Contributions to Zoology, 65 (4) 201-231 (1995)

SPB Academic Publishing bv, Amsterdam

The phylogeny and taxonomic status of the Chlorocystini (sensu stricto)

(Homoptera, Tibicinidae)

A.J. de Boer

Institute for Systematics and Population Biology (Zoölogisch Museum), University of Amsterdam, P.O.

Box 94766, 1009 GT Amsterdam, The Netherlands

Keywords: Homoptera, Chlorocystini, Prasiini, phylogeny, taxonomy

Abstract PAUP 3.1.1 (Swofford, 1993) a été utilisé pour analyser les don-

dans les Prasia Muda été choisis nées; cetteanalyse genres et one

comme “outgroups”. Les résultats obtenus lors de cette analyse The “Baeturia and related genera complex”, as defined earlier par ordinateur ont été légèrement modifiés a posteriori, pour (De Boer, 1990) by shared aedeagal characters, is identified as favoriser certains caractères supposés phylogénétiquementim- the tribe Chlorocystini (sensu stricto). The Prasiini (sensu stric- fluctuants. portants, au dépens de caractères puissamment Ces to) are identified as the sister group of the Chlorocystini (sensu modifications finales été réalisées à l’aide du ont programme stricto), while the genus Muda is recognized as the nearest out- d’ordinateur MacClade 3.0 (Maddison & Maddison, 1992). Une The and of the sister and group. phylogeny biogeography group matrice complète des données ainsi qu’uneliste des caractères et outgroup is briefly discussed. Baeturia kuroiwae Matsumura is

des “character states” figurent dans un Appendix; on se rap- A transferred to the genus Muda. phylogeneticreconstruction of porte à des publications précédentes pour la description et pour all 147 species of the Chlorocystini (sensu stricto) is presented,

l’illustration de ces caractères. based on 154 characters and 409 character states. The computer program PAUP 3.1.1 (Swofford, 1993) was used for analysing the data; the genera Prasia and Muda were used as outgroups in this analysis. The results obtained from the computer analysis Introduction were slightly modified a posteriori, favouring some presumably phylogenetically important characters over strongly fluctuating This paper presents a cladogram for the “Baeturia ones.These final modifications were carried out with the aid of

and related complex", a monophyletic the computer program MacClade 3.0 (Maddison & Maddison, genera

A of tibicinid cicadas from the New Guinean- 1992). complete data matrix and a list of characters and group character in for states are given an appendix; descriptions and western Pacific This here identified region. group, illustrations of these characters oneis referred to previous publi- the selected for as Chlorocystini (sensu stricto), was cations. area-cladistic study of an area comprising Maluku,

New Guinea, and East-Melanesia. A well-solved

Résumé cladogram is essential for such a study. The

phylogenetic and biogeographic data of the genus

Le “complexe de Baeturia et des apparentés”, tel and its sister genres que complex presumed group, combined précédemment défini sur la base de caractères aedéagaux par- with such data of the Cosmopsaltriaria (a similarly tagés (De Boer, 1990), est ici identifié comme tribu Chlorocystini distributed of monophyletic group cicadas) are (sensu stricto). La tribu Prasiini (sensu stricto) est considérée used in cladistic of New Guineaand des tandis an area analysis comme groupe-frère Chlorocystini(sensu stricto), que of Indo-Melanesian le genre Muda est pris en qualité de plus proche “outgroup”. adjacent areas the region (De

Sont brièvement discutées la phylogénieet la biogéographie du Boer, 1995d). In that paper the area cladograms of groupe-frère et du “outgroup”. L’espèce Baeturia kuroiwae these groups are related to the palaeogeographical Matsumura est transférée au genre Muda. Est proposée une data of the region. reconstruction phylogénétiquede l’ensemble des 147 espèces de

The genus under 147 Chlorocystini (sensu stricto), reconstruction basée sur 154 carac- complex study comprises

into 14 Taxonomie revi- tères et 409 “character states”. Le programme d’ordinateur species, classified genera. 202 A.J. de Boer - Phytogeny and taxonomicstatus of Chlorocystini s. s.

sions, with descriptions of the species, have been are presumably not monophyletic, most of the

in for most of these in tribe are close- published separate papers generaplaced any single probably

Aedeastria 1990 Boer, related to each and the classifica- genera: De Boer, (De 1990; ly other, present

1993a), Guineapsaltria De Boer, 1993 (De Boer, tion into tribes can certainly form the basis for

1993b), Gymnotympana Stài, 1861 (De Boer, modern phylogenetic systematic work. The current

1995a), Mirabilopsaltria De Boer, 1995 (De Boer, subdivision of cicadas in six families (Cicadidae,

1995b), Papuapsaltria De Boer, 1995 (De Boer, Platypedidae, Plautillidae, Tettigadidae, Tettigarc-

1995c), Scottotympana De Boer, 1991 (De Boer, tidae, and Tibicinidae) is even more disputed than

1991), and Thaumastopsaltria Kirkaldy, 1900 (De the classification in tribes, although the two largest

Boer, 1992a). The large genus Baeturia Stài, 1866 families, the Cicadidae and the Tibicinidae, and the was subdivided into seven monophyletic species smallfamily Tettigarctidae are generally considered groups that were discussed in separate papers: the to be sound groups (see Duffels, 1993). bloetei group (De Boer, 1989), conviva group (De Several tibicinid genera from New Guinea and

Boer, 1986), exhausta group (De Boer, 1994a), gut- Eastern Australia were recognized to form one tulinervis group (De Boer, 1994b), loriaegroup (De monophyletic group, based on apomorphous ae-

Boer, 1994c), nasuta group(De Boer, 1982; 1994d), deagal characters (De Boer, 1990). This group, of

viridis Boer, the Australian which the in four and group (De 1992b); genera were previously placed genera of the complex (Chlorocysta West wood, different tribes (viz., the Chlorocystini, Gym-

1851, Cystopsaltria Goding & Froggatt, 1904, Cys- notympanini, Hemidictyini, and Taphurini), was tosoma Westwood, 1842, Glaucopsaltria Goding & not given the official status of tribe or subtribe,

since its Froggatt, 1904, Owra Ashton, 1912, and Venustria sister-group relationships were not known.

will Goding & Froggatt, 1904) be treated in a single Pending the discovery of its sister group, the group

in discussion of the the of “Baeturia and paper (De Boer, prep.). For a was given preliminary name characters used in the present phylogenetic analysis related genera complex." one is referred to these taxonomie revisions.

In order to polarise the characters, two out-

have been selected. These dis- groups outgroups are A quest for the sister group of the “Baeturia and cussed preceding the phylogenetic analysis proper. related genera complex”

Some short remarks on their phylogeny and bio- geography are made and some of the consequences The species of the “Baeturia and related genera of the presumed relationships between ingroup and complex" possess, apart from their apomorphous

for the in- which outgroups traditional classifications are aedeagal shape, one character separates them dicated. from the vast majority of other tibicinid cicadas:

they lack a medial uncus lobe. The presence of a

distinct and posteriorly directed medial uncus lobe,

The existing classification of cicadas often larger than either the lateral uncus lobes or

the claspers, is found throughout the cicadas, both

The higher classification of cicadas, as presented in in the Cicadidae and in the Tibicinidae. This the catalogues of Metcalf (1963a, b) and Duffels & character is very unstable within the Cicadidae

Van der Laan is (1985) mainly based on the work of since presence and absence often occur among

between related it be Distant, performed roughly 1888 and closely taxa, but can possibly regarded

1920. Many of the "divisions" proposed by Distant plesiomorphous in the Tibicinidaewhere a total ab- survive, virtually unchanged, as tribes in the sence occurs less frequently. The mere size of this modern classification. This classification thus pre- lobe varies strongly within both families, which

tribes in dates phylogenetic systematics and the thus makes it questionable whetherthe lobes as found formed rather are phenetic than phylogenetic these families all have a monophyletic origin. groups. Nevertheless, though most of these tribes The “Baeturia and related genera complex" Contributions to Zoology, 65 (4) - 1995 203

of cicadas. The first five shares the absence of a medial uncus lobe with the tered in other groups

Southeast Asian Muda 1897 and characters all be for the genera Distant, might apomorphies

Kumanga Distant, 1905, while this lobeis either ab- “Baeturia and related genera complex" and the sent or strongly reduced in size in the oriental Prasiini together (4-8 in Fig. la), but then each of

must have reversed several times in Prasiini (consisting of the genera Arfaka Distant, the characters

swollen 1905, Jacatra Distant, 1905, Lembeja Distant, these two groups. The angularly post-

is also found in Muda. One character in 1892, and Prasia Stài, 1863, see De Jong, 1985). clypeus

cruciform eleva- Tibicinid species without a distinct medial uncus particular, viz., the very narrow

foundin several far- tion the close lobe can also be geographically on mesonotum, strongly suggests a

Prasiini and the ther removed genera, for example Fractuosella relationship between the oriental

Boulard, 1979, AbromaStài, 1866 (Boulard, 1979), “Baeturia and related genera complex". That and Dinarobia Mamet, 1957(Orian, 1954) from the remarkable feature is shared by the Prasiini and the

Mascarene Isles, Abricta Stài, 1866 (Orian, 1964) genera Cystopsaltria and Cystosoma. The distribu- from Australia, and the African genera Trismarcha tions of the above-discussed characters are regard-

Karsch, 1891 (Dlabola, 1960) and Lamotialna ed to be convincing enough to propose the oriental

Prasiini the sister of the “Baeturia and Boulard, 1975 (Boulard, 1976) all lack a distinct as group

that is medial uncus lobe. This list does not pretend to be related genera complex", provided complex

of Some of these characters could also complete; it was based on comparison genital monophyletic.

Prasiini included drawings of a selection of the more recent litera- indicatethat the oriental are to be

" ture. in the “Baeturia and related genera complex" and

Taking into account the above-discussed instabil- that the latter [exclusively] is not monophyletic. ity of the character, it is far from certain that the This problem can only be tackled by a phylogenetic

that includes the nearest and shared absence of a medial uncus lobe forms a syn- analysis outgroups,

for all the that lobeand these at not known with apomorphy genera lacking are present any certainty.

For the of the “Baetur- a thorough phylogenetic revision of the higher taxa now unique aedeagal shape of cicadas is needed to come to any definite conclu- iaand related genera complex" is regarded as suffi- sions. However, it is supposed here that the im- cient evidence for the monophyly of that group (9 mediate relatives of the “Baeturia and related in Fig. la).

must for It is that based its genera complex" be looked among the supposed Muda, on post-

in forms the sister taxa lacking a distinct medial uncus lobe. Based on clypeus shape (2 Fig. la), group

of lobe in of theoriental Prasiini and the “Baeturia and relat- the absence a distinctmedial uncus (1 Fig.

ed combined. Several characters la) the above-listed genera might well form (part genera complex" of) a monophyletic group. of Muda, however, indicate a relationship with the

The oriental Prasiini are the most likely candi- oriental Prasiini. The large and inwardly curved

of date to be the sister group of the “Baeturia and lateral lobes of the male pygofer Muda are

from with but related genera complex". Apart the reduced presumably homologous similar, posteri- or absent medialuncus lobe, the species of that tax- orly directed, lobes in the Prasiini, while the ex- on share several characters that also occur in several tremely long ovipositor, which is found in Muda,

and also indicate rela- species of the “Baeturia and related genera com- Jacatra, Lembeja, might a plex" and represent possible apomorphies: a very tionship betweenMuda and the Prasiini. This latter narrow hyaline border along the hind margin of the character is not considereda strong synapomorphy, tegmen, a medially furrowed pronotum, a continu- since it occurs in many other cicadas. Based on the ous lateral ridge on the pronotum (only in Prasia), pygofer lobe (1 in Fig. lb), Muda and the Prasiini

otherwise this lobe a very narrow cruciform elevation on the mesono- might be sister groups, must be tum, a strongly bent proximal spine on the fore synapomorphous for Muda, the Prasiini, and the femur in of and “Baeturia and related (only Arfaka and part Lembeja generacomplex" together (3

Prasia), and a ventrally angularly swollen post- in Fig. la) and must then have reversed in the latter

These clypeus. characters are but rarely encoun- group. 204 A.J. de Boer - Phylogeny and taxonomic status of Chlorocystini s. s.

Goding & Froggatt. Most of these genera, viz. those

of which the descriptions predate that of the

“Baeturia and related genera complex", are either

included in the tribe Chlorocystini (Chlorocysta,

Glaucopsaltria, Owra, Thaumastopsaltria, and, re-

cently transferred from the Taphurini by Moulds

(1990), Venustria), or in the Gymnotympanini

(Baeturia and Gymnotympana). The tribe Gym-

notympanini was separated from the Chlorocystini

by Boulard (1979), though he regards these tribes as

sister groups (Boulard, 1990). Cystosoma is includ-

ed in the Hemidictyini and Cystopsaltria was re-

cently transferred from the Chlorocystini to that Fig. 1. (a), Cladogram showing the possible relationships of the same tribe (Moulds, 1990). Chlorocystini (sensu stricto), (b), Alternative cladogram of the The presumed monophyletic “Baeturia and possible relationships of the Chlorocystini (sensu stricto) with

related should be included in the Muda and the Prasiini as sister groups. Numbers refer tocharac- genera complex" ters discussed in the text. tribe Chlorocystini since the complex contains the

type genus, Chlorocysta, of that tribe. In this con-

The Southeast Asian genus Kumanga and some struction the Gymnotympanini must be regarded as

above-listed medial since of the genera lacking a uncus a junior synonym of the Chlorocystini, the lobe and included in the of la of the polytomy Fig. might type genus Gymnotympanini (Gymnotym-

in of possibly have to be included the sister group pana) also belong to the “Baeturia and related the Prasiini and the “Baeturia and related and thereforeto the genera genera complex" Chlorocystini

combined. complex" in their new concept. This also implies that Cysto-

In a tentative of the Mauritian cica- cladogram soma and Cystopsaltria must be transferred from

Boulard does not the relation- das, (1990) recognize the Hemidictyini to the Chlorocystini. ship between the genera of the “Baeturia and relat- ed genera complex" (viz., the tribes Chlorocystini and and the oriental Gymnotympanini, see below) The Chlorocystini

Prasiini (the latter do not occur on Mauritius and are not included in the cladogram anyway), but he The Chlorocystini (sensu lato) do not form a does indicate the genera Abricta and Abroma as monophyletic group in the new concept proposed the sister The belonging to possible group. phylo- here; the 14 genera of the “Baeturia and related genetic reasoning behind his cladogram remains genera complex" (the Chlorocystini sensu stricto) obscure. Prasiini are more closely related to the oriental than

to theremaining generaincluded in the tribe. These

remaining genera of the Chlorocystini, 15 in num- Tribal allocation of the “Baeturia and related ber, will have to be removed to other tribes. Six of genera complex” Dis- these genera (Akamba Distant, 1905, Bavea

1905, Decebalus Distant, 1920, Musoda The “Baeturia and related genera complex" com- tant, Karsch, 1890, Nablistes Karsch, 1891, and Stagira prises 14 genera: AedeastriaDe Boer, BaeturiaStài,

Chlorocysta Westwood, Cystopsaltria Goding & Stài, 1861) occur in Africa, two ((Fractuosella and

Froggatt, Cystosoma Westwood, Glaucopsaltria Dinarobia) are endemic to Mauritius and Réunion,

Goding & Froggatt, Guineapsaltria De Boer, Gym- two (Muda and Kumanga) come from Southeast

is restrict- notympana Stài, Mirabilopsaltria De Boer, Owra Asia, one ((Toxopeusella Schmidt, 1926)

Ashton, Papuapsaltria De Boer, Scottotympana De ed to Buru Islandin Maluku, New Guinea, the Bis-

Boer, Thaumastopsaltria Kirkaldy, and Venustria marck Archipelago, and the Solomon Islands, and Contributions to Zoology, 65 (4) - 1995 205

1905 and Dis- two ((Conibosa Distant, Durangona Karsch, 1890 and Iruana Distant, 1905) are Afri-

tant, 1911) are endemic to Middle and South can, and one (Sapantanga Distant, 1905) comes

America, respectively. from South America. These non-oriental genera

the As explained above, genus Muda alone might were placed in the Prasiini because of their narrow

be the sister group of the Chlorocystini (sensu stric- head and protruding postclypeus. However, closer

to) and the Prasiini (sensu stricto) together (Fig. examination shows that there are large differences

la), but Dinarobia, Fractuosella and Kumanga between these and the oriental genera. The oriental

be of that sister be might part group, or closely Prasiini have a much narrower head than either

related that the to sister group. Lacetas, Iruana, or Sapantanga. Furthermore,

Several of the African genera have been recently postclypeus of these latter three genera is not angu-

treated in literature. Bavea, Musoda, and Stagira larly swollen as is the case in the oriental Prasiini;

share medial a large tube-like uncus lobe (Boulard, Lacetas and Sapantanga have a conically protrud-

1979; Villet, 1993). They are supposed to be related ing postclypeus, while Iruana has a much smaller,

to the South African generaXosopsaltria Kirkaldy, hardly swollen, postclypeus. Iruana (Dlabola,

1904 and Tettigomia Amyot & Serville, 1843, and 1960; Boulard, 1981b, 1990) and Lacetas (Boulard,

should be included in the African distinct medial possibly mainly 1975a, 1975b) have a very uncus

tribe Tettigomyiini. Nablistes likewise has a very lobe. Lacetas is presumably related to Hovana Dis-

large medial uncus lobe (Dlabola, 1958) and is tant, 1905 and Hemidictya Burmeister, 1835, shar-

also related streamlined presumably more closely to the Tet- ing a very similar head and thorax (De

tigomyiini. No genital drawings have been pub- Boer, in prep.) and should probably be placed in the

lished for Akamba. tribe Hemidictyini. Only two females of Sapantan-

Conibosa is reported to share the total absence of ga were examined. That genus is presumed to be

with the South opercula American genera Coata related to the South American genera Prosotettix

Distant, 1906 and Prunasis Stài, 1862 (Boulard, Jacobi, 1907 and Selymbria Stài, 1861 of the

1979). Coata and Prunasis are at present placed in Taphurini (De Jong, pers. comm.).

two different families: Coata belongs to the Tet-

tigadidae and Prunasis to the tribe Parnisini of the

Tibicinidae. The correct tribal allocation of Phylogeny and biogeography of the outgroups

Conibosa is therefore uncertain. No recent litera-

ture is available on Durangona, but Distant states The Prasiini (sensu stricto), the supposed sister

in its first description that the genus is allied to group of the Chlorocystini (sensu stricto), and the

Conibosa. genusMuda serve as outgroups in the phylogenetic

Toxopeusella presumably belongs to the tribe analysis of the Chlorocystini. In this section some

Cicadettini (De Boer, 1990). Its species share the of the most characteristic features and possible typical trifid aedeagus and a reddish costal area of apomorphies of the various genera of these out- the with the of that tribe tegmen genera (see groups are discussed inrelation to a tentative clado-

Boulard, of the Prasiini. remarks 1981a). gram Furthermore, some

on the geographical distributions of these genera

will be made.

The Prasiini

The Prasiini (sensu lato) are likewise not mono- The Prasiini (sensu stricto)

The four oriental phyletic. genera (the oriental

Prasiini the Prasiini close- or sensu stricto) are more Thanks to the revisionary work of De Jong (1982, ly related to the Chlorocystini (sensu stricto) and to 1985, 1986, 1987) and De Jong & Duffels (1981) the the genus Muda than to the three other genera of oriental Prasiini form a fairly well-known group. the tribe. Two of these other genera (Lacetas Regrettably the revision of this tribe has not been 206 A.J. de Boer - Phytogeny and taxonomic status of Chlorocystini s. s.

ed to Sulawesi and the nearby island of Muna. Ac-

cording to De Jong (1985) the monophyly of Prasia

"the is based on a single apomorphy: degrading

lateral margin of the male pygofer between the

lateral lobes and the caudodorsal beak" (5 in Fig.

2). Inaddition to this, a long and spine-shaped male

to caudodorsal beak is presumed here be apomor-

for well in Other phous the genus as (6 Fig. 2).

apomorphies for Prasia can possibly be found in

the shape of the lateral lobesof the pygofer and in

the structure of the base of the aedeagal append-

is from the other ages. Prasia easily separated

of Prasiini Prasiini lateral of the Fig. 2. Tentative cladogram the (L. = Lembeja). by a sharply edged margin

refer to characters discussed in the text. Numbers pronotum. This character is also foundin Cystoso-

ma, Cystopsaltria, three species of Mirabilopsal-

solved and three of It completed and a fully phylogenetic recon- tria, species Gymnotympana. is as-

struction is not available. The following remarks sumed that Prasia forms the sister group of.Arfaka,

about the phylogeny and the biogeography of the Jacatra, and Lembeja combined. The latter three

oriental Prasiini are principally based on De Jong's genera share a carinated male abdomen as a pre-

publications and on unpublished observations sumed synapomorphy (7 in Fig. 2).

made author. is small with 3 of by the same A tentative cladogram of Arfaka a genus only species,

the Prasiini is presented in Fig. 2. which one is still undescribed. The genus is

The oriental Prasiini form a supposedly mono- presumably endemic to Cendrawasih (the Vogelkop

of the islands phyletic group, containing the genera Arfaka, Peninsula New Guinea) and adjacent

is Jacatra, Lembeja, and Prasia (De Jong, 1985), on of Waigeu and Salawati. Arfaka easily recog-

the basis of the following presumed apomorphies: nized by the long and slender apical areas of the teg-

(1) a very narrow vertex, with the ocelli close mina and wings, a large laminiform protuberance

of and together; (2) a large obconical and triangularly pro- on the lateral lobes the pygofer, a peculiar

first truding postclypeus; (3) a very narrow hyaline narrow and bilobate aedeagal apex. The

border of and also in several of the Chlo- along the hind margin the wing; (4) character occurs genera

well-developed and posteriorly projecting pro- rocystini; the latter two characters presumably

tuberances on the lateral lobes of the male pygofer represent good apomorphies for this genus(8 & 9 in

(1-4 in Fig. 2). The latter character is presumably Fig. 2). Arfaka presumably forms a monophyletic

with but its homologous with similarly shaped, though incurv- group Jacatra and Lembeja, phylo-

ing, lobes in Muda. genetic relationships are not unambiguous. Arfaka

Other characters, that either have a very limited and Prasia share a sexual dimorphism in the anal

distribution outside the oriental Prasiini (a very areas of the wings and might be sister groups,

narrow cruciform elevation on the mesonotum), or together forming the sister group of Jacatra and

shared all of fu- similar sexual are not by members the group (a Lembeja together. A dimorphism

sion of wing veins Cu 2 and A 1, a dorsal aedeagal was found in Gymnotympana (De Boer, 1995a) and

appendage) also seem to support a close relation- the Baeturia loriae group (De Boer, 1994c) and

ship between the species of the oriental Prasiini. therefore the character cannot be regarded a strong

The in oriental Prasiini are distributed Sumatra, synapomorphy. Jacatra and Lembeja share an

Java, Borneo, the Lesser Sunda Islands, Philip- elongate first tergite in male individuals as a pre-

pines, New Guinea, and northern Queensland, but sumed synapomorphy (10 in Fig. 2).

have most species on Sulawesi. Jacatra contains only 2 species, occurring in Java

The is described. The genusPrasia contains seven species, restrict- and Sumatra. Only one species Contributions to Zoology, 65 (4) - 1995 207

three genus can be recognized by apomorphies: an tion of the first tergite of the male as presumed syn-

aberrant the in vein partly dividing 4th ulnar area of apomorphy (14 Fig. 2).

the tridentate first tegmen, a proximally weakly

tergite in males, and a peculiar coalescence of the The genus Muda apical lobes of the aedeagus (characters 11-13 in

Fig. 2). Muda presently contains three species, but several Lembeja, with about 40 species, forms by far the new species can presumably be separated from of the oriental Prasiini. Five largest genus mono- these. Muda kuroiwae (Matsumura, 1913) n. comb, phyletic species groups can be recognized within from the Ryukyu and Loochoo Islands of southern Lembeja, and three such groups have been revised: Japan, is hereby transferred from Baeturia to (1) the foliata group, with 13 species endemic to Muda. Both other species are distributed over the and Sulawesi Sangihe Island, can be recognized by Greater Sunda Islands (Java, Sumatra, and Bor- the first sterniteof a longitudinal ridge on males(15 neo), and the Malay Peninsula. Muda is easily in Fig. 2) (De Jong, 1986); (2) the fatiloqua group recognized by its peculiar pygofer, being sharply

with 10 species, most of which are endemic to foldedalong its dorsal margin and, in consequence,

Sulawesi, also occurs in Mindanao, northern Bor- nearly triangular in cross section (20 in Fig. 2). The

neo, Sumba, southeastern New Guinea and north- laminiform large and rounded protuberances on

ern and has a Queensland, strongly developed the laterallobes of the pygofer, which are probably medial dent in the first of males and in- tergite an homologous with the large protuberances in the

flatable male abdomen as its telescoping presumed Prasiini (sensu stricto), curve or fold mesiad along

in apomorphies (16-17 Fig. 2) (De Jong, 1987); the base of the anal valves so that their apices are and (3) the small parvula group with 2 species en- adjacent. These lobes possibly replace the strongly demic Sulawesi has hindwards directed distal to a reduced claspers in function. Muda shares a some- margin of the male pygofer and five apical areas in what similarly shaped postclypeus with the oriental the wing as apomorphies (18-19 in Fig. 2) (De Prasiini, and with Cystopsaltria, Cystosoma, and

Jong, 1987). Two other groups, the harderi group Thaumastopsaltria. from the Lesser Sunda Islands and the robusta group from New Guinea, Obi Island, and northern The phylogenetic analysis Queensland, can be recognized (De Jong, pers. comm.), but these groups still await description. Phylogeny and parsimony, the value and restric- The phylogeny of Lembeja is not resolved, but tions of a computer analysis the genus is assumed to be non-monophyletic in its present concept. Lembeja harderi Schmidt, 1925 The method of phylogenetic reconstruction, as de- and several related, undescribed, species forming veloped by Hennig (1950; 1966), is based on the as- the harderi relat- group, are probably more closely thatif character sumption a state is uniquely shared ed to Prasia and Arfaka than to other species of by two species, that character state did not evolve

Lembeja. These species lack the carinatedmale ab- independently in these two species, but developed domenand first of elongate tergite males, common in the only once common ancestor of those species. to the other Lembeja species, and share a fairly Based on this principle, dichotomous family trees small and sexual in with tymbal dimorphism wings or cladograms can be constructed, in which all the

Prasia and The four various Arfaka. remaining species groupings of species which share a direct

of Lembeja together are thought to form a common ancestor groups (monophyletic groups) are sup- monophyletic group (De Jong, pers. comm.), ported by one or more shared, but otherwise

character has although no synapomorphous as yet unique, character states that developed in that com- been The and published. fatiloqua parvula groups mon ancestor. are supposedly sister groups, with a medial indenta- This may sound very simple in theory, but in 208 A.J. de Boer - Phytogeny and taxonomic status of Chlorocystini s. s.

into trouble when has six and that this number is practice one immediately runs cies ridges, provided character state distributions conflict. Suppose that not the most plesiomorphous state, it could have a species "A" shares one character uniquely with acquired that number by gaining a sixth or loosing

with If it this have species "B" and two characters uniquely spe- a seventh ridge. gained a ridge, can cies "C". In that case the most parsimonious solu- happened by the duplication of any of its previous tion is chosen as the most likely. five ridges; ifit lost a ridge, this can be due to the

The principle of parsimony is nothing but apply- loss of any of its previous seven ridges, or to the fu-

sion of successive In if ing Ockham's razor, choosing for the simplest any two ridges. short, any solution in view of all the facts. Translated to two species share the same number of tymbal phylogenetic analysis, this means that the shortest ridges, unless these species are closely related, it is cladogram is opted for, the cladogram requiring the unlikely that a shared number of ridges represents

still is character "number of smallest number of character changes that a homology. The tymbal compatible with the data set. In our example, arela- ridges" should thereforeonly be allowed to play a tionship between "A" and "C" is favouredsince in role within species groups and should be entirely that option only one character need be supposed to disregarded at a more basal level. Presumably there

of of have developed twice. are many more these types characters; they

If evolution often be low we accept as being a parsimonious can recognized a posteriori by a con- process, a statement that is by no means proven, sistency index. then it is clear that the "true" cladogram of the Another consideration should be that one

in evolutionary relationships between species of a character simply does not equal another. While,

is also the substitution of group the most parsimonious one. Regretta- genetic terms, one base-pair

reversal of this that sufficient character from bly the statement, the most par- might be to change one a simoniouscladogram is also the "true" cladogram, state "A" to a state "B" and back, a much more is not necessarily true, perhaps even unlikely. The drastic change in the DNA structure might be re- reason is thatany given data set is per definitionin- quired to change another character from state "A"

of for One in the latter character complete. In thepresent study cicadas, exam- to state "B". change ple, only morphological characters are used, but might (in evolutionary sense) be less parsimonious song, periodicity, food preference, ecological con- than five changes, gains and losses, in the former. straints, and even distributionare all characters of Such character differences cannot be recognized a the species. Furthermore, of these morphological priori, though an education guess can be madethat

characters of adults the and characters, only were used, as more complex structures many genital

sometimes limited number of characters and characters for the represented by a very responsible mutu- specimens, and only those characters were used that al recognition between the sexes belong to the latter were recognized as such by the author. In short, the type. Characters of the former, easily changing, data set contains only a fraction of the total number type can be recognized a posteriori by a low con- of characters and that fraction is far from random- sistency index, but there is no way of telling how ly chosen. many changes in one character equal a change in

Apart from being incomplete, the data set is another. Even when possible differences in genetic

in bound to be inaccurate as well. Many of the complexity are not madeapparent by differences

obvious func- homoplasous characters (character states that ap- consistency index, characters with an

tion in and pear to develop two or more times in the clado- reproduction (as clasper aedeagal gram) that were recognized, and therefore scored, characters) must presumably prevail over charac- as one and the same character (state) are in fact two ters with an obscure function (as a pointed or trun- or more character(state)s. Often this cannot be cate caudodorsal beak on the pygofer). However,

in since if do know recognized a priori, it can only be deduced a we must be reluctant this, we not

is posteriori. Consider for example the number of the function, this does not mean that there no sclerotized of cicadas. function. the former characters are to ridges on the tymbal If a spe- Still, likely Contributions to Zoology, 65 (4) - 1995 209

have been more subjected to natural selection than shared characters and, unless we know exactly how

the latter. the apomorphous character changed in the descen-

The PAUP 3.1.1 have computer program (Swofford, dants, we no convincing arguments to regard

1993) offers many options to adjust the data set a that group as monophyletic. On the other hand, if

priori, in order to counteract some of the above- we do recognize groups of species that share a

discussed Characters be do have problems. can ordered, or unique character, we a strong argument to

made irreversible, or given a weight (for explana- regard that group as monophyletic. In that case,

tions see the manual of the program), but, since unless this group contradicts with other equally

is need there is, at least in morphological studies as the well-argued monophyletic groups, there no

method to determine for the for anal- present one, no objective parsimony criterion, nor computer

what character should be adjusted in what way, it ysis. One should not allow to let the computer com-

could is feared that such manipulations are only per- pute what is already known. The computer

solution based formed to bring the computer results closer to the come up with a more parsimonious

pre-conceived ideas of the scientist. Alternatively, on characters, none of which are distinctive for the

PAUP gives the opportunity to weight characters a monophyletic groups they are supposed to support.

posteriori, with a character weight relative to the Only when the scientist does not have any strong ar-

index retention index of that charac- in the form of shared consistency or guments apomorphous ter (Farris' reweighting method; Farris, 1969). characters in favourof certain groupings, the most

However, then the weights given are dependent on parsimonious solution is the best he can come up the behaviour of in with. character is a character a cladogram that When a regarded as apomor- was unsatisfactory to us in the first place. In large phous for a certain species group by argument of its data sets as the one presented here, such a succes- exclusivity and considered important in view of its

sive weighting runs the risk of loosing valuable in- supposed genetic complexity, and/or its supposed

formation. When character sound role in sexual and that character does a supports a isolation, not monophyletic group in one part of the cladogram conflict with another equally well-argued possible but is highly inconsistent inother parts, it will have apomorphy, then the final cladogram should show

index thus will in not- a low consistency and get a low the species group question as monophyletic,

result of which lose the conflict with characters of weight, as a we might sup- withstanding any sup- port for the monophyletic group. Such a group posed minor importance. When there are such should then be protected by a constraint before re- strong arguments in favour of a certain decision,

is for weighting. there no need parsimony. Only when no clear-

ly apomorphous characters can be found, or when

Ockham's razor is used in philosophy to decide be- two equally well-supported apomorphies con- tween questions where there are no convincing ar- tradict, parsimony offers a last resource and the guments in favour of any one solution. The sim- most parsimonious solutionshould be chosen as the plest solution is chosen by lack of good arguments most likely one. pro or contra. I plea for a similar use of parsimony Although the relative importance of characters in phylogenetic analysis. A monophyletic group is can often be made plausible, there is, at least in recognized in principle by one or more apomor- morphological studies, no way of quantifying that phous characters, characters that developed in the importance and we thereforehave to accept that the

and inherited the descen- science of will contain common ancestor are by phylogeny always an element dants. Of course, a common ancestor could have of subjectivity.

developed an apomorphy that changed or reversed A computer analysis as discussed below has no

of its of in some, or many, descendants, as a result other objective thanto demonstrate the amount of which the monophyletic group cannot easily be conflict between the supposed apomorphies and the recognized as such. The monophyly, though exist- remaining characters. The amount of consistency ing in reality, is then apparently not supported by between the computer tree and a tree solely based 210 A.J. de Boer - Phytogeny and taxonomicstatus of Chlorocystini s. s.

on the supposed apomorphies can be taken as a topological constraint on the monophyly of the kind of measure for the soundness of these apomor- Chlorocy stini. phies. Only when the computer tree deviates strong- A second constraint, on the monophyly of a

from the tree based the ly on the recognized apomor- groupcontaining generaBaeturia, Chlorocysta, phies, the value of these apomorphies must be seri- Glaucopsaltria, Guineapsaltria, Gymnotympana, ously reconsidered. Since these deviations are not Owra, Papuapsaltria, Scottotympana, and Venus- easily quantified, it must be left to the scientist to tria, appeared necessary to force some single and

what decide deviations are acceptable. deviating species into their correct genus. Although

a definite segregation, based on several presumed

apomorphies (see Fig. 4), appears to exist between

The computer analysis the above nine genera and theremaining five genera

of the Chlorocystini, some deviations occur in both

Material. - A cladistic analysis was performed for these subgroups which had a negative effect on the the 147 species of the Chlorocystini (sensu stricto), whole tree. using 154 characters with a total of 409 character

states. The search. - An heuristic search was made (in

view of the size of the data set) with the options ran-

Characters. - All characters are introduced as un- domadding sequence by stepwise addition, holding orderedand no character weights have been applied one tree at each step, keeping minimal trees only for reasons discussed above, that is: all characters and collapsing zero length branches, and the option have an equal weight. A matrixof the data and a list Mulpars off. Swapping was performed on minimal of the characters are included in an Appendix. trees only with the swapping algorithm tree bisec-

tion - reconnection. The trees found had a length

- in Outgroups. The character states as foundin Pra- of 875 steps; these were used as starting trees a sia and Muda are added to the data matrix, as new search, now with the option Mulpars on, and

the These maximum of representing presumed outgroup states. a set 600 equally parsimonious trees two genera were defined as outgroup in the search. was reached.

Prasia, as representing the sister group Prasiini, was chosen for several reasons: (1) Prasia is Results. - The strict consensus tree of these 600

the basal of the oriental is level in presumably most genus trees given to a generic or species group

Prasiini (see Fig. 2), (2) a taxonomie revision of Fig. 3. The total strict consensus tree has a length

Prasia with genital drawings is available, (3) of all of 821 steps; consistency index = 0.29; retention in-

Prasiini, Prasia shows the greatest similarity to the dex = 0.82; rescaled consistency index = 0.24. The

of the species Chlorocystini (sensu stricto), and has 50% majority rule consensus tree is one of the 600 therefore retained of the with of 875 index presumably more plesio- trees a length steps; consistency morphous states, but an analysis with a third out- = 0.27; retention index = 0.80; rescaled consisten- group is necessary to check this point. cy index = 0.22. This tree shows nearly all previ-

ously recognized genera and species groups as

Constraints. - Two topological constraints were monophyletic groups. Parts of this 50% majority built in. The male genitalia (which comprise 47% of rule consensus tree are given in Figs. 5, 6, 7, 9, 12a, the of the Muda and 13. Inother this is identical the characters used) genus are so com- parts tree to one pletely differentfrom those of the Prasiini and the presented in Fig. 4. Several of the 600 trees found

Chlorocystini that the remaining characters of that give amost interesting alternativefor the phylogeny

insufficient force of genus carry weight to Muda to an the genus Papuapsaltria (see Fig. 12b). outgroup position within the tree. If a third out- group were known, this problem would probably Modifications. - The strict, and 50% majority

from not exist. The problem was solved by putting a rule consensus trees differ in several instances Contributions to Zoology, 65 (4) - 1995 211

and will be from left to cies groups genera discussed

right (Fig. 4), starting with the genus Baeturia.

Discussion of modifications

Baeturia

The ingroup phylogeny of the Baeturia nasuta

of distinct group is very uncertain, due to a lack

apomorphies, which is illustrated by the fact that

in the strict this groupappears as a single polytomy

consensus tree. The 50% majority rule tree of this

group (Fig. 5) differs from the tentativecladogram

published earlier (De Boer, 1994d) in that B. mar- Fig. 3. Simplified version of the Paup reconstruction for the morata and not B. guttulipennis is placed at the Chlorocystini s. s. (strict consensus) (B. = Baeturia; P. = base of the The nasuta minus B. gut- Papuapsaltria). group. group considered based tulipennis was monophyletic on

two apomorphies: a narrowly protruding post- the tentative cladograms published earlier (De clypeus and four tymbal ridges. A transparent ab-

nasuta and Boer, 1994a, d; 1995a, b, c). These differences are domen is apomorphous for the group,

abdomen of B. discussed below and some minor modifications are the untransparent marmorata, made. The "costs" of each of these modifications which is responsible for the basal position of that

reversal. The and of some proposed alternatives, expressed in the species, was considered a character

four number of extra steps needed to construct the tree, apomorphy of tymbal ridges was not recog-

nized in the caused the fact that were calculated with the computer program Mac- present analysis, by

all characters introduced unordered Clade 3.0 (Maddison & Maddison, 1992). To en- were as

within sure that the most parsimonious tree that takes the (though the number of ridges reduces Bae-

numberof required modificationsinto account was reached, a turiafrom 7-8 to 4 ridges; the ridges can

increase in other B. new run was made in PAUP, but now using a con- genera). Removing guttulipen- straint tree enforcing the proposed changes. The nis to the base of the nasuta group, as proposed

The of resulting alternative shortest acceptable tree is here, costs 1 extra step (Fig. 4). phylogeny

the of shown in Fig. 4. This tree has a length of 906 steps. monophyletic subgroup arabuensis, bipunc-

Where several equally parsimonious solutions were tata, gibberosa, laminifer, mamillata, and retracta

earlier. possible, the tree of Fig. 4 was madeto resemblethe agrees with the cladogram published

of the and 50% majority rule consensus tree as much as possi- The phylogeny viridis, conviva, gut-

alter- tulinervis and their relative ble. Modifiedin the same way, a tree with the groups, positions, agree

earlier native phylogeny for Papuapsaltria yields a short- with the cladograms published (Duffels &

Their est acceptable tree of 905 steps (cf. Figs. 12c and De Boer, 1990; De Boer, 1994a, d). phyloge-

14). ny as given in Fig. 4 is identical to that of the 50°7o

in 4 indicate rule tree for the conviva and The black squares Fig. one or more majority and, gut-

the strict characters that are considered phylogenetically tulinervis groups, to consensus tree.

differences between the of thebloetei meaningful; these are given in Appendix 2. The species

the are often small, which has resulted in an Branches without a black square just represent group very

with most parsimonious way to deal with all characters only partly resolved cladogram, several polyt- concerned. omies. Even this cladogram shouldbe considered as

The phylogenies and positions of the various spe- doubtful. Only the monophyly of the Moluccan 212 A.J. de Boer Phytogeny and taxonomic status of Chlorocystini s. s.

4. Fig. Most parsimonious McClade reconstruction for the Chlorocystini s. s. after including the modifications suggested. Tree length

= 906; consistency index = 0.26; retention index = 0.79; rescaled consistency index = 0.21. Black squares refer to the apomorphies

listed in Appendix 2.

and New Guinean B. B. species macgillavryi, subgroups was suggested (De Boer, 1994c). B.

bloetei, and B. papuensis and of the east Melanesi- daviesi, B.fortuini, B. hartonoi, B. loriae, B. pigra-

an species B. brandti, B. cristovalensis, B. edauber- mi, and B. silveri were supposed to form a

maddisoni, sedlaceko- ti, B. B. reynhoudti, and B. monophyletic group based on a shared clasper pro-

assumed be sound The latter tuberance. The rum are to (Fig. 6). position of B. tenuispina, as part of

does not in the strict group appear consensus tree, either the exhausta group or of the loriae group,

however. The species from the Bismarck Islands (B. was a problem. Its present position between these

bismarckensis, B. manusensis, B. mussauensis), two species groups is thereforeconsidered quite ac-

which were based on presumed to be monophyletic, ceptable (Figs. 3, 4). B. tenuispina, however, ap-

shared and related a aedeagal shape, closely to B. parently has the same clasper protrusion as found papuensis (cf. De Boer, 1989), appears as para- in the loriae group and due to the position of that phyletic at the base of the bloeteigroup in the 50% species, the most parsimonious reconstruction does

majority rule tree (Fig. 6). Joining these three not recognize that character as apomorphous for

to species a monophyletic group, as proposed here, the B. loriae subgroup so that the species of that costs 1 extra B. should in strict step (Fig. 4). papuensis pos- subgroup appear as paraphyletic the con- sibly be included in this group. The suggested sensus tree (Fig. 7). Consequently, this character monophyly of the species of the bloetei group ex- must have been lost in the other subgroup of the

these Bismarck cluding species is based on a shared loriaegroup. Several other presumed apomorphies, broad aedeagal pore. The aedeagus of the Bismarck shared by B. fortuini, B. hartonoi, and B. loriae,

is in from that of other also such. the species quite different aspect are not recognized as In regard of un-

Baeturia species. The narrow pore of the Bismarck certain position of B. tenuispina it is quite possible species might well be a reversal, and the basal posi- that its clasper protrusion is not homologous with tion of this within subgroup the bloetei group is that of the above-listed loriae subgroup species. regarded far from certain. Converting these species into a monophyletic group

The phylogenetic reconstruction of the exhausta and making B. hartonoithe sister species of B. for- group presented in Fig. 4 closely matches the pre- tuini and B. loriae, as proposed here costs no less sumed relationships as discussed earlier (De Boer, than 6 extra steps, however (Fig. 4).

1994a). This phylogenetic reconstruction is identi- The relative positions of the bloetei, exhausta, cal to that of the strict and loriae considered uncertain in consensus tree. groups was

In earlier of the an discussion on the phylogeny previous publications and a tentative cladogram loriae subdivision into two has been in which these form group, a monophyletic published groups a Contributions to Zoology, 65 (4) - 1995 213

Fig. 4. (Cont.).

polytomy with the remaining part of Baeturia (De tree and in accordance with earlier discussions (De

Boer, 1994 a, b, c). The here presented most par- Boer, 1994a, 1995a).

reconstruction is simonious regarded as good as

any. None of the branches responsible for this par- Gymnotympana

ticular solution are supported by distinct and un- The phylogeny of Gymnotympana as represented in

differs ambiguous apomorphies. The weakness of this part the 50% majority rule tree (Fig. 9) slightly

of the cladogram is illustrated by the fact that swap- from the analysis presented earlier (De Boer,

ping the position of any two of the three groups 1995a), which was based on an incomplete data set.

would cost only 1 step. A cladogram for the species Although the earlier recognized subgroups still ap-

of in the their relative groups Baeturiaas presented Fig. 8 seems pear as monophyletic groups, posi-

most realistic in view of the data available.

B. bloetei

B. macgillavryi Scottotympana B. papuensis The phylogeny and position of Scottotympana B. gressitti (Fig. 4) is identical to that in the strict consensus B. bilebanarai

B. mendanai

B. marmorata B. marginata

B. nasuta B. boulardi

B. guttulipennis B. rotumae

B. parva B. reynhoudti

B. mamillata B. brandti

arabuensis B. B. edauberti

B. gibberosa B. sedlacekorum

B. bipunctata B. cristovalensis

B. laminifer B. maddisoni

B. retracta B. mussauensis

B. intermedia B. bismarckensis

B. splendida B. manusensis

Fig. 5. Section of the 50% majority rule tree dealing with the Fig. 6. Section of the 50% majority rule tree dealing with the

Baeturia nasuta group. Numbers refer to percentages of trees Baeturia bloetei group. Numbers refer to percentages of trees supporting the branch. supporting the branch. 214 A.J. de Boer - Phytogeny and taxonomic status of Chlorocystini s. s.

B. silveri tions have slightly changed. The tree presented here

B. daviesi is, as to these relative positions, regarded as equally

B. pigrami acceptable as the one published before. Four spe-

B. loriae cies (G. dahli, G. rubricata, G. strepitans, and G.

B. fortuini form viridis) that are supposed to a monophyletic

B. hamiltoni based shared characters of the group on clasper, B. wegeneri and that appeared as paraphyletic in the earlier

B. bemmeleni cladogram (De Boer, 1995a) now appear as being B. hartonoi less closely related. Converting these four species

into a monophyletic in a then most 7. group, par- Fig. Section of the strict consensustree dealing with the simonious reconstruction of Gymnotympana as Baeturia loriae group.

proposed here (Fig. 4) costs 4 extra steps.

Venustria

The position of the monotypic genus Venustria as

sister taxon of Gymnotympana, which also appears

in the strict is in consensus tree (Fig. 3), agreement

with earlier proposals and here accepted as most

likely. Nevertheless, two unique characters con-

tradict this solution and suggest that Venustria

should be includedin Gymnotympana as in Fig. 10.

V. G. and G. varicolor share 8. Alternative for the of Baeturia. superba, rufa, a very Fig. cladogram species groups

similar basal part of the male operculum (1 in Fig.

G. strepitans 10), while V. superba and G. rufa share a distinct

G. nigravirgula ventral thorn on the female genital segment (2 in

G. parvula Fig. 10) (De Boer, 1995a). The inclusion of Venus- G. stenocephalis tria in Gymnotympana, as is considered possible, G. stridens would cost 5 extra steps. G. obiensis

G. subnotata Chlorocysta, Glaucopsaltria, and Owra G. membrana The phylogenetic reconstruction of Chlorocysta, G. minoramembrana

and in Glaucopsaltria, Owra, as presented Fig. 4, is G. phyloglycera

identical to that in the strict consensus tree and ac- G. langeraki

as correct. G. rubricata cepted

G. hirsuta In an earlier discussion on the phylogeny of the

the relative G. olivacea Chlorocystini (De Boer, 1995a, b, c, d)

G. verlaani positions of the Australian genera complex (Chlo-

G. montana rocysta, Glaucopsaltria, and Owra) and of the sis-

G. viridis ter genera Guineapsaltria and Papuapsaltria re-

G. dahli mained uncertain; either of these groups could be

G. rufa sister of contain- the group the monophyletic group

G. varicolor ing Baeturia, Gymnotympana, Scottotympana,

V. superba and Venustria. The most parsimonious solution in-

dicates, also in the strict consensus tree, that the

Fig. 9. Section of the 50% majority rule with tree dealing Gym- Australian sister of the latter genera form the group notympana. Numbers refer to percentages of trees supporting This is solution, however, not based on con- the branch. group.

vincing synapomorphies. The weakness of this part Contributions to Zoology, 65 (4) - 1995 215

of the cladogram is expressed by the fact that swap- G. nigravirgula

the of the Australian with that ping position genera G. parvula of Guineapsaltria and Papuapsaltria would make G. stenocephalis

the total tree only 2 steps longer. In view of the G. membrana available data, a generic cladogram with a polyto-

minoramembrana in is realistic of G. my as Fig. 11 a more way represent-

G. ing these uncertainties. phyloglycera

G. stridens

Guineapsaltria G. obiensis

The most parsimonious reconstruction of Guine- G. subnotata

apsaltria in the 50% majority rule tree (Fig. 12a) is G. langeraki largely in agreement with the earlier discussion on

G. hirsuta the of that The phylogeny genus (De Boer, 1993b).

G. olivacea only problem forms the ambiguous position of G.

chinai. This species shares a similar clasper with G. G. verlaani pallida, G. pallidula, and G. viridula, but a similar G. montana

postclypeus with G. flava, G. flaveola, and G. G. rubricata

stylata, and has a, presumably plesiomorphous, G. strepitans narrower vertex than other species of Guineapsal-

G. viridis tria. Consequently, G. chinai is placed at the base

G. dahli of the cladogram of Guineapsaltria. The striking

G. rufa similaritiesin the unique clasper shape of G. chinai,

G. pallida, G. pallidula, and G. viridula, however, V. superba

evidence for the are regarded as convincing mono- G. v aricolor

phyly of that group, though possibly including the

G. the of G. deviating pennyi. Changing position Fig. 10. Cladogram of Gymnotympanawith Venustria included.

Numbers refer characters discussed in the chinaito that group, as proposed here (Fig. 4), costs to text.

2 extra steps.

Baeturia

Papuapsaltria Scottotympana

The monophyly of the genus Papuapsaltria was Gymnotympana considered doubtful, since its presumed apomor- Venustria phy, a reduced femoral spine, is not found in all

Chlorocysta species of the genus (De Boer, 1995c). Several

Owra monophyletic groups were recognized within Pa- puapsaltria and it was stated that a subgroup with Glaucopsaltria a mainly northwestern New Guinea distribution Guineapsaltria showed similarities in venation with tegmen Papuapsaltria

Two P. and Guineapsaltria. species, phyllophora Aedeastria

the P. woodlarkensis, were tentatively placed at Mirabilopsaltria base of the cladogram since their characters did not Thaumastopsaltria give a clue to a possible sister-group relationship Cystopsaltria with other species (groups) of the genus. In this Cystosoma light, it is remarkablethat the computer analysis in-

all characters with volving comes up two equally 11. Generic ofthe with Fig. cladogram Chlorocystini a polytomy parsimonious solutions. In Fig. 12b, Papuapsaltria for the groups Baeturia-Venustria, Chlorocysta-Glaucopsaltria, is paraphyletic, with the northwestern New Guinea and Guineapsaltria-Papuapsaltria. Number refers to character species as sister group of Guineapsaltria and the re- discussed in the text. 216 A.J. de Boer - Phylogeny and taxonomic status of Chlorocystini s, s.

G. flava G. flava

flaveola G. G. flaveo la

G. G. stylata stylata

G. pallidula G. viridula

G. i penny G. pallida

G. pallida G. pennyi

G. viridula G. pallidula

G. chinai G. chinai

P. phyllophora P. toxopei

niodes P. P. baasi go

P. brassi P. ustulata

P. angulata P. novariae

P. lachiani P. dolabrata

P. spinigera P. phyllophora

P. stoliodes P. goniodes

P. plicata P. brassi

P. ungula P. angulata

P. bidigitula P. lachlani

baasi P. P. spini gera

P. nana P. stoliodes

P. toxopei P. plicata

riae P. n ova P. ungula

P. dolabrata P. nana

P. ustulata P. bidigitula

P. woodlarkensis P. woodlarkensis

P. dioedes P. dioedes

Fig. 12. (a), Section of the 50% majority rule tree dealingwithGuineapsaltria andPapuapsaltria, numbers refer to percentages of trees supporting the branch; (b), alternative phylogeneticreconstruction for the generaGuineapsaltria andPapuapsaltria,with a paraphyletic genus Papuapsaltria.

maining species as sister group of these two groups however. This is demonstratedby the fact that its combined. In Fig. 12a, Papuapsaltria is practically position in the paraphyletic construction as either monophyletic. the sister group of Guineapsaltria, or of toxopei-

in In both these options, which are principle dolabrata, or of phylophora, or of Guineapsal-

dioedes is excluded of regarded equally plausible, P. tria-phyllophora, or of goniodes-ustulata, or from Papuapsaltria and figures as sister taxon to all goniodes-woodlarkensis, is equally parsimonious. the above-discussed genera together (cf. Fig. 3). P. The two most parsimonious alternatives (Figs. 12a, dioedes differs strongly from other Papuapsaltria b) show several other differences with the earlier species by several presumably autapomorphous cladogram (De Boer, 1995c). In both alternatives, male genital characters, but the species does share and in the strict consensus tree, P. goniodes and P. the reduced femoral and broader presumed apomorphous spine brassi (based on five tymbal ridges a and certainly shouldbe includedsomewhere within vertex) and P. angulata and P. lachlani (based on

Papuapsaltria. Besides, the suggested monophyly a slender male caudodorsal beak) are sister species, of the group including all genera discussed above while in the previous cladogram (De Boer, 1995c) and dioedes based the similar excluding P. is on presumed P. goniodes and P. angulata (based on a apomorphous but strongly fluctuating character of male operculum) and P. lachlani and P. brassi a broadly rounded pygofer opening, which certain- (based on an extremely long aedeagus) are sup- ly does not characterise that group as a whole and posed to be sister species. The latter character is is considered meaningless at this level of the clado- difficult to quantify, and was not included in the

dioedes in in matrix. is gram. Including P. Papuapsaltria a Still, this latter option preferred here, most parsimonious position costs 3 extra steps in although it is two steps longer in both alternatives both the monophyletic alternative (Fig. 4) and the (Figs. 4, 12c). paraphyletic solution (Fig. 12c). The northwesternNew Guineasubgroup that was

The position of P. dioedes is very uncertain, recognized before (P.i baast, P. dolabrata, P. nova- Contributions to Zoology, 65 (4) - 1995 217

the most of P. G. flava changes, parsimonious position

G. fla veola is the base of phyllophora no longer at Guineapsal-

G. stylata tria, but at the base of Papuapsaltria. G. pennyi

S. pallida

G. pallidula Aedeastria G. viridula

G. chinai The most parsimonious reconstruction of Aedeas-

P. toxopei tria in the50% majority rule tree (Fig. 13) is almost P. novariae

but P. dolabrata identical to that in the strict consensus tree,

P. baasi completely differs from earlier conclusions on its P. ustulata

P. phyllophora phylogeny (De Boer, 1993a). The earlier proposed

P. dioedes monophyly of a group consisting ofA. digitata, A. P. goniodes and P. angulata cobrops, A. kaiensis, A. latifrons, A. sepia, A.

P. brassi waigeuensis which was recognized on the basis of a P. lachlani

is not P. spinigera unique bicuspidate clasper (De Boer, 1993a),

P. stoliodes recognized in the computer analysis (in that earlier P. plicata discussion A. cheesmanae was abu- P. nana phylogenetic

P. ungula sively exchanged for A. waigeuensis). Instead, the

P. bidigitula of the form a P. woodlarkensis remaining species genus monophylet-

and ic group, based on a hook-shaped apically

12. Most McClade reconstruction for the Fig. (c), parsimonious which pointed clasper, a character frequently oc- generaGuineapsaltria and Papuapsaltriain the alternative ver- curs in many other genera. Converting the above- sion with a paraphyletic genus Papuapsaltria,and after includ- listed species into a monophyletic as ing the modifications suggested. group pro- in the then posed here costs 7 extra steps most par- riae, P. ustulata) is recognized in the paraphyletic simonious solution (Fig. 4), but only 6 extra steps option but not in the monophyletic one. In the lat- in the reconstruction with a paraphyletic genus ter option P. nana and P. bidigitula are included in Papuapsaltria which gives a slightly different solu-

is in most that group, based on an in other parts of the tree tion (Fig. 14). A. hastulata a parsimoni- strongly variable character of the postclypeus. In ous solution part of this group in both cases. both these options, the likewise northwestern New The position of Aedeastria, either as sister group

Guinea species P. toxopei is included in this group. of Mirabilopsaltria and Thaumastopsaltria (based

shared wrinkled with distinct In the cladogram presented earlier (De Boer, 1995c) on a vertex diverging

included in 1 in sister of all above- P. nana and P. bidigitula were a fissures, Fig. 11), or as group monophyletic group with P. plicata, P. spinigera, discussed genera together (based on a shared erect

with sister femoral in discussed in several P. stoliodes, and P. ungula, P. nana as spine, 1 Fig. 15) was species of P. ungula, which was based on several previous publications (De Boer, 1995a, b, c, d). The convincing apomorphies in the clasper. Removing most parsimonious tree unambiguously recognized

credi- of these species, as proposed here, to their more Aedeastria as the sister group Mirabilopsaltria

inboth the and but Aedeastria ble position minimally costs 2 extra steps Thaumastopsaltria, changing to monophyletic option (Fig. 4) and the paraphyletic a sister-group position of all above-discussed gen-

make the total one (Fig. 12c). era (see Fig. 15) will tree only 1 step

The phylogenetic position of P. woodlarkensis longer and should still be considered a possibility. remains dubious; in the paraphyletic option the

is included in the and in the species Papuan group, Thaumastopsaltria monophyletic option in the northwestern New The cladogram of Thaumastopsaltria in the 50%

its rule is solved and in Guinea group, but after the proposed changes majority tree (Fig. 13) fully most parsimonious position is in the Papuan group agreement with previous discussions on the phy- in the latteroption as well. Furthermore, after these togeny of this genus (De Boer, 1992a). de - and taxonomic 218 A.J. Boer Phytogeny status of Chtorocystini s. s

A. cobrops Baeturia

A. sepia Scottotympana

A. latifrons Gymnotympana

A. kaiensis

Venustria A. hastulata

Chlorocysta A. waigeuensis

A. cheesmanae Owra

A. bullata Glaucopsaltria

A. dilobata Guineapsaltria

A. moluccensis Papuapsaltria

A. obiensis Aedeastria

A. digitata Mirabilopsaltria T. pneumatica Thaumastopsaltria T. adipata

Cystopsaltria T. spelunca

T. sicula Cystosoma

T. globosa Fig. 15. Generic cladogram of the Chlorocystini with an alterna- T. sarissa tive position for Aedeastria. Number refers to character dis-

T. lanceola cussed in the text.

M. viridicata

M. globulata

M. inflata morphy in clasper shape is not present in M.

M. inconspicua viridicata and, though possibly changed beyond

M. humilis This recognition, in M. globulata. presumed apo- M. toxopeusi morphy is not recognized in the most parsimonious

whole and for Fig. 13. Section ofthe 50% majority rule tree dealingwith Aede- tree, not for Mirabilopsaltria as a not astria,Thaumastopsaltria, and Mirabilopsaltria. Numbers refer the species that do have that character. In the 50% to percentages of trees supporting the branch. majority rule tree (Fig. 13) the species of Mirabi-

sister of lopsaltria appear as a paraphyletic group

A. cobrops Thaumastopsaltria. The characters in support of A. sepia this paraphyletic reconstruction are, with a possible A. latifrons exception of the one determining M. viridicata and A. kaiensis M. globulata as the immediate sister group of A. hastulata Thaumastopsaltria (an extremely long 1st apical A. waigeuensis considered less than areain the tegmen), convincing A. digitata the clasper character. M. humilis, M. inconspicua, A. bullata

M. inflata, and M. toxopeusi are still supposed to A. dilobata

form a This is A. moluccensis monophyletic group (Fig. 4). option

A. cheesmanae proposed here and makes the tree 2 steps longer.

A. obiensis Although the inclusion of M. viridicata and M.

globulata into a monophyletic genus Mirabilopsal- Fig. 14. Most parsimonious McClade reconstruction of Aede- tria is still a possibility to be considered, depending astria in the alternative version with a paraphyletic genus Papu- on the possible homology of the M. globulata apsaltria, and after including the modifications suggested. clasper with that of the other species of Mirabi-

lopsaltria, their sister-group relationship to Thau-

Mirabilopsaltria mastopsaltria now seems more likely. Including

The monophyly of Mirabilopsaltria was considered these species in Mirabilopsaltria wouldcost no less uncertain (De Boer, 1995b). The presumed apo- than 4 extra steps. Contributions to Zoology, 65 (4) - 1995 219

Cystopsaltria and Cystosoma in the Chlorocystini (sensu stricto) are caused by

the of The basal position of Cystopsaltria and Cystosoma vicariance following fragmentation a histori- in the most parsimonious reconstruction (Fig. 3) is cal volcanic island arc (De Boer, 1995d). The phy- caused by the several characters these genera share logenetic reconstruction with a paraphyletic genus with the outgroup Prasia. The there suggested Papuapsaltria (Fig. 12c) which separates the genus

of western monophyly the Chlorocystini excluding Cys- roughly in a and eastern species group,

and is based two to best with the historical topsaltria Cystosoma on very un- appears agree explana- convincing characters that cannot be regarded as tions provided in that publication. Similarly, a representative for that group. In previous discus- paraphyletic option for the genus Mirabilopsaltria, sions of which the solu- on the phylogeny the Chlorocystini (De turns out to be most parsimonious

Boer, 1995a, b, c, d) it was suggested that Cys- tion (Fig. 4), better fits the historical explanations.

topsaltria and Cystosoma formed the sister group Furthermore, a position of Guineapsaltria-Papua- of to Mirabilopsaltria and Thaumastopsaltria, either psaltria as sister group the group Baeturia-

Aedeastria based of which the is discussed or not including (see above) on Venustria, possibility the narrow tymbal cavity shared by Cystosoma and above (cf. Fig. 11) is in biogeographical sense most species of Mirabilopsaltria and Thaumas- slightly more parsimonious thanthe option given in

this since the latter the extinction of topsaltriar; solution is still regarded as more Fig. 4, requires a credible. Moving Cystopsaltria and Cystosoma to group in a missing area (De Boer, 1995d). the base ofAedeastria, Mirabilopsaltria, and Thau- mastopsaltria as proposed here costs only 1 step more (Fig. 4). The ingroup phylogeny of Cystopsal- Acknowledgements tria and Cystosoma given in Fig. 4 is identical to

I that of the strict consensus tree. am most gratefulto Dr. W.N. Ellis for running the PAUP ana-

lyses on his computer. Prof. F.R. Schram, Dr. R. Sluys, and Dr.

J.P. Duffels are thanked for their critical reading and comments

I Mr. D.A. on the manuscript. thank Langerak for his help in Conclusions preparing the figures.

The final tree which contains all of the presumed synapomorphies and deals with theremaining char- References acters in a most parsimonious way is still considered to have two equally likely solutions for the genus Boer, A.J. de, 1982. The taxonomy and biogeography of the

The with Papuapsaltria (Figs. 4, 12c). option a nasuta group of the genus Baeturia Stài, 1866 (Homoptera,

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these alternatives differ in most bloetei of the Baeturia 1866 gram only slightly a group genus Stài, (Homoptera, parsimonious reconstruction of Aedeastria and the Tibicinidae). Beaufortia, 39(1): 1-43.

A.J. 1990. cicada from Boer, de, Aedeastria, a new genus New Baeturia nasuta group. Guinea, its phylogeny and biogeography (Homoptera, The results of this phylogenetic analysis, together Tibicinidae), preceded by a discussion on the taxonomy of with the biogeographical data discussed in previous New Guinean Tibicinidae. Beaufortia, 40(3): 63-72.

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New Guinea, with the description of three new species, their vide the data for an area cladistic study of the New

and Guinea-West taxonomy biogeography (Homoptera, Tibicinidae). Pacific region. A comparison of area

Beaufortia, 42(1): 1 — 11. cladograms with data on the palaeogeology of the Boer, A.J. de, 1992a. The taxonomy and biogeography of the area in question suggests that the main speciation genus Thaumastopsaltria Kirkaldy, 1900 (Homoptera, Tibi- events that led to the present-day generic diversity cinidae). Beaufortia, 43(3): 17-44. - 220 A.J. de Boer Phytogeny and taxonomic status of Chlorocystini s. s.

Boer, A.J. de, 1992b. The taxonomy and biogeography of the norrhyncha). Acta ent. Mus. natn. Pragae, 32: 317-352.

1866 viridis group of the genus Baeturia Stài, (Homoptera, Dlabola, J., 1960. Cicadidae (Hemiptera, Homoptera, Auche-

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of the Aedeastria Witte, 59(2): 9-37. Boer, A.J. de, 1993a. Ten new species genus

with the and 1993. The of Moana De Boer, 1990, notes on taxonomy biogeography Duffels, J.P., systematic position expansa

(Homoptera, Tibicinidae). Beaufortia, 43(9): 140-167. (Homoptera,Cicadidae), with reference to sound organs and

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63(1): 15-41. composite areas in East Malesia. In: P. Baas, C. Kalkman &

Boer, A.J. de, 1994a. The taxonomy and biogeography of the R. Geesink (eds.), The plant diversity of East Malesia;

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Tibicinidae). Beaufortia, 44(5): 127-158. memorating Professor Dr. C.G.G.J, van Steenis, Leiden,

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Boer, A.J. de, 1994c. The taxonomy and biogeography of the Cicadoidea (Homoptera, Auchenorhyncha) 1956-1980. Se-

loriae group of the genus Baeturia Stài, 1866 (Homoptera, ries Entomologica, 33: i-xvi, 1-414.

Tibicinidae). Tijdschr. Ent., 137: 1-26. Farris, J.S., 1969. A successive approximations approach to

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tera, Tibicinidae). Tijdschr. Ent., 137: 161-172. schen Systematik: 1-370 (Deutscher Zentralverlag, Berlin).

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phy of the cicada genus Gymnotympana Stài, 1861 (Homop- of Illinois Press, Urbana).

tera: Tibicinidae). Invert. Taxon., 9(1): 1—81. Jong, M.R. de, 1982. The Australian species of the genus

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New Guinea, its taxonomy and biogeography (Homoptera, Dierk., 52(2): 175-185.

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Boer, A.J. de, 1995c. The taxonomy and biogeography of the Prasiini 1: The genus Prasia Stài, 1863 (Homoptera, Tibicini-

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Cicadidae from the Belgian Congo (Homoptera, Auche- Mag. nat. Hist., (12) 7: 233-237, pl. V. Contributions to Zoology, 65 (4) - 1995 221

Orian, A.J. E., 1964. The morphology of the male genitalia of 15 Pronotum: (1) without coloured medial band, (2)with dark

Abricta ferruginosa(Stài) (Homoptera, Cicadidae). Proc. R. streaks along light medial band, (3) with darkened medial

ent. Soc. London, (A) 39(1-3): 1-4. band

Swofford, D.L., 1993. PAUP: Phylogenetic Analysis Using 16 Pronotum: (1) without distinct medial furrow, (2) with dis-

version 3.1.1 Parsimony, (computer program distributed by tinct medial furrow

the Illinois Natural History Survey, Champaign, Illinois). 17 Anterolateral side of pronotal collar: (1) rounded, (2)

Bavea Villet, M.H., 1993. The cicada genus Distant, 1905 swollen, (3) angularly bent, (4) forming continuous ridge

(Homoptera, Tibicinidae): redescription, distribution and

phylogenetic affinities. Trop. Zool., 6(2): 435-440. Mesonotum

18 Mesonotal colouring: (1) without spots in front of cruci-

Received: 24 1995 July form elevation, (2) two black spots in front of cruciform

Revised: 12 October 1995 elevation

19 Cruciform elevation: (1) wider than long, (2) narrower than

long

Appendix 1. List of characters Legs

20 Proximal spine of fore femur: (1) erect, (2) bent adjacent to

Colour femur

1 colour: with brown 21 Body (1) unspeckled, (2) speckled Proximal spine of fore femur: (1) of normal length, (2) ex-

2 Colour of female: (1) uniformly coloured, (2) bicoloured: tremely long, (3) reduced

and head thorax green, abdomen ochraceous 22 Proximal spine of fore femur: (1) not shorter than distance

to middle spine, (2) distinctly shorter than distance to mid-

Head dle spine, (3) slightly shorter than distance to middle spine

3 with anterior Postclypeus: (1) anteriorly rounded, (2) sharp 23 Middle spine of fore femur: (1) narrow spiny, (2) broad tri-

edge angular

4 Anterior margin of postclypeus in dorsal view: (1) con-

tinuous with anterior margins of vertex lobes, (2) convexly Tegmina and wings

vertex protruding beyond lobes, (3) strongly curving back 24 Colour of tegmen: (1) hyaline, (2) opaque greenish or red-

laterally and making almost right angle with vertex lobes dish, (3) slightly reddish but still hyaline, (4) bronzed

5 Postclypeus in lateral view: (1) not swollen, (2) angularly 25 Colour markings on tegmen: (1) immaculate, (2) brown

swollen, (3) globularlyswollen, (4) protrudingin obtuse an- spots along veins, (3) triangular brown spots at ends of api-

gle, (5) with convex anterior margin cal areas

6 Lateral sides of without 26 of rounded postclypeus: (1) posterior ridge Shape tegmen: (1) at apex, (2) pointed at apex

along lorum, (2) with swollen posterior ridge along lorum 27 Tegmen venation: (1) not reticulate, (2) reticulate

7 28 Lateral sides of postclypeus: (1) with distinct rasps of rows Hyaline hind margin of tegmen: (1) very narrow, (2)narrow

of short parallel ridges, (2) with very short and indistinct but distinct, (3) broad

rows of without of 29 of parallel ridges, (3) rows parallel ridges Costal area tegmen: (1) narrow to apex, (2) widening to

8 Vertex: (1) without dark spots, (2) with dark spots between apex

and lateral ocelli eyes 30 Number of apical areas in tegmen: (1) eight, (2) eight or

9 Medial part of vertex: (1) distinctly rising from postclypeus, nine, (3) nine, (4) nine or ten, (5) variable between nine and

indented (2) very flat, (3) slightly fifteen, (6) more than twenty

10 Surface of vertex: (1) smooth, (2) with distinct fissures 31 Ultimate apical area of tegmen: (1) quadrangular, (2) tri-

from centre to diverging of head corners of postclypeus angular

11 Distance between lateral ocelli: 32 (1) hardly exceeding width Subapical areas of tegmen: (1) absent, (2) present but not

of frontal ocellus, (2) distinctly larger than width of frontal forming a continuous band, (3) forming a continuous band,

ocellus (?) not clear due to reticulate venation

12 Distance 33 between lateral ocelli: (1) about equalling distance Base of first apical area of tegmen: (1) distally of base of

between lateral ocellus and eye, (2) generallyslightly larger third apical area, (2) slightly proximallyof base of third api-

than distance between lateral ocellus and eye, (3) generally cal area, (3) distinctly proximally of base of third apical

than distinctly larger distance between lateral ocellus and area

eye 34 Veins of tegmen: (1) without or practically without setae,

(2) densely set with setae

Pronotum 35 Number of apical areas in wing: (1) six, (2) five, (3) six or

13 Pronotal surface: (1) smooth, (2) mottled seven, (4) seven or more

14 Pronotum: without (1) dark medial spot, (2) with dark 36 Sexual dimorphism in anal fields of wing: (1) absent, (2)

medial spot at pronotal collar present 222 A.J. de Boer - Phytogeny and taxonomic status of Chlorocystini s. s.

Tymbal organ swollen but folded inwards along tymbal, (3) forming a dis-

37 Tymbal: (1) parallel to lateral side of abdomen, (2) strongly tinct ridge along tymbal

curved inwards towards its ventral corner 61 Second tergite of male: (1) forming a distinct ridge along

38 Number of tymbal ridges: (1) four ridges or less, (2) five tymbal cavity, (2) not forming a distinct ridge along tymbal

cavity ridges, (3) six ridges, (4) sevenridges, (5) eight ridges, (6)ten

62 Second tergite of male: not membranous, (2) mem- ridges, (7) eleven ridges or more (1)

branous 39 Tymbal ridges: (1) parallel, (2) aberrant opposite tymbal

63 Second tergite of male: (1) not swollen, (2) swollen along 40 Tymbal ridges: (1) not curved around intercalary ridges, (2)

curved around intercalary ridges tymbal cavity

64 Eighth tergite of male: (1) not forming ventral lobe, (2) 41 Tymbal cavity: (1) broad, (2) narrow forming small posteriorly projecting ventral lobe

65 Male swollen and Opercula auditory capsule: (1) protruding, (2) not

swollen and not protruding 42 Male opercula: (1) widely separated, (2) juxtaposed

66 Pygofer and 9th sternite: (1) resting on 8th sternite, (2) medially elevated respective to 8th sternite 43 Male operculum: (1) longer than meracanthus, (2) shorter 67 Connectingbar between tymbal and abdomen: (1) not elon- than meracanthus

gate, (2) strongly elongate 44 Male operculum: ( 1 ) notcompletelycovering tymbal cavity, 68 Connecting bar between tymbal and abdomen: (1) parallel (2) completely covering tymbal cavity to body axis, (2) directed mesiad 45 Male operculum: (1) not elongate at medial margin, (2)

elongate at medial margin

Male 46 Basal part of male operculum: (1) distinctly vaulted, (2) pygofer

strongly curved down medially, (3) hardly vaulted 69 Pygofer: (1) not curved to ventral position, (2) curved to

ventral abdomen 47 Basal part of male operculum: (1) almost oblong, (2) later- position on

70 without lateral with distinct fold ally longer than medially,(3) laterally shorter than medially Pygofer: (1) fold, (2) at

base of caudodorsal beak and abruptlylengthening,(4)laterally shorter than medially

71 Male caudodorsal beak 1 and anal and graduallylengthening : ( ) erect not reaching over

valves, (2) curved and extending over anal valves, (3) absent 48 Lateral crest of operculum base: (1)normally developed,(2)

72 Male caudodorsal beak: (1) not covering anal valves, (2) very long and slender strongly curved and covering anal valves 49 of distal ofmale very Shape part operculum: (1) angularlyoval,

73 Male caudodorsal beak: (1) erect, (2) angularly bent and (2) squarely rounded, (3)blunt triangularand flat, (4) blunt dorsally flattened, (3) bent down towards apices of lateral triangular with straight lateral margin, (5) triangular and lobes, (4) strongly rounded; continuously rounded with domed, (6) small and rounded, (7) small and angular

dorsal margin of pygofer, (5) strongly rounded but not con- 50 Lateral margin of male operculum: (1) not continuous to tinuous with dorsal margin of pygofer, (6) weakly curved distolateral corner of operculum base, (2)continuous to dis- 74 Male caudodorsal beak: (1) straight to apex, (2) concaveto tolateral corner of operculum base

apex 1 51 Distomedial margin ofmale operculum: ( ) without recurv- 75 Apex of male caudodorsal beak: (1) sharply pointed, (2) ing rim, (2) with recurving rim rounded, (3) truncate or weakly concave, (4) bicuspidate, 52 Medial margin ofmale operculum: (1)medially ofmeracan- (5) broadly rounded, (6) beak missing thus, (2) laterally of meracanthus 76 Male caudodorsal beak in lateral view: (1) stout, (2) slender 53 Meracanthus in male: (1) normally developed, (2) strongly 77 Male caudodorsal beak: (1) broad at base, (2) narrow ob- reduced long, (3) spine-shaped 54 Distal part of female operculum: (1) rectangularoblong, (2) 78 Male caudodorsal beak: (1) hollow,reversed gutter-shaped, trapezoid, (3) narrow sickle-shaped, (4) broad sickle- (2) massive continuous distolateral of basal shaped, (5) to corner part, 79 Male caudodorsal beak in dorsal view: (1) oblong with (6) triangular almost lateral with parallel margins, (2) triangular converg-

ing margins Abdomen 80 Protuberance on lateral lobe of pygofer: (1) narrow lobate 55 Male abdomen: (1) not transparent, (2) transparent or laminiform,(2) extremely broad and lobate 56 Male abdomen: (1) rounded lateroventrally, (2) tergites 81 Protuberance onlateral lobe of pygofer: (1) not conically sharply folded lateroventrally elongate beyond pygofer margin, (2) conically elongate be-

57 Abdominal sternites: without medial with red (1) band, (2) yond pygofer margin

medial band 82 Protuberance on lateral lobe of pygofer: (1) not triangularly

58 First sternite of male: (1)short; broader than long, (2) long; swollen, (2) triangularly swollen .ìutoi

longer than broad 83 Protuberance on lateral lobe of pygofer: (1) rouitdéü, (2)

59 Eighth sternite of male: (1) not sharply incised, (2) sharply with sharp lateral edge -oil

incised at distal margin 84 Protuberance on lateral lobe of pygofer: (1) straightly

60 Second tergite of male: (1) swollen along tymbal, (2) not directed posteriad, (2) upcurving towards apex Contributions to Zoology, 65 (4) - 1995 223

85 Distal margin ofpygofer: (1) curving to base of lateral pro- 110 Proximal parts of claspers: (1) not merged between aedea-

of lateral and anal between and anal tuberance, (2) curving to apex protuberance gus valves, (2) merged aedeagus

86 Pygofer: (1)not broadeningbetween lateral protuberances, valves

(2) broadening between lateral protuberances 111 Dorsal part of clasper: (1) not forming a shaft around ae-

87 Pygofer: (1) not narrowing between lateral protuberances, deagus, (2) forming a short shaft around aedeagus

(2) narrowing between lateral protuberances 112 Claspers: (1)without sharp and converging dorsal edges, (2)

88 Distal margin of pygofer: (1) weakly convex, (2) strongly with sharp and converging dorsal edges

convexinto a distinct posteriorlobe, (3) weakly concave, (4) 113 Dorsal crest of clasper: (1) notangularlyprotruding, (2) an-

strongly concave gularly protrudingat distal corner

89 Ventral part of pygofer: (1) not or hardly protruding, (2) 114 Dorsal margin of clasper: (1) not rectangularly bent, (2)

forming a rectangular corner under lateral protuberance, rectangularly bent at half-length

(3) forming a thorn-shapedprotuberance under lateral pro- 115 Clasper: (1) without dorsal crest, (2) with short dorsal crest

tuberance on proximal half only, (3) with dorsal crest over almost

90 Pygofer: (1) not swollen at ventral margin, (2) swollen at whole length

ventral margin 116 Clasper: (1) with no or straight dorsal crest, (2) with convex

91 Ventral margin of pygofer: (1) straightto base, (2) strongly dorsal crest

117 concave to base Dorsal crest of clasper: (1) erect or absent, (2) curving

92 Pygofer opening: (1) without straight basal margin,(2) with laterad

straight basal margin 118 Dorsal crest of clasper: (1) absent or not forming dorsal

93 half of Ventral pygofer opening: (1) not bordered by a con- protrusion, (2) forming dorsal protrusion

tinuous sharp edge, (2) bordered by a continuous sharp 119 Dorsal protrusion of clasper: (1) absent, (2) broad lamini-

edge form, (3) narrow finger-shaped, (4) humped

94 Ventral half of pygofer opening: (1) not horseshoe-shaped, 120 Dorsal edge of clasper: (1) not curved around aedeagus, (2)

(2) horseshoe-shaped with incurving lateral corners forminga rectangular corner around aedeagus, (3) forming

95 Ventral half of pygofer opening: (1) triangular and pointed, a rectangular crest around aedeagus, (4) forming a distinct-

(2)broad and rounded, (3) with concavemargins and point- ly swollen protuberance around aedeagus, (5) globularly

ed, (4) very long and slender rounded

96 Basal margin of pygofer opening: (1) not lowered, (2) 121 Apical part of clasper: (1) without proximal protrusion,

lowered with proximal protrusion

97 Anal valves: (1) not swollen, (2) swollen 122 Clasper apex: (1) not swollen, (2) swollen

123 Clasper apex: (1) not globularly swollen, (2) globularly

Clasper swollen

98 Claspers: (1) fused or practically fused to a more or less 124 Clasperapex: (1) broad in dorsal view, (2) abruptly narrow-

continuous ring around base of anal valves, (2) distinctly ing in dorsal view

separated at base 125 Clasper apex: (1) pointed or rounded, (2) bicuspidate or

99 Clasper: (1) without lateral plate, (2) with lateral plate dentate

100 Clasper: (1) strongly bent and hook-shaped, (2) nearly 126 Apical part of clasper in lateral view: (1) not broader than

straight and posteriorly directed basal part, (2) broader than basal part

101 Clasper: (1) not triangular, (2) triangle-shaped in lateral 127 Clasper: (1) without small triangle-shaped apical part, (2)

view with small very triangle-shaped apical part

102 around 128 Clasper: (1) not forminga shaft aedeagus, (2) form- Clasper hollow: (1) long and strongly bent, (2) long and

ing a shaft around aedeagus straight, (3) short and straight, (4) short and strongly bent,

103 Claspers: (1) parallel, (2) diverging towards apices (5) short and downcurved, (6) different

104 Clasper: (1) not rectangular laminiform, (2) rectangular 129 Clasper: (1) without ventromedial protuberance, (2) with

laminiform broad and laminiform ventromedial protuberance, (3) with

105 Clasper: (1) not transparent ventromedially, (2) ven- narrow and spiny ventromedial protuberance

tromedially transparent 130 Clasper: (1) without distinct rim around ventral hollow, (2)

rim 106 Clasper: (1) without distinct cross fold, (2) with distinct with distinct around ventral hollow

fold cross between basal part and apical part 131 Clasper: (1) without lateral process, (2) with lobate lateral

107 base: without with lateral with lateral Clasper (1) crest protruding beyond pygofer process, (3) thorn, (4) crest ending

margin, (2) with distinct lateral crest protruding beyond in small thorn

pygofer margin

108 Clasper base: (1) not forming clasper heel, (2) forming dis- Aedeagus

tinct angular dorsal corner or clasper heel 132 Aedeagus: (1) S-curved, (2) C-curved

109 Clasper: (1) without dorsally protruding clasper heel, (2) 133 Aedeagus: (1) not Z-curved in apical part, (2) Z-curved in

with dorsally protrudingclasper heel apical part 224 A.J. de Boer - Phytogeny and taxonomic status of Chlorocystini s. s.

the 134 Surface of aedeagus: (1) smooth, (2) with pattern of con- Appendix 2. List of presumed apomorphies, black squares

centric ridges in Fig. 4.

135 Orientation of aedeagus: (1) more or less erect between

nr. Homoplasies (H) and claspers, (2) directed posteriad parallel to and between Internode Character reversals

claspers and state (R)

with with lateral 136 Aedeagus: (1) lateral crests, (2) lobes, (3)

4-3 119-2 with elongate lateral lobes

5-4 119—(2-1-3+ 4) 137 Lateral lobes of aedeagus: (1) straight or absent, (2) strong-

153-4 H in B. O. ly curving laterad edauberti, insignis

7-5 118-2 138 Lateral crest of aedeagus: (1) graduallytaperingtowards ae-

9-8 5-2 H in ,B. bemmeleni, B. daviesi, deagalapex, (2) ending in rectangular corner

B. guttulinervis, B. hamiltoni, 139 Aedeagus: (1) without crest between lateral lobes, (2) with

B. pigrami, B. wauensis crest between lateral lobes

38-1 H in internode 66-65 140 Aedeagus: (1) not swollen between lobes, (2) swollen be-

16-9 55-2 R in B. marmorata tween lobes

H in internode 67-66 141 Aedeagus part distally of basal curve: (1) not long and

73-2 slender, (2) extremely long and slender

141-2 H in internode 52-51 142 Apical part of aedeagus: (1) not abruptly narrowing, (2)

15-11 116-2 H in internode 53-50 abruptly narrowing and upcurving, (3) abruptly narrowing

13-12 75-4 H in internode 99-97 and downcurving

14-13 140-2 143 Aedeagal apex: (1) not elongated dorsally, (2) distinctly

15-14 73-3 H in B. elongated dorsally, (3) slightly elongated dorsally furcillata

16-15 83-2 H in B. inconstans 144 Aedeagus: (1) without distinct subapical lobe, (2) with dis-

107-2 H in B. inconstans tinct subapical lobe

25-16 79-1 H in B. guttulipennis 145 Aedeagus: (1) without dorsal crests, (2) with single dorsal

with dorsal with of dor- 20-19 137-2 crest, (3) paired crests, (4) pair

24-20 38-3 H in solateral protuberances many groups

23-22 130-2 146 Aedeagus: (1) without ventral crest, (2) with ventral crest or

24-23 123-2 keel

25-24 73-4 H in A. digitata, B. furcillata, 147 Aedeagus from behind: (1) not bottle-shaped, (2) bottle-

B. & internode 35-34 towards gressitti shaped; abruptly narrowing apex

122-2 148 curved and Aedeagus: (1) strongly tapering to apex, (2) 153-3 H in B. gibberosa strongly curved and not taperingto apex, (3)weakly curved

40-25 81-2 R in node 18-17 149 Aedeagus: (1) not strongly swollen proximally, (2) strongly

35-34 73-4 H in A. B. swollen proximally digitata, furcillata,

B. gressitti & internode 25-24 150 Aedeagalpore: (1) narrow oval, (2) triangular and widening

36-35 124-2 to broadly rounded proximally, (3) apically pointedbut not

39-38 150-1 H in B. and widening proximally, (4) broad oval, (5) round papuensis many

groups of Baeturia 151 Margin of aedeagalpore in lateral view: (1) weakly concave,

40-39 49-2 H in fi. rossi continuous with curve of aedeagus, (2) strongly concave, 113-2 not to 44-43 continuous with curve of aedeagus, (3) convex, due 30-4 dorsoventral incision, (4) straight 46-45 22-3 H in B. karkarensis, B. rufa,

B. turgida

47-46 87-2

Female genitalia 117-2

H 152 Ovipositor sheaths: (1) reaching to apex of caudodorsal 53-50 116-2 in internode 15-11

beak, (2) reaching just beyond apex of caudodorsal beak, 52-51 25-3

H (3) reaching far beyond apex of caudodorsal beak 144-2 in internode 16—9

153 Apex of female caudodorsal beak: (1) sharply pointed, (2) 53-52 126-2

broadlyrounded, (3) narrowly rounded, (4)truncate, (5) bi- 136-3

cuspidale 56-53 109-2 H in B. tenuispina

154 Femalepygofer: (1) without ventral thorn, (2) with ventral 56-55 49-3 H in G. hirsuta, G. olivacea

thorn 57-56 36-2 H in internode 80-79 & Prasia Contributions to Zoology, 65 (4) - 1995 225

Appendix 2. Continued.

Internode Character nr. Homoplasies (H) and reversals Internode Character nr. Homoplasies (H) and reversals

and state (R) and state (R)

91-2 H in G. obiensis, G. olivacea, P. goniodes & internodes

S. huibregtsae, V. superba 73-72 & 131-129

60-57 23-2 H in G. rubricata 32-3

136-(2+3) 47-4 H in C. immaculata

148-1 65-2 H in internode 131-129

150—(1-4) H in A. waigeuensis, A. 120-2 H in internode 104-103

obiensis, internodes 73-72 & 93-88 95-3

127-126 110-2 H in A. latifrons

151—(1 +2) 135-2

59-58 88-2 H in internode 110-109 92-91 6-2 H in V. superba & internode

99-2 138-137

60-59 46-2 93-92 101-2 R in G. pennyi

64-2 111-93 144-2

121-2 96-94 45-2 H in G. chinai, P. dioedes

138-2 103-96 94-2

81-60 98-1 H in A. dilobata 104-2

69-62 120-4 H in G. varicolor, A. hastula- 99-97 52-1 H in M. globulata& inter-

la & internode 99-98 node 86-81

64-63 60-3 70-2

62-2 75-4 H in internode 13-12

67-64 89-3 in woodlarkensis H P. 99-98 120-4 H in A. hastulata, G.

66-65 38-1 H in internode 8-9 varicolor & internode 69-62

40-2 100-99 105-2 R in P. plicata

67-66 48 -2 102-101 131-3 H in internode 129-124

55-2 H in internode 16-9 104-103 88-2 H in internode 59-58

106-2 120-2 H in internode 86—85

69-68 142-3 109-104 88-2 H in internode 59-58

71-70 R in P. 49-4 P. bidigitula, nana, P.

72-71 17-4 H in M. humilis Prasia & in- woodlarkensis ,

ternodes 131-130 & 139-138 108-107 111-2

73-72 B. R P. P. 30-5 H in inconstans, M. infla- 110-109 21-3 in P. bidigitula, nana,

ta, P. goniodes& internodes woodlarkensis

86-85 & 131-129 112-111 49-6 R in several species of Papua-

31-2 R in G. olivacea psaltria

32-2 H in M. inflata, M. globulata 140-112 16-1 H in M. toxopeusi

& internode 129-128 20-1 H in A edeast ria, M. viridica-

150-4 H in A. obiensis, A. ta, T. sarissa

waigeuensis, G. varicolor 28-3 R in G. stylata, O. insignis,

76-75 127-2 many Papuapsaltria species

77-76 148-2 further evolved? in internode

79-77 37-2 R in G. dahli, G. olivacea 60-57. H in Muda

79-78 47-3 H in V. superba 114-113 71-3 H in A. cheesmanae

57-2 134-2

80-79 36-2 H in Prasia & internode 149-2

57-56 123-118 125-2 R in A. hastulata

86-81 52-1 H in M. globulata, P. plicata, H in T. sicula

P. stoliodes 136-123 17-2

83-82 38-7 127-126 129-2

84-83 133-2 129-128 129—(2 + 3) H in A. hastulata

85-84 35-2 H in P. dolabrata 32-2 H in M. inflata, M. globulata

86-85 30-5 H in B. inconstans, M. infla- & internode 73-72

ta, 226 A.J. de Boer - and Phytogeny taxonomicstatus of Chlorocystini s. s.

Appendix 2. Continued.

Internode Character nr. Homoplasies (H) and reversals Internode Character nr. Homoplasies (H) and reversals

and state (R) and state (R)

R in T. sicula 17-4 H in M. humilis, Prasia, inter-

& 131-129 5-4 H in Prasia & internode nodes 72-71 131-130

13-2 139-138 H in Prasia

19-2 H in Prasia 30-5 H in B. inconstans, M. inflata,

26-2 P. goniodes& internodes

27-2 73-72 & 86-85

140-139 41-2 R in T. spelunca, T. sicula, 90-2 R in T. adipata, T. lanceola M.

C. immaculata & in- 152-3 H in G. rubricata, M. globula- globulata,

ternode 136-123 ta, Muda

131-130 17-4 H in M. humilis, Prasia, inter- Remarks. - Character 41-2 is H in the PAUP reconstruction; nodes 72-71 & 139-138 character 43-3 is placed at the base of Gymnotympana and 135-131 33-3

Venustria; character 73-2 is placed oninternode 47-40 by the 135-134 112-2 PAUP reconstruction; character 150-1,regarded apomorphous 136-135 61-2 R in M. globulata, M. toxo- for the Bismarck species ofthe bloetei group, is regarded plesio- peusi & internode 127-126 for that in the PAUP character morphous group reconstruction; 139-136 10-2 R in A. bullata, A. digitata, A. 153-4 is placed on internode 9-8 by the PAUP reconstruction, dilobata, M. toxopeusi, M. presumably due to the interpretation of lacking information. viridicata. H in G.

stenocephalis

139-138 5-4 H in Prasia & internode

131-129

9-3 - Contributions to Zoology, 65 (4) 1995 227

= = = = Appendix 3. Data matrix (? unknown; — structure absent; capitals represent the following polymorphies: A 1, 4; B 1, 2;

= E = F = C 5, 6; D = 1, 3; 3, 4, 5, 6; 1, 4)

1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 3 3 3 3 3 3 3 3 3 3 5 5 5 5 5 5 5 5 5 5 6 6 6 6 6 6 6

1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 12 3 4 5 6 7 8 9 0 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6

1 B guttulipennis 2 1 1 2 2 1 1 1 1 1 1 1 2 1 1 1 1 2 1 2 1 1 2 1 11112 1 1 2 1 1 2 1 1 1 1 1 3 2 1 1 1 1 1 1 1

2 B.marmorata 2 1 1 2 2 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1 1 1 2 1 11112 1 1 1 1 1 2 1 1 1 1 1 3 1 1 1 1 1 1 1 1

B. 1 1 1 1 1 1 3 nasuta 2 1 1 2 2 1 1 1 1 1 1 1 2 1 1 1 1 2 1 2 1 1 2 1 11112 1 1 1 1 1 2 1 1 1 1 3 2 1 1

1 1 1 1 1 4 B.intermedia 2 ? 1 2 2 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1 1 1 2 1 11112 1 1 1 1 1 2 1 1 1 1 1 7 2 1 1

5 B.splendida 1 2 1 2 2 1 1 1 1 1 1 1 2 1 1 1 î 2 1 1 1 1 2 1 1111 2 1 1 1 1 1 2 1 1 1 1 1 3 2 1 1 1 1 1 1 1

6 B. parva 2 1 1 2 2 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1 1 1 2 1 11112 1 1 1 1 1 2 1 1 1 1 1 3 2 1 1 1 1 1 1 1

1 1 7 B.mamillata 2 1 1 2 2 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1 1 1 2 1 11112 1 1 1 1 2 1 1 1 1 1 3 2 1 1 1 1 1

1 8 B arabuensis 2 1 1 2 2 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1 1 1 2 1 1111 2 1 1 1 1 1 1 1 1 1 1 3 2 1 1 1 1 1 1

1 1 1 1 1 1 1111 9 B gibberosa 2 1 1 2 2 1 1 1 1 1 1 1 1 1 1 2 2 1 2 1 1 1 1 2 1 1 1 1 1 3 2 1 1 1 1 1 1 1

1 0 B.bipunctata 2 1 1 2 2 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1 1 1 2 1 11112 1 1 1 1 2 1 1 1 1 1 3 2 1 1 1 1 1 1 1

1 1 1 1 1 1 1 B.laminifer 2 1 1 2 2 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1 1 1 2 1 1111 2 1 1 1 1 2 1 1 1 1 3 2 1 1 1

1 1 1 1 2 1 2 1 1 1 2 B. retracta 2 1 1 2 2 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1111 2 1 1 1 1 1 1 3 2 1 1 1 1 1 1 1

1 3 B.brongersmai 1 2 1 2 5 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 1 1 1 1 1 1 1 1 1 4 1 1 1 1 1 1 1 1

1 4 1 1 1 1 1 1 1 1 4 B. vi ri di s 1 2 1 2 5 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 1 1 1 1 1 1 1 1 1

1 1 1 1 5 B.lorentzi 1 2 1 2 5 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 1 1 1 1 1 1 4 1 1 1 1 1 1 1 1

1 6 B.rufula 1 ? 1 2 3 1 1 1 1 1 1 1 1 1 1 1 3 2 1 1 1 1 2 1 1111 2 1 1 1 1 2 1 1 1 1 1 ? 1 1 1 1 1 1 1 1

1 1 1 1 1 7 B. turgida 2 1 1 2 3 1 1 1 1 1 1 1 2 1 1 1 3 2 1 1 1 1 2 1 1111 2 1 1 3 1 1 1 1 1 1 1 1 3 1 1 1 1

1 1 1 1 8 B.furcillata 2 ? 1 2 3 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1 1 1 2 1 1111 2 1 1 3 1 1 2 1 1 1 1 ? 1 1 1 1 1 1

1 9 B karkarensis 2 1 1 2 3 1 1 1 1 1 1 1 2 1 1 1 3 2 1 1 1 1 2 1 3 111 2 1 1 3 1 1 1 1 1 1 1 1 3 1 1 1 1 1 1 1 1

1 1 20 B.conviva 2 1 1 2 3 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 2 1 1 3 1 1 1 1 1 1 1 1 3 1 1 1 1 1 1

1 1 21 B. laureti 2 1 1 2 3 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 2 1 1 3 1 1 1 1 1 1 1 1 3 1 1 1 1 1 1

2 2 B.schulzi 2 1 1 2 3 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 2 1 1 3 1 1 1 1 1 1 1 1 3 1 1 1 1 1 1 1 1

1 1 1 1 1 2 1 1 2 3 B. quadrifida 1 1 1 2 3 1 1 1 1 1 1 1 1 1 2 1 1 1111 2 1 3 1 1 2 1 1 1 1 1 3 1 1 1 1 1 1 1 1

1 1 24 B.hardyi 2 1 1 2 3 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 2 1 1 3 1 1 2 1 1 1 1 1 3 1 1 1 1 1 1

2 5 B guttulinervis 1 1 1 2 2 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 2 1 1111 2 1 1 4 1 1 1 1 1 1 1 1 3 1 1 1 1 1 1 1 1

1 1 1 2 6 B.roonensis 1 1 1 2 5 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 2 1 1 6 1 2 1 1 1 1 3 1 1 1 1 1 1 1

27 B.inconstans 2 1 1 2 3 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 5 111 2 1 1 2 1 1 2 1 1 1 1 1 3 1 1 1 1 1 1 1 1

1 1 1 i 1 1 1 3 1 1 1 1 1 1 1 2 8 B.biroi 2 1 1 2 5 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 2 2 2 1 1 1 1 1

1 4 1 1 1 1 1 4 2 9 B.bloetei 1 2 1 2 5 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 2 1 2 1 1 1 1 1 1 1 1 1

1 3 0 B macgillavryi 1 2 1 2 5 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 1 4 1 1 1 2 1 1 2 1 1 4 1 1 1 1 1 1 1

1 4 4 1 1 3 1 B papuensis 1 2 1 2 5 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 1 1 1 2 1 1 2 1 1 1 1 1 1 1 1

4 1 1 1 1 1 1 1 1 1 3 2 B.bismarckensis 2 2 1 2 5 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 1 1 2 2 1 1 3 1 1 1

3 3 B manusensis 2 1 1 2 5 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 1 4 1 1 1 2 1 1 2 1 1 3 1 1 1 1 1 1 1 1

34 B.mussauensis 2 1 1 2 5 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 1 4 1 1 1 2 1 1 2 1 1 3 1 1 1 1 1 1 1 1

3 5 B.brandti 2 1 1 2 5 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 1 4 1 1 1 2 1 1 2 1 1 3 1 1 1 1 1 1 1 1

1 1 1 2 1 4 1 1 1 1 2 1 1 1 1 1 3 6 B.sedlacekorum 2 1 1 2 5 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1111 1 1 1 1 2 1 3 1 1 1 1

1 37 B.reynhoudti 2 1 1 2 5 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 1 4 1 1 1 2 1 1 2 1 1 3 1 1 1 1 1 1 1

1 1 2 1 1 1 3 1 1 1 1 3 8 B.cristovalensis 2 1 1 2 5 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 1 4 1 1 2 1 1 1 1

1 1 1 1 3 9 B.edauberti 2 1 1 2 5 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 1 4 1 1 1 2 1 1 2 1 1 3 1 1 1 1

4 0 B maddisoni 2 1 1 2 5 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 1 4 1 1 1 2 1 1 2 1 1 3 1 1 1 1 1 1 1 1

1 4 1 B.gressitti 1 2 1 2 5 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 1 4 1 1 1 2 1 1 2 1 1 3 1 1 1 1 1 1 1 1

1 4 2 B bilebanarai 1 2 1 2 5 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 1 4 1 1 1 2 1 1 2 1 1 3 1 1 1 1 1 1 1

4 3 B mendanai 1 2 1 2 5 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 1 4 1 1 1 2 1 1 2 1 1 3 1 1 1 1 1 1 1 1

1 1 4 4 B. marginata 1 2 1 2 5 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 1 4 1 1 1 2 1 1 2 1 1 3 1 1 1 1 1 1

45 B.boulardi 2 1 1 2 5 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 1 4 1 1 1 2 1 1 2 1 1 3 1 1 1 1 1 1 1 1

4 6 B rotumae 2 1 1 2 5 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 1 4 1 1 1 2 1 1 2 1 1 3 1 1 1 1 1 1 1 1

1 1 1 1 1 1 1 1 4 7 B exhausta 2 1 1 2 5 1 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 1 4 1 1 1 2 1 1 1 1 3 1 1 1 1 1 1 1 1

4 8 B.bicolorata 2 2 1 2 5 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 1 4 1 1 1 2 1 1 1 1 1 4 1 1 1 1 1 1 1 1

1 1 4 1 1 1 1 1 1 1 1 1 1 1 1 49 B colossea 2 1 1 2 5 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 1 3 1 1 1

1 1 1 1 1 1 1 1 50 B. rossi 2 1 1 2 3 1 1 1 1 1 1 1 1 2 2 1 1111 1 1 1 4 1 1 1 2 1 1 2 1 1 3 1 1 1 1 1 1 1

51 B.vandertiammeni 2 1 1 2 5 1 1 1 1 1 1 1 1 1 1 1 3 2 1 1 1 1 2 1 1111 2 1 1 4 1 1 1 1 1 1 1 1 1 3 1 1 1 1 1 1 1 1

5 2 Bwauensis 2 1 1 2 2 1 1 1 1 1 1 1 1 1 1 1 3 2 1 1 1 1 2 1 4 111 2 1 1 4 1 1 2 1 1 1 1 1 1 3 1 1 1 1 1 1 1 1

111 1 1 4 1 1 2 1 1 1 1 1 1 1 1 53 B.versicolor 1 1 1 2 3 1 1 1 1 1 1 1 1 1 1 1 3 2 1 1 1 1 2 1 4 2 3 1 1 1 1 1 1

1 5 4 B.maai 2 1 1 2 5 1 1 1 1 1 1 1 1 1 1 1 3 2 1 1 1 1 2 1 1111 1 1 1 3 1 1 1 1 1 1 1 1 1 ? 1 1 1 1 1 1 1

55 B.tenuispina 2 1 1 2 5 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 2 1 1 5 1 1 1 1 1 1 1 1 1 3 1 1 1 1 1 1 1 1

1 1 1 4 1 1 1 1 1 1 1 56 B.silveri 2 1 1 2 3 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1 2 1 1111 2 1 1 1 1 1 1 1 3 1 1 1

57 B.daviesi 2 1 1 2 2 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 2 1 3 1 1 1 2 1 1 5 1 2 ? 1 2 1 1 1 1 1 1

5 8 B.pigrami 1 1 1 2 2 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1 2 1 1111 1 1 2 1 4 1 1 1 2 1 1 5 1 2 3 1 2 1 1 1 1 1 1

59 B.hartonoi 2 1 1 2 5 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1 1 1 2 1 1111 1 1 2 1 4 1 1 1 1 1 1 1 1 1 7 1 2 1 1 1 1 1 1

60 B.loriae 1 1 1 2 3 1 1 2 1 1 1 1 1 1 2 1 1 1 1 2 1 3 1 1 3 1 1111 1 1 2 1 3 1 1 1 2 1 1 1 1 1 ? 1 ? 1 2 1 1 1 1 1 1

61 B.fortuini 1 1 1 2 3 1 1 2 1 1 1 1 1 1 2 1 1 1 1 2 1 3 1 1 3 1 1111 1 1 2 1 3 1 1 1 2 1 2 1 5 1 2 3 1 2 1 2 1 1 1 1 1 1

6 2 B.hamiltoni 1 1 1 2 2 1 1 2 1 1 1 1 2 1 2 1 1 1 1 2 1 1 1 1 2 1 1112 1 2 1 4 1 1 1 2 1 2 1 3 1 2 3 1 1 1 2 1 1 1 1 1 1

63 B.bemmeleni 1 1 1 2 2 1 1 2 1 1 1 1 2 1 2 1 1 1 1 2 1 1 1 1 2 1 1112 1 1 2 1 4 1 1 1 2 1 2 1 3 1 2 3 1 2 1 2 1 1 2 1 1 1

6 4 B.wegeneri 1 1 2 3 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1 1 1 2 1 1112 1 2 1 4 1 1 1 2 1 2 1 3 1 2 ? 1 1 1 2 1 1 1 1 1 1

6 5 S.sahebdivanii 1 1 1 1 1 1 1 2 1 1 1 3 1 1 2 1 1 2 1 1 2 1 1 1 1 1 1 3 1 1111 1 1 1 1 4 1 1 1 1 1 2 1 1 1 1 1 1 1 2 1 1 1 1 1 2 1 1

66 S.biardae 1 1 1 1 1 1 1 2 1 1 1 3 1 1 2 1 1 2 1 1 2 1 1 1 1 1 1 3 1 1111 1 1 1 1 4 1 1 1 1 1 2 1 1 1 1 1 ? 1 2 1 1 1 1 1 2 1 1

67 S.huibregtsae 1 1 1 1 1 1 1 2 1 1 1 3 1 1 2 1 1 2 1 1 2 1 1 1 1 1 1 3 1 1111 1 1 1 1 3 1 1 1 1 1 2 1 1 1 1 1 ? 1 2 1 1 1 1 1 2 1 1

1111 1 1 2 1 68 G.strepitane 1 1 1 1 1 1 1 2 1 1 2 3 1 2 2 1 3 2 1 1 2 1 1 1 1 1 1 3 1 2 3 2 2 1 2 1 1 2 1 5 2 2 5 1 2 1 2 1 1 1 1 1

6 9 G.viridis 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1 3 1 1111 1 1 2 2 3 1 1 1 2 1 1 1 1 1 1 1 3 1 2 1 2 1 1 1 1 1 1 1 1

70 G.daNi 1 1 1 1 1 1 1 2 1 1 2 3 1 1 1 1 3 2 1 1 1 2 1 1 1 1 1 3 1 1111 1 1 2 1 3 2 1 1 1 1 1 2 1 1 2 2 3 1 2 1 1 1 1 1 1 1 1 1 1

1 1 1 1 1 1 71 G.rubricata 1 1 1 1 1 1 1 1 1 1 2 3 1 2 1 1 3 2 1 1 1 1 1 1 1 1 3 1 1111 1 1 2 2 1 2 1 1 2 1 1 2 1 5 1 2 3 2 1 1 1 1 1

72 G.stridere 1 1 1 1 1 1 1 2 1 1 2 3 1 2 2 1 1 2 1 1 2 1 1 1 1 1 1 3 2 1111 1 1 2 2 4 2 1 1 2 1 1 2 2 5 2 2 1 2 2 1 1 1 1 1 1 2 1 1 1

73 G.obienss 1 1 1 1 1 1 1 2 1 1 2 3 1 2 2 1 1 2 1 1 2 1 1 1 1 1 1 3 2 1111 1 1 2 2 1 1 2 1 1 2 1 1 2 2 5 2 2 1 2 1 2 2 1 1 1 1 1 1 2 1 1 1

2 1 74 G.subnotata 1 1 1 1 1 1 1 2 1 1 2 3 1 2 2 1 1 2 1 1 2 1 1 1 1 1 1 3 2 1111 1 1 2 2 1 1 2 1 1 2 1 1 2 2 5 2 2 1 2 1 2 1 1 1 1 1 2 1 1 1

75 G.nigravirgula 1 1 1 1 1 1 1 2 1 1 2 3 1 2 3 1 1 2 1 1 2 1 1 1 1 1 1 3 2 1111 1 1 2 2 1 1 1 1 1 2 1 1 2 1 3 1 2 1 2 1 1 2 1 2 1 2 1 1 2 1 1 1 228 A.J. de Boer - Phytogeny and taxonomicstatus of Chlorocystini s. s.

Appendix 3. Continued.

1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 3 3 3 3 3 3 3 3 3 5 5 5 5 5 5 5 5 5 5 6 6 6 6 6 6 6

4 6 8 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 5 7 0 2 3 4 5 6

? 2 1 2 2 76 G stanocephais 1 1 1 1 1 1 1 2 1 3 1 2 2 1 2 1 1 2 1 4 1 1 3 1 1 1 1 1 1 1 2 2 1 1 1 1 2 1 2 1 2 1 5 2 2 1 2 1 1 1 1 1

77 G párvula 1 1 1 1 1 2 1 1 2 3 1 2 1 1 2 1 2 1 1 1 1 1 3 1 1 1 1 1 1 1 2 2 1 1 1 1 2 1 2 1 2 1 5 2 2 1 2 ? 1 2 1 2 2 1 2 1 1

1 1 2 1 1 2 1 2 3 2 1 78 G.membrana 1 1 1 1 1 2 1 1 2 3 1 2 1 2 1 2 1 1 1 1 1 3 2 1 1 1 1 1 1 2 2 1 1 1 2 1 2 1 2 1 5 2 2 1 1

1 1 1 79 G.minoramambrant 1 1 1 1 1 1 1 2 2 1 1 1 1 1 2 1 1 1 1 1 3 1 1 1 1 1 1 1 2 2 1 1 1 2 1 2 1 2 1 5 2 2 1 2 1 1 2 1 2 3 2 1 1

80 G.phyloglycara 1 1 1 1 1 2 1 1 2 3 1 1 1 2 1 2 1 1 1 1 1 3 2 1 1 1 1 1 1 2 2 1 1 1 1 1 1 2 1 2 1 5 2 2 1 2 1 1 2 1 2 2 1 1 1

? 81 G.langaraki 1 1 1 1 1 2 1 1 2 3 1 3 1 2 1 2 1 1 1 1 1 3 1 1 1 1 1 1 1 2 2 1 1 1 1 2 1 2 1 2 1 5 2 2 1 2 1 2 1 2 2 1 1 1

1 1 1 1 1 1 1 82 G hirsuta 1 2 3 1 3 1 1 1 1 2 1 1 1 1 1 1 4 1 1 3 1 5 2 2 1 1 1 2 2 1 1 2 1 2 1 3 1 1 1 2 3 1 2 1 2

8 3 G.olivaoaa 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1 1 1 1 3 1 5 1 2 1 1 1 2 1 5 1 1 1 1 1 2 1 2 1 3 2 1 1 2 5 1 2 1 2 1 1

5 1 8 4 G.varlaani 1 1 1 1 1 1 1 1 1 3 1 1 1 1 1 1 1 1 1 1 3 1 5 2 2 1 1 1 2 2 1 1 1 1 1 1 2 1 2 1 4 2 1 1 2 1 2 2 1 1

8 5 G.montana 1 1 1 1 1 1 1 1 1 3 1 1 1 1 1 1 1 1 1 1 3 1 5 2 2 1 1 1 2 2 1 1 1 1 1 1 2 1 2 1 4 2 1 1 2 5 1 2 1 2 1 1

1 1 3 2 1 1 1 1 86 G.rufa 1 1 1 1 3 1 1 2 ? 1 3 1 1 1 1 1 1 1 1 1 1 2 1 2 1 1 1 1 1 1 1 3 1 1 2 1 1 2 1 2 2 1 1 1

87 G.varicolor 1 1 1 1 3 1 1 1 2 3 1 1 1 1 1 2 1 1 1 1 3 2 1 1 1 1 1 1 2 1 3 1 1 1 1 1 1 1 3 1 1 1 1 1 2 3 1 2 2 1 1 1

1 1 1 3 2 3 1 1 1 1 1 1 3 1 1 1 1 1 1 1 3 1 1 1 1 1 2 3 1 1 1 1 1 8 8 V.superba 1 1 1 2 1 2 1 1 2 1 1 1 1 2 1 4 1

C 1 1 1 89 C.vilripannis 1 1 1 1 1 1 2 1 2 1 1 1 1 1 1 2 1 1 1 1 3 2 5 1 3 1 1 2 1 1 1 1 1 1 1 1 1 7 1 1 2 1 3 1 2

90 C.suffusa 1 1 1 1 1 2 1 1 2 1 1 2 1 1 1 2 1 1 1 1 3 2 5 1 3 1 1 2 1 7 1 1 1 1 1 1 1 1 7 1 1 2 1 3 1 1 1 1 2

1 1 ? 91 C.lumea ? 1 1 2 1 1 2 1 1 2 1 1 1 2 1 1 1 1 3 2 5 1 3 1 1 2 1 7 1 1 1 1 1 1 1 1 7 1 1 2 1 1 1 1 1 2

9 2 0.insignia 1 1 1 1 1 1 2 1 2 2 1 1 1 1 1 1 1 1 1 1 1 2 5 1 3 1 1 2 1 3 1 1 1 1 2 1 1 1 7 1 1 2 1 3 1 1 1 1 2

93 Gl.viridi» ? 1 1 1 1 1 2 1 2 2 1 1 1 1 1 2 1 1 1 1 3 2 5 1 3 1 1 1 4 1 1 1 1 1 1 1 1 7 1 1 2 1 1 1 1 1 2

1 1 1 1 1 1 1 6 1 94 G.(lava 1 2 3 1 1 2 1 1 2 3 1 1 1 1 1 2 1 1 1 3 1 1 1 3 1 1 2 1 1 1 1 1 2 1 1 1 1 1

9 5 GHavaola 1 2 3 1 1 2 1 1 1 2 3 1 1 1 1 1 2 1 1 1 1 3 1 1 1 1 1 1 3 1 1 1 1 2 1 1 1 6 1 1 2 1 1 1 1 1 1

9 6 G.styiata 1 2 3 1 1 1 1 1 1 2 3 1 1 1 1 1 2 1 1 1 1 2 1 1 1 1 1 1 3 1 1 1 1 2 1 1 1 6 1 1 2 1 1 1 1 1 1

4 1 1 1 1 9 7 G.viridula 1 1 1 2 2 1 1 1 2 3 1 1 1 1 1 2 1 1 1 1 3 1 1 1 1 1 1 1 1 1 1 2 1 1 1 6 1 1 2 1 1

9 8 G pallida 1 1 1 2 3 1 1 1 2 3 1 1 1 1 1 2 1 1 1 1 3 1 1 1 1 1 1 3 1 1 1 1 2 1 1 1 6 1 1 2 1 1 1 1 1 1

1 1 1 9 9 G pallidula 1 1 1 2 1 1 1 1 2 3 1 1 1 1 1 1 1 1 1 1 3 1 1 1 1 1 1 3 1 1 1 1 2 1 6 1 1 2 1 1 1 1 1

1 6 1 1 1 1 1 100 G chinai 1 3 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 3 1 1 1 1 1 1 3 1 1 1 1 2 2 1 1 2 1 1

101 Gparmyi ? 1 3 2 3 1 1 1 2 1 1 1 1 1 2 1 1 1 1 3 1 1 1 1 1 1 3 1 1 1 1 2 1 1 1 6 1 1 2 1 1 1 1 1 1

102 P.phyllophora 1 1 1 1 2 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 3 1 1 1 1 1 1 5 1 1 1 1 2 1 1 1 6 1 1 2 1 1 1 1 1 1

103 Pgoniodes 1 1 1 1 1 1 1 1 1 1 1 1 1 3 2 1 1 1 1 3 1 1 1 1 1 1 3 1 1 1 1 2 1 2 1 6 1 1 2 1 1 1 1 1 1

1 4 1 1 1 1 6 104 P.angulata 1 1 1 1 2 1 1 1 1 1 1 1 1 1 3 2 1 1 1 1 3 1 1 1 1 1 2 1 2 1 1 1 2 1 1 1 1 1 1

105 P.lachlani 1 3 1 1 1 1 1 1 1 1 1 1 1 1 3 2 1 1 1 1 3 1 1 1 1 1 1 4 1 1 1 1 2 1 1 6 1 1 2 1 1 1 1 1 1

106 P.brassi 1 3 1 1 1 1 1 1 1 1 1 1 1 3 2 1 1 1 1 3 1 1 1 1 1 1 3 1 1 1 1 2 1 1 6 1 1 2 1 1 1 1 1 1

107 P.stoliodes 1 1 1 1 1 1 1 1 1 1 1 1 1 1 3 2 1 1 1 1 3 1 1 1 1 1 1 4 1 1 1 1 2 1 1 7 1 1 1 1 1 1 1 1 1

108 P.plicata 1 1 1 1 1 1 1 1 1 1 1 1 1 1 3 2 1 1 1 1 3 1 1 1 1 1 1 4 1 1 1 1 1 1 1 7 1 1 1 1 1 1 1 1 1

1 1 109 P.spinigera 1 1 1 1 1 1 1 1 1 1 1 1 3 2 1 1 1 1 2 1 1 1 1 1 1 4 1 1 1 1 2 1 1 6 1 1 2 1 1 1 1 1 1

110 P.bidigitula 1 3 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1 1 1 1 1 4 1 1 1 1 2 1 1 7 1 1 2 1 1 1 1 1 1

111 P ungula 1 1 1 1 1 1 1 1 1 1 1 1 1 1 3 2 1 1 1 1 1 1 1 1 1 1 1 5 1 1 1 1 2 1 1 7 1 1 2 1 1 1 1 1 1

1 1 1 1 1 2 6 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1 3 1 1 1 1 2 1 1 1 1 1 1 11 2 P nana 1 3 1 1 1 1 1 1 1 1 1

113 P.woodlarkansis 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1 1 4 1 1 1 1 2 1 1 6 1 1 2 1 1 1 1 1 1

114 P.ustulata 1 1 1 1 1 1 1 1 1 1 1 1 1 1 3 2 1 1 1 1 2 1 1 1 1 1 1 4 1 1 1 1 2 1 1 6 1 1 2 1 1 1 1 1 1

115 P. novaría* 1 3 1 1 1 1 1 1 1 1 1 1 1 1 3 2 1 1 1 1 2 1 1 1 1 1 1 3 1 1 1 1 2 1 1 6 1 1 2 1 1 1 1 1 1

116 P.baasi 1 3 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1 1 1 1 1 3 1 1 1 1 2 1 1 7 1 1 2 1 1 1 1 1 1

11 7 P dolabrata 1 3 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1 6 1 1 2 1 1 1 1 1 1

11 8 P.toxopai 1 3 1 1 1 1 1 1 1 1 1 1 1 1 3 2 1 1 1 1 1 1 1 1 1 1 3 1 1 1 1 2 1 1 6 1 1 2 1 1 1 1 1 1

1 4 1 1 1 2 1 1 119 P dioedee 1 1 1 1 1 1 1 1 1 1 1 1 1 1 3 2 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1 1 1 1

1 20 A.oobrops 1 1 1 1 1 1 1 2 1 1 1 1 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 4 1 1 1 1 2 1 1 1 1 2 1 1 1 1 1 1

121 Asapia 1 1 1 1 1 1 1 2 1 1 1 1 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 5 1 1 1 1 2 1 1 1 1 2 1 1 1 1 1 1

122 A.latifrona 1 1 1 1 1 1 1 2 2 2 1 1 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 3 1 1 1 1 2 1 1 1 1 2 1 1 1 2 1 1 1

123 A kaiansis 1 1 1 1 1 1 1 2 1 2 1 1 2 2 1 1 2 1 1 1 1 1 1 1 1 1 1 3 1 1 1 1 2 1 1 1 1 2 1 1 1 2 1 1 1

1 24 A.waigauensjs 1 1 1 1 1 1 1 2 1 2 1 1 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 3 1 1 1 1 2 1 1 1 1 2 1 1 1 2 1 1 1

125 A digitata 1 1 1 1 2 1 1 1 1 2 1 1 2 2 2 1 2 1 1 1 1 1 1 1 1 1 1 4 1 1 1 1 2 1 1 1 1 2 1 1 1 1 1 1 1

126 A.hastulata 1 1 1 1 1 1 1 2 1 2 1 1 2 2 1 1 2 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1 1 1 2 1 1 1 2 1 1 1

12 7.A.cheesmanae 1 1 1 1 1 1 1 2 1 2 1 1 2 2 1 1 2 1 1 1 1 1 1 1 1 1 1 3 1 1 1 1 2 1 1 1 1 2 1 1 1 2 1 1 1

128 A moluccansis 1 1 1 1 1 1 1 2 1 2 1 1 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 3 1 1 1 1 2 1 1 1 1 2 1 1 1 2 1 1 1

129 A.obiansis 1 1 1 1 1 1 1 2 1 2 1 1 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1 1 1 2 1 1 1 1 1 1 1

1 30 A búllala 1 1 1 1 2 1 1 1 1 2 1 1 2 2 1 1 2 1 1 1 1 1 1 1 1 1 1 3 1 1 1 1 2 1 1 1 1 2 1 1 1 2 1 1 1

1 1 1 1 1 1 1 131 A dilobata 1 1 1 2 1 1 1 1 2 1 1 2 2 1 2 1 1 1 1 1 2 1 1 1 1 2 1 1 1 1 2 1 1 1 2 1 1 1

132 Tadipata 1 1 4 1 1 1 2 2 1 2 1 1 2 1 2 1 1 1 1 1 1 1 1 1 3 1 4 1 3 1 1 2 1 2 1 1 1 1 2 1 1 1 1 1 1 2

133 T.pnaumabca 1 2 1 4 1 1 1 2 2 1 1 1 1 2 1 2 1 2 1 3 1 1 1 1 1 3 1 4 i 3 1 1 2 1 2 1 1 1 1 2 1 1 1 1 1 1 2

1 1 1 34 T speiunca 1 2 1 4 1 1 1 2 2 1 2 1 1 2 1 2 1 1 1 3 1 1 1 1 2 3 1 3 1 3 1 1 1 2 1 1 1 2 1 1 1 1 1 1 1

135 T.gtobosa 1 2 1 4 1 1 1 2 2 1 2 1 1 2 1 2 1 2 1 3 1 1 1 1 2 3 1 3 1 3 1 1 2 1 2 1 1 1 1 2 1 1 1 1 1 1 2 2

1 36 T.sicula 1 2 1 4 1 1 1 2 2 1 2 1 1 2 1 2 1 2 1 1 1 1 1 1 1 1 1 1 4 1 1 1 1 2 1 1 1 1 2 1 1 1 1 1 1 1 1

1 37 T lanceola 1 2 1 4 1 1 1 2 2 1 2 1 1 2 1 2 1 2 1 3 1 1 1 1 2 3 1 1 3 1 1 2 1 2 1 1 1 1 2 1 1 1 1 2 1 1 2 2

138 T.sarissa 1 2 1 4 1 1 1 2 2 1 1 1 1 2 1 1 1 2 1 3 1 1 1 1 2 3 1 1 3 1 1 2 1 2 1 1 1 1 2 1 1 1 1 2 1 1 2 2

139 M. vi ridicala 1 2 3 1 1 1 1 2 1 1 2 1 1 2 4 1 1 2 1 2 1 1 1 1 1 3 1 1 2 1 1 2 1 2 1 1 1 1 2 1 1 1 1 2 1 1 1 2

1 1 1 140 M globulata 1 2 3 1 1 1 1 1 2 1 2 1 1 2 4 2 1 2 1 2 1 1 1 1 2 3 1 1 3 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

14 1 M inconspicua 1 2 1 1 1 1 1 2 2 1 2 1 1 2 1 2 1 2 1 1 1 1 1 1 1 1 1 1 3 1 1 2 1 2 1 1 1 1 2 1 1 1 1 2 1 1 1

142 M.humilia 1 2 1 1 1 1 1 2 2 1 2 1 1 2 4 2 1 2 1 1 1 1 1 1 1 1 1 1 3 1 1 2 1 2 1 1 1 1 2 1 1 1 1 2 1 1 1

143 M. infiala 1 2 1 1 1 2 1 2 2 1 2 1 1 2 1 2 1 2 1 2 1 1 1 5 1 2 1 1 1 2 1 1 2 1 2 1 1 1 1 2 1 1 1 1 2 1 1 1

1 1 1 1 1 1 144 M.toxopeusi 1 2 1 1 1 2 1 1 1 1 1 1 1 1 1 2 1 2 1 1 1 1 1 1 1 1 1 1 1 3 1 1 2 1 2 1 1 2 1 1 1 1

145 C.schmeltzi 1 1 1 4 2 3 1 3 1 1 1 2 1 2 4 2 2 1 2 1 2 1 2 2 1 6 1 ? 1 1 1 4 1 1 2 1 1 2 3 1 1 1 2 1 1 1 1 1 1 1 1

146 C.saundersii 1 2 1 4 2 3 1 3 1 1 1 2 1 2 4 2 2 1 2 1 2 1 2 2 1 6 1 ? 1 1 1 4 1 1 2 1 1 2 3 1 1 1 1 1 1 1 1 1 1 1 1

147 C immaculata 1 1 1 4 1 3 1 3 1 1 1 2 1 2 4 2 2 1 2 1 2 1 2 2 1 6 1 ? 1 1 1 6 1 1 1 1 1 1 3 1 1 1 1 1 1 1 1 1 1 1 1

148 1 1 1 1 1 1 1 1 1 1 1 1 1 Praaia 1 1 3 4 1 1 1 1 1 1 1 2 1 2 4 2 2 1 2 1 D 1 1 1 2 E 1 1 1 1 1 1 2 1 1 1 1 1 1

149 Muda 1 1 2 A 1 3 1 1 1 1 1 1 1 B 1 1 B 1 2 B 1 1 1 1 1 1 1 1 1 1 1 1 1 F 1 1 1 1 2 1 1 1 1 1 2 1 1 2 1 1 1 1 1 Contributions to Zoology, 65 (4) - 1995 229

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

8 8 8 8 8 8 8 8 8 9 9 9 9 9 9 9 9 9 9 1 1 1 character nr 6 6 6 7 7 7 7 7 7 7 7 7 7 8 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 3 3

7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 2 3 4 5 6 0 2 3 4 5 6 7 8 9 0 1

1 B guttulipennis 1 1 1 2 2 2 1 2 1 1 1 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1 1 1 1 1 1 2 B marmo rata 1 1 1 2 2 2 2 3 1 1 2 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 3 1 1 1 1 1 1

3 B nasuta 1 1 1 2 2 2 2 2 1 1 2 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 3 1 1 1 1 1 1 1 1 1 1

4 B intermedia 1 1 1 2 2 2 2 3 1 1 2 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 3 1 1 1 1 1 1 1 1 1 1

5 B splendida 1 1 1 2 2 2 2 3 1 1 2 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1 1 1 1 1 1

6 1 1 B. parva 1 1 2 2 2 2 3 1 1 2 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 3 1 1 1 1 1 1 1 1 1

7 B.mamillata 1 1 1 2 2 2 2 3 1 1 2 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 3 1 2 3 1 1 1 1 1 1 1

8 B arabuensis 1 1 1 2 2 2 2 3 1 1 2 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 3 1 2 3 1 1 1 1 1 1 1

9 B gibberosa 1 1 1 2 2 2 2 3 1 1 2 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 1 1 1 1 1 1 3 1 2 4 1 1 1 1 1 1 1

1 0 B.bipunctata 1 1 1 2 2 2 2 3 1 1 2 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1 2 4 1 1 1 1 1 1 1

1 1 B laminifer 1 1 1 2 2 2 2 3 1 1 2 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 1 1 1 1 1 1 3 1 2 2 1 1 1 1 1 1 1

1 2 B retracta 1 1 1 2 2 2 2 3 1 1 2 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 ? 1 1 1 1 1 3 1 2 2 1 1 1 1 1 1 1

1 3 B brongersmai 1 1 1 2 2 5 2 3 1 2 2 1 2 1 2 2 2 1 1 4 1 1 1 1 2 1 2 1 1 1 1 2 2 1 1 1 1 1 3 2 1 1 1 1 1 4 1

1 4 B.viridis 1 1 1 2 2 5 2 3 1 2 2 1 2 1 2 2 2 1 1 4 1 1 1 1 2 1 2 1 1 1 1 2 2 1 1 1 1 1 3 2 1 1 1 1 1 4 1

1 5 B lorentzi 1 1 1 2 2 5 2 3 1 2 2 1 2 1 2 2 2 1 1 4 1 1 1 1 2 1 2 1 1 1 1 2 ? 1 1 1 1 1 3 2 1 1 1 1 1 4 1

1 6 B.rufula 1 1 1 2 2 3 2 3 1 2 2 1 2 1 2 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 ? ? 1 1 1 1 1 ? 1 1 1 1 1 1 2 1

1 7 B. turgida 1 1 1 2 2 3 2 4 1 2 2 1 2 1 2 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 7 1 1 1 1 1 1 3 1 1 1 1 1 1 2 1

1 8 B.furcillata 1 1 1 2 2 4 2 4 1 2 2 1 2 1 2 2 1 1 4 1 1 1 1 2 1 2 1 1 1 1 2 1 1 1 1 1 1 3 1 1 1 1 1 1 2 1

1 9 B karfcarensis 1 1 1 2 2 3 2 3 1 2 2 1 2 1 2 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 2 1 1 1 1 1 1 3 1 1 1 1 1 1 2 1

1 1 1 1 1 1 1 1 1 1 1 1 2 0 B.conviva 1 1 1 2 1 4 1 2 1 1 3 1 2 2 1 1 1 1 1 1 1 1 1 3 1 2 1 1 1 1 3 1

21 B laurei) 1 1 1 2 1 4 1 2 1 1 1 1 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 1 1 1 1 1 1 3 1 2 1 1 1 1 3 1

22 Bschulzi 1 1 1 2 1 4 1 2 1 1 2 1 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 1 1 1 1 1 1 3 1 2 1 1 1 1 3 1

23 B. quadrifida 1 1 1 2 1 4 1 2 1 1 2 1 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 3 1 2 1 1 1 1 3 1

24 B.hardyi 1 1 1 2 1 4 1 2 1 1 2 1 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 3 1 2 1 1 1 1 3 1

25 Bguttulinervis 1 1 1 2 1 4 2 2 1 1 2 1 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 3 1 2 2 1 1 1 1 3 1

1 1 1 26 B.roonensis 1 1 1 2 1 4 2 2 1 1 2 1 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 1 1 1 1 1 3 1 2 2 1 1 3 1

27 B.inconstans 1 1 1 2 1 4 1 2 1 1 2 1 2 1 1 1 4 1 1 1 1 2 1 2 1 1 1 1 2 2 1 1 1 1 1 3 1 2 2 1 1 1 1 3 1 2 1

28 B.biroi 1 1 1 2 1 4 1 2 1 1 2 1 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 3 1 2 2 1 1 1 1 3 1 2 1

29 B bloetei 1 1 1 2 1 6 1 2 1 1 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 2 1 1 1 1 1 1 1

30 B.macgillavryi 1 1 1 2 1 6 1 2 1 1 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 2 1 1 1 1 1 1 1

3 1 B papuensis 1 1 1 2 1 6 1 2 1 1 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 2 1 1 1 1 1 1 1

32 B.bismarckensis 1 1 1 2 1 6 1 2 1 1 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 3 1 1 1 1 1 1 1

3 3 B manusensis 1 1 1 2 1 6 1 2 1 1 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 3 1 1 1 1 1 1 1

3 4 B mussauensis 1 1 1 2 1 6 1 2 1 1 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 3 1 1 1 1 1 1 1

35 B brandii 1 1 1 2 1 4 1 2 1 1 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 3 1 2 1 1 1 1 1

3 6 B sedlacekorum 1 1 1 2 1 4 1 2 1 1 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 2 1 2 1 1 1 1 1

3 7 B.reynhoudti 1 1 1 2 1 6 1 2 1 1 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 2 1 2 1 1 1 1 1

3 8 B.cristovalensis 1 1 1 2 1 6 1 2 1 1 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 3 1 2 1 1 1 1 1

3 9 B edaubert 1 1 1 2 1 4 1 2 1 1 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 3 1 2 1 1 1 1 1

1 1 1 1 1 1 1 2 1 2 1 1 1 1 1 1 1 1 1 4 0 B maddisoni 1 1 1 2 1 6 1 2 1 1 2 1 3 1 1 1 1 1 1 2 1 3 1 2

4 1 B gressitti 1 1 1 2 1 4 1 2 1 1 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 3 1 1 1 1 1 1

4 2 B bilebanarai 1 1 1 2 1 6 1 2 1 1 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 3 1 1 1 1 1 1

4 3 B mendanai 1 1 1 2 1 6 1 2 1 1 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 2 1 1 1 1 1 1

4 4 B marginata 1 1 1 2 1 6 1 2 1 1 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 2 1 1 1 1 1 1

4 5 Bboutardi 1 1 1 2 1 6 1 2 1 1 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 2 1 1 1 1 1 1

46 B rotumae 1 1 1 2 1 6 1 2 1 1 1 2 1 1 1 1 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 2 1 1 1 1 1 1

4 7 B exhausta 1 1 1 2 1 1 1 1 2 1 1 2 1 1 1 2 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 3 1 2 1 1 1 4 1

4 8 B bicolorata 1 1 1 2 1 1 1 1 2 1 1 2 1 1 1 2 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 3 1 2 1 1 1 4 1

49 B oolossea 1 1 1 2 1 1 3 1 1 1 2 1 1 1 2 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 3 1 2 1 1 1 4 1

50 B rossi 1 1 1 2 1 1 1 1 1 1 2 1 1 1 2 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 3 1 2 1 1 1 4 1

5 1 B vanderhammeni 1 1 1 2 1 1 2 1 2 1 1 2 1 1 1 2 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 3 1 2 1 1 1 4 1

5 2 B wauensis 1 1 1 2 1 1 2 1 2 1 1 2 1 1 1 2 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 2 1 3 1 2 1 1 1 4 1

53 B. versicolor 1 1 1 2 1 1 1 1 1 1 2 1 1 1 2 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 2 1 3 1 2 1 1 1 4 1

54 B.maai 1 1 1 2 1 1 1 1 1 1 2 1 1 1 2 3 1 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 3 1 2 1 1 1 4 1

5 5 B.tenuisptna 1 1 1 2 1 1 1 1 1 1 2 1 1 1 1 1 1 1 1 1 2 1 2 1 1 1 1 1 2 2 1 1 1 1 3 1 1 1 1 1 1

56 B.ailveri 1 1 1 2 1 6 1 3 1 1 1 2 1 1 1 1 1 1 1 2 1 1 1 2 1 2 1 1 1 1 1 1 2 1 1 1 1 3 2 1 1 1 1 1

57 B daviesi 1 1 1 2 1 6 1 1 1 1 1 2 1 1 1 1 3 1 1 2 1 1 1 2 1 2 1 1 1 1 1 1 2 1 1 1 1 3 2 1 1 1 1 1

5 8 B . pi grami 1 1 1 2 1 6 1 1 1 1 1 2 1 1 1 1 3 1 1 2 1 1 1 2 1 2 1 1 1 1 1 1 2 1 1 1 1 3 2 1 1 1 1 1

5 9 B hartonoi 1 1 1 2 1 6 1 1 1 1 1 2 1 1 1 1 3 1 1 2 1 1 1 2 1 2 1 1 1 1 1 1 2 1 1 1 1 1 1 2 1 5 1

60 B loriae 1 1 1 2 1 6 1 3 1 1 1 2 1 1 1 1 1 1 1 2 1 1 1 2 1 2 1 1 1 1 1 1 ? 1 1 1 1 1 1 2 1 5 1

61 B. fortumi 1 1 1 2 1 6 1 3 1 1 1 2 1 1 1 1 1 1 1 2 1 1 1 2 1 2 1 1 1 1 1 1 2 1 1 1 1 1 1 2 1 5 1

62 B.hamiltoni 1 1 1 2 1 6 1 3 1 1 1 2 1 1 1 1 3 1 1 2 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 2 3 1 1 1 1 1

6 3 B bemmeleni 1 1 1 2 1 6 1 1 1 1 1 2 1 1 1 1 3 1 1 2 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 2 3 1 1 1 1 1

6 4 Bwegeneri 1 1 1 2 1 1 1 1 1 1 2 1 1 1 1 3 1 1 2 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 2 3 1 1 1 1 1

6 5 S sahebdivanii 1 1 1 2 1 1 1 1 1 1 2 1 1 1 1 1 1 1 1 2 2 1 2 1 1 1 1 1 1 1 1 1 1 2 1 1 1 6 1

66 S btardae 1 1 1 1 1 1 1 1 1 1 1 1 1 2 2 1 1 1 1 2 2 2 2 1 1 1 1 1 1 1 1 1 1 2 1 1 1 6 1

67 S huibregtsae 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 2 1 1 1 2 2 2 1 1 1 1 1 1 1 1 1 1 2 1 1 1 6 1

6 8 G.strepitane 2 2 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1 2 1 2 2 1 1 2 1 1 2 1 1 1 1 1 1 1 1 1 3 1 1 1 2 6 1

6 9 G.viridis 2 1 1 1 1 1 2 1 1 1 1 1 2 1 1 1 1 1 2 1 2 2 1 1 1 1 1 2 1 1 1 1 1 1 1 1 1 3 1 1 1 2 6 1

70 G dahii 1 1 1 1 1 1 2 1 1 1 1 1 2 1 1 1 1 1 2 1 2 2 1 1 1 1 1 2 1 1 1 1 1 1 1 1 1 3 1 1 1 2 6 1

1 71 G. rubrìcata 1 2 1 1 1 1 1 1 1 1 1 1 2 1 3 1 1 1 2 2 2 1 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 6 1

72 G.stride ns 1 2 1 1 1 1 1 1 1 1 1 1 2 1 1 2 1 1 2 1 2 2 2 1 1 1 1 1 2 1 1 2 1 1 1 1 1 1 5 1 1 1 1 6 1

1 2 1 2 7 3 G obtenais 2 2 1 1 1 1 1 1 1 1 1 1 2 1 3 2 1 2 2 2 2 1 1 1 1 1 1 2 1 1 1 1 1 1 5 1 1 1 1 6 1

74 G subnotata 2 2 1 1 1 1 1 1 1 1 1 1 2 1 3 2 1 1 2 1 2 1 2 1 1 1 1 1 2 1 1 2 1 1 1 1 1 1 5 1 1 1 1 6 1

75 G.nigravirgula 2 2 1 1 1 1 1 1 1 1 1 1 2 1 1 2 1 1 2 2 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 1 1 1 1 4 1 1 1 1 6 1 1 4 230 A.J. de Boer - Phylogeny and taxonomic status of Chlorocystini s. s.

1 1 1 1 1 1 1 1 1 1 1111 1111 1 1 1111 1111 1 1

6 6 6 7 7 7 7 7 7 7 7 7 7 8 8 8 8 8 8 8 8 8 8 9 9 9 9 9 9 9 9 9 9 0 0 0 0 0 0 0 0 0 0 1111 1111 2 2 2 2 2 2 2 2 2 2 3 3

7 8 9 0 1 2 3 4 5 6 7 8 9 0 12 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 2 3 4 5 6 7 8 9 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1

76 G stenocephais 2 2 1 1 1 1 1 1 1 1 1 1 1 1111 1 2 1 1 1 1 1 2 1 1 2 1 2 1 1 1 2 1 1111 11114 1 1111 116 1

77 G.párvula 2 2 1 1 1 1 1 1 1 1 1 1 1 1111 1 2 1 1 1 1 1 2 2 1 2 1 2 1 1 1 1 1 1111 11114 1 1111 116 1

1 1111 1 1 1 1 3 78 G.membrana 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 2 1 1111 1111 3 1 1111 116 1

79 G.minoramembram2 2 1 1 1 1 1 1 1 1 1 1 1 1111 1 1 1 1 3 1 1 1 1 1 1 1 2 1 1 1 2 1 1111 11113 1 1111 116 1

80 G.phyloglycara 2 2 1 1 1 1 1 1 1 1 1 1 1 1111 1 1 1 1 3 1 1 1 1 1 1 1 2 1 1 1 2 1 1111 11113 1 1111 116 1

81 G langaraki 1 1 1 1 1 1 1 1 1 1 1 1 1 1111 1 2 1 1 2 1 1 2 2 1 2 1 2 1 1 1 2 1 1111 11113 1 1111 116 1

82 G.hirsuta 1 2 1 1 1 1 1 1 1 1 1 1 1 1111 1 2 1 1 1 1 1 1 1 1 2 1 2 1 1 1 2 1 1111 11113 1 1111 116 1

83 G.olrvaoaa 1 1 1 1 1 1 1 1 1 1 1 1 1 1111 1 2 1 1 1 1 2 1 1 1 2 1 2 1 1 1 2 1 1111 11113 1 1111 116 1

1111 3 84 G.vartaani 1 2 1 1 1 1 1 1 1 1 1 1 1 1111 1 2 1 1 1 1 1 1 1 1 2 1 2 1 1 1 2 1 1111 1 1111 116 1

85 G montana 2 1 1 1 1 1 1 1 1 1 1 1 1111 1 2 1 1 1 1 1 1 1 1 2 1 2 1 1 1 2 1 1111 1111 3 1 1111 116 1

86 G rufa 1 1 1 1 1 1 1 1 1 1 1 1 1 1111 1 2 1 3 1 1 1 2 2 1 2 1 1 1 1 2 1 1111 1111 3 1 1111 116 1

87 G.varicolor 1 1 1 2 1 6 1 1 1 1 1 1 1 1111 1 2 1 3 1 1 1 2 2 1 2 1 1 1 1 2 1 1111 1111 4 1 1111 116 1

8 8 V.supertoa 1 1 1 1 1 1 1 1 1 1 1 1 1 1111 1 1 1 1 2 1 2 2 1 1 1 1 1 1 1 1 1111 1111 1 1 1111 116 1

89 C.vitripannia 1 1 1 1 1 1 1 2 1 1 1 1 1 12 11 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1111 1111 2 1 1111 116 1

90 C.suffusa 1 1 1 1 1 1 1 2 1 1 1 1 1 12 11 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 1 1111 1111 2 1 1111 116 1

9 1 C.lumea 1 1 1 1 1 1 1 2 1 1 1 1 1 12 11 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1111 1111 1 1 1111 116 1

92 O.insignis 1 1 1 1 1 1 1 3 1 1 1 1 1 12 11 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1111 1111 2 1 1111 116 1

93 Gl.virídis 1 1 1 1 1 2 1 1 1 1 1 12 11 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1111 1111 2 1 1111 116 1

94 G.flava 1 1 1 1 1 1 1 2 1 1 1 1 1 1111 1 1 1 3 1 1 1 1 1 3 1 1 1 2 2 1 1 2 1111 1111 1 1 1111 116 1

95 G.flavaola 1 1 1 1 1 1 1 2 1 1 1 1 1 1111 1 1 1 3 1 1 1 1 1 3 1 1 1 2 2 1 1 2 1111 1111 1 1 1111 116 1

96 G stylata 1 1 1 1 1 1 1 2 1 1 1 1 1 1111 1 1 1 1 1 1 1 1 1 3 1 1 1 2 2 1 1 2 1111 1111 1 1 1111 116 1

97 G.viridula 1 1 1 1 1 1 1 2 1 1 1 1 1 1111 1 3 1 1 1 1 1 1 1 1 1 2 1 2 1 1111 1111 1 1 1111 116 1

9 8 Gpallida 1 1 1 1 1 1 1 2 1 1 1 1 1 1111 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1111 1111 1 1 1111 116 1

99 G.pallidula 1 1 1 1 1 1 1 2 1 1 1 1 1 1111 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1111 1111 1 1 1111 116 1

100 G.chinai 1 1 1 1 1 1 1 2 1 1 1 1 1 1111 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1111 1111 1 1 1111 116 1

101 Gpanriyi 1 1 1 1 1 1 1 2 1 1 1 1 1 1111 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1111 1111 1 1 1111 116 1

102 P.phyllophora 1 1 1 1 1 1 1 2 1 1 1 1 1 1111 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1111 1111 1 1 1111 116 1

103 P goniodas 1 1 1 1 1 1 1 2 1 1 1 1 1 1111 1 1 1 2 2 1 1 1 2 1 1 1 1 1 1 2 2 1111 1111 2 1 1111 116 1

104 P.angulata 1 1 1 1 1 1 1 2 2 1 1 1 1 1111 1 1 1 2 2 1 1 1 2 1 1 1 1 1 1 2 2 1111 1111 2 1 1111 116 1

105 P.lachlani 1 1 1 1 1 1 1 2 2 1 1 1 1 1111 1 2 2 1 1 1 2 2 1 1 1 1 1 2 2 1111 1111 2 1 1111 116 1

106 P.brassi 1 1 1 1 1 1 1 2 1 1 1 1 1 1111 1 1 1 2 2 1 1 1 2 1 1 1 1 1 1 2 2 1111 1111 2 1 1111 116 1

107 P.stoliodas 1 1 1 2 2 2 1 1 4 1 1 1 1 1 1111 1 1 1 2 1 1 1 1 1 1 1 1 1 1 1 2 1 1111 1111 2 1 1111 116 1 1 3

108 P.plicata 1 1 1 2 2 2 1 1 4 1 1 1 1 1 1111 1 1 1 2 1 1 1 1 1 2 1 1 1 1 1 2 1 1111 1111 2 1 1111 116 1 1 3

1 1 1 109 P.spinigera 1 1 1 1 1 6 1 2 1 1 1 1 1 1111 1 1 1 2 1 1 1 1 1 2 1 1 1 2 1111 1111 2 1 1111 116 1 1 3

11 0 P.bidigitula 1 1 1 1 1 1 1 2 2 1 1 1 1 1111 1 1 1 1 2 1 1 1 1 1 1 1 1 1 1 2 1 1111 1111 2 1 1111 116 1

111 P,ungula 1 1 1 1 1 1 1 2 1 1 1 1 1 1111 1 1 1 2 1 1 1 1 1 2 1 1 1 1 1 2 1 2 1111 1111 4 1 1111 116 1

11 1 1 1 1 1 1 1 1 1 1111 1 1 1 1 1 1 1 1 1 2 P nana 1 2 1 1 1 2 1 1 1 1 2 2 1111 1111 4 1 1111 116 1

113 P.woodlarVensis 1 1 1 1 1 1 1 5 1 1 1 1 1 1111 1 1 1 3 1 1 1 1 1 1 1 1 1 1 2 1 1111 1111 2 1 1111 116 1

114 P.ustulata 1 1 1 1 1 1 1 2 2 1 1 1 1 1111 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 2 1111 1111 3 1 1111116 1

115 P.novariaa 1 1 1 1 1 1 1 2 2 1 1 1 1 1111 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1111 1111 1 1 1111116 1

116 P baasi 1 1 1 1 1 1 1 2 2 1 1 1 1 1111 1 1 1 2 1 1 1 1 1 1 1 1 1 1 2 1 2 1111 1111 3 1 1111 116 1

117 P.dolabrata 1 1 1 1 1 1 1 2 2 1 1 1 1 1111 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1111 1111 1 1 1111 116 1

11 8 P toxopai 1 1 1 1 1 1 1 2 2 1 1 1 1 1111 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1111 1111 2 1 1111116 1

11 9 P.dioedaa 1 1 1 1 1 1 1 2 2 1 1 1 1 1111 1 1 1 1 1 1 1 1 2 1 1 1 1 1 1 1 1111 1111 1 1 1111116 1

120 A.oobrops 1 1 1 3 1 - 1 6 1 1 1111 1 1 1 1 1 1 1 1 4 1 1 1 2 1 1 2 1 1111 1111 1 1 1112116 1

1 121 A sepia 1 1 1 3 1 - 1 6 1 1 1111 1 2 1 2 1 1 1 1 1 1 1 1 2 1 1 2 1 1111 1111 1 1 1112116

1 22 A.latifrons 1 1 1 1 1 1 1 2 1 1 1 1 1 1111 1 2 1 1 1 1 1 1 2 1 1 1 2 1 1 1 1 1111 1111 1 1 1112116 1

123 A.kaiansis 1 1 1 1 1 1 1 5 1 1 1 1 1 1111 1 2 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1111 1111 3 1 1112116 1

124 A.waigeuensis 1 1 1 1 1 1 1 5 1 1 1 1 1 1111 1 1 1 1 1 1 1 1 4 1 1 1 2 1 1 1 1 1111 1111 3 1 1112116 1

125 A digitata 1 1 1 4 1 2 1 1 1 1 1 1111 1 1 1 2 1 1 2 1 1 1 2 1 1 1 1 1111 1111 1 1 1112116 1

126 A.hastulata 1 1 1 1 1 1 1 5 1 1 1 1 1 1111 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1111 11114 1 1111 116 3

12 7 A chaasmanae 1 1 1 3 1 - 1 6 - 1 1 1 1111 1 1 1 1 1 1 1 1 4 1 1 1 1 1 2 1 1111 1111 3 1 1111 116 1

128 A moluccansis 1 1 1 1 1 1 1 5 1 1 1 1 1 1111 1 1 1 1 1 1 1 1 2 1 1 1 1 1 2 1 1111 1111 3 1 1111 116 1

129 A.obianais 1 1 1 1 1 1 1 5 1 1 1 1 1 1111 1 1 1 2 1 1 1 1 2 1 1 1 1 1 1 1 1111 1111 5 1 1111 116 1

130 A.bullata 1 1 1 2 1 1 1 2 1 1 1 1 1 1111 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 11111111 5 1 1111 116 1

1 6 1 1 131 A dilobata 1 1 1 2 2 1 1 1 1 1 1111 2 1 1 1 1 1 1 1 1 2 1 1 1 2 1 11111111 5 1 1111 116 1

132 Tadipala 1 1 1 1 1 1 1 2 1 1 1 1 1 12 11 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 11111111 5 1 1111 116 1 1 3

T 133 pneumatica 1 1 1 1 1 1 1 3 1 1 1 1 1 12 11 1 1 1 1 2 1 1 1 1 1 1 1 1 1 2 1 11111111 5 1 1111 116 1 1 3

134 T.spalunca 1 1 1 1 1 1 1 3 1 1 1 1 1 1111 1 1 1 1 2 1 1 1 1 1 1 1 1 1 1 1 11111111 1 1 1111 116 2 1 2

135 T.0oboea 1 1 1 1 1 1 1 2 1 1 1 1 1 1111 1 1 1 1 2 1 1 1 1 1 1 1 2 1 1 1 1 11111111 1 1 1111 116 2 1 2

136 T.siciia 1 1 1 1 1 1 1 3 1 1 1 1 1 12 11 1 1 1 1 2 1 1 1 1 1 1 1 2 1 1 1 1 11111111 1 1 1112 116 2 1 1

1 1111 137 T.lancaola 1 1 1 1 1 1 1 2 1 1 1 1 1 1111 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 11111111 1 116 3 1 1

138 T.sarissa 1 1 1 1 1 1 1 2 1 1 1 1 1 1111 1 1 1 1 2 1 1 1 1 1 1 1 2 1 1 2 1 11111111 1 1 1111 116 3 1 1

139 M.viridicata 1 1 1 1 1 1 1 2 1 1 1 1 1 12 11 1 1 1 1 1 1 1 1 2 1 1 1 1 1 1 2 1 11111111 1 1 1111 116 1 1 1

1 1 1 1 1 140 M.globulata 1 1 5 1 1 1 12 11 1 1 1 1 1 1 1 1 2 1 1 1 1 1 1 2 1 11111111 1 1 1111 116 1 1 1

141 M.inconspicua 1 1 1 1 1 1 1 5 1 1 1 1 1 12 11 1 1 1 1 1 1 1 1 2 1 1 1 2 1 1 1 1 2 111 1111 1 1 1111 116 1 1 1

142 M.humilis 1 1 1 1 1 1 1 5 1 1 1 1 1 12 11 1 1 1 1 1 1 1 1 2 1 1 1 2 1 1 1 1 2 111 1111 1 1 1111 116 1 1 1

143 M infla ta 1 1 1 1 1 1 1 2 1 1 1 1 1 12 11 1 1 1 1 1 1 1 1 2 1 1 1 1 1 1 1 1 2 111 1111 1 1 1111 116 1 1 1

144 1 1 1 1 M.toxopausi 1 1 1 1 1 2 1 1 1 12 11 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 2 111 1111 1 1 1111 116 1 1 1

145 C.schmettzi 1 1 2 1 1 1 1 2 1 1 1 1 1 1111 1 1 1 1 2 1 1 1 1 1 1 1 1 11111111 1 1 1111 116 1 1 1

146 C saundarsii 1 1 2 1 1 1 1 2 1 1 1 1 1 1111 1 1 1 1 1 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1111 1 1 1111 116 1 1 1

147 Cimmaculata 1 1 2 1 1 1 1 2 1 1 1 1 1 1111 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1111 1111 1 1 1111 116 1 1 1

148 Prasia 1 1 1 1 1 1 1 1 1 3 1 1 2 1111 1 1 1 1 1 1 1 1 2 1 1 1 1 1 1 2 1 11111111 1 1 1111 116 1 1 1

149 1 1 1 1 1 - - 6 1 2 1111 1 1 1 • KAtda 1 1 3 1 1 2 1 - - - Contributions to Zoology, 65 (4) 1995 231

Appendix 3. Continued.

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 11111111 1 1 1 1 1 11111 1

5 5 5 5 3 3 3 3 3 3 3 4 4 4 4 4 4 character nr 3 3 3 3 3 3 3 5 character nr 3 4 4 4 4 5 5 5 5 5

2 3 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 12 3 4

1 B.guttulipennis 1 1 1 2 1 1 2 1 2 1 1 1 1 1 1 2 2 1 76 G.stenocephalis 1 11111111 1 1 1 1 3 15 4?? 1

2 B.marmorata 1 1 1 2 1 1 2 1 2 1 ? 1 1 1 1 1 4 1 77 G.parvula 1 11111111 1 1 1 1 3 15 4?? 1

4 3 B nasuta 1 1 1 2 1 1 2 1 2 1 2 1 1 1 1 1 1 1 78 G.membrana 1 11111131 1 2 2 1 3 15 11 1

4 B intermedia 1 1 1 2 1 1 2 1 2 1 1 1 1 1 1 ? ? 1 79 G.minoramembrant 1 1 11111131 1 2 1 1 3 15 4 11 1

5 B.splendida 1 1 1 2 1 1 2 1 2 1 1 1 1 1 1 1 4 1 8 0 G.phyloglycera 1 11111131 1 2 1 1 3 15 4 11 1

6 B. parva 1 1 1 2 1 1 2 1 2 1 2 1 1 1 1 1 1 1 8 1 G.langeraki 1 11111131 1 2 1 1 3 15 4 11 1

7 B.mamillata 1 1 1 2 1 1 2 1 2 1 2 1 1 1 1 1 4 1 8 2 G.hirsuta 1 11111113 1 2 1 1 3 14 4 11 1

8 Barabuensis 1 1 1 2 1 1 2 1 2 1 2 1 1 1 1 1 G 1 8 3 G.olivácea 1 11111113 1 1 1 1 3 14 4 11 1

9 B.gibberosa 1 1 1 2 1 1 2 1 2 1 2 1 1 1 1 1 3 1 8 4 G.verlaani 1 11111113 1 1 1 1 3 14 4 11 1

1 0 B bipunctata 1 1 1 2 1 1 2 1 2 1 1 1 1 1 1 1 4 1 8 5 G.montana 1 11111113 1 1 1 1 3 14 4 11 1

11 B laminifer 1 1 1 2 1 1 2 1 2 1 1 1 1 1 1 1 4 1 8 6 G.rufa 1 11111111 1 3 1 1 3 15 4 11 2

1 2 B. retracta 1 1 1 2 1 1 2 1 2 1 1 1 1 1 1 1 4 1 87 G.varicolor 1 11111111 1 1 1 1 3 14 4 11 1

1 3 B.brongersmai 1 1 1 2 1 1 1 2 1 1 1 1 1 2 1 1 1 8 8 V.superba 1 11111111 1 1 2 1 3 15 4 11 2

1 4 B.viridis 1 1 1 2 1 1 1 2 1 1 1 1 2 1 2 1 8 9 C.vitripennis 2 11111111 1 4 1 1 3 1 5 4 2 1 1

1 5 B.lorentzi 1 1 1 2 1 1 1 2 1 1 1 1 1 2 1 1 1 9 0 C.suffusa 2 11111111 1 1 1 1 3 15 3 11 1

16 B.rufula 1 1 1 2 1 1 1 2 1 2 1 1 1 ? ? 1 9 1 C fumea 2 11111111 1 1 1 1 3 15 3?? 1

1 7 B. turgida 1 1 1 2 1 2 1 2 1 1 2 1 1 1 1 5 1 9 2 O.insignis 11111111 1 1 1 1 3 15 3 14 1

1 1 1 8 B.furcillata 1 2 1 2 1 2 1 1 2 1 1 1 ? ? 1 93 Gl.viridis 11111111 1 1 1 1 3 15 4?? 1

1 9 B.karkarensis 1 1 1 2 1 2 1 2 1 1 2 1 1 1 1 2 1 9 4 G.flava 2 11111111 2 1 1 1 3 15 3 12 1

20 B.convfva 1 1 1 2 2 1 2 1 1 2 1 1 1 1 3 1 9 5 G.flaveola 2 11111111 2 1 1 1 3 15 3 12 1

2 1 B laureli 1 1 1 2 2 1 2 1 1 2 1 1 1 1 3 1 9 6 G.stylata 2 11111111 2 1 1 1 3 15 4 12 1

22 B schulzi 1 1 1 2 2 1 2 1 1 2 1 1 1 1 3 1 9 7 G.viridula 11111111 2 1 1 1 3 15 3 12 1

23 B. quadrifida 1 1 1 2 2 1 2 1 1 1 1 1 1 3 1 9 8 G pallida 11111111 2 1 1 1 3 15 3 11 1

4 2 B.hardyi 1 1 1 2 1 1 2 1 1 1 1 1 1 3 1 9 9 G.pallidula 11111111 2 1 1 1 3 15 3 11 1

25 B.guttulinervis 1 1 1 2 1 1 2 1 1 1 1 1 1 1 1 100 G.chinai 11111111 2 2 1 1 3 13 4 12 1

2 6 B roonensis 1 1 1 2 1 1 2 1 1 1 1 1 1 1 1 1 01 G.permyi 11111111 2 1 1 1 3 1 5 3 ? ? 1

27 B inconstans 1 1 1 2 1 1 2 1 1 1 1 1 1 1 1 P 1111111 1 1 1 1 15 4 102 phyllophora 2 - 12 1

28 B.biroi 1 1 1 2 1 1 2 1 1 1 1 1 1 1 1 103 Pgonktdes 1111111 1 1 2 1 1 3 15 3 11 1

29 B bloetei 1 1 1 2 1 1 2 1 1 1 2 1 1 1 104 P.angulata 11111111 1 2 1 1 3 15 4 12 1

30 B.macgillavryi 1 1 1 2 1 1 2 1 1 1 1 1 1 1 1 105 P.lachlani 11111111 1 1 1 1 3 15 4?? 1

3 1 B papuensis 1 1 1 2 1 1 2 1 1 2 1 1 2 1 1 1 106 P.brassi 11111111 1 2 1 1 3 15 4 12 1

32 B bismarckensis 1 1 1 2 1 1 2 1 1 2 1 1 2 1 1 1 107 P.stoliodes 11111111 1 3 1 1 3 15 4 12 1

33 Bmanusensis 1 1 1 2 1 1 2 1 1 1 1 1 2 1 1 1 108 P.plicata 11111111 1 2 1 1 3 15 4 12 1

3 4 Bmussauensis 1 1 1 2 1 1 2 1 1 1 1 1 2 1 2 1 109 P.spinigera 11111111 1 1 1 1 3 15 4 12 1

3 5 B.brandti 1 1 1 2 1 1 2 1 1 1 1 2 2 1 2 1 110 P.bidigitula 11111111 1 2 1 1 3 15 4 12 1

3 6 B.sedlacekorum 1 1 1 2 1 1 2 1 1 1 1 2 2 1 1 1 111 P. ungula 11111111 1 2 1 1 3 15 4?? 1

3 7 1 1 1 1 1 2 1 1 1 1 2 2 1 1 B.reynhoudti 2 2 112 P nana 11111111 1 2 1 1 3 15 4 11 1

3 8 B.cristovalensis 1 1 1 2 1 1 2 1 1 1 1 2 2 1 2 1 113 P.woodlarkensis 11111111 1 2 1 1 3 15 4 12 1

1 4 1 3 9 B edauberti 1 1 1 2 1 1 2 1 1 1 2 2 1 114 P us tulata 11111111 1 2 1 1 3 15 4 11 1

4 0 1 1 1 1 1 1 1 B maddisoni 1 2 1 2 1 2 2 1 2 115 P.novariae 11111111 1 2 1 1 3 15 3?? 1

4 1 B.gressitti 1 1 1 2 1 1 2 1 1 1 1 2 2 1 2 1 116 P baasi 11111111 1 2 1 1 3 15 4 11 1

4 2 B.bilebanarai 1 1 1 2 1 1 2 1 1 1 1 2 2 1 2 1 117 P dolabrata 11111111 1 2 1 1 3 15 3 11 1

4 3 1 1 1 2 1 2 1 1 1 2 B mendanai 1 1 2 1 2 1 118 P.toxopei 11111111 1 2 1 1 3 15 4 11 1

4 4 B marginata 1 1 1 2 1 1 2 1 1 1 1 2 2 1 2 1 119 P dioedes 1111111 1 1 1 1 1 3 15 4 12 1

4 5 B boulardi 1 1 1 2 1 1 2 1 1 1 1 2 2 1 2 1 120 A.oobrops 2 1111111 1 1 1 1 1 3 2 5 3 1 1 1

4 6 rotumae 1 1 1 2 1 B 1 1 2 1 1 1 2 2 1 2 1 121 A. sepia 2 1111111 1 1 1 1 1 3 2 5 4 ? ? 1

47 B. exhausta 1 1 1 2 1 1 2 1 1 1 1 1 1 1 1 122 A.latifrons 1111111 1 1 3 1 1 3 15 4 12 1

4 8 B.bicolorata 1 1 1 2 1 1 2 1 1 1 1 1 1 1 1 123 A.kaiensis 1111111 1 1 2 1 1 3 1 5 ? 1 2 1

4 9 B cotos 1 1 1 1 1 2 1 1 1 1 se a 2 1 1 1 1 124 A.waigeuensis 1111111 1 1 2 1 1 3 14 4?? 1

50 B rossi 1 1 1 2 1 1 2 1 1 1 1 1 1 1 1 125 A.digitata 1111111 1 1 3 1 1 3 15 4?? 1

51 B vanderhammeni 1 1 1 2 1 1 2 1 1 1 1 1 1 1 1 1 26 A.hastulata 1111111 1 1 2 1 1 3 15 4 1? 1

52 B.wauensis 1 1 1 2 1 1 2 1 1 1 1 1 1 1 127 Acheesmanae 1111111 1 1 2 1 1 3 1 5 4 2 1 1

5 3 B versicolor 1 1 1 2 1 1 2 1 1 1 1 1 1 1 1 128 A moluccensis 1111111 1 1 2 1 1 3 1 5 4 2 1 1

? ? 54 B maai 1 1 1 2 1 1 2 1 1 1 1 1 1 129 A.obiensis 11111111 1 2 1 1 3 14 4 11 1

5 5 B tenuispina 1 1 1 2 1 1 2 1 1 1 1 1 1 1 130 A.bullata 11111111 1 3 1 1 3 15 1?? 1

5 6 B.silveri 1 1 1 2 1 2 1 2 1 1 1 1 1 1 1 1 1 31 A.dilobata 1111111 1 1 3 2 1 3 15 3?? 1

1 57 Bdaviesi 1 1 1 2 2 1 2 1 1 1 1 1 ? ? 1 132 T.adipata 11111111 1 1 1 1 3 15 4?? 1

5 8 B pigrami 1 1 1 2 1 2 1 2 1 1 1 1 1 1 1 1 133 Tpneumabca 11111111 1 3 1 1 3 1 5 4 2 2 1

59 B hartonoi 1 1 1 3 1 2 1 2 1 1 1 1 ? 7 1 1 34 T spelunca 11111121 1 3 2 1 3 1 3 4 2 2 1

60 1 1 1 1 1 1 B.loriae 1 3 2 2 2 1 1 1 1 ? ? 135 T globosa 11111121 1 3 1 1 3 1 3 4 2 1 1

61 B.fortuini 1 1 1 3 1 2 2 1 2 1 1 1 1 1 2 1 1 1 1 36 T.sicula 11111121 1 3 1 1 3 1 3 4 2 2 1

6 2 B.hamiltoni 1 1 1 2 1 2 1 2 1 1 1 1 1 2 1 1 1 137 "Nanceóla 11111111 1 2 1 1 3 1 5 4 2 1 1

6 3 B.bemmeleni 1 1 1 2 1 2 1 2 1 1 1 1 1 2 1 1 1 138 T.sarissa 11111111 1 3 1 1 3 1 5 4 2 2 1

6 4 1 1 2 1 2 1 1 1 1 2 B wegeneri 1 2 1 1 7 ? 1 139 M.viridicata 11111111 1 2 1 1 3 15 4 12 1

6 5 S sahebdivanii 1 1 1 1 1 2 1 1 1 1 1 3 1 5 4 2 1 1 140 M.globulata 11111111 1 1 1 1 3 1 5 4 3 2 1

66 Sbiardae 1 1 1 1 1 2 1 1 1 1 1 3 1 5 4 ? ? 1 141 M.inconspicua 11111111 1 1 1 1 3 15 4 12 1

6 1 1 2 1 1 4 ? ? 1 7 S.huibregtsae 1 1 1 1 1 1 1 3 5 142 M.humilie 11111111 1 1 1 1 3 15 4 12 1

68 G.strepitans 1 1 1 1 1 1 1 1 2 1 2 1 5 4 1 1 1 143 M.inflata 11111111 1 2 1 1 3 15 4 12 1

G.viridis 69 1 1 1 1 1 1 1 1 2 1 2 1 5 4 1 2 1 144 M.toxopeusi 11111111 1 1 1 1 3 1 5 ? 1 2 1

1 1 70 Gdahli 1 1 1 1 1 1 2 1 2 1 5 4 1 1 1 145 C.schmeltzi 11111111 1 1 1 1 3 15 4?? 1

71 G.rubricata 1 1 1 1 1 1 1 1 2 1 2 1 5 4 3 1 1 146 C.saundersii 11111111 1 3 1 1 3 15 4 12 1

G stridere 1 1 1 72 1 1 1 1 1 3 1 1 1 1 3 1 5 4 1 1 1 147 C immaculata 1111111 3 1 3 15 3 12 1

73 G obiensis 1 1 1 1 1 3 1 1 2 1 1 3 1 5 4 1 1 1 148 Prasia 2 1 1 1 1 1 - 15 3 12 1

74 G subnotata 1 1 1 1 1 1 ? 1 1 2 1 1 3 1 5 3 2 2 149 Muda 2 1 1 1 1 1 - 1 5 4 3 - 1

7 5 G.nigravirgula 1 1 1 1 1 1 1 1 1 1 1 3 1 5 4 1 2 1