

Polhemus & Polhemus

(1998)

. Taxon-areasubtreefor Metrobatiniderivedfrom the largertaxon-areacladogramshownin Fig. 3. Areas

Figure

tectonicunits in the NewGuinearegionas definedby Polhemus& Polhemus(1998)and herein (see Fig.

represent

4-16).The notation 'Craton (Australia)'indicates

the Australiancontinentalcraton exclusiveof the portioncom-

southernNew Guinea;the notation 'Craton (New Guinea)' indicatesthe portion of southernNew Guinea

prising

derivedfromthe Australiancontinentalcratonbut exclusiveof the currentAustraliancontinentsouth of the Torres

Strait.

noted that the distribution

of Thetibates is

4.) Stenobates is widespread, being recorded

from Singapore, Celebes, the southern Philippines

(Palawan, Mindanao), the northern Moluccas,

New Guinea and the Raja Ampat Islands, and

northern Australia (Fig. 11). It is unknown from

pattern

terranes recently

derived from the Miocene Solomons Arc (Kro-

enke 1984), and

that the

with

congruent

geological

suggested

genus might

have evolved in isolation on this arc system prior

to accretion of the arc's western sector into the the Greater Sunda Islands, the Lesser Sunda

modem island of New Guinea.

Islands, the northeastern coast of New Guinea

Cenderawasih

the Bismarcks, or the

.) Pseudohalobates occupies roughly

rectan-

Mindanao on the north,

the Minahasa Peninsula of Celebes on the west,

beyond

Bay,

bounded

Solomons. This distribution indicates that the

genus never occupied the Miocene Solomons Arc,

since its current distribution is completely

gular range

and the

Peninsula of New Guinea on the

Vogelkop

9). Since it occurs on

east

different

with that of Thetibates, which occupies

(Fig.

many

and

differing ages

allopatric

terranes derived from that arc system (compare

1 and 16). Stenobates instead shows its

highest species diversity in the Vogelkop region

island arc

systems

trajecto-

ries, any linkage between its distribution and geo-

Fig. 9,

events is difficult to assess, and suggests

logical

that this

have

12), suggesting that it evolved on the

(Fig.

genus may

relatively high dispersal

abilities.

Vogelkop microcontinent or on arc systems such

as the Halmahera Arc lying immediately to the

.) Rheumatometroides occupies

east-west distribution

elongate,

the center of the north and west. The absence of Stenobates on

through

and into Melanesia. It is re-

Borneo is odd given that its distribution surrounds

Malay Archipelago

corded from

and

southern

adjacent

this island, and we predict that future collecting

Singapore

peninsular Malaysia, Borneo, Celebes,

Guinea and the

Islands, the Bis- Rheumatometroides, this genus is also unknown to

New

will

it to be

there. Like

prove

present

Raja Ampat

marcks, the Solomons, and northern Australia

It is unknown from Sumatra, Java, the

date from the

islands of the Sunda Arc.

5.) Stenobatopsis is confined to the northern

Moluccas, being known from Bacan and

(

Fig. 10).

Lesser Sunda Islands, the Moluccas or the

is much

Halmahera

9). Both are of a Miocene

parts

Philippines. Species diversity

along

higher

(Fig.

arc that formed at the southeastern comer of the

also

the coasts of New Guinea than in any other

area, a pattern of distribution suggesting that the Philippine Sea Plate, although Bacan

appears

have evolved in the New Guinea

to contain a small continental fragment (Mt.

Sabela) derived from the Vogelkop microcontinent

via westward translation along the Sorong Fault

(Malaihollo & Hall, 1996). The position of Steno-

batopsis as the most basal member of the Steno-

genus may

and later

region

westward into the central Malay

spread

reinforced

the

appar-

Archipelago, hypothesis

ent absence of this

islands of the

any

species

Sunda Arc east of Sumatra.

. Distributionsof three

of Stenobatiniin the New Guinearegion.

genera

Figure

batini

it and the

on arc

2) further

remaining

systems

of the Caroline and

the

that

Pseudohalobates inobonto. It

two distinctive subareas:

la.) Bomberai Peninsula and Raja Ampat

Islands, which contain the endemic species

to contain

appears

(Fig.

supports

hypotheses

members of the tribe evolved

that formed

the southern

along

Philippine

incorporated

Sea

margins

remants of which are now

Moluccas and New Guinea.

plates,

into the Rheumatometroides sele and

Stenobates fakfak.

Peninsula and the Cenderwasih

such a

lb.)

Following

the broader distributions now dis-

Vogelkop

islands of Biak and

which contain the

hypothesis,

played by

result of

west

following

isolated arc

Within the

distributions

Stenobatini are indicative of several

Bay

within the tribe are the endemic

Yapen,

Rheumatometroides wabon and

certain

genera

subsequent dispersal

tectonic

species

to the south and Stenobates

kamojo.

Within this area of endemism, the distributions

of Stenobatini are near the

convergence

formerly

systems.

regional

of individual

particularly complex

Peninsula

that this has been a zone of

context of New Guinea, the western

of the

tip

indicating

Vogelkop

(Fig.

12),

secondary

in the

areas

species

contact for a number of

general

previously separate

of marine endemism

13):

stenobatine faunas. Salawati, for exam-

(

Fig.

.) Vogelkop Peninsula and associated islands -

this area is defined the distribution of another Stenobates

species, kamojo,

regional

ple, supports the endemic Stenobates kasim, has

shared with

broadly

0. Distributionof Rheumatonzetroide si n the New Guinea

The

distributionof this

genus

Figure

extendswestwardoff the

region.

peninsularMalaysia

larger

and

to includeCelebes,Borneo,

map

Singapore.

the

Peninsula, Biak and

and

eastward into the Coral Sea, and westward

ranges

Vogelkop

Yapen,

shares Rheumatometroides sele with the

Peninsula, the Bomberai Peninsula, Misool,

as far as Misool Island.

Vogelkop

3.) Northeastern New Guinea, Bismarcks, and

Gam and Batanta. Misool,

contrast, is Solomons - this area is defined

of the same distribution of Rheumatometro-

the distribu-

Waigeo,

part

tions of Rheumatometroides browni, Thetibates

ides sele noted above, but shares its

Steno-

serena, and Thetibates matawa. It

the remnants of the Miocene Solomons Arc

single

S. australicus, with southern New

corresponds

bates

species,

Guinea and northeastern Australia.

system

as defined

Kroenke

the

(1984), including

.) Torres Strait - this area is defined by Adelbert-Finisterre terrane of northeastern New

Rheumatometroides kikori, Rheumatometro ides

Guinea. Taxa

not known from

mouth of the

of this area's

typical assemblage

are

carpentaria and Stenobates australicus. It basical-

northern New Guinea west of the

River.

Sepik

to the warm, shallow

ly corresponds

tal seas of the Australian craton,

epicontinen-

the

including

coasts of southern New Guinea and northern

Australia

13). One member of the

Summary

(Fig.

species

of this area, S. australicus,

To summarize the above discussion, we

assemblage typical

hypothe-

1.Distributionof Stenobatesin the New Guinea

The

distributionof this

extendswest-

genus

Figure

ward off the

region.

2.)

larger

map Singapore.

size the

following

Trepobatinae present

histories for the three tribes of

The Naboandelini are

in New

confined

represented

in the New Guinea

Guinea

region:

by single genus, Calyptobates,

.) The Metrobatini have one component of to the cratonic portion of the island south of the

genera (Metrobatopsis, Metrobatoides,

ella) that on the basis of cladistic and biogeo-

Andersen-

central mountains. This

Australia, suggesting a vicariance pattern across

the Torres Strait.

3.) The Stenobatini are similar to the Metro-

batini in

is shared with

genus

evidence

to have evolved

clearly appear

graphic

on terranes north and east of New Guinea,

partic-

those linked to the Solomons Arc. Another

one (Thetibates) whose

ularly

exhibiting

genus

to have evolved on distribution mirrors that of the Solomons Arc ter-

genus (Stygiobates) appears

terranes north and west of New Guinea linked to ranes, another

either the current Halmahera Arc or the

microcontinent. Several other

confined to the Hal-

widespread genera

(Stenobatopsis)

mahera Arc, and three more

Vogelkop

(Ciliometra,

(Pseudohalobates,

Stenobates, Rheumatometro-

ides) with numerous locally endemic species that

but no clear

provide intriguing regional patterns

genera

lobates) are widespread in the island and, though

endemic

are not

tectonic

possessing regionally

informative

distributions.

species,

concerning

indication of

distributions.

tectonic control in their

overriding

unequivocally

controls on

past

2.Distributionsof Stenohates

on New Guinea and

islands.Note the

nearby complexjuxtaposition

Figure

species

rangesin the area surroundingthe VogelkopPeninsula.

species

World checklist of

somare J. & D. Polhemus, 1993:256. Indonesia (Irian

NewGuinea

Trepobatinae

Jaya),Papua

MetrobatesUhler, 1871: 108.

TREPOBATINAE Matsuda, 1960: 330.

Subfamily

Uhler.

Metrobates

Type-species,

Type-genusTrepobates

hesperiusUhler,

871.

Tribe METROBATINIJ. & D. Polhemus, 1993: 244. = Tr-epobatopsisChampion, 1898: 157. Type-species,

MetrobatesUhler.

Type-genus

denticornis

Trepobatopsis

1898.

Champion,

Syn. by

Andersenella J. & D. Polhemus, 1993: 247.

Anderson,1932b:300.

Type-

Andersenella binotata J. & D. Polhemus, t aeternalis Scudder, 1890: 353.

FlorissantBeds,Colorado

fossil,

species

Oligocene

993.

binotata J. & D. Polhemus, 1993: 248. Papua New alacris Drake, 1955:130.U. S. (La.,Tx.)

Guinea

Nieser, 1993:23. Trinidad

amblydonti

anomalusanomalus

1948:123.U. S. (Fl.)

Hussey,

CiliometraJ. & D. Polhemus, 1993:249.

anomalus comatipesHussey & Herring, 1949: 169.U.

S. (Fl.)

Type-species

CiliometrakiungaJ. & D. Polhemus,1993.

femorata J.

D. Polhemus, 1993: 252.

Papua

New artus Anderson, 1932a:56. Mexico,U. S. (N. M., Tx.)

curracis Drake& Roze, 1954:228.Venezuela

Guinea

hirsuta

kiunga

1996:59.

New Guinea

Papua

denticornis

1898: 158.

[Trepobatopsis].

Lansbury,

Papua

(Champion),

Mesoamerica,Mexico,U. S. (Az.,N. M., Tx.)

945:180.Bolivia,Peru

J. & D. Polhemus,1993:251.

NewGuinea

minajerwiJ. & D. Polhemus,2000:296.Indonesia(Irian fugientis Drake& Harris,

hesperiu.rhesperiusUhler,

1871:108.U. S. (east)

Jaya)

996:55.

NewGuinea

Papua

(Ashmead),1897:56.

[Halobatopsis].

Syn. by

priori Lansbury,

Papua

= beginii

J. & D. Polhemus,1993:252.

NewGuinea

Torre-Bueno,1911 b 2: 43.

sepik

setosaJ. & D. Polhemus,2000:292.

NewGuinea hesperius

1949: 168.U.S.

1949:167.U. S.

Hussey Herring,

depilatusHussey Herring,

J. & D. Polhemus,2000: 295. Indonesia(Irian

(Fl.)

waigeo

ocalensis

Jaya)

hesperius

(Fl.)

Iobates J. & D. Polhemus,1993:253.

Me- laetus Drake, 1954:50. Brazil

Type-species

trobatopsisaffinisEsaki,

926.

laudatus Drake & Harris, 1937: 357. Costa Rica,

Panama,PuertoRico

affinis (Esaki), 1926:

IrianJaya), NorthernPapuaNewGuinea.

ivimka J. & D. Polhemus, 2002: 287. South Central

New Guinea

46.

[Metrobatopsis].

Indonesia

(

plaumanniplaumanni Hungerford,1951:72. Argentina

(north),Brazil

plaumanni genikos Nieser, 1993: 23. Brazil (Minas

Papua

salawati J. & D. Polhemus, 1993:255. Indonesia(Irian

SalawatiIsl.)

Gerais)

porcus Anderson,1932a:59. Belize,Honduras,Mexico,

Jaya,

Nicaragua

Queensland)

Drake & Harris, 1932b:88.

Syn. by'

Drake & kamoroJ. & D. Polhemus,2000: 308. Indonesia(Irian

Jaya)

spissus

Harris, 1937:357.

tumidusAnderson, 1932a:57. Cuba, Haiti,Jamaica

kopiJ. & D.Polhemus,2000:306.Indonesia(Irian

Jaya)

cubanusDrake& Harris, 1932b:86.

Harris, 1937:358.

Drake& minimus J. & D. Polhemus, 1994: 340. Australia

(NorthernTerritory)

Syn.by

trux trux (TorreBueno), 1921:274.

S. (west)

U. rubidus J. & D. Polhemus, 1994:342. Australia

Queensland)

[Trepobatopsis].

(north

trux

1953:179.U. S. (Ca.,Or.)

samarinda J. & D. Polhemus, 1994: 344. Indonesia

(Borneo;East Kalimantan,Sumba)

simplex J. & D. Polhemus, 1994: 345. Papua New

Guinea

infuscatusUsinger,

vigilis Drake, 1958: 100. Argentina (north), Brazil

MinasGerais,SantaCatarina)

(

Metrobatoides J. & D. Polhemus, 1993: 259.

Type-

Metrobatoides

J. & D. Polhemus, Hynesionella Poisson, 1949: 83. Type-species

HynesionellaaethiopicaPoisson, 1949.

species

genitalis

993.

bifurcatusJ. &D. Polhemus,2002:284.Indonesia(Irian aethiopicaPoisson,

949:83.

Ethiopia

1955: 130. [Naboandelus]. South

Jaya)

capensis (Poisson),

Africa

= omercooperiHungerford& Matsuda, 1959a:37. Syn.

J. & D. Polhemus, 1993: 261.

New

Papua

genitalis

Guinea

J. & D. Polhemus,1994:357.

MetrobatopsisEsaki,

926: 144.

Type-species,

Metro- cobbeniLinnavuori,1975:46. CentralAfricanRepublic

karatara J. Polhemus,1998:191.SouthAfrica

batopsis flavonotatu, Ef saki, 1926.

browniJ. & D. Polhemus,1993:266. SolomonIslands

flavonotatus Esaki, 1926: 146. Indonesia (Irian Jaya),

schuhiJ. Polhemus,1998:192.SouthAfrica

Naboandelus Distant, 1910a: 151.

NaboandelussignatusDistant, 1910.

Type-species

New Guinea

Papua

insularisJ. & D. Polhemus,1993:266. Indonesia(Irian africanusHoberlandt,1950:14.Angola

BiakIsl.)

bandama Linnavuori,1981:18.Ivory Coast

Jaya;

lannae J. & D. Polhemus,1993:269. SolomonIslands

mussau J. & D. Polhemus, 1993: 269. Bismarck

bergevini bergevini Bergroth, 1911: 256. Egypt, Iraq,

Israel (Palestine, Sinai), Lybia, Palestine, Sudan,

Zaire

Syria,

Archipelago(MussauIsl.)

solomonensis

& Matsuda, 1959b: 35. bergeviniboukeanusLinnavuori,1975:45. IvoryCoast,

Hungerford

SolomonIslands

west andcentralAfrica

orientalisZettel, 1996:250.

bergevini Iraq

bergevini

popovi

Brown, 1951: 227. Israel, Jordan,

Rheumatometra

Kirkaldy,

Rheumatometra

1902: 280.

Type-species,

Kirkaldy,1902,monobasic.

SaudiArabia,Yemen

philarete

Andersen & Weir, 1998: 519. Eastern

Linnavuori,1971:360. Sudan

dimorpha

bergevinipygmaea

Australia

borneensis J. & D. Polhemus, 1994: 350. Indonesia

902:280.EasternAustralia.

(Borneo;East Kalimantan)

dentipesLinnavuori,1981:21. IvoryCoast

philarete Kirkaldy,

StygiobatesJ. & D. Polhemus,

993:272.

Type-species hynesiPoisson, 1949:83. Ethiopia,Somalia

iriana J. & D. Polhemus, 1993.

Polhemus,

1994:351.West

Malaysia

Stygiobates

johorensis

J. & D.

iriana J. & D. Polhemus, 1993: 273. Indonesia (Irian

Jaya)

madacascariensisPoisson, 1952:31.

Madagascar

nionodiPoisson,1928:73. Cameroon,Ghana,

Nigeria

iwekaJ.&D.Polhemus ,2 000:297.Indonesia(Irian

1939:311.

Ethiopia,Somalia,

Jaya) patrizii patrizii Mancini,

moratai J. & D. Polhemus, 1993: 275. Indonesia

Moratai,Moluccas)

mubiJ. & D. Polhemus,2000:300.

Sudan

patrizii

(

danae Linnavuori,1981:20.

Coast

Ivory

NewGuinea

1971:361. Sudan

Papua

J. & D. Polhemus, 2000: 301. Indonesia (Irian

patrizii migransLinnavuori,

rajana

Jaya)

signatus Distant, 1910a: 152. India, Myanmar,

SriLanka,Thailand,Vietnam

Nepal,

taprobanicusJ. & D. Polhemus,1994:351. Sri Lanka

TalaudiaJ. & D. Polhemus,2002:289.

An-

Type-species

umuahia

Linnavuori,

1981:20.

Nigeria

dersenella nilsi Chen& Nieser,2000.

nilsi (Chen & Nieser), 2000:70. [Andersenella]

Indonesia(Talaud

witteiPoisson, 1950:72. Zaire

Archipelago)

TribeSTENOBATIN IJ . & D.

Polhemus,

1993:245.

StenobatesEsaki.

Type-genus,

TribeNABOANDELIN IJ . & D. Polhemus,1993:244.

Type-genusNaboandelusDistant.

Pseudohalobates J. & D. Polhemus, 1996:292. Type-

PseudohalobatesinobontoJ. & D. Polhemus,

species,

1996.

species Calyptobatesjourama J. & D. Polhemus, inobonto J. & D. Polhemus, 1996: 294. Indonesia

994.

J. & D. Polhemus, 1994: 334.

Calyptobates

Type-

Irian Kri

amboina J. & D. Polhemus, 1994: 337. Indonesia

Ambon)

(

Celebes, Moluccas,

Jaya: Biak,

Misool,

Yapen,Vogelkop) ,P hilippines(Mindanao)

(

andaman J. & D. Polhemus, 1994: 338. Andaman Rheumatometroides Hungerford & Matsuda, 1958a:

Islands

203. Type-species Rheumatometroides browni

Hungerford& Matsuda, 1958.

jourama J. & D. Polhemus,

1994:339. Australia(north

3.Areasof marineendemismin theNewGuinearegionasdefined thedistributionsof Stenobatini

by species

Figure

forfurtherdiscussionsee text).

(

browni

& Matsuda, 1958a:204. Solomon biroi (Esaki), 1926:119.[Stenometra].

Singapore

Hungerford

Islands,PNG

fakfak J. & D. Polhemus, 1996: 317. Indonesia (Irian

carpentaria (J.

D. Polhemus), 1991:5. [Stenobates].

Jaya;

kamajo

Jaya;

southwest,Batanta,

Waigeo)

N. Australia

J. & D. Polhemus, 1996:319. Indonesia(Irian

Chen & Nieser, 1992:160.Celebes

Biak,Salawatiand

drepanephoros

nearby,Yapen)

insularis insularis (Polhemus &

1982: 225. kasim J. & D. Polhemus, 1996: 320. Indonesia (Irian

Jaya)

Cheng),

[

Stenobates].Singapore,W.Malaysia

insularis

J. & D. Polhemus, 1996:303. Sabah

labuha J. & D. Polhemus,1996:324.Indonesia(Bacan,

Halmahera)

papar

kikoriJ. & D. Polhemus,1996:305.

New Guinea

Papua

seleJ. & D. Polhemus,1996:307. Indonesia(Irian

langoban J. & D. Polhemus, 1996: 326.

(Palawan)

Jaya;

Philippines

Batanta,BomberaiPeninsula,Gam,Misool,Salawati,

makraitos(Chen& Nieser), 1992: 159. [Rheumatomet-

Waigeo)

wabonJ. & D. Polhemus, 1996:311.Indonesia(Biak)

roides].Indonesia(Celebes)

J. & D. Polhemus, 1996: 333. Indonesia

sangihe

(SangiheArchipelago)

J. & D.

StenobatesEsaki, 1927: 184.

Stenometra

Type-species,

biroiEsaki, 1926,monobasic;nom.n. forStenometra

Esaki, 1926.

1996:335.

Philippines

zamboanga

Mindanao)

Polhemus,

(

Stenometra

1926: 118.

Type-species

Esaki,

Stenometrabiroi Esaki, 1926,monobasic.Preocc.

StenometraClark, 1909,in Echinodermata.

Stenobatopsis J. & D. Polhemus, 1996: 338.

Type-

species, Stenobatopsis stygius J. & D. Polhemus,

1996.

australicus J. & D. Polhemus, 1991:3. Australia(north

Queensland), Indonesia (Irian Jaya; south central, stygiusJ. &

D. Polhemus,1996:340.Indonesia(Halma-

Misool),PapuaNewGuinea

hera,MamboaIsl.)

ThetibatesJ. & D. Polhemus, 1996:341.

rozeboomi

1937: 358.

[Telmat-

Type-species,

Rheumatometroide ss erena Lansbury,1992.

matawa

1989: 67.

[Rheumatometroides].

(Drake

Harris),

ometra].Colombia,CostaRica,Ecuador,Panama

(

Lansbury),

TrepobatesUhler,

894: 213.

Halobates

Type-species,

SolomonIslands

pictus Herrich-Schaeffer ,1 847;nom.n. forStephania

aqaaqa Lansbury, 1989: 65. Syn. by J. & D.

Polhemus,1996:343.

B. White, 1883.

- Stephania

Halobates

B. White, 1883: 79.

Type-species,

pictus

serena

1992: 1.

[Rheumatometroides].

(

Lansbury),

Herrich-Schaeffer, 1847.

New Guinea

Papua

Preoccupied by Stephania Leseur, 1813, in

Coelenterata.

TribeTREPOBATIN IM atsuda, 1960:330.

TrepobatesUhler.

Type-genus

beckiDrake & Harris, 1932c:120.Mexico,U. S.

Ca.,N. M.)

(Az.,

Cryptobates Esaki, 1929a: 412. Type-species,Gerris

carri Kittle, 1982: 157. Belize, Cuba, Guatemala,

Honduras,Jamaica,Mexico,U. S. (Tx.)

raja Distant, 1910.

johorensisJ. & D. Polhemus,1995:102.WestMalaysia, floridensis

Drake& Harris, 1928c:273. U. S. (Al., Ga.,

Thailand

Fl., Ms.)

obscurus

1967: 253. Brunei,

inermis Esaki, 1926: 140. Canada (southeast), U. S.

(east)

Miyamoto,

Malaysia

(

Sabah)

raja (Distant), 1910a:142. [Gerris]. India (Kerala),Sri knighti

Drake& Harris, 1928a:28. U. S. (central)

Lanka

panamensis Drake & Hottes, 1952: 35. Mesoamerica,

Colombia,Ecuador,Venezuela

rufus J. & D. Polhemus, 1995: 105. West Malaysia,

pictus (Herrich-Schaeffer),

847: 111 [Halobates].

Singapore

Canada(southeast,U. S. (east)

J. Polhemus, 1991: 82.

polhemiKittle, 1982:161.Mexico(central,western)

Type-species,

Cryptobatoides

CryptobatoidesbrunneusJ. Polhemus,1991.

brunneusJ. Polhemus,1991:84. Brazil

subnitidusEsaki, 1926: 141. Canada

(central,east, N. M., Tx.)

(southeast),

1953:111.

Kittle,

Chapman, Syn. by

U. S.

citatus Drake&

1991: 945.

GnomobatesJ. & D. Polhemus,1995:107.

Type-species

CryptobateskuitertiHungerford& Matsuda, 1958.

kuiterti

1958c: 246.

taylori (Kirkaldy), 1899: 28 [Kallistometra].

Matsuda),

Mesoamerica,South

America,U. (Tx.),

WestIndies

(

Hungerford

[

Cryptobates].India,Myanmar[Burtna],Thailand

= comitialisDrake& Harris, 1928b:7. Syn.by Drake&

Harris, 1932a:112.

Halobatopsis Bianchi, 1896: 70. Type-species,Halo-

Drake & Harris, 1928a: 27. U. S.

trepidus

(Az.),

bates

platensis Berg,1879.

Mesoamerica,

Colombia,Ecuador,

Venezuela

Nieser& Lanede Melo,1999:98. Brazil

chrysocastanis

Drake &

Hottes,

1951: 141. Mexico

(south-

vasquezae

west)

delectusDrake& Harris,1941:237. Brazil

platensis (Berg), 1879: 24. [Halobates]. Argentina,

Brazil

Matsuda,

1958b: 125.

Trepobatoides Hungerford

spiniventris Drake & Harris, 1936: 107. Argentina,

Type-speciesTrepobatoidesboliviensisHungerford&

Brazil

Matsuda,1958.

boliviensis

Peru

& Matsuda,1958b:125.Bolivia,

Hungerford

Ovatametra

1942: 136.

Halo-

Type-species,

Kenaga,

Drake& Harris,1935.

batopsisparvulus

amnica Drake, 1957:112.Panama

bella Drake, 1957:111. Bolivia

of the

Bibliography

The

subfamily Trepobatinae

fusca Kenaga, 1942:139.Brazil,Guyana

lists all

works known

to the authors information

containing important

following bibliography

gualeguayBachmann,1966a:87.Argentina,Brazil

minima

1942:137.Brazil

Kenaga,

obesa

1942:138.Brazil

concerning the subfamily Trepobatinae. Incidental

Kenaga,

parvula (Drake

Brazil

thaumanaDrake, 1959:107.Brazil

Harris), 1935: 3.

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the northern

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