19. Tribe OXYRHACHINI Distant 1908 Old World: Afrotropical ...

19. Tribe OXYRHACHINI Distant 1908 

Old World: Afrotropical, Indomalayan, and Palearctic Regions 

Figs. 19.1-19.3 

Type genus: Oxyrhachis Germar, 1833a 

Oxyrhachisaria Distant, 1908g [new division]: first treated as subfamily Oxyrrhachinae 

[sic: for Oxyrhachinae] and tribe Oxyrrhachini (Haupt 1929c); tribe Oxyrhachisini [sic: 

for Oxyrhachini] Goding 1930b; subfamily Oxyrhachinae equals Centrotinae and tribe 

Oxyrhachini moved to Centrotinae (Dietrich et al. 2001a). 

Xiphistesini Goding, 1930a [new division]: first treated as tribe Xiphistini and equals 

Oxyrhachini (Capener 1962a). 

Diagnostic characters.—Frontoclypeal lobes indistinct, head with large foliate lobes. 

Posterior pronotal process concealing scutellum. Pleuron with propleural lobe present and 

mesopleural lobe enlarged. Forewing with Cu1 vein abutting clavus (not marginal vein), with 

m-cu1 and m-cu2 crossveins in at least one wing, M and Cu veins adjacent at base, base of 

R2+3 and R4+5 veins truncate. Hind wing with R4+5 and M1+2 veins fused or not (3 or 4 apical 

cells). Tibiae foliaceous. Mesothoracic and metathoracic femora without ab- and adlateral 

cucullate setae. Metathoracic tibial rows I and III without cucullate setae (row II without 

cucullate setae in some species). Female second valvulae short with undulating dorsal 

margin, narrow near base, not curved, dorsal margin with fine teeth. Male style clasp 

oriented laterally, apex membranous, cylindrical, angled ventrally. Abdomen with paired 

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dorsal swellings, larger in posterior segments; acanthae distinct, bases not heightened, 

acanthae without ornamentation. 

Description.—Length 5-6.3 mm. Color tan to dark brown, or combinations thereof. 

HEAD (Fig. 19.1 I): frontoclypeal margins parallel or slightly converging ventrally, 

frontoclypeal lobes indistinct; with large foliate lobes; ocelli about equidistant from each 

other and eyes; vertex without toothlike projections. THORAX: PRONOTUM (Figs. 19.1 A- 

H): suprahumeral horns present or absent; posterior process straight at base, appressed 

against scutellum, significantly extending past m-cu3 crossvein in forewing. SCUTELLUM: 

emarginate with apices acute, concealed by posterior process; shortened--with abdomen 

removed, notch and apices not visible. PLEURON: propleural lobe present, mesopleural lobe 

enlarged. FOREWING (Figs. 19.1 J): hyaline; apical limbus broad; s crossvein distad of r-m2 

crossvein; Cu1 vein abutting clavus (not marginal vein); m-cu1 and m-cu2 crossveins present 

in at least 1 wing; M and Cu veins adjacent at base; R and M veins not confluent preapically; 

R1 vein not perpendicular to marginal vein; r-m1 crossvein originating anterior to first 

division of R vein, parallel to longitudinal veins or bent towards R vein; R, M, and Cu veins 

not parallel apically; R4+5 vein shape prior to s crossvein significantly angled or not; base of 

R2+3 and R4+5 veins truncate. HIND WING (Fig. 19.1 K): R4+5 and M1+2 veins fused or not (3 or 

4 apical cells). PRO- AND MESOTHORACIC LEGS: tibiae foliaceous; mesothoracic tibia without 

row(s) of cucullate setae; mesothoracic femur without ab- and adlateral cucullate setae. 

METATHORACIC LEG (Figs. 19.2 A-B): ventral margin of coxa, trochanter, and femur without 

enlarged setal bases; femur without ab- and adlateral cucullate setae; femur without ablateral 

cucullate setae ventrolaterally; tibia foliaceous, rows I and III without cucullate setae; row II 

with or without cucullate setae, if present, in single row; tarsomere I with 1 cucullate seta or 

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none. ABDOMEN: in anterior aspect (abdomen removed) nearly triangular; anterior tergal 

borders not modified; sternal longitudinal carina absent; paired dorsal swellings present, 

larger in posterior segments; tergum III ventrolateral margin carinate; abdominal setal bases 

not enlarged. FEMALE GENITALIA (Figs. 19.3 A-D): second valvulae short with undulating 

dorsal margin, narrow near base, not curved, dorsal teeth fine in size, acute projections on 

dorsal margin absent; third valvulae without small ventral conelike projections. MALE 

GENITALIA (Figs: 19.3 E-K) lateral plate with short dorsoapical lobe extending laterally (Fig. 

19.3 J) or vertically, without ventral lobe; subgenital plate without distinct division; style 

clasp (Fig. 19.3 E-F) oriented laterally, membranous, cylindrical, angled ventrally; style 

shank arched, apex at midpoint or past midpoint. ABDOMINAL FINE STRUCTURE (Figs. 19.2 

C-D): acanthae distinct, bases not heightened, acanthae without ornamentation. 

Chromosome numbers.—Male 2n= 21 (Table 26.3). 

Distribution.—The tribe Oxyrhachini is recorded from the Afrotropical, 

Indomalayan, and Palearctic Regions (McKamey 1998a). 

Ecology.—Members of the tribe Oxyrhachini are reported from the host plant 

families Amaranthaceae, Balanitaceae, Bignoniaceae, Buddlejaceae, Casuarinaceae, 

Compositae, Ebenaceae, Euphorbiaceae, Gramineae, Lauraceae, Leguminosae, Moraceae, 

Myrtaceae, Proteaceae, Rhamnaceae, Santalaceae, Solanaceae, Sterculiaceae, Tamaricaceae, 

Ulmaceae, and Verbenaceae. Oxyrhachis is the only centrotine genus reported from the 

family Balanitaceae (Table 26.2). Oxyrhachis is reported to be tended by ants (Table 26.1). 

Some species of Oxyrhachis are gregarious intra- and interspecifically as both nymphs and 

adults (Ananthasubramanian 1996a). In addition, certain species of Oxyrhachis show 

285

parental care by guarding egg masses and nymphs from predators and parasitoids 

(Ananthasubramanian 1996a). 

Discussion.—The tribe Oxyrhachini, represented by the genus Oxyrhachis, is a very 

morphologically distinct, geographically widespread, and speciose group. The genus 

Oxyrhachis is among the four largest membracid genera, with 117 species (McKamey 

1998a). Oxyrhachini, although monotypic, is here retained as a valid tribe due to the 

numerous character changes on the phylogenetic tree (Fig. 24.1) and the size of the lineage. 

The genus Oxyrhachis is perhaps best known morphologically for its large foliate lobes on 

the head that closely border the frontoclypeus. Oxyrhachini was historically placed in the 

Membracinae by many workers including Stål (1866a), Distant (1908g), Goding (1931a), and 

Metcalf and Wade (1965a). Funkhouser (1951a), however, included the Oxyrhachini within 

the Centrotinae. 

Distant’s (1908g) key characteristics are still valuable diagnostic features for the 

tribe: oxyrachine treehoppers have a long and narrow posterior process, the tibiae are 

foliaceous, and the pro- and mesosterna have distinct tooth-like processes. Along with these 

features, all oxyrhachines examined have dorsal abdominal swellings which are larger in the 

posterior portion of the abdomen. This is in contrast to nessorhinines and gargarines where 

the dorsal abdominal processes are more distinct in the anterior portion of the abdomen. 

Unlike other centrotines, the oxyrhachines examined here have Cu1 vein abutting the clavus 

in the forewing, or the junction between the clavus and the marginal vein, rather than clearly 

abutting the marginal vein. Additionally, M and Cu veins in the forewing are clearly 

adjacent in oxyrhachine forewings and m-cu1 and m-cu2 crossveins are present. 

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Although most oxyrhachine species are morphologically homogenous, certain 

characters vary among species. Some species, including O. carinata and O. sulicornis, have 

R4+5 and M1+2 veins in the hind wing fused (3 apical cells) while in other species these veins 

are not fused (4 apical cells). Apparently, according to Capener (1962a), the number of 

species with 3 apical cells and 4 apical cells in the hind wing is about equal. The presence of 

cucullate setae in metathoracic tibial row II is also variable among species of Oxyrhachis. 

Indeed, Oxyrhachis formerly was split into 6 genera to accommodate the differences in hind 

wing venation and pronotal features, including the presence or absence of suprahumeral 

horns. Based on a phylogenetic analysis (Fig. 24.12) of three former genera (Gongroneura 

Jacobi, Kombazana Distant, and Xiphistes Stål) and Oxyrhachis, however, there are too few 

morphological differences to defend splitting Oxyrhachis into multiple genera at this time. A 

phylogenetic analysis of Oxyrhachis, using generic morphological characters and molecular 

methods, is recommended to better determine its taxonomic and phylogenetic limits. 

The reduction in rank from subfamily Oxyrhachinae to tribe Oxyrhachini (within the 

Centrotinae) by Dietrich et al. (2001a) based on a morphological phylogenetic analysis is 

supported by the phylogenetic analysis of the Centrotinae presented here (Fig. 24.1). The 

Oxyrhachini are closely related to the tribes Ebhuloidesini, Hypsaucheniini, and Terentiini. 

Apparently, the Oxyrhachini are sister group to the Hypsaucheniini. The Ebhuloidesini, 

Hypsaucheniini, and Oxyrhachini share similarly shaped female second valvulae and male 

clasps, and all have enlarged pleural projections. This diagnostic male clasp was described 

as resembling the “head of a snake” by Capener (1962a). Although the Oxyrhachini are 

sister group to the Nessorhinini in the phylogenetic analysis of Dietrich et al. (2001a), this 

relationship was thought to be an artifact of the small number of centrotines sampled. It is 

287

critical that future molecular higher level phylogenetic analyses of the Membracidae include 

the Oxyrhachini in order to investigate further their relationship with the morphologically 

extreme tribe Hypsaucheniini and predominantly Australian Terentiini. 

Genera of the tribe Oxyrhachini 

Oxyrhachis Germar, 1833a (type species: Membracis taranda Fabricius by monotypy). 

Specimens examined. —Oxyrhachis carinata (Funkhouser), det. A.L. Capener, 

AMNH, #00-175b%, #00-175c&; O. delalendei Fairmaire, det. A.L. Capener, AMNH, #00- 

175d%, #00-175e&; O. sulcicornis (Thunberg), det. A.L. Capener, NCSU, #81-48a%, #81- 

48b&; O. taranda (Fabricius), det. Z.P. Metcalf, NCSU, #99-82a&, #99-97a%, #99-97b&, 

#01-260a& —as det. in NCSU, #99-82b%—as det. in USNM, #01-54c%. 

288

Fig. 19.1. Oxyrhachini: pronota (lateral aspects, A-D; and anterior aspects, E-H), heads (I), and 

wings (J-K). Bars = 3 mm. A, Oxyrhachis carinata (Funkhouser), #00-175c&. B, O. delalendei 

Fairmaire, #00-175d%. C, O. sulcicornis (Thunberg), #81-48a%. D, O. taranda (Fabricius), #99- 

97a%. E, O. carinata (Funkhouser), #00-175c&. F, O. delalendei Fairmaire, #00-175d%. G, O. 

sulcicornis (Thunberg), #81-48a%. H, O. taranda (Fabricius), #99-82a&. I, O. taranda 

(Fabricius), #99-82a&. J, O. taranda (Fabricius), #99-82b%, right forewing. K, O. taranda 

(Fabricius), #99-82b%, left hind wing (inverted). fo, foliate lobes. 

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Fig. 19.2. Oxyrhachini: metathoracic legs (A-B) and maximum 

development of abdominal fine-structure (C-D). All scanning 

electron micrographs near tergum III. A, Oxyrhachis delalendei 

Fairmaire, #00-175e&. B, O. taranda (Fabricius), #99-82a&. 

C-D, O. taranda (Fabricius), #01-54c%. i, inornate pit. l, 

lateral seta. 

290

Fig. 19.3. Oxyrhachini: female second valvulae (A-D, lateral aspects and closeup of apex), 

and male styles (E-F, lateral aspects), aedeagi (G-H, lateral aspects), lateral plates (I-J, 

lateral aspects), and subgenital plate (K, ventral aspect). A, Oxyrhachis carinata 

(Funkhouser), #00-175c&. B, O. delalendei Fairmaire, #00-175e&. C-D, O. taranda 

(Fabricius), #99-82a&. E, O. carinata (Funkhouser), #00-175b%. F, O. taranda 

(Fabricius), #99-82b%. G, O. carinata (Funkhouser), #00-175b%. H, O. taranda 

(Fabricius), #99-82b%. I, O. carinata (Funkhouser), #00-175b%. J, O. taranda (Fabricius), 

#99-82b%. K, O. carinata (Funkhouser), #00-175b%. c, clasp. dl, dorsoapical lobe. 

291

20. Tribe PIELTAINELLINI, new tribe 

New World: Neotropical Region 

Figs. 20.1-20.2 

Type genus: Pieltainellus Peláez, 1970a 

Diagnostic characters.— Frontoclypeal lobes indistinct and not extending to apex of 

frontoclypeus. Posterior pronotal process not appressed against scutellum, not significantly 

extending past or not reaching m-cu3 crossvein in forewing. Scutellum not shortened--with 

abdomen removed, notch and apices visible. Forewing with R1 vein perpendicular to 

marginal vein. Hind wing with R4+5 and M1+2 veins not fused (4 apical cells). Mesothoracic 

femur without ab- and adlateral cucullate setae. Metathoracic femur with ab- and adlateral 

cucullate setae; tibia with cucullate setal row II single. Female second valvulae not 

broadened, narrow near base, curved, with many fine dorsal teeth extending to apex, acute 

projections present. Abdominal acanthae (Pieltainellus) distinct, bases heightened, acanthae 

multidentate. Anterior tergal borders modified into irregular ridges. 

Description.—Length 4.3-5.4 mm. Color tan to dark brown, or combinations 

thereof. HEAD (Figs. 20.1 E-F): frontoclypeal margins parallel or slightly converging 

ventrally, frontoclypeal lobes indistinct and not extending to apex of frontoclypeus; ocelli 

about equidistant from each other and eyes; vertex without toothlike projections. THORAX: 

PRONOTUM (Figs. 20.1 A-D): suprahumeral horns present in Spathocentrus (Fig. 20.1 D) and 

polymorphic in Pieltainellus; posterior process straight (Fig. 20.1 A) or curved at base (Fig. 

20.1 B), not appressed against scutellum, not significantly extending past or not reaching m- 

cu3 crossvein in forewing. SCUTELLUM: emarginate with apices acute, not concealed by 

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posterior process, 1 lateral apex or none visible from dorsolateral view; not shortened--with 

abdomen removed, notch and apices visible, only slightly extending beyond thorax. 

PLEURON: propleural lobe absent, mesopleural lobe not enlarged. FOREWING (Figs. 20.1 G- 

H): hyaline or opaque; apical limbus broad; s crossvein distad of r-m2 crossvein; m-cu1 and 

m-cu2 crossveins absent; M and Cu veins fused at base; R and M veins not confluent 

preapically; R1 vein perpendicular to marginal vein; forewing without pterostigma; r-m1 

crossvein originating near or distad of first division of R vein, bent towards R vein; R, M, 

and Cu veins not parallel apically; discoidal cells similar in length; R4+5 vein shape prior to s 

crossvein significantly angled or not; base of R2+3 and R4+5 veins truncate. HIND WING: R4+5 

and M1+2 veins not fused (4 apical cells). PRO- AND MESOTHORACIC LEGS: tibiae not 

foliaceous; mesothoracic tibia without row(s) of cucullate setae; mesothoracic femur without 

ab- and adlateral cucullate setae. METATHORACIC LEG (Fig 20.1 I): ventral margin of coxa, 

trochanter, and femur without enlarged setal bases; femur with ab- and adlateral cucullate 

setae, adlateral cucullate seta preapical; femur without ablateral cucullate seta ventrolaterally; 

tibial row I with 19-26 cucullate setae, row II with 16-25 cucullate setae in single row, row 

III with 23-32 cucullate setae; tarsomere I with 2 or more cucullate setae (usually 2). 

ABDOMEN: in anterior aspect (abdomen removed) nearly triangular or flattened; anterior 

tergal borders modified into irregular ridges; sternal longitudinal carina absent; paired dorsal 

swellings absent; tergum III ventrolateral margin carinate; abdominal setal bases at anterior 

tergal borders not enlarged. FEMALE GENITALIA (Figs. 20.2 A-D): second valvulae not 

broadened, narrow near base, curved, with many fine dorsal teeth extending to apex, acute 

projections present; third valvulae without ventral projections. MALE GENITALIA 

(Pieltainellus, Figs. 20.2 E-J): lateral plate with short dorsoapical lobe extending laterally, 

293

without ventral lobe; subgenital plate without distinct division; style clasp oriented laterally, 

thickened, elliptical, not angled; style shank without significant arch. ABDOMINAL FINE 

STRUCTURE (Pieltainellus) (Fig. 20.2 K): acanthae distinct, bases heightened, acanthae 

multidentate. 

Chromosome numbers.—Unknown. 

Distribution.—The tribe Pieltainellini is recorded from the Neotropical Region 

(Maes 1998a, McKamey 1998a). 

Ecology.— Host plant information for the tribe Pieltainellini is unknown. 

Discussion.—The new tribe Pieltainellini is a monophyletic group in the 

phylogenetic analysis (Fig. 24.1) and is closely related to the tribes Beaufortianini 

Nessorhinini, and Platycentrini. The females of Pieltainellus and Spathocentrus both have 

curved second valvulae with acute projections on the dorsal margin and have R1 vein 

perpendicular to the marginal vein in the forewing. Moreover, unlike many of their relatives, 

the scutellum is fully exposed and is not appressed by the posterior process. In his 

description of Pieltainellus, Peláez (1970a) remarked on the morphological similarities 

between Pieltainellus and Spathocentrus. The genus Pieltainellus was chosen as the type 

genus because both female and male specimens were examined. 

Both Pieltainellus and Spathocentrus were placed in the Boocerini by Deitz (1975a). 

The Pieltainellini lack extra m-cu crossveins and have an exposed scutellum, characteristics 

Deitz used to define the Boocerini. The Pieltainellini, however, lack other boocerini features 

including mesothoracic ablateral cucullate setae on the femur and the long ventral lobe of the 

male lateral plate. 

294

Genera of the tribe Pieltainellini 

Pieltainellus Peláez, 1970a (type species: P. boneti Peláez by original designation) 

[previously placed in Boocerini (McKamey 1998a)]. 

Spathocentrus Fowler, 1896d (type species: S. intermedius Fowler by monotypy) [previously 

placed in Boocerini (McKamey 1998a)]. 

Specimens examined.—Pieltainellus sp., det. S.H. McKamey, SHMC, #01-235f&; 

P. boneti Peláez, det. L.L. Deitz, AMNH, #72-97c&, #72-129a%; Spathocentrus intermedius 

Fowler, holotype, OXUM, #72-296a&—det. S.H. McKamey, SHMC, #00-195a&—as det. in 

CNCI, #01-39b&. 

295

Fig. 20.1. Pieltainellini: pronota (lateral aspects, A-B; and anterior aspects, C-D), heads 

(E-F), wings (G-H), and metathoracic leg (I). Bars = 3 mm. A, Pieltainellus boneti Peláez, 

#72-97c&. B, Spathocentrus intermedius Fowler, #00-195a&. C, P. boneti, #72-129a%. 

D, S. intermedius, #00-195a&. E, P. boneti, #72-97c&. F, S. intermedius, #00-195a&. G, 

P. boneti, #72-129a%, left forewing (inverted). H, S. intermedius, #00-195a&, left 

forewing (inverted). I, P. boneti, #72-97c&, left metathoracic leg. fc, frontoclypeus. 

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Fig. 20.2. Pieltainellini: female second valvulae (lateral aspects and 

closeup of apex, A-D), and male styles (lateral aspect, E; dorsal aspect, 

F; and posterior aspect, G), aedeagus (lateral aspect, H), lateral plate 

(lateral aspect, I), and subgenital plate (ventral aspect, J), and maximum 

development of abdominal fine-structure (K). Scanning electron 

micrograph near tergum III. A-B, Pieltainellus boneti Peláez, #72-97c&. 

C-D, Spathocentrus intermedius Fowler, #00-195a&. E-J, P. boneti, 

#72-129a%. K, P. boneti, #01-235f&. c, clasp. a, acanthus. d, 

dorsoapical lobe. i, inornate pit. l, lateral seta. 

297

21. Tribe PLATYCENTRINI Haupt, 1929 

New World: Nearctic and Neotropical Regions 

Figs. 21.1-21.3 

Type genus: Platycentrus Stål, 1869 

Platycentrini Haupt, 1929c [new tribe]: equals Hebesini (Peláez 1970a)[error]; reinstated 

as tribe Platycentrini, within subfamily Centrotinae (Deitz 1975a); move to 

Membracinae (Kosztarab 1982a)[error]; returned to Centrotinae (Deitz and Dietrich 

1993a). 

Diagnostic characters.—Frontoclypeal lobes indistinct and not extending to apex of 

frontoclypeus. Posterior pronotal process appressed against scutellum, not significantly 

extending past or not reaching m-cu3 crossvein in forewing. Scutellum not concealed by 

posterior process, shortened--with abdomen removed, at most scutellar apices visible. 

Forewing hyaline, with m-cu2 crossvein present and base of R2+3 and R4+5 veins truncate. 

Hind wing with R4+5 and M1+2 veins not fused (4 apical cells). Mesothoracic femur with 

ablateral cucullate setae. Female second valvulae with gradual broadening before midpoint 

and gently tapering after midpoint; narrow near base, not curved, dorsal teeth fine and not 

extending to apex. Abdominal acanthae distinct, bases heightened, acanthae multidentate. 

Description.—Length 4.5-6 mm. Color brown, tan, mottled or combinations thereof. 

HEAD (Fig. 21.1): frontoclypeal margins parallel or slightly converging ventrally, 

frontoclypeal lobes indistinct and not extending to apex of frontoclypeus; ocelli about 

equidistant from each other and eyes; vertex without toothlike projections. THORAX: 

298

PRONOTUM (Figs. 21.1 A-E): suprahumeral horns present or absent; posterior process straight 

at base, appressed against scutellum, not significantly extending past or not reaching m-cu3 

crossvein in forewing. SCUTELLUM: emarginate with apices acute, not concealed by posterior 

process, 1 lateral apex visible from dorsolateral view; shortened--with abdomen removed, at 

most only apices visible. PLEURON: propleural lobe absent, mesopleural lobe not enlarged. 

FOREWING (Figs. 21.1 H, 21.2 A): hyaline; apical limbus broad; s crossvein distad of r-m2 

crossvein; m-cu1 crossvein present (Fig. 21.2 A) or absent; m-cu2 crossvein present; M and 

Cu veins fused at base; R and M veins not confluent preapically; veins reticulate in 

Tylocentrus (Fig. 21.2 A); R1 vein perpendicular to marginal vein (Tylocentrus, Fig. 21.2 A) 

or not (Platycentrus, Fig. 21.1 H), never parallel to longitudinal veins; forewing without 

pterostigma; r-m1 crossvein originating anterior to first division of R vein, bent towards R 

vein; R, M, and Cu veins parallel apically (Fig. 21.1 H) or not (Fig. 21.2 A); discoidal cells 

similar in length (Fig. 21.1 H) or not (Fig. 21.2 A); R4+5 vein shape prior to s crossvein 

significantly angled (Fig. 21.1 H) or not (Fig. 21.2 A); base of R2+3 and R4+5 veins truncate. 

HIND WING (Fig. 21.1 I): R4+5 and M1+2 veins not fused (4 apical cells). PRO- AND 

MESOTHORACIC LEGS: tibiae not foliaceous; mesothoracic tibia without row(s) of cucullate 

setae; mesothoracic femur with ablateral cucullate setae, adlateral cucullate setae present in 

Tylocentrus but absent in Platycentrus. METATHORACIC LEG (Fig. 21.2 B): ventral margin of 

coxa, trochanter, and femur without enlarged setal bases; femur with ab- and adlateral 

cucullate setae; femur with ablateral cucullate setae ventrolaterally in Platycentrus; tibia not 

foliaceous, row I with 15-21 cucullate setae, row II with 15-36 cucullate setae in single 

(Tylocentrus) or irregular or double row (Platycentrus), row III with 17-33 cucullate setae 

(irregular in Platycentrus); tarsomere I with 1 cucullate seta in Platycentrus, 2 or more 

299

cucullate setae (usually 2) in Tylocentrus. ABDOMEN: in anterior aspect (abdomen 

removed) nearly triangular (Tylocentrus) or dorsoventrally flattened (Platycentrus); anterior 

tergal borders modified into irregular ridges in Platycentrus; sternal longitudinal carina 

absent; paired dorsal swellings absent; tergum III ventrolateral margin carinate; abdominal 

setal bases at anterior tergal borders not enlarged. FEMALE GENITALIA (Figs. 21.2 C-F): 

second valvulae with gradual broadening before midpoint and gently tapering after midpoint; 

narrow near base, not curved, dorsal teeth fine and not extending to apex, acute projections 

absent; third valvulae without ventral projections. MALE GENITALIA (Figs. 21.3 A-I): lateral 

plate with (Platycentrus, Fig. 21.3 G) or without (Tylocentrus, Fig. 21.3 H) short dorsoapical 

lobe extending dorsally, without ventral lobe; subgenital plate (Fig. 21.3 I) without distinct 

division; style clasp oriented laterally, thickened, expanding dorsally and laterally with a 

sclerotized ridge (Platycentrus) or truncate with an acuminate projection (Tylocentrus), not 

angled, without basal thickening; style shank without significant arch. ABDOMINAL FINE 

STRUCTURE (Fig. 21.3 J-K): acanthae distinct, bases heightened, acanthae multidentate. 


Distribution.—The tribe Platycentrini is recorded from the Nearctic and Neotropical 

Regions (McKamey 1998a), with all members recorded from the Northern Hemisphere. 

Ecology.—Members of the tribe Platycentrini are reported only from the host plant 

family Leguminosae (Table 26.2). Platycentrus acuticornis is listed as subsocial by Hinton 

(1977a). 

Discussion.— Haupt (1929c) originally placed the Platycentrini within the subfamily 

Stegaspinae (sic: for Stegaspidinae) but misidentified the type genus Platycentrus. Metcalf 

and Wade (1965a), Evans (1948b), and Deitz (1975a) assigned the Platycentrini to the 

300

Centrotinae. Deitz (1975a) included those genera with multiple m-cu crossveins in the 

forewing and with the posterior process abutting a laterally exposed scutellum in the tribe 

Platycentrini. Here, these features are also considered diagnostic. A monophyletic 

Platycentrini (Figs. 24.1, 24.3), as defined here, is supported by the synapomorphy of the 

female second valvulae broadening gradually at midpoint and tapering gently towards the 

apex. A phylogenetic analysis of the family Membracidae using two nuclear genes also 

resulted in a monophyletic Platycentrini (Platycentrus and Tylocentrus) with high bootstrap 

and Bremer support (Cryan et al. 2000a). The Platycentrini are closely related to the New 

World tribes Nessorhinini and Pieltainellini and the Old World tribe Beaufortianini. 

Two genera formerly placed in the Platycentrini are here referred to the tribes 

Monobelini (Monobelus) and Nessorhinini (Orthobelus). See the discussions of these tribes 

for evidence supporting the new placement. 

Genera of the tribe Platycentrini 

Platycentrus Stål, 1869c (type species: P. acuticornis Stål by subsequent designation). 

Tylocentrus Van Duzee, 1908a (type species: T. reticulatus Van Duzee by monotypy). 

Specimens examined.— Platycentrus acuticornis Stål, as det. in USNM, #71-82e&, 

#71-82f%, #99-167e%—det. W.D. Funkhouser, USNM, #71-299a%, #01-221b%—det. L.M. 

Russell, USNM, #99-167d&; Tylocentrus quadricornis Funkhouser, as det. in USNM, #01- 

232b&; T. reticulatus Van Duzee, as det. in USNM, #00-195b&, #00-195c%—det. L.M. 

Russell, USNM, #00-195k[n]. 

301

Fig. 21.1. Platycentrini: pronota (lateral aspects, A-B; anterior aspects, C-E), 

heads (F-G), and wings (H-I). Bars = 3 mm. A, Platycentrus acuticornis Stål, 

#71-82f%. B, Tylocentrus reticulatus Van Duzee, #00-195b&. C, P. acuticornis, 

#71-82f%. D, T. reticulatus, #00-195b&. E, T. reticulatus, #00-195c%. F, P. 

acuticornis, #71-82f%. G, T. reticulatus, #00-195b&. H, P. acuticornis, #99- 

167d&, right forewing. I, P. acuticornis, #99-167d&, left hind wing (inverted). 

fcl, frontoclypeal lobes. 

302

Fig. 21.2. Platycentrini: wings (A), metathoracic leg (B), and 

female second valvulae (lateral aspect and closeup of apex, C-F). A, 

Tylocentrus reticulatus Van Duzee, #00-195b&, left forewing 

(inverted). B, Platycentrus acuticornis Stål, #71-82f%, right 

metathoracic leg (inverted). C-D, P. acuticornis, #99-167d&. E-F, 

T. reticulatus, #00-195b&. 

303

Fig. 21.3. Platycentrini: male styles (lateral aspects, A-B; and dorsal 

aspects, C-D), aedeagi (lateral aspects, E-F), lateral plates (lateral aspects, 

G-H), and subgenital plate (ventral aspect, I), and maximum development 

of abdominal fine-structure (J-K). All scanning electron micrographs near 

tergum III. A, Platycentrus acuticornis Stål, #99-167e%. B, Tylocentrus 

reticulatus Van Duzee, #00-195c%. C, P. acuticornis, #99-167e%. D, T. 

reticulatus, #00-195c%. E, P. acuticornis, #99-167e%. F, T. reticulatus, 

#00-195c%. G, P. acuticornis, #99-167e%. H, T. reticulatus, #00-195c%. 

I, T. reticulatus, #00-195c%. J, P. acuticornis #01-221b%. K, T. 

quadricornis Funkhouser, #01-232b&. a, acanthus. dl, dorsoapical lobe. 

i, inornate pit. c, clasp. 

304

22. Tribe TERENTIINI Haupt, 1929 

Old World: Australasian and Oceanian, Indomalayan, and Palearctic Regions 

Figs. 22.1-22.24 

Type genus: Terentius Stål 1866a 

Terentiinae Haupt, 1929c [new subfamily] and Terentiini Haupt, 1929c [new tribe]: 

subfamily Terentiinae equals Centrotinae and tribe Terentiini moved to Centrotinae 

(Metcalf and Wade 1965a); elevated to subfamily Terentiinae (Evans 1966a) [error]; 

equals Centrotinae (Evans 1966a). 

Bulbaucheniini Goding, 1931a [new tribe]: herein equals Terentiini, NEW SYNONYMY. 

Funkhouserellini Yuan and Zhang, in Yuan and Chou 2002a [new tribe]: herein equals 

Terentiini, NEW SYNONYMY. 

Diagnostic characters.—Frontoclypeal margins parallel or slightly converging 

ventrally. Frontoclypeal lobes distinct. Posterior pronotal process straight at base, appressed 

against scutellum and significantly extending past m-cu3 crossvein in forewing (exception: 

posterior process not extending past m-cu3 crossvein in Pyrgonota). Scutellum shortened-- 

with abdomen removed, at most only scutellar apices visible. Forewing with or without m- 

cu1 and m-cu2 crossveins; r-m1 crossvein originating anterior to first split of R vein, parallel 

to longitudinal veins (exception: r-m1 crossvein bent strongly towards R vein in Alocebes); 

base of R2+3 and R4+5 veins truncate (exceptions: base of R2+3 and R4+5 veins truncate or acute 

in Sextius and some species of Bulbauchenia). Hind wing with R4+5 and M1+2 veins not fused 

(4 apical cells) (exceptions: R4+5 and M1+2 veins fused in Bucktoniella). Mesothoracic femur 

305

without ab- and adlateral cucullate setae. Metathoracic tibial row I cucullate or not; row II in 

single row (exceptions: row II irregular or double in Otinotoides, Sertorius and Yangupia). 

Male style clasp laterally oriented, thickened dorsally and membranous ventrally, quadrate 

(with acuminate apex in Bulbauchenia and Pyrgonota), angled ventrally; style shank with 

significant arch following midpoint, and with ventral preapical broadening. 

Description.—Length 2.6-8.0 mm. Color black, dark brown, light green, light 

yellow, or combinations thereof. HEAD (Figs. 22.5-22.7): frontoclypeal margins parallel or 

slightly converging ventrally, frontoclypeal lobes distinct, not extending to apex of 

frontoclypeus (exceptions: frontoclypeal lobes extending nearly to apex in Bulbauchenia, 

Fig. 22.5 O, Funkhouserella, Figs. 22.6 I-J, and Pyrgonota, Fig. 22.7 C); ocelli about 

equidistant from each other and eyes (exception: ocelli closer to eyes than each other in 

Pyrgonota, Fig. 22.7 C); vertex without toothlike projections. THORAX: PRONOTUM (Figs. 

22.1-22.5): suprahumeral horns present dorsolaterally, absent, or present at apex of median 

anterior horn; median anterior horn present in Bulbauchenia (Fig. 22.3 L), Eutryonia (Fig. 

22.4 E), Funkhouserella (Figs. 22.4 G-H), and Pyrgonota (Fig. 22.5 A); posterior process 

straight at base, appressed against scutellum, signficantly extending past m-cu3 crossvein 

(exception: posterior process not extending past m-cu3 crossvein in Pyrgonota). SCUTELLUM 

(Fig. 22.23 B): emarginate with apices acute, not concealed by posterior process (exceptions: 

scutellum concealed by posterior process in Bulbauchenia, Fig. 22.1 H, and polymorphic in 

Sextius); 1 lateral apex visible from dorsolateral view; shortened--with abdomen removed, at 

most only apices visible, only slightly extending beyond thorax. PLEURON: propleural lobe 

present or absent, mesopleural lobe enlarged or not. FOREWING (Figs. 22.8-22.11): hyaline 

or opaque; apical limbus broad (exceptions: apical limbus narrow in Anzac, Fig. 22.9 C, and 

306

some species of Funkhouserella, Fig. 22.10 F, and Neosextius); R vein initial division R1+2+3 

and R4+5 in Cebes (Fig. 22.9 G), Ceraon (Fig. 22.9 H), Matumuia, and Sarantus (Fig. 22.11 

E); s crossvein distad of r-m2 crossvein; m-cu1 and m-cu2 crossveins present or absent; M and 

Cu veins fused, adjacent, or separate at base; R and M veins not confluent preapically; R1 

perpendicular to marginal vein or not; forewing without pterostigma; r-m1 crossvein 

originating anterior to first split of R vein, parallel to longitudinal veins (exception: r-m1 

crossvein bent strongly towards R in Alocebes); R, M, and Cu veins parallel apically or not; 

R4+5 vein shape prior to s crossvein variable; base of R2+3 and R4+5 veins truncate (exceptions: 

base of R2+3 and R4+5 veins truncate or acute in Sextius and some species of Bulbauchenia). 

HIND WING (Fig. 22.8 B): R4+5 and M1+2 veins not fused (4 apical cells) (exception: hind wing 

with R4+5 and M1+2 fused in Bucktoniella, 3 apical cells). PRO- AND MESOTHORACIC LEGS: 

tibiae foliaceous or not; mesothoracic tibia without row(s) of cucullate setae; mesothoracic 

femur without ab- and adlateral cucullate setae. METATHORACIC LEGS (Fig. 22.12): ventral 

margin of coxa, trochanter, and femur without enlarged setal bases; femur with ab- and 

adlateral cucullate setae (exceptions: femur without ab- and adlateral cucullate setae in 

Anzac, Goddefroyinella, Pogonotypellus, Sextius; adlateral cucullate setae absent in Cebes, 

some species of Funkhouserella, and Pyrgonota, Fig. 22.12 B); femur with or without 

ablateral cucullate setae ventrolaterally; tibiae foliaceous or not, row I with 13-40 cucullate 

setae or without cucullate setae, row II with 4-56 cucullate setae in single row (exceptions: 

row II irregular or double in Otinotoides, Sertorius, and Yangupia), row III with 5-30 

cucullate setae; tarsomere I with 1 cucullate seta or none. ABDOMEN: in anterior aspect 

(abdomen removed) nearly triangular; anterior tergal borders not modified; sternal 

longitudinal carina present or absent; paired dorsal swellings absent; tergum III ventrolateral 

307

margin carinate or shelflike; abdominal setal bases at anterior tergal borders enlarged in 

Terentius, not dispersed on terga. FEMALE GENITALIA (Figs. 22.13-22.17): second valvulae 

shape variable; narrow or broadening near base, curved or not, dorsal teeth variable in size, 

acute projections on dorsal margin present or absent, third valvulae without ventral 

projections. MALE GENITALIA (Figs. 22.17-22.21): lateral plate with or without (Fig. 22.20 

D) short (Fig. 22.20 F) or long (Fig. 22.20 E) dorsoapical lobe extending dorsally or laterally, 

without ventral lobe; subgenital plate without distinct division; style clasp laterally oriented, 

thickened dorsally and membranous ventrally, quadrate (with acuminate apex in 

Bulbauchenia Figs. 22.17 I-J, and Pyrgonota, Fig. 22.18 F), angled ventrally; style shank 

with significant arch following midpoint, and with ventral preapical broadening. 

ABDOMINAL FINE STRUCTURE (Figs. 22.21-22.24): inornate pits with lateral setae present 

(exceptions: inornate pits indistinct in Bulbauchenia, Fig. 22.22 B-D, Funkhouserella, Figs 

22.23 C-D, and Pyrgonota, Fig. 22.23 H), acanthae distinct or not; bases heightened or not; 

acanthae multidentate or divided into threadlike microtrichia. 

Chromosome numbers.—Male 2n= 21 (Table 26.3). 

Distribution.—The tribe Terentiini is recorded from the Australasian and Oceanian, 

Indomalayan, and Palearctic Regions, although they are primarily Australasian in distribution 

(McKamey 1998a). Melichar’s (1905a) records of Acanthuchus trispinifer (Fairmaire) [as 

Ophicentrus trispinifex Fairmaire] in Tanzania needs to be verified and is not here included 

in the distribution of Acanthuchus. 

Ecology.—Members of the tribe Terentiini are reported from the host plant families 

Casuarinaceae, Chenopodiaceae, Euphorbiaceae, Gramineae, Lauraceae, Lecythidaceae, 

Leguminosae, Malvaceae, Moraceae, Myrtaceae, Plumbaginaceae, Polygonaceae, Proteaceae, 

308

Rosaceae, Rutaceae, Solanaceae (Table 26.2). The genus Sextius and the nymphs of 

Australian treehoppers are reported to be tended by ants (Table 26.1). In addition, Terentius 

and Pyrgonota have been observed providing maternal care in the form of egg guarding 

(Stegmann and Linsenmair 2002a). 

Discussion.—Haupt (1929c) divided the Old World Membracidae into three 

subfamilies: the Terentiinae, Centrotinae, and Oxyrhachinae (as Oxyrrhachinae). He 

originally included the tribes Hypsaucheniini and Terentiini within his subfamily Terentiinae. 

Haupt defined the Terentiinae as treehoppers with M and Cu veins in the forewing usually 

connected for a short distance and sometimes united by crossveins, and with the forewing 

often net-like. The tribe Terentiini, as defined here, is monophyletic in the phylogenetic 

analysis (Figs. 24.1, 24.14). All terentiine males observed to this point have a quadrate style 

clasp, a synapomorphy for the tribe. Day (1999a) also commented on the homogenous 

nature of the male genitalia in Australian membracids. Additionally, all terentiines have the 

posterior process appressed against a shortened scutellum, similar to the Gargarini. As seen 

in some Gargarini, a few terentiines, including Bulbauchenia and some Sextius species, have 

the scutellum concealed by the posterior process. The Terentiini are closely related to the 

Ebhuloidesini, Oxyrhachini, and Hypsaucheniiini, and are primarily distributed in Australia. 

The classification presented here largely confirms previous studies on the Australian 

treehopper fauna. Both Evans (1966a) and Day (1999a) examined the Australian genera and 

believed they represent a distinctive group, long isolated from other membracids. Day cited 

the distinctive frontoclypeal lobes, the similar shapes in the male and female genitalia, and 

uniform wing venation, among other features, as evidence for the monophyly of Australian 

genera. A phylogeny of treehoppers based on two nuclear genes (Cryan et al. 2000a) 

309

supported the grouping of the Australian genera Ceraon, Eufairmairia, and Sextius (although 

included in the analysis, Pyrgonota was not part of this group) using both parsimony and 

maximum likelihood methods. 

The names Bulbaucheniini Goding, 1931a and Funkhouserellini Yuan and Zhang, in 

Yuan Chou, 2002a, are here considered junior synonyms of Terentiini Haupt, 1929c, NEW 

SYNONYMIES, based on the phylogenetic analysis (Fig. 24.1). The tribe Bulbaucheniini 

was originally placed in the subfamily Membracinae by Goding (1931a) because of the 

concealed scutellum and folicaeous tibiae of the type genus Bulbauchenia. Goding 

characterized the tribe, consisting only of the type genus, based on the broadly elevated 

pronotum with suprahumeral horns at the apex. Strümpel (1972a) believed the 

Bulbaucheniini belonged in the subfamily Centrotinae, where it was subsequently placed, 

although he was not explicit in his justification. Due to the quadrate shape of the male style 

clasp, Bulbauchenia is here placed in the Terentiini. 

Yuan and Zhang, in Yuan and Chou (2002a) erected the Funkhouserellini, including 

the genera Funkhouserella and Hybandoides, for those treehoppers with the median pronotal 

horn inclined anteriorly and without teeth on the mesonotum Hybandoides is here referred 

back to the tribe Hypsaucheniini, based on the phylogenetic analysis (Fig. 24.12). Although 

no males of Funkhouserella were examined, according to the phylogenetic analysis (Fig. 

24.1) this genus is closely related to Bulbauchenia and Pyrgonota based on several 

morphological features, including the indistinct abdominal inornate pits and absence of 

metathoracic tibia cucullate setal row I. The male style clasp of Pyrgonota, similar to 

Bulbauchenia, is quadrate with an acuminate apex. This shape differs significantly from the 

cylindrical style clasp of the hypsaucheniines where Pyrgonota was previously placed. In 

310

addition, Pyrgonota has cucullate setae in rows II and III of the metathoracic tibia. 

Conversely, these rows are non-cucullate in the Hypsaucheniini. 

The genera Crito and Otinotoides were formerly placed in the Centrotypini for 

reasons that are not clear. They differ from centrotypines, nonetheless, in the style clasp 

shape and leg chaetotaxy. Arimanes, Polonius, and Sarantus were previously placed in the 

Leptocentrini. The male style clasp of Sarantus is distinctly quadrate and not triangular. 

Although males of Arimanes and Polonius were not examined, characteristics of the head, 

pronotum, and leg chaetotaxy differentiate them from leptocentrine genera. The remaining 

terentiini genera were previously assigned to Centrotinae, incertae sedis (McKamey 1998a, 

Day 1999a). 

Four NEW COMBINATIONS are proposed here, all referred from the genus 

Emphusis Buckton: Bulbauchenia bakeri (Funkhouser), B. rugosa (Funkhouser), B. globosa 

(Funkhouser), and B. kurosawai (Hayashi and Endo). These species were originally placed 

in Emphusis (Centrotinae: Centrotypini) based on the elevated pronotum and shape of the 

posterior process. The genus Emphusis, however, has ab- and adlateral cucullate setae on the 

mesothoracic femur and an elliptical clasp as opposed to the characteristic terentiine quadrate 

clasp. 

311

Genera of the tribe Terentiini 

† no specimen examined 

* placement based on morphological similarity 

Acanthucalis Evans, 1966a (type species: A. macalpini Evans by original designation) 

[previously incertae sedis (McKamey 1998a)].* 

Acanthuchus Stål, 1866c (type species: Centrotus trispinifer Fairmaire by subsequent 

designation) [previously incertae sedis (McKamey 1998a)]. 

Alocanthella Evans, 1966a (type species: A. fulva Evans by original designation) [previously 

incertae sedis (McKamey 1998a)]. 

Alocebes Evans, 1966a (type species: A. dixoni Evans by original designation), see figs. 72- 

74, 196 of Day (1999a: 649, 733) [previously incertae sedis (McKamey 1998a)].†* 

Alosextius Evans, 1966a (type species: Acanthuchus carinatus Funkhouser by original 


Anzac Distant, 1916d (type species: Membracis bipunctata Fabricius by original designation) 

[previously incertae sedis (McKamey 1998a)]. 

Arimanes Distant, 1916e (type species: A. doryensis Distant by monotypy) [previously 

placed in the Leptocentrini (McKamey 1998a)].* 

Bucktoniella Evans, 1966a (type species: Acanthuchus pyramidatus Funkhouser by original 


Bulbauchenia Schumacher, 1915b (type species: B. taiwanensis Schumacher by original 

designation) [previously placed in Bulbaucheniini (McKamey 1998a)]. 

312

Bunyella Day, 1999a (type species: Acanthuchus dromedarius Kirkaldy by original 

designation), see figs. 23, 84-86, 200 of Day (1999a: 633, 657, 734) [previously incertae 

sedis (Day 1999a)]. †* 

Cebes Distant, 1916d (type species: Centrotus transiens Walker by subsequent designation) 


Ceraon Buckton, 1903a (type species: Centrotus tasmaniae Fairmaire by subsequent 

designation) [previously incertae sedis McKamey 1998a)]. 

Crito Distant, 1916d (type species: C. festivum Distant by monotypy) [previously placed in 

Centrotypini (McKamey 1998a)].* 

Dingkana Goding, 1903a (type species: D. borealis Goding by original designation) 


Eufairmairia Distant, 1916d (type species: Centrotus decisus Walker by original designation) 


Eufairmairiella Evans, 1966a (type species: Sertorius curvicaudus Goding by original 

designation) [previously incertae sedis (McKamey 1998a)].* 

Eufrenchia Goding, 1903a (type species: Centrotus falcatus Walker by original designation) 


Eutryonia Goding, 1903a (type species: Centrotus monstrifer Walker by monotypy) 


Evansiana McKamey, 1994a (type species: Acanthuchus iasis Kirkaldy by original 


Funkhouserella Schmidt, 1926d (type species: Pyrgonota pinguiturris Funkhouser by 

original designation) [previously placed in Funkhouserellini (Yuan and Chou 2002a)]. 

313

Goddefroyinella Distant, 1916d (type species: G. indicans Distant by monotypy, junior 

synonym of G. neglecta (Buckton)) [previously incertae sedis (McKamey 1998a)]. 

Lubra Goding, 1903a (type species: Oxyrhachis spinicornis Walker by subsequent 


Matumuia Day, 1999a (type species: M. laura Day by original designation), see figs. 24, 36, 

123-125, 213 of Day (1999a: 633, 684, 736) [previously incertae sedis (Day 1999a)]. †* 

Neocanthuchus Day, 1999a (type species: N. tropicus Day by original designation) 

[previously incertae sedis (Day 1999a)].* 

Neosextius Day, 1999a (type species: N. longinotum Day by original designation), see figs. 

26, 30, 43, 132-137, 215 of Day (1999a: 633, 634, 691-692, 736) [previously incertae 

sedis (Day 1999a)].† 

Otinotoides Distant, 1916c (type species: Centrotus pallipes Walker by original designation) 

[previously placed in Centrotypini (McKamey 1998a)].* 

Pogonella Evans, 1966a (type species: Centrotypus minutus Goding by original designation) 


Pogonotypellus Evans, 1966a (type species: Pogontypus australis Goding by original 


Polonius Distant, 1916e (type species: P. biseratensis Distant by monotypy) [previously 

placed in Leptocentrini (McKamey 1998a)].* 

Protinotus Day, 1999a (type species: Otinotus doddi Distant by original designation), see 

figs. 12, 150-152, 219 of Day (1999a: 632, 703, 737) [previously incertae sedis (Day 

1999a)].†* 

314

Pyrgonota Stål, 1870c (type species: Centrotus bifoliatus Westwood by subsequent 

designation) [previously placed in Hypsaucheniini (Yuan and Chou 2002a)]. 

Rentzia Day, 1999a (type species: R. yarla Day by original designation), see figs. 10, 26, 32, 

42, 153-158, 220 of Day (1999a: 632-634, 704, 707, 737) [previously incertae sedis (Day 

1999a)].†* 

Rigula Day, 1999a (type species: R. calperum Day by original designation), see figs. 18, 34, 

40, 159-164, 221 of Day (1999a: 633-634, 708, 710-711, 737) [previously incertae sedis 

(Day 1999a)].†* 

Sarantus Stål, 1863c (type species: Sarantus wallacei Stål by monotypy) [previously placed 

in Leptocentrini (McKamey 1998a)]. 

Sertorius Stål, 1866a (type species: Centrotus australis Fairmaire by subsequent designation) 


Sextius Stål, 1866c (type species: Centrotus virescens Fairmaire by subsequent designation) 


Strzeleckia Day, 1999a (type species: S. montanus Day by original designation), see figs. 26, 

30, 41, 174-176, 224 of Day (1999a: 633-634, 719, 738) [previously incertae sedis (Day 

1999a)].†* 

Terentius Stål, 1866a (type species: T. convexus Stål by subsequent designation). 

Undarella Day, 1999a (type species: U. storeyi Day by original designation) [previously 

incertae sedis (Day 1999a)].* 

Yangupia Day, 1999a (type species: Centrotypus occidentalis Goding by original 

designation) [previously incertae sedis (Day 1999a)].* 

315

Specimens examined.—Acanthucalis macalpini Evans as det. in ANIC, #00-136i&; 

Acanthuchus trispinifer (Fairmaire), as det. in USNM, #00-80g&, #00-80h%—as det. in 

ANIC, #00-181o%, #01-43a&; Alocanthella fulva Evans, as det. in ANIC, #01-225b%; 

Alosextius carinatus (Funkhouser), as det. in AMSA, #00-136a&; Anzac bipunctatum 

(Fabricius), as det. in ANIC, #00-136d&, #00-136e%; Arimanes doryensis Distant, as det. in 

USNM, #83-334a&; Bucktoniella pyramidatus (Funkhouser), as det. in CNCI, #01-39a%; 

Bulbauchenia sp. (probably mirablis), det. M.S. Wallace, USNM, #00-230g♂; B. bakeri 

(Funkhouser), [holotype of Emphusis bakeri Funkhouser], USNM—det. D. Flynn, NCSU, 

#02-67b(sex?) —det. Z.P. Metcalf, NCSU, #00-221m♀, #02-67a♀, #02-67f♀ —as det. in 

NCSU, #01-277a♂, #01-278a♀, #02-67c♀, #02-67d♂, #02-67e(sex?), #02-67g♀, #02- 

67h♀, #02-67i♂, #02-67j(sex?)—as det. in USNM, #01-232c%; B. globosa (Funkhouser), 

[holotype of Emphusis globosus Funkhouser], USNM; B. mirabilis (Funkhouser), [holotype 

of Clonauchenia mirablis Funkhouser], USNM, #01-54a♂; B. rugosa (Funkhouser), 

[holotype of Emphusis rugosus Funkhouser], USNM; Cebes transiens (Walker), as det. in 

ANIC, #00-80c%, #00-80d&; Ceraon tasmaniae (Fairmaire), det. W.D. Funkhouser, USNM, 

#00-80a&—as det. in USNM, #00-80b%, #01-232e%; Crito festivum Distant, holotype [as 

Crito festivus Distant], BMNH; Dingkana borealis Goding, det. W.D. Funkhouser, USNM, 

#00-87d&, #00-87e%—as det. in USNM, #01-219c&; Eufairmairia decisa (Walker), as det. 

in USNM, #83-333c%, #83-333d&; E. fraterna Distant, as det. in ANIC, #00-87k&—as det. 

in USNM, #01-22b&; Eufairmairiella sp, as det. in ANIC, #00-87j%; E. curvicauda 

(Goding), as det. in USNM, #00-87g%; Eufrenchia falcata (Walker), as det. in ANIC, #00- 

87h%, #00-87i&; Eutryonia monstrifera (Walker), as det. in USNM, #00-87a%, #00-87b&, 

316

#01-219b&; Evansiana iasis (Kirkaldy), as det. in ANIC, #01-225a&; Funkhouserella 

arborea (Funkhouser), [holotype of Pyrgonota arborea Funkhouser], USNM; F. binodis 

(Funkhouser), [holotype of Pyrgonota binodis Funkhouser], USNM; F. brevifurca 

(Funkhouser), [holotype of Pyrgonota brevifurca Funkhouser], USNM, #01-089e& —as det. 

in USNM, #00-228c&; F. bulbiturris (Funkhouser), [holotype of Pyrgonota bulbiturris 

Funkhouser], USNM; F. pinguiturris (Funkhouser), [holotype of Pyrgonota pinguiturris 

Funkhouser], USNM, #01-89f&; F. sinuata (Funkhouser), [holotype of Pyrgonota sinuata 

Funkhouser], USNM; Goddefroyinella neglecta (Buckton), paralectotype, BMNH, #01-69e& 

—as det. in ANIC, #01-3b%; Lubra spinicornis (Walker), as det. in ANIC, #00-181l&—as 

det. in AMSA, #00-122f%—as det. in USNM, #00-122e&—det. M.S. Wallace, USNM, #00- 

122j%; Neocanthuchus tropicus Day, paratype, #01-225c&; Otinotoides sp., as det. in ANIC, 

#00-122k&, #00-122l%; O. pallipes (Walker), det. W.D. Funkhouser, AMSA, #00-122g%, 

#00-122h&; Pogonella minutus (Goding), as det. in ANIC, #00-136f&, #00-136j%—as det. in 

USNM, #00-136g%; Pogonotypellus australis (Goding), as det. in ANIC, #00-136h&; 

Polonius froggatti Goding, holotype, USNM, #01-89j&; Pyrgonota sp., USNM, #01- 

47b(sex?)—det. M.S. Wallace, USNM, #01-47 (sex?); P. bifoliata (Westwood), as det. in 

USNM, #00-230a&, #00-230b%—det. D. Flynn, DJFC, #00-230j&; Sarantus nobilus 

Kirkaldy, as det. in ANIC, #00-136b&, #00-136c&; S. wallacei Stål, as det. W.D. 

Funkhouser, USNM, #00-122c&—as det. in USNM, #00-122d%, #01-247b&; Sertorius sp., 

as det. in ANIC, #00-122m%; S. australis (Fairmaire), as det. in USNM, #00-122a&—det. 

F.W. Goding, USNM, #00-122b%; Sextius kurandae Kirkaldy, as det. in USNM, #01-235b%; 

S. virescens (Fairmaire), det. W.D. Funkhouser, USNM, #00-80e%—as det. in USNM, #00- 

80f&—as det. in ANIC, #00-80i%; Terentius convexus Stål, as det. in USNM, #99-100h&, 

317

#99-100i%, #01-29a%—as det. in ANIC, 00-12a&, 00-12b%; Undarella storeyi Day, 

holotype, QMBA—paratype, QMBA, #01-250a&; Yangupia occidentalis (Goding), as det. in 

BMNH, #01-296a%, #01-296b%. 

318

Fig. 22.1. Terentiini: pronota (lateral aspects). Bars = 3 mm. A, Acanthucalis macalpini Evans, 

#00-136i&, reversed from right lateral aspect. B, Acanthuchus trispinifer (Fairmaire), #00-181o%. 

C, Alocanthella fulva Evans, #01-225b%. D, Alosextius carinatus (Funkhouser), #00-136a&. E, 

Anzac bipunctatum (Fabricius), #00-136e%. F, Arimanes doryensis Distant, #83-334a&. G, 

Bucktoniella pyramidatus (Funkhouser), #01-39a%. H, Bulbauchenia sp., #00-230g%. I, B. bakeri 

(Funkhouser), #00-221m&. J, Cebes transiens (Walker), #00-80d&. K, Ceraon tasmaniae 

(Fairmaire), #00-80a&. L, Dingkana borealis Goding, #00-87d&. M, Eufairmairia fraterna 

Distant, #00-87k&. N, Eufairmairiella sp., #00-87j%. O, Eufrenchia falcata (Walker), #00-87i&. 

319

Fig. 22.2. Terentiini: pronota (lateral aspects). Bars = 3 mm. A, Eutryonia monstrifera (Walker), #00- 

87a%. B, Evansiana iasis (Kirkaldy), #01-225a&. C, Funkhouserella brevifurca (Funkhouser), #00- 

228c&. D, F. pinguiturris (Funkhouser), holotype, #01-89f&. E, Goddefroyinella neglecta (Buckton), 

#01-3b%. F, Lubra spinicornis (Walker), #01-122e&. G, Neocanthuchus tropicus Day, paratype, #01- 

225c&. H, Otinotoides pallipes (Walker), #00-122h&. I, Pogonella minutus (Goding), #00-136f&. J, 

Pogonotypellus australis (Goding), #00-136h&. K, Polonius froggatti Goding, holotype, #01-89j&. L, 

Pyrgonota bifoliata (Westwood), #00-230j&. M, Sarantus nobilus Kirkaldy, #00-136c&. N, S. 

wallacei Stål, #00-122c&. O, Sertorius australis (Fairmaire), #00-122a&. 

320

Fig. 22.3. Terentiini: pronota (lateral aspects A-D, and anterior aspects, E-O). Bars = 3 mm. A, 

Sextius virescens (Fairmaire), #00-80f&. B, Terentius convexus Stål, #00-12b%. C, Undarella storeyi 

Day, holotype. D, Yangupia occidentalis (Goding), #01-296a%. E, Acanthucalis macalpini Evans, 

#00-136i&. F, Acanthuchus trispinifer (Fairmaire), #00-181o%. G, Alocanthella fulva Evans, #01- 

225b%. H, Alosextius carinatus (Funkhouser), #00-136a&. I, Anzac bipunctatum (Fabricius), #00- 

136e%. J, Arimanes doryensis Distant, #83-334a&. K, Bucktoniella pyramidatus (Funkhouser), #01- 

39a%. L, Bulbauchenia sp., #00-230g%. M, B. bakeri (Funkhouser), #00-221m&. N, Cebes transiens 

(Walker), #00-80d&. O, Ceraon tasmaniae (Fairmaire), #00-80a&. Copyrights: D © 2003, The Natural 

History Museum, London. 

321

Fig. 22.4. Terentiini: pronota (anterior aspects). A, Dingkana borealis Goding, #00- 

87d&. B, Eufairmairia fraterna Distant, #00-87k&. C, Eufairmairiella sp., #00-87j%. 

D, Eufrenchia falcata (Walker), #00-87i&. E, Eutryonia monstrifera (Walker), #00- 

87a%. F, Evansiana iasis (Kirkaldy), #01-225a&. G, Funkhouserella brevifurca 

(Funkhouser), #00-228c&. H, F. pinguiturris (Funkhouser), holotype, #01-89f&. I, 

Goddefroyinella neglecta (Buckton), #01-3b%. J, Lubra spinicornis (Walker), #01- 

122e&. K, Neocanthuchus tropicus Day, paratype, #01-225c&. L, Otinotoides pallipes 

(Walker), #00-122h&. M, Pogonella minutus (Goding), #00-136f&. N, Pogonotypellus 

australis (Goding), #00-136h&. O, Polonius froggatti Goding, holotype, #01-89j&. 

322

Fig. 22.5. Terentiini: pronota (anterior aspects, A-H) and heads (I-O). A, Pyrgonota bifoliata 

(Westwood), #00-230j&. B, Sarantus nobilus Kirkaldy, #00-136c&. C, S. wallacei Stål, #00-122c&. 

D, Sertorius australis (Fairmaire), #00-122a&. E, Sextius virescens (Fairmaire), #00-80f&. F, 

Terentius convexus Stål, #99-100h&. G, Undarella storeyi Day, holotype. H, Yangupia occidentalis 

(Goding), #01-296a%. I, Acanthuchus trispinifer (Fairmaire), #00-181o%. J, Alocanthella fulva Evans, 

#01-225b%. K, Alosextius carinatus (Funkhouser), #00-136a&. L, Anzac bipunctatum (Fabricius), #00- 

136d&. M, Arimanes doryensis Distant, #83-334a&. N, Bucktoniella pyramidatus (Funkhouser), #01- 

39a%. O, Bulbauchenia sp., #00-230g%. Copyrights: H © 2003, The Natural History Museum, 

London. fcl, frontoclypeal lobes. 

323

Fig. 22.6. Terentiini: heads. A, Cebes transiens (Walker), #00-80c%. B, Ceraon tasmaniae 

(Fairmaire), #00-80a&. C, Dingkana borealis Goding, #00-87d&. D, Eufairmairia fraterna 

Distant, #00-87k&. E, Eufairmairiella sp., #00-87j%. F, Eufrenchia falcata (Walker), #00-87i&. 

G, Eutryonia monstrifera (Walker), #00-87a%. H, Evansiana iasis (Kirkaldy), #01-225a&. I, 

Funkhouserella brevifurca (Funkhouser), #00-228c&. J, F. pinguiturris (Funkhouser), holotype, 

#01-89f&. K, Goddefroyinella neglecta (Buckton), #01-3b%. L, Lubra spinicornis (Walker), #01- 

122e&. M, Neocanthuchus tropicus Day, paratype, #01-225c&. N, Otinotoides pallipes (Walker), 

#00-122h&. O, Pogonella minutus (Goding), #00-136f&. fcl, frontoclypeal lobes. 

324

Fig. 22.7. Terentiini: heads. A, Pogonotypellus australis (Goding), #00- 

136h&. B, Polonius froggatti Goding, holotype, #01-89j&. C, 

Pyrgonota bifoliata (Westwood), #00-230j&. D, Sarantus nobilus 

Kirkaldy, #00-136c&. E, S. wallacei Stål, #00-122c&. F, Sertorius 

australis (Fairmaire), #00-122a&. G, Sextius virescens (Fairmaire), #00- 

80f&. H, Terentius convexus Stål, #99-100h&. I, Undarella storeyi Day, 

holotype. fcl, frontoclypeal lobes. 

325

Fig. 22.8. Terentiini: wings. A, Terentius convexus Stål, #99-100h&, 

left forewing (inverted). B, T. convexus, #99-100h&, left hind wing 

(inverted). C, Bulbauchenia sp., #00-230g%. D, Acanthucalis macalpini 

Evans, #00-136i&, left forewing (inverted). E, Acanthuchus trispinifer 

(Fairmaire), #00-80g&. 

326

Fig. 22.9. Terentiini: wings. A, Alocanthella fulva Evans, #01-225b%. B, 

Alosextius carinatus (Funkhouser), #00-136a&. C, Anzac bipunctatum 

(Fabricius), #00-136d&. D, Arimanes doryensis Distant, #83-334a&. E, 

Bucktoniella pyramidatus (Funkhouser), #01-39a%, left forewing (inverted). 

F, Bulbauchenia bakeri (Funkhouser), #00-221m&. G, Cebes transiens 

(Walker), #00-80dc%. H, Ceraon tasmaniae (Fairmaire), #00-80a&, left 

forewing (inverted). I, Dingkana borealis Goding, #00-87d&. J, Eufairmairia 

decisa (Walker), #83-333d&, left forewing (inverted). 

327

Fig. 22.10. Terentiini: wings. A, Eufairmairiella sp., #00-87g%. B, Eufrenchia 

falcata (Walker), #00-87h%. C, Eutryonia monstrifera (Walker), #00-87b&. D, 

Evansiana iasis (Kirkaldy), #01-225a&, left forewing (inverted). E, 

Funkhouserella brevifurca (Funkhouser), #00-228c&. F, F. pinguiturris 

(Funkhouser), holotype, #01-89f&. G, Goddefroyinella neglecta (Buckton), 

paralectotype, #01-69e&, left forewing (inverted). H, Lubra spinicornis (Walker), 

#01-122e&, left forewing (inverted). I, Neocanthuchus tropicus Day, paratype, 

#01-225c&. J, Otinotoides pallipes (Walker), #00-122h&. 

328

Fig. 22.11. Terentiini: wings. A, Pogonella minutus (Goding), #00-136f&. B, 

Pogonotypellus australis (Goding), #00-136h&. C, Polonius froggatti Goding, 

holotype, #01-89j&, left forewing (inverted). D, Pyrgonota bifoliata 

(Westwood), #00-230a&. E, S. wallacei Stål, #00-122c&. F, Sertorius australis 

(Fairmaire), #00-122a&. G, Sextius virescens (Fairmaire), #00-80f&. H, 

Undarella storeyi Day, holotype, left forewing (inverted). I, Yangupia 

occidentalis (Goding), #01-296a%. 

329

Fig. 22.12. Terentiini: metathoracic legs. A, Bulbauchenia bakeri 

(Funkhouser), #00-221m&. B, Pyrgonota bifoliata (Westwood), 

#00-230a&. C, Terentius convexus Stål, #99-100h&, reversed from 

right lateral aspect. 

330

Fig. 22.13. Terentiini: female second valvulae (lateral aspects and 

closeup of apex). A, Acanthucalis macalpini Evans, #00-136i&. B, 

Anzac bipunctatum (Fabricius), #00-136d&. C-D, Acanthuchus 

trispinifer (Fairmaire), #00-80g&. E-F, Alosextius carinatus 

(Funkhouser), #00-136a&. G-H, Bulbauchenia bakeri (Funkhouser), 

#00-221m&. I-J, Cebes transiens (Walker), #00-80d&. 

331


closeup of apex). A-B, Ceraon tasmaniae (Fairmaire), #00-80a&. C- 

D, Dingkana borealis Goding, #00-87d&. E, Eufairmairia decisa 

(Walker), #83-333d&. F, Pogonella minutus (Goding), #00-136f&. G- 

H, Eufrenchia falcata (Walker), #00-87i&. I-J, Eutryonia monstrifera 

(Walker), #00-87b&. 

332


closeup of apex). A-B, Funkhouserella brevifurca (Funkhouser), #00- 

228c&. C, F. pinguiturris (Funkhouser), holotype, #01-89f&. D-E, 

Goddefroyinella neglecta (Buckton), paralectotype, #01-69e&. F-G, 

Lubra spinicornis (Walker), #00-122e&. H-I, Otinotoides pallipes 

(Walker), #00-122h&. Copyrights: D-E © 2003, The Natural History 

Museum, London. 

333


closeup of apex). A-B, Pogonotypellus australis (Goding), #00-136h&. 

C-D, Pyrgonota bifoliata (Westwood), #00-230a&. E-F, Sarantus 

nobilus Kirkaldy, #00-136b&. G-H, S. wallacei Stål, #00-122c&. I-J, 

Sertorius australis (Fairmaire), #00-122a&. 

334

Fig. 22.17. Terentiini: female second valvulae (lateral aspects and closeup 

of apex, A-D) and male styles (lateral aspects, E-P). A-B, Sextius virescens 

(Fairmaire), #00-80f&. C-D, Terentius convexus Stål, #99-100h&. E, 

Acanthuchus trispinifer (Fairmaire), #00-80h%. F, Alocanthella fulva 

Evans, #01-225b%. G, Anzac bipunctatum (Fabricius), #00-136e%. H, 

Bucktoniella pyramidatus (Funkhouser), #01-39a%. I, Bulbauchenia sp., 

#00-230g%. J, B. bakeri (Funkhouser), #00-277a%. K, Cebes transiens 

(Walker), #00-80c%. L, Ceraon tasmaniae (Fairmaire), #00-80b%. M, 

Dingkana borealis Goding, #00-87e%. N, Eufairmairia decisa (Walker), 

#83-333c%. O, Eufairmairiella curvicauda (Goding), #00-87g%. P, 

Eufrenchia falcata (Walker), #00-87h%. c, clasp. 

335

Fig. 22.18. Terentiini: male styles (lateral aspects, A-K) and aedeagi (lateral aspects, L-Q). A, 

Eutryonia monstrifera (Walker), #00-87a%. B, Goddefroyinella neglecta (Buckton), #01-3b%. 

C, Lubra spinicornis (Walker), #01-122j%. D, Otinotoides pallipes (Walker), #00-122g%. E, 

Pogonella minutus (Goding), #00-136g%. F, Pyrgonota bifoliata (Westwood), #00-230b%. G, 

Sarantus wallacei Stål, #00-122d%. H, Sertorius australis (Fairmaire), #00-122b%. I, Sextius 

virescens (Fairmaire), #00-80e%. J, Terentius convexus Stål, #99-100i%. K, Yangupia 

occidentalis (Goding), #01-296a%. L, Alocanthella fulva Evans, #01-225b%. M, Bucktoniella 

pyramidatus (Funkhouser), #01-39a%. N, Bulbauchenia bakeri (Funkhouser), #00-277a%. O, 

Eufairmairiella curvicauda (Goding), #00-87g%. P, Eutryonia monstrifera, #00-87a%. Q, L. 

spinicornis, #01-122j%. Copyrights: K © 2003, The Natural History Museum, London. c, 

clasp. 

336

Fig. 22.19. Terentiini: male aedeagi (lateral aspects, A-F) and lateral 

plates (lateral aspects, G-L). A, Otinotoides pallipes (Walker), #00- 

122g%. B, Pyrgonota bifoliata (Westwood), #00-230b%, reversed from 

right lateral aspect. C, Sarantus wallacei Stål, #00-122d%. D, Sextius 

virescens (Fairmaire), #00-80e%. E, Terentius convexus Stål, #99- 

100i%. F, Yangupia occidentalis (Goding), #01-296a%. G, 

Bucktoniella pyramidatus (Funkhouser), #01-39a%. H, Bulbauchenia 

sp., #00-230g%. I, Bulbauchenia bakeri (Funkhouser), #00-277a%. J, 

Cebes transiens (Walker), #00-80c%. K, Ceraon tasmaniae 

(Fairmaire), #00-80b%. L, Dingkana borealis Goding, #00-87e%. 

Copyrights: F © 2003, The Natural History Museum, London. dl, 

dorsoapical lobe. 

337

Fig. 22.20. Terentiini: male lateral plates (lateral aspects). A, 

Eufairmairia decisa (Walker), #83-333c%. B, Eufairmairiella 

curvicauda (Goding), #00-87g%. C, Eufrenchia falcata (Walker), 

#00-87h%. D, Eutryonia monstrifera (Walker), #00-87a%. E, 

Goddefroyinella neglecta (Buckton), #01-3b%. F, Lubra spinicornis 

(Walker), #01-122j%. G, Otinotoides pallipes (Walker), #00-122g%. 

H, Sarantus wallacei Stål, #00-122d%. I, Sertorius australis 

(Fairmaire), #00-122b%. J, Sextius virescens (Fairmaire), #00-80e%. 

K, Terentius convexus Stål, #99-100i%. L, Yangupia occidentalis 

(Goding), #01-296a%. Copyrights: L © 2003, The Natural History 

Museum, London. dl, dorsoapical lobe. 

338

Fig. 22.21. Terentiini: male subgenital plates (ventral aspects, A- 

D) and maximum development of abdominal fine-structure (E-H). 

All scanning electron micrographs near tergum III. A, 

Bucktoniella pyramidatus (Funkhouser), #01-39a%. B, Cebes 

transiens (Walker), #00-80c%. C, Eufrenchia falcata (Walker), 

#00-87h%. D, Otinotoides pallipes (Walker), #00-122g%. E-F, 

Acanthuchus trispinifer (Fairmaire), #01-43&. G-H, Anzac 

bipunctatum (Fabricius), #00-136e%. a, acanthae. i, inornate pit. 

l, lateral seta. m, microtrichia. 

339

Fig. 22.22. Terentiini: maximum development of abdominal finestructure. 


Bucktoniella pyramidatus (Funkhouser), #01-39a%. B, Bulbauchenia 

mirabilis (Funkhouser), holotype, #01-54a%. C-D, B. bakeri 

(Funkhouser), #01-278a&. E, Ceraon tasmaniae (Fairmaire), #01- 

232e%. F, Dingkana borealis Goding, #01-219c&. G-H, Eufairmairia 

fraterna Distant, #01-22b&. i, inornate pit. m, microtrichia. 

340

Fig. 22.23. Terentiini: maximum development of abdominal and 

pronotal (B) fine-structure. All scanning electron micrographs near 

tergum III. A, Eutryonia monstrifera (Walker), #01-219b&. B-D, 

Funkhouserella brevifurca (Funkhouser), #01-89e&. E, Lubra 

spinicornis (Walker), 00-181l&. F, Otinotoides sp., 00-122k&. G, 

Pogonella minutus (Goding), #00-136j%. H, Pyrgonota sp., #01- 

47b(sex?). i, inornate pit. m, microtrichia. sc, scutellum. 

341

Fig. 22.24. Terentiini: maximum development of abdominal finestructure. 


Sarantus wallacei Stål, #01-247b&. B, Sertorius sp., #00-122m%. 

C, Sextius kurandae Kirkaldy, #01-235b%. D, Terentius convexus 

Stål, #01-29a%. i, inornate pit. m, microtrichia. 

342

Xiphopoeini Capener, 1966a [new tribe]. 

23. Tribe XIPHOPOEINI Capener, 1966 

Old World: Afrotropical Region 

Figs. 23.1-23.3 

Type genus: Xiphopoeus Stål, 1866c 

Diagnostic characters.—Frontoclypeal lobes distinct, not extending to apex of 

frontoclypeus; vertex with toothlike projections. Pronotum with numerous acute projections 

(spines). Mesopleural lobe enlarged. Forewing with m-cu2 crossvein present, discoidal cells 

similar in length, r-m1 crossvein originating anterior to first division of R vein, with 

pterostigma at or near R1 vein, base of R2+3 and R4+5 veins truncate. Hind wing with R4+5 and 

M1+2 veins not fused (4 apical cells). Mesothoracic femur with ab- and adlateral cucullate 

setae. Male lateral plate without dorsoapical lobe; clasp of style oriented laterally, apex 

membranous, elliptical or circular, angled dorsally. Abdominal tergum III ventrolateral 

margin with upcurved groove, abdominal setal bases enlarged, numerous, and dispersed on 

terga. 

Description.—Length 4.7-6.3 mm. Color brown to dark brown. HEAD (Figs. 23.1 

E-F): frontoclypeal margins parallel or slightly converging ventrally, frontoclypeal lobes 

distinct and not extending to apex of frontoclypeus; ocelli about equidistant from each other 

and eyes; vertex with toothlike projections. THORAX: PRONOTUM (Figs. 23.1 A-D): 

pronotum with numerous acute projections (spines); suprahumeral horns present; posterior 

process curving dorsally (Xiphopoeus) or straight at base (Negus), appressed against 

343

scutellum (Negus) or not (Xiphopoeus), significantly extending past m-cu3 vein in forewing. 

SCUTELLUM: emarginate with apices acute, not concealed by posterior process, 2 lateral 

apices or 1 visible from dorsolateral view; not shortened--with abdomen removed, notch and 

apices visible, only slightly extending beyond thorax or posterior half extending past thorax. 

Pleuron: propleural lobe present or absent, mesopleural lobe enlarged. FOREWING (Figs. 

23.1 G, I): sub-opaque; apical limbus broad; s crossvein distad of r-m2 crossvein; m-cu1 

crossvein absent and m-cu2 crossvein present in at least one wing; M and Cu veins fused at 

base; R and M veins not confluent preapically; forewing with pterostigma near R1 vein; R1 

vein not perpendicular to marginal vein; r-m1 crossvein originating anterior to first split of R 

vein, parallel to longitudinal veins or bent towards R vein; R, M, and Cu veins not parallel 

apically; discoidal cells similar in length; base of R2+3 and R4+5 veins truncate. HIND WING 

(Fig. 23.1 H): R4+5 and M1+2 veins not fused (4 apical cells). PRO- AND MESOTHORACIC LEGS: 

tibiae not foliaceous; mesothoracic tibia without rows of cucullate; mesothoracic femur with 

ab- and adlateral cucullate setae. METATHORACIC LEG (Fig. 23.1 J): ventral margin of coxa, 

trochanter, and femur without enlarged setal bases; femur with ab- and adlateral cucullate 

setae; femur without ablateral cucullate setae ventrolaterally; tibia not foliaceous, row I with 

7-18 cucullate setae, row II with 19-33 cucullate setae in irregular or double row, row III 

with 13-23 cucullate setae; tarsomere I with 2 or more cucullate setae (usually 2). 

ABDOMEN: in anterior aspect (abdomen removed) nearly triangular; anterior tergal borders 

not modified; sternal transverse carina present in Negus; paired dorsal swellings absent; 

tergum III ventrolateral margin with upcurved groove, abdominal setal bases enlarged, 

numerous, and dispersed on terga. FEMALE GENITALIA (Figs. 23.2 A-D): second valvulae 

broadened slightly, tapering unevenly to apex, curved or not, dorsal teeth fine, acute 

344

projections on dorsal margin present or absent; third valvulae without ventral projections. 

MALE GENITALIA (Figs. 23.2 E-K): lateral plate without dorsoapical lobe (Figs. 23.2 I-K), 

without ventral lobe; subgenital plate without distinct division; style clasp (Figs. 23.2 E-F) 

oriented laterally, membranous, elliptical or circular, angled dorsally; style shank without 

significant arch. ABDOMINAL FINE STRUCTURE (Figs. 23.3 B): acanthae indistinct, bases not 

heightened, acanthae divided into threadlike microtrichia. 


Distribution.—The tribe Xiphopoeini is recorded from numerous Afrotropical 

countries (McKamey 1998a). 

Ecology.—Members of the tribe Xiphopoeini are reported from the host plant 

families Euphorbiaceae, Gramineae, and Leguminosae (Table 26.2). 

Discussion.—Capener (1966a) placed those African treehoppers with toothlike 

projections on the lower margins of the vertex and enlarged mesopleural lobes in the tribe 

Xiphopoeini. This tribe is monophyletic in the phylogenetic analysis (Figs. 24.1, 24.7) and is 

characterized by Capener’s diagnostic features as well as the presence of numerous enlarged 

setal bases dispersed on the abdominal terga. The Xiphopeoini are closely related to the 

predominantly Afrotropical tribes Leptocentrini and Centrotini, and the Micreunini. The 

Centrotini, like the Xiphopoeini, also have enlarged setal bases dispersed on the abdominal 

terga although they are less numerous. In addition, both xiphopoeines and centrotines have 

an elliptical, dorsally angled, male clasp. 

345

Genera of the tribe Xiphopoeini 

Negus Jacobi, 1910b (type species: N. asper Jacobi by original designation). 

Xiphopoeus Stål, 1866a (type species: Centrotus phantasma Signoret by subsequent 

designation). 

Specimens examined.—Negus asper Jacobi, det. A.L. Capener, USNM, #99-93c&— 

det. A.L. Capener, AMNH, #00-175a%, #00-175j[n]—det. A.L. Capener, PPRI, #01-256e%; 

Xiphopoeus sp., det. M.S. Wallace, USNM, #01-61a&, #02-10e[n]; X. erectus Distant, det. 

W.D. Funkhouser, USNM, #83-333h%; X phantasma Signoret, det. A.L. Capener, PPRI, 

#99-315c%, #99-315d&. 

346

Fig. 23.1. Xiphopoeini: pronota (lateral aspects, A-B; and anterior aspects, C-D), 

heads (E-F), wings (G-I), and metathoracic leg (J). Bars = 3 mm. A, Negus asper 

Jacobi, #99-93c&. B, Xiphopoeus phantasma Signoret, #99-315c%. C, N. asper, 

#99-93c&. D, X. phantasma, #99-315c%. E, N. asper, #99-93c&. F, X. phantasma, 

#99-315c%. G, X. phantasma, #99-315d&, right forewing. H, X. phantasma, #99- 

315d&, right hind wing. I, N. asper, #99-93c&, right forewing. J, X. phantasma, 

#99-315d&. tp, toothlike processes. 

347

Fig. 23.2. Xiphopoeini: female second valvulae (lateral aspects and 

closeup of apex, A-D), and male styles (lateral aspects, E-F), aedeagi 

(lateral aspects, G-H), lateral plates (lateral aspects, I-J), and subgenital 

plate (ventral aspect, K). A-B, Negus asper Jacobi, #99-93c&. C-D, 

Xiphopoeus phantasma Signoret, #99-315d&. E, N. asper, #00-175a%. 

F, X. phantasma, #99-315c%. G, N. asper, #00-175a%. H, X. 

phantasma, #99-315c%. I, N. asper, #00-175a%. J, X. phantasma, #99- 

315c%. K, N. asper, #00-175a%. c, clasp. 

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Fig. 23.3 Xiphopoeini: maximum development of pronotal 

(A) and abdominal fine structure (B). All abdominal 

scanning electron micrographs near tergum III. A-B, 

Xiphopoeus sp., #01-61a&. l, lateral seta. m, microtrichia. 

349

Introduction 

24: PHYLOGENETIC RELATIONSHIPS WITHIN THE SUBFAMILY 

CENTROTINAE 

The treehopper subfamily Centrotinae accounts for roughly half of the worldwide 

membracid diversity at the species and higher-levels. It is the only subfamily with genera 

distributed in the Old World (the Afrotropical, Palearctic, Indomalayan, and Australasian/ 

Oceanian Regions) and New World (Nearctic and Neotropical Regions), but no centrotine 

tribe (or genus) occurs in both the Old and New Worlds. Historically, workers have focused 

on taxonomic revisions of centrotines within a particular geographic region (for example, 

Capener 1962a, 1968a; Evans 1966a; Ananthasubramanian 1996a; Day 1999a; Yuan and 

Chou 2002a). Furthermore, relatively few workers have justified centrotine classifications at 

any taxonomic level with quantitative phylogenetic analyses (Strümpel 1972a; Ahmad 

1988a; Dietrich and Deitz 1993a; Cryan et al. 2000a; Dietrich et al. 2001a; Yuan and Chou 

2002a). Consequently, a large number of centrotine tribes, many described in the early 

1900’s, are based largely on symplesiomorphies, and thus, the monophyly of the subfamily 

and tribes has not been tested. 

These disparities have impeded the development of a stable higher classification and 

taxonomic studies of centrotines at the generic and species levels. In addition, the lack of 

information on the evolutionary relationships between the Old and New World centrotines 

has hindered investigations on biogeographic patterns within the Membracidae and in 

determining the geographic origins of the Membracidae and the Centrotinae. The lack of 

common taxa between the New World and Old World has long fueled a debate concerning 

the geographic origin of the Membracidae (Dietrich and Deitz 1993a, Wood 1993a, Dietrich 

350

et al. 2001a). The first comprehensive phylogenetic analysis of generic representatives from 

all 24 centrotine tribes (20 Old World, 4 New World), (sensu McKamey 1998a and Yuan and 

Chou 2002a) using morphological characters is presented here. 

The objectives of this study are to establish the phylogenetic limits of the Centrotinae 

and its included tribes and to determine the evolutionary relationships among these tribes in 

order to provide a comprehensive classification and to aid in the investigation of 

biogeographical patterns and life history traits. The great number of centrotine genera-- 

together with the limited morphological data for many--precluded a single comprehensive 

phylogenetic analysis. To circumvent this problem, 10 analyses were performed on subsets 

of taxa as follows: (1) an overall phylogenetic analysis based on genera representing overall 

tribal diversity within Centrotinae; (2-8) phylogenetic analyses based on genera representing 

larger tribes; and (9-10) phenetic analyses based on all scorable genera representing larger 

tribes. 

Methods 

Taxon sampling. Overall, 222 taxa representing 213 genera (Table 24.13) were coded 

for their morphological characters in the taxonomic database DELTA (DEscription Language 

for TAxonomy) version 1.03e (Dallwitz 1980a; Dallwitz et al. 1993a, 1999a). Ingroup taxa 

included 208 genera (178 centrotines and 4 centrodontines coded from specimens; 26 

centrotine genera coded from published descriptions and illustrations). Taxa in Table 24.13 

labeled with only a generic name are represented by more than one species. Knowledge of 8 

centrotine genera was insufficient for coding: Aspasiana, Centrobelus, Insitor, Insitoroides, 

Megalocentrus, Megaloschema, Saudaraba, and Sinocentrus (all but Saudaraba placed as 

351

incertae sedis). Outgroup taxa included the New World genera: Nicomia and Tolania 

(Nicomiinae), Microcentrus Stål (Stegaspidinae), and Centronodus Funkhouser and 

Paracentronodus Sakakibara (Centronodini). The only possible Old World outgroup is the 

genus Darthula Kirkaldy, in the family Aetalionidae (sister group to the Membracidae). 

Darthula, however, is not useful for polarizing because it is too distantly related to 

centrotines. Generic representatives from 23 of the 24 valid tribes were examined; only the 

tribe Choucentrini (Choucentrus) was coded entirely from the literature. In order to lessen 

over-generalization of character data, an effort was made to examine the type species of each 

available genus. The type species (at least one sex) was examined in 164 of the 178 

centrotine genera coded. 

Analyses. The number of taxa and characters included and the characters excluded in 

each analysis are shown in Table 24.2. In relatively few cases where only one sex of the type 

species was available for examination, or where significant morphological diversity occurred 

within a genus, another species was coded in addition to the type species and included in the 

dataset. With three exceptions, the type species in these instances was used in all of the 

phylogenetic analyses. In the Boocerini/Centrodontini analysis, Campylocentrus hamifer 

was used as the generic representative. In the Choucentrini/Leptocentrini/Maarbarini 

analysis, Otinotus bantuantus and Nilautama minutaspina were chosen as the generic 

representatives. 

Question marks (?) in the data matrix indicate missing data, inapplicable character 

states, and cases where character scoring was ambiguous. With some characters, 

polymorphism (sexual, interspecific, or intraspecific) was designated as a hypothesized 

intermediate character state. Characters listed in the “other” column of Table 24.2 were 

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excluded because they were highly homoplasious and not informative as tribal or generic 

characters for the analyzed taxa. In the analysis of the entire subfamily Centrotinae, six 

autapomorphic characters that helped define monotypic tribes or tribes represented by one 

genus were retained in the analysis. 

In the first analysis, generic representatives of 22 of the 24 valid tribes plus the New 

World subfamily Centrodontinae were included in an overall phylogenetic analysis of the 

subfamily Centrotinae. Where possible, at least two genera from each valid tribe were 

included in this analysis. The generic representatives and institutional acronyms used in the 

subfamily analysis are given in Table 24.5. 

All of the centrotine genera could not be included in this overall analysis due to the 

large number of taxa and limited morphological data for some genera (often only one sex 

was available). Therefore, separate phylogenetic analyses (2-8) were also completed for 

several large tribes and for the generic representatives of the 2 remaining tribes not included 

in the overall phylogenetic analysis of the Centrotinae. Thus, many of the remaining 

centrotine genera could be confidently placed into tribes. The phylogenetic trees from these 

analyses were intended to indicate the monophyly of the tribes and preliminary relationships 

among their genera. It should be noted, however, that many of the characters used in the 

analyses were selected to delineate tribes and infer relationships among them, rather than 

infer generic relationships. Therefore, the addition of further morphological characters that 

vary among genera would likely provide even better resolution of generic relationships. 

Analyses 2-3 (Table 24.2) investigated the relationships among several basal 

centrotine tribes, mostly from the New World. The closely related tribes Beaufortianini (new 

tribe), Nessorhinini, Pieltainellini (new tribe), and Platycentrini were studied in Analysis 2. 

353

All of these tribes except the Beaufortianini are found in the New World. The generic 

relationships of Boocerini and Centrodontini, two New World tribes, were examined in 

Analysis 3. 

The remaining phylogenetic analyses examined relationships of Old World 

centrotines: Analysis 4, included two primarily Afrotropical tribes, the Centrotini and 

Xiphopoeini; 5, the predominantly Palearctic and Indomalayan tribes Choucentrini, 

Leptocentrini, and Maarbarini (new tribe); 6, the Gargarini; 7 the Oxyrhachini and 

Hypsaucheniini; and 8, Terentiini (Table 24.2). 

DELTA datasets were converted to NEXUS files in DELTA. Phylogenetic analyses 

(analyses 1-8) were performed using PAUP* (Phylogenetic Analysis Using Parsimony) 

version 4.0b10 for Windows (Swofford 2002a). Character change lists and apomorphy lists 

were generated using PAUP*. Due to the size of all the analyses, heuristic analyses were 

performed using the tree-bisection-reconnection routine (TBR) with 50 random addition 

replicates. Five trees were held at each step of cladogram construction. The number of 

changes assigned per branch under ACCTRAN optimization was determined by PAUP*. 

Branch lengths are proportional to the number of changes per branch and are used here as 

measure of node support. 

Two similarity analyses (Analyses 9-10) of several larger tribes using UPGMA were 

performed to show overall morphological similarity among genera in a tribe, including those 

in which data were too limited for inclusion in the phylogenetic studies. UPGMA was 

performed in PAUP*. The distance measure or branch lengths for the UPGMA trees are 

proportional to the mean character distance. 

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Morphological Characters. The dataset for the phylogenetic and phenetic analyses 

consists of 116 morphological characters (82 binary and 34 multistate) (Table 24.1). Only 

adult males and females were coded because too few nymphal specimens were examined to 

include characters from the immatures. Many of the characters, especially those of the 

forewings, are based on those in Dietrich et al. (2001a). For all analyses, characters were 

treated as unordered because polarizations were ambiguous. Characters were assigned equal 

weight in all analyses. See the “Introduction” section for a discussion of morphological 

characters. 

Results and Discussion 

Phylogeny of the Centrotinae and character evolution. The results presented here 

represent the first comprehensive morphological phylogenetic analysis of the subfamily 

Centrotinae. The overall phylogenetic analysis (Analysis 1) resulted in a single most 

parsimonious tree (Fig. 24.1) of 665 steps, consistency index (CI) of 0.23, and retention 

index (RI) of 0.60. Table 24.3 gives descriptive tree statistics of the other phylogenetic 

analyses (Analyses 2-8). In many cases, nodes defining tribes and relationships among tribes 

were well supported with numerous character changes-- for example: node 79 (Terentiini), 

nodes 82-83 (Ebhuloidesini, Oxyrhachini, Hypsaucheniini, and Terentiini), node 91 

(Choucentrini + Maarbarini), node 102 (Leptocentrini), and node 116 (Nessorhinini). 

Nevertheless, although only 1 most parsimonious tree was found, in basal portions of the tree 

branch support was either low (i.e., nodes 113 and 111) or nodes were supported by mostly 

homoplasious characters (i.e., node 118). Indeed, in general, homoplasy was high (CI=0.23) 

in this analysis, as one might expect with 69 taxa (Sanderson and Donoghue 1989a). Certain 

355

characters, however, such as the shape of the female second valvulae, were homoplasious in 

some tribes (i.e., Centrotini) but consistent and informative in others (i.e., Hypsaucheniini). 

In a separate analysis using the dataset of Analysis 1, all of the Old World genera 

were constrained for monophyly to determine the extra number of steps needed for a 

monophyletic Old World fauna. Analysis parameters were equal to the above unconstrained 

studies. Constraining a monophyletic Old World centrotine fauna resulted in 65 equally most 

parsimonious trees with 671 steps, 6 steps longer than the most parsimonious tree without the 

constraint. According to a winning-sites test between the most parsimonious tree 

(unconstrained for Old World monophyly) and the 65 equally parsimonious trees 

(constrained for Old World monophyly), the data did not provide significantly less support 

for the constrained analysis compared to the unconstrained analysis (p-values ranged from 

0.42-0.09). Despite this result, the unconstrained analysis is favored here not only because it 

is more parsimonious, but also because most of the Old World phylogeny was unresolved in 

the constrained study. 

As a result of these analyses, 23 centrotine tribes are recognized, including 6 new 

tribes; 11 tribal synonymies and 1 subfamily synonymy are proposed, and the tribal 

placements of 108 genera are changed. Based on the present phylogeny, Fig. 24.2 and Table 

24.14 contrast the existing centrotine tribal classification (sensu Ananthasubramanian 1996a, 

McKamey 1998a, and Yuan and Chou 2002a) with the revised classification. The tribal 

descriptions give detailed morphological descriptions and updated placements of each 

centrotine genus, while the tribal synonymies and discussions give details on the taxonomic 

status of each tribal name. 

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The Centrotinae, as defined here, are a monophyletic group based on the phylogenetic 

analysis (Fig. 24.1), supporting the findings of Dietrich and Deitz (1993a) and Dietrich et al. 

(2001a). All centrotines have abdominal inornate pits, each with an associated lateral seta. 

This feature is independently derived in three other treehopper genera: Nicomia 

(Nicomiinae), Endoiastus Fowler (Endoiastinae), and Eunusa Conseca (Membracini). 

Dietrich et al. (2001a) also listed these pits as a synapomorphy for centrotines. Moreover, 

with the exception of the centrodontines, centrotines have a truncate clavus while R, M, and 

Cu forewing veins lack “extra” branches. The remaining apomorphies of the Centrotinae 

(Table 24.4) are not as reliable due to the limited number of outgroups presented and the 

homoplastic nature of these characters. These include r-m1 crossvein bent towards R vein in 

the forewing, cucullate setae on dorsal femur absent, and abdominal shape in cross-section 

nearly triangular. 

Additional apomorphies of Centrotinae listed by Dietrich et al. (2001a) were not 

retrieved in the present analyses. Although the initial division of R vein in the forewing as 

R1 and Rs and the presence of r-m1 crossvein are diagnostic centrotine characters, they appear 

to be plesiomorphic based on the outgroups used here. Of these however, only Microcentrus 

has R1 and Rs as the initial division of R vein in the forewing like most centrotines--this 

character state is ambiguous for the remaining outgroup taxa. 

The Centrotinae topology presented by Dietrich et al. (2001a: Fig. 10) roughly 

resembles the tree presented here (Fig. 24.1). In both analyses, the Boocerini, as defined 

here, and Tricentrus, are basal to the remaining centrotines. With the exceptions of the 

placements of Oxyrhachini and Gargarini in Dietrich et al. (2001a), the two phylogenies are 

very similar. The Nessorhinini and Platycentrini form a monophyletic group in both analyses 

357

and are the sister group to the Leptocentrini and Centrotini, two closely related tribes in both 

trees. 

In contrast to morphological studies, a molecular analysis of the Membracidae using 

nuclear genes did not consistently result in a monophyletic Centrotinae (Cryan et al. 2000a). 

The Centrodontinae (here considered a tribe within Centrotini) and Stegaspidinae often 

grouped with the Centrotinae into one of two major membracid lineages. In a combined 

analysis with EF-1" and 28S rDNA, the New World centrotines arose from the Old World 

centrotines, contradictory to the results presented here. Moreover, many of the relevant 

nodes were not highly supported based on the molecular data (Cryan et al. 2000a). 

Suprahumeral horns are gained and lost numerous times in the Centrotinae. Although 

these pronotal horns are sometimes diagnostic at lower levels, they are rarely useful in 

defining tribes. This is not surprising considering the polymorphic nature of these horns in 

many species. Nevertheless, several centrotine genera have interesting modifications of the 

suprahumeral horns. For example, certain genera in the Centrotini (Euceropsila, 

Eumocentrulus, Eumonocentrus, Flatyperphyma, Foliatrotus, Mitranotus, Monocentrus, and 

Zanzia), Nessorhinini (Nessorhinus and Orekthophora), and Terentiini (Neosextius) have 

suprahumeral horns partially fused into a median anterior pronotal horn. These partially 

united horns appear to be an intermediate state between separate suprahumeral horns situated 

dorsolaterally on the pronotum (as in Centrotus) and a single median anterior pronotal horn 

(as in Micreune). Despite these modifications, the current phylogenetic analysis provides 

little to confirm this hypothesis. One would expect taxa with the intermediate state to be 

found basally on the phylogenetic tree with respect to taxa with a median anterior horn, but 

this apparently is never the case. This hypothesized intermediate state arose multiple times 

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in the Centrotinae and even within the tribe Centrotini (Fig. 24.7), but no genera of the 

Centrotini have the fully developed median anterior horn. The terentiine genera Eutyronia, 

Bulbauchenia, Funkhouserella, and Pyrgonota have a median anterior horn but they are not 

closely related to Neosextius (Fig. 24.14). In constrast, the presence of a median anterior 

horn is a synapomorphy of the tribes Hypsaucheniini, Micreunini, and Leptobelini, and there 

is no evidence of the intermediate state. 

An exposed scutellum has long helped to distinguish centrotines from other 

membracids, however, Dietrich et al. (2001a) found this condition to be plesiomorphic for 

the subfamily. The present study supports this finding and indicates that pronotal 

concealment of the scutellum was independently derived in the tribes Centrodontini, 

Oxyrhachini, and most Nessorhinini, and also in Insitor (incertae sedis), Monobeloides 

(Monobelini), Bulbauchenia and Neosextius (Terentiini), and several gargarine genera 

(Cryptaspidia, Gargarina, Madlinus, Mesocentrina). The condition is polymorphic in 

Orthobelus (Nessorhinini), Centrotypus (Centrotypini), and Sextius (Terentiini). 

Centrotinae tribal relationships. Several suites of characters proved especially 

significant in determining centrotine phylogeny. Characters important in elucidating tribal 

relationships include the characteristics, especially shape, of the male style clasp; shape of 

the female second valvulae; forewing and hind wing characteristics; features of the 

scutellum; leg chaetotaxy; and abdominal characteristics. 

The centrotines are arranged in two major clades plus the basal tribe Centrodontini 

(Fig. 24.1). The New World subfamily Centrodontini is the first lineage of the Centrotinae 

(Figs. 24.1, 24.5). Thus, Centrodontinae is here considered a junior synonym of the 

Centrotinae. Cryan et al.’s (2000a) recent molecular analyses using nuclear data consistently 

359

placed centrodontines near centrotine genera in the phylogenetic tree and recovered the 

Centrodontini clade consistently with 98-100% bootstrap support. Nevertheless, the 

Centrodontini are here tentatively placed within the Centrotinae. Although the 

centrodontines have inornate pits, each with an associated lateral seta (the synapomorphy for 

the Centrotinae), they differ significantly from other centrotines in forewing venation and leg 

chaetotaxy. Relationships within the Centrodontini are well resolved (Fig. 24.5). 

Apparently, Nodonica, the only centrodontine found in South America, is the sister group to 

the North American centrodontines, differing significantly from the latter in leg chaetotaxy 

and features of the male and female genitalia. Character 113 (abdominal inornate pits with 

lateral setae) was coded as ambiguous (?) for Nodonica because a specimen was not 

examined, and it is appropriate to score this character with a high power dissecting scope or 

scanning electron microscope. 

The next higher clade following the Centrodontini (Figs. 24.1-24.2) contains genera 

most recently placed in the Abelini, Antialcidini, Boocerini, Coccosterphini, Gargarini, 

Platycentrini, Nessorhinini, Leptocentrini, Madlinini, and Tricentrini (Ananthasubramanian 

1996a, McKamey 1998a, and Yuan and Chou 2002a). Based on the present results, the 

existing classification is clearly unacceptable, with numerous polyphyletic and paraphyletic 

tribes, forcing the placement of many genera into different tribes and the creation of new taxa 

and new synonymies. In most of the genera of this clade, the male subgenital plate has a 

distinct division near the base. 

The new tribe Monobelini contains genera previously placed in Boocerini and 

Nessorhinini (both sensu McKamey 1998a) (Figs. 24.1-24.2). The tribe Monobelini is the 

basal lineage of this first major centrotine clade and is one of two tribes confined to the 

360

Caribbean Islands. Monobelus and Monobeloides both have extra cucullate setae at the distal 

end of the femur and are more similar morphologically to each other than to 

Brachycentrotus. Brachycentrotus, the sister group to Monobelus and Monobeloides, lacks 

extra cucullate setae at the end of the hind femur. The Abelini Goding, 1930a, and Boocerini 

Goding, 1892a (sensu McKamey 1998a, in part), together form a monophyletic group (Figs. 

24.1-24.2, 24.5), rendering Abelini a junior synonym of Boocerini. The boocerines are 

defined by the long ventral lobe of the male lateral plate. A similar relationship, with Bremer 

support of 2, was recovered in the morphological analysis of Dietrich et al (2001a). 

However, phylogenetic analyses using nuclear data (Cryan et al. 2000a) did not result in a 

monophyletic Abelini and Boocerini sensu McKamey (1998a). Although generic 

relationships are well resolved, the strict consensus tree (Fig. 24.5) of the Boocerini indicates 

a basal polytomy. This lack of resolution may be due to missing data, because no males of 

Centriculus were available. Amblycentrus and Brachybelus, a monophyletic group, differ 

from other boocerines in hind wing venation, features of the male genitalia, and leg 

chaetotaxy. 

The monophyletic group of genera forming a sister group to the Boocerini (Fig. 24.1) 

includes a large number of tribes under the existing classification (Ananthasubramanian 

1996a, McKamey 1998a, and Yuan and Chou 2002a), but with the exceptions of the 

monotypic tribes Aleptocentrini and Madlinini, none of these are monophyletic (Figs 24.1- 

24.2). Therefore, based on the phylogenetic analyses (Figs. 24.1-24.2, 24.10), five tribes-- 

Antialcidini Yuan and Zhang, in Yuan and Chou, 2002a; Aleptocentrini Thirumalai and 

Ananthasubramanian, 1985a; Madlinini Boulard, 1995d; Tricentrini Ahmad and Yasmeen, 

1972a; and Coccosterphini Distant, 1908g--are all junior synonyms of Gargarini Distant, 

361

1908g. Molecular analyses of the Membracidae (Cryan et al. 2000a) consistently resulted in 

a monophyletic relationship between Gargara and Tricentrus with high bootstrap support. In 

contrast, Dietrich et al.’s (2001a) morphological analysis did not produce a monophyletic 

Gargara and Tricentrus. A UPGMA analysis including the remaining genera placed in the 

Gargarini (Fig. 24.16) also produced a single gargarine group based on overall similarity. 

Many of the generic relationships of the Gargarini are still unresolved (Fig. 24.10). 

Three morphologically distinct gargarines, however, Aleptocentrus, Yasa, and Parayasa, are 

consistently positioned at the base of the tree. The Gargarini, although morphologically 

heterogenous in some features, are united based on the expanding frontoclypeus, by the 

posterior process appressed against the scutellum, and the shortened scutellum (Figs. 24.1- 

24.2, 24.10). In the hind wing, R4+5 and M1+2 veins are fused (3 apical cells) in all genera 

except Aleptocentrus and Yasa. The Gargarini include the two largest centrotine genera, 

Tricentrus with 223 species, and Gargara with 184 (McKamey 1998a). Primarily distributed 

in the Indomalayan and Palearctic regions, the Gargarini are the descendants of one of the 

two major invasions of centrotines from the New World to the Old World. See the 

“Biogeography” chapter for a more detailed discussion of this dispersal. 

The next major clade accounts for most of the centrotine diversity at the tribal and 

generic levels. Similar to the other major clade, many of the tribes based on McKamey 

(1998a) and Yuan and Chou (2002a) are para- or polyphyletic resulting in new synonymies 

and the reassignment of genera. Furthermore, the recognition of several monotypic tribes is 

supported from the analysis in this major clade. The Centrocharesini, Oxyrhachini, 

Leptobelini, Ebhuloidesini, and Micreunini all have strong support based on convincing 

synapomorphies justifying their retainment as tribes (see the relevant tribal descriptions). 

362

As in its sister clade, the basal lineage of this second large assembly of tribes is a 

New World clade. The phylogenetic analyses agree with the recent placement of the 

Nessorhinini (formerly a subfamily) as a tribe within Centrotinae (Dietrich et al. 2001a). 

Furthermore, Callicentrus and Nessorhinus were monophyletic in the molecular phylogenetic 

analysis of the Membracidae using two nuclear genes (Cryan et al. 2000). Here, the 

nessorhinines (sensu McKamey 1998a) are polyphyletic (Fig. 24.1-24.2), with several genera 

formerly placed in the Platycentrini and Boocerini forming a monophyletic group with the 

type genus Nessorhinus and several other genera previously placed in the Nessorhinini (Fig. 

24.3). All of these genera, along with the Monobelini, are restricted to the Caribbean Islands. 

The internal phylogeny of the Nessorhinini is well resolved (Fig. 24.3) and is split 

into two groups. A morphologically homogenous group is represented by Nessorhinus and 

Goniolomus while a more heterogenous group is represented by Callicentrus and Orthobelus 

(Figs. 24.1, 24.3). These two groups differ in forewing characteristics and shape of the male 

and female genitalia. Despite the two clades, all nessorhines have anterior dorsal swellings 

on the abdomen and long blade-shaped female second valvulae with large dorsal teeth. A 

UPGMA analysis (Fig. 24.17) including the genus Spathenotus resulted in the same two 

groupings. The Platycentrini sensu McKamey (1998a) are also polyphyletic but Platycentrus 

and Tylocentrus (Fig. 24.3) form a monophyletic group supported by the shape of the female 

ovipositor, and are the sister group to the Nessorhinini. This grouping of Platycentrus and 

Tylocentrus is also supported by molecular phylogenetic analysis (Cryan et al. 2000a). The 

Pieltainellini, a new tribe proposed to accommodate elements of the Boocerini sensu 

McKamey (1998a), are found in Mexico and are the sister group to a majority of the Old 

World centrotines. 

363

The first Old World lineage in the second major centrotine clade are a group of 

genera formerly placed in the Leptocentrini sensu McKamey (1998a) (Fig. 24.2), here 

considered polyphyletic. This group of genera form the Beaufortianini (Figs. 24.2-24.3), a 

new tribe based on the phylogenetic analysis. Imporcitor, distributed in the Palearctic and 

Indomalayan Regions, is basal to Centrolobus, Centruchus, Beaufortiana, Mabokiana, and 

Dukeobelus, a clade primarily distributed in the Afrotropical Region but with a few 

Indomalayan and Palearctic components. Beaufortianines have a long dorsoapical lobe on 

the male lateral plate, although the lobe is lost in the genus Centruchus. Many of the 

characters of this group are intermediate between features found in the New World tribes and 

a majority of the Old World centrotines. A UPGMA analysis (Fig. 24.16) that included the 

genus Centrotusoides grouped the beaufortianines together based on overall similarity. 

A substantial clade of genera including the Centrocharesini, Ebhuloidesini, 

Oxyrhachini, Hypsaucheninii, and the Terentiini (Fig. 24.1) all lack mesothoracic ab- and 

adlateral cucullate setae on the femur, and with the exception of the Centrocharesini, they all 

have a reduced number (0 or 1) of cucullate setae on the first metathoracic tarsomere. The 

Centrocharesini is a highly derived tribe with numerous autapomorphies, including acute 

projections on the pronotum and abdomen, and foliaceous tibiae. The monophyly of the 

clade Oxyrhachini + Ebhuloidesini + Hypsaucheniini is strongly supported by a number of 

characters. The female second valvulae are all short and broad with an undulating dorsal 

margin, the male style clasp is cylindrical in most genera, they lack metathoracic ab- and 

adlateral cucullate setae on the hind femur, and their mesopleural lobes are enlarged. 

The tribe Ebhuloidesini Goding, 1931a, senior synonym of Ebhulini Yuan, in Yuan 

and Chou 2002a, consists of only its nominative genus Ebhul. Sister group to the 

364

Oxyrhachini + Hypsaucheniini (Fig. 24.1), this tribe has the first division of R vein in the 

forewing as R1+2+3 and R4+5 rather than R1 and Rs. The Ebhuloidesini are a distinct lineage in 

the phylogenetic analysis (Fig. 24.1), supporting Yuan and Chou’s (2002a) findings. The 

placement of Oxyrhachini as a monotypic tribe within the Centrotinae is consistent with the 

findings of Dietrich et al. (2001a) (Fig 24.1). Oxyrhachines have paired dorsal swellings on 

the abdomen that are larger on the posterior segments and have Cu1 vein abutting the clavus 

in the forewing, not the marginal vein. Several Oxyrhachis species, representatives of 

generic synonyms of Oxyrhachis, form a monophyletic group (Fig. 24.12) in the analysis, 

partly justifying these synonymies. 

The Hypsaucheniini, as defined here, are monophyletic in the phylogenetic analysis, 

and largely correspond to McKamey’s (1998a) catalog with the exception of Pyrgonota, here 

placed in the Terentiini. Yuan and Chou (2002a) placed the genus Hybandoides in the 

Funkhouserellini (here a synonym of Terentiini) but here it is referred back to the 

Hypsaucheniini (McKamey 1998a). All hypsaucheniines have a median anterior pronotal 

horn. With the exception of Gigantorhabdus, hypsaucheniines have an anomalous 

longitudinal vein in the forewing that may represent a distinct branch of R vein. The generic 

relationships of the Hypsaucheniini are largely unresolved (Fig. 24.12) with the exception of 

the clade including Hybanda, Gigantorhabdus, and Pyrgauchenia. This group of genera has 

conelike projections on the female third valvulae and have the m-cu3 crossvein in the 

forewing basad of the fork of M vein. 

Bulbaucheniini Goding, 1931a, and Funkhouserellini Yuan and Zhang, in Yuan and 

Chou 2002a, are junior synonyms of Terentiini Haupt, 1929c, based on the phylogenetic 

analysis (Figs. 24.1-24.2, 24.14). The type genera of the two former tribes form a 

365

monophyletic group with Terentius (Figs. 24.1, 24.14). Other genera here placed in 

Terentiini were previously incertae sedis or placed in the tribes Centrotypini or 

Leptocentrini, which in the sense of McKamey (1998a) are here considered polyphyletic 

(Fig. 24.2). The Terentiini, as defined here, form a sister group to the Ebhuloidesini + 

Oxyrhachini + Hypsaucheniini (Fig. 24.1). The Australasian genera Eufairmairia, Ceraon, 

and Sextius were a monophyletic group in a molecular analysis of the Membracidae (Cryan et 

al. 2000a). A UPGMA analysis (Fig. 24.17) grouped all of the genera here listed within 

Terentiini together based on morphological similarity. Day’s (1999a) WPGMA analysis 

grouped the Australian membracids into 3 clusters plus the genus Goddefroyinella. Many of 

the genera from the present UPGMA analysis cluster in a similar manner. 

Terentiines have the posterior pronotal process appressed against the scutellum and 

all males have a quadrate style clasp, a unique clasp shape among all centrotines. Although 

many of the the basal relationships of the Terentiini are unresolved (Fig. 24.14), some 

terminal generic relationships are well defined. Anzac, Neosextius, Goddefroyinella, and 

Sextius are a closely related group of genera with reticulate wing venation. Cebes, Sarantus, 

and Ceraon all have the first division of R vein as R1+2+3 and R4+5 rather than R1 and Rs in the 

forewing and lack cucullate setae in row I of the metathoracic tibia. The inornate abdominal 

pits of Bulbauchenia, Funkhouserella, and Pyrgonota are not distinct and the apex of their 

male style clasp is acuminate. 

The next large monophyletic clade contains the tribes (as defined here) Boccharini, 

Leptobelini, Choucentrini, Leptocentrini, Lobocentrini, Xiphopoeini, Micreunini, 

Centrotypini, and Centrotini. With the exception of some Centrotini genera, all the members 

366

of this clade have indistinct abdominal acanthae with their bases not significantly heightened. 

All of these tribes, in addition, except the Lobocentrini, have membranous male style clasps. 

Four genera previously placed in the Leptocentrini sensu McKamey (1998a) (Fig. 

24.2), form the new tribe Lobocentrini. Three of these genera, Lobocentrus, Arcuatocornum, 

and Truncatocornum, form a monophyletic group in the phylogenetic analysis (Fig. 24.1- 

24.2) and Amphilobocentrus grouped with these genera based on morphological similarity in 

a UPGMA analysis (Fig. 24.17). Lobocentrines have numerous cucullate setae, arranged 

irregularly, in row II of the metathoracic tibia and have a dorsoventrally oriented male style 

clasp that is rounded with an acuminate projection. Elaphiceps and Tyrannotus, here placed 

Centrotinae, incertae sedis, form a monophyletic assemblage with the Lobocentrini (Figs. 

24.1-24.2) because their clasps are similar, but are not included in the Lobocentrini because 

they differ significantly in leg chaetotaxy and features of the female genitalia. 

The new tribe Boccharini consists of two genera, Bocchar and Lanceonotus, formerly 

of Leptocentrini sensu McKamey (1998a) (Fig. 24.2). The monophyly of the Boccarhini 

(Figs. 24.1-24.2) is supported by a unique male clasp that is elliptical or circular with a 

preapical ventral extension. The sister group of the Boccharini is composed of a large 

number of genera (8 tribes), all with ab- and adlateral cucullate setae on the metathoracic 

femur. The Leptobelini, a monotypic tribe (Figs. 24.1-24.2), are a distinct lineage with a 

cucullate setal row on the mesothoracic tibia and an acuminate scutellum. Choucentrus, 

Evanchon, and Dograna comprise the Choucentrini, a monophyletic assemblage (Fig. 24.9) 

of centrotines lacking crossvein s in the forewing. They are closely related to the Maarbarini, 

a new tribe consisting of genera formerly incertae sedis or placed in the Centrotini, 

Centrotypini, or Leptocentrini (Fig. 24.2). Males of Dograna (no other choucentrine males 

367

were examined) and the Maarbarini have a triangular style clasp with a basal thickening. 

Most maarbarines have a long acuminate scutellum and parallel, curving longitudinal veins 

in the forewing. The phylogeny of this monophyletic group of genera (Figs. 24.1-24.2, 24.9) 

is well resolved. The most basal genus, Telingana, differs from other maarbarines in 

scutellar characteristics and forewing venation. 

Members of the tribes Leptocentrini, Xiphopoeini, Micreunini, Centrotypini, and 

Centrotini, as defined here, have: a scutellum that extends only slightly beyond the thorax, 

metathoracic tibia with cucullate setal row II double or irregular in most genera, abdominal 

tergal borders not modified into irregular ridges, and the abdominal setal bases are usually 

enlarged. Indeed, previous molecular (Cryan et al. 2000a) and morphological analyses 

(Dietrich et al. 2001a) of the Membracidae also indicated a close relationship between the 

Leptocentrini and Centrotini. 

As mentioned previously, Leptocentrini sensu McKamey (1998a) is polyphyletic in 

the present analysis (Fig. 24.2). The Leptocentrini defined here, however, are a 

monophyletic group of genera (Figs. 24.1-24.2, 24.9). Included in this tribe is the genus 

Demanga, type genus of the Demangini Yuan and Zhang, in Yuan and Chou, 2002a, here 

considered a junior synonym of the Leptocentrini Distant, 1908g. The UPGMA analysis 

(Fig. 24.17) produced a single leptocentrine group based on overall morphological similarity. 

Leptocentrines have a triangular style clasp (except Periaman) and broadened second 

valvulae. 

Members of the monophyletic group of tribes including the Xiphopoeini, Micreunini, 

Centrotypini, and Centrotini, all have elliptical or circular male style clasps. The 

Xiphopoeini, found only in the Afrotropical Region, are a monophyletic group (Fig. 24.7), 

368

characterized by the presence of toothlike projections on the lower margin of the vertex and 

enlarged setal bases dispersed on the abdominal terga. The Micreunini, a monotypic tribe 

(Fig. 24.1), have a long median anterior pronotal horn with suprahumeral horns at its tip. 

The Centrotypini, a polyphyletic group sensu McKamey (1998a), consists here of two 

genera, Centrotypus and Emphusis. Their monophyly is supported by rounded or blunt 

scutellar apices, a condition unique to this tribe. 

The Centrotini is the largest tribe in the subfamily Centrotinae in terms of genera, 

with 47. It is monophyletic, as defined here, based on phylogenetic analyses of selected 

genera (Figs. 24.1, 24.7). The UPGMA analysis of all genera of Centrotini resulted in a 

single group. The phylogenetic tree of the tribe (Fig. 24.7) is fairly well resolved. 

Predominantly Afrotropical, the Centrotini are characterized by their reduced hind wing 

venation where R4+5 and M1+2 veins are fused (3 apical cells). Furthermore, many members 

of the tribe have R1 vein represented by a pterostigma in the forewing and many lack distinct 

frontoclypeal lobes. Centrotus, the type genus, is somewhat enigmatic in its morphology. 

Occupying a basal position in the phylogenetic tree (Fig. 24.7), its frontoclypeal lobes are 

distinct and it lacks a pterostigma on the forewing. 

Summary 

An overall phylogenetic analysis of the subfamily Centrotinae using 116 

morphological characters resulted in a single most parsimonious tree showing numerous 

poly- or paraphyletic tribes as delimited by existing classifications (Ananthasubramanian 

1996a, McKamey 1998a, and Yuan and Chou 2002a). Based on this overall analysis, 11 

tribal synonymies and 1 subfamily synonymy are proposed, 6 new tribes are described, and 

369

the included genera are placed into a total of 23 centrotine monophyletic tribes (see tribal 

descriptions). Tribal relationships were supported with many character changes. The 

subfamily Centrotinae is a monophyletic group supported by the synapomorphy of the 

presence of abdominal inornate pits, each with a lateral seta. The phylogenetic analysis of 

the subfamily resulted in two major clades, each with New World and Old World 

components, plus the New World tribe Centrodontini. 

Apparently, centrotines invaded the Old World twice (Fig. 24.1). One invasion 

included the ancestors of the tribe Gargarini while the other invasion included the ancestors 

of the remaining Old World centrotine tribes. Characters important in elucidating tribal 

relationships include features of the male and female genitalia, the wings, the scutellum, the 

abdomen; and leg chaetotaxy. 

A significant number of the 216 known centrotine genera are poorly represented in 

collections--some being known from only one sex or even a single specimen. Here, all but 9 

genera were placed in tribes supported by quantitative phylogenetic analyses of 

morphological features, or in cases where data were limited, on phenetic overall similarity. 

370

Table 24.1. Character list. 

Head: 

1. Width: (1) less than distance between humeral angles of pronotum; (2) greater than or equal to distance 

between humeral angles of pronotum. 

2. Frontoclypeus: (1) without median longitudinal carina; (2) with median longitudinal carina. 

3. Frontoclypeal lobes: (1) indistinct (Fig. 0.1 C); (2) distinct (Fig. 0.1 D). 

4. Frontoclypeal lobes: (1) not extending to apex of frontoclypeus (Fig. 0.1 D); (2) extending to near apex of 

frontoclypeus (Fig. 0.1 C). 

5. Frontoclypeal margins: (1) distinctly converging (Fig. 4.1 C); (2) parallel or converging only slightly 

ventrally (Fig. 0.1 D); (3) broadly expanding towards apex (Fig. 10.4 G); (4) abruptly expanding near 

apex ( Fig. 11.2 B). 

6. Ocelli: (1) closer to eyes than each other (Fig. 18.2 F); (2) about equidistant from each other and eyes (Fig. 

0.1 C). 

7. Vertex: (1) without multiple toothlike projections on lower margins (Fig. 0.1 D); (2) with multiple toothlike 

projections on lower margins (Figs 23.1 E-F). 

Pronotum/Scutellum 

8. Posterior pronotal process: (1) lacking (Fig. 3.1 A); (2) produced posteriorly (Fig. 3.1 C). [Coding state 1 

makes character #’s 9-12, and 14 inapplicable.] 

9. Posterior process: (1) not originating from median anterior horn (not significantly raised above scutellum) 

(Fig. 0.1 A); (2) originating from median anterior horn (significantly raised above scutellum) (Fig. 0.11 

A). 

10. Posterior process: (1) not significantly extending past m-cu3 crossvein in forewing (Fig. 1.1 A); (2) 

significantly extending past m-cu3 crossvein (Fig. 1.1 D). 

11. Posterior process (shape at base): (1) straight (Fig. 1.1 A); (2) curved dorsally (Fig. 1.1 B). 

12. Posterior process (contact with scutellum): (1) not appressed against scutellum (Fig. 0.1 A); (2) appressed 

against scutellum (for the entire length of the scutellum) (Fig. 0.1 B). 

371

Table 24.1 cont’d. 

13. Posterior process (dorsolateral concealment of scutellum by posterior process): (1) both lateral scutellar 

apices or median acuminate point clearly visible from dorsolateral view (Fig. 0.2 E); (2) 1 lateral apex 

clearly visible from dorsolateral view (Fig. 0.1 B). 

14. Extent of posterior pronotal process relative to scutellum: (1) extended over but not completely concealing 

scutellum (Fig. 0.3 A); (2) polymorphic; (3) completely concealing scutellum (Fig. 0.3 C). [Character 

state #2 is here considered an intermediate state between states #1 and #3. In the genus Sextius 

(Terentiini), for example, some specimens in the same species have the scutellum concealed, while in 

others it is visible. Coding state 2 makes character #13 inapplicable.] 

15. Suprahumeral horns: (1) absent (Fig. 1.1 M); (2) polymorphic; (3) present at base of pronotum (Fig. 1.1 K); 

(4) present but partially fused into a median anterior horn (Figs. 6.5 D-E, G); (5) present on tip of 

median anterior horn (Fig. 16.1 B). [The bifurcate process at the tip of the median anterior horn in 

some treehoppers, for example in many Hypsaucheniini, is here considered homologous to 

suprahumeral horns (character state #5); see discussions within phylogenetic analysis for 

morphological progression of this character.] 

16. Acute pronotal projections or spines: (1) absent (Fig. 0.1 B); (2) present (Figs. 23.1 B, D; Figs. 4.2 I-J). 

[Acute projections, or spines, on the pronotum, here considered homologous, have been derived 

independently in Centrocharesini, Xiphopoeini, several Centrotini (Anchonobelus, Anchonastes, 

Anchonomonoides, Barsumas, Barsumoides, Eumocentrulus, Flatyperphyma, Hamma, Mitranotus), 

several gargarines (Madlinus, Eucoccosterphus, Coccosterphus), and the genera Daimon, Jingkara, 

and Maguva. With the exception of Daimon, the presence of pronotal spines is confined to Old World 

centrotines.] 

17. Median anterior pronotal horn: (1) absent; (2) present (Fig. 16.1 B). 

18. Scutellar keel: (1) present; (2) absent. 

19. Scutellar posterior margin (if visible): (1) acuminate (always exposed, and without median groove (Fig. 0.2 

E); (2) emarginate (notched) (Fig. 0.3 B); (3) acuminate with posterior medial groove . [Coding state 1 

or 3 makes character #20 inapplicable.] 

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20. Scutellar apices (posterior margin notched): (1) acute (Fig. 0.3 B); (2) rounded or blunt (Fig. 0.2 B). 

21. Scutellum length (viewed ventrally, with abdomen removed): (1) not shortened--full notch or acuminate 

point visible (Figs. 0.2 A-B); (2) shortened, at most apices visible (Figs. 0.2 C-D) 

22. Scutellar extension: (1) only slightly extending past thorax (Fig. 0.3 A); (2) posterior half of scutellum 

Pleuron 

extending past thorax (Fig. 0.2 E). 

23. Propleural lobe: (1) absent; (2) present (Fig. 0.3 D). 

24. Mesopleural lobe: (1) not enlarged; (2) enlarged (Fig. 0.3 D). [In centrotines the mesopleural lobe is 

Forewing 

consistently present, but is substantially enlarged in some.] 

25. Clavus: (1) acuminate (Fig. 5.2 E); (2) truncate (Fig. 0.4 A). 

26. Pigmentation of wing: (1) mostly hyaline (Fig. 0.1 A); (2) mostly translucent with darker areas (Fig. 11.1 A- 

B). 

27. Apical limbus: (1) narrow (Fig. 5.2 A); (2) broad (Fig. 0.4 A). 

28. Degree that wing (in repose) is concealed by posterior process: (1) not concealed (Fig. 0.1 A); (2) wing 

partially concealed. 

29. Ratio of wing width to length: (1) < 0.30 (Fig. 11.1 H); (2) 0.30 > 0.40 (Fig. 0.4 A); (3) ≥ 0.40 (Fig. 10.6 

A). 

30. M and Cu (number of branches): (1) 3 veins reaching marginal vein (i.e., M 2-branched, Cu unbranched) 

(Fig. 0.4 A); (2) with 4 branches reaching marginal vein (M apparently 3-branched) (Fig. 2 D of 

Dietrich et al. 2001a); (3) with 5 or more branches reaching marginal vein. 

31. Crossvein s: (1) present (Fig. 0.4 A); (2) absent (Fig. 8.1 G). [Coding state 2 makes character #32 

inapplicable.] 

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32. Position of s-crossvein relative to r-m2 crossvein: (1) distad (Fig. 0.4 A); (2) directly dorsal to or very close 

to r-m2 crossvein (Fig. 7.1 G). 

33. Vein R (number of branches): (1) with 3 or fewer branches reaching marginal vein (Fig. 0.4 A); (2) with 4 

branches reaching marginal vein (Fig. 2 D of Dietrich et al. 2001a); (3) with 5 or more branches 

reaching marginal vein. 

34. Vein R initial division: (1) R1 and RS (Fig. 0.4 A); (2) R1+2+3 and R4+5 (Fig. 9.1 D). 

35. Cu1 vein: (1) distally abutting clavus (Fig. 0.4 A); (2) distally abutting marginal vein (Fig. 1.2 I). 

36. Crossvein r-m1: (1) present (Fig. 0.4 A); (2) absent. [Coding state 2 makes character #’s 49 and 50 


37. Crossvein m-cu1: (1) present on at least one wing (Fig. 0.4 A); (2) absent. [This crossvein, as well as m-cu2 

crossvein, is sometimes only found on either the right or left wing.] 

38. Crossvein m-cu2: (1) present in at least one wing (Fig. 0.4 A); (2) absent. [This crossvein, usually closely 

associated with crossvein r-m1 in centrotines, is equivalent to “m-cu1 crossvein” of Dietrich et al. 

2001a.] 

39. Position of m-cu3 crossvein: (1) distad of fork of vein M (Fig. 0.4 A); (2) basad of fork of vein M (Fig. 10.8 

I). [This crossvein is equivalent to “m-cu2 crossvein” of Dietrich et al. 2001a.] 

40. M and Cu veins: (1) fused, often for considerable distance (Fig. 2.1 H); (2) adjacent with distinct line 

between veins (Fig. 0.4 A); (3) separate (Fig. 7.1 G). 

41. R and M veins (apical fusion): (1) not confluent preapically (Fig. 0.4 A); (2) R4+5 confluent with M distad of 

its fork (Fig. 4.1 D). [Coding state 2 makes character #32 inapplicable because r-m2 crossvein is 

absent when R and M veins are confluent]. 

42. Additional r-m crossveins: (1) present (Fig. 11.3 C); (2) absent. 

43. Anomalous basal r-m crossvein: (1) absent; (2) present (Fig. 11.3 C). 

44. Reticulate venation: (1) absent; (2) present (Fig. 5.2 A). 

45. Pterostigma: (1) absent (Fig. 0.4 A); (2) present (Fig. 4.1 D). [Coding state 1 makes character #46 


374


46. Pterostigma placement: (1) at or near R1 (Fig. 4.1 D); (2) distad of R1 (marginal vein often ambiguous) (Fig. 

10.8 F). 

47. R1 vein represented by large pterostigma: (1) absent; (2) present (Fig. 0.4 A). [Coding state #2 makes 

character #48 inapplicable because R1 vein is not visible.] 

48. R1 vein: (1) parallel for considerable distance with longitudinal veins (Fig. 15.2 G); (2) weakly bent towards 

marginal vein (Fig. 0.4 A); (3) perpendicular to marginal vein (Fig. 3.4 A). 

49. Origin of r-m1 crossvein: (1) arising before initial division of R vein (Fig. 0.4 A); (2) arising near or distad 

of initial division of R vein in at least one wing (Fig. 7.1 G). 

50. Shape of r-m1 crossvein: (1) bent towards R vein (Fig. 10.5 F); (2) parallel to longitudinal veins (Fig. 1.2 I); 

(3) bent nearly to a right angle (Fig. 11.3 C). 

51. Longitudinal veins: (1) strongly curving together in unison apically (Fig. 15.3 E); (2) not strongly curving 

together in unison apically (Fig. 0.4 A). 

52. Longitudinal veins: (1) not parallel apically (Fig. 8.1 G); (2) parallel apically (Fig. 13.4 A). 

53. Relative length of discoidal cells: (1) not similar in length (Fig. 1.2 I); (2) similar in length (Fig. 10.5 F). 

54. R4+5 vein shape prior to s: (1) not significantly angled (Fig. 0.4 A); (2) significantly angled (Fig. 10.5 F). 

55. Base of R2+3 and R4+5 veins: (1) truncate (Fig. 0.4 A); (2) polymorphic; (3) acute (Fig. 10.5 F). 

Hind wing 

56. Veins: (1) R4+5 and M1+2 veins not fused (4 apical cells) (Fig. 0.4 C); (2) R4+5 and M1+2 vein fused for short 

Legs 

distance basally (4 apical cells) (Fig. 5.2 F); (3) R4+5 and M1+2 veins free near apex (4 apical cells) (Fig. 

4.1 E); (4) r-m and m-cu crossveins absent (2 apical cells); (5) R4+5 and M1+2 veins fused (3 apical 

cells)(Fig. 0.4 B); (6) R4+5 vein apparently absent (2 apical cells) (Fig. 3.3 F); (7) polymorphic (genus 

Oxyrhachis only). [State #7 is polymorphic between #1 and #5.] 

57. Front, middle, and hind tibia shape: (1) not foliaceous (Fig. 0.6 A); (2) foliaceous (Fig. 0.7 A). 

375


58. Prothoracic femur ablateral and adlateral cucullate setae: (1) absent; (2) present. 

59. Mesothoracic tibia with longitudinal row(s) of cucullate setae: (1) absent; (2) present (Fig. 3.4 H). 

60. Mesothoracic femur ablateral cucullate setae: (1) absent; (2) present (Fig. 0.7 B). 

61. Mesothoracic femur adlateral cucullate setae: (1) absent; (2) present (Fig. 0.7 B). [Coding state 1 makes 

character #62 inapplicable.] 

62. Mesothoracic leg femur adlateral cucullate setae: (1) apical (Fig. 0.7 B); (2) preapical. 

Metathoracic leg 

63. Ventral setal bases of coxa: (1) small and raised or nearly flat with little or no projection; (2) enlarged, 

raised, and often spinelike. 

64. Ventral setal bases of trochanter: (1) small and raised or nearly flat with little or no projection (Fig. 0.6 A); 

(2) enlarged, raised, and often spinelike; (3) very large spines (Fig. 0.6 D). 

65. Ventral setal bases of femur: (1) small and raised or nearly flat with little or no projection (Fig. 0.6 A); (2) 

enlarged, raised, and often spinelike (Fig. 0.6 D). 

66. Cucullate setae on dorsal femur: (1) absent; (2) present (Fig. 17.3 A). [Coding state 1 makes character #67 


67. Cucullate setae position on dorsal femur: (1) apical (Fig. 17.3 A); (2) in a longitudinal row; (3) scattered. 

68. Femur ablateral cucullate setae: (1) absent; (2) present (Fig. 0.7 B). [Coding state 1 makes character #69 


69. Femur ablateral cucullate setae position: (1) apical (Fig. 0.7 B); (2) preapical. 

70. Femur adlateral cucullate setae: (1) absent; (2) present (Fig. 0.7 B-C). [Coding state 1 makes character #71 


71. Femur adlateral cucullate setae position: (1) apical (Fig. 0.7 B); (2) preapical (Fig. 14.2 A). 

72. Femur ablateral ventrolateral cucullate setae in male: (1) absent; (2) polymorphic; (3) present. 

73. Femur ablateral ventrolateral cucullate setae in female: (1) absent; (2) polymorphic; (3) present (Fig. 17.3 

A). 

376


74. Tarsomere I apical cucullate setae: (1) absent (Fig. 19.2 B); (2) 1 seta (Fig. 0.7 E, Fig. 11.4 F); (3) 2 or more 

setae (Fig. 0.7 D, Fig. 2.1 J). 

75. Tibia setal row I: (1) non-cucullate (Fig. 22.12 B, Fig. 0.6 C, Fig. 0.7 A); (2) cucullate (Fig. 0.5 A-C). 

76. Tibia setal row II: (1) non-cucullate (Fig. 0.7 A); (2) cucullate (Figs. 0.6 A-B). [Coding state 1 makes 

character #77 inapplicable.] 

77. Tibia setal row II: (1) mostly single row (Fig. 0.6 A); (2) polymorphic; (3) irregular or double row (Fig. 0.6 

B). 

78. Tibia setal row III: (1) cucullate (Fig. 0.5 A); (2) non-cucullate. 

Abdomen 

79. Shape (cross-section): (1) dorsoventrally flattened; (2) nearly triangular. 

80. Sternum III and/or IV transverse carina: (1) absent; (2) present (Fig. 6A of Dietrich et al. 2001a). 

81. Sternum longitudinal median carina: (1) absent; (2) polymorphic; (3) present on at least one segment (Fig. 

0.8 B). 

82. Number of inornate pits along side of abdominal segments: (1) 3 or fewer per segment (Fig. 5.4 G); (2) 4 or 

more per segment (Fig. 0.8 E). 

83. Paired dorsal swellings or remnants of swellings: (1) absent (Fig. 0.8 E); (2) larger anteriorly (Fig. 0.8 C); 

(3) larger posteriorly (Fig. 0.8 D). 

84. Enlarged setal bases: (1) absent; (2) present (Figs. 0.8 A, E; Fig. 10.17 D). [Coding state 1 makes character 

#85 inapplicable.] 

85. Enlarged setal bases position: (1) only on tergal borders (Fig. 0.8 E, Fig. 10.17 D); (2) extending to tergal 

segment, sparse; (3) extending to tergal segment, numerous (Fig. 0.8 A). 

86. Tergum III ventrolateral margin: (1) carinate (Fig. 5 A of Dietrich et al. 2001a); (2) shelflike (Fig. 5 B of 

Dietrich et al. 2001a); (3) upcurved groove (Fig. 5 C of Dietrich et al. 2001a). 

87. Tergal borders: (1) not extensively modified (Fig. 0.8 E); (2) modified into irregular ridges (Fig. 0.3 A). 

377


Female 

88. Second valvulae shape: (1) significant broadening absent (Fig. 5.3 A); (2) short and broad with undulation 

on dorsal margin (Fig. 11.5); (3) abrupt slight broadening (Fig. 22.13 A); (4) gradual broadening (Fig. 

22.13 C); (5) broad throughout with very slight increase and decrease after midpoint (Fig. 21.2 C-F). 

[Coding states 1 or 2 makes character #’s 89 and 90 inapplicable.] 

89. Second valvulae shape (broadening): (1) widest before or near midpoint (Fig. 22.13 C); (2) widest past 

midpoint (Fig. 4.2 A). 

90. Second valvulae dorsal margin: (1) tapering evenly after broadening (Fig. 0.9 D); (2) tapering unevenly 

after broadening (Fig. 22.13 A). 

91. Second valvulae width near base: (1) narrow (Fig. 5.3 A); (2) broad or broadening (0.9 D). 

92. Second valvulae curvature: (1) not curved (Fig. 3.5 E); (2) curved (concave) (Fig. 3.5 I). 

93. Second valvulae teeth development: (1) teeth absent or indiscernible (Fig. 0.9 D); (2) fine and distinct (Fig. 

1.4 D); (3) large (Fig. 18.5 J). [Coding state 1 makes character #94 inapplicable]. 

94. Second valvulae teeth: (1) present to tip of apex (Fig. 3.6 F); (2) absent apically (Fig. 22.14 H). 

95. Second valvulae acute projections: (1) absent (Fig. 18.5 H); (2) present (Fig. 18.5 J). [Acute projections on 

the second valvulae are usually triangular and apparently not homologous to dorsal teeth.] 

96. Third valvulae large ventral projections: (1) absent; (2) present (Fig. 11.6 A-D). 

97. Third valvulae apical cleft or groove: (1) not distinct; (2) distinct (Fig. 0.9 D). 

Male 

98. Pygofer with dorsal projection: (1) absent; (2) present (Fig. 11.7 H). 

99. Pygofer with lateral plate: (1) apparently absent; (2) free distally (Fig. 6 D of Dietrich et al. 2001a); (3) 

entirely free (Fig. 0.9 A). 

100. Lateral plate dorsoapical posterior lobe: (1) absent (Fig. 1.6 B); (2) present (Fig. 0.9 A). [Coding state 1 

makes character #’s 101 and 102 inapplicable.] 

101. Lateral plate dorsoapical lobe length: (1) short (Fig. 0.9 A); (2) long (Fig. 1.6 A). 

378


102. Lateral plate dorsoapical lobe shape: (1) angled dorsally (Fig. 0.9 A); (2) angled laterally (Fig. 11.7 D); (3) 

angled ventrally (Fig. 3.8 F). 

103. Lateral plate ventral lobe: (1) absent; (2) small lobe present; (3) long and large lobe present (Fig. 3.8 C). 

104. Subgenital plate: (1) distinct division absent (Fig. 2.2 I); (2) distinct division present (Fig. 3.8 I). 

105. Style clasp overall shape: (1) rounded with acuminate projection (Fig. 10.13 N); (2) truncate with 

acuminate projection (Figs. 18.7. A-B); (3) elliptical or circular (Figs. 7.3 A-B); (4) expanding 

dorsoventrally and laterally with a sclerotized ridge (Figs. 1.5 C-K); (5) cylindrical (Fig. 3.7 H); (6) 

quadrate (Fig. 22.18 A-K); (7) triangular (Fig. 0.9 A); (8) rounded at apex with preapical ventral 

extension (2.2 E-F). [Here, 

the clasp is defined as the apical portion of the style, which is often flattened or expanded.] 

106. Style clasp thickness: (1) thickened (Figs. 18.6 E, M); (2) thickened dorsally, membranous ventrally; (3) 

membranous (Fig. 0.9 B-C). 

107. Style clasp orientation: (1) dorsoventrally flattened or curving dorsally (Fig. 0.9 C); (2) oriented laterally 

(Figs. 0.9 A-B). 

108. Style clasp basal thickening: (1) absent; (2) present (Fig. 15.5 A-D). 

109. Style clasp angle (viewed laterally): (1) angled ventrally (Fig. 1.5 D); (2) angled dorsally (Fig. 7.3 A); (3) 

not angled (Fig. 18.6 J). 

110. Style clasp with acuminate apex: (1) absent; (2) blunt (Fig. 0.9 A); (3) acute (Fig. 15.5 D). 

111. Style shank ventral margin: (1) without preapical broadening; (2) with preapical broadening (Figs. 0.9 A, 

2.2 E, 22.18 A). 

112. Style shank shape (viewed laterally): (1) without significant arch (Fig. 0.9 C); (2) apex of arch at midpoint 

of style shank (Fig. 0.9 A); (3) apex of arch just prior to style clasp (Fig. 9.2 A). 

379


Abdominal fine structure 

113. Inornate pits and associated lateral setae: (1) absent (Fig. 6 C of Dietrich et al. 2001a); (2) present (Figs. 

0.8 E, 0.10 B-D); (3) setae present, pits indistinct (Fig. 22.22 C). [Coding state 1 makes character #82 

inapplicable]. 

114. Acanthae development: (1) distinct (Figs. 0.10 A, 0.10 C); (2) indistinct, acanthae blending together (Fig. 

0.10 B). 

115. Acanthae base development: (1) heightened and broad (Fig. 0.10 C); (2) not significantly heightened or 

broadened (Fig. 0.10 A). 

116. Acanthae/microtrichia development: (1) single acanthae without ornamentation (Fig. 0.10 D); (2) acanthae 

multidentate (Fig. 0.10 A); (3) acanthae divided into threadlike microtrichia (Fig. 0.10 B). 

380

Table 24.2. Number of taxa and characters for phylogenetic (PAUP*) and phenetic (UPGMA) analyses. 

Characters in the “other” column were excluded because they were highly homoplasious and not informative as 

tribal or generic characters for the analyzed taxa. 

INCLUDED EXCLUDED 

Analysis no. no. Constant Parsimony- Other 

1. PAUP*: 

Centrotinae 

(overall) 

2. PAUP*: 

Beaufortianini, 

Nessorhinini, 

Pieltainellini, 

Platycentrini 

3. PAUP*: 

Boocerini, 

Centrodontini 

4. PAUP*: 

Centrotini, 

Xiphopoeini 

5. PAUP*: 

Choucentrini, 

Leptocentrini, 

Maarbarini 

6. PAUP*: 

Gargarini 

7. PAUP*: 

Hypsaucheniini, 

Oxyrhachini 

8. PAUP*: 

Terentiini 

9. UPGMA: 

Centrotini, 

Beaufortianini, 

Gargarini, 

Lobocentrini 

10. UPGMA: 


Nessorhinini, 

Terentiini 

genera 

characters 

381 

uninformative 

69 109 69, 98 (see text) 42, 44, 

57, 93, 

30 77 7, 9, 17, 20, 31, 35, 

39, 41, 43, 46, 47, 51, 

57, 58, 62, 63, 65, 68, 

69, 76, 78, 85, 96, 98, 

108, 110, 111 

23 86 7, 9, 16, 17, 20, 31, 32, 

35, 41, 43, 46, 51, 52, 

69, 85, 90, 96, 98, 108, 

110, 111 

41 81 5, 9, 10, 17, 20, 28, 31, 

32, 35, 39, 43, 46, 51, 

57, 58, 63, 65, 68, 69, 

76, 78, 82, 96, 98, 103 

104, 108 

32 74 5, 7, 14, 16, 20, 24, 28, 

35, 39, 43, 46, 57, 58, 

63, 65, 68-70, 76, 78, 

80, 85, 96, 98 

35 71 7, 9, 17, 20, 22, 24, 28, 

31, 32, 35, 37, 43, 51, 

52, 57, 68, 69, 70, 76, 

78, 80, 82, 85, 90, 96, 

98, 108, 110, 111 

17 68 7, 9, 13, 20, 22, 28, 31, 

32, 41, 45-47, 51, 52, 

55, 58, 62, 63, 69, 71- 

73, 80, 81, 84-87, 89, 

90, 95, 103, 104, 108, 

110, 111 

28 73 5, 7, 9, 13, 16, 20, 22, 

31, 32, 39, 41, 43, 46, 

47, 51, 58, 62, 63, 65, 

69, 71, 72, 76, 85, 96, 

98, 103, 104, 108 

22-24, 32, 36, 

56, 82, 84, 

89, 103 

22, 28, 36, 

46, 62, 63, 

80, 84 

6, 14, 36, 62, 

70, 71, 74, 

102 

9, 17, 23, 37, 

41, 44, 47, 

56, 71, 75, 

82, 83, 89, 

90, 103, 116 

12, 23, 36, 

41, 44, 47, 

49, 60, 61, 

62, 72-74, 75, 

102, 112 

4, 16, 36, 53, 

54, 65, 77, 

82, 92, 100, 

114, 115 

12, 36, 45, 

50, 55, 73, 

78, 80, 81, 

83, 84, 87 

91 115 -- -- 94 

76 115 -- -- 94 

94 

42, 44 

42 

-- 

93, 94 

-- 

-- 

42, 57

Table 24.3. Summary of phylogenetic analyses 1-8. 

Analysis Fig. #’s 

SCT (Strict consensus tree) 

MPT (Most parsimonious 

tree) 

EPT (Equally parsimonious 

tree) 

1. Centrotinae 

(overall) 

2. Beaufortianini, 

Nessorhinini, 

Pieltainellini, 

Platycentrini 

3. Boocerini, 

Centrodontini 

4. Centrotini, 

Xiphopoeini 

382 

no. 

of 

MPT 

trees 

Steps Consistency 

index (CI) 

24.1 (MPT) 1 665 0.23 0.60 

24.3 (SCT), 24.4 (EPT) 2 268 0.36 0.57 

24.5 (SCT), 24.6 (EPT) 2 209 0.50 0.64 

24.7 (SCT), 24.8 (EPT) 4 277 0.37 0.62 

5. Choucentrini, 


Maarbarini 

24.9 (MPT) 1 207 0.43 0.69 

6. Gargarini 24.10 (SCT), 24.11 

(EPT) 

28 224 0.38 0.60 

7. Hypsaucheniini, 24.12 (SCT), 24.13 25 150 0.55 0.71 

Oxyrhachini 

(EPT) 

8. Terentiini 24.14 (SCT), 24.15 

(EPT) 

4 224 0.42 0.60 

Retention 

index (RI)

Fig. 24.1. Phylogenetic relationships within Centrotinae (Analysis 1, PAUP*). 

Single most parsimonious tree with branch lengths indicating numbers of assigned 

character changes. Nodes are labeled by number below or above each branch 

(CI=0.23, RI=0.60, length 665). 

383

Table 24.4. List of apomorphies for Analysis 1 (Fig. 24.1). 

Characters are listed with states in parentheses; non-homoplastic 

changes are marked by an asterisk (*). 

Node 134 (Centrotinae) 50(1), 66(1), 79(2), 113(2) 

Node 133 25(2), 26(1), 27(2), 74(3), 

87(2), 101(1) 

Node 132 49(2), 54(2), 67(1)*, 81(3), 

90(2), 104(2) 

Node 131 (Monobelini) 4(2), 100(1), 102(2), 105(2), 

114(2) 

Node 130 10(2), 37(1), 53(1), 55(3), 

66(2), 72(3), 73(3) 

Node 129 60(2), 61(2), 89(2), 92(2), 

106(3) 

Node 128 (Boocerini) 13(1), 21(1), 48(3), 88(3), 

103(3)*, 109(1), 115(1) 

Node 127 4(2), 39(2)*, 56(5), 58(2)*, 

95(2) 

Node 126 12(1), 71(2), 105(5) 

Node 125 6(1), 45(2), 47(2), 59(2), 88(1) 

Node 124 (Gargarini) 5(3)*, 55(3), 77(3), 87(1) 

Node 123 3(2), 56(5) 

Node 122 46(2),* 83(2), 84(2), 86(2), 

95(2) 

Node 121 29(3), 97(2) 

Node 120 63(2), 64(2), 65(2), 101(2), 

115(1) 

Node 119 16(2), 26(2), 53(1), 116(1) 

Node 118 15(3), 105(4), 107(2), 115(1), 

116(2) 

Node 117 38(1), 52(2), 97(2) 

Node 116 (Nessorhinini) 10(2), 14(3), 40(2), 50(2), 

83(2), 86(2), 105(2), 112(2) 

Node 115 4(2), 6(1), 45(2), 70(1), 80(2), 

86(3), 107(1) 

Node 114 37(1), 38(2), 81(3), 92(2), 

116(3) 

Node 113 12(1), 21(1) 

Node 112 (Pieltainellini) 48(3), 49(2), 71(2), 92(2), 

95(2), 102(2), 105(3) 

Node 111 3(2), 53(1) 

Node 110 (Beaufortianini) 101(2), 109(1) 

Node 109 50(2), 83(2), 95(2) 

Node 108 10(2), 88(3), 105(7), 116(3) 

Node 107 13(1), 22(2), 54(2) 

Node 106 55(3), 106(3), 109(2), 114(2), 

115(2) 

Node 105 (Boccharini) 88(4), 91(2), 100(1), 105(8)*, 

111(2) 

Node 104 60(2), 61(2), 77(3), 112(2) 

Node 103 22(1), 50(2), 54(1), 55(1), 

84(2), 87(1) 

Node 102 (Leptocentrini) 40(2), 48(3), 52(2), 81(3), 

86(2), 91(2), 109(1), 110(2), 

111(2) 

Node 101 88(4), 97(2) 

Node 100 11(2), 53(2), 90(2), 105(3) 

Node 99 (Centrotini) 13(2), 56(5), 85(2)*, 114(1) 

Node 98 3(1), 4(2), 48(3), 49(2), 50(1), 

54(2), 55(2) 

Node 97 45(2), 47(2) 

Node 96 26(2), 45(2), 86(3), 89(2), 

92(2), 95(2), 97(2), 112(1) 

Node 95 3(1), 11(1), 32(2), 53(1), 

54(2), 81(3), 88(1) 

384 

Node 94 (Centrotypini) 13(2), 20(2)*, 37(1), 40(2), 

45(1), 49(2), 84(1), 86(2), 

112(2) 

Node 93 32(2), 49(2), 101(2) 

Node 92 3(1), 11(2), 77(1), 88(1), 

92(2), 108(2)*, 109(1), 110(3) 

Node 91 (Maarbarini) 27(1), 48(1), 50(2), 51(2)*, 

52(2) 

Node 90 3(2), 26(2), 55(1), 92(1), 95(2) 

Node 89 11(1), 13(2), 97(2) 

Node 88 81(2), 86(2), 97(2), 104(2), 

105(1), 107(1) 

Node 87 48(1), 50(2), 54(1), 60(2), 

61(2), 88(1) 

Node 86 (Lobocentrini) 10(1), 40(2), 48(3), 53(2), 

62(2), 71(2), 77(3), 86(1), 

95(2), 114(2), 115(2) 

Node 85 13(2), 26(2), 74(2) 

Node 84 24(2), 26(2), 70(1), 75(1), 

82(1), 87(1), 90(2) 

Node 83 21(2), 37(1), 74(2), 105(5), 

106(2), 109(1), 112(3), 115(2) 

Node 82 3(1), 15(1), 68(1), 76(1), 

88(2)*, 102(2), 106(3), 116(2) 

Node 81 23(2), 38(1), 50(3)*, 78(2), 

82(2), 112(1) 

Node 80 (Hypsaucheniini) 3(2), 5(4)*, 17(2), 43(2)*, 

48(1) 

Node 79 (Terentiini) 12(2), 48(1), 50(2), 70(2), 

91(2), 105(6)*, 111(2) 

Node 78 40(3), 52(2), 97(2) 

Node 77 26(1), 86(2), 88(1) 

Node 76 52(1), 66(2), 75(2), 82(2), 

91(1), 95(2), 114(2) 

Node 75 38(1), 54(2), 74(1) 

Node 74 26(1), 28(2), 68(1), 70(1) 

Node 73 4(2), 15(5), 17(2), 24(1), 

110(3), 113(3)*, 114(2) 

Node 72 23(2), 37(2) 

Microcentrus 15(2), 38(1), 48(1), 88(3), 

91(2), 97(2), 102(3), 116(2) 

Centronodus 2(2), 10(2), 15(3), 21(1), 

23(2), 30(3), 33(2), 60(2), 

61(2), 74(3), 79(2), 91(1), 

92(2), 99(2) 

Paracentronodus 2(2), 30(3), 33(3)*, 38(1), 

100(1) 

Nicomia 4(2), 25(2), 27(2), 59(1), 

75(1), 99(3), 105(5), 113(2), 

115(1) 

Tolania 15(3), 36(2)*, 40(1), 77(3), 

114(2) 

Centrodontus, Centrodontini 14(3), 29(3), 56(5), 63(2), 

64(2), 65(2), 68(1), 70(1), 

75(1), 76(1), 78(2), 82(1), 

92(2), 102(2) 

Brachycentrotus 29(3), 56(4)*, 82(1) 

Monobelus 21(1), 22(2), 83(2), 88(3), 

97(2) 

Monobeloides 14(3), 28(2), 49(1), 81(1), 

92(2), 100(2), 104(1) 

Amblycentrus 13(2), 29(3) 

Brachybelus 56(6)*, 65(2) 

Boocerus 10(2), 11(2), 15(3), 55(3), 

92(1), 112(3) 

Abelus 2(1), 81(2), 101(2), 107(2)


Ischnocentrus 71(1), 102(3) 

Aleptocentrus 6(1), 48(1) 

Parayasa 45(2), 81(1), 101(2), 104(1), 

105(3), 107(2) 

Pantaleon 15(3), 26(2), 54(1), 81(2), 

106(1), 114(2) 

Gargara -- 

Tricentrus 15(2), 45(2), 64(3), 100(1), 

106(1) 

Coccosterphus -- 

Madlinus 3(1), 14(3), 27(1), 77(1) 

Platycentrus, Platycentrini 60(2), 71(2), 72(3), 73(3), 

74(2), 77(3), 79(1), 88(5)* 

Callicentrus 26(2), 48(1), 97(1), 105(4) 

Orthobelus 14(2), 53(1), 100(1), 109(2), 

112(1) 

Goniolomus 28(2), 49(2), 50(1), 101(2) 

Nessorhinus 15(4), 52(1), 95(2) 

Pieltainellus 15(2), 79(1) 

Spathocentrus 11(2), 26(2), 54(2) 

Imporcitor 21(2), 27(1), 48(3), 49(2), 

92(2), 97(2) 

Beaufortiana 37(1) 

Dukeobelus 15(1), 26(2) 

Centrochares, Centrocharesini 5(1)*, 11(2), 16(2), 41(2)*, 

45(2), 47(2), 49(2), 56(3)*, 

64(2), 65(2), 77(3), 80(2), 

83(2), 86(2), 89(2), 96(2)*, 

100(1), 110(2) 

Ebhul, Ebhuloidesini 11(2), 21(1), 34(2), 37(2), 

49(2), 56(5) 

Oxyrhachis, Oxyrhachini 12(2), 14(3), 15(2), 26(1), 

35(2), 40(2), 56(7)*, 83(3)*, 

116(1) 

Hypsauchenia 15(5), 21(1), 29(1), 97(2) 

Hypsolyrium 49(2) 

Ceraon 34(2), 48(2), 90(1), 101(2) 

Eufairmairia 74(1), 88(4) 

Sertorius 54(2), 77(3) 

Terentius 15(1), 24(1), 29(3), 48(2), 

84(2), 92(2), 97(1) 

Anzac 15(1), 27(1), 29(3), 88(4), 

90(1), 115(1), 116(2) 

Sextius 14(2), 35(2), 55(2), 89(2), 

91(1), 92(2), 101(2) 

Bulbauchenia 14(3), 29(1), 74(2) 

Funkhouserella 27(1), 88(4), 97(2) 

Pyrgonota 6(1), 10(1), 70(1) 

Elaphiceps 3(1), 4(2), 9(2), 15(5), 17(2), 

26(2), 32(2), 55(2), 89(2), 

101(2), 103(2) 

Tyrannotus 11(2), 15(2), 72(3), 87(1), 

88(4), 91(2), 102(3) 

Lobocentrus 22(1), 49(2), 81(1), 92(2), 

97(1), 101(2) 

Arcuatocornum 11(2), 53(1), 87(1) 

Truncatocornum 95(1) 

Bocchar 15(2), 37(1), 53(2) 

Lanceonotus 11(2), 26(2), 48(1), 55(2) 

Leptobelus, Leptobelini 9(2), 15(5), 17(2), 19(1), 

59(2), 72(3), 73(3), 97(2), 

103(2), 105(3) 

Dograna, Choucentrini 13(2), 22(1), 31(2)*, 45(2), 

60(1), 61(1), 111(2) 

385 

Maarbarus 4(2), 19(1) 

Indicopleustes 10(1) 

Pogon 3(1), 88(3), 95(1) 

Pogontypus 29(3), 49(1), 92(2) 

Awania 10(1), 15(1), 72(3) 

Leptocentrus 29(1), 73(2), 81(2) 

Umfilianus 13(2), 15(2), 38(1), 49(2), 

53(2) 

Xiphopoeus, Xiphopoeini 7(2)*, 16(2), 24(2), 38(1), 

48(1), 85(3)*, 100(1) 

Micreune, Micreunini 4(2), 6(1), 15(5), 17(2), 52(2), 

62(2), 71(2), 101(2), 105(7), 

110(2) 

Centrotypus 12(2), 14(2), 40(3), 53(2), 

100(1) 

Emphusis 54(1), 95(1), 97(1) 

Centrotus 72(2)*, 73(2), 91(2), 101(2), 

115(1), 116(2) 

Capeneralus 40(2) 

Anchon 13(1), 86(2), 88(1), 92(2), 

95(2), 114(2) 

Takliwa 116(1)

Table 24.5. List of taxa in the overall phylogenetic analysis (1) of the Centrotinae 

(alphabetized by genus). 

Abelus inermis (Lethierry), USNM; A. luctuosus Stål, NCSU, USNM; Amblycentrus 

pubescens Fowler, USNM; Anchon sp., USNM; A. limbatum Schmidt, USNM; A. nodicornis 

(Germar), USNM, PPRI, A. ulniforme Buckton, USNM; A. ximenes Capener, MNHN; Anzac 

bipunctatum (Fabricius), ANIC; Arcuatocornum sp., LBOB; Awania sp., CASC; A. typica 

Distant, PPRI, MNHN; Beaufortiana distanti (Funkhouser), PPRI; B. viridis (Capener), 

AMNH, USNM; Bocchar confusus (Distant), USNM; B. montanum Jacobi, SMTD, USNM; 

Boocerus gilvipes Stål, NCSU, USNM; Brachycentrotus punctatus (Metcalf and Bruner), 

NCSU; B. rufinervis Ramos, USNM; Brachybelus sp., NCSU; B. cruralis Stål, USNM; 

Bulbauchenia sp. (probably mirablis), USNM; B. bakeri (Funkhouser), USNM, NCSU; B. 

globosa (Funkhouser), USNM; B. mirabilis (Funkhouser), USNM; B. rugosa (Funkhouser), 

USNM; Callicentrus ignipes (Walker), BMNH, USNM; Capeneralus lobatus (Capener), 

PPRI; C. subnodosus (Jacobi), PPRI; Centrochares horrifica (Westwood), USNM. 

Centrodontus atlas (Goding), NCSU, USNM; C. atlas atlas (Goding), USNM; C. atlas 

paucivenosus Cook, NCSU; Centronodus denticulus Funkhouser, NCSU; C. rochalimai 

Fonseca, NCSU; Centrotus cornutus (Linnaeus), LSUK, NCSU, USNM; Centrotypus sp., 

NCSU; C. assamensis (Fairmaire), SHMC; C. flexuosus (Fabricius), USNM; Ceraon 

tasmaniae (Fairmaire), USNM; Coccosterphus sp., SHMC; C. minutus (Fabricius), USNM, 

BMNH; C. obscurus Distant, USNM; Dograna suffulta Distant, PPRI, CASC; Dukeobelus 

simplex (Walker), USNM, PPRI; Ebhul varium (Walker), USNM; Elaphiceps cervus 

Buckton, USNM; E. javanensis Funkhouser, USNM; Emphusis obesa (Fairmaire), NCSU, 

USNM; Eufairmairia decisa USNM; E. fraterna Distant, ANIC, USNM; Funkhouserella 

arborea (Funkhouser), USNM; F. binodis (Funkhouser), USNM; F. brevifurca 

(Funkhouser), USNM; F. bulbiturris (Funkhouser), USNM; F. pinguiturris (Funkhouser), 

USNM; F. sinuata (Funkhouser), USNM; Gargara aenea Distant, NCSU; G. fraterna 

Distant, NCSU; G. genistae (Fabricius), NCSU, AMNH, USNM; G. nyanzai Funkhouser, 

NCSU; Goniolomus tricorniger Stål, USNM; Hypsauchenia hardwickii (Kirby), USNM, 

MNHN; Hypsolyrium uncinatum (Stål), USNM; Imporcitor typicus Distant, BMNH; 

Indicopleustes albomaculata Distant, BMNH; Ischnocentrus sp., NCSU; I. inconspicuous 

Buckton, USNM; I. niger Stål, USNM; Lanceonotus basilicus Capener, PPRI; L. defloccatus 

Capener, PPRI; Leptobelus dama (Germar), USNM; L. metuendus (Walker), USNM; 

Leptocentrus sp., USNM; L. bos (Signoret) USNM, PPRI, MNHN; L. reponens (Walker), 

NCSU; L. taurus (Fabricius), NCSU. Lobocentrus falco (Buckton), USNM; L. zonatus Stål, 

USNM; Maarbarus sp., USNM; M. bubalus (Kirby), BMNH; Micreune formidanda Walker, 

USNM; Microcentrus caryae, NCSU; Monobeloides stuarti Ramos, NCSU, SHMC; 

Monobelus sp., NCSU; M. biguttatus (Fabricius), USNM; M. flavidus (Fairmaire), USNM; 

Nessorhinus gibberulus Stål, USNM, N. gracilis Metcalf and Bruner, NCSU; N. vulpes 

Amyot and Serville, USNM; Nicomia sp., BMNH; N. cicadoides (Walker), BMNH, MNHN; 

Orthobelus sp., det., USNM, O. urus (Fairmaire), USNM; Oxyrhachis carinata 

(Funkhouser), AMNH; O. delalendei Fairmaire, AMNH; O. sulcicornis (Thunberg), NCSU; 

O. taranda (Fabricius), NCSU; Pantaleon dorsalis (Matsumura), USNM, SHMC; P. 

montiferum (Walker), BMNH; Paracentronodus sp., NCSU; Parayasa elegantula Distant, 

USNM; P. typica Distant, USNM; Pieltainellus sp., SHMC; P. boneti Peláez, AMNH; 

386

Platycentrus acuticornis Stål, USNM; Pogon incurvatum Buckton, BMNH; Pogontypus sp., 

USNM; P. complicatus (Melichar), BMNH; P. horvathi Distant, BMNH; Pyrgonota sp., 

USNM; P. bifoliata (Westwood), USNM, DJFC; Sertorius sp., ANIC; S. australis 

(Fairmaire), USNM; Sextius kurandae Kirkaldy, USNM; S. virescens (Fairmaire), USNM, 

ANIC; Spathocentrus intermedius, OXUM, SHMC, CNCI; Takliwa carteri Funkhouser, 

PPRI, MNHN; Terentius convexus Stål, USNM, ANIC; Tolania sp., NCSU, MNHN; 

Tricentrus curvicornis Funkhouser, NCSU; T. fairmairei (Stål), USNM; Truncatocornum 

sp., LBOB; Tyrannotus tyrannicus Capener, PPRI, MNHN; Umfilianus declivis Distant, 

USNM; Xiphopoeus sp., USNM; X. erectus Distant, USNM; X phantasma Signoret, PPRI. 

387

Fig. 24.2. Phylogenetic relationships within Centrotinae (Analysis 1, PAUP*) with 

“existing” tribal names based on Ananthasubramanian (1996a), McKamey (1998a), and 

Yuan and Chou (2002a). “Revised” tribal classification based on the current study are 

shown at far right. 

388

Fig. 24.3. Phylogenetic relationships within Beaufortianini, Nessorhinini, 

Pieltainellini, and Platycentrini (Analysis 2, PAUP*). Strict consensus of 2 equally 

parsimonious trees (CI=0.36, RI=0.57, length 268). 

389

Fig. 24.4. Phylogenetic relationships within Beaufortianini, Nessorhinini, 

Pieltainellini, and Platycentrini (Analysis 2, PAUP*). One of 2 equally 

parsimonious trees with branch lengths indicating numbers of assigned character 

changes. Nodes are labeled by number below or above each branch. 

390




Node 51 15(3), 105(4), 107(2), 115(1) 

Node 50 38(1), 49(1) 

Node 49 (Platycentrini) 60(2), 88(5)* 

Node 48 (Nessorhinini) 10(2), 14(3), 40(2), 83(2), 

86(2), 93(3), 112(2) 

Node 47 4(2), 6(1), 45(2), 80(2), 86(3), 

105(2), 107(1) 

Node 46 15(1), 29(1), 48(1) 

Node 45 53(1), 71(2) 

Node 44 70(1)*, 97(2) 

Node 43 24(2), 37(1), 38(2), 50(2), 

52(2), 81(3), 92(2) 

Node 42 53(1), 97(2) 

Node 41 75(1), 81(1) 

Node 40 12(1), 21(1), 48(3), 92(2) 

Node 39 (Pieltainellini) 71(2), 94(1), 95(2), 102(2), 

105(3) 

Node 38 (Beaufortianini) 3(2), 53(1), 101(2), 109(1) 

Node 37 48(2), 49(1), 50(2), 83(2) 

Node 36 92(1), 95(2) 

Node 35 10(2), 116(3) 

Node 34 94(1) 

Node 33 71(2), 81(2), 86(2), 104(2) 

Pieltainellus 15(2), 79(1) 

Spathocentrus 11(2), 26(2), 54(2) 

Imporcitor 21(2), 27(1), 93(3), 97(2) 

Maguva 11(2), 16(2), 26(2), 105(3) 

Mabokiana 10(2), 54(2) 

Beaufortiana 37(1), 93(3) 

Dukeobelus 15(1), 26(2) 

Centrolobus 11(2), 13(1) 

Centruchus 5(3), 38(1), 53(2), 91(2), 

97(2), 100(1) 

Platycentrus 52(2), 71(2), 72(3), 73(3), 

74(2), 77(3), 79(1), 97(2) 

Tylocentrus 15(2), 48(3), 53(1), 54(2), 

61(2), 87(1), 100(1), 105(2) 

Callicentrus 48(1) 

Orthobelus 14(2)*, 26(1), 94(1), 100(1), 

105(2), 109(2)*, 112(1), 

116(3) 

Daimon 4(2), 14(1), 16(2), 45(2), 

54(2), 55(2)*, 86(3) 

Marshallella 15(2), 37(2) 

Goniolomus 28(2), 49(2), 52(2), 101(2) 

Nessorhinus 15(4), 50(2), 95(2) 

Spinodarnoides -- 

Orekthoptera 15(4), 26(2), 40(1), 80(1), 

81(3), 100(1) 

Paradarnoides 6(2), 29(2), 50(2), 112(1) 

391

Fig. 24.5. Phylogenetic relationships within Boocerini and Centrodontini 

(Analysis 3, PAUP*). Strict consensus of 2 equally parsimonious trees 

(CI=0.50, RI=0.64, length 209). 

392

Fig. 24.6. Phylogenetic relationships within Boocerini and Centrodontini 

(Analysis 3, PAUP*). One of 2 equally parsimonious trees with branch 

lengths indicating numbers of assigned character changes. Nodes are 

labeled by number below each branch 

393




Node 42 50(1), 54(2), 66(1), 67(1)*, 

79(2), 101(1), 102(2), 113(2) 

Node 41 25(2), 26(1), 27(2), 74(3), 

81(3), 87(2), 104(2) 

Node 40 4(2), 93(3), 100(1), 105(2)*, 

114(2) 

Node 39 10(2), 37(1)*, 53(1), 55(3), 

66(2), 72(3)*, 73(3)*, 94(2) 

Node 38 60(2), 61(2), 92(2), 102(1), 

106(3) 

Node 37 (Boocerini) 21(1), 48(3)*, 88(3), 103(3)*, 

109(1)*, 115(1) 

Node 36 13(1), 89(2)* 

Node 35 4(2), 39(2)*, 56(5), 58(2)*, 

95(2) 

Node 34 12(1)*, 88(1), 105(5) 

Node 33 10(2), 11(2), 15(3), 71(2), 

94(2), 112(3) 

Node 32 3(2), 49(1), 50(2), 53(1), 

93(1), 107(2) 

Node 31 6(1), 45(2), 47(2), 59(2) 

Node 30 81(2), 107(2) 

Node 29 5(3)*, 55(3), 77(3), 83(2), 

86(2), 87(1), 95(2), 97(2) 

Node 28 (Centrodontini) 14(3), 15(3), 29(3), 44(2), 

56(5), 57(2)*, 75(1), 82(1) 

Node 27 68(1)*, 70(1)*, 76(1)*, 

78(2)*, 92(2) 

Node 26 23(2), 24(2)*, 56(2) 

Nodonica 4(2), 40(2), 99(1)* 

Centrodontus 15(1), 49(1), 64(2)*, 65(2), 

101(2) 

Multareis 15(2), 29(2), 56(3)* 

Multareoides 6(1) 

Brachycentrotus 29(3), 56(4)*, 82(1) 

Monobelus 21(1), 83(2), 88(3), 93(2), 

97(2) 

Monobeloides 14(3), 49(1), 81(1), 92(2), 

100(2), 104(1) 

Gargara 3(2), 29(3), 56(5) 

Aleptocentrus 6(1), 48(1) 

Centriculus 29(1), 59(2), 81(1), 92(1) 

Amblycentrus 13(2), 29(3) 

Brachybelus 56(6)*, 65(2) 

Ischnocentrus 102(3) 

Abelus 8(1), 71(2), 101(2) 

Psilocentrus 6(2), 95(2), 112(2) 

Boocerus 55(3), 88(3), 92(1) 

Campylocentrus 11(1), 13(2), 54(1), 86(2), 

106(1), 112(2) 

Ophicentrus 6(1), 15(1), 26(2), 45(2), 

47(2), 71(1), 81(1) 

394

Fig. 24.7. Phylogenetic relationships within Centrotini and Xiphopoeini 

(Analysis 4, PAUP*). Strict consensus of 4 equally parsimonious trees 

(CI=0.37, RI=0.62, length 277). 

395

Fig. 24.8. Phylogenetic relationships within Centrotini and Xiphopoeini 

(Analysis 4, PAUP*). One of 4 equally parsimonious trees with branch 

lengths indicating numbers of assigned character changes. Nodes are 

labeled by number below or above each branch. 

396




Node 68 11(2), 85(2)*, 88(3) 

Node 67 (Centrotini) 56(5)*, 100(2), 114(1) 

Node 66 3(1), 4(2), 48(3), 49(2), 50(1), 

54(2), 55(2), 91(1) 

Node 65 45(2), 47(2)* 

Node 64 13(1), 86(2), 114(2) 

Node 63 90(1) 

Node 62 94(1) 

Node 61 112(1) 

Node 60 89(2) 

Node 59 11(1), 86(1) 

Node 58 13(2), 38(1) 

Node 57 55(1) 

Node 56 101(2) 

Node 55 55(3) 

Node 54 41(2), 97(2) 

Node 53 40(2) 

Node 52 88(1), 90(2), 92(2) 

Node 51 13(2), 15(4), 93(3) 

Node 50 45(1), 114(1) 

Node 49 95(2), 112(2) 

Node 48 26(2), 93(1) 

Node 47 26(2), 55(1) 

Node 46 13(2), 15(4), 16(2), 86(1) 

Node 45 16(2), 55(3) 

Node 44 29(3), 41(2), 112(1) 

Node 43 13(2) 

Node 42 (Xiphopoeini) 7(2)*, 16(2), 24(2)*, 26(2), 

38(1), 45(2), 85(3)*, 86(3), 

112(1) 

Xiphopoeus 13(1), 48(1), 89(2), 91(1), 

92(2), 94(1), 95(2), 97(2) 

Negus 11(1), 12(2), 22(2), 23(2), 

29(3), 50(1), 54(2), 80(2) 

Centrotus 72(2)*, 73(2)*, 101(2), 

115(1), 116(2) 

Capeneralus 40(2) 

Takliwa 93(1), 116(1)* 

Anchonomonoides 88(1) 

Hamma 53(1), 94(1) 

Barsumas 26(2), 41(1), 42(1), 44(2), 

114(1) 

Barsumoides 11(1), 37(1) 

Stalobelus 37(1) 

Monanchon 41(2) 

Mitranotus -- 

Eumocentrulus -- 

Tricoceps 4(1), 29(3) 

Anchon -- 

Anchonastes 16(2), 86(1) 

Paraxiphopoeus 29(1), 77(1), 88(3) 

Monocentrus 26(2), 77(2)*, 94(2), 

Eumonocentrus 11(1), 112(2) 

Zanzia 29(3), 75(1), 95(2) 

Bleccia 81(2) 

Platybelus -- 

Jacobiana 29(3), 112(2) 

Tiberianus 4(1), 15(2), 86(2), 

Capeneriana 13(2) 

Dagonotus -- 

Vecranotus -- 

Farcicaudia 53(1) 

Promitor 13(2), 15(1), 29(3), 81(3), 

89(1), 91(2), 94(2) 

397

Fig. 24.9. Phylogenetic relationships within Choucentrini, Leptocentrini, 

and Maarbarini (Analysis 5, PAUP*). Single most parsimonious tree with 

branch lengths indicating numbers of assigned character changes. Nodes 

are labeled by number below each branch (CI=0.43, RI=0.69, length 207). 

398




Node 57 15(3), 49(2), 105(7)*, 

106(3)*, 107(2)*, 109(1)*, 

110(2), 111(2), 112(2)*, 

114(2) 

Node 56 (Leptocentrini) 40(2), 48(3)*, 77(3), 84(2), 

101(1) 

Node 55 3(2), 52(2), 54(1) 

Node 54 49(1) 

Node 53 53(2), 81(3) 

Node 52 72(3) 

Node 51 49(1) 

Node 50 15(1) 

Node 49 40(1), 73(3)* 

Node 48 48(2) 

Node 47 13(2), 15(3), 53(1) 

Node 46 97(1) 

Node 45 13(2) 

Node 44 10(1), 53(1), 88(3) 

Node 43 32(2), 86(1), 87(2), 88(1), 

91(1), 92(2), 108(2)*, 110(3)* 

Node 42 (Maarbarini) 22(2), 111(1) 

Node 41 27(1), 51(2)*, 52(2) 

Node 40 13(2), 26(2) 

Node 39 11(2), 19(1), 97(1) 

Node 38 92(1), 95(2) 

Node 37 3(2), 26(2) 

Node 36 10(1), 

Node 35 (Choucentrini) 11(2), 31(2)*, 50(1), 55(3), 

60(1), 61(1), 114(1) 

Node 34 45(2)*, 48(2) 

Choucentrus -- 

Evanchon 3(2), 6(1), 27(1), 74(2), 86(2) 

Dograna 13(2), 97(1) 

Telingana 6(1), 29(3), 36(2) 

Pogon 88(3), 92(1) 

Pogontypus 3(2), 29(3), 49(1), 95(2) 

Maarbarus 4(2), 55(3) 

Parapogon 29(3), 50(1) 

Pogonotus 6(1), 49(1) 

Bathoutha -- 

Indicopleustes 19(2) 

Hemicentrus 8(1), 50(1) 

Leptocentrus 29(1), 73(2)*, 81(2) 

Otinotus 13(2), 21(2), 86(1) 

Umfilianus 13(2), 15(2), 38(1) 

Nilautama 10(1), 50(1), 81(2) 

Dacaratha 62(2) 

Awania -- 

Occator 6(1) 

Joveriana 29(1) 

Uroxiphus 40(1), 42(1), 81(1) 

Demanga 49(2) 

Yaponotus 11(2) 

Periaman 12(2), 40(2), 49(2), 84(1), 

100(1), 105(3)*, 110(1), 

111(1) 

Trioxiphus 62(2), 72(2)*, 81(2) 

399

Fig. 24.10. Phylogenetic relationships within Gargarini (Analysis 6, 

PAUP*). Strict consensus of 28 equally parsimonious trees (CI=0.38, 

RI=0.60, length 224). 

400

Fig. 24.11. Phylogenetic relationships within Gargarini (Analysis 6, 

PAUP*). One of 28 equally parsimonious trees with branch lengths 

indicating numbers of assigned character changes. Nodes are labeled by 

number below, above, or to the side of each branch. 

401 

Node 38




Node 60 (Gargarini) 5(3)*, 21(2), 55(3), 88(1) 

Node 59 3(2), 29(3) 

Node 58 48(2), 56(5), 77(3) 

Node 57 83(2)*, 84(2) 

Node 56 15(3), 114(2) 

Node 55 40(2), 88(3), 106(1) 

Node 54 81(3), 95(2) 

Node 53 86(2), 97(2) 

Node 52 16(2)*, 26(2), 53(1), 93(3), 

115(1), 116(1)* 

Node 51 63(2), 64(2)*, 65(2) 

Node 50 3(1), 14(3), 77(1) 

Node 49 45(2), 46(2)* 

Node 48 100(1) 

Node 47 15(3), 29(2), 40(2) 

Node 46 3(1), 100(2) 

Node 45 45(1), 106(1) 

Node 44 81(2), 101(1), 114(2) 

Node 43 40(1), 54(1), 93(1) 

Node 42 3(2) 

Node 41 54(1) 

Node 40 55(1), 83(3)* 

Node 39 86(2), 93(1) 

Node 38 -- 

Node 37 64(3), 106(1), 115(1) 

Node 36 3(1), 14(3), 26(2), 59(2), 

114(2) 

Aleptocentrus 6(1), 42(1), 48(1) 

Yasa 39(2), 93(3), 94(2), 95(2) 

Parayasa 29(2), 42(1), 45(2), 104(1), 

105(3)*, 107(2) 

Maurya 54(1), 92(1) 

Antialcidas 45(2), 77(2), 81(3) 

Machaerotypus 97(2) 

Gargara 101(1) 

Eucoccosterphus 27(1), 42(1), 45(2), 48(1), 

58(2), 84(1), 95(1), 104(1), 

105(4)*, 106(1), 107(2), 

109(1) 

Coccosterphus -- 

Madlinus 27(1) 

Gargarina 29(2) 

Xanthosticta 63(2), 65(2), 77(1), 93(3), 

95(1) 

Kanada 77(1), 81(1) 

Cryptaspidia 29(2), 53(1), 86(1) 

Mesocentrina 48(3), 95(1) 

Sipylus 29(2) 

Tricentrus 15(2), 63(2), 65(2), 97(2) 

Butragulus 55(2), 115(1) 

Tricentroides 86(2), 94(2) 

Cryptoparma 10(2), 53(1), 64(3) 

Thelicentrus 101(1) 

Nondenticentrus 55(2) 

Subrincator 77(1) 

Tribulocentrus 53(1) 

Pantaleon 26(2), 86(2) 

Tsunozemia 55(2), 81(1), 97(2) 

402

Fig. 24.12. Phylogenetic relationships within Hypsaucheniini and 

Oxyrhachini (Analysis 7, PAUP*). Strict consensus of 25 equally 

parsimonious trees (CI=0.55, RI=0.71, length 150). 

403

Fig. 24.13. Phylogenetic relationships within Hypsaucheniini and Oxyrhachini 

(Analysis 7, PAUP*). One of 25 equally parsimonious trees with branch lengths 

indicating numbers of assigned character changes. Nodes are labeled by number 

below each branch. 

404




Node 29 21(2), 23(2), 37(1), 78(2)* 

Node 28 (Oxyrhachini) 12(2), 14(3)*, 15(3), 26(1), 

35(2)*, 40(2), 42(2), 49(1), 

83(3)*, 93(2), 97(2), 112(2)*, 

116(1)* 

Node 27 50(2), 56(5) 

Node 26 76(1) 

Node 25 (Hypsaucheniini) 3(2), 5(3)*, 17(2)*, 43(2), 

50(3)*, 57(1), 76(1) 

Node 24 5(4), 44(2), 48(1), 112(1) 

Node 23 15(5)*, 21(1), 49(1) 

Node 22 29(1), 94(1) 

Node 21 5(2), 10(1), 12(2), 21(2), 

39(2)*, 96(2) 99(2) 

Node 20 6(1)*, 34(2), 43(1), 57(2), 

98(2)* 

Oxyrhachis delalandei 94(1), 102(1) 

Oxyrhachis sulcicornis -- 

Oxyrhachis 15(2), 56(7)* 

Oxyrhachis carinata -- 

Hybandoides 10(1), 27(1), 34(2), 96(2) 

Hypsolyrium -- 

Jingkara 11(2), 34(2), 40(3)*, 48(2), 

93(2) 

Hypsauchenia 97(2) 

Hybanda 44(1), 105(3)* 

Gigantorhabdus 5(4), 10(2), 37(2), 50(2), 

93(2), 116(1) 

Pyrgauchenia 3(1), 101(2), 102(1), 113(3)* 

405

Fig. 24.14. Phylogenetic relationships within Terentiini (Analysis 8, 

PAUP*). Strict consensus of 4 equally parsimonious trees (CI=0.42, 

RI=0.60, length 224). 

406

Fig. 24.15. Phylogenetic relationships within Terentiini (Analysis 8, 

PAUP*). One 4 equally parsimonious trees with branch lengths 

indicating numbers of assigned character changes. Nodes are labeled by 

number below, above, or to the side of each branch. 

407 

Node 33




Node 49 (Terentiini) 3(2)*, 54(2), 70(2), 86(2), 

92(2), 105(6)*, 111(2)* 

Node 48 15(3), 38(2), 102(1), 116(3) 

Node 47 66(2), 114(2) 

Node 46 37(1), 40(3), 95(2) 

Node 45 48(1), 86(1) 

Node 44 24(2), 88(4), 91(2), 92(1) 

Node 43 29(3) 

Node 42 54(1) 

Node 41 37(1) 

Node 40 38(1), 75(1), 82(1), 88(3), 

94(2) 

Node 39 28(2)*, 44(2), 68(1), 70(1), 

74(1), 97(1) 

Node 38 35(2), 54(1), 89(2)*, 90(2), 

91(1), 101(2) 

Node 37 15(1), 27(1), 29(3), 88(4), 

93(1), 115(1), 116(2) 

Node 36 94(2) 

Node 35 40(2), 52(2), 54(1) 

Node 34 34(2), 38(2), 48(2), 101(2) 

Node 33 -- 

Node 32 4(2), 15(5), 17(2), 24(1), 

26(2), 90(2), 110(3)*, 

113(3)*, 114(2) 

Node 31 23(2), 37(2), 74(1) 

Dingkana 95(2) 

Alosextius 53(2), 67(3) 

Sertorius 24(2), 48(1), 77(3), 92(1) 

Terentius 15(1), 29(3), 54(1), 97(1) 

Eutryonia 15(5), 17(2), 40(2), 100(1) 

Bucktoniella 40(2), 53(2), 56(5), 82(1), 

115(1) 

Acanthuchus -- 

Alocanthella -- 

Lubra 40(3), 52(2) 

Anzac -- 

Neosextius 14(3), 15(4), 38(2) 

Goddefroyinella 23(2), 40(3), 48(2), 97(2) 

Sextius 14(2)*, 92(2) 

Bulbauchenia 14(3), 29(1)* 

Funkhouserella 27(1), 44(2), 88(4), 93(3) 

Pyrgonota 6(1), 10(1), 70(1), 97(1) 

Eufrenchia -- 

Cebes 52(1), 70(1), 88(4) 

Ceraon 26(2), 40(3) 

Sarantus 56(2)*, 90(2), 94(1) 

408

Fig. 24.16. Phenetic relationships within Lobocentrini, Beaufortianini, 

Gargarini, and Centrotini (Analysis 9, UPGMA). Branch lengths 

indicate mean character distance. Asterisks indicate genera not included 

in the phylogenetic analyses because of limited data. 

409

Fig. 24.17. Phenetic relationships within Nessorhinini, Leptocentrini, and 

Terentiini (Analysis 10, UPGMA). Branch lengths indicate mean 

character distance. Asterisks indicate genera not included in the 

phylogenetic analyses because of limited data. 

410

Table 24.13. Data matrix. 

1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 3 

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 

Abelus 1 1 1 1 2 1 1 1 ? ? ? ? ? ? 1 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Acanthophyes decens 1 1 1 1 2 2 1 2 1 2 1 1 2 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Acanthucalis macalpini 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 2 1 1 1 2 1 2 1 2 1 

Acanthuchus trispinifer 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 2 1 1 2 2 1 2 1 3 1 

Afraceronotus quinquefasciatus 1 1 1 2 2 2 1 2 1 2 1 1 1 1 1 1 1 2 2 1 1 ? 1 1 2 1 2 1 2 1 

Aleptocentrus notabilis 1 1 1 1 3 1 1 2 1 1 1 2 2 1 1 ? 1 2 2 1 ? 1 ? ? 2 ? 2 1 2 1 

Alocanthella fulva 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 2 1 1 2 2 1 2 1 3 1 

Alocebes dixoni 1 1 2 1 2 2 1 2 ? 2 1 2 2 1 3 ? 1 2 ? ? ? 1 ? ? 2 1 2 1 2 1 

Alosextius carinatus 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 2 1 1 1 2 1 2 1 2 1 

Amblycentrus pubescens 1 1 1 2 2 2 1 2 1 1 1 2 2 1 1 1 1 2 2 1 1 1 1 1 2 1 2 1 3 1 

Amphilobocentrus bifasciatus 1 1 ? 1 2 2 1 2 1 1 2 1 ? 1 3 ? 1 2 2 1 1 2 ? ? 2 ? 2 1 2 ? 

Ananthasubramanian tomentosus 1 1 2 1 2 2 1 2 1 2 1 1 ? 1 3 ? 1 2 2 1 1 ? ? ? 2 1 2 1 2 1 

Anchon 1 1 1 2 2 2 1 2 1 2 2 1 1 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Anchonastes hastatus 1 1 1 2 2 2 1 2 1 2 2 1 1 1 3 2 1 2 2 1 1 1 1 1 2 2 2 1 2 1 

Anchonobelus aries 1 1 1 2 2 2 1 2 1 2 2 1 1 1 3 2 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Anchonomonoides expansus 1 1 1 2 2 2 1 2 1 2 2 1 1 1 3 2 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Antialcidas 1 1 2 1 3 2 1 2 1 1 1 2 2 1 3 1 1 2 2 1 2 1 1 1 2 1 2 1 ? 1 

Anzac bipunctatum 1 1 2 1 2 2 1 2 1 2 1 2 2 1 1 1 1 2 2 1 2 1 1 2 2 1 1 2 3 1 

Arcuatocornum sp. 1 1 2 1 2 2 1 2 1 1 2 1 2 1 3 1 1 2 2 1 1 2 1 1 2 2 2 1 2 1 

Arimanes doryensis 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 2 1 1 1 2 2 2 1 2 1 

Aurinotus auricornis 1 1 1 1 2 2 1 2 1 2 1 1 2 1 3 1 1 2 2 1 1 1 1 1 2 2 2 1 2 1 

Awania 1 1 2 1 2 2 1 2 1 1 1 1 1 1 1 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Barsumas 1 1 1 2 2 2 1 2 1 2 2 1 2 1 3 2 1 2 2 1 1 1 1 1 2 2 ? 1 3 1 

Barsumoides 1 1 1 2 2 2 1 2 1 2 1 1 2 1 3 2 1 2 2 1 1 1 1 1 2 1 2 1 3 1 

Bathoutha indicans 1 1 2 1 2 2 1 2 1 1 2 1 1 1 3 1 1 2 1 ? 1 2 1 1 2 2 1 1 ? 1 

Beaufortiana 1 1 2 1 2 2 1 2 1 1 1 1 2 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Bleccia fastidiosus 1 1 1 2 2 2 1 2 1 2 1 1 1 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Bocchar 1 1 2 1 2 2 1 2 1 2 1 1 1 1 2 1 1 2 2 1 1 2 1 1 2 1 2 1 2 1 

Boocerus gilvipes 1 1 1 1 2 2 1 2 1 2 2 1 1 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Brachybelus 1 1 1 2 2 2 1 2 1 1 1 2 1 1 1 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Brachycentrotus 1 1 1 2 2 ? 1 2 1 1 1 2 2 1 1 1 1 2 2 1 2 1 1 1 2 1 2 1 3 1 

Bucktoniella pyramidatus 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 2 1 1 2 2 1 2 1 3 1 

Bulbauchenia sp. 1 1 2 2 2 2 1 2 1 2 1 2 ? 3 5 1 2 2 2 ? 2 1 1 1 2 2 2 1 1 1 

Bulbauchenia bakeri 1 1 2 2 2 2 1 2 ? 2 1 2 ? 3 3 1 1 2 2 1 2 ? 1 1 2 1 2 1 2 1 

Bunyella dromedarius 1 1 2 1 2 2 1 2 ? 2 1 2 ? 1 3 ? 1 2 ? ? ? 1 ? ? 2 ? ? 1 ? 1 

Butragulus flavipes 1 1 2 1 3 2 1 2 1 1 1 2 2 1 3 1 1 2 2 1 2 1 1 1 2 1 2 1 2 1 

Callicentrus ignipes 1 1 1 1 2 2 1 2 1 2 1 2 ? 3 3 1 1 2 ? ? 2 1 1 1 2 2 2 1 2 1 

Camelocentrus yunnanensis 1 1 2 1 2 2 1 2 1 2 2 1 ? 1 3 ? 1 2 ? ? ? ? ? ? 2 ? 2 1 3 1 

Campylocentrus hamifer 1 1 2 1 2 2 1 2 1 2 1 1 2 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Campylocentrus obscuripennis 1 1 2 1 2 2 1 2 1 2 1 1 1 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 ? 1 

Capeneralus 1 1 1 2 2 2 1 2 1 ? 2 1 2 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Capeneriana tenuicornis 1 1 1 2 2 2 1 2 1 2 1 1 2 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Cebes transiens 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 2 1 1 2 2 1 2 1 2 1 

Centriculus 1 1 1 1 2 2 1 2 1 1 1 2 2 1 1 1 1 2 2 1 1 1 1 1 2 1 2 1 1 1 

Centrochares horrifica 1 1 2 1 1 2 1 2 1 2 2 1 2 1 3 2 1 2 2 1 1 1 1 2 2 2 2 1 2 1 

Centrodontus 1 1 1 1 2 2 1 2 1 ? 1 2 ? 3 1 1 1 2 ? ? 2 1 1 1 1 2 1 1 3 ? 

Centrolobus africanus 1 1 2 1 2 2 1 2 1 ? 2 1 1 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 ? 1 

Centronodus 1 2 1 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 1 1 2 1 1 2 1 1 2 3 

Centrotosuoides muiri 1 1 2 1 3 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 1 1 1 2 2 1 2 1 3 1 

Centrotus cornutus 1 1 2 1 2 2 1 2 1 2 2 1 2 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Centrotypus 1 1 1 1 2 2 1 2 1 2 1 2 ? 2 3 1 1 2 2 2 1 1 1 1 2 2 2 1 2 1 

Centruchus 1 1 2 1 3 2 1 2 1 2 1 1 2 1 3 1 1 2 2 1 1 1 1 2 2 1 2 1 2 1 

Ceraon tasmaniae 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 2 1 1 2 2 2 2 1 2 1 

Choucentrus 1 1 ? 1 2 2 1 2 1 2 2 1 ? 1 3 ? 1 2 2 1 ? ? ? ? 2 ? 2 1 2 1 

Coccosterphus 1 1 2 1 3 2 1 2 1 1 1 2 2 1 1 2 1 2 2 1 2 1 1 1 2 2 2 1 3 1 

Cornutobelus mutabilis 1 1 1 2 2 2 1 2 1 2 1 1 2 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Crito festivum 1 1 2 1 2 2 1 2 1 2 1 2 ? 1 3 ? 1 2 ? ? ? 1 ? ? 2 ? 2 1 2 1 

Cryptaspidia pubera 1 1 1 1 3 2 1 2 1 1 1 2 ? 3 1 1 1 2 2 1 2 1 1 1 2 2 2 1 2 1 

Cryptoparma parva 1 1 1 1 3 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 2 1 1 1 2 1 2 1 2 1 

Dacaratha 1 1 2 1 2 2 1 2 1 2 1 1 1 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Daconotus projectus 1 1 1 2 2 2 1 2 1 2 1 1 2 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 3 1 

Dagonotus lectus 1 1 1 2 2 2 1 2 1 1 1 1 1 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Daimon serricorne 1 1 1 2 2 2 1 2 1 2 1 2 2 1 3 2 1 2 2 1 2 1 1 1 2 2 2 1 2 1 

Demanga sookana 1 1 2 1 2 2 1 2 ? ? 1 1 1 1 1 1 1 2 ? ? 1 ? 1 1 2 1 2 1 ? 1 

Dingkana borealis 1 1 2 1 2 2 1 2 1 2 1 2 2 1 1 1 1 2 2 1 2 1 1 1 2 1 2 1 2 1 

Distanobelus sericeus 1 1 1 2 2 2 1 2 1 2 1 1 2 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Dograna suffulta 1 1 1 1 2 2 1 2 1 2 2 1 2 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Dukeobelus simplex 1 1 2 1 2 2 1 2 1 1 1 1 2 1 1 1 1 2 2 1 1 1 1 1 2 2 2 1 2 1 

Ebhul varium 1 1 1 1 2 2 1 2 1 ? 2 1 2 1 1 1 1 2 2 1 1 1 1 2 2 2 2 1 2 1 

Elaphiceps 1 1 1 2 2 2 1 2 2 2 1 1 1 1 5 1 2 2 2 1 1 2 1 1 2 2 2 1 2 1 

Emphusis obesa 1 1 1 1 2 2 1 2 1 2 1 1 2 1 3 1 1 2 2 2 1 1 1 1 2 2 2 1 2 1 

411

Table 24.13 cont’d. Data matrix. 

3 3 3 3 3 3 3 3 3 4 4 4 4 4 4 4 4 4 4 5 5 5 5 5 5 5 5 5 5 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 

Abelus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 1 1 1 1 2 2 

Acanthophyes decens 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 3 5 1 1 1 2 

Acanthucalis macalpini 1 1 1 1 1 1 2 2 1 1 1 2 1 1 ? ? 1 2 1 2 1 1 2 2 1 1 1 1 1 1 

Acanthuchus trispinifer 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 1 1 2 1 1 1 1 1 1 1 1 1 1 

Afraceronotus quinquefasciatus 1 1 1 1 1 1 2 2 1 2 1 1 1 1 2 1 2 ? 2 1 1 1 2 2 3 5 1 1 1 2 

Aleptocentrus notabilis 1 1 1 1 ? 1 2 2 1 1 1 1 1 1 1 ? 1 1 2 1 1 1 ? 2 3 1 ? ? ? ? 

Alocanthella fulva 1 1 1 1 1 1 2 2 1 1 1 1 1 1 1 ? 1 1 1 2 1 1 1 1 1 1 1 1 1 1 

Alocebes dixoni 1 1 1 1 ? 1 1 2 1 3 1 2 1 1 1 ? 1 1 1 1 1 ? 2 1 1 1 ? ? ? ? 

Alosextius carinatus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 2 1 2 1 1 2 2 1 1 1 1 1 1 

Amblycentrus pubescens 1 1 1 1 1 1 2 2 2 1 1 2 1 1 1 ? 1 3 2 1 1 1 2 2 1 5 1 2 1 2 

Amphilobocentrus bifasciatus 1 1 1 1 1 1 2 2 1 ? 1 2 1 1 1 ? 1 3 1 1 1 1 2 1 1 1 ? ? ? ? 

Ananthasubramanium tomentosum 1 1 1 1 1 1 2 2 1 ? 1 2 1 1 1 ? 1 3 2 1 1 1 2 2 1 1 ? ? ? ? 

Anchon 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 2 5 1 1 1 2 

Anchonastes hastatus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 2 5 1 1 1 2 

Anchonobelus aries 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 3 5 1 1 2 2 

Anchonomonoides expansus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 3 5 1 1 1 2 

Antialcidas 1 1 1 1 1 1 2 2 1 2 1 2 1 1 2 1 1 2 2 1 1 1 2 2 3 5 1 1 1 2 

Anzac bipunctatum 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 ? 1 1 1 2 1 1 1 2 1 1 2 1 1 1 

Arcuatocornum sp. 1 1 1 1 1 1 2 2 1 2 1 2 1 1 1 ? 1 3 1 2 1 1 1 1 1 1 1 1 1 2 

Arimanes doryensis 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 1 1 2 1 1 1 2 1 1 1 1 1 1 

Aurinotus auricornis 1 1 1 1 1 1 2 1 1 2 1 2 1 1 2 1 2 ? 1 1 1 1 2 2 1 5 1 1 1 2 

Awania 1 1 1 1 1 1 2 2 1 2 1 2 1 1 1 ? 1 3 1 2 1 2 1 1 1 1 1 1 1 2 

Barsumas 1 1 1 1 1 1 2 ? 1 1 1 1 1 2 2 1 2 ? 2 1 1 1 2 2 3 5 1 1 1 2 

Barsumoides 1 ? 1 1 1 1 1 ? 1 1 2 2 1 1 2 1 2 ? 2 1 1 1 2 2 3 5 1 1 1 2 

Bathoutha indicans 1 2 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 1 2 2 2 2 1 2 1 1 1 1 1 2 

Beaufortiana 1 1 1 1 1 1 1 2 1 1 1 2 1 1 1 ? 1 2 1 2 1 1 1 1 1 1 1 1 1 ? 

Bleccia fastidiosus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 2 5 1 1 1 2 

Bocchar 1 1 1 1 1 1 1 2 1 1 1 2 1 1 1 ? 1 2 1 1 1 1 2 2 3 1 1 1 1 1 

Boocerus gilvipes 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 3 2 1 1 1 2 2 3 1 1 1 1 2 

Brachybelus 1 1 1 1 1 1 2 2 2 1 1 2 1 1 1 ? 1 3 2 1 1 1 2 2 1 6 1 2 1 2 

Brachycentrotus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 2 2 1 1 1 2 2 1 4 1 1 1 1 

Bucktoniella pyramidatus 1 1 1 1 1 1 ? 2 1 2 1 1 1 1 1 ? 1 1 1 2 1 1 2 2 1 5 1 1 1 1 

Bulbauchenia sp. 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 ? 1 1 1 2 1 1 1 2 1 1 2 1 1 1 

Bulbauchenia bakeri 1 1 1 1 1 1 1 1 1 1 1 2 1 1 ? ? 1 1 1 2 1 ? ? 2 2 1 1 1 1 1 

Bunyella dromedarius 1 1 1 1 ? 1 2 2 1 2 1 2 1 1 1 ? 1 ? 1 2 1 1 2 2 1 1 ? ? ? ? 

Butragulus flavipes 1 1 1 1 1 1 2 2 1 2 1 2 1 1 2 2 1 2 2 1 1 1 2 2 2 5 1 1 1 2 

Callicentrus ignipes 1 1 1 1 1 1 1 2 1 2 1 2 1 1 1 ? 1 1 1 2 1 2 2 1 1 1 1 1 1 1 

Camelocentrus yunnanensis 1 1 1 1 1 1 2 2 1 ? 1 2 1 1 1 ? 1 3 2 1 1 1 2 2 1 1 ? ? ? ? 

Campylocentrus hamifer 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 3 1 2 1 1 1 1 1 1 1 1 1 2 

Campylocentrus obscuripennis 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 2 2 1 1 1 1 2 3 1 1 1 1 2 

Capeneralus 1 1 1 1 1 1 2 2 1 2 1 2 1 1 1 ? 1 3 2 1 1 1 2 2 2 5 1 1 1 2 

Capeneriana tenuicornis 1 1 1 1 1 1 2 2 1 2 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 2 5 1 1 ? 2 

Cebes transiens 1 ? 1 2 1 1 1 2 1 2 1 1 1 1 1 ? 1 ? 1 2 1 1 1 1 1 1 2 1 1 1 

Centriculus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 3 2 1 1 1 2 2 1 1 1 1 2 2 

Centrochares horrifica 1 ? 1 ? 1 1 2 2 1 1 2 1 1 1 2 1 2 ? 2 1 1 1 ? ? ? 3 2 1 1 1 

Centrodontus 1 ? ? ? 1 1 2 2 1 1 1 1 1 2 1 ? 1 ? 1 ? 1 1 ? ? ? 5 2 1 1 1 

Centrolobus africanus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 2 1 2 1 1 1 1 1 1 1 1 1 1 

Centronodus 1 ? 2 ? 1 1 2 2 1 1 1 1 1 2 1 ? 1 ? ? ? 1 1 ? ? ? 1 1 1 1 2 

Centrotosuoides 1 1 1 1 1 1 2 1 1 1 1 1 1 1 1 ? 1 2 2 1 1 1 2 2 1 1 1 1 1 1 

Centrotus cornutus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 2 1 2 1 1 2 1 1 5 1 1 1 2 

Centrotypus 1 2 1 1 1 1 ? 2 1 3 1 2 1 1 1 ? 1 2 2 2 1 1 2 2 1 1 1 1 1 2 

Centruchus 1 1 1 1 1 1 2 1 1 1 1 2 1 1 1 ? 1 2 1 2 1 1 2 1 1 1 1 1 1 1 

Ceraon tasmaniae 1 1 1 2 1 1 1 2 1 3 1 1 1 1 1 ? 1 2 1 2 1 2 1 1 1 1 1 1 1 1 

Choucentrus 2 ? 1 1 1 1 2 2 1 ? 1 2 1 1 1 ? 1 1 2 1 1 1 ? ? 3 1 ? ? ? ? 

Coccosterphus 1 ? 1 1 1 1 2 2 1 1 ? 2 1 1 1 ? 1 2 2 1 1 1 1 2 3 5 1 1 1 2 

Cornutobelus mutabilis 1 1 1 1 1 1 2 2 1 2 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 2 5 1 1 1 2 

Crito festivum 1 1 1 1 1 1 1 2 1 3 1 2 1 1 1 ? 1 1 1 2 1 1 1 2 1 1 ? ? ? ? 

Cryptaspidia pubera 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 2 1 2 2 1 1 1 1 2 3 5 1 1 2 2 

Cryptoparma parva 1 1 1 1 1 1 2 2 1 2 1 2 1 1 2 2 1 2 2 1 1 1 1 1 1 5 1 1 1 2 

Dacaratha 1 1 1 1 1 1 2 2 1 2 1 2 1 1 1 ? 1 3 1 2 1 2 2 1 1 1 1 1 1 2 

Daconotus projectus 1 1 1 1 1 1 1 1 1 1 1 2 1 1 2 1 2 ? 1 1 1 1 2 2 2 5 1 1 1 2 

Dagonotus lectus 1 1 1 1 1 1 2 2 1 2 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 2 5 1 1 1 2 

Daimon serricorne 1 1 1 1 1 1 1 2 1 2 1 2 1 1 2 1 1 2 1 2 1 2 1 2 2 1 1 1 1 1 

Demanga sooknana 1 1 1 1 1 1 2 2 1 1 1 2 1 1 ? ? 1 3 2 ? 1 ? ? ? ? 1 1 1 1 2 

Dingkana borealis 1 1 1 1 1 1 2 1 1 1 1 2 1 1 1 ? 1 2 1 2 1 1 1 2 1 1 1 1 1 1 

Distanobelus sericeus 1 1 1 1 1 1 2 1 1 1 1 2 1 1 2 1 2 ? 1 1 1 1 2 2 2 5 1 1 1 2 

Dograna suffulta 2 ? 1 1 1 1 2 2 1 1 1 2 1 1 2 1 1 2 2 1 1 1 ? ? ? 1 1 1 1 1 

Dukeobelus simplex 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 2 1 2 1 1 1 1 1 1 1 1 1 1 

Ebhul varium 1 1 1 2 1 1 2 2 1 1 1 1 1 1 1 ? 1 2 2 1 1 1 1 1 1 5 2 1 1 1 

Elaphiceps 1 2 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 2 1 1 1 1 1 2 2 1 1 1 1 1 

Emphusis obesa 1 2 1 1 1 1 1 2 1 2 1 2 1 1 1 ? 1 2 2 2 1 1 1 1 1 1 1 1 1 2 

412


6 6 6 6 6 6 6 6 6 7 7 7 7 7 7 7 7 7 7 8 8 8 8 8 8 8 8 8 8 9 

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 

Abelus 2 1 1 1 1 1 ? 2 1 2 2 1 1 3 2 2 1 1 2 1 2 2 1 1 ? 1 2 1 ? ? 

Acanthophyes decens 2 1 1 1 1 1 ? 2 1 2 1 ? 1 3 2 2 3 1 2 1 1 2 1 2 2 2 1 3 2 1 

Acanthucalis macalpini 1 ? 1 1 1 1 ? 2 1 2 1 ? 1 2 2 2 1 1 2 1 1 ? ? ? ? ? 1 3 1 2 

Acanthuchus trispinifer 1 ? 1 1 1 1 ? 2 1 2 1 1 1 2 2 2 1 1 2 1 1 2 1 1 ? 1 1 4 1 1 

Afraceronotus quinquefasciatus 2 1 1 1 1 1 ? 2 1 2 1 1 ? 3 2 2 3 1 2 ? ? 2 ? ? ? 2 1 ? ? ? 

Aleptocentrus notabilis ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Alocanthella fulva 1 ? 1 1 1 1 ? 2 1 2 1 1 ? 2 2 2 1 1 2 1 1 ? 1 1 ? 1 1 ? ? ? 

Alocebes dixoni ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Alosextius carinatus 1 ? 1 1 1 2 3 2 1 2 1 ? 1 2 2 2 1 1 2 1 1 ? 1 1 ? ? 1 1 ? ? 

Amblycentrus pubescens 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 1 1 2 1 3 2 1 1 ? 1 2 3 2 2 

Amphilobocentrus bifasciatus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Ananthasubramanium tomentosus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Anchon 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 2 2 1 1 ? ? 

Anchonastes hastatus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 2 1 1 1 ? ? 

Anchonobelus aries 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 2 2 1 3 2 2 

Anchonomonoides expansus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 2 2 1 1 ? ? 

Antialcidas 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 2 1 2 1 3 2 2 2 1 1 1 3 2 2 

Anzac bipunctatum 1 ? 1 1 1 1 ? 1 ? 1 ? 1 1 1 1 2 1 1 2 1 1 ? 1 1 ? 1 1 4 1 1 

Arcuatocornum sp. 2 2 1 1 1 1 ? 2 1 2 2 1 1 2 2 2 3 1 2 1 2 2 1 1 ? 1 1 1 ? ? 

Arimanes doryensis 1 ? 1 1 1 1 ? 2 1 2 1 ? 1 2 2 2 1 1 2 1 1 2 1 1 ? ? 1 4 ? ? 

Aurinotus auricornis 2 1 1 1 1 1 ? 2 1 2 1 1 ? 3 2 2 3 1 2 ? ? 2 ? ? ? 2 1 ? ? ? 

Awania 2 1 1 1 1 1 ? 2 1 2 1 3 1 3 2 2 3 1 2 1 3 2 1 2 1 2 1 3 1 1 

Barsumas 2 1 1 1 1 1 ? 2 1 2 1 ? 1 3 2 2 3 1 2 1 1 2 1 2 2 2 1 3 1 2 

Barsumoides 2 1 1 1 1 1 ? 2 1 2 1 ? 1 3 2 2 3 1 2 1 1 2 1 2 ? 2 1 3 1 2 

Bathoutha indicans 2 1 1 1 1 1 ? 2 1 2 1 1 ? ? 2 2 1 1 2 1 1 2 1 1 ? 1 2 ? ? ? 

Beaufortiana 1 ? 1 1 1 1 ? 2 1 ? ? 1 1 3 2 2 1 1 2 1 1 2 2 1 ? 1 2 1 ? ? 

Bleccia fastidiosus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 2 2 1 2 2 1 1 3 2 1 

Bocchar 1 ? 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 1 1 2 1 1 2 1 1 ? 1 2 4 1 1 

Boocerus gilvipes 2 1 1 1 1 1 ? 2 1 2 2 1 1 3 2 2 1 1 2 1 3 2 1 1 ? 1 2 3 2 2 

Brachybelus 2 1 1 1 2 1 ? 2 1 2 1 1 1 3 2 2 1 1 2 1 3 2 1 1 ? 1 2 3 2 2 

Brachycentrotus 1 ? 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 1 1 2 1 3 1 1 1 ? 1 2 1 ? ? 

Bucktoniella pyramidatus 1 ? 1 1 1 1 ? 2 1 2 1 1 ? 2 2 2 1 1 2 1 1 1 1 1 ? 1 1 ? ? ? 

Bulbauchenia sp. 1 ? 1 1 1 1 ? 2 1 2 1 1 ? 2 1 2 1 1 2 1 1 ? 1 1 ? 1 1 ? ? ? 

Bulbauchenia bakeri 1 ? 1 1 1 1 ? 2 1 2 1 1 1 ? 1 2 1 1 2 1 1 ? 1 1 ? 1 1 3 2 2 

Bunyella dromedarius ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 2 ? 1 ? ? ? ? ? ? ? ? ? ? ? ? 

Butragulus flavipes 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 3 2 2 2 1 1 1 1 ? ? 

Callicentrus ignipes 1 ? 1 1 1 1 ? 2 1 2 1 1 1 3 ? 2 1 1 2 1 3 2 2 1 ? 2 2 1 ? ? 

Camelocentrus yunnanensis ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Campylocentrus hamifer 2 2 1 1 1 1 ? 2 1 2 2 1 1 ? 2 2 1 1 2 1 3 2 1 1 ? 2 2 1 ? ? 

Campylocentrus obscuripennis 2 2 1 1 1 1 ? 2 1 2 2 1 ? 3 2 2 1 1 2 1 1 2 1 1 ? 2 2 ? ? ? 

Capeneralus 2 1 1 1 1 1 ? 2 1 2 1 ? 1 3 2 2 3 1 2 1 1 2 1 2 2 1 1 3 1 2 

Capeneriana tenuicornis 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 ? 1 1 ? ? ? 

Cebes transiens 1 ? 1 1 1 1 ? 2 1 1 ? 1 1 2 1 2 1 1 2 1 1 ? 1 1 ? ? 1 4 1 1 

Centriculus 2 1 1 1 1 1 ? 2 1 2 1 ? 1 3 2 2 1 1 2 1 1 2 1 1 ? 1 2 3 1 2 

Centrochares horrifica 1 ? ? 2 2 1 ? 2 1 1 ? 1 1 3 1 2 3 1 2 2 1 1 2 1 ? 2 1 3 2 2 

Centrodontus 1 ? 2 2 2 1 ? 1 ? 1 ? 1 1 2 1 1 ? 2 2 1 1 1 1 1 ? 1 1 1 ? ? 

Centrolobus africanus 1 ? 1 1 1 1 ? 2 1 2 1 ? 1 3 2 2 1 1 2 1 1 2 2 1 ? 1 2 1 ? ? 

Centronodus 2 1 1 1 1 2 2 2 1 2 1 1 1 3 2 2 1 1 2 1 1 ? 1 1 ? 1 1 1 ? ? 

Centrotosuoides muiri 1 ? 1 1 1 2 2 2 1 2 2 ? 1 3 2 2 1 1 2 1 1 2 2 1 ? 2 2 1 ? ? 

Centrotus cornutus 2 1 1 1 1 1 ? 2 1 2 1 2 2 3 2 2 3 1 2 1 1 2 1 2 2 1 1 3 1 2 

Centrotypus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 3 2 1 1 ? 2 1 1 ? ? 

Centruchus 1 ? 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 1 1 2 1 1 2 2 1 ? 1 2 1 ? ? 

Ceraon tasmaniae 1 ? 1 1 1 1 ? 2 1 2 1 1 1 2 1 2 1 1 2 1 1 1 1 1 ? 1 1 3 1 1 

Choucentrus ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 2 ? 1 ? ? ? ? ? ? ? ? ? ? ? ? 

Coccosterphus 2 1 2 2 2 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 3 2 2 2 1 2 1 1 ? ? 

Cornutobelus mutabilis 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 2 1 1 3 1 2 

Crito festivum ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 2 ? 1 ? ? ? ? ? ? ? ? ? ? ? ? 

Cryptaspidia pubera 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 3 2 2 2 1 1 1 1 ? ? 

Cryptoparma parva 2 1 1 3 1 1 ? 2 1 2 1 3 ? 3 2 2 3 1 2 1 3 2 3 2 1 1 1 ? ? ? 

Dacaratha 2 2 1 1 1 1 ? 2 1 2 1 ? 1 3 2 2 3 1 2 1 3 2 1 2 1 2 1 4 1 1 

Daconotus projectus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 2 1 1 3 1 2 

Dagonotus lectus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 2 1 1 ? ? ? 

Daimon serricorne 1 ? 1 1 1 1 ? 2 1 2 1 ? 1 3 1 2 1 1 2 1 1 2 2 1 ? 3 2 1 ? ? 

Demanga sooknana 2 1 1 1 1 1 ? 2 1 2 1 3 ? 3 2 2 3 1 2 ? ? ? ? ? ? 2 ? ? ? ? 

Dingkana borealis 1 ? 1 1 1 1 ? 2 1 2 1 1 3 2 2 2 1 1 2 1 2 2 1 1 ? 2 1 1 ? ? 

Distanobelus sericeus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 2 2 1 1 ? ? 

Dograna suffulta 1 ? 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 1 1 2 1 1 2 1 1 ? 1 2 1 ? ? 

Dukeobelus simplex 1 ? 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 1 1 2 1 1 2 2 1 ? 1 2 1 ? ? 

Ebhul varium 1 ? 1 1 1 1 ? 1 ? 1 ? 1 1 2 1 ? ? 1 2 1 1 1 1 1 ? 1 1 2 ? ? 

Elaphiceps 1 ? 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 1 1 2 1 2 2 1 1 ? 2 2 3 2 1 

Emphusis obesa 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 3 2 1 1 ? 2 1 1 ? ? 

413


1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 

9 9 9 9 9 9 9 9 9 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 

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 

Abelus 1 2 2 1 1 1 1 1 3 2 2 1 3 2 5 3 2 1 1 1 1 1 2 1 1 3 

Acanthophyes decens 2 1 2 1 1 1 1 1 3 2 1 1 1 ? 3 3 2 1 2 1 1 1 2 2 2 3 

Acanthucalis macalpini 2 1 1 ? 1 1 2 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Acanthuchus trispinifer 2 1 2 1 1 1 2 1 3 2 1 1 1 1 6 2 2 1 1 1 2 3 2 1 2 3 

Afraceronotus quinquefasciatus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Aleptocentrus notabilis ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Alocanthella fulva ? ? ? ? ? ? ? 1 3 2 1 1 1 1 6 2 2 1 1 1 2 3 2 ? ? ? 

Alocebes dixoni ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Alosextius carinatus 1 2 2 1 1 1 2 ? ? ? ? ? ? ? 6 2 2 1 1 1 2 3 2 ? ? ? 

Amblycentrus pubescens 1 2 2 1 2 1 1 1 3 2 1 1 3 2 1 3 1 1 1 1 1 1 2 1 1 3 

Amphilobocentrus bifasciatus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Ananthasubramanium tomentosus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Anchon 1 2 2 1 2 1 1 1 3 2 1 1 1 1 3 3 2 1 2 1 1 2 2 2 2 3 

Anchonastes hastatus 1 2 1 ? 2 1 1 1 3 2 1 1 1 1 3 3 2 1 2 1 1 2 2 2 2 3 

Anchonobelus aries 1 1 2 1 1 1 1 1 3 2 2 1 1 1 3 3 2 1 2 1 1 1 2 2 2 3 

Anchonomonoides expansus 1 1 2 2 1 1 1 1 3 2 1 1 1 1 3 3 2 1 2 1 1 2 2 ? ? ? 

Antialcidas 1 2 2 1 1 1 1 1 3 2 2 1 1 2 1 1 1 1 3 1 1 1 2 ? ? ? 

Anzac bipunctatum 2 1 1 ? 1 1 1 1 3 2 1 1 1 ? 6 2 2 1 1 1 2 3 2 1 1 2 

Arcuatocornum sp. 1 1 1 ? 2 1 2 1 3 2 1 1 1 2 1 1 1 1 3 1 1 1 2 ? ? ? 

Arimanes doryensis 2 1 ? ? 1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Aurinotus auricornis ? ? ? ? ? ? ? 1 3 2 1 1 1 ? 3 3 2 1 2 1 1 ? 2 ? ? ? 

Awania 2 1 1 ? 1 1 1 1 3 2 1 1 1 1 7 3 2 1 ? 2 2 ? 2 2 2 3 

Barsumas 1 1 2 2 1 1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 1 2 3 

Barsumoides 1 1 2 2 1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 2 2 3 

Bathoutha indicans ? ? ? ? ? ? ? 1 3 2 2 1 1 1 7 3 2 2 1 3 1 2 2 ? ? ? 

Beaufortiana 1 1 3 1 2 1 1 1 3 2 2 1 1 1 4 1 2 1 1 1 1 1 2 1 1 2 

Bleccia fastidiosus 1 1 2 1 1 1 1 1 3 2 1 1 1 1 3 3 2 1 2 1 1 1 2 ? ? ? 

Bocchar 2 1 2 2 1 1 1 1 3 1 ? ? 1 1 8 3 2 1 2 1 2 1 2 ? ? ? 

Boocerus 1 1 2 2 1 1 1 1 3 2 1 1 3 2 5 3 1 1 1 1 1 3 2 1 1 3 

Brachybelus 1 2 2 1 2 1 1 1 3 2 1 1 3 2 1 3 1 1 1 1 1 1 2 ? ? ? 

Brachycentrotus 1 1 3 1 1 1 1 1 3 1 ? ? 1 2 2 1 1 1 3 1 1 1 2 ? ? ? 

Bucktoniella pyramidatus ? ? ? ? ? ? ? 1 3 2 1 1 1 1 6 2 2 1 1 1 2 3 2 1 1 3 

Bulbauchenia ? ? ? ? ? ? ? 1 3 2 1 1 1 1 6 2 2 1 1 3 2 3 3 2 2 3 

Bulbauchenia bakeri 1 1 2 2 1 1 1 1 3 2 1 1 1 1 6 2 2 1 1 3 2 3 3 2 2 3 

Bunyella dromedarius ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Butragulus flavipes 1 2 2 1 2 1 1 1 3 1 ? ? 1 2 1 3 1 1 3 1 1 1 2 1 1 3 

Callicentrus ignipes 1 2 3 2 1 1 1 1 3 2 1 1 1 1 4 1 2 1 3 1 1 2 2 ? ? ? 

Camelocentrus yunnanensis ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Campylocentrus hamifer 1 2 1 ? 1 1 1 1 3 2 1 1 3 2 5 1 2 1 1 1 1 2 2 ? ? ? 

Campylocentrus obscuripennis ? ? ? ? ? ? ? 1 3 2 1 1 3 2 5 3 2 1 1 1 1 3 2 ? ? ? 

Capeneralus 1 1 2 2 1 1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? 3 

Capeneriana tenuicornis ? ? ? ? ? ? ? 1 3 2 2 1 1 1 3 3 2 1 2 1 1 1 2 2 2 3 

Cebes transiens 2 1 2 2 1 1 2 1 3 2 2 1 1 1 6 2 2 1 1 1 2 3 2 ? ? ? 

Centriculus 1 1 2 1 1 1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 1 1 3 

Centrochares horrifica 1 1 2 2 1 2 1 1 3 1 ? ? 1 1 7 1 2 1 3 2 1 1 2 1 1 3 

Centrodontus 1 2 2 1 1 1 1 1 3 2 2 2 1 1 1 1 1 1 3 1 1 1 2 1 2 3 

Centrolobus africanus 1 1 2 2 2 1 1 ? 3 2 2 1 1 1 4 1 2 1 1 1 1 1 2 ? ? ? 

Centronodus 1 2 2 2 1 1 1 1 2 2 2 1 1 1 1 1 1 1 3 1 1 1 1 ? ? ? 

Centrotosuoides muiri 2 1 2 2 1 1 2 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Centrotus cornutus 2 1 2 2 1 1 1 1 3 2 2 1 1 1 3 3 2 1 2 1 1 2 2 1 1 2 

Centrotypus 1 2 2 2 2 1 2 1 3 1 ? ? 1 1 3 3 2 1 2 1 1 2 2 2 2 3 

Centruchus 2 1 2 2 2 1 2 1 3 1 ? ? 1 1 4 1 2 1 1 1 1 1 2 1 1 3 

Ceraon tasmaniae 2 1 2 2 1 1 2 1 3 2 2 1 1 1 6 2 2 1 1 1 2 3 2 1 2 3 

Choucentrus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Coccosterphus 1 2 3 1 2 1 2 1 3 2 2 1 1 2 1 3 1 1 3 1 1 1 2 1 1 1 

Cornutobelus mutabilis 1 1 2 2 1 1 2 1 3 2 1 1 1 1 3 3 2 1 2 1 1 2 2 ? ? ? 

Crito festivum ? ? ? ? ? ? ? ? ? ? ? ? ? ? 6 2 2 ? 1 ? 2 ? ? ? ? ? 

Cryptaspidia pubera 1 2 1 ? 2 1 1 1 3 1 ? ? 1 2 1 3 1 1 3 1 1 1 2 2 2 3 

Cryptoparma parva ? ? ? ? ? ? ? 1 3 2 2 1 1 2 1 3 1 1 3 1 1 1 2 ? ? ? 

Dacaratha 2 1 2 2 1 1 2 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 2 2 3 

Daconotus projectus 1 1 1 ? 2 1 1 1 3 2 1 1 1 1 3 3 2 1 2 1 1 1 2 2 2 3 

Dagonotus lectus ? ? ? ? ? ? ? 1 3 2 2 1 1 1 3 3 2 1 2 1 1 1 2 ? ? ? 

Daimon serricorne 1 2 3 2 1 1 2 1 3 2 1 1 1 1 4 1 2 1 3 1 1 2 2 1 1 2 

Demanga sooknana ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Dingkana borealis 1 2 2 1 2 1 2 1 3 2 1 2 1 1 6 2 2 1 1 1 2 3 2 1 2 2 

Distanobelus sericeus 1 1 2 1 1 1 1 1 3 2 1 1 1 1 3 3 2 1 2 1 1 1 2 ? ? ? 

Dograna suffulta 1 2 2 1 1 1 1 1 3 2 2 1 1 1 7 3 2 2 1 3 2 2 2 ? ? ? 

Dukeobelus simplex 1 1 2 1 2 1 1 1 3 2 2 1 1 1 4 1 2 1 1 1 1 1 2 1 1 2 

Ebhul varium 1 1 1 ? 1 1 1 1 3 2 1 2 1 1 5 3 2 1 1 1 1 3 2 1 2 2 

Elaphiceps 1 1 1 ? 1 1 2 1 3 2 2 1 2 2 1 1 1 1 3 1 1 1 2 1 1 3 

Emphusis obesa 1 2 2 2 1 1 1 1 3 2 1 1 1 1 3 3 2 1 2 1 1 2 2 2 2 3 

414


1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 3 

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 

Erecticornia 1 1 1 1 3 2 1 2 1 1 1 2 ? 1 3 ? 1 2 2 1 ? 1 ? ? ? 1 2 1 2 1 

Euceropsila primus 1 1 1 2 2 2 1 2 1 2 1 1 1 1 4 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Eucoccosterphus 1 1 2 1 3 2 1 2 1 1 1 2 2 1 1 2 1 2 2 1 2 1 1 1 2 2 1 1 3 1 

Eufairmairia 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 2 1 1 2 2 1 2 1 2 1 

Eufairmairiella 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 2 1 1 2 2 1 2 1 2 1 

Eufrenchia falcata 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 2 1 1 2 2 1 2 1 2 1 

Eumocentrulus 1 1 1 2 2 2 1 2 1 2 2 1 2 1 4 2 1 2 2 1 1 1 1 1 2 2 2 1 2 1 

Eumonocentrus 1 1 1 2 2 2 1 2 1 2 1 ? 2 1 4 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Eutryonia monstrifera 1 1 2 1 2 2 1 2 1 2 1 2 2 1 5 1 2 2 2 1 2 1 1 1 2 1 2 1 2 1 

Evanchon 1 1 2 1 2 1 1 2 1 2 2 1 1 1 3 1 1 2 2 1 1 1 1 1 2 1 1 1 2 1 

Evansiana iasis 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 2 1 1 2 2 1 2 1 2 1 

Farcicaudia nitida 1 1 1 2 2 2 1 2 1 2 1 1 1 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Flatyperphyma flavocristatus 1 1 1 2 2 2 1 2 1 2 1 1 2 1 4 2 1 2 2 1 1 1 1 1 2 2 2 1 2 1 

Flexanotus albescens 1 1 1 2 2 2 1 2 1 2 2 1 1 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Foliatrotus elephas 1 1 1 2 2 2 1 2 1 2 1 1 2 1 4 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Funkhouserella brevifurca 1 1 2 2 2 2 1 2 ? ? 1 2 2 1 5 1 2 2 2 1 2 ? 2 1 2 2 2 1 1 1 

Funkhouserella pinguiturris 1 1 2 2 2 2 1 2 1 2 1 ? 2 1 5 1 2 2 2 1 2 1 2 1 2 2 1 1 2 1 

Gargara 1 1 2 1 3 2 1 2 1 1 1 2 2 1 1 1 1 2 2 1 2 1 1 1 2 1 2 1 3 1 

Gargarina carinata 1 1 1 1 3 2 1 2 1 1 1 2 ? 3 1 ? 1 2 ? ? ? 1 ? ? 2 ? 2 1 2 1 

Gigantorhabdus enderleini 1 1 2 1 4 1 1 2 1 2 1 2 2 1 5 1 2 2 2 1 2 1 2 2 2 2 2 1 1 1 

Goddefroyinella neglecta 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 2 1 2 2 2 1 2 2 2 1 

Goniolomus tricorniger 1 1 1 2 2 1 1 2 1 2 1 2 ? 3 3 1 1 2 ? ? 2 1 1 1 2 1 2 2 2 1 

Hamma 1 1 1 2 2 2 1 2 1 2 2 1 1 1 3 2 1 2 2 1 1 1 1 1 2 1 2 1 3 1 

Hemicentrus 1 1 1 1 2 2 1 1 ? ? ? ? ? ? 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Hybanda anodonta 1 1 2 1 2 2 1 2 1 1 1 2 2 1 ? 1 2 2 2 1 2 1 2 2 2 2 2 1 1 1 

Hybandoides 1 1 2 1 3 2 1 2 1 1 1 1 2 1 1 1 2 2 2 1 2 1 2 2 2 2 1 1 2 1 

Hypsauchenia hardwickii 1 1 2 1 4 2 1 2 1 2 1 1 2 1 5 1 2 2 2 1 1 1 2 2 2 2 2 1 1 1 

Hypsolyrium uncinatum 1 1 2 1 4 2 1 2 1 ? 1 1 2 1 1 1 2 2 2 1 2 1 2 2 2 2 2 1 2 1 

Imporcitor typicus 1 1 2 1 2 2 1 2 1 1 1 1 2 1 3 1 1 2 2 1 2 1 1 1 2 1 1 1 2 1 

Indicopleustes albomaculata 1 1 2 1 2 2 1 2 1 1 2 1 1 1 3 1 1 2 2 1 1 2 1 1 2 2 1 1 2 1 

Ischnocentrus 1 1 1 1 2 1 1 2 1 1 1 1 1 1 1 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Jacobiana 1 1 1 2 2 2 1 2 1 2 1 1 2 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 3 1 

Jingkara hyalipunctata 1 1 2 1 4 2 1 2 1 2 2 1 2 1 5 2 2 2 2 1 1 1 2 2 2 2 2 1 2 1 

Joveriana 1 1 2 1 2 2 1 2 1 2 1 1 2 1 1 1 1 2 2 1 1 1 1 1 2 1 2 1 1 1 

Kallicrates bellicornis 1 1 1 2 2 2 1 2 1 2 1 1 1 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 3 1 

Kanada irvinei 1 1 2 1 3 2 1 2 1 1 1 2 2 1 1 1 1 2 2 1 2 1 1 1 2 1 2 1 3 1 

Lanceonotus 1 1 2 1 2 2 1 2 1 2 2 1 1 1 3 1 1 2 2 1 1 2 1 1 2 2 2 1 2 1 

Leprechaunus 1 1 1 2 2 2 1 2 1 1 2 1 2 1 1 1 1 2 2 1 1 1 1 1 2 1 2 1 3 1 

Leptobelus 1 1 2 1 2 2 1 2 2 2 1 1 1 1 5 1 2 2 1 ? 1 2 1 1 2 1 2 1 2 1 

Leptocentrus 1 1 2 1 2 2 1 2 1 2 1 1 1 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 1 1 

Leptoceps viniculum 1 1 2 1 2 2 1 2 1 2 2 1 1 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Lobocentrus 1 1 2 1 2 2 1 2 1 1 1 1 1 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Lubra spincornis 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 2 1 1 2 2 1 2 1 2 1 

Maarbarus 1 1 1 2 2 2 1 2 1 2 2 1 1 1 3 1 1 2 1 ? 1 2 1 1 2 1 1 1 2 1 

Mabokiana teocchii 1 1 2 1 2 2 1 2 1 2 1 1 2 1 3 1 1 2 2 1 1 ? 1 1 2 1 2 1 2 1 

Machaerotypus sibiricus 1 1 2 1 3 2 1 2 1 1 1 2 2 1 3 1 1 2 2 1 2 1 1 1 2 1 2 1 3 1 

Madlinus seychellensis 1 1 1 1 3 2 1 2 1 1 1 2 ? 3 1 2 1 2 2 1 2 1 1 1 2 2 1 1 3 1 

Maguva nigra 1 1 2 1 2 2 1 2 1 1 2 1 2 1 3 2 1 2 2 1 1 1 1 1 2 2 2 1 2 1 

Marshallella rubripes 1 1 1 1 2 2 1 2 1 2 1 2 ? 3 2 1 1 2 ? ? 2 1 1 1 2 ? 2 1 2 1 

Matonotus 1 1 1 2 2 2 1 2 1 2 2 1 2 1 3 ? 1 2 2 1 ? ? ? ? ? ? 2 1 2 1 

Matumuia 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 ? 1 2 ? ? ? 1 ? ? 2 ? 2 1 1 1 

Maurya 1 1 2 1 3 2 1 2 1 1 1 2 2 1 3 1 1 2 2 1 2 1 1 1 2 1 2 1 3 1 

Menthogonus badhami 1 1 1 2 2 2 1 2 1 2 2 1 1 1 3 1 1 2 2 1 1 1 1 1 2 2 2 1 2 1 

Mesocentrina pyramidata 1 1 1 1 3 2 1 2 1 1 1 2 ? 3 1 1 1 2 ? ? 2 1 1 1 2 2 2 1 3 1 

Micreune formidanda 1 1 1 2 2 1 1 2 1 2 1 1 1 1 5 1 2 2 2 1 1 1 1 1 2 2 2 1 2 1 

Microcentrus caryae 1 1 1 1 2 2 1 2 1 1 1 2 2 1 2 1 1 2 2 1 2 1 1 1 1 2 1 1 2 1 

Mitranotus albofascipennis 1 1 1 2 2 2 1 2 1 2 2 1 2 1 4 2 1 2 2 1 1 1 1 1 2 2 2 1 2 1 

Monanchon monanchonus 1 1 1 2 2 2 1 2 1 2 2 1 1 1 3 1 1 2 2 1 1 1 1 1 2 2 2 1 2 1 

Monobeloides stuarti 1 1 1 2 2 2 1 2 1 2 1 2 ? 3 1 1 1 2 2 1 2 1 1 1 2 1 2 2 2 1 

Monobelus 1 1 1 2 2 2 1 2 1 2 1 2 2 1 1 1 1 2 2 1 1 2 1 1 2 1 2 1 2 1 

Monocentrus 1 1 1 2 2 2 1 2 1 2 2 1 2 1 4 1 1 2 2 1 1 1 1 1 2 2 2 1 2 1 

Multareiodes 1 1 1 1 2 1 1 2 1 ? 1 2 ? 3 3 1 1 2 ? ? 2 1 2 2 1 2 1 1 3 ? 

Multareis 1 1 1 1 2 2 1 2 1 ? 1 2 ? 3 2 1 1 2 ? ? 2 1 2 2 1 2 1 1 2 ? 

Negus asper 1 1 2 1 2 2 2 2 1 2 1 2 2 1 3 2 1 2 2 1 1 2 2 2 2 2 2 1 3 1 

Neocanthuchus tropicus 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 2 1 1 2 2 1 2 1 2 1 

Neocentrus rufus 1 1 2 ? 2 2 1 2 1 1 1 1 ? 1 1 ? 1 2 2 1 ? ? ? ? 2 ? 2 1 2 1 

Neomachaerotypus eguchii 1 1 2 ? 3 2 ? 2 1 1 1 2 ? 1 3 ? 1 2 ? ? ? ? ? ? 2 ? 2 1 ? 1 

Neosextius 1 1 2 1 2 2 1 2 1 2 1 2 ? 3 4 ? 1 2 ? ? ? 1 ? ? 2 1 1 ? ? 1 

Nessorhinus 1 1 1 2 2 1 1 2 1 2 1 2 ? 3 4 1 1 2 2 1 2 1 1 1 2 1 2 1 2 1 

Nicomia 2 1 1 2 2 2 1 1 ? ? ? ? ? ? 1 1 1 1 3 ? 1 1 1 1 2 1 2 1 2 2 

Nilautama minutaspina 1 1 2 1 2 2 1 2 1 1 1 1 1 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Nilautama typica 1 1 2 1 2 2 1 2 1 ? 1 1 1 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Nodonica bispinigera 1 1 1 2 2 2 1 2 1 1 1 2 ? 3 3 1 1 2 ? ? 2 ? ? ? 1 2 1 1 ? ? 

Nondenticentrus 1 1 1 1 3 2 1 2 1 1 1 2 2 1 3 1 1 2 2 1 2 1 1 1 2 1 2 1 2 1 

415


3 3 3 3 3 3 3 3 3 4 4 4 4 4 4 4 4 4 4 5 5 5 5 5 5 5 5 5 5 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 

Erecticornia 1 1 1 1 1 1 2 2 1 ? 1 2 1 1 2 2 1 2 2 1 1 1 2 2 2 5 ? ? ? ? 

Euceropsila primus 1 ? 1 1 1 1 2 2 1 2 ? 2 1 1 2 1 2 ? 2 1 1 1 2 2 2 5 1 1 1 2 

Eucoccosterphus 1 1 1 1 1 1 2 2 1 1 1 1 1 1 2 1 1 1 2 1 1 1 1 2 3 5 1 2 1 2 

Eufairmairia 1 1 1 1 1 1 1 2 1 3 1 2 1 1 1 ? 1 1 1 2 1 2 1 1 1 1 2 1 1 1 

Eufairmairiella 1 1 1 1 1 1 1 2 1 3 1 2 1 1 1 ? 1 1 1 2 1 1 1 1 1 1 ? 1 1 1 

Eufrenchia falcata 1 1 1 1 1 1 1 1 1 2 1 2 1 1 1 ? 1 1 1 2 1 2 1 1 1 1 2 1 1 1 

Eumocentrulus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 1 5 1 1 1 2 

Eumonocentrus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 2 ? 2 1 1 1 2 2 2 5 1 1 1 2 

Eutryonia monstrifera 1 1 1 1 1 1 2 2 1 2 1 2 1 1 1 ? 1 1 1 2 1 1 1 2 1 1 1 1 1 1 

Evanchon 2 ? 1 1 1 1 2 2 1 1 2 2 1 1 2 1 2 ? 2 1 1 1 ? 2 3 5 1 1 1 1 

Evansiana iasis 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 2 1 2 1 1 1 2 1 1 1 1 1 1 

Farcicaudia nitida 1 ? 1 1 1 1 2 2 1 1 2 2 1 1 2 1 2 ? 2 1 1 1 1 2 3 5 1 1 1 2 

Flatyperphyma flavocristatus 1 1 1 1 1 1 2 1 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 1 5 1 1 1 2 

Flexanotus albescens 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 2 5 1 1 1 2 

Foliatrotus elephas 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 2 5 1 1 1 2 

Funkhouserella brevifurca 1 1 1 1 1 1 1 1 1 1 1 1 1 1 ? ? 1 1 1 ? 1 ? ? ? ? 1 1 1 1 1 

Funkhouserella pinguiturris 1 1 1 1 1 1 2 1 1 1 1 1 1 2 1 ? 1 1 1 2 1 1 ? ? 1 1 2 1 1 1 

Gargara 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 2 2 1 1 1 2 2 3 5 1 1 1 2 

Gargarina carinata 1 1 1 1 1 1 2 2 1 ? 1 2 1 1 1 ? 1 2 2 1 1 1 1 2 3 5 ? ? ? ? 

Gigantorhabdus enderleini 1 1 1 2 1 1 2 1 2 1 1 1 1 2 1 ? 1 1 1 2 1 1 ? 1 1 1 2 1 1 1 

Goddefroyinella neglecta 1 1 ? 1 ? 1 1 1 1 3 1 1 1 2 1 ? 1 2 1 2 1 1 1 1 1 1 2 1 1 1 

Goniolomus tricorniger 1 1 1 1 1 1 2 1 1 2 1 2 1 1 2 1 1 2 2 1 1 2 2 1 1 1 1 1 1 1 

Hamma 1 ? 1 1 1 1 2 2 1 1 2 2 1 1 2 1 2 ? 2 1 1 1 1 2 3 5 1 1 1 2 

Hemicentrus 1 1 1 1 1 1 2 2 1 2 1 2 1 1 1 ? 1 3 2 1 1 1 1 2 1 1 1 1 1 2 

Hybanda anodonta 1 1 1 1 1 1 1 1 2 1 1 1 2 1 1 ? 1 1 1 3 1 1 1 1 1 1 1 1 1 1 

Hybandoides 1 1 1 2 1 1 1 1 ? 1 1 1 2 1 1 ? 1 2 2 3 1 1 1 1 1 1 1 1 1 1 

Hypsauchenia hardwickii 1 1 1 1 1 1 1 1 1 1 1 1 2 2 1 ? 1 1 1 3 1 1 ? 1 1 1 1 1 1 1 

Hypsolyrium uncinatum 1 1 1 1 1 1 1 1 1 1 1 1 2 2 1 ? 1 1 2 3 1 1 ? 1 1 1 1 1 1 1 

Imporcitor typicus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 3 2 1 1 1 1 1 1 1 1 1 1 1 

Indicopleustes albomaculata 1 2 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 1 2 2 2 2 1 2 1 1 1 1 1 2 

Ischnocentrus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 1 1 1 1 2 2 

Jacobiana 1 1 1 1 1 1 2 1 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 1 5 1 1 1 2 

Jingkara hyalipunctata 1 1 1 2 1 1 1 1 1 3 1 1 2 2 ? ? 1 2 1 3 1 1 ? ? ? 1 1 1 1 1 

Joveriana 1 1 1 1 1 1 2 2 1 2 1 2 1 1 1 ? 1 3 ? 2 1 2 2 1 1 1 1 1 1 2 

Kallicrates bellicornis 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 2 5 1 1 1 2 

Kanada irvinei 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 2 1 2 2 1 1 1 2 2 3 5 1 1 ? ? 

Lanceonotus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 1 1 1 1 1 1 2 2 1 1 1 1 1 

Leprechaunus 1 1 1 1 1 1 2 1 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 2 5 1 1 1 2 

Leptobelus 1 ? 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 2 2 1 1 1 1 2 3 1 1 1 2 2 

Leptocentrus 1 1 1 1 1 1 2 2 1 2 1 2 1 1 1 ? 1 3 1 2 1 2 1 1 1 1 1 1 1 2 

Leptoceps viniculum 1 1 1 1 1 1 2 2 1 2 1 2 1 1 1 ? 1 2 1 1 1 1 2 2 2 1 1 1 1 2 

Lobocentrus 1 1 1 1 1 1 2 2 1 2 1 2 1 1 1 ? 1 3 2 2 1 1 2 1 1 1 1 1 1 2 

Lubra spinicornis 1 1 1 1 1 1 1 2 1 3 1 2 1 1 1 ? 1 1 1 2 1 2 1 2 1 1 1 1 1 1 

Maarbarus 1 2 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 1 2 2 2 2 1 2 3 1 1 1 1 2 

Mabokiana teocchii 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 2 1 2 1 1 1 2 1 1 1 1 1 1 

Machaerotypus sibiricus 1 1 1 1 1 1 2 2 1 2 1 2 1 1 1 ? 1 2 2 1 1 1 2 2 3 5 1 1 1 2 

Madlinus seychellensis 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 2 2 1 1 1 1 2 3 5 1 1 1 2 

Maguva nigra 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 2 1 2 1 1 1 1 1 1 1 1 1 1 

Marshallella rubripes 1 1 1 1 1 1 2 2 1 2 1 2 1 1 1 ? 1 2 1 2 1 2 1 1 1 1 1 1 1 1 

Matonotus 1 1 1 1 1 1 2 2 1 ? 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 3 5 ? ? ? ? 

Matumuia 1 1 1 2 ? 1 1 2 1 3 1 2 1 1 1 ? 1 2 1 2 1 2 2 1 1 1 ? ? ? ? 

Maurya 1 1 1 1 1 1 2 ? 1 1 1 2 1 1 1 ? 1 2 2 1 1 1 2 1 3 5 1 1 1 2 

Menthogonus badhami 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 2 5 1 1 1 2 

Mesocentrina pyramidata 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 2 1 3 2 1 1 1 2 2 3 5 1 1 2 2 

Micreune formidanda 1 2 1 1 1 1 2 2 1 1 1 2 1 1 2 1 1 2 1 2 1 2 1 2 1 1 1 1 1 2 

Microcentrus caryae 1 1 1 1 1 1 2 1 1 1 1 2 1 1 1 ? 1 1 1 2 1 1 2 1 1 1 1 1 1 1 

Mitranotus albofascipennis 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 1 5 1 1 1 2 

Monanchon monanchonus 1 ? 1 1 1 1 2 2 1 1 2 2 1 1 2 1 2 ? 2 1 1 1 2 2 1 5 1 1 1 2 

Monobeloides stuarti 1 1 1 1 1 1 1 2 1 1 1 2 1 1 1 ? 1 2 1 1 1 1 1 2 3 1 1 1 1 1 

Monobelus 1 1 1 1 1 1 1 2 1 1 1 2 1 1 1 ? 1 2 2 1 1 1 1 2 3 1 1 1 1 1 

Monocentrus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 2 5 1 1 1 2 

Multareiodes 1 ? ? ? 1 1 ? ? 1 1 1 1 1 2 1 ? 1 ? ? ? 1 1 ? ? ? 2 2 1 1 1 

Multareis 1 ? ? ? 1 1 ? ? 1 1 1 1 1 2 1 ? 1 ? 2 ? 1 1 ? ? ? 3 2 1 1 1 

Negus asper 1 1 1 1 1 1 2 1 1 1 1 2 1 1 2 1 1 2 1 1 1 1 2 2 1 1 1 1 1 2 

Neocanthuchus tropicus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 1 1 2 1 1 1 2 1 1 1 1 1 1 

Neocentrus rufus 1 1 1 1 1 1 2 2 1 ? 1 2 1 1 1 ? 1 2 1 2 1 1 2 1 1 1 ? ? ? ? 

Neomachaerotypus eguchii 1 1 1 1 1 1 2 1 ? ? 1 2 1 1 1 ? 1 1 1 1 1 1 2 2 3 5 ? ? ? ? 

Neosextius 1 ? 1 1 1 1 1 2 ? 1 1 1 1 2 1 ? 1 1 1 2 1 ? ? ? ? 1 ? ? ? ? 

Nessorhinus 1 1 1 1 1 1 ? 1 1 2 1 2 1 1 2 1 1 2 1 2 1 1 2 1 1 1 1 1 1 1 

Nicomia 1 1 2 2 1 1 2 2 1 2 1 2 1 1 ? ? 1 2 2 2 1 1 ? ? ? 1 1 1 1 2 

Nilautama minutaspina 1 1 1 1 1 1 2 2 1 2 1 2 1 1 1 ? 1 3 2 1 1 2 2 1 1 1 1 1 1 2 

Nilautama typica 1 1 1 1 1 1 2 2 1 2 1 2 1 1 2 2 1 3 2 1 1 2 2 1 1 1 1 1 1 2 

Nodonica bispinigera 1 ? ? ? ? ? 2 2 1 2 1 1 ? 2 1 ? ? ? ? ? 1 1 ? ? ? 5 ? ? ? ? 

Nondenticentrus 1 1 1 1 1 1 2 2 1 2 1 2 1 1 1 ? 1 2 2 1 1 1 2 2 2 5 1 1 1 2 

416


6 6 6 6 6 6 6 6 6 7 7 7 7 7 7 7 7 7 7 8 8 8 8 8 8 8 8 8 8 9 

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 

Erecticornia ? ? 1 1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Euceropsila primus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 3 2 1 2 2 1 1 ? ? ? 

Eucoccosterphus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 3 2 2 1 ? 2 1 1 ? ? 

Eufairmairia 1 ? 1 1 1 1 ? 2 1 2 1 1 1 1 1 2 1 1 2 1 1 1 1 1 ? 2 1 4 1 2 

Eufairmairiella 1 ? 1 1 1 1 ? 2 1 2 1 1 1 2 2 2 1 1 2 1 1 1 1 1 ? 2 1 ? ? ? 

Eufrenchia falcata 1 ? 1 1 1 1 ? 2 1 2 1 1 1 2 1 2 1 1 2 1 1 ? 1 1 ? ? 1 3 1 1 

Eumocentrulus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 2 1 1 3 1 1 

Eumonocentrus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 2 2 1 1 ? ? 

Eutryonia monstrifera 1 ? 1 1 1 1 ? 2 1 2 1 1 1 2 2 2 1 1 2 1 1 2 1 1 ? 1 1 1 ? ? 

Evanchon 1 ? 1 1 1 1 ? 2 1 2 1 ? 1 2 2 2 1 1 2 1 1 1 2 1 ? 2 2 1 ? ? 

Evansiana iasis 1 ? 1 1 1 1 ? 2 1 2 1 ? 1 2 2 2 1 1 2 ? ? ? ? ? ? 1 1 ? ? ? 

Farcicaudia nitida 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 2 1 1 3 2 1 

Flatyperphyma flavocristatus 2 1 1 1 1 1 ? 2 1 2 1 ? 1 3 2 2 3 1 2 ? ? ? ? ? ? 1 ? ? ? ? 

Flexanotus albescens 2 1 1 1 1 1 ? 2 1 2 1 ? 1 ? 2 2 3 1 2 1 1 2 1 2 2 2 1 3 1 1 

Foliatrotus elephas 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 2 2 1 1 ? ? 

Funkhouserella brevifurca 1 ? 1 1 1 1 ? 2 1 1 ? ? 1 2 1 2 1 1 2 1 1 ? 1 1 ? 1 ? 3 1 2 

Funkhouserella pinguiturris 1 ? 1 1 1 1 ? 2 1 2 1 ? 1 1 1 2 1 1 2 1 1 ? 1 1 ? 1 1 4 1 2 

Gargara 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 3 2 2 2 1 2 1 1 ? ? 

Gargarina carinata ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Gigantorhabdus enderleini 1 ? 1 1 1 1 ? 1 ? 1 ? 1 1 2 1 1 ? 2 2 1 1 2 1 1 ? 1 1 2 ? ? 

Goddefroyinella neglecta 1 ? 1 1 1 1 ? 1 ? 1 ? 1 1 1 1 2 1 1 2 1 1 1 1 1 ? 1 1 3 2 2 

Goniolomus tricorniger 1 ? 1 1 1 1 ? 2 1 1 ? 1 1 3 2 2 1 1 2 2 1 2 2 1 ? 3 2 1 ? ? 

Hamma 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 2 2 1 3 1 2 

Hemicentrus 2 1 1 1 1 1 ? 2 1 2 1 1 ? 3 2 2 3 1 2 1 1 2 1 2 1 2 1 4 1 1 

Hybanda anodonta 1 ? 1 1 1 1 ? 1 ? 1 ? 1 1 2 1 1 ? 2 2 1 1 2 1 1 ? 1 1 2 ? ? 

Hybandoides 1 ? 1 1 2 1 ? 1 ? 1 ? 1 1 2 1 1 ? 2 2 1 1 2 1 1 ? 1 1 2 ? ? 

Hypsauchenia hardwickii 1 ? 1 1 1 1 ? 1 ? 1 ? 1 1 2 1 1 ? 2 2 1 1 2 1 1 ? 1 1 2 ? ? 

Hypsolyrium uncinatum 1 ? 1 1 1 1 ? 1 ? 1 ? 1 1 2 1 1 ? 2 2 1 1 2 1 1 ? 1 1 2 ? ? 

Imporcitor typicus 1 ? 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 1 1 2 1 1 2 1 1 ? 1 2 1 ? ? 

Indicopleustes albomaculata 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 1 1 2 1 1 2 1 1 ? 1 2 1 ? ? 

Ischnocentrus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 1 1 2 1 3 2 1 1 ? 1 2 1 ? ? 

Jacobiana 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 2 1 1 3 2 1 

Jingkara hyalipunctata 1 ? 1 1 1 1 ? 1 ? 1 ? ? 1 2 1 1 ? 2 2 1 1 2 1 1 ? 1 1 2 ? ? 

Joveriana 2 1 1 1 1 1 ? 2 1 2 1 3 ? 3 2 2 3 1 2 1 3 2 1 2 1 2 1 4 1 1 

Kallicrates bellicornis 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 2 2 1 3 1 1 

Kanada irvinei ? ? 1 1 1 1 ? 2 1 2 1 ? 1 ? 2 2 1 1 2 1 1 2 2 2 1 2 1 1 ? ? 

Lanceonotus 1 ? 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 1 1 2 1 1 2 1 1 ? 1 2 4 1 1 

Leprechaunus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 1 1 2 1 1 2 1 2 ? 2 1 3 2 2 

Leptobelus 2 1 1 1 1 1 ? 2 1 2 1 3 3 3 2 2 3 1 2 1 1 2 1 1 ? 1 2 3 1 1 

Leptocentrus 2 1 1 1 1 1 ? 2 1 2 1 1 2 3 2 2 3 1 2 1 2 2 1 2 1 2 1 4 1 1 

Leptoceps viniculum 2 1 1 1 1 1 ? 2 1 2 1 ? ? 3 2 2 3 1 ? ? ? ? ? ? ? ? ? ? ? ? 

Lobocentrus 2 2 1 1 1 1 ? 2 1 2 2 1 1 3 2 2 3 1 2 1 1 2 1 1 ? 1 2 1 ? ? 

Lubra spinicornis 1 ? 1 1 1 1 ? 2 1 2 1 1 1 2 2 2 1 1 2 1 1 2 1 1 ? 1 1 4 1 1 

Maarbarus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 1 1 2 1 1 2 1 1 ? 1 2 1 ? ? 

Mabokiana teocchii 1 ? 1 1 1 1 ? 2 1 2 2 1 ? 3 2 2 1 1 ? ? ? 2 ? ? ? 2 ? ? ? ? 

Machaerotypus sibiricus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 2 2 1 1 1 3 2 2 

Madlinus seychellensis 2 1 2 2 2 1 ? 2 1 2 1 1 1 3 2 2 1 1 ? ? ? ? ? ? ? 2 1 ? ? ? 

Maguva nigra 1 ? 1 1 1 1 ? 2 1 2 2 1 1 3 2 2 1 1 2 1 2 2 2 1 ? 2 2 1 ? ? 

Marshallella rubripes 1 ? 1 1 1 1 ? 2 1 2 1 1 1 3 1 2 1 1 2 1 1 2 2 1 ? 2 2 1 ? ? 

Matonotus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Matumuia ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 2 ? 1 ? ? ? ? ? ? ? ? ? ? ? ? 

Maurya 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 2 2 1 1 1 1 ? ? 

Menthogonus badhami 2 1 1 1 1 1 ? 2 1 2 1 ? 1 3 2 2 1 1 2 1 1 2 1 2 ? 2 ? 3 2 1 

Mesocentrina pyramidata 2 1 1 1 1 1 ? 2 1 2 1 ? 1 3 2 2 3 1 2 1 3 2 2 2 1 2 1 1 ? ? 

Micreune formidanda 2 2 1 1 1 1 ? 2 1 2 2 1 1 3 2 2 3 1 2 1 3 2 1 2 1 3 1 1 ? ? 

Microcentrus caryae 1 ? 1 1 1 2 2 2 1 2 1 1 1 2 2 2 1 1 1 1 1 ? 1 1 ? 1 1 3 1 ? 

Mitranotus albofascipennis 2 1 1 1 1 1 ? 2 1 2 1 ? 1 3 2 2 3 1 ? ? ? 2 ? ? ? 1 ? ? ? ? 

Monanchon monanchonus 2 1 1 1 1 1 ? 2 1 2 1 ? 1 3 2 2 3 1 ? ? ? 2 ? ? ? 2 ? ? ? ? 

Monobeloides stuarti 1 ? 1 1 1 2 1 2 1 2 1 3 3 3 2 2 1 1 2 1 1 2 1 1 ? 1 2 1 ? ? 

Monobelus 1 ? 1 1 1 2 1 2 1 2 1 3 3 3 2 2 1 1 2 1 3 2 2 1 ? 1 2 3 1 2 

Monocentrus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 2 1 2 1 1 2 1 2 2 2 1 1 ? ? 

Multareiodes 1 ? 1 1 1 1 ? 1 ? 1 ? 1 1 2 1 1 ? 2 2 1 1 1 1 1 ? 1 1 1 ? ? 

Multareis 1 ? 1 1 1 1 ? 1 ? 1 ? 1 1 2 1 1 ? 2 2 1 1 1 1 1 ? 1 1 1 ? ? 

Negus asper 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 2 1 2 1 2 3 3 1 3 1 2 

Neocanthuchus tropicus 1 ? 1 1 1 1 ? 2 1 2 1 ? 1 2 2 2 1 1 2 ? ? ? ? ? ? 1 1 ? ? ? 

Neocentrus rufus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Neomachaerotypus eguchii ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Neosextius ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Nessorhinus 1 ? 1 1 1 1 ? 2 1 1 ? 1 1 3 2 2 1 1 2 2 1 2 2 1 ? 3 2 1 ? ? 

Nicomia 2 1 1 3 1 2 3 2 1 2 1 1 1 3 1 2 1 1 1 1 1 2 1 1 ? 1 1 4 1 ? 

Nilautama minutaspina 2 1 1 1 1 1 ? 2 1 2 1 3 1 3 2 2 3 1 2 1 2 2 1 2 1 2 1 4 1 1 

Nilautama typica 2 1 1 1 1 1 ? 2 1 2 1 3 1 3 2 2 3 1 ? ? ? 2 ? 2 1 ? 1 ? ? ? 

Nodonica bispinigera ? ? ? ? ? ? ? 2 ? 2 ? ? ? ? 1 2 ? 1 ? ? ? ? ? ? ? ? ? 1 ? ? 

Nondenticentrus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 3 2 2 2 ? 1 1 1 ? ? 

417


1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 

9 9 9 9 9 9 9 9 9 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 

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 

Erecticornia ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Euceropsila primus ? ? ? ? ? ? ? 1 3 2 1 1 1 1 3 3 2 1 2 1 1 2 2 ? ? ? 

Eucoccosterphus 1 2 3 1 1 1 2 1 3 2 2 1 1 1 4 1 2 1 1 1 1 1 2 1 1 1 

Eufairmairia 2 1 2 2 1 1 2 1 3 2 1 1 1 1 6 2 2 1 1 1 2 3 2 1 2 3 

Eufairmairiella ? ? ? ? ? ? ? 1 3 2 2 1 1 1 6 2 2 1 1 1 2 3 2 ? ? ? 

Eufrenchia falcata 2 1 2 2 1 1 2 1 3 2 1 1 1 1 6 2 2 1 1 1 2 3 2 ? ? ? 

Eumocentrulus 1 1 2 1 1 1 1 1 3 2 1 1 1 1 3 3 2 1 2 1 1 2 2 ? ? 3 

Eumonocentrus 1 2 3 1 1 1 1 1 3 2 1 1 1 1 3 3 2 1 2 1 1 2 2 1 2 3 

Eutryonia monstrifera 1 2 2 1 1 1 2 1 3 1 ? ? 1 1 6 2 2 1 1 1 2 3 2 1 2 3 

Evanchon 1 2 3 2 1 1 2 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 1 2 3 

Evansiana iasis ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Farcicaudia nitida 1 1 2 1 1 1 2 1 3 2 2 1 1 1 3 3 2 1 2 1 1 1 2 2 2 3 

Flatyperphyma flavocristatus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Flexanotus albescens 1 1 2 1 1 1 1 ? ? ? ? ? ? ? 3 ? ? ? ? ? ? ? 2 ? ? ? 

Foliatrotus elephas 1 1 3 1 2 1 1 1 3 2 1 1 1 1 3 3 2 1 2 1 1 2 2 ? ? ? 

Funkhouserella brevifurca 1 1 2 2 1 1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 3 2 2 3 

Funkhouserella pinguiturris 2 1 3 2 1 1 2 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 3 ? ? ? 

Gargara 1 2 2 1 2 1 2 1 3 2 1 1 1 2 1 3 1 1 3 1 1 1 2 1 2 3 

Gargarina carinata ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Gigantorhabdus enderleini 1 1 2 1 1 2 1 2 2 2 1 2 1 1 5 3 2 1 1 1 1 1 2 1 2 1 

Goddefroyinella neglecta 1 1 2 1 1 1 2 1 3 2 2 1 1 1 6 2 2 1 1 1 2 ? 2 ? ? ? 

Goniolomus tricorniger 1 1 3 2 1 1 2 1 3 2 2 1 1 1 2 1 1 1 3 1 1 2 2 ? ? ? 

Hamma 1 1 2 1 1 1 1 1 3 2 1 1 1 1 3 3 2 1 2 1 1 1 2 2 2 3 

Hemicentrus 2 1 ? ? 1 ? ? 1 3 2 1 1 1 1 7 3 2 1 1 2 2 2 2 2 2 3 

Hybanda anodonta 1 1 1 ? 1 2 1 1 2 2 1 2 1 1 3 3 2 1 1 1 1 1 2 ? ? ? 

Hybandoides 1 1 1 ? 1 2 1 1 3 2 1 2 1 1 5 3 2 1 1 1 1 3 2 1 2 2 

Hypsauchenia hardwickii 1 1 1 ? 1 1 2 1 3 2 1 2 1 1 5 3 2 1 1 1 1 1 2 1 2 2 

Hypsolyrium uncinatum 1 1 1 ? 1 1 1 1 3 2 1 2 1 1 5 3 2 1 1 1 1 1 2 1 2 2 

Imporcitor typicus 1 2 3 2 1 1 2 1 3 2 2 1 1 1 4 1 ? 1 1 1 1 1 2 ? ? ? 

Indicopleustes albomaculata 1 1 2 2 2 1 1 1 3 2 2 1 1 1 7 3 2 2 1 3 1 2 2 ? ? ? 

Ischnocentrus 1 2 2 1 1 1 1 1 3 2 1 3 3 2 5 3 1 1 1 1 1 1 2 1 1 3 

Jacobiana 1 1 2 1 1 1 1 1 3 2 1 1 1 1 3 3 2 1 2 1 1 2 2 2 2 3 

Jingkara hyalipunctata 1 1 2 2 1 1 1 ? ? ? ? ? ? ? 5 3 2 ? ? ? 1 ? 2 ? ? ? 

Joveriana 2 1 2 2 1 1 ? 1 3 2 1 1 1 1 7 3 2 1 1 2 2 2 2 ? ? ? 

Kallicrates bellicornis 1 1 2 1 1 1 1 1 3 2 1 1 1 1 3 3 2 1 2 1 1 2 2 2 2 3 

Kanada irvinei 1 2 1 ? 2 1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Lanceonotus 2 1 2 1 1 1 1 1 3 1 ? ? 1 1 8 3 2 1 2 1 2 1 2 2 2 3 

Leprechaunus 1 1 2 1 1 1 1 1 3 2 1 1 1 1 3 3 2 1 2 1 1 1 2 ? ? ? 

Leptobelus 1 1 2 2 1 1 2 1 3 2 2 1 2 1 3 3 2 1 2 1 1 2 2 2 2 3 

Leptocentrus 2 1 2 2 1 1 2 1 3 2 1 1 1 1 7 3 2 1 1 2 2 2 2 2 2 3 

Leptoceps viniculum ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Lobocentrus 1 2 1 ? 2 1 1 1 3 2 2 1 1 2 1 1 1 1 3 1 1 1 2 2 2 3 

Lubra spinicornis 2 1 2 1 1 1 2 1 3 2 1 1 1 1 6 2 2 1 1 1 2 3 2 1 2 3 

Maarbarus 1 2 2 2 1 1 1 1 3 2 2 1 1 1 7 3 2 2 1 3 1 2 2 ? ? 3 

Mabokiana teocchii ? ? ? ? ? ? ? ? ? ? ? ? ? ? 4 1 2 1 ? 1 1 ? 2 ? ? ? 

Machaerotypus sibiricus 1 2 2 1 1 1 2 1 3 2 2 1 1 2 1 1 1 1 3 1 1 1 2 ? ? ? 

Madlinus seychellensis ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Maguva nigra 1 2 2 2 1 1 1 1 3 2 2 1 1 2 3 1 2 1 1 1 1 1 2 ? ? ? 

Marshallella rubripes 1 2 3 2 1 1 2 1 3 2 1 1 1 1 4 1 2 1 3 1 1 2 2 1 1 2 

Matonotus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Matumuia ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Maurya 1 1 2 1 1 1 1 1 3 2 2 1 1 2 1 3 1 1 3 1 1 1 2 2 2 3 

Menthogonus badhami 1 1 2 1 1 1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Mesocentrina pyramidata 1 2 1 ? 1 1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Micreune formidanda 1 2 2 2 2 1 2 1 3 2 2 1 1 1 7 3 2 1 2 2 1 1 2 2 2 3 

Microcentrus caryae 2 1 1 ? 1 1 2 1 3 2 2 3 1 1 1 1 1 1 3 1 1 1 1 1 2 2 

Mitranotus albofascipennis ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Monanchon monanchonus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Monobeloides stuarti 1 2 3 2 1 1 1 1 3 2 1 2 1 1 2 1 1 1 3 1 1 1 2 ? ? ? 

Monobelus 1 1 2 2 1 1 2 1 3 1 ? ? 1 2 2 1 1 1 3 1 1 1 2 2 2 3 

Monocentrus 1 2 3 2 1 1 1 1 3 2 1 1 1 1 3 3 2 1 2 1 1 1 2 2 2 3 

Multareiodes 1 2 2 1 1 1 1 1 3 2 1 2 1 1 1 1 1 1 3 1 1 1 2 1 2 3 

Multareis 1 2 2 1 1 1 1 1 3 2 1 2 1 1 1 1 1 1 3 1 1 1 2 1 2 3 

Negus asper 2 1 2 2 1 1 1 1 3 1 ? ? 1 1 3 3 2 1 2 1 1 1 2 2 2 3 

Neocanthuchus tropicus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Neocentrus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Neomachaerotypus eguchii ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Neosextius ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Nessorhinus 1 1 3 2 2 1 2 1 3 2 1 1 1 1 2 1 1 1 3 1 1 2 2 1 1 2 

Nicomia 2 1 2 2 1 1 1 1 3 2 2 1 1 ? 5 1 1 1 3 1 1 1 2 1 1 3 

Nilautama minutaspina 2 1 1 ? 1 1 2 1 3 2 1 1 1 1 7 3 2 1 1 2 2 2 2 2 2 3 

Nilautama typica ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Nodonica bispinigera 1 1 2 ? 1 ? ? ? 1 ? ? ? ? ? 1 ? 1 1 ? ? ? ? ? ? ? ? 

Nondenticentrus 1 2 2 1 2 1 1 1 3 2 2 1 1 2 1 1 1 1 3 1 1 1 2 1 2 3 

418


1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 3 

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 

Occator erectus 1 1 ? ? 2 1 1 2 1 1 1 1 ? 1 1 ? 1 2 ? ? ? ? ? ? 2 ? 2 1 2 1 

Ophicentrus notandus 1 1 2 1 2 1 1 2 1 2 2 1 1 1 1 1 1 2 2 1 1 1 1 1 2 2 2 1 2 1 

Orekthophora cornuta 1 1 1 2 2 1 1 2 1 2 1 2 ? 3 4 1 1 2 ? ? 2 1 1 1 2 2 2 1 1 1 

Orthobelus 1 1 1 1 2 2 1 2 1 2 1 2 2 2 3 1 1 2 2 1 2 1 1 1 2 1 2 1 2 1 

Otinotoides 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 2 1 1 1 2 1 2 1 1 1 

Otinotus ammon 1 1 1 1 2 2 1 2 1 1 1 1 2 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 ? 1 

Otinotus bantuantus 1 1 2 1 2 2 1 2 1 2 1 1 2 1 3 1 1 2 2 1 2 1 1 1 2 1 2 1 2 1 

Oxyrhachis 1 1 1 ? 2 2 1 2 1 2 1 2 ? 3 2 1 1 2 2 1 2 1 2 2 2 1 2 1 2 1 

Oxyrhachis carinata 1 1 1 ? 2 2 1 2 1 2 1 2 ? 3 3 1 1 2 ? ? ? 1 2 2 2 1 2 1 2 1 

Oxyrhachis delalandei 1 1 1 ? 2 2 1 2 1 2 1 2 ? 3 3 1 1 2 ? ? ? 1 2 2 2 1 2 1 2 1 

Oxyrhachis sulcicornis 1 1 1 ? 2 2 1 2 1 2 1 2 ? 3 3 1 1 2 ? ? ? 1 2 2 2 1 2 1 2 1 

Pantaleon 1 1 2 1 3 2 1 2 1 1 1 2 2 1 3 1 1 2 2 1 2 1 1 1 2 2 2 1 2 1 

Paracentronodus 1 2 1 1 2 2 1 2 1 2 2 2 2 1 1 1 1 2 2 1 1 1 1 1 1 1 1 1 2 3 

Paradarnoides 1 1 1 2 2 2 1 2 1 2 1 2 ? 3 1 1 1 2 ? ? 2 1 1 1 2 1 2 1 2 1 

Parapogon kandyiana 1 1 1 1 2 2 1 2 1 2 2 1 1 1 3 1 1 2 1 ? 1 2 1 1 2 1 1 1 3 1 

Paraxiphopoeus 1 1 1 2 2 2 1 2 1 2 2 1 1 1 3 1 1 2 2 1 1 1 1 1 2 2 2 1 1 1 

Parayasa 1 1 2 1 3 2 1 2 1 1 1 2 2 1 1 1 1 2 2 1 2 1 1 1 2 1 2 1 2 1 

Peltzerella borneensis 1 1 2 1 2 2 1 2 1 2 1 ? 2 1 3 1 1 2 2 1 ? 1 1 1 2 1 2 1 ? 1 

Periaman 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Pieltainellus 1 1 1 1 2 2 1 2 1 1 1 1 2 1 2 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Platybelus flavus 1 1 1 2 2 2 1 2 1 2 1 1 2 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Platycentrus acuticornis 1 1 1 1 2 2 1 2 1 1 1 2 2 1 3 1 1 2 2 1 2 1 1 1 2 1 2 1 2 1 

Pogonella minutus 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 2 1 1 1 2 1 2 1 2 1 

Pogon incurvatum 1 1 1 1 2 2 1 2 1 ? 1 1 2 1 3 1 1 2 2 1 1 2 1 1 2 2 1 1 2 1 

Pogonotus indicus 1 1 2 ? 2 1 1 2 1 2 2 1 ? 1 3 ? 1 2 1 ? 1 2 ? ? 2 ? 1 1 2 1 

Pogonotypellus australis 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 2 1 1 2 2 1 2 1 3 1 

Pogontypus 1 1 2 1 2 2 1 2 1 2 1 1 2 1 3 1 1 2 2 1 1 2 1 1 2 2 1 1 3 1 

Polonius froggatti 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 ? 1 1 1 2 2 2 1 2 1 

Promitor nominatus 1 1 1 2 2 2 1 2 1 1 1 1 2 1 1 1 1 2 2 1 1 1 1 1 2 1 2 1 3 1 

Protinotus doddi 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 ? 1 2 ? ? ? 1 ? ? 2 1 2 1 2 1 

Psilocentrus 1 1 1 1 2 2 1 2 1 1 1 1 1 1 1 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Pyrgauchenia 1 1 1 1 2 1 1 2 1 1 1 2 2 1 5 1 2 2 2 1 2 1 2 2 2 2 2 1 1 1 

Pyrgonota 1 1 2 2 2 1 1 2 1 1 1 2 2 1 5 1 2 2 2 1 2 1 2 1 2 2 2 1 2 1 

Rachinotus marshalli 1 1 2 1 2 2 1 2 1 2 1 1 2 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 1 1 

Rentzia 1 1 2 1 2 2 1 2 1 2 1 2 ? 1 3 ? 1 2 ? ? ? 1 ? ? 2 2 2 1 1 1 

Rexicornia elegans 1 1 1 2 2 2 1 2 1 2 1 1 1 1 3 1 1 2 2 1 1 1 1 1 2 2 2 1 2 1 

Rigula 1 1 2 1 2 2 1 2 1 2 1 2 ? 1 3 ? 1 ? ? ? ? 1 ? ? 2 1 2 1 2 1 

419


3 3 3 3 3 3 3 3 3 4 4 4 4 4 4 4 4 4 4 5 5 5 5 5 5 5 5 5 5 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 

Occator erectus 1 1 1 1 ? 1 2 2 1 ? 1 2 1 1 1 ? 1 3 1 2 1 2 1 1 1 1 ? ? ? ? 

Ophicentrus notandus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 1 2 ? 1 2 1 1 1 2 1 1 1 1 1 2 

Orekthophora cornuta 1 1 1 1 1 1 2 1 1 1 1 2 1 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 

Orthobelus 1 1 1 1 1 1 1 ? 1 2 1 2 1 1 1 ? 1 2 1 2 1 2 1 1 1 1 1 1 1 1 

Otinotoides 1 1 1 1 1 1 1 2 1 2 1 2 1 1 1 ? 1 1 1 2 1 1 2 2 1 1 1 1 1 1 

Otinotus ammon 1 1 1 1 1 1 2 2 1 2 1 2 1 1 1 ? 1 2 1 1 1 2 1 2 2 1 1 1 1 2 

Otinotus bantuantus 1 1 1 1 1 1 2 2 1 2 1 2 1 1 1 ? 1 3 1 2 1 2 1 1 1 1 1 1 1 2 

Oxyrhachis 1 1 1 1 2 1 1 1 1 2 1 2 1 1 1 ? 1 2 1 ? 1 1 ? ? 1 7 2 1 1 1 

Oxyrhachis carinata 1 1 1 1 2 1 1 1 1 2 1 2 1 1 1 ? 1 2 1 2 1 1 1 1 1 5 2 1 1 1 

Oxyrhachis delalandei 1 1 1 1 2 1 1 1 1 2 1 2 1 1 1 ? 1 2 1 1 1 1 1 2 1 1 2 1 1 1 

Oxyrhachis sulcicornis 1 1 1 1 2 1 1 1 1 2 1 2 1 1 1 ? 1 2 1 2 1 1 1 1 1 5 2 1 1 1 

Pantaleon 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 2 2 1 1 1 2 1 3 5 1 1 1 2 

Paracentronodus 1 ? 3 ? 1 1 2 1 1 2 1 1 1 1 1 ? 1 ? ? ? 1 1 ? ? ? 1 1 1 2 2 

Paradarnoides 1 1 1 1 1 1 2 1 1 2 1 2 1 1 2 1 1 1 1 2 1 1 1 1 1 1 1 1 1 1 

Parapogon kandyiana 1 2 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 1 2 1 2 2 1 2 1 1 1 1 1 2 

Paraxiphopoeus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 2 5 1 1 1 2 

Parayasa 1 1 1 1 1 1 2 2 1 1 1 1 1 1 2 1 1 2 2 1 1 1 ? 2 3 5 1 1 1 2 

Peltzerella borneensis 1 1 1 1 1 1 2 2 1 2 1 2 1 1 1 ? 1 3 2 1 1 2 2 1 1 1 1 1 1 2 

Periaman 1 1 1 1 1 1 2 2 1 2 1 2 1 1 1 ? 1 2 2 2 1 2 1 1 1 1 1 1 1 2 

Pieltainellus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 3 2 1 1 1 2 1 1 1 1 1 1 1 

Platybelus flavus 1 1 1 1 1 1 2 1 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 2 5 1 1 1 2 

Platycentrus acuticornis 1 1 1 1 1 1 ? 1 1 1 1 2 1 1 1 ? 1 2 1 1 1 2 2 1 1 1 1 1 1 2 

Pogonella minutus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 1 1 2 1 1 1 2 1 1 1 1 1 1 

Pogon incurvatum 1 2 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 1 2 2 2 2 1 2 1 1 1 1 1 2 

Pogonotus indicus 1 2 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 1 1 2 2 2 1 2 1 1 ? ? ? ? 

Pogonotypellus australis 1 1 1 1 1 1 1 2 1 3 1 2 1 1 1 ? 1 1 1 2 1 1 2 1 1 ? 1 1 1 1 

Pogontypus 1 2 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 1 1 2 2 2 1 2 1 1 1 1 1 2 

Polonius froggatti 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 1 1 2 1 1 1 2 1 1 1 1 1 1 

Promitor nominatus 1 ? 1 1 1 1 2 2 1 1 2 2 1 1 2 1 2 ? 2 1 1 1 2 2 3 5 1 1 1 2 

Protinotus doddi 1 1 1 1 ? ? 2 2 1 2 1 2 1 1 1 ? 1 2 ? 2 1 1 1 2 1 1 ? ? ? ? 

Psilocentrus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 1 1 1 1 2 2 

Pyrgauchenia 1 1 1 ? 1 1 1 1 2 1 1 1 ? 2 1 ? 1 1 1 3 1 1 ? ? 1 1 2 1 1 1 

Pyrgonota 1 1 1 1 1 1 2 1 1 1 1 2 1 1 1 ? 1 1 1 2 1 1 1 2 1 1 1 1 1 1 

Rachinotus marshalli 1 1 1 1 1 1 1 2 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 2 5 1 1 1 2 

Rentzia 1 1 1 ? 1 1 1 2 1 3 1 2 1 1 1 ? 1 1 1 2 1 2 1 2 1 1 ? ? ? ? 

Rexicornia elegans 1 ? 1 1 1 1 2 2 1 2 2 2 1 1 2 1 2 ? 2 1 1 1 2 2 3 5 1 1 1 2 

Rigula 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 1 1 2 1 1 1 1 1 1 ? ? ? ? 

420


6 6 6 6 6 6 6 6 6 7 7 7 7 7 7 7 7 7 7 8 8 8 8 8 8 8 8 8 8 9 

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 

Occator erectus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Ophicentrus notandus 2 1 1 1 1 1 ? 2 1 2 1 1 ? 3 2 2 1 1 2 2 1 2 1 1 ? 1 2 ? ? ? 

Orekthophora cornuta 1 ? 1 1 1 1 ? 2 1 2 2 1 1 3 2 2 1 1 2 1 3 2 2 1 ? 3 2 ? ? ? 

Orthobelus 1 ? 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 1 1 2 1 3 2 2 1 ? 2 2 1 ? ? 

Otinotoides 1 ? 1 1 1 1 ? 2 1 2 1 1 1 2 2 2 3 1 2 1 1 ? 1 1 ? ? 1 1 ? ? 

Otinotus ammon 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 1 1 2 1 1 2 1 1 ? 1 2 1 ? ? 

Otinotus bantuantus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 1 1 1 4 1 1 

Oxyrhachis 1 ? 1 1 1 1 ? 1 ? 1 ? 1 1 ? 1 ? ? 2 2 1 1 2 3 1 ? 1 1 2 ? ? 

Oxyrhachis carinata 1 ? 1 1 1 1 ? 1 ? 1 ? 1 1 2 1 1 ? 2 2 1 1 2 3 1 ? 1 1 2 ? ? 

Oxyrhachis delalandei 1 ? 1 1 1 1 ? 1 ? 1 ? 1 1 2 1 2 1 2 2 1 1 2 3 1 ? 1 1 2 ? ? 

Oxyrhachis sulcicornis 1 ? 1 1 1 1 ? 1 ? 1 ? 1 1 2 1 2 1 2 2 1 1 2 3 1 ? 1 1 2 ? ? 

Pantaleon 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 2 2 2 2 1 2 1 1 ? ? 

Paracentronodus 2 1 1 1 1 2 3 2 1 2 1 1 1 3 2 2 1 1 1 1 1 ? 1 1 ? 1 1 4 ? ? 

Paradarnoides 1 ? 1 1 1 1 ? 2 1 2 2 1 1 3 2 2 1 1 2 2 1 2 2 1 ? 3 2 1 ? ? 

Parapogon kandyiana 2 1 1 1 1 1 ? 2 1 2 1 ? 1 3 2 2 1 1 2 1 1 2 1 1 ? 1 2 1 ? ? 

Paraxiphopoeus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 1 1 2 1 1 2 1 2 2 2 1 3 2 2 

Parayasa 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 1 ? 1 1 1 ? ? 

Peltzerella borneensis 2 1 1 1 1 1 ? 2 1 2 1 ? 1 3 2 2 3 1 2 ? ? 2 ? ? ? 2 ? ? ? ? 

Periaman 2 1 1 1 1 1 ? 2 1 2 1 3 3 3 2 2 3 1 2 1 3 2 1 1 ? 2 1 4 1 2 

Pieltainellus 1 ? 1 1 1 1 ? 2 1 2 2 1 1 3 2 2 1 1 1 1 1 2 1 1 ? 1 2 1 ? ? 

Platybelus flavus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 2 1 1 3 2 1 

Platycentrus acuticornis 1 ? 1 1 1 1 ? 2 1 2 2 3 3 2 2 2 3 1 1 1 1 2 1 1 ? 1 2 5 1 1 

Pogonella minutus 1 ? 1 1 1 1 ? 2 1 2 1 1 1 2 2 2 1 1 2 1 1 1 1 1 ? 1 1 3 1 1 

Pogon incurvatum 2 1 1 1 1 1 ? 2 1 2 1 ? 1 3 2 2 1 1 2 1 1 2 1 1 ? 1 2 3 1 1 

Pogonotus indicus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Pogonotypellus australis 1 ? 1 1 1 1 ? 1 ? 1 ? ? 1 2 1 2 1 1 2 1 1 ? 1 1 ? ? 1 3 1 1 

Pogontypus 2 1 1 1 1 1 ? 2 1 2 1 ? 1 3 2 2 1 1 2 1 1 2 1 1 ? 1 2 1 ? ? 

Polonius froggatti 1 ? 1 1 1 1 ? 2 1 2 1 ? 1 2 2 2 1 ? 2 ? ? ? ? ? ? 1 ? ? ? ? 

Promitor nominatus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 3 2 1 2 2 1 1 3 1 1 

Protinotus doddi ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Psilocentrus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 1 1 2 1 2 2 1 1 ? 1 2 1 ? ? 

Pyrgauchenia 1 ? 1 1 1 1 ? 1 ? 1 ? 1 1 2 1 1 ? 2 2 1 1 ? 1 1 ? 1 1 2 ? ? 

Pyrgonota 1 ? 1 1 1 1 ? 2 1 1 ? 1 1 ? 1 2 1 1 2 1 1 ? 1 1 ? 1 1 3 ? ? 

Rachinotus marshalli 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 3 2 1 2 2 2 1 3 2 2 

Rentzia ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 2 ? 1 ? ? ? ? ? ? ? ? ? ? ? ? 

Rexicornia elegans 2 1 1 1 1 1 ? 2 1 2 1 ? 1 3 2 2 3 1 2 ? ? ? ? ? ? 2 ? ? ? ? 

Rigula ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 2 ? 1 ? ? ? ? ? ? ? ? ? ? ? ? 

421


1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 

9 9 9 9 9 9 9 9 9 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 

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 

Occator erectus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Ophicentrus notandus ? ? ? ? ? ? ? 1 3 2 1 1 3 2 5 3 2 1 1 1 1 3 2 ? ? ? 

Orekthophora cornuta ? ? ? ? ? ? ? 1 3 1 ? ? 1 1 2 1 1 1 3 ? 1 2 2 ? ? ? 

Orthobelus 1 2 3 1 1 1 2 1 3 1 ? ? 1 1 2 1 2 1 2 1 1 1 2 1 1 3 

Otinotoides 1 1 2 2 1 1 1 1 3 2 2 1 1 1 6 2 2 1 1 1 2 3 2 1 1 3 

Otinotus ammon 1 2 2 1 2 1 2 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Otinotus bantuantus 2 1 2 2 1 1 2 1 3 ? ? ? ? ? 7 3 2 1 1 2 2 2 2 ? ? ? 

Oxyrhachis 1 1 2 ? 1 1 ? 1 3 2 1 ? 1 1 5 3 2 1 1 1 1 ? 2 1 2 1 

Oxyrhachis carinata 1 1 2 2 1 1 2 1 3 2 1 2 1 1 5 3 2 1 1 1 1 2 2 ? ? ? 

Oxyrhachis delalandei 1 1 2 1 1 1 2 1 3 2 1 1 1 1 5 3 2 1 1 1 1 2 2 ? ? ? 

Oxyrhachis sulcicornis 1 1 2 2 1 1 ? 1 3 2 1 2 1 1 5 3 2 1 1 1 1 2 2 ? ? ? 

Pantaleon 1 2 1 ? 2 1 1 1 3 2 1 1 1 2 1 1 1 1 3 1 1 1 2 2 2 3 

Paracentronodus 2 1 2 2 1 1 1 1 2 1 ? ? 1 1 1 1 1 1 3 1 1 1 1 ? ? ? 

Paradarnoides 1 1 3 2 1 1 1 1 3 2 1 1 1 1 2 1 1 1 3 1 1 1 2 ? ? ? 

Parapogon kandyiana 1 1 2 2 2 1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? 1 ? 2 ? ? ? 

Paraxiphopoeus 1 2 1 ? 2 1 1 1 3 2 1 1 1 1 3 3 2 1 2 1 1 2 2 ? ? ? 

Parayasa 1 2 2 1 1 1 1 1 3 2 2 1 1 1 3 3 2 1 3 1 1 1 2 ? ? ? 

Peltzerella borneensis ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Periaman 2 1 2 2 1 1 2 1 3 1 ? ? 1 1 3 3 2 1 1 1 1 2 2 ? ? ? 

Pieltainellus 1 2 2 1 2 1 1 1 3 2 1 2 1 1 3 1 2 1 3 1 1 1 2 1 1 2 

Platybelus flavus 1 1 2 1 1 1 1 1 3 2 1 1 1 1 3 3 2 1 2 1 1 1 2 2 2 3 

Platycentrus acuticornis 1 1 2 2 1 1 2 1 3 2 1 1 1 1 4 1 2 1 3 1 1 1 2 1 1 2 

Pogonella minutus 1 1 3 2 1 1 2 1 3 2 1 1 1 1 6 2 2 1 1 1 2 3 2 1 2 3 

Pogon incurvatum 1 1 2 1 1 1 2 ? ? ? ? ? ? ? ? ? ? ? ? ? 1 ? 2 ? ? ? 

Pogonotus indicus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Pogonotypellus australis 2 1 2 2 1 1 2 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Pogontypus 1 2 2 2 2 1 2 ? ? ? ? ? ? ? 7 3 2 2 1 3 1 2 2 2 2 3 

Polonius froggatti ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Promitor nominatus 2 1 2 2 1 1 2 1 3 2 2 1 1 1 3 3 2 1 2 1 1 1 2 ? ? ? 

Protinotus doddi ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Psilocentrus 1 2 2 1 2 1 1 1 3 2 1 1 3 2 5 3 2 1 1 1 1 2 2 ? ? ? 

Pyrgauchenia 1 1 1 ? 1 2 1 2 2 2 2 1 1 1 5 3 2 1 1 1 1 1 3 1 2 2 

Pyrgonota 2 1 2 2 1 1 1 1 3 2 1 1 1 1 6 2 2 1 1 3 2 3 3 2 2 3 

Rachinotus marshalli 1 2 2 2 1 1 2 1 3 2 2 1 1 1 3 3 2 1 2 1 1 1 2 2 2 3 

Rentzia ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Rexicornia elegans ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Rigula ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

422


1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 3 

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 

Sarantus wallacei 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 2 1 1 2 2 1 2 1 2 1 

Sarantus nobilus 1 2 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 2 1 1 2 2 1 2 1 1 1 

Sertorius 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 2 1 1 2 2 1 2 1 2 1 

Sextius 1 1 2 1 2 2 1 2 1 2 1 2 2 2 3 1 1 2 2 1 2 1 1 2 2 1 2 2 2 1 

Sinodemanga 1 1 2 ? 2 2 1 2 1 1 1 1 1 1 1 ? 1 2 2 1 ? ? ? ? 2 ? 2 1 2 1 

Sipylus 1 1 2 1 3 2 1 2 1 1 1 2 2 1 1 1 1 2 2 1 2 1 1 1 2 1 2 1 2 1 

Spalirises rugosa 1 1 1 2 2 2 1 2 1 2 2 1 2 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Spathenotus tridentatus 1 1 1 ? 2 1 1 2 1 2 1 2 ? 3 ? 1 1 2 ? ? ? 1 ? ? ? ? 2 2 ? 1 

Spathocentrus intermedius 1 1 1 1 2 2 1 2 1 1 2 1 2 1 3 1 1 2 2 1 1 1 1 1 2 2 2 1 2 1 

Spinodarnoides typus 1 1 1 2 2 1 1 2 1 2 1 2 ? 3 1 1 1 2 2 1 2 1 1 1 2 1 2 1 1 1 

Stalobelus 1 1 1 2 2 2 1 2 1 2 2 1 1 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Streonus tenebrosus 1 1 1 2 2 2 1 2 1 2 1 1 1 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Strzeleckia montanus 1 1 2 1 2 2 1 2 1 2 1 2 ? 1 1 ? 1 2 ? ? ? 1 ? ? 2 ? 2 1 2 1 

Subrincator tonkinensis 1 1 1 1 3 2 1 2 1 1 1 2 2 1 3 1 1 2 2 1 2 1 1 1 2 1 2 1 2 1 

Takliwa carteri 1 1 1 2 2 2 1 2 1 2 2 1 2 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Telingana 1 1 1 1 2 1 1 2 1 2 1 1 1 1 3 1 1 2 2 1 1 2 1 1 2 1 2 1 3 1 

Terentius convexus 1 1 2 1 2 2 1 2 1 2 1 2 2 1 1 1 1 2 2 1 2 1 1 1 2 1 2 1 3 1 

Thelicentrus xizangensis 1 1 ? ? ? ? ? 2 1 1 1 2 ? 1 3 ? 1 2 ? ? ? ? ? ? 2 ? 2 1 2 1 

Tiberianus 1 1 1 1 2 2 1 2 1 1 1 1 2 1 2 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Tolania 2 1 1 1 2 2 1 1 ? ? ? ? ? ? 3 1 1 1 3 ? 1 1 1 1 1 1 1 1 2 2 

Tribulocentrus zhenbaensis 1 1 1 ? ? 2 ? 2 1 1 1 2 2 1 3 ? 1 2 ? ? ? ? ? ? 2 1 2 1 2 1 

Tricentroides orcus 1 1 1 1 3 2 1 2 1 1 1 2 2 1 3 1 1 2 2 1 2 1 1 1 2 1 2 1 ? 1 

Tricentrus 1 1 2 1 3 2 1 2 1 1 1 2 2 1 2 1 1 2 2 1 2 1 1 1 2 1 2 1 3 1 

Tricoceps 1 1 1 1 2 2 1 2 1 2 2 1 1 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 3 1 

Trioxiphus 1 1 2 1 2 2 1 2 1 2 1 1 2 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Truncatocornum sp. 1 1 2 1 2 2 1 2 1 1 1 1 2 1 3 1 1 2 2 1 1 2 1 1 2 2 2 1 2 1 

Tsunozemia paradoxa 1 1 2 1 3 2 1 2 1 1 1 2 2 1 3 1 1 2 2 1 2 1 1 1 2 1 2 1 2 1 

Tylocentrus 1 1 1 1 2 2 1 2 1 1 1 2 2 1 2 1 1 2 2 1 2 1 1 1 2 1 2 1 2 1 

Tyrannotus tyrannicus 1 1 2 1 2 2 1 2 1 2 2 1 1 1 2 1 1 2 2 1 1 2 1 1 2 1 2 1 2 1 

Umfilianus declivis 1 1 2 1 2 2 1 2 1 ? 1 1 2 1 2 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Undarella storeyi 1 1 2 1 2 2 1 2 1 2 ? ? 2 1 3 1 1 2 2 1 2 1 1 2 2 1 2 1 2 1 

Uroxiphus maculiscutum 1 1 2 1 2 2 1 2 1 2 1 1 2 1 1 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Vecranotus sinuatus 1 1 1 2 2 2 1 2 1 2 1 1 1 1 3 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Xanthosticta pygmaea 1 1 2 1 3 2 1 2 1 1 1 2 2 1 1 1 1 2 2 1 2 1 1 1 2 1 2 1 3 1 

Xiphopoeus 1 1 2 1 2 2 2 2 1 2 2 1 1 1 3 2 1 2 2 1 1 1 1 2 2 2 2 1 2 1 

Yangupia occidentalis 1 1 2 1 2 2 1 2 1 2 1 2 2 1 3 1 1 2 2 1 2 1 1 2 2 1 2 2 ? 1 

Yaponotus villiersi 1 1 2 1 2 2 1 2 1 2 2 1 1 1 1 1 1 2 2 1 1 1 1 1 2 1 2 1 2 1 

Yasa greeni 1 1 2 1 3 2 1 2 1 1 1 2 2 1 1 1 1 2 2 1 2 1 1 1 2 1 2 1 3 1 

Zanzia vanderplanki 1 1 1 2 2 2 1 2 1 1 2 1 2 1 4 1 1 2 2 1 1 1 1 1 2 1 2 1 3 1 

Zigzagicentrus bannaensis 1 1 2 ? 2 2 1 2 1 2 2 1 ? 1 3 ? 1 2 ? ? ? ? ? ? 2 ? 2 1 2 1 

423


3 3 3 3 3 3 3 3 3 4 4 4 4 4 4 4 4 4 4 5 5 5 5 5 5 5 5 5 5 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 

Sarantus wallacei 1 1 1 2 1 1 1 2 1 2 1 2 1 1 1 ? 1 2 1 2 1 2 ? 1 1 2 1 1 1 1 

Sarantus nobilus 1 1 1 2 1 1 1 2 1 3 1 2 1 1 1 ? 1 ? 1 2 1 2 2 1 1 1 1 1 1 1 

Sertorius 1 1 1 1 1 1 1 2 1 3 1 2 1 1 1 ? 1 1 1 2 1 1 1 2 1 1 1 1 1 1 

Sextius 1 1 1 1 2 1 1 1 1 1 1 1 1 2 1 ? 1 1 1 2 1 1 1 ? 2 1 2 1 1 1 

Sinodemanga 1 1 1 1 1 1 2 2 1 ? 1 2 1 1 1 ? 1 3 2 1 1 1 2 1 1 1 ? ? ? ? 

Sipylus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 2 1 2 2 1 1 1 2 2 3 5 1 1 1 2 

Spalirises rugosa 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 2 5 1 1 1 2 

Spathenotus tridentatus 1 1 1 1 ? 1 2 1 ? ? 1 1 1 1 ? ? ? ? 1 2 1 2 1 1 1 1 ? ? ? ? 

Spathocentrus intermedius 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 3 2 1 1 1 2 2 1 1 1 1 1 1 

Spinodarnoides typus 1 2 1 1 1 1 2 1 1 2 1 2 1 1 2 1 1 1 1 1 1 1 2 1 1 1 1 1 1 1 

Stalobelus 1 1 1 1 1 1 1 2 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 2 5 1 1 1 2 

Streonus tenebrosus 1 1 1 1 1 1 2 2 1 2 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 2 5 1 1 1 2 

Strzeleckia montanus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 2 1 2 1 1 1 2 1 1 ? ? ? ? 

Subrincator tonkinensis 1 1 1 1 1 1 2 2 1 2 1 2 1 1 1 ? 1 2 2 1 1 1 2 2 3 5 1 1 1 2 

Takliwa carteri 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 2 5 1 1 1 2 

Telingana 1 2 1 1 1 2 2 2 1 1 1 2 1 1 1 ? 1 1 ? 2 1 1 1 2 1 1 1 1 1 2 

Terentius convexus 1 1 1 1 1 1 1 2 1 3 1 2 1 1 1 ? 1 2 1 2 1 1 1 1 1 1 1 1 1 1 

Thelicentrus xizangensis 1 1 1 1 1 1 2 2 1 ? 1 2 1 1 2 2 1 2 2 1 1 1 2 1 1 5 ? ? ? ? 

Tiberianus 1 1 1 1 1 1 2 ? 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 1 5 1 1 1 2 

Tolania 1 1 2 2 1 2 2 2 1 1 1 2 1 1 1 ? 1 2 ? ? 1 1 ? ? ? 1 1 1 2 2 

Tribulocentrus zhenbaensis 1 1 1 1 1 1 2 2 1 ? 1 2 1 1 1 ? 1 2 2 1 1 1 1 1 3 5 ? ? ? ? 

Tricentroides orcus 1 1 1 1 1 1 2 2 1 2 1 2 1 1 2 2 1 2 2 1 1 ? 2 1 3 5 1 1 1 2 

Tricentrus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 2 1 2 2 1 1 1 2 2 3 5 1 ? 1 2 

Tricoceps 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 2 5 1 1 1 2 

Trioxiphus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 ? ? 1 2 1 2 1 2 1 1 1 1 1 1 1 2 

Truncatocornum sp. 1 1 1 1 1 1 2 2 1 2 1 2 1 1 1 ? 1 3 1 2 1 1 2 1 1 1 1 1 1 2 

Tsunozemia paradoxa 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 2 2 1 1 1 2 1 2 5 1 ? ? 2 

Tylocentrus 1 1 1 1 1 1 2 1 1 1 1 2 1 2 1 ? 1 3 1 1 1 1 1 2 1 1 1 1 1 2 

Tyrannotus tyrannicus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 1 1 2 1 1 1 1 1 1 1 1 1 2 

Umfilianus declivis 1 1 1 1 1 1 2 1 1 2 1 2 1 1 1 ? 1 3 2 2 1 2 2 1 1 1 1 1 1 2 

Undarella storeyi 1 1 1 1 1 1 2 1 1 1 1 1 1 1 1 ? 1 1 1 2 1 1 1 2 1 1 1 1 1 1 

Uroxiphus maculiscutum 1 1 1 1 1 1 1 2 1 1 1 1 1 1 1 ? 1 3 1 2 1 2 2 1 1 1 1 1 1 2 

Vecranotus sinuatus 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 1 2 ? 2 1 1 1 2 2 3 5 1 1 1 2 

Xanthosticta pygmaea 1 1 1 1 1 1 2 2 1 1 1 2 1 1 2 2 1 2 2 1 1 1 2 2 3 5 1 1 1 2 

Xiphopoeus 1 1 1 1 1 1 2 1 1 1 1 2 1 1 2 1 1 1 1 2 1 1 2 1 1 1 1 1 1 2 

Yangupia occidentalis 1 1 1 1 1 1 2 2 1 3 1 2 1 1 1 ? 1 1 1 2 1 1 1 1 1 1 1 1 1 1 

Yaponotus villiersi 1 1 1 1 1 1 2 2 1 1 1 2 1 1 1 ? 1 2 1 2 1 2 2 1 1 1 1 1 1 2 

Yasa greeni 1 ? 1 1 1 1 2 2 2 1 2 2 1 1 ? ? 1 3 2 1 1 1 2 2 ? 1 1 1 1 2 

Zanzia vanderplanki 1 1 1 1 1 1 2 ? 1 1 1 2 1 1 1 ? 2 ? 2 1 1 1 2 2 2 5 1 1 1 2 

Zigzagicentrus bannaensis 1 1 1 1 1 1 2 2 1 ? 1 2 1 1 1 ? 1 3 1 2 1 1 1 1 1 1 ? ? ? ? 

424


6 6 6 6 6 6 6 6 6 7 7 7 7 7 7 7 7 7 7 8 8 8 8 8 8 8 8 8 8 9 

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 

Sarantus wallacei 1 ? 1 1 1 1 ? 2 1 2 1 1 1 2 1 2 1 1 2 1 1 1 1 1 ? 1 1 3 1 2 

Sarantus nobilus 1 ? 1 1 1 1 ? 2 1 2 1 ? 1 2 2 2 1 1 2 1 1 ? 1 1 ? ? 1 4 1 1 

Sertorius 1 ? 1 1 1 2 2 2 1 2 1 1 1 2 2 2 3 1 2 1 1 2 1 1 ? 2 1 1 ? ? 

Sextius 1 ? 1 1 1 1 ? 1 ? 1 ? 1 1 1 1 2 1 1 2 1 1 1 1 1 ? 1 1 3 2 2 

Sinodemanga ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Sipylus 2 1 1 3 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 3 2 2 2 1 2 1 1 ? ? 

Spalirises rugosa 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 2 1 1 3 1 1 

Spathenotus tridentatus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 3 ? ? ? ? 

Spathocentrus intermedius 1 ? 1 1 1 1 ? 2 1 2 2 ? 1 3 2 2 1 1 2 1 1 2 1 1 ? 1 2 1 ? ? 

Spinodarnoides typus 1 ? 1 1 1 1 ? 2 1 2 1 ? 1 ? 2 2 1 1 2 ? ? 2 ? ? ? 3 2 ? ? ? 

Stalobelus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 2 2 1 3 1 1 

Streonus tenebrosus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 2 2 1 1 ? ? 

Strzeleckia montanus ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 2 ? 1 ? ? ? ? ? ? ? ? ? ? ? ? 

Subrincator tonkinensis 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 1 1 2 1 2 2 2 2 1 1 1 1 ? ? 

Takliwa carteri 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 2 1 1 3 1 2 

Telingana 2 1 1 1 1 1 ? 2 1 2 1 ? ? 3 2 2 1 1 2 1 1 2 1 1 ? 1 2 ? ? ? 

Terentius convexus 1 ? 1 1 1 2 2 2 1 2 1 1 1 2 2 2 1 1 2 1 1 2 1 2 1 2 1 1 ? ? 

Thelicentrus xizangensis ? ? 1 1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 ? ? 

Tiberianus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 2 2 1 3 2 1 

Tolania 2 1 1 3 1 2 3 2 1 2 1 1 1 3 2 2 3 1 1 1 1 ? 1 1 ? 1 1 4 1 ? 

Tribulocentrus zhenbaensis ? ? 1 1 1 ? ? ? ? ? ? ? ? ? 2 2 3 1 ? ? ? ? ? ? ? ? ? ? ? ? 

Tricentroides orcus 2 2 1 1 1 1 ? 2 1 2 1 ? 3 3 2 2 3 1 2 1 3 2 2 2 1 2 1 1 ? ? 

Tricentrus 2 1 2 3 2 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 3 2 2 2 1 2 1 1 ? ? 

Tricoceps 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 2 2 1 3 1 1 

Trioxiphus 2 2 1 1 1 1 ? 2 1 2 2 2 3 3 2 2 3 1 2 1 2 2 1 2 1 2 1 4 1 1 

Truncatocornum sp. 2 2 1 1 1 1 ? 2 1 2 2 ? 1 2 2 2 3 1 2 ? ? 2 1 1 ? 1 2 1 ? ? 

Tsunozemia paradoxa 2 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 1 1 2 2 2 1 1 1 1 ? ? 

Tylocentrus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 1 1 2 1 1 2 1 1 ? 1 1 5 1 1 

Tyrannotus tyrannicus 2 1 1 1 1 1 ? 2 1 2 1 3 1 3 2 2 1 1 2 1 2 2 1 1 ? 2 1 4 1 1 

Umfilianus declivis 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 3 2 1 2 1 2 1 4 1 1 

Undarella storeyi 1 ? 1 1 1 1 ? 2 1 2 1 ? 1 2 2 2 1 1 2 ? ? ? ? ? ? 2 1 ? ? ? 

Uroxiphus maculiscutum 2 1 1 1 1 1 ? 2 1 2 1 ? 1 3 2 2 3 1 2 1 1 2 1 2 1 2 1 4 1 1 

Vecranotus sinuatus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 2 1 1 3 2 1 

Xanthosticta pygmaea 2 1 2 1 2 1 ? 2 1 2 1 1 1 3 2 2 1 1 2 1 3 2 2 2 1 1 1 1 ? ? 

Xiphopoeus 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 2 2 3 1 2 1 1 2 1 2 3 3 1 3 2 2 

Yangupia occidentalis 1 ? 1 1 1 1 ? 2 1 2 1 1 ? 2 2 2 3 1 2 1 1 2 1 1 ? 2 1 ? ? ? 

Yaponotus villiersi 2 1 1 1 1 1 ? 2 1 2 1 3 3 3 2 2 3 1 2 1 3 2 1 2 1 2 1 4 1 1 

Yasa greeni 2 1 1 1 1 1 ? 2 1 2 1 ? 1 3 2 2 1 1 2 1 1 2 1 1 ? 1 1 1 ? ? 

Zanzia vanderplanki 2 1 1 1 1 1 ? 2 1 2 1 1 1 3 1 2 3 1 2 1 1 2 1 2 2 2 1 1 ? ? 

Zigzagicentrus bannaensis ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

425


1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 

9 9 9 9 9 9 9 9 9 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 

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 

Sarantus wallacei 2 1 2 1 1 1 2 1 3 2 2 1 1 1 6 2 2 1 1 1 2 3 2 1 2 3 

Sarantus nobilus 2 1 2 2 1 1 2 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Sertorius 1 1 2 1 2 1 2 1 3 2 1 1 1 1 6 2 2 1 1 1 2 3 2 2 2 3 

Sextius 1 2 2 1 1 1 1 1 3 2 2 1 1 1 6 2 2 1 1 1 2 3 2 1 2 3 

Sinodemanga ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Sipylus 1 2 1 ? 2 1 1 1 3 1 ? ? 1 2 1 1 1 1 3 1 1 1 2 ? ? ? 

Spalirises 1 1 2 1 1 1 1 1 3 2 2 1 1 1 3 3 2 1 2 1 1 1 2 ? ? ? 

Spathenotus tridentatus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Spathocentrus intermedius 1 2 2 1 2 1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Spinodarnoides typus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Stalobelus 1 1 2 2 1 1 1 1 3 2 1 1 1 1 3 3 2 1 2 1 1 2 2 ? ? 3 

Streonus tenebrosus 1 2 3 2 2 1 1 1 3 2 1 1 1 1 3 3 2 1 2 1 1 1 2 2 2 3 

Strzeleckia montanus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

Subrincator tonkinensis 1 2 2 1 2 1 1 1 3 2 1 1 1 2 1 1 1 1 3 1 1 1 2 2 2 3 

Takliwa carteri 1 1 1 ? 1 1 1 1 3 2 1 1 1 1 3 3 2 1 2 1 1 2 2 1 2 1 

Telingana ? ? ? ? ? ? ? 1 3 2 2 1 1 1 7 3 2 2 1 3 1 2 2 2 2 3 

Terentius convexus 1 2 2 1 2 1 1 1 3 2 1 1 1 1 6 2 2 1 1 1 2 3 2 2 2 3 

Thelicentrus xizangensis 1 2 2 1 2 ? ? ? 3 2 1 1 1 2 1 ? 1 1 3 ? 1 1 ? ? ? ? 

Tiberianus 1 1 2 1 1 1 1 1 3 2 1 1 1 1 3 3 2 1 2 1 1 1 2 2 2 3 

Tolania 2 1 2 1 1 1 1 1 2 2 2 1 1 ? 1 1 1 1 3 1 1 1 1 2 2 3 

Tribulocentrus zhenbaensis ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 ? 1 1 3 ? 1 1 ? ? ? ? 

Tricentroides orcus 1 2 2 2 2 1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Tricentrus 1 2 1 ? 2 1 2 1 3 1 ? ? 1 2 1 1 1 1 3 1 1 1 2 1 1 3 

Tricoceps 1 1 2 1 1 1 1 1 3 2 1 1 1 1 3 3 2 1 2 1 1 1 2 2 2 3 

Trioxiphus 2 1 2 2 1 1 2 1 3 2 1 1 1 1 7 3 2 1 1 2 2 2 2 2 2 3 

Truncatocornum sp. 1 1 1 ? 1 1 2 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Tsunozemia paradoxa 1 2 1 ? 2 1 2 ? ? ? ? ? ? ? 1 ? 1 1 3 1 1 1 2 ? ? ? 

Tylocentrus 1 1 2 2 1 1 1 1 3 1 ? ? 1 1 2 1 2 1 3 1 1 1 2 1 1 2 

Tyrannotus tyrannicus 2 1 2 2 1 1 2 1 3 2 1 3 1 2 1 1 1 1 3 1 1 1 2 1 1 3 

Umfilianus declivis 2 1 1 ? 1 1 2 1 3 2 1 1 1 1 7 3 2 1 1 2 2 2 2 ? ? ? 

Undarella storeyi ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Uroxiphus maculiscutum 2 1 2 2 1 1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Vecranotus sinuatus 1 1 2 1 1 1 1 1 3 2 2 1 1 1 3 3 2 1 2 1 1 1 2 2 2 3 

Xanthosticta pygmaea 1 2 3 1 1 1 1 1 3 2 2 1 1 2 1 3 1 1 3 1 1 1 2 ? 2 3 

Xiphopoeus 1 2 2 1 2 1 2 1 3 1 ? ? 1 1 3 3 2 1 2 1 1 1 2 2 2 3 

Yangupia occidentalis ? ? ? ? ? ? ? 1 3 2 1 1 1 1 6 2 2 1 1 1 2 3 2 ? ? ? 

Yaponotus villiersi 2 1 2 2 1 1 2 1 3 2 1 1 1 1 7 3 2 1 1 2 2 2 2 ? ? ? 

Yasa greeni 1 2 3 2 2 1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? ? ? 

Zanzia vanderplanki 1 2 3 1 2 1 1 1 3 2 1 1 1 1 3 3 2 1 2 1 1 1 2 ? ? ? 

Zigzagicentrus bannaensis ? ? ? ? ? ? ? ? 3 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 

426

Table 24.14. Summary of taxonomic changes 

based on phylogenetic analyses 1-8. 

Existing classification (Dietrich et al. 2001a, 

Yuan and Chou 2002a) 

CENTRODONTINAE 

ABELINI 

BOOCERINI 

BULBAUCHENIINI 

FUNKHOUSERELLINI 

TERENTIINI 

ANTIALCIDINI 

COCCOSTERPHINI 

MADLININI 

TRICENTRINI 

GARGARINI 

DEMANGINI 

LEPTOCENTRINI 

CENTROCHARESINI 

CENTROTINI 

CENTROTYPINI 

CHOUCENTRINI 

EBHULINI 

HYPSAUCHENIINI 

LEPTOBELINI 

MICREUNINI 

NESSORHININI 

OXYRHACHINI 

PLATYCENTRINI 

XIPHOPOEINI 

427 

Revised classification presented here 

Centrotinae: 

Centrodontini 

Boocerini 

Terentiini 

Gargarini 

Leptocentrini 

Ebhuloidesini 

Centrocharesini 

Centrotini 

Centrotypini 

Choucentrini 

Hypsaucheniini 

Leptobelini 

Micreunini 

Nessorhinini 

Oxyrhachini 

Platycentrini 

Xiphopoeini 

Lobocentrini 

Maarbarini 

Monobelini 

Pieltainellini 

Beaufortianini 

Boccharini 

moved to 

Centrotinae 

new 

synonymies 

names 

unchanged 

new tribes

25: BIOGEOGRAPHY OF THE CENTROTINAE AND MEMBRACIDAE 

Introduction: Origins of the Membracidae 

Where and when did treehoppers originate? This question is an outstanding and 

controversial topic for three reasons: (1) the current geographic distribution of treehoppers, 

with one large cosmopolitan subfamily, but the eight other subfamilies all restricted to the 

New World; (2) the limited fossil record of treehoppers; and, (3) the lack of phylogenetic 

understanding for the Membracidae (only recently elucidated) and the Centrotinae 

(elucidated in this work). Following a review of our knowledge in these three areas, the 

biogeographic patterns within the subfamily Centrotinae are, for the first time, placed in a 

phylogenetic context. 

Geographic patterns. The current distribution of major treehopper lineages is 

intriguing. Except for a few introductions by man--Spissistilus to Hawaii (Zimmerman 

1948a); Centrotus and Gargara to North America (Metcalf and Wade 1965a, McKamey 

1998a); and Stictocephala to Europe (Goidanich 1946a, McKamey 1998a)--no membracid 

tribe (and thus no species or genus) occurs in both the Old World and the New. 

Nevertheless, the largest membracid subfamily, Centrotinae, occurs in all major regions of 

the Old World (The Afrotropical, Australasian/Oceanian, Indomalayan, and Palearctic 

Regions) as well as the New World (Nearctic and the Neotropical Regions). All of the other 

eight membracid subfamilies occur only in the New World. This overall pattern lead Wood 

(1993) to believe that treehoppers arose in tropical Gondwana prior to its breakup, with the 

Old World and New World membracids diversifying after the continents separated. 

Similarly, Strümpel (1972a) postulated that the membracids arose in Gondwana, but that the 

428

Centrotinae reached the New World from Asia by way of the Bering Land Bridge in the 

Pleistocene. 

In contrast, the center of origin theory, a controversial method based solely on 

distribution (Futuyma 1998a), suggests a New World origin of the family. Areas of origin 

include regions that contain the largest number of species and the most morphological 

diversity (Cranston and Naumann 1991a). South Asia, for example, is believed to be the 

center of origin for vespid wasps (Hymenoptera: Vespidae) because it is the only area where 

all the subfamilies are found (Briggs 1995a). With treehoppers, the Neotropical Region 

contains more species, genera, tribes, and subfamilies than any other region (McKamey 

1998a) and thus is arguably the most morphologically diverse, and, under the center of origin 

theory, the most likely area of origin. Nonetheless, this method can not distinguish between 

primary and secondary areas of diversification. 

Fossils. While a number of hemipteran families have a relatively rich fossil record 

(Labandeira and Seposki 1993a), the only known fossils assignable to the family 

Membracidae with certainty are undescribed representatives of the subfamily Stegaspidinae 

from amber of the Dominican Republic (Schlee 1990a, Poinar 1992a, McKamey 1998a). 

Dominican amber was deposited during the Eocene-Miocene, 57.8 to 5.3 million years ago 

(mya), long after the vicariance of Gondwana. Perhaps further membracid fossils await 

discovery, but meanwhile, the limited records provide a minimum age of 57.8 to 5.3 million 

years for the Stegaspidinae, and suggest a Tertiary origin for Membracidae, perhaps in the 

New World. 

Phylogeny. Recent phylogenetic studies provide a broad outline of evolutionary 

relationships among New World membracids, but phylogeny within the cosmopolitan 

429

subfamily Centrotinae--so critical to addressing biogeographical patterns among treehoppers- 

-was largely neglected until the present study. Both morphological and molecular data (Deitz 

and Dietrich 1993a, Dietrich and Deitz 1993a, Cryan et al. 2000a, Dietrich et al. 2001a) 

suggest that the Membracidae arose in the Neotropics, but the time of origin is difficult to 

pinpoint with neither a substantial fossil record nor divergence times calculated from 

molecular data. Phylogenetic analyses have consistently placed four subfamilies as the first 

membracid lineages. These groups--Endoiastinae, Stepaspidinae, Nicomiinae, and 

Centronodinae--are largely confined to the Neotropics. 

To summarize, evidence from phylogeny, fossils, and geography (region of greatest 

morphological diversity) all favor a Neotropical origin for the family Membracidae. 

Furthermore, fossils provide a minimum age of 57.8 to 5.3 million years for the 

Stegaspidinae (one of four basal membracid lineages), pointing toward a Tertiary origin for 

the family. 

Biogeographic patterns of the Centrotinae 

The Centrotinae account for roughly half of all treehopper diversity at the tribal, 

generic, and species levels. As noted above, centrotines are found in all major 

zoogeographic regions. While a few tribes are widely distributed, many occur primarily in 

one or two major regions. The distinctive fauna of the Afrotropical, Indomalayan, 

Australasian/Oceanian, and Caribbean Regions are especially notable. Indeed, all but one of 

the Old World centrotine tribes have representative genera found in the Indomalayan Region 

(Fig. 25.1). 

430

How did treehoppers acquire their Old World distribution if it is assumed that 

membracids arose in the Tertiary Neotropics, when the southern continents had already 

drifted far apart? If the centrotine distribution were to be explained by vicariance, their 

phylogeny should, to some extent, reflect the historical splitting of the land masses and their 

historical proximity to one another. The geologic history of the continents followed here is 

based on McLoughlin (2001a). North America and Africa separated roughly 180-165 mya. 

Madagascar + India separated from Africa approximately 165 mya, and from Antarctica and 

Australia about 132 mya. Africa diverged from South America 135-105 mya and India and 

Madagascar separated from each other 95-84 mya. By the early Tertiary (65 mya), New 

Zealand was widely separated from Australia. South America, Australia, and Antarctica 

were more or less connected from 60-35 mya. 

The evolutionary relationships of the Centrotinae (Fig. 25.1) do not coincide with the 

historical relationships of the continents. Centrotines found in former Gondwanan regions 

such as Africa, Australia, and South America, are not closely related (Fig. 25.1). The 

Terentiini, placed relatively basally in the phylogeny (Fig. 25.1) and primarily Australasian 

in distribution, show no close relationship with South American centrotines. Aside from a 

single record of the genus Gargara (Gargarini) (Day 1999a), the only tribe on continental 

Australia is the Terentiini. Terentiines are the sister group to a primarily Palearctic and 

Indomalayan clade of centrotines, Ebhuloidesini + Oxyrhachini + Hypsaucheniini. 

Furthermore, the immediate basal lineage of the Terentiini are the Centrocharesini, found in 

the Palearctic and Indomalayan Regions (Fig. 25.1). With the exception of the 

Centrodontini, the only tribe with South American elements, the Boocerini, are the sister 

group of the Gargarini, which are predominantly Palearctic and Indomalayan in distribution. 

431

The highly derived (Fig. 25.1) and primarily Afrotropical tribes Centrotini and Xiphopoeini 

are distantly related to the Australasian Terentiini and to South American centrotines. 

Additionally, no centrotines are apparently native to Madagascar or New Zealand 

based on extensive collecting in recent times (Capener 1968a; Eyles 1970a, 1971a; Wise 

1977a), suggesting either that they went extinct on both islands or perhaps that these islands 

were isolated before treehoppers reached the Old World. Had treehoppers originated before 

the breakup of South America and Africa, however, they would likely have colonized both 

New Zealand and Madagascar. Moreover, centrotines of several tribes have been described 

from India. If treehoppers were present in Gondwana prior to its breakup, they would likely 

be present in Madagascar also, because of its historical proximity to India up to 85 mya. 

Treehoppers likely invaded India in the Cenozoic either when it collided with Asia 

approximately 45 mya ago (McLoughlin 2001a) or when it is thought to have briefly 

contacted Africa (Hedges 2001a). The cuckoo wasps (Hymenoptera: Chrysididae) show a 

similar relationships to centrotines. No higher chrysidid taxa found on different Gondwanan 

continents are closely related, and thus it is believed that the cuckoo wasps evolved following 

the Gondwanan breakup (Briggs 1995a). 

Based on the phylogeny presented here (Fig. 24.1, Fig. 25.1), centrotines originated in 

the New World, dispersed to the Old World twice, and subsequently underwent explosive 

radiations. All suitable outgroups for the Centrotinae (Fig. 24.1) are from the New World. 

Furthermore, the first centrotines are apparently from the New World, ancestors of the 

Centrodontini (Fig. 25.1), a disjunct tribe located in South and North America. Apparently, 

one centrotine invasion of the Old World eventually gave rise to the predominantly 

Indomalayan and Palearctic Gargarini--the other invasion gave rise to most of the remaining 

432

centrotine species, with more than half distributed in the Afrotropical and the 

Australasian/Oceanian Regions. Although the first Old World invasion (Fig. 24.1: node 124, 

Gargarini) is supported by 4 character changes, the second (Fig 24.1: node 111) is supported 

by only 2 character changes. 

Centrotinae biogeographical patterns and scenarios 

Biogeographical patterns of New World centrotines. The first centrotines were from 

the New World, ancestors of the Centrodontini (Fig. 25.1), a disjunct tribe located in South 

and North America. The genera Multareis, Multareoides, and Centrodontus are confined to 

creosote bush in the southwestern United States and northern Mexico, the apparent center of 

diversification of the centrodontines, while Nodonica has been found only in the South 

American countries of Brazil, Ecuador, and Peru. Most New World centrotines are located 

in Central America and Mexico (31 spp.), and the Caribbean islands (66 spp.) (McKamey 

1998a, Dietrich et al. 2001a). Just 5 genera and 12 centrotine species, all in the Boocerini 

and Centrodontini, are found in South America. Based on these numbers, the early 

centrotines likely arose in North America and later invaded South America. Nonetheless, 

even though South American centrotines are few, phylogenetic analyses of the Boocerini and 

Centrodontini (Fig. 24.5) do not rule out a South American origin for centrotines. Nodonica, 

the basal genus of Centrodontini, is found in South America, and the basal relationships of 

the Boocerini are unresolved. Ramos (1988a), however, noted a close relationship among 

Central American, Mexican, and Caribbean centrotines. 

Apparently, there were two centrotine invasions to what are now the Caribbean 

Islands from mainland Neotropical centrotines, one eventually giving rise to the Monobelini 

433

and the other giving rise to the Nessorhinini, both tribes endemic to the Caribbean Region 

(Fig. 25.1). The distribution of these centrotines is likely explained by dispersal rather than 

vicariance due to the high endemism of the Caribbean fauna and the absence of nessorhinines 

and monobelines in the mainland Neotropics. The Caribbean membracid fauna would likely 

resemble a cross-section of the mainland Neotropical fauna if there had been an ancient 

vicariance of land masses, with a higher diversity at the generic and subfamily levels. 

Therefore, it is more likely that ancestors of the Centrodontini dispersed into the Caribbean 

Region giving rise to the Monobelini (Fig. 25.1). The same ancestors apparently also gave 

rise to the Boocerini. The common ancestor of the Mexican tribe Platycentrini likely made 

the second invasion into the Antillean region, eventually giving rise to the Nessorhinini (Fig. 

25.1). 

The mechanism for these dispersals is unclear due to the controversial geologic 

history of the Antillean region (Iturralde-Vinent and MacPhee 1999a, Hedges 2001a). It is 

generally agreed that the Caribbean Islands were formed by volcanism as a result of 

subduction of the North American plate beneath the Caribbean plate in the mid-Cretaceous 

(Hedges 2001a). As a result of this event, it is likely that North America and South America 

were connected by a “proto Antillean” land arc or land bridge consisting largely of present 

day Antillean land masses in the late Cretaceous, approximately 70-80 mya. This land bridge 

could have been an avenue of dispersal to the Caribbean islands. Researchers disagree on the 

timing and permanency of this land bridge and which islands were continuously above water. 

If this was a mechanism for dispersal, however, why didn’t more centrotine higher level taxa 

disperse to these areas? One possibility is that the ancestral centrotines were isolated in 

various areas of North America. 

434

Another possibility is an early Tertiary dispersal following the extraterrestrial impact 

believed responsible for massive terrestrial and marine extinctions 65 mya. At that time, the 

continuous land bridge connecting North and South America was beginning to dissolve 

leaving an island archipelago (Briggs 1995a). Dispersal via more temporary land bridges, 

rafting, or other means, to these newly formed islands would better explain the endemic 

Caribbean centrotine fauna. While a permanent land bridge would permit more animals to 

disperse, island chains would act as filter, allowing some taxa to colonize while acting as a 

barrier to others (Hedges 2001a). Vertebrate fauna are also low in taxonomic diversity at 

higher levels in the Caribbean but some genera have a large number of species, similar to 

centrotines (Hedges 2001a). Many researchers believe that approximately 65 mya, at the 

border of the Cretaceous and Tertiary, an extraterrestrial object impacted the Yucatan 

peninsula of Mexico resulting in a mass extinction of numerous marine organisms, dinosaurs, 

mammals, insects, and plants (Labandeira et al. 2002a). The side effects to the fauna would 

have been widespread, including massive tsunamis and hurricanes, perhaps destroying all 

Antillean fauna (Hedges 2001a). It is possible that any Caribbean centrotines that had 

dispersed via a permanent land bridge prior to the impact would have also been exterminated. 

This would favor a dispersal following the 65 mya impact event. 

Biogeographic patterns of Old World centrotines. Apparently, there were two 

centrotine colonizations of the Old World (Fig. 25.1); one eventually giving rise to the 

predominantly Indomalayan and Palearctic Gargarini and another giving rise to most of the 

remaining centrotine species, with more than half of these distributed in the Afrotropical and 

the Australasian/Oceanian Regions. Interestingly, the most basal Old World groups, the 

tribes Gargarini and Beaufortianini, are also very widely distributed (Fig. 25.1). If the Old 

435

World Centrotinae had reached the Old World by dispersal after the extraterrestrial impact 

(based on the earlier dispersal times to Caribbean and the fossil record), how was this 

accomplished seeing that the Gondwanan continents by this time were well separated? Four 

possible scenarios are considered here. 

Firstly, as a result of a sea level drop in the late Cretaceous, southern South America, 

Antarctica, and Australia were joined from approximately 60 to 35 mya (Briggs 1995a, 

McLoughlin 2001a). This passage has been hypothesized as the Tertiary dispersal route for 

marsupials, birds, reptiles, invertebrates, and angiosperms from southern South America to 

Australia (Briggs 1995a). There is no evidence, however, from current centrotine 

distributions or the present phylogenetic analysis (Fig. 25.1) to support centrotines dispersing 

to the Old World via South America to Australia. The South American centrotines 

(Centrodontini and Boocerini) are not closely related to Australasian centrotines (all 

Terentiini except one record of Gargara) (Fig. 25.1). Additionally, as noted above, most 

New World centrotines are located in Central and North America and the Caribbean, not 

southern South America. This scenario is therefore unlikely for either of the two invasions. 

Dispersal to the Afrotropical Region from the New World is a second possibility. 

Still, there are only a few species of Gargarini in the Afrotropical Region, a primarily 

Indomalayan and Palearctic tribe (Fig. 25.1). The first Old World tribe of the other 

centrotine invasion is the Beaufortianini (Fig. 25.1). Their immediate ancestor is the New 

World tribe Pieltainellini, found in the mainland Neotropics. The tribe Beaufortianini 

consists of Afrotropical and Indomalayan genera but the basal lineage is the genus 

Imporcitor, found in the Palearctic and Indomalayan Regions. It is possible that both 

invasions dispersed from the Africa to India, which is thought to have been joined with 

436

Somalia in northeast Africa in the late Cretaceous (Briggs 1995). Both invasions could have 

colonized the Indomalayan and Palearctic Regions following India’s collision with Asia, 

approximately 45 mya (McLoughlin 2001a). 

The distribution of the gargarine genus Madlinus provides some support for this 

scenario. Madlinus, which is closely related to Coccosterphus, has been found only on the 

Seychelle Islands. The Seychelles were closely associated with India until 65 mya. 

Coccosterphus and its relatives are mostly confined to India. Based on the above scenario, 

the ancestors of Madlinus were subsequently confined to the Seychelles after its split with 

India 65 mya. The presence of Madlinus in the Seychelles is evidence for an ancient 

dispersal of the gargarine lineage to India via Africa, considering the great distance between 

India and the Seychelles today and the relative proximity of the Seychelles to Madagascar, 

which is devoid of membracids. Nevertheless, the presence of Madlinus on the Seychelles-- 

the only membracid collected there besides Leptocentrus--could also be explained by a recent 

long distance dispersal from India. 

Other factors discredit an initial dispersal through the Afrotropical Region. In the 

Tertiary, the New World and Africa were separated by a long distance seemingly making 

dispersal by any means, whether rafting or by wind, difficult. The largest centrotine genus, 

Tricentrus, here placed in the Gargarini, is absent from the Afrotropical Region. Presumably 

some Tricentrus would be present in the Afrotropical Region, if treehoppers migrated from 

the New World to India through Africa. Also, the late Cretaceous connection between India 

and northeastern Africa may have been earlier than the hypothesized origin of membracids 

following the Cretaceous/Tertiary boundary. Moreover, the immediate Old World lineages 

following the Beaufortianini in the tree (Fig. 25.1) are primarily distributed in the 

437

Indomalayan, Palearctic, and Australasian/Oceanian Regions with very few genera present in 

Afrotropical Region. Finally, as mentioned previously, centrotines distributed in Gondwanan 

areas are not closely related, further excluding a Gondwanan vicariant event and dispersal 

through Africa. 

Thirdly, it is possible that both Old World invasions were from east to west over the 

Bering Land Bridge to Asia. This bridge connected North America and Asia beginning in 

the mid- to late Cretaceous, and from time to time throughout the Tertiary, as a result of 

continental convergence (Briggs 1995a). The geographic distributions of the larger 

centrotine lineages support this scenario. The Gargarini, the eventual descendants of the one 

invasion, are predominantly distributed in the Palearctic and Indomalayan Regions. The 

genus Imporcitor, the first lineage of the Beaufortianini (Fig. 24.1) (the first Old World tribe 

resulting from the other invasion), has been recorded in India, Taiwan, and Japan (McKamey 

1998a). As noted previously, all but one of the Old World centrotine tribes have 

representative genera found in the Indomalayan Region (Fig. 25.1). Evans (1966a) noted that 

the Indomalayan Region may be the center of origin for the Centrotinae considering the large 

number of species found there. Therefore, based on the phylogeny and known distributions 

of the centrotine tribes (Fig. 25.1), the Indomalayan Region appears to be a likely center of 

diversification and “jumping-off point” for centrotine dispersals to other zoogeographic 

regions, notably Australia and the Afrotropical Region. 

Strümpel (1972a) also believed that centrotines used the Bering Land Bridge as a 

dispersal route. However, he thought the Membracidae arose in Gondwana and that the 

Centrotinae reached the New World from Asia by way of the Bering Land Bridge in the 

438

Pleistocene. This would imply a more derived position for the New World centrotine tribes 

which is not supported by the current phylogenetic analysis (Fig. 25.1). 

The center of diversification for the Gargarini, the immediate descendants of one of 

the Old World Invasions, is clearly the Indomalayan Region (Fig. 25.1). All gargarine 

genera except one, Butragulus (McKamey 1998a) are found in the Indo-Malaysian Region. 

Furthermore, 17 of the 28 gargarine genera are restricted to this Region (McKamey 1998a). 

Based on the phylogenetic tree (Fig. 25.1), ancestors of the Australian treehoppers 

(Terentiini) likely arrived from the Indomalayan and Palearctic Regions. Apparently 

numerous angiosperm families now present in Australia invaded from southeast Asia in the 

early Tertiary (Briggs 1995a). It is possible that treehoppers followed these angiosperms into 

Australia. Other Australian insects, including the ground beetles (Coleoptera: Carabidae) 

and some scarabs (Scarabaeidae: Dynastinae), likely dispersed into Australia from southeast 

Asia (Briggs 1995a). According to Hall (1998a), northern Australia and Asia (present day 

Philippines) collided approximately 25 mya, which probably resulted in a temporary land 

connection. The ant genus Tetraponera likely invaded Australia from Asia 20 mya using this 

land bridge (Ward 2001a). It is possible that terentiine ancestors all used this temporary land 

bridge. Nonetheless, it is clear that Australia was the center of diversification for the 

Terentiini; all but 3 of the 40 terentiine genera are found in the Australasian/Oceanian 

Region. 

The center of diversification for the tribe Maarbarini is the Indomalayan Region (Fig. 

25.1) with all of the genera recorded from either India or Sri-Lanka or both (McKamey 

1998a). The derived position of this tribe on the phylogeny (Fig. 24.1, 25.1) and its 

immediate Indomalayan and Palearctic ancestors and relatives provide further evidence that 

439

treehoppers invaded India after it collided with Asia in the mid-Cretaceous. One would 

expect a closer relationship between maarbarines and Afrotropical and/or Australian tribes if 

maarbarine ancestors had colonized India when it was closer to or contiguous with the other 

Gondwanan continents in the late Cretaceous. 

Ancestors of the Afrotropical centrotines (Xiphopoeini and Centrotini, Fig. 25.1) 

likely would have arrived more recently from Indomalayan ancestors. The ancestors of these 

derived tribes may have dispersed into Africa approximately 23 mya in the Miocene when 

Africa collided with Asia. The Leptocentrini, the lineage basal to both these tribes, is widely 

distributed in the Old World and has significant components in the Afrotropical and 

Indomalayan Regions. All of the genera in the Centrotini, the largest centrotine tribe, are 

found in the Afrotropical Region, with a few genera (notably the basal genus Centrotus) also 

occurring in the Australasian/Oceanian, Indomalayan, and Palearctic Regions (McKamey 

1998a). The Afrotropical Region, therefore, appears to be the center of diversification for 

this large tribe. The Centrotini are a derived lineage with reduced hind wing venation (Fig. 

25.1). Their derived position and large distance from New World centrotines in the 

phylogenetic tree (Fig. 24.1, 25.1) is evidence against a historic Gondwanan relationship with 

Neotropical centrotines. 

A fourth possibility involves dispersal from west to east across a North Atlantic Land 

Bridge that is thought to have connected the Laurasian continents in the North Atlantic from 

the Mesozoic to the Eocene (57.8 mya) (Briggs 1995a). This route was a possible alternative 

to the Bering Land Bridge or perhaps a parallel dispersal route to the Old World. 

Nevertheless, if ancestral centrotines arose in tropical North America, dispersal to the Old 

440

World--particularly to the Indomalayan Region--across the North Atlantic route would 

involve a far greater distance than the Bering Land Bridge. 

Discussion. Of the four scenarios presented here, dispersal across the Bering Land 

Bridge is most compelling. A number of other organisms are thought to have migrated 

between North America and Asia via the Bering Land Bridge in the late Cretaceous and early 

Tertiary. The use of this bridge as a dispersal route to explain disjunct distributions has been 

hypothesized for placental mammals, salamanders, dinosaurs, and freshwater fish (Briggs 

1995a). Although most dispersals over the Bering Land Bridge are thought to have occurred 

from Asia to North America, there are several examples of organisms dispersing from North 

America to Asia (Briggs 1995a). The freshwater fish family Esocidae (pikes), distributed 

today in eastern North America and eastern Asia, apparently migrated a long distance from 

North America to Asia based on fossil evidence (Briggs 1995a). Researchers believe that 

numerous aquatic insects lineages used the land bridge as a connection between Asia and 

North America. The trichopteran genus Chimarra is thought to have dispersed from South 

America to Asia in the early Tertiary (Briggs 1995a). 

Further evidence in support of dispersal across the Bering Land Bridge includes 

records of similar floras in eastern Asia and eastern North America (Briggs 1995a). 

Numerous plant genera and families show disjunct distributions among Asia and North 

America. Apparently, angiosperms migrated back and forth between the two continents 

using the land bridge as a dispersal route. The initial dispersal was apparently eastern but 

later there were migrations of plants back to Asia from North America (Briggs 1995a). The 

presence of angiosperms would have provided a constant food source for dispersing 

treehoppers from North America to Asia. 

441

Evidence from plant biogeography supports a possible treehopper migration from 

North America to the Bering Land Bridge. In the early Tertiary, the dominant vegetation in 

temperate latitudes consisted mainly of deciduous plants. At the beginning of the Eocene, 

the time when the earliest known treehoppers (fossil stegaspidines) have been found, the 

climate was the warmest of the entire Cenozoic, allowing tropical animals, such as 

treehoppers, to invade to the north. Tropical and temperate vegetation spread northward due 

to this worldwide warming trend (Briggs 1995a). Several plant families, including 

Cactaceae, Liliaceae, Loasaceae, Nyctaginaceae, Martyniaceae, Tecophilaceae, and 

Zygophyllaceae, migrated northward from South America in the early Tertiary representing 

the beginnings of colonization of North America by South American tropical flora (Briggs 

1995a). Indeed, centrotines have been recorded from the families Liliaceae, Nyctaginaceae, 

and most notably the Zygophyllaceae (Table 26.2). The tribe Centrodontini, the first 

centrotine lineage (Fig. 25.1), is the only treehopper group known to feed on the 

Zygophyllaceae. A northward migration of treehoppers from North America to the Bering 

Land Bridge is supported by these plant migrations. 

Assuming the Bering Land Bridge was the dispersal route for both Old World 

invasions (Fig. 25.1), the treehoppers likely dispersed to the Old World and diversified in 

isolation without migrating back to North America. No centrotines are currently found in 

northern North America; the predominant North American treehopper subfamily is the 

Smiliinae. Centrotines may have gone extinct in these areas or migrated to southern refugia 

as a result of Pleistocene glaciations. Similarly, modern day continental Europe has only 

three native centrotine species but may have had more prior to the Pleistocene glaciations. 

Depauperate fauna in Europe may be due to extreme climatic conditions during the 

442

Pleistocene glaciation. Both eastern and western North America have close to 20% more 

genera of trees and shrubs endemic to their area than Europe. This discrepancy in floral 

diversity has been attributed to the east/west oriented mountains in Europe that may have 

prevented the migration of trees into lower, warmer latitudes during the glaciations of the 

Quaternary. In North America, the major mountain ranges run north/south. Here, the woody 

plants could migrate ahead of the glaciers without any barriers (Reid 1935a). According to 

Huntley (1993a), the harsh environments and climates of the Quaternary period eliminated 

many European forest taxa. These climatic effects could also have severely reduced 

treehopper numbers. 

Summary and Conclusions 

According to the fossil record and recent phylogenetic analyses, Membracidae likely 

originated in the New World during the Tertiary, possibly near the time of the hypothesized 

extraterrestrial impact 65 mya. Other evidence for a Tertiary origin include the absence of 

treehoppers in Madagascar and New Zealand, the historic connection of India with 

Madagascar and the high species richness of India (including the tribe Maarbarini), and the 

distantly related centrotine faunas of the Afrotropical Region, Australia, and South America. 

These observations do not correspond to patterns that would be expected if centrotines 

originated in Gondwana, prior to the splitting of the land masses. The highly endemic 

Monobelini and Nessorhinini of the Caribbean occupy a basal position in the phylogenetic 

analysis. The hypothesized dispersal of their ancestors to the Caribbean over an island 

archipelago at the Cretaceous/Tertiary border also supports a treehopper origin near this time. 

443

Centrotines are found in all major zoogeographic regions: The Afrotropical, the 

Nearctic, Neotropical, Palearctic, Indomalayan, and Australasian/Oceanian Regions. While a 

few tribes are widely distributed, many occur primarily in one or two major zoogeographic 

regions. The distinctive fauna of the Indomalayan, Australian/Oceanian, Afrotropical, and 

Caribbean regions are especially notable. All of the Old World centrotine tribes, except the 

Xiphopoeini, have representatives in the Indomalayan Region. 

Based on the phylogenetic analysis presented here, the early centrotines, apparently 

widely abundant in North America, dispersed twice to the Caribbean and twice to the Old 

World. Indeed, each of the two major centrotine clades has a basal lineage that dispersed to 

the Caribbean Region and also a more derived clade that dispersed to the Old World. 

Although the pathways and methods of dispersal are unclear, the repetitive pattern of 

dispersals in the two groups suggests that the timing, routes, and mechanisms may have been 

similar in each clade. It is possible that the dispersals to the Old World occurred over the 

Bering Land Bridge, accounting for the Indomalayan and Palearctic distributions of basal 

centrotine lineages. Based on the phylogeny and known distributions of the centrotine tribes, 

the Indomalayan Region is the most plausible center of diversification and “jumping-off 

point” for centrotine dispersals to other zoogeographic regions, notably Australasian and 

Afrotropical Regions. 

444

Figure 25.1. Distributions of centrotine tribes. Geographic regions are noted on 

the tribal phylogeny (modified from Fig. 24.1). Asterisks (*) denote obvious 

centers of generic diversification for those taxa known from multiple geographic 

regions. Plus signs (+) denote the region of the basal genus, if apparent. 

445

26: CENTROTINE ANT ASSOCIATIONS, HOST PLANTS, AND CHROMOSOME 

Introduction 

NUMBERS 

Although primarily recognized for their exaggerated morphological characteristics, 

treehoppers are also known for their complex life history patterns including ant-mutualisms, 

maternal care, sound communication, and host plant specialization. Wood summarized 

behavioral patterns in the New World Membracidae (1993a), and Ananthasubramanian and 

Ananthakrishnan (1975a, 1975b), Ananthasubramanian (1996a), and Capener (1962a, 1968a) 

discussed similar characteristics for Indian and African membracids, respectively. 

Ant-membracid mutualisms, where ants feed on the excreted treehopper honeydew 

and in exchange provide the homopterans defense against natural enemies, are well 

documented (Ananthasubramanian 1996a, Hölldobler and Wilson 1990a, Panda 1968a). 

Ant-attendance is common in both Old World (Table 26.1) and New World Membracidae 

(Wood 1993a). 

Wood (1993a) described different levels of maternal care in treehoppers. Maternal 

care occurs in four of the seven New World subfamilies and in approximately half of the 

tribes in the highly derived subfamilies Membracinae and Smiliinae. In addition to egg 

guarding, some New World groups aggressively defend their offspring from predators and 

parasitoids (Wood 1993a, McKamey and Deitz 1996a). 

The life history traits of the Centrotinae, a group primarily distributed in the Old 

World, have not been examined in an evolutionary context. Firstly, it is unclear if ant- 

attendance is an ancestral trait for centrotines and if ant-attendance and maternal care (egg 

446

guarding) are correlated. Secondly, there are many questions regarding the relationship 

between centrotines and their host plants. What were the original centrotine host plant 

families and what are the subsequent patterns of host plant exploitation? Baseline 

information on centrotine host plant families is presented here. Finally, centrotine 

chromosome data was gathered to determine the likely ancestral chromosome number, what 

derivations occurred, and the usefulness of chromosomes as phylogenetic characters. 

In order to examine these evolutionary patterns, centrotine behavioral and host plant 

patterns, as well as male chromosome numbers, are tabulated by genus and tribe and are 

optimized as unweighted characters on a tribal phylogeny (modified from Fig. 24.1). 

Although Ananthasubramanian (1996a) provided a summary of aggregation behaviors 

among the Centrotinae, data are lacking for too many taxa to establish evolutionary trends. 

Methods 

Published records of centrotine genera reported to be ant-attended, host plant 

families, and male chromosome numbers are summarized in Tables 26.1-26.3. Sources are 

given in each table. The phylogeny of the 23 centrotine tribes was modified from the tree in 

Fig. 24.1 to serve as a framework for mapping the ecological, behavioral, and chromosomal 

features. Ant-attendance, maternal care in the form of egg guarding, host plant families, and 

chromosome numbers were treated as unweighted characters (i.e., each host plant family was 

treated as a separate character in the analysis) and scored for each tribe in DELTA (Dallwitz 

et al. 1999a). Characters were scored as present if at least one genus in a tribe was reported 

as having the trait in the literature. For example, although the genus Coccosterphus is the 

only gargarine genus listed to feed on the family Nyctaginaceae, this host character was 

447

scored as present for the tribe Gargarini. Host plant information is unknown for the tribes 

Choucentrini, Micreunini, and Pieltainellini and these were scored with a question mark (?). 

Otherwise, if a host plant family is not recorded for a tribe, it was scored as absent. Tribes 

and genera without published information on ant-attendance or chromosome numbers were 

scored with a question mark (?). The presence or absence of each of these characters for the 

23 centrotine tribes was optimized in the parsimony program Winclada (Nixon 1999a) using 

the fast optimization procedure. The gain and loss of these characters were then mapped 

onto the tribal phylogeny (Figs. 26.1-26.3). 

Chromosome numbers for Chinese taxa follow Tian and Yuan (1997a: Table 1) 

except that Tricentrus acuticornis Funkhouser should be male 2n=10 (1997a: 156); some 

numbers in Yuan and Chou’s (2002a) review are ambiguous. 

Results and Discussion 

Ant-attendance and Maternal Care. Ant-attendance has been reported from 28 

centrotine genera and 11 of 23 centrotine tribes: Beaufortianini, Boocerini, Centrocharesini, 

Ebhuloidesini, Hypsaucheniini, Maarbarini, Oxyrhachini, and the four largest tribes in terms 

of genera, Centrotini, Gargarini, Leptocentrini, and Terentiini (Table 26.1). The tribe 

Gargarini has the largest number of ant-attended genera with 8, although “all” Australasian 

(Terentiini) and South African (including some Beaufortianini, Boccharini, Centrotini, 

Leptocentrini, Oxyrhachini, and Xiphopoeini) nymphs are said to be ant-attended. The New 

World tribe Centrodontini is the only centrotine group reported in the literature as not 

attended by ants (Dietrich et al. 2001a). 

448

Ancestral state reconstruction using parsimony resulted in a single derivation of ant- 

attendance in the common ancestor of Monobelini + Boocerini + Gargarini and the remaining 

19 centrotine tribes (Fig. 26.1). Ant-attendance, however, is an ambiguous trait in the 11 

tribes (denoted with a “?’ in Fig. 26.1) where attendance information is unknown. 

Therefore, although ant-attendance is ambiguous for many centrotine tribes, it appears to be 

an ancestral character for a majority of the Centrotinae. 

Lin (2003a), in a study of the treehopper subfamily Membracinae, concluded that ant- 

attendance precedes the derivation of maternal care (egg guarding) but that the gain of 

maternal care is accompanied by the loss of ant-attendance. These findings supported the 

hypothesis of Wood (1984a) that egg guarding in treehoppers is correlated with the absence 

of ant-attendance. Conversely, according to Tallamy and Schaefer (1997a), maternal care in 

the Hemiptera is frought with increased costs to the mother and young including increased 

exposure to predators and lower fecundity. Thus, many hemipteran groups have acquired 

traits such as ant-attendance to lower these costs. 

The results presented here for the Centrotinae (Fig. 26.1) do not reflect either of these 

conclusions. Maternal care in the form of egg guarding appears to be apomorphic in the 

Centrotinae. Tribes within the well supported clade Ebhuloidesini + Oxyrhachini + 

Terentiini + Hypsaucheniini + Centrocharesini all have ant-attended genera (Table 26.1) and 

are the only centrotines known to exhibit maternal care in the form of egg-guarding (Fig. 

26.1) (Stegmann and Linsenmair 2002a). (Hinton 1977a listed Platycentrus acuticornis as 

subsocial but it is unclear from the figure if the female is guarding nymphs or eggs. Thus, for 

the purposes of this work, Platycentrus is not considered to show maternal care in the form 

of egg guarding). This common trait, unique to these tribes, provides independent support of 

449

their relatedness. In centrotines, therefore, ant-attendance and maternal care in the form of 

egg guarding appear to occur concurrently. Cryan (1999a) also found sociality and ant- 

attendance to occur together on a phylogenetic tree of the Membracidae derived from 

combined morphological and molecular data. Nevertheless, as found in the Membracinae 

(Lin 2003a), maternal care is preceded by ant-attendance on the centrotine phylogenetic tree 

(Fig. 26.1). 

Apparently, females of some Membracinae, a derived membracid subfamily, 

developed the trait of defending eggs and did not require additional ant defense (Lin 2003a), 

but the females of some Centrotinae, a more primitive membracid subfamily, guard their 

eggs and have ant herders that provide additional defense. Clearly, further data on the life 

history of centrotines is needed to further elucidate the relationships between maternal care 

and ant-attendance. 

Host plants. Centrotines are documented from 105 different host plant families 

(Table 26.2). Based on ancestral state reconstruction by parsimony, most of the plant family 

derivations in the phylogenetic tree (Fig. 26.2) are concentrated at the tips, suggesting many 

centrotine taxa only recently adapted to these hosts. Apparent losses of host plants could 

reflect the absence of a plant family in a tribe’s geographic range. These patterns, especially 

in the large tribes Gargarini, Terentiini, Leptocentrini, and Centrotini, suggest an increasing 

trend towards generalization in host use although there are many examples of novel 

adaptations of hosts in centrotine tribes. Despite this observable trend in host generalization 

in centrotine tribes, these patterns should be examined based on generic-level data to avoid 

over-generalizations on centrotine/host relationships. 

450

Excluding the New World tribe Centrodontini, most centrotine tribes are extremely 

polyphagous. The North American genera of the Centrodontini, Centrodontus, Multareis, 

and Multareoides, feed only on a single plant species, the creosote bush, Larrea divaricata 

tridentata (DC) Felger and Lowe, in the family Zygophyllaceae. Twelve of 23 centrotine 

tribes, however, have at least 5 or more known host plant families. Three of the four largest 

centrotine tribes in numbers of genera have the most reported host plant families (Table 26.2, 

Fig. 26.2). Sixty plant families are listed for the Gargarini, 54 for the Leptocentrini, and 40 

for the Centrotini. Within the Gargarini, the genus Gargara is known from 35 host plant 

families and Tricentrus is known from 44 families. The species Centrotus cornutus 

(Centrotini) is recorded from 16 different plant families: Aceraceae, Betulaceae, Compositae, 

Corylaceae, Cornaceae, Ericaceae, Euphorbiaceae, Fagaceae, Juglandaceae, Leguminosae, 

Moraceae, Onagraceae, Pinaceae, Rhamnaceae, Rosaceae, and Salicaceae (Table 26.2). 

Wood (1993a) also noted the widespread polyphagy in tropical Old World centrotines and 

Central and South American membracids. In contrast, most North American temperate 

membracids are found on a single host genus (Wood 1993a). 

In addition to being polyphagous, many centrotines show adaptations to novel plant 

families. Forty plant families are hosts for only a single centrotine genus or tribe. Centrotus 

(Centrotini) is the only centrotine genus found on the plant families Aceraceae, Cornaceae, 

Corylaceae, Onagraceae, while Gargarini is the only centrotine tribe reported to feed on 

Alangiaceae, Cannabaceae, Coriariaceae, Elaeagnaceae, Hamamelidaceae, Myristicaceae, 

Nyctaginaceae, Oleaceae, Rhizophoraceae, Saxifragaceae, and Thymelaeaceae. The tribe 

Nessorhinini, a primitive group based on the phylogenetic analysis (Fig. 24.1), is the only 

centrotine group found on the Malpighiaceae. These patterns suggest recent adaptations by 

451

tribes to plant hosts, however, in most cases, the same tribes include genera that also feed on 

a number of other host plant families. Conversely, the Leptocentrini and Gargarini share 37 

host plant families even though they are not closely related (Fig. 24.1), while the more 

closely related Centrotini and Leptocentrini share 23 host plant families. These patterns may 

reflect convergent adaptations to host plants. 

The Leguminosae (hosts for 42 centrotine genera, 13 tribes), Solanaceae (22 genera, 

10 tribes), Euphorbiaceae (19 genera, 8 tribes), and Compositae (18 genera, 7 tribes), are 

exploited at more basal positions in the tree and appear to be favored centrotine hosts. 

Moreover, the Solanaceae and Leguminosae are likely the two original host plant families 

(Fig. 26.2) for most centrotines except the Centrodontini. More than half of the centrotine 

tribes have genera reported to feed on the host plant family Leguminosae. Ancestral state 

reconstruction by parsimony indicates 2 independent gains of legume feeding and 5 losses, 

with the most prominent gain in the common ancestor of Monobelini + Boocerini + 

Gargarini and the remaining 19 centrotine tribes (Fig. 26.2). With the exception of 

Ebhuloidesini, all tribes in the clade Centrocharesini + Terentiini + Ebhuloidesini + 

Oxyrhachini + Hypsaucheniini include some genera that feed on the Euphorbiaceae. 

Several closely related tribes feed on the same host plant family(s), providing 

independent support of their relatedness. For example, the related tribes Lobocentrini, 

Leptobelini, Maarbarini, Leptocentrini, and Centrotini all have genera that feed on plants 

within the family Fagaceae. Furthermore, only the Gargarini and the related Monobelini are 

known to feed on the family Araceae. 

Male Chromosome Numbers. Chromosome numbers have only been examined in 8 

centrotine tribes, the male number being reported universally. Therefore, many ambiguous 

452

gains were plotted using ancestral state optimization. Data is insufficent to conclude on the 

usefulness of male chromosome numbers as phylogenetic characters in Centrotinae. 

Nevertheless, 2n=21 may have been the original centrotine chromosome number (Fig. 26.3). 

Indeed, Kirillova’s (1987a) review gave 2n=21 as the mode among all treehoppers and 17 as 

the mode among leafhoppers (Cicadellidae). This chromosome number is apparently lost in 

the Ebhuloidesini, Hypsaucheniini, and Centrotini. Furthermore, the large tribe Gargarini 

independently acquired 5 different male chromosome numbers. The closely related tribes 

Centrotini, Maarbarini, and Leptocentrini all have genera with a male chromosome number 

of 2n=19, while the Ebhuloidesini and Hypsaucheniini have a number of 2n=17, providing 

independent corroboration of their relatedness. The sex chromosome system in centrotines is 

primarily XO:XX although Tian and Yuan (1997a) report a XY:XX system in two gargarine 

species: Nondenticentrus curvispineus Chou and Yuan and Tricentrus acuticornis 

Funkhouser. 

453

Table 26.1. Centrotine genera reported to be tended by ants with citation. Note: not all species within the 

genus are necessarily ant attended. 

Tribe: Genus (citation) 

BEAUFORTIANINI: Dukeobelus (Capener 1952b). 

BOOCERINI: Ischnocentrus (Loye 1992a; Olmstead and Wood 1990a). 

CENTROCHARESINI: Centrochares: (Stegmann and Linsenmair 2002a). 

CENTROTINI: Anchon (Ananthasubramanian 1984a, 1987a; Ayyar 1937a; Capener 1953b; Lamborn, 

1914a); Centrotus (Green 1900a); Leprechaunus (Capener 1950a); Monocentrus (Kenne and Dejean 

1997a). 

EBHULOIDESINI: Ebhul (Azhar 1992a, Funkhouser 1951a). 

GARGARINI: Butragulus (Hayashi and Endo 1985b); Coccosterphus (Ananthasubramanian and 

Ananthakrishnan 1975a); Eucoccosterphus (Ananthasubramanian and Ananthakrishnan 1975a); 

Chitra and Ananthasubramanian 1999a); Gargara (Hayashi and Endo 1985b; Ananthasubramanian 

and Ananthakrishnan 1975a); Enslin 1911a, 1911b; Funkhouser 1919d, 1951a; Panda 1968a; Weiss 

and Dickerson 1921a); Machaerotypus (Hayashi and Endo 1985b); Parayasa (Ananthasubramanian 

and Ananthakrishnan 1975a; Ananthasubramanian 1987a); Tsunozemia (Hayashi and Endo 1985b); 

Tricentrus (Ananthasubramanian and Ananthakrishnan 1975a); Funkhouser 1919d). 

HYPSAUCHENIINI: Gigantorhabdus (Ushijima and Nagai 1979a); Hybandoides (Stegmann and 

Linsenmair 2002a); Hypsauchenia (Funkhouser 1951a); Pyrgauchenia (Melichar 1914b). 

LEPTOCENTRINI: Hemicentrus (Melichar 1914b); Leptocentrus (Ananthasubramanian and 

Ananthakrishnan 1975a; Ananthasubramanian and Ramachandran 1990a; Boulard 1969a; Dejean and 

Bourgoin 1998a; Funkhouser 1951a; Kenne and Dejean 1997a; Panda 1968a; Panda and Behura 

1957a; Lamborn 1914a); Otinotus (Ananthasubramanian and Ananthakrishnan 1975a; Behura 1951a, 

1955a, 1962a; Behura and Panda 1959a; Behura and Sengupta 1951a; Behura and Sinha 1951a; 

Panda 1968a; Panda and Behura 1956a). 

MAARBARINI: Telingana (Ananthasubramanian and Ananthakrishnan 1975a). 

OXYRHACHINI: Oxyrhachis (Adenuga and Adeboyeku 1987a; Gersani and Degen 1988a; Lamborn 

1914a; Panda 1968a; Panda and Behura 1957a; Singh 1986a; Thakur 1973a). 

TERENTIINI: Cebes (Cookson and New 1980a); Sextius (Buckley 1982a, 1983a; Cookson and New 

1980a; Froggatt 1902a; Goding 1903a; Kitching and Filshie 1974a; Hölldobler and Wilson 1990a); 

Terentius (Kitching 1987a); all Australian nymphs and adults (Carver et al. 1991a; Evans 1966a; 

Tillyard 1926d). 

South African nymphs (all ?) (Jacobs 1985a). 

454

Table 26.2. List of host plant families and treehopper genera. Numbers correspond to labels in Fig. 25.2. Host 

plant families are based on the website: http://www.rbgkew.org.uk/data/vascplnt.html and Brummitt (1992a). 

Treehopper host plant data taken from: Ahmad 1975a, 1976a, 1978a, 1988a; Alma 1999a; Ananthasubramanian 

1987a, 1996a; Ananthasubramanian and Ananthkrishnan 1975a; Ananthasubramanian et al. 1990a; Ayyanna et 

al. 1978a; Ballou 1935a, 1936b; Behura 1951a, 1962a; de Bergevin 1934b; Boulard 1966a, 1968b, 1968d, 

1969a, 1969c, 1971c, 1979i, 1979j, 1983b; Capener 1951a, 1962a, 1966a, 1968a, 1968b, 1968c, 1971a, 1972a, 

1972b, 1972c; Chatterjee 1933c; Chatterjee and Bose 1933a; Cheo 1935b; Davli et al. 1992a; Day 1999a; 

Dietrich et al. 2001a; Funkhouser 1919a, 1919d, 1927b, 1935b; Goding 1893d; Gunji and Nagai 1994a; 

Günthart 1987b; Hargreaves 1937a; Hayashi and Endo 1985b; Helmore 1982a; Hoffmann 1942a; Jankovic 

1975a; Kirkaldy 1906c; Koningsberger 1915a; Krauss 1965a; Lamborn 1914a; Lodos and Kalkandelen 1981a; 

Loye 1992a; Matsumura 1912a; Melichar 1914b; Mohammad and Ahmad 1991a, 1995a; Morley 1905b; Okáli 

and Janský 1998a; Panda and Behura 1957a; Peláez 1941b; Plummer 1935a; Ramos 1957a, 1979a; Rao et al. 

1988a; Raut and Bhattacharya 1999a; Richter 1942c; Smithers 1985a; Swezey 1942a; Wolcott 1941a; Yousuf et 

al. 1997a; Yuan and Chou 2002a. 

Host plant family Tribe (Genus) 

1. Aceraceae Centrotini (Centrotus) 

2. Actinidiaceae Leptobelini (Leptobelus) 

3. Adiantaceae Leptocentrini (Leptocentrus) 

4. Alangiaceae Gargarini (Tricentrus) 

5. Alliaceae Leptocentrini (Otinotus); Maarbarini (Telingana) 

6. Amaranthaceae Gargarini (Gargara, Tricentrus); Leptocentrini (Leptocentrus); Oxyrhachini 

(Oxyrhachis) 

7. Anacardiaceae Boocerini (Campylocentrus); Centrotypini (Centrotypus); Gargarini (Tricentrus); 

Leptocentrini (Hemicentrus, Leptocentrus, Otinotus); Monobelini (Monobelus); 

Nessorhinini (Nessorhinus) 

8. Annonaceae Hypsaucheniini (Gigantorhabdus); Leptocentrini (Leptocentrus, Otinotus) 

9. Apocynaceae Centrotini (Monocentrus); Gargarini (Tricentrus); Leptocentrini (Leptocentrus, 

Otinotus) 

10. Araceae Gargarini (Tricentrus); Monobelini (Monobelus) 

11. Araliaceae Gargarini (Machaerotypus); Leptocentrini (Leptocentrus) 

12. Aristolochiaceae Leptobelini (Leptobelus) 

13. Asclepiadaceae Boocerini (Campylocentrus); Gargarini (Tricentrus); Leptocentrini (Leptocentrus, 

Otinotus) 

14. Balanitaceae Oxyrhachini (Oxyrhachis) 

15. Balsaminaceae Leptocentrini (Otinotus) 

16. Betulaceae Centrotini (Centrotus); Gargarini (Butragulus, Gargara, Machaerotypus, Tricentrus) 

17. Bignoniaceae Gargarini (Coccosterphus, Gargara, Tricentrus); Leptocentrini (Otinotus); Oxyrhachini 

(Oxyrhachis) 

18. Bixaceae Leptocentrini (Leptocentrus) 

19. Bombacaceae Leptocentrini (Hemicentrus, Leptocentrus); Gargarini (Tricentrus) 

20. Bromeliaceae Centrotini (Hamma) 

21. Buddlejaceae Oxyrhachini (Oxyrhachis) 

22. Cannabaceae Gargarini (Gargara) 

23. Capparaceae Leptocentrini (Leptocentrus, Otinotus) 

24. Caprifoliaceae Maarbarini (Telingana) 

25. Caricaceae Centrotini (Hamma, Monocentrus) 

26. Casuarinaceae Gargarini (Tricentrus); Leptocentrini (Leptocentrus, Otinotus); Oxyrhachini 

(Oxyrhachis); Terentiini (Acanthuchus, Terentius) 

27. Celastraceae Gargarini (Gargara, Parayasa); Leptocentrini (Periaman) 

28. Chenopodiaceae Terentiini (Acanthucalis) 

29. Combretaceae Centrotini (Distanobelus, Platybelus); Gargarini (Gargara); Leptocentrini 

(Leptocentrus, Otinotus, Umfilianus); Monobelini (Monobelus) Continued. 

455


Host plant family Genus 

30. Compositae Centrotini (Anchon, Centrotus, Distanobelus, Leprechaunus); Gargarini (Butragulus, 

(Asteraceae) Coccosterphus, Gargara, Machaerotypus, Tricentrus, Tsunozemia); Hypsaucheniini 

(Hypsauchenia, Hypsolyrium, Jingkara); Leptobelini (Leptobelus); Leptocentrini 

(Leptocentrus, Otinotus); Monobelini (Monobeloides); Oxyrhachini (Oxyrhachis) 

31. Connaraceae Leptocentrini (Leptocentrus) 

32. Convolvulaceae Boocerini (Campylocentrus); Centrotini (Anchon); Leptobelini (Leptobelus), 

Leptocentrini (Leptocentrus) 

33. Coriariaceae Gargarini (Tricentrus) 

34. Cornaceae Centrotini (Centrotus) 

35. Corylaceae Centrotini (Centrotus) 

36. Cruciferae Gargarini (Gargara); Leptocentrini (Otinotus) 

37. Cucurbitaceae Boocerini (Campylocentrus) 

38. Cupressaceae Maarbarini (Telingana) 

39. Dryopteridaceae Maarbarini (Telingana) 

40. Ebenaceae Gargarini (Gargara, Machaerotypus); Oxyrhachini (Oxyrhachis) 

41. Elaeagnaceae Gargarini (Erecticornia, Gargara, Maurya, Tricentrus) 

42. Ericaceae Centrotini (Centrotus); Gargarini (Butragulus, Machaerotypus) 

43. Euphorbiaceae Centrocharesini (Centrochares); Centrotini (Anchon, Centrotus, Eumonocentrus, 

Hamma, Monocentrus); Gargarini (Coccosterphus, Eucoccosterphus, Gargara, Sipylus, 

Tricentrus); Hypsaucheniini (Hypsauchenia, Hypsolyrium); Leptocentrini 

(Leptocentrus, Otinotus); Oxyrhachini (Oxyrhachis); Terentiini (Pyrgonota, Terentius); 

Xiphopoeini(Xiphopoeus) 

44. Fagaceae Centrotini (Centrotus); Gargarini (Erecticornia, Gargara, Machaerotypus, Maurya, 

Tricentrus); incertae sedis (Elaphiceps); Leptobelini (Leptobelus); Leptocentrini 

(Leptocentrus); Lobocentrini (Arcuatocornum, Truncatocornum); Maarbarini 

(Telingana) 

45. Flacourtiaceae Ebhuloidesini (Ebhul); Leptocentrini (Hemicentrus, Otinotus) 

46. Gnetaceae Leptobelini (Leptobelus) 

47. Gramineae Boocerini (Campylocentrus); Centrotini (Anchon); Gargarini (Tricentrus); Leptocentrini 

(Leptocentrus); Maarbarini (Telingana); Oxyrhachini (Oxyrhachis); Terentiini 

(Pogonella, Strzeleckia); Xiphopoeini (Xiphopoeus) 

48. Guttiferae Boocerini (Ischnocentrus); Gargarini (Tricentrus); Leptocentrini (Otinotus) 

49. Hamamelidaceae Gargarini (Tricentrus) Continued. 

50. Hernandiaceae Leptocentrini (Leptocentrus) 

51. Juglandaceae Centrotini (Centrotus); Gargarini (Machaerotypus, Tricentrus) 

52. Labiatae Leptocentrini (Leptocentrus, Otinotus); Nessorhinini (Nessorhinus) 

53. Lauraceae Boccharini (Lanceonotus); Ebhuloidesini (Ebhul); Gargarini (Gargara); Leptocentrini 

(Hemicentrus, Otinotus); Oxyrhachini (Oxyrhachis); Terentiini (Acanthuchus) 

54. Lecythidaceae Terentiini (Terentius) 

55. Leguminosae Beaufortianini (Centruchus); Centrocharesini (Centrochares); Centrotini 

(Fabaceae); (Acanthophyes, Anchon, Anchonobelus, Centrotus, Cornutobelus, Distanobelus, 

(Leguminosae- Eumonocentrus, Hamma, Monocentrus, Rachinotus, Stalobelus, Tiberianus, Tricoceps); 

Caesalpinioideae); Gargarini (Butragulus, Coccosterphus, Eucoccosterphus, Gargara, Machaerotypus, 

(Leguminosae- Parayasa, Tricentrus); Hypsaucheniini (Jingkara); Leptobelini (Leptobelus); 

Mimosoideae); Leptocentrini (Hemicentrus, Leptocentrus, Otinotus); Monobelini (Monobelus); 

(Leguminosae- Nessorhinini (Nessorhinus); Oxyrhachini (Oxyrhachis); Platycentrini (Platycentrus, 

Papilionoideae) Tylocentrus); Terentiini (Acanthuchus, Anzac, Cebes, Ceraon, Eufairmairia, Eufrenchia, 

Pogonella, Sarantus, Sextius); Xiphopoeini (Xiphopoeus) 

56. Liliaceae Centrotini (Anchon, Tricoceps); Gargarini (Gargara); Leptocentrini (Leptocentrus) 

57. Lythraceae Gargarini (Eucoccosterphus, Gargara, Tricentrus); Leptocentrini (Otinotus) 

58. Magnoliaceae Hypsaucheniini (Hypsauchenia); Leptocentrini (Leptocentrus, Otinotus, Periaman) 

59. Malpighiaceae Nessorhinini (Nessorhinus, Orthobelus) Continued. 

456



60. Malvaceae Centrotini (Hamma); Gargarini (Gargara, Tricentrus); Hypsaucheniini (Jingkara); 

Leptocentrini (Leptocentrus); Nessorhinini (Nessorhinus); Terentiini (Alosextius) 

61. Melastomataceae Boocerini (Ischnocentrus); Ebhuloidesini (Ebhul), Gargarini (Gargara, Sipylus, 

Tricentrus); Hypsaucheniini (Pyrgauchenia); Leptocentrini (Leptocentrus, 

Nilautama) 

62. Meliaceae Gargarini (Gargara, Tricentrus); Leptocentrini (Leptocentrus) 

63. Moraceae Centrotini (Centrotus, Eumonocentrus); Centrotypini (Centrotypus); Ebhuloidesini 

(Ebhul); Gargarini (Eucoccosterphus, Gargara, Machaerotypus, Tricentrus); 

Leptocentrini (Leptocentrus, Otinotus); Monobelini (Monobelus); Nessorhinini 

(Nessorhinus); Oxyrhachini (Oxyrhachis); Terentiini (Pogonotypellus) 

64. Moringaceae Gargarini (Gargara); Leptocentrini (Leptocentrus, Otinotus) 

65. Myristicaceae Gargarini (Tricentrus) 

66. Myrtaceae Gargarini (Tricentrus); Leptocentrini (Otinotus); Nessorhinini (Nessorhinus); 

Oxyrhachini (Oxyrhachis); Terentiini (Sextius, Acanthuchus, Ceraon, Eufairmairia, 

Eufairmairiella, Eufrenchia) 

67. Nyctaginaceae Gargarini (Coccosterphus) 

68. Olacaceae Centrotini (Anchon, Monocentrus) 

69. Oleaceae Gargarini (Gargara, Tricentrus) 

70. Onagraceae Centrotini (Centrotus) 

71. Orchidaceae Leptocentrini (Leptocentrus) 

72. Oxalidaceae Centrotini (Zanzia); Gargarini (Tricentrus) 

73. Palmae Centrotini (Monocentrus); Leptocentrini (Leptocentrus) 

74. Passifloraceae Leptocentrini (Leptocentrus) 

75. Phytolaccaceae Boocerini (Campylocentrus); Nessorhinini (Nessorhinus) 

76. Pinaceae Centrotini (Centrotus); Gargarini (Gargara, Machaerotypus, Maurya, Tricentrus) 

77. Piperaceae Centrotini (Monocentrus); Gargarini (Tricentrus); Leptocentrini (Leptocentrus) 

78. Plumbaginaceae Terentiini (Pogonella) 

79. Polygonaceae Centrotini (Tricoceps); Gargarini (Gargara, Tricentrus); Leptocentrini 

(Leptocentrus); Terentiini (Cebes) 

80. Proteaceae Beaufortianini (Dukeobelus); Oxyrhachini (Oxyrhachis); Terentiini (Acanthuchus, 

Otinotoides, Pogonella, Sertorius, Terentius) 

81. Ranunculaceae Centrotini (Anchon) 

82. Rhamnaceae Centrotini (Acanthophyes, Centrotus, Distanobelus); Centrotypini (Centrotypus); 

Gargarini (Gargara); Leptocentrini (Leptocentrus, Otinotus); Oxyrhachini 

(Oxyrhachis) 

83. Rhizophoraceae Gargarini (Tricentrus) 

84. Rosaceae Centrotini (Anchon, Centrotus, Daconotus); Gargarini (Butragulus, Machaerotypus, 

Maurya, Pantaleon, Tricentrus); Leptobelini (Leptobelus); Maarbarini (Telingana); 

Terentiini (Acanthuchus) 

85. Rubiaceae Boocerini (Campylocentrus); Centrotini (Euceropsila, Eumonocentrus, Hamma, 

Monocentrus); Centrotypini (Centrotypus); Ebhuloidesini (Ebhul); Gargarini 

(Eucoccosterphus, Tricentrus); Leptocentrini (Leptocentrus, Otinotus); Nessorhinini 

(Nessorhinus) 

86. Rutaceae Centrotini (Hamma, Monocentrus); Gargarini (Gargara, Machaerotypus); 

Leptocentrini (Leptocentrus, Otinotus); Nessorhinini (Nessorhinus); Terentiini 

(Pogonella) 

87. Salicaceae Centrotini (Centrotus); Gargarini (Gargara, Machaerotypus, Tricentrus); 

Hypsaucheniini (Hypsauchenia) 

88. Salvadoraceae Leptocentrini (Leptocentrus) 

89. Santalaceae Centrotypini (Centrotypus); Gargarini (Eucoccosterphus, Coccosterphus, Gargara, 

Tricentrus); Leptocentrini (Leptocentrus, Otinotus); Maarbarini (Pogon); 

Oxyrhachini (Oxyrhachis) Continued. 

457



90. Sapindaceae Centrotini (Anchon); Gargarini (Tricentrus) 

91. Sapotaceae Monobelini (Brachycentrotus) 

92. Saxifragaceae Gargarini (Antialcidas, Pantaleon) 

93. Solanaceae Beaufortianini (Beaufortiana); Boocerini (Ischnocentrus); Centrotini (Anchon, 

Eumonocentrus); Gargarini (Coccosterphus, Eucoccosterphus, Gargara, Parayasa, 

Tricentrus); Leptocentrini (Leptocentrus, Otinotus); Maarbarini (Telingana); 

Monobelini (Monobelus); Nessorhinini (Nessorhinus); Oxyrhachini (Oxyrhachis); 

Terentiini (Neocanthuchus, Oxyrhachis, Pogonella, Pogonotypellus, Rentzia, Sarantus, 

Terentius) 

94. Sterculiaceae Centrotini (Eumonocentrus, Hamma, Monocentrus, Stalobelus); Leptocentrini 

(Leptocentrus); Oxyrhachini (Oxyrhachis) 

95. Styracaceae Hypsaucheniini (Hypsauchenia); Leptobelini (Leptobelus) 

96. Tamaricaceae Gargarini (Gargara); Leptocentrini (Leptocentrus, Otinotus); Oxyrhachini (Oxyrhachis) 

97. Theaceae Centrotini (Stalobelus); Gargarini (Machaerotypus, Tricentrus) 

98. Thymelaeaceae Gargarini (Gargara); Hypsaucheniini (Jingkara) 

99. Tiliaceae Centrotini (Anchon, Eumonocentrus, Hamma, Leprechaunus, Monocentrus); Gargarini 

(Butragulus, Gargara, Tricentrus); Leptocentrini (Leptocentrus, Otinotus) 

100. Ulmaceae Centrotini (Anchon, Leprechaunus, Monocentrus); Gargarini (Butragulus, Gargara, 

Machaerotypus, Tricentrus); Leptocentrini (Otinotus); Oxyrhachini (Oxyrhachis) 

101. Urticaceae Centrotini (Anchon, Mitranotus, Monocentrus); Leptocentrini (Leptocentrus, Otinotus) 

102. Verbenaceae Gargarini (Gargara, Tricentrus); Leptocentrini (Leptocentrus, Otinotus); Oxyrhachini 

(Oxyrhachis) 

103. Vitaceae Gargarini (Gargara, Tricentrus); Nessorhinini (Nessorhinus) 

104. Zingiberaceae Centrotini (Kallicrates) 

105. Zygophyllaceae Centrodontini (Centrodontus, Multareis, Multareoides) 

458

Table 26.3. Chromosome numbers (2n) of male centrotines. 

Male 2n Taxa and citations 

2n=10 Gargarini: Tricentrus (Tian and Yuan 1997a). 


2n=17 Centrotini: Anchon (Kirillova 1988a; Parida and Dalua 1981a). 

Ebhuloidesini: Ebhul (Tian and Yuan 1997a). 

Hypsaucheniini: Jingkara (Tian and Yuan 1997a). 

2n=19 Centrotini: Centrotus (Halkka 1959a, 1962a; Kirillova 1988a). 

Gargarini: Coccosterphus (Bhattacharya and Manna 1973a; Kirillova 1988a); Gargara 

(Bhattacharya and Manna 1967a, 1973a; Halkka 1959a, 1962a; Kirillova 1988a; 

Menon 1958a; Parida and Dalua 1981a; Ray-Chaudhuri et al. 1967a; Tian and 

Yuan 1997a); Tricentrus (Ahmad and Yasmeen 1980a; Bhattacharya and Manna 

1973a; Kirillova 1988a; Parida and Dalua 1981a; Tian and Yuan 1997a). 

Ebhuloidesini: Ebhul (Tian and Yuan 1997a). 

Leptocentrini: Leptocentrus substitutus (Halkka 1962a; Menon 1959a). 

Maarbarini: Telingana (Kirillova 1988a; Sharma et al. 1964a). 

2n=20 Gargarini: Nondenticentrus (Tian and Yuan 1997a). 

2n=21 Boocerini: Boocerus (Halkka 1964a; Kirillova 1988a). 

Gargarini: Coccosterphus (Bhattacharya and Manna 1967a); Pantaleon (Tian and Yuan 

1997a); Tricentrus (Kirillova 1988a; Parida and Dalua 1981a; Tian and Yuan 

1997a). 

Leptocentrini: Hemicentrus (Tian and Yuan 1997a); Leptocentrus (Banerjee 1958a; 

Bhattacharya and Manna 1967a, 1973a; Halkka 1959a; Kirillova 1988a; Parida 

and Dalua 1981a; Rao 1956a; Tian and Yuan 1997a); Otinotus (Bhattacharya and 

Manna 1967a, 1973a; Halkka 1959a, 1962a; Kirillova 1988a; Menon 1958a; 

Parida and Dalua 1981a). 

Maarbarini: Pogon (Sharma et al. 1964a); Telingana (Sharma et al. 1964a). 

Oxyrhachini: Oxyrhachis (Abrar and Ahmad 1975a; Banerjee 1958a; Bhattacharya and 

Manna 1967a, 1973a; Biswas and Bhattacharya 1989a; Halkka 1959a, 1962a; 

Kirillova 1988a; Menon 1958a; Parida and Dalua 1981a; Rao 1956a; Sharma et 

al. 1964a; Tian and Yuan 1997a). 

Terentiini: Eufairmairia (Whitten 1965a; Kirillova 1988a); Sextius (Kirillova 1988a; 

Whitten 1965a). 


459

Fig. 26.1. Ant attendance and maternal care in centrotines. The gain or loss of each 

trait was mapped on the tribal phylogeny tree using fast optimization in Winclada. 

Question marks (?) represent missing data. 

460

Fig. 26.2. Host plant families of centrotines. The gain (indicated below branches) or loss 

(indicated above branches) of each host plant family was mapped on the tribal phylogeny 

using fast optimization in Winclada. Numbers on the branches identify host plant families 

in Table 26.2. Question marks represent missing data. 

461

Fig. 26.3. Male chromosome numbers in centrotines. The gain (indicated below 

branches) or loss (indicated above branches) of each was mapped on the tribal 

phylogeny using fast optimization in Winclada. Question marks (?) represent 

missing data. 

462

CONCLUSIONS 

The treehopper subfamily Centrotinae is the only treehopper group found worldwide. 

Predominantly Old World in distribution, this subfamily accounts for roughly half of the 

membracid diversity at the tribal, generic, and species levels. Although centrotines are 

cosmopolitan in distribution, there is no centrotine tribe (or genus) found in both the Old and 

New Worlds. 

Despite their diversity, the Centrotinae have been poorly studied. Historically, 

workers have focused their work on centrotines within a particular geographic region (for 

example, Capener 1962a, 1968a; Evans 1966a; Ananthasubramanian 1996a; Day 1999a; 

Yuan and Chou 2002a) and few have justified classifications based on quantitative 

phylogenetic analyses. These disparities have impeded the development of a stable higher 

classification and taxonomic studies of centrotines at the generic and species levels. The 

objectives of this study were to establish the phylogenetic limits of the Centrotinae and its 

included tribes, to determine the evolutionary relationships among these tribes in order to 

provide a sound and comprehensive classification, to advance investigations of 

biogeographical patterns and life history traits, and to develop a tribal key to facilitate 

identification of the centrotine assemblage. 

An overall phylogenetic analysis of the 24 existing tribes plus 5 outgroups, using 116 

morphological characters, resulted in a single most parsimonious tree with 2 major clades 

(each with New and Old World components) plus the basal New World tribe Centrodontini. 

Numerous tribes, as defined in earlier classifications, were rendered polyphyletic or 

paraphyletic. Eight further phylogenetic analyses confirmed the monophyly of the larger 

463

tribes, and, along with two phenetic analyses, helped to place the remaining genera. The 

subfamily Centrotinae is a monophyletic group supported by the synapomorphy of the 

presence of abdominal inornate pits, each with a lateral seta. Characters important in 

elucidating tribal relationships include features of: the male and female genitalia, the fore- 

and hind wings, the scutellum, leg chaetotaxy; and abdominal characteristics using scanning 

electron microscopy. 

Based on the overall analysis, 11 tribal synonymies and 1 subfamily synonymy are 

proposed: Abelini, junior synonym of Boocerini; Acanthophyesaria, junior synonym of 

Centrotini; Ebhulini, junior synonym of Ebhuloidesini; Aleptocentrini, Antialcidini, 

Coccosterphini, Madlinini, and Tricentrini, all junior synonyms of Gargarini; Demangini, 

junior synonym of Leptocentrini; Bulbaucheniini and Funkhouserellini, junior synonyms of 

Terentiini; and Centrodontinae, junior synonym of Centrotinae. Furthermore, 6 new tribes 

are described: the Beaufortianini, Boccharini, Lobocentrini, and Maarbarini, all from the Old 

World, and Monobelini, and Pieltainellini from the New World. The 216 included centrotine 

genera are placed into a total of 23 centrotine monophyletic tribes. Two genera from the 

phylogenetic analysis, Elaphiceps and Tyrannotus, are placed as Centrotinae, incertae sedis, 

although they are closely related to the Lobocentrini. Seven genera that were not examined 

are placed as Centrotinae, incertae sedis: Aspasiana, Centrobelus, Insitor, Insitoroides, 

Megalocentrus, Megaloschema, and Sinocentrus. 

Brachytalis, formerly in Nessorhinini, is placed as Membracidae, incertae sedis. 

Additionally, a lectotype of Butragulus flavipes (Uhler) is designated. The new combinations 

Hybanda bulbicornis (Funkhouser), referred from Funkhouserella, and Bulbauchenia bakeri 

(Funkhouser), B. rugosa (Funkhouser), B. globosa (Funkhouser), and B. kurosawai (Hayashi 

464

and Endo), all referred from the Emphusis Buckton, are proposed. Deitzius 

Ananthasubramanian is a junior objective synonym of Ananthasubramanium McKamey, 

both of which are replacement names for Paranotus Ananthasubramanian, preoccupied. A 

taxonomic key to the 23 centrotine tribes is included. The descriptions of the 23 tribes 

include diagnoses, descriptions, notes on ecology and distribution, and discussions of 

phylogeny and morphological characters. 

Numerous centrotine genera are poorly represented in collections with some known 

from only one sex or even a unique specimen. All but 9 of the 216 genera were here placed 

in tribes based on phylogenetic analyses of morphological features, or in cases where data 

were limited, based on overall morphological similarity. 

The broader impacts of this work extend beyond systematics to related areas of 

biological inquiry. The phylogenetically based classification presented here has greater 

predictive value than prior classifications that included numerous para-or polyphyletic tribes. 

Furthermore, the phylogenetic hypotheses outlined provide a sound foundation for exploring 

patterns in host plant associations, ant-attendance, maternal care, acoustic communication, 

and biogeography. 

Based on the phylogenetic analyses, centrotines colonized the Old World twice. One 

invasion included the ancestors of the tribe Gargarini while the other invasion included the 

ancestors of the remaining 16 Old World centrotine tribes. It seems possible that these 

dispersals from the New World to the Old occurred over the Bering Land Bridge, accounting 

for the Indomalayan and Palearctic distributions of the basal centrotine lineages. According 

to the fossil record and recent phylogenetic analyses of the Membracidae, treehoppers likely 

originated in the New World in the Tertiary, possibly near the time of the hypothesized 

465

extraterrestrial impact 65 mya. Other evidence for a Tertiary origin includes the absence of 

treehoppers in Madagascar and New Zealand, the historic connection of India with 

Madagascar and the high species richness of India, and the distantly related centrotine faunas 

of Africa, Australia, and South America. These observations do not favor an ancient 

vicariant event, for example the splitting of Gondwana, that isolated the Old and New World 

treehopper faunas. The highly endemic Monobelini and Nessorhinini of the Caribbean 

occupy a basal position in the phylogenetic analysis. The hypothesized dispersal of their 

ancestors to the Caribbean over an island archipelago at the Cretaceous/Tertiary border also 

supports a treehopper origin near this time. 

Centrotines have been reported from 105 different host plant families, notably the 

Leguminosae, Solanaceae, Compositae, and Euphorbiaceae. Eleven of 23 centrotine tribes 

have genera that are ant-attended and the Centrocharesini + Ebhuloidesini + Oxyrhachini + 

Hypsaucheniini + Terentiini are the only centrotine group known to exhibit maternal care by 

egg guarding. Although male chromosome numbers of many centrotine tribes are unknown, 

many of the tribes reported have 2n=21. The reported range in males is 2n= 10-23. 

Armed with the revised classification presented here and the distributional data 

summarized by recent workers (McKamey 1998a, Day 1999a, Yuan and Chou 2002a), it is 

possible to identify hotspots of centrotine diversity or endemism--areas that merit special 

attention in efforts to preserve global biodiversity. The New World tribes Nessorhinini and 

Monobelini, for example, are endemic to the Caribbean. Likewise, Queensland, Australia, is 

a hotspot of diversity for the Terentiini. It is hoped that the improved identification tools 

provided here will encourage further collecting, stimulate systematic studies at the generic 

and species levels, and facilitate molecular research at all taxonomic levels. 

466

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19. Tribe OXYRHACHINI Distant 1908 Old World: Afrotropical ...

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