phylogenetic evaluation of the tropical camptotypus genus-group (hymenoptera: ichneumonidae), with a...

This article was downloaded by: [University of Saskatchewan Library]On: 05 October 2012, At: 19:27Publisher: Taylor & FrancisInforma Ltd Registered in England and Wales Registered Number: 1072954Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK

Journal of Natural HistoryPublication details, including instructions for authors andsubscription information:http://www.tandfonline.com/loi/tnah20

Phylogenetic evaluation of thetropical Camptotypus genus-group(Hymenoptera: Ichneumonidae), witha key to the world generaIlari E. Sääksjärvi a , Ian D. Gauld b & Jukka Salo aa The Zoological Museum, Centre for Biodiversity, Departmentof Biology, FIN-20014, University of Turku, Finland E-mail:b The Natural History Museum of London, Cromwell Road,London SW7 5BD, UK E-mail:

Version of record first published: 19 May 2010.

To cite this article: Ilari E. Sääksjärvi, Ian D. Gauld & Jukka Salo (2004): Phylogeneticevaluation of the tropical Camptotypus genus-group (Hymenoptera: Ichneumonidae), with a keyto the world genera , Journal of Natural History, 38:21, 2759-2778

To link to this article: http://dx.doi.org/10.1080/0022293032000140949

PLEASE SCROLL DOWN FOR ARTICLE

Full terms and conditions of use: http://www.tandfonline.com/page/terms-and-conditions

This article may be used for research, teaching, and private study purposes. Anysubstantial or systematic reproduction, redistribution, reselling, loan, sub-licensing,systematic supply, or distribution in any form to anyone is expressly forbidden.

The publisher does not give any warranty express or implied or make anyrepresentation that the contents will be complete or accurate or up to date. Theaccuracy of any instructions, formulae, and drug doses should be independentlyverified with primary sources. The publisher shall not be liable for any loss, actions,claims, proceedings, demand, or costs or damages whatsoever or howsoever causedarising directly or indirectly in connection with or arising out of the use of thismaterial.

http://www.tandfonline.com/loi/tnah20

http://dx.doi.org/10.1080/0022293032000140949

http://www.tandfonline.com/page/terms-and-conditions

http://www.tandfonline.com/page/terms-and-conditions

Phylogenetic evaluation of the tropical Camptotypus genus-group(Hymenoptera: Ichneumonidae), with a key to the world genera

ILARI E. SAAKSJARVI{, IAN D. GAULD{ and JUKKA SALO{

{The Zoological Museum, Centre for Biodiversity,

Department of Biology, FIN-20014, University of Turku, Finland;

e-mail: [email protected]

{The Natural History Museum of London, Cromwell Road,

London SW7 5BD, UK; e-mail: [email protected]

(Accepted 10 May 2003)

The phylogeny of the disjunct pantropical Camptotypus genus-group (Hymenoptera,Ichneumonidae, Pimplinae) was reconstructed using 40 morphological charac-ters.The monophyly of many established genera (Parvipimpla, Clydonium,Odontopimpla) was found to be supported, but Hemipimpla was found to nestwithin Camptotypus and has thus been treated as a junior synonym of it (syn. n.). Agroup of undescribed Neotropical species were found to form a separatemonophyletic group, herein referred to as Genus A. Species of the Neotropicalgenus Zonopimpla formed a basal paraphyletic grade which could not satisfactorilybe resolved into discrete monophyletic units. Rather than subsuming all the morederived genera within one large group, or attempting to erect numerous new andprobably ephemeral genera (for isolated species of Zonopimpla), we propose theretention of this grade-group as a separate genus at present. The host utilizationpatterns of the genus-group, and the biogeography of the genus-group are discussed.The group is hypothesized as having originated and radiated in the Neotropics, asectoparasitoids of weakly concealed lepidopterous hosts. One lineage, Clydonium,has become specialized at attacking hosts in woody galls where it utilizescoleopterous as well as lepidopterous hosts. The Old World genera (Camptotypusand Parvipimpla) comprise a monophyletic group and are suggested to haveoriginated from a single dispersal event across the widening Atlantic. No evidencefor either a southern or northern circum-polar dispersal was observed in thisentirely tropical genus-group. One lineage in the Afrotropical region has switchedfrom attacking lepidopterous hosts, to attacking pre-pupal and pupal polistinevespids.

KEYWORDS: Biogeography, classification, parasitoid, host utilization, pantropical.

Introduction

The Camptotypus genus-group is one of the five generic groups currently

recognized in the tribe Ephialtini within the ichneumonid subfamily Pimplinae

Journal of Natural HistoryISSN 0022-2933 print/ISSN 1464-5262 online # 2004 Taylor & Francis Ltd

http://www.tandf.co.uk/journalsDOI: 10.1080/0022293032000140949

JOURNAL OF NATURAL HISTORY, 2004, 38, 2759–2778

Dow

nloa

ded

by [

Uni

vers

ity o

f Sa

skat

chew

an L

ibra

ry]

at 1

9:27

05

Oct

ober

201

2

(Gauld, 1991; Gauld et al., 2002). The species of this group are usually brightly

coloured and often strikingly patterned ichneumonids, and they are found

throughout all tropical regions of the world, but not elsewhere. Most are

encountered in lowland wet forests, where they can be found flying in sunlit

clearings, often in the company of similarly coloured and generally much more

numerous braconids. As many of these braconids might be distasteful (Buckingham

and Sharkey, 1988) and models for mimicry complexes (Quicke, 1986; Gauld,

1991), we suspect that these pimplines are Batesian mimics of them. Other species

apparently mimic aggressive vespids, such as the Costa Rican Clydonium cabrerai

Gauld that with black body and white-tipped black wings closely resembles

Parachartergus species (Gauld, 1991).The monophyly of the Camptotypus genus-group has long been recognized

(Townes and Townes, 1960) and is attested by the marked reduction of the occipital

carina, which at the most is only present on the lower part of the head. Current

phylogenetic investigation places the Camptotypus genus-group very close to the

Sericopimpla and Ephialtes genus-groups, which together form a derived group with

respect to the Alophosternum and Pseudopimpla genus-groups (Gauld et al., 2002)

thus: Pseudopimpla g-g z (Alophosternum g-g z (Camptotypus g-g z Ephialtes g-g

z Sericopimpla g-g)). As currently defined, the Camptotypus genus-group

comprises five genera: Camptotypus, Parvipimpla, Clydonium, Odontopimpla and

Zonopimpla (including Cenodontis), whilst a sixth taxon, Hemipimpla, has

sometimes been recognized as a separate genus (e.g. by Gupta and Tikar, 1978),

and at other times treated as a subgenus of Camptotypus (e.g. by Baltazar, 1961;

Townes, 1969). Biogeographically, the Camptotypus genus-group is of particular

interest because all species are entirely tropical. Camptotypus, Hemipimpla and

Parvipimpla occur in the Old World and northern Australia, and the remaining

three genera are restricted to tropical America. This is a particularly unusual

distribution pattern for the Ephialtini, as the other genus-groups are well-

represented in temperate habitats, and in cases where they are species-rich in the

tropics, highly derived and separate lineages occur in the Old and New World

(Gauld et al., 2002). For example, in the Ephialtes genus-group, Anastelgis is

restricted to the New World, whilst the structurally similar Xanthophenax only

occurs in the Old World, where both apparently parasitize concealed lepidopterous

borers (Townes, 1969). Similarly, within the Polysphincta group of genera,

Acrotaphus and Hymenoepimecis are endemic New World genera, whereas a

quite separate lineage, Eriostethus, fills a similar niche in the Old World, attacking

orb-web spiders (Townes, 1969; Gauld, 1984b, 1991).

In recent years, as exhaustive collecting has been undertaken in tropical

America, it has become apparent that a plethora of undescribed species of the

Camptotypus genus-group occur in the region. Some of these cannot be placed to

genera, and many other superficially dissimilar species seem to belong in

Zonopimpla, making the monophyly of the genus as it is presently delineated,

improbable. The aims of the present study are to resolve the phylogeny of the

species-groups within the Camptotypus genus-group so as to provide a basis for

more rigorously defining the genera, and to discover the phylogenetic relationships

between Old and New World taxa.

2760 I. E. Saaksjarvi et al.

Dow

nloa

ded

by [

Uni

vers

ity o

f Sa

skat

chew

an L

ibra

ry]

at 1

9:27

05

Oct

ober

201

2

Materials and terminology

The specimens examined in this study were borrowed from or deposited in the

following collections: The Natural History Museum, London, UK (BMNH);

Department of Primary Industries, Indooroopilly, Queensland, Australia; Instituto

Nacional de Biodiversidad, Santo Domingo, Costa Rica; The Zoological Museum,

University of Turku, Turku, Finland. The Peruvian specimens of Genus A (see

below), currently deposited at the University of Turku, will be deposited in the

Museum of Natural History of San Marcos, Lima, Peru. The morphological

terminology is mostly that of Gauld (1991).

Taxa

For the purpose of this study a representative of the basal pimpline lineage,

Delomerista mandibularis (Gravenhorst) [collected in Finland] and relatively

unspecialized member of the Ephialtini, Iseropus stercorator (Fabricius) [England]

were used as out-groups (see Wahl and Gauld, 1998 and Gauld et al., 2002 for

preliminary analysis of Pimplinae). Representatives of all described genera in the

Camptotypus genus-group have been included. Clydonium was represented by three

species: C. analuisae Gauld, Ugalde Gomez and Hanson [Costa Rica], C. cabrerai

Gauld [Costa Rica] and C. sp. 1 [Peru] (in BMNH) and the small genus

Odontopimpla by two: O. fasciata (Brulle) [Peru] and O. maxima (Morley) [Peru].

The very large and morphologically heterogeneous genus Zonopimpla was

represented by 12 species that encompass the morphological variation present in

the taxon: Z. aquilari Gauld [Costa Rica], Z. atriceps (Cresson) [El Salvador], Z.

aurae Gauld [Costa Rica], Z. bermudezi Gauld [Costa Rica], Z. carolinae Gauld

[Costa Rica], Z. fasciata (Townes) (the type-species of Cenodontis) [Brazil], Z.

mirandai Gauld [Costa Rica], Z. snortumi Gauld, Ugalde Gomez and Hanson

[Costa Rica], Z. tenorioi Gauld, Ugalde Gomez and Hanson [Costa Rica], Z. sp. 2

[Peru] (in BMNH), Z. sp. 4 [Peru] (in BMNH) and Z. sp. 10 [Peru] (in BMNH).

The Indo-Australian genus Camptotypus was represented by four species: C.

olynthius (Cameron) [India], C. pseudostigmaticus Gupta and Tikar [Solomon Isles],

C. rugosus (DeGeer) [Sarawak] and C. sellatus Kriechbaumer [Australia], and the

small Australian genus, Parvipimpla, was represented by the only described species,

P. petita Gauld [Australia]. Hemipimpla, an Afrotropical group of species, was

represented by three species: H. flavicaput Morley [Uganda], H. sp. 2 [Angola] (in

BMNH) and H. superba (Szepligeti) [Malawi]. In addition we included five

Neotropical species, Genus A 1–5 [Brazil, Colombia, Guyana and Peru] that could

not be placed into any of the described genera.

Characters

The following set of 40 characters was used in the analysis. The presumed

plesiomorphic condition is denoted by a ‘0’ and derived states by other integers.

The data matrix for character distributions is shown in table 1.

(1) Occipital carina: (0) complete; (1) dorsally absent, ventrally present joining

hypostomal carina; (2) completely absent. We treat this as being a

transformation series, 1.0p1.1p1.2.

(2) Mediodorsal notch of occiput: (0) present, more or less in the form of an

inverted V; (1) absent entirely; (2) with lateral swellings present but broadly

separated.

Phylogenetic evaluation of the tropical Camptotypus 2761

Dow

nloa

ded

by [

Uni

vers

ity o

f Sa

skat

chew

an L

ibra

ry]

at 1

9:27

05

Oct

ober

201

2

(3) Anterior margin of pronotum, dorsal view: (0) centrally reflexed; (1)

transverse.

(4) Anterior part of pronotum, lateral view: (0) anteriorly horizontal, shelf-like,

forming a more or less right angle with the hind margin (figures 1, 2); (1)

more or less vertical in entirety (figure 3).

(5) Epomia: (0) complete; (1) present centrally as small, sharp remnant; (2)

absent or its position indicated by an obscure, low rounded swelling.

(6) Mesoscutal sculpture: (0) uniformly closely punctate, the punctures

separated from each other by about their own diameters; (1) uniformly

sparsely punctate, the punctures separated by about three or more times

their own diameters; (2) smooth and without punctures over most of surface

(7) Notauli: (0) distinct on anterior 0.5 of mesoscutum; (1) shallow, present only

anteriorly as a vestige.

(8) Lateral margin of mesoscutum in front of scutellum: (0) narrowly carinate; (1)

rounded, without a carina or ridge.

(9) Epicnemial carina: (0) dorsal end reaching far above level of lower corner

Table 1. Data matrix.

1111111111 2222222222 3333333333 4123456789 0123456789 0123456789 0123456789 0

Delomerista mandibularis 000000000 0000200000 0010000000 0000100000 0Iseropus stercorator 000000000 0000001000 0011103010 0000000000 1Odontopimpla maxima 211022011 1022202110 0011103031 2000100300 0Odontopimpla fasciata 211022011 1021202111 0011120031 2000100300 0Clydonium sp. 1 110022011 0021202110 0001123031 2001020222 0Clydonium analuisae 100022012 0031202100 0101103032 0001000222 0Clydonium cabrerai 100022011 0011202110 0001123030 2001020222 0Genus A sp. 1 110022011 2112202100 0101123031 0000021100 0Genus A sp. 2 110022011 0132202100 0111121031 0000021100 0Genus A sp. 3 110022011 0102202100 0001103031 0000020100 0Genus A sp. 4 110022012 0101202100 0001113031 0000020110 0Genus A sp. 5 110012011 0131202100 0001113031 0000010100 0Zonopimpla tenorioi 200012010 3002202100 0111120031 0000000000 0Zonopimpla snortumi 200012110 0010202100 0111113032 1000000000 0Zonopimpla aquilari 200012010 0010202100 0011103010 0000000100 0Zonopimpla aurae 200012011 0000202100 0001103030 0000010100 0Zonopimpla atriceps 200012011 0002202100 0001123032 0000000000 0Zonopimpla bermudezi 200011010 0000202100 0011102032 0000000000 0Zonopimpla carolinae 200011010 0000202100 0011103032 0000000000 0Zonopimpla mirandai 200012011 0000202100 0001122030 1000000001 0Zonopimpla sp. 2 200011010 0010202100 0111113032 0000000000 0Zonopimpla sp. 4 200012010 0000202100 0001103032 0000000000 0Zonopimpla sp. 10 200012011 0010202100 0001103032 0000000100 0Camptotypus rugosus 111112110 0042112100 0011120111 0110000000 2Camptotypus pseudostigmatus 111112010 0042102100 1011120010 0010000000 1Camptotypus olynthius 111122010 0042112100 0011121121 0010000000 1Camptotypus sellatus 111122010 0042102100 0111120021 0010000000 1Hemipimpla flavicaput 120122010 0003002100 0011100010 0010000000 1Hemipimpla sp. 2 120122010 0003002100 0001100010 0010000000 1Hemipimpla superba 120122010 0003002100 0011100010 0020000000 1Cenodontis fasciatus 200012012 0030202100 0011113031 0000000001 0Parvivipimpla petita 211112012 0032102100 0211113031 0020000000 1


Dow

nloa

ded

by [

Uni

vers

ity o

f Sa

skat

chew

an L

ibra

ry]

at 1

9:27

05

Oct

ober

201

2

of pronotum; (1) dorsal end more or less level with lower corner of

pronotum; (2) laterally entirely absent.

(10) Medioventral part of epicnemial carina: (0) not raised into a conspicuous

flange-like structure; (1) expanded into a flange-like structure: (2) swollen,

tooth-like; (3) forming a pair of flatten lobes.

(11) Epicnemium: (0) without a distinct secondary carina, at most with a minute

tooth present on upper edge; (1) with a distinct secondary carina present

near lower corner of pronotum (figure 13).

(12) Submetapleural carina: (0) complete, anteriorly very sharp and posteriorly

bifurcate with an inner and outer branch; (1) outer branch absent; (2)

present anteriorly, but both branches absent; (3) entirely absent; (4) inner

branch absent.

(13) Pleural carina: (0) complete; (1) represented by sharp groove; (2) more or

FIGS. 1–3. Anterior part of pronotum, lateral view. (1) Zonopimpla atriceps. (2) Clydoniumsp. 1. (3) Camptotypus flaviceps. FIG. 4. Clydonium sp. 1: antero-median part oftergite II, dorsal view.


Dow

nloa

ded

by [

Uni

vers

ity o

f Sa

skat

chew

an L

ibra

ry]

at 1

9:27

05

Oct

ober

201

2

less absent; (3) complete posteriorly, but absent in front of propodeal

spiracle.

(14) Sculpture of propodeum: (0) laterally conspicuously punctate; (1) laterally

sparsely punctate; (2) smooth to quite smooth.

(15) Propodeal spiracle: (0) oval; (1) elongate, more than 2.5 times as long as broad.

(16) Propodeum dorsally with carinae: (0) more or less complete; (1) reduced

with only lateromedian longitudinal carina present anteriorly; (2) carinae

absent dorsally. We treat this as being a transformation series,

16.0p16.1p16.2.

(17) Insertion of metasoma with respect to insertion of hind coxa: (0) clearly

separated by a strongly sclerotized cuticular band; (1) with a very thin,

almost unsclerotized division, or without an apparent division.

(18) Structure of front coxae: (0) simple; (1) posteriorly horizontal, shelf-like.

FIG. 5. Camptotypus flaviceps: antero-median part of tergite II, dorsal view.FIGS. 6–8. Apico-lateral corners of tergites III–V. (6) Clydonium sp. 1. (7) Camptotypusflaviceps. (8) Camptotypus sp.


Dow

nloa

ded

by [

Uni

vers

ity o

f Sa

skat

chew

an L

ibra

ry]

at 1

9:27

05

Oct

ober

201

2

(19) Hind femur: (0) simple; (1) with a ventral tooth.(20) Pterostigma: (0) normal; (1) conspicuously enlarged.(21) Fore wing with areolet: (0) subrhombic; (1) subtriangular; (2) absent, i.e.

with vein 3rs-m missing.(22) Hind wing with cu-a: (0) short, about 0.3 or less of length of abscissa of Cu1

between M and cu-a ; (1) longer than 0.5 times length of abscissa of Cu1

between M and cu-a.(23) Hind tarsal claw of female with base: (0) simple; (1) lobate.(24) Hind tarsal claw of female: (0) with a small internal membranous vesicle;

(1) without a vesicle.(25) Tergite I with lateromedian carinae: (0) weak, only reaching back to level of

spiracle; (1) absent; (2) very strong, reaching to or past centre of tergite.(26) Tergite I with lateral longitudinal carinae: (0) strong, reaching to or past

centre of tergite, above spiracle; (1) strong, reaching to or past centre of tergite,

below spiracle; (2) weak, reaching back to level of spiracle; (3) absent.

FIGS. 9–12. Apex of lower valve of ovipositor. (9) Zonopimpla atriceps. (10) Odonto-pimpla fasciata. (11) Clydonium sp. 1. (12) Camptotypus flaviceps.


Dow

nloa

ded

by [

Uni

vers

ity o

f Sa

skat

chew

an L

ibra

ry]

at 1

9:27

05

Oct

ober

201

2

(27) Sternite II: (0) without prominent flap-like fold; (1) with prominent flap-

like fold.

(28) Tergite II with raised part: (0) coriaceous; (1) closely regularly punctate; (2)

with few scattered punctures centrally; (3) impunctate.

(29) Tergite II with impressed part: (0) closely regularly punctate; (1) with few

scattered punctures centrally; (2) impunctate.

(30) Antero-median part of tergite II: (0) simple (figure 5); (1) with a broad,

shallow indentation; (2) with a strong, deep indentation (figure 4).

(31) Tergites II–V: (0) without a longitudinal carina; (1) with a discontinuous

median longitudinal carina.

(32) Apico-lateral corners of tergites III–V: (0) slightly rounded (figure 6); (1) a

sharp right-angle (figure 7); (2) with a small acute tooth (figure 8). We treat

this as being a transformation series, 32.0p32.1p32.2.

FIG. 13. Genus A sp. 1: epicnemium. FIGS. 14, 15. Apex of lower valve of ovipositor.(14) Genus A sp. 1. (15) Genus A sp. 2.


Dow

nloa

ded

by [

Uni

vers

ity o

f Sa

skat

chew

an L

ibra

ry]

at 1

9:27

05

Oct

ober

201

2

(33) Apical part of ovipositor: (0) subcylindrical; (1) strongly laterally

compressed.(34) Form of upper valve of ovipositor tip: (0) slender; (1) stout and apically

abruptly rounded.(35) Apex of upper valve of ovipositor: (0) simple (figures 9, 10); (1) with low

serrations (figure 11); (2) with clearly discernible teeth (figures 14, 15).(36) Extreme apex of upper valve of ovipositor: (0) simple; (1) with a small apical

swelling (figures 14, 15).(37) Relationship of upper and lower valves of ovipositor: (0) lower valve not

enclosing the upper, and with upper part of the central teeth of lower valve

exposed (figures 9, 12); (1) lower valve with clear lobe partially enclosing

upper valve; (2) lower valve apically more or less entirely enclosing upper

valve (figure 11); (3) upper valve partly enclosing the lower, so dorsal

margins of central teeth of lower valve are obscured (figure 10).(38) Lower valve of ovipositor tip just basal to apex: (0) without scabrous area(s);

(1) with a single long scabrous area proximal to basal tooth; (2) with two

scabrous areas.(39) Apex of lower valve of ovipositor: (0) with distinctly oblique, moderately

interspaced teeth (figures 9, 10); (1) without discernible teeth; (2) with quite

closely interspaced, vertical teeth (figure 11).(40) Surface of ovipositor sheath: (0) with fine short hairs; (1) with long coarse

hairs; (2) with exceptionally dense thick pubescence. We treat this as being

a transformation series, 40.0p40.1p40.2.

Analytical technique

Data were manipulated using Winclada Ver. 0.9.99 (Nixon, 2000). Data were

analysed searching for minimum-length trees under equal character weights using

the NONA program (version 1.6) of Goloboff (1997). NONA was run with all

multistate characters treated as non-additive, except for #1, 16, 32 and 40, all of

which are obviously transformation series. The heuristic search command mult*

(random addition sequence with tree bisection and reconnection) was implemented

with 80 replications, with 12 trees held during each replication (hold/); the resulting

trees were then subjected to branch swapping using tree bisection/reconnection

(max*). The concave function selected was 8. NONA’s successive weighting option

(swt.run) was used to reduce the number of trees.

Results of analysis

Analysis of the data set yielded 92 cladograms with a length of 136 steps, a

consistency index of 0.50, and a retention index of 0.73. Successive weighting

reduced this to three very similar trees differing only in the relationships of the

species of Zonopimpla. One of these is illustrated in figure 16. In the alternative trees

either Z carolinae z Z. bermudezi forms an unresolved trichotomy with Z. snortumi

z Z. sp. 2 and Z. sp. 4z all higher taxa, or Z. carolinae and Z. bermudezi form an

unresolved tetrachotomy with the other taxa.

It can be seen that the Old World species form a strongly supported

monophyletic group (figure 16) which is defined by the following synapomorphies:

anterior part of pronotum in lateral view centrally subvertical (character 4-1);

propodeum laterally sparsely punctate (character 14-1); apico-lateral corners of

tergites III–V angulate (character 32-1). Within this group Parvipimpla is


Dow

nloa

ded

by [

Uni

vers

ity o

f Sa

skat

chew

an L

ibra

ry]

at 1

9:27

05

Oct

ober

201

2

monophyletic, supported by the autapomorphy fore wing with areolet absent

(character 21-2), but Camptotypus is paraphyletic with respect to the Hemipimpla

species. The monophyletic Neotropical genus Odontopimpla is the sister lineage to

the Old World genera.

FIG. 16. Arrangement of species of Camptotypus genus-group in NONA cladogram, and thecharacters supporting this hierarchy. Black circles are uniquely derived characters,white circles indicate putative parallelisms or reversals.


Dow

nloa

ded

by [

Uni

vers

ity o

f Sa

skat

chew

an L

ibra

ry]

at 1

9:27

05

Oct

ober

201

2

The unplaced group of species from South America form a distinct

monophyletic group, which is supported by the presence of a distinct secondary

carina on the epicnemium near lower corner of pronotum (character 11-1). This

group may be the sister lineage to Odontopimpla z Old World genera. The species

of Clydonium form a strongly supported monophyletic group defined by the

following synapomorphies: ovipositor subapically strongly compressed (character

33-1); lower valve of ovipositor apically entirely enclosing upper valve (character

37-2); lower valve of ovipositor tip just basal to apex with two scabrous areas

(character 38-2); apex of lower valve of ovipositor with closely interspaced vertical

teeth (character 39-2). It is the sister lineage to the unplaced group of species from

South America z Odontopimpla z Old World genera.

The major problem remaining concerns the species of Zonopimpla which

together comprise a paraphyletic grade showing little robust structure. Type-species

of Cenodontis (Zonopimpla fasciata) nests amongst the other Zonopimpla species,

supporting the synonymy advocated by Gauld (1991).

Classification of the Camptotypus genus-group

The results of this analysis present a problem which is quite commonly

encountered within the Ichneumonidae, and indeed the Hymenoptera as a whole

(Belshaw et al., 2001). That is, within a clearly definable higher group a number of

discrete monophyletic groups of species have been accorded generic status, whilst

one genus remains paraphyletic. This is known to be the case within the Pimplini

(Ichneumonidae, Pimplinae) where Theronia is a basal paraphyletic assemblage, in

the Glyptini (Ichneumonidae, Banchinae) where Glypta is paraphyletic, and within

the Atrophini (Ichneumonidae, Banchinae) where Lissonota is paraphyletic (Gauld,

1991, 2002). A similar situation almost certainly exists in most other groups of

ichneumonids, and probably most families of Hymenoptera.

Establishing a formal classification for groups such as the Camptotypus genus-

group presents a real problem. Although we accept that strictly monophyletic

groups are desirable, the only way this could be achieved in the present case is

either to include the entire group of species into a single genus, or to erect

numerous new genera for various species of Zonopimpla. The disadvantages of

either approach are obvious. First, creating numerous new genera for species of

Zonopimpla at present is unlikely to produce nomenclatural stability, because the

complex includes a large number of rarely encountered species that are difficult to

group in any meaningful way. Obvious species-groups have not been found and

any proposal of new genera would almost certainly have to be revised repeatedly

because numerous additional species are being discovered all the time as the

fauna of the Andean–Amazonian interface becomes better known. Many of

these species are likely to comprise additional separate lineages that ultimately

would require new genera or blur the distinction between newly postulated ones.

Second, amalgamating all the currently recognized genera would result in a loss

of biological meaning. For example, the very distinctive form of the ovipositor of

the Clydonium species is an adaptation for penetrating tough plant tissue to oviposit

on to hosts living deeply concealed inside the plant (Gauld, 1991). Placing

Clydonium into the same genus together with other species of the Camptotypus

genus-group would make the use and prediction of biological information more

difficult.

We propose a compromise classification. Hemipimpla is here treated as a


Dow

nloa

ded

by [

Uni

vers

ity o

f Sa

skat

chew

an L

ibra

ry]

at 1

9:27

05

Oct

ober

201

2

synonym of Camptotypus eliminating one incidence of paraphyly and recognizing a

single distinctive genus for most Old World species. Parvipimpla, Odontopimpla and

Clydonium are treated as distinct genera, as is the group of species (Genus A sp. 1–

sp. 5) from South America. This new genus has been lately described formally in

another paper that also delineates all the species (Saaksjarvi et al., 2003). We

continue to recognize Zonopimpla as a plesiomorphic basal genus, at least until such

time as the overwhelming majority of species are described, when a more exhaustive

analysis might reveal clear groups of species that could be accorded separate generic

status. Details of the classification proposed, together with a key to genera and

diagnoses of genera is given in Appendices 1 and 2.

The biology and patterns of host-utilization by the Camptotypus genus-group

Little is known about the biology of the Camptotypus genus-group. From

information available (Sonan, 1930; Seyrig, 1932; Beeson and Chatterjee, 1935;

Costa Lima, 1945; LePelley, 1954; Keeping and Crewe, 1983; Brooks and Wahl,

1987; Gauld, 1991) it is apparent that the species are idiobiont ectoparasitoids of

the pupae and prepupae of holometabolous insects. Because many of the more

basal taxa in closely related groups utilize hosts concealed in plant tissue—including

Scambus species in the Ephialtes genus-group (Fitton et al., 1988), Pseudopimpla in

the Pseudopimpla genus-group (Bruzzese, 1982; Matsumoto and Saigusa, 1999),

Alophosternum in the Alophosternum genus-group (Cushman, 1933) and Perithous

in the Delomeristini (Danks, 1971)—we suspect that the group may have originated

attacking similar hosts. Although it is a less parsimonious assumption, the

alternative scenario that the ancestors of the group arose attacking cocooned hosts,

cannot be entirely discounted. This is because some basal taxa in related groups—

including Delomerista in the Delomeristini (Furniss and Dowden, 1941) and

Acropimpla and Iseropus in the Sericopimpla genus-group (Fitton et al., 1988)—only

attack such hosts. Both habits occur widely in the Camptotypus genus-group,

although species of one genus, Clydonium, are apparently specialists in attacking

hosts in rather woody galls (Costa Lima, 1945; Gauld, 1991).

The majority of host records for the group are from immature stages of

Lepidoptera. Species of Zonopimpla, Clydonium and Camptotypus all are known to

parasitize such hosts, an observation that suggests Lepidoptera may be the host

group that the Camptotypus lineage diversified using. However, as with many other

idiobiont ectoparasitoids, ordinal-level host switching has occurred. We speculate

that some species of Clydonium have switched from attacking lepidopterous larvae

in galls to attacking gall-forming curculionid larvae (Costa Lima, 1945), and a few

species of Afrotropical Camptotypus have probably switched from Lepidoptera to

attacking the prepupae or pupae of species of polistine Vespidae, such as

Belonogaster and Rhopalidia, in paper nests (Keeping and Crewe, 1983; Brooks and

Wahl, 1987).

The biogeography of the Camptotypus genus-groupWhen trying to understand the biogeographic relationships between Old and

New World species-groups of the Camptotypus genus-group, several things must be

taken into account:

(1) Without exception, all basal lineages (figure 16) are present only in the

Neotropical region and the generic and species richness of the group is


Dow

nloa

ded

by [

Uni

vers

ity o

f Sa

skat

chew

an L

ibra

ry]

at 1

9:27

05

Oct

ober

201

2

highest in New World tropics, where four genera are present (Genus A,

Clydonium, Odontopimpla and ‘Zonopimpla’), which together probably

include more than 100 species.(2) The entire modern Camptotypus genus-group is tropical. There is no

evidence any species has ever spread into adjacent subtropical areas, and

unlike many other predominantly tropical American Pimplinae (such as

Acrotaphus and Anastelgis), none are present in southern USA (vide Townes

and Townes, 1960).(3) The group has a disjunct (non-contiguous) distribution. Four genera are

restricted to the New World tropics, while two genera occur in the Old

World (Camptotypus and Parvipimpla), where they probably comprise fewer

than 50 species.(4) The Old World species form a strongly monophyletic group, and are derived

with respect to the Neotropical fauna (figure 16).

These data suggest that the group originated and primarily radiated in what is

today tropical America. Because the more basal groups of Pimplinae are

predominantly northern (Gauld et al., 2002), we speculate that the ancestor of

the Camptotypus group may have reached South America from North America

during the late Mesozoic or early Paleogenic, by dispersal across the putative

archipelagic connection (Coney, 1982). This Neotropical origin scenario poses the

question of how to explain the disjunct distribution of the group.

Although there is evidence that the family Ichneumonidae existed at the time of

the break-up of Gondwanaland (see discussion in Gauld and Wahl, 2002), we

doubt that the disjunct distribution pattern of the Camptotypus genus-group

resulted from a vicariance event, the separation of Africa from South America. If

such were the case, one might expect to find several lineages of the genus-group

present in both the Old World and the New—just as one finds many lineages of the

Labeninae (a putative southern vicariant group) present in both Australia and

South America (Gauld and Wahl, 2000). The most plausible and parsimonious

explanation of the phylogenetic pattern observed is that the Old World fauna

results from a single dispersal event from tropical America. But how was such

dispersal accomplished?

As nothing has been published on the phylogenetic differentiation rates of

Ichneumonidae, it is difficult to formally link the projected dispersal event of

the genus-group with the New World discrete faunal units (Marshall and Sempere,

1993) and the suggested dispersion routes. We consider the southern Paleogenic

dispersal (sensu Axelrod and Raven, 1978) of the Camptotypus genus-group to

be improbable because the group is absent from temperate South America as

well as from temperate Australia. The groups of ichneumonids that are believed

to have originated in and/or dispersed across Gondwanaland, such as the

Labeninae (Gauld and Wahl, 2000) and Eucerotinae (Gauld and Wahl, 2002),

have basal species present today in the temperate parts of South America and

Australasia. Furthermore, in the Australasian region, such taxa usually show a

drastic reduction in species richness westwards across Wallacea (Whitmore, 1981),

or are even restricted to the Australian tectonic plate (Gauld, 1984a; see also

Crisci et al., 1991 for many other examples of southern distributions). This is

not the case in Camptotypus. All basal taxa are tropical American species

(figure 16), and the genus-group is well-represented in the continental Old


Dow

nloa

ded

by [

Uni

vers

ity o

f Sa

skat

chew

an L

ibra

ry]

at 1

9:27

05

Oct

ober

201

2

World tropics but shows a marked decrease in species richness east of Wallace’s

line (Gupta and Tikar, 1978). Furthermore, the Paleogene–Neogene southern realm

has most probably always remained cold-temperate (Coetzee, 1993), without

evident tropical corridors providing an avenue for the dispersal of the Camptotypus

genus-group.

A Neogene northern dispersal route (via North America and across the Bering

Strait) also appears to be improbable for Camptotypus. Although there is a

considerable body of evidence showing that the climate of the circum-arctic region

was far warmer in the past than it is at present, and that at various times between

the middle Eocene and late Pliocene the region supported a rich plant and mammal

biota (McKenna, 1980; Hsu, 1983; Funder et al., 1985; Woodburne and Swisher,

1995), groups of ichneumonids that have putatively dispersed across this bridge are

mainly cool-tolerant taxa. Even amongst predominantly tropical groups which

putatively dispersed across Beringia (such as the Thyreodon genus-group), one

today finds a few species present as far as 50‡N in Asia and in North America

(Gauld, 1985). There are no modern representatives of the Camptotypus genus-

group in either the Eastern Palaearctic or in North America.

The most plausible scenario seems to be that the group dispersed from tropical

South America to Old World tropics via an archipelagic connection in the

spreading Atlantic. In this, the Camptotypus genus-group may resemble the paper

wasps (Polistinae) that also are absent from southern South America and are

considered to have dispersed via tropical South America–tropical Africa before the

full formation of the Mid-Atlantic Seaway (Carpenter, 1993). It is noteworthy that

Camptotypus is one of the few pimpline genera that is apparently good at dispersing

across water into archipelagos, as today one highly derived species (C.

pseudostigmaticus) is endemic to the Solomon Islands.

Appendix 1. Key to the genera of the Camptotypus genus-group

Using the key below it should be possible to place virtually all specimens of the

Camptotypus genus-group into one or other genus. However, it should be noted

that many of the most reliable characters for separating the genera are found in the

structure of the ovipositor. Whenever possible the identification of the males should

be checked by comparing the specimens to females.

1 Anterior part of pronotum in lateral view centrally vertical, not shelf-like (figure 3);Old World species . . . . . . . . . . . . . . . . . . . . 2

– Anterior part of pronotum in lateral view centrally horizontal, shelf-like (figures 1, 2);New World species . . . . . . . . . . . . . . . . . . . . 3

2 Occipital carina present ventrally, as a short vestige joining hypostomal carina; epicne-mial carina present; fore wing with 3rs-m present . . . . . . . Camptotypus

– Occipital carina entirely absent; epicnemial carina absent; fore wing with 3rs-mabsent . . . . . . . . . . . . . . . . . . . . Parvipimpla

3 Anterior margin of pronotum mediodorsally transverse or slightly reflexed but thenwithout a distinct backwardly direct point present; upper valve of ovipositor unusuallystout, towards the apex with its lower margin partly overhanging lower valve, so thatdorsal margins of central teeth of lower valve are obscured (figure 10) Odontopimpla

– Anterior margin of pronotum mediodorsally reflexed, forming a small backwardlydirected point; upper valve of ovipositor not unusually stout, never partly enclos-ing the lower, so that the dorsal margins of the central teeth of the lower valveare not occluded (figures 9, 11, 12) . . . . . . . . . . . . . . 4


Dow

nloa

ded

by [

Uni

vers

ity o

f Sa

skat

chew

an L

ibra

ry]

at 1

9:27

05

Oct

ober

201

2

4 Epicnemium with a distinct secondary carina (figure 13), and upper valve of ovipo-sitor not distinctly enclosed by lower valve (figures 14, 15) . . . . . Genus A

– Epicnemium without a distinct secondary carina, or if with trace of a secondary carina

then upper valve of ovipositor more or less entirely enclosed by lower valve

(figure 11) . . . . . . . . . . . . . . . . . . . . . . 5

5 Pronotum, in lateral view, with epomia present as small sharp remnant in centre;tergite II mediodorsally simple, without a sharp indentation (cf. figure 5); ovipositorapically subcylindrical, lower valve either not laterally expanded to enclose the uppervalve (figure 9) or with a small lobe partially enclosing upper valve Zonopimpla

– Pronotum, in lateral view, with epomia vestigial or absent; tergite II mediodor-

sally usually with a sharp indentation (figure 4); ovipositor conspicuously com-

pressed, the lower valve of ovipositor more or less entirely enclosing the upper valve

(figure 11) . . . . . . . . . . . . . . . . . . . . Clydonium

Appendix 2. Classification of the Camptotypus genus-group

Genus Camptotypus Kriechbaumer

Camptotypus Kriechbaumer, 1889: 311. Type-species: Camptotypus sellatus

Kriechbaumer, by subsequent designation, Viereck, 1914: 27.

Hemipimpla Saussure, 1892: 13. Type-species: Hemipimpla pulchripennis Saussaure,

by monotypy. Syn. n.

Erythropimpla Ashmead, 1900: 57. Type-species: Erythropimpla abbottii Ashmead

(~Pimpla olynthia Cameron), by original designation.Trichiothecus Cameron, 1903: 136. Type-species: Trichiothecus ruficeps Cameron

(~Ichneumon rugosus DeGeer), by monotypy.

Cosmiopimpla Cameron, 1905: 198. Type-species: Cosmiopimpla ferruginea

Cameron, by monotypy.

Pleurodontoplax Enderlein, 1914: 228. Type-species: Pleurodontoplax congoensis

Enderlein (~Pimpla vipioides Brulle), by original designation.

Phruropimpla Benoit, 1964: 387. Type-species: Hemipimpla flavicaput Morley, by

original designation.

Diagnosis. Camptotypus can be easily separated from the New World genera by

the form of the anterior part of pronotum which is in lateral view centrally more or

less vertical. In this feature it resembles the other Old World genus, Parvipimpla,

but unlike this genus all species of Camptotypus have 3rs-m present in the fore wing.Remarks. Camptotypus is a moderately large genus containing 47 described spe-

cies in the Indo-Australian and Afrotropical regions (Yu and Horstmann, 1997) and

a rather few undescribed species in museum collections. The Indo-Australian species

have been revised by Gupta and Tikar (1978), but there is no modern treatment of

the Afrotropical species. There are few reliable host records for species of this

genus, but what few there are suggests species attack cocooned pupae of Macrolepi-

doptera (e.g. Sonan, 1930; Beeson and Chatterjee, 1935; LePelley, 1954). One puta-

tively monophyletic Afrotropical group attack the prepupae or pupae of polistine

Vespidae (Keeping and Crewe, 1983; Brooks and Wahl, 1987).

Genus Clydonium Townes

Clydonium Townes in Townes and Townes, 1966: 324. Type-species: Hemipimpla

recta Morley, by original designation.


Dow

nloa

ded

by [

Uni

vers

ity o

f Sa

skat

chew

an L

ibra

ry]

at 1

9:27

05

Oct

ober

201

2

Diagnosis. Clydonium can be identified by the following set of characters: occi-

pital carina present ventrally, joining the hypostomal carina; epomia vestigial or

absent; ovipositor subapically strongly compressed; upper valve of ovipositor

with subapical serrations; lower valve of ovipositor apically entirely or more or

less enclosing upper valve; lower valve of ovipositor tip just basal to apex with

two scabrous areas; apex of lower valve of ovipositor with closely interspaced

vertical teeth; tergite II usually with a median indentation.Remarks. Clydonium is a rather small Neotropical genus comprising 10

described species (Yu and Horstmann, 1997; Gauld et al., 1998) and a similar

number of undescribed species in museums. Most species are very rarely col-

lected. Recorded hosts are woody gall-forming Thryididae and Curculionidae

(Costa Lima, 1945; Gauld, 1991).

Genus Odontopimpla Cameron

Odontopimpla Cameron, 1886: 272. Type-species: Pimpla pulcherrima Cresson

(~Pimpla fasciata Brulle), by monotypy.

Diagnosis. Odontopimpla is characterized by the following set of characters:

occipital carina entirely absent ventrally; epomia discernible as a vestige; upper

valve of ovipositor not compressed and without serrations; lower valve of ovipo-

sitor apically partly enclosed by the upper valve; lower valve of ovipositor with-

out scabrous areas; apex of lower valve of ovipositor with strong and widely

spaced oblique teeth; tergite II with a weak median indentation.Remarks. Odontopimpla is a very small Neotropical genus containing only two

described species, the type species and O. maxima (Morley). Nothing is known

about the biology of either.

Genus Parvipimpla Gauld

Parvipimpla Gauld, 1984a: 67. Type-species: Parvipimpla minuta Gauld, by original

designation.

Diagnosis. Parvipimpla is easily discernible from other genera of the Campto-

typus genus-group by the absence of the areolet in the fore wing.Remarks. This very small genus is represented only by one described species,

Parvipimpla minuta Gauld, occurring in tropical Australia. Nothing is known

about its biology.

Genus ‘Zonopimpla’ Ashmead

Zonopimpla Ashmead, 1900: 55. Type-species: Zonopimpla albicincta Ashmead, by

original designation.

Cenodontis Townes, 1969: 94. Type-species: Cenodontis fasciata Townes, by original

designation.

Diagnosis. Zonopimpla can be distinguished from other genera of the Campto-

typus genus-group by using the following set of characters: occipital carina

entirely absent ventrally; epomia distinct but short; upper valve of ovipositor

subcylindrical to weakly compressed and usually without serrations; lower valve

of ovipositor apically not or only partly enclosing the upper valve; lower valve


Dow

nloa

ded

by [

Uni

vers

ity o

f Sa

skat

chew

an L

ibra

ry]

at 1

9:27

05

Oct

ober

201

2

of ovipositor without scabrous areas; apex of lower valve of ovipositor with

weak; strong and oblique; or sparse and subvertical teeth; or with teeth absent;

tergite II without median indentation.Remarks. Zonopimpla is a large Neotropical genus containing about 30

described species (Yu and Horstmann, 1997; Gauld et al., 1998) and rather large

numbers of undescribed ones. Many of the species are rather infrequently col-

lected, and even in areas that have been intensively sampled, such as Costa

Rica, many new species have been found by additional collecting (compare

Gauld, 1991 and Gauld et al., 1998). We expect many additional species to be

discovered as the fauna of the Amazonian region is better sampled. Such an

incomplete knowledge of the diversity in this genus precludes us from attempting

to delineate monophyletic groups at present.

Very little is known about the biology of species of this group. One or two

species have been reared from cocooned Lepidopterous hosts, and one species is

known to be gregarious within a large Macrolepidopteran host (Gauld, 1991).

Genus A

Diagnosis. Genus A can be distinguished from other genera of the Camptotypus

genus-group by the following set of characters: occipital carina present ventrally as

a vestige, joining hypostomal carina; epicnemium with a distinct secondary carina

near lower corner of pronotum; ovipositor subapically subcylindrical; apex of upper

valve of ovipositor with low serrations or clearly discernible teeth.Remarks. The genus is restricted to the Amazonian region of South America,

and despite very intensive collecting, it has never been found further north, in

Costa Rica. Many new species found during recent studies made in northern

South America belong to this genus, and we now know of species from Brazil,

Ecuador, French Guyana and Peru. Our colleague, Edgard Palacio, has drawn

our attention to species in Colombia. Given the species richness of this genus,

and the fact that all included species were undescribed, the taxon and its compo-

nent species have been lately described formally in another paper (Saaksjarvi

et al., 2003).

Acknowledgements

We are most grateful to Sondra Ward and Reijo Jussila for their assistance in

preparing the figures. Part of the study was financed by the European

Commission’s Large Scale Facility Programme (travel grant to Natural History

Museum of London for I.E.S.) and Graduate School of Biological Interactions,

Ministry of Education, Finland (grant for I.E.S.). Ministry of Agriculture of Peru

provided the collecting and export permits for the Peruvian material. Samuli

Haataja, Andrew Polaszek, Leif Schulman and anonymous referee gave

constructive comments on the manuscript. Finally we would like to thank all

those Costa Rican and Peruvian parataxonomists, biologists and students for their

most valuable help during the study.

References

ASHMEAD, W. H., 1900, Classification of the ichneumon-flies, or the superfamilyIchneumonoidea, Proceedings of the United States National Museum, 23, 1–220.

AXELROD, D. I. and RAVEN, P. H., 1978, Late Cretaceous and Tertiary vegetation history of


Dow

nloa

ded

by [

Uni

vers

ity o

f Sa

skat

chew

an L

ibra

ry]

at 1

9:27

05

Oct

ober

201

2

Africa, in M. J. A. Werger (ed.) Biogeography and Ecology of Southern Africa (TheHague: W. Junk), pp. 77–130.

BALTAZAR, C. R., 1961, The Philippine Pimplini, Poemeniini, Rhyssini, and Xoridini(Hymenoptera, Ichneumonidae, Pimplinae), Monographs of the National Institute ofScience and Technology, 7, 1–130.

BEESON, C. F. C. and CHATTERJEE, S. N., 1935, On the biology of the Ichneumonidae(Hymenoptera), Indian Forest Records, 1, 151–168.

BELSHAW, R., LOPEZ-VAAMONDE, C., DEGERLI, N. and QUICKE, D. L. J., 2001, Paraphyletictaxa and taxonomic chaining: evaluating the classification of braconine wasps(Hymenoptera: Braconidae) using 28S D2-3rDNA sequences and morphologicalcharacters, Biological Journal of the Linnean Society, 73, 411–424.

BENOIT, P. L. G., 1964, Ichneumonidae africains du Naturhistorisches Museums de Vienne(Insecta-Hymenoptera), Annalen des Naturhistorischen Museums in Wien, 67, 377–388.

BROOKS, R. W. and WAHL, D. B., 1987, Biology and mature larva of Hemipimplapulchripennis (Saussure), a parasite of Rhopalidia (Hymenoptera: Ichneumonidae,Vespidae), Journal of the New York Entomological Society, 95, 547–552.

BRUZZESE, E., 1982, Observations on the biology of Pseudopimpla pygidiator Seyrig (Hym.,Ichneumonidae), a parasite of the blackberry stem-mining sawfly Hartigiaalbomaculatus (Stein) (Hym., Cephidae), Entomologist’s Monthly Magazine, 118,249–252.

BUCKINGHAM, G. R. and SHARKEY, M. J., 1988, Abdominal exocrine glands in Braconidae(Hymenoptera) in V. K. Gupta (ed.) Advances in Parasitic Hymenoptera Research(Leiden: Brill), pp. 199–242.

CAMERON, P., 1886, Biologia Centrali-Americana. Insecta. Hymenoptera (FamiliesTenthredinidae-Chrysididae), Vol. 1 (London), pp. 265–288.

CAMERON, P., 1903, Description of new genera and species of Hymenoptera taken by MrRobert Shelford at Sarawak, Borneo, Journal of the Straits Branch of the RoyalAsiatic Society, 39, 89–181.

CAMERON, P., 1905, On some new genera and species of Hymenoptera from CapeColony and Transvaal, Transactions of the South African Philosophical Society, 15,195–257.

CARPENTER, J. A., 1993, Biogeographic patterns in the Vespidae (Hymenoptera): two viewsof Africa and South America, in P. Goldblatt (ed.) Biological Relationships BetweenAfrica and South America (New Haven, CT and London: Yale University Press),pp. 139–155.

COETZEE, J. A., 1993, South American paleofloras in P. Goldblatt (ed.) BiologicalRelationships Between Africa and South America (New Haven, CT and London:Yale University Press), pp. 37–61.

CONEY, P. J., 1982, Plate tectonic constraints on the biogeography of Middle America andthe Caribbean region, Annals of the Missouri Botanical Garden, 69, 432–443.

COSTA LIMA, A. DA, 1945, Sobre dois insetos galicolas (Curculionidae—Cryptorhynchinae eIchneumonidae—Pimplinae), Boletim da Sociedade Brasileira de Agronomia, 8,189–192.

CRISCI, J. V., CIGLIANO, M. M., MORRONE, J. J. and ROIG-JUNENT, S., 1991, Historicalbiogeography of southern South America, Systematic Zoology, 40, 152–171.

CUSHMAN, R. A., 1933, Descriptions of new ichneumon-flies with taxonomic notes,Proceedings of the United States National Museum, 82(2955), 1–16.

DANKS, H. V., 1971, Biology of some stem-nesting aculeate Hymenoptera, Proceedings of theRoyal Entomological Society of London, 122, 323–399.

ENDERLEIN, G., 1914, Hymenoptera IV, Ichneumonidae in W. Michaelsen (ed.) Beitrage zurKenntnis der Land-und Susswasserfauna Deutsch-Sudwestafrikas, Band I (Hamburg),452 pp.

FITTON, M. G., SHAW, M. R. and GAULD, I. D., 1988, Pimpline ichneumon-flies Handbooksfor the Identification of British Insects, 7(1), 1–110.

FUNDER, S., ABRAHAMSEN, N., BENNIKE, O. and FEYLING-HANSSEN, R. W., 1985, ForestedArctic: evidence from North Greenland, Geology, 13, 542–546.

FURNISS, R. L. and DOWDEN, P. B., 1941, Western hemlock sawfly, Neodiprion tsugaeMiddleton, and its parasitoids in Oregon, Journal of Economic Entomology, 34, 46–52.


Dow

nloa

ded

by [

Uni

vers

ity o

f Sa

skat

chew

an L

ibra

ry]

at 1

9:27

05

Oct

ober

201

2

GAULD, I. D., 1984a, An Introduction to the Ichneumonidae of Australia (London: BritishMuseum (Natural History)), 413 pp.

GAULD, I. D., 1984b, The Pimplinae, Xoridinae, Acaenitinae and Lycorininae of Australia(Hymenoptera: Ichneumonidae), Bulletin of the British Museum (Natural History)(Entomology), 49, 235–339.

GAULD, I. D., 1985, The phylogeny, classification and evolution of parasitic wasps of thesubfamily Ophioninae (Ichneumonidae), Bulletin of the British Museum (NaturalHistory) (Entomology), 51, 61–185.

GAULD, I. D., 1991, The Ichneumonidae of Costa Rica, 1, Memoirs of the AmericanEntomological Institute, 47, 1–589.

GAULD, I. D. (ed.), 2002, The Ichneumonidae of Costa Rica, 4, Memoirs of the AmericanEntomological Institute, 66, 1–768.

GAULD, I. D. and WAHL, D., 2000, The Labeninae (Hymenoptera: Ichneumonidae): a studyin phylogenetic reconstruction and evolutionary biology, Zoological Journal of theLinnean Society, 129, 271–347.

GAULD, I. D. and WAHL, D., 2002, The Eucerotinae: a Gondwanan origin for acosmopolitan group of Ichneumonidae?, Journal of Natural History, 36,2229–2248.

GAULD, I. D., UGALDE GOMEZ, J. A. and HANSON, P., 1998, Guıa de los Pimplinae de CostaRica (Hymenoptera: Ichneumonidae) Revista de Biologia Tropical, 46(1), 1–189.

GAULD, I. D., WAHL, D. and BROAD, G. R., 2002, The suprageneric groups of the Pimplinae(Hymenoptera: Ichneumonidae): a cladistic re-evaluation and evolutionary biologicalstudy, Zoological Journal of the Linnean Society, 136, 421–485.

GOLOBOFF, P., 1997, NONA, version 1.6, MS-DOS program (San Miguel de Tucuman,Argentina: published by the author).

GUPTA, V. K. and TIKAR, D. T., 1978 (1976), Ichneumonologia Orientalis. Part 1. The tribePimplini Oriental Insects Monographs, 1, 1–302.

HSU, J., 1983, Late Cretaceous and Cenozoic vegetation in China, emphasizing theirconnections with North America, Annals of the Missouri Botanic Garden, 70,490–508.

KEEPING, M. G. and CREWE, R. M., 1983, Parasitoids, commensals and colony size in nestsof Belonogaster (Hymenoptera: Vespidae), Journal of the Entomological Society ofSouthern Africa, 46, 309–323.

KRIECHBAUMER, J., 1889, Nova genera et species Pimplidarum, Entomologische Nachrichten,15, 307–312.

LEPELLEY, R. H., 1954, Agricultural Insects of East Africa (Nairobi), 307 pp.MARSHALL, L. G. and SEMPERE, T., 1993, Evolution of the Neotropical Cenozoic land

mammal fauna and its geochronologic, stratigraphic, and tectonic context in P.Goldblatt (ed.) Biological Relationships Between Africa and South America (NewHaven, CT and London: Yale University Press), pp. 329–392.

MATSUMOTO, R. and SAIGUSA, T., 1999, Biology of Pseudopimpla glabripropodeum He andChen (Hymenoptera, Ichneumonidae, Pimplinae) with description of final instar larvaand a redescription of adults Entomological Science, 2, 467–471.

MCKENNA, M. C., 1980, Eocene paleolatitude, climate, and mammals of Ellesmere IslandPalaeogeography, Palaeoclimatology, Palaeoecology, 30, 349–362.

NIXON, K. C., 2000, Winclada, version 0.9.99, documentation and software, http://www.cladistics.com.

QUICKE, D. L. J., 1986, Preliminary notes on homeochromatic associations within andbetween the Afrotropical Braconinae (Hym., Braconidae) and Lamiinae (Col.,Cerambycidae) Entomologist’s Monthly Magazine, 122, 97–109.

SAAKSJARVI, I. E., PALACIO, E., GAULD, I. D., JUSSILA, R. and SALO, J., 2003, A new genusand six new species of the tropical Camptotypus genus-group (Hymenoptera:Ichneumonidae; Pimplinae) from northern South America, Zootaxa, 197, 1–18.

SAUSSURE, H. DE, 1892, Hymenopteres in A. Grandidier (ed.) Histoire Physique Naturelle etPolitique de Madagascar, 20 (Paris), 590 pp.

SEYRIG, A., 1932, Les Ichneumonides de Madagascar, I. Ichneumonidae PimplinaeMemoires de l’Academie Malgache, 11, 1–183.


Dow

nloa

ded

by [

Uni

vers

ity o

f Sa

skat

chew

an L

ibra

ry]

at 1

9:27

05

Oct

ober

201

2

SONAN, J., 1930, A few host known Ichneumonidae found in Japan and Formosa (Hym.),Transactions of the Natural History Society of Formosa, 20, 268–273.

TOWNES, H., 1969, Genera of Ichneumonidae, 1, Memoirs of the American EntomologicalInstitute, 11, 1–300.

TOWNES, H. and TOWNES, M., 1960, Ichneumon-flies of America north of Mexico: 2.Subfamilies Ephialtinae, Xoridinae, Acaenitinae, United States National MuseumBulletin, 216(2), 1–676.

TOWNES, H. and TOWNES, M., 1966, A catalogue and reclassification of NeotropicIchneumonidae, Memoirs of the American Entomological Institute, 8, 1–367.

VIERECK, H. L., 1914, Type species of the genera of Ichneumon flies, Bulletin of the UnitedStates National Museum, 83, 1–186.

WAHL, D. B. and GAULD, I. D., 1998, The cladistics and higher classification of thePimpliformes (Hymenoptera: Ichneumonidae), Systematic Entomology, 23, 265–298.

WHITMORE, T. C. (ed.), 1981, Wallace’s Line and Plate Tectonics (Oxford: Clarendon Press),91 pp.

WOODBURNE, M. O. and SWISHER, C. C., 1995, Land mammal, high resolutiongeochronology, intercontinental overland dispersals, sea-level, climate and vicariancein W. A. Berggren and D. V. Kent (eds) Geochronology, Time Scales and GlobalStratigraphic Correlation (Tulsa, OK: SEPM), pp. 335–364.

YU, D. and HORSTMANN, K., 1997, Catalogue of world Ichneumonidae (Hymenoptera),Memoirs of the American Entomological Institute, 58, 1–1558.

2778 Phylogenetic evaluation of the tropical Camptotypus

Dow

nloa

ded

by [

Uni

vers

ity o

f Sa

skat

chew

an L

ibra

ry]

at 1

9:27

05

Oct

ober

201

2

phylogenetic evaluation of the tropical camptotypus genus-group (hymenoptera: ichneumonidae), with a...

Documents