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A new genus of Caribbean Odontiinae with Palaeotropical affinities (Lepidoptera: Crambidae)

December 2009
Journal of the Lepidopterists’ Society 63(4)

December 2009
63(4)

Authors:

Florida Department of Agriculture and Consumer Services

Suinoorda maccabei gen. et sp. nov. is described from the Bahama Islands and Cuba. Cladistic analysis of the Eurrhypini (39 terminals, 48 characters) tests the phylogenetic relationships of the species and confirms that it is the earliest-diverging member of an otherwise Palaeotropical clade characterized by a modified plectrum of the eurrhypine genitalic stridulatory apparatus. The new species is not closely re-lated to other Neotropical Eurrhypini with similar but plesiomorphic or convergent maculation. The nomenclatural status of other Eurrhypini is revised. Clupeosoma orientalale (Viette) comb. nov. and Clupeosoma vohilavale (Marion & Viette) comb. nov. are transferred from Au-tocharis Swinhoe, Noordodes magnificalis (Rothschild), comb. nov. is transferred from Noorda Walker, and Hyalinarcha hyalinalis (Hampson), comb. nov. is transferred from Boeotarcha Meyrick. Metrea Grote is synonymized with Cliniodes Guenée, syn. nov., and the synonymy of Ba-songa Möschler with Cliniodes is reinstated. Missing data for female genitalia and larval feeding habit are predicted with a simple method.

. Species and specimen preparations included in the analysis. Slide numbers refer to preparations by the author (JEH) unless indicated by preparator's initials.

…

. Predicted missing data. The following states add no extra steps to the cladograms when analyzed together.

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Cladogram (187 steps, CI = 0.35, RI = 0.67). Numbers above branches refer to nodes (Appendix C), and numbers below are Bremer support values (suboptimality of trees lacking the node). NA, Nearctic; NT, Neotropical; PA, Palaearctic; PT, Palaeotropical; E, Eurrhypini; L, leaf-feeding clade; S, sepiform clade. *A. margarita is also known from Madagascar (Munroe 1974). **C. ostreonalis is Eastern Nearctic.

…

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Journal of the Lepidopterists’ Society

63(4), 2009,185-208

A NEW GENUS OF CARIBBEAN ODONTIINAE WITH PALAEOTROPICAL AFFINITIES

(LEPIDOPTERA: CRAMBIDAE)

JAMES E. HAYDEN

2144 Comstock Hall, Cornell University, Ithaca, NY 14853 USA, email: jeh63@cornell.edu

ABSTRACT. Suinoorda maccabei gen. et sp. nov.is described from the Bahama Islands and Cuba. Cladistic analysis of the Eurrhypini (39

terminals, 48 characters) tests the phylogenetic relationships of the species and conﬁrms that it is the earliest-diverging member of an otherwise

Palaeotropical clade characterized by a modiﬁed plectrum of the eurrhypine genitalic stridulatory apparatus. The new species is not closely re-

lated to other Neotropical Eurrhypini with similar but plesiomorphic or convergent maculation. The nomenclatural status of other Eurrhypini

is revised. Clupeosoma orientalale (Viette) comb. nov. and Clupeosoma vohilavale (Marion & Viette) comb. nov. are transferred from Au-

tocharis Swinhoe, Noordodes magniﬁcalis (Rothschild), comb. nov. is transferred from Noorda Walker, and Hyalinarcha hyalinalis (Hampson),

comb. nov. is transferred from Boeotarcha Meyrick. Metrea Grote is synonymized with Cliniodes Guenée, syn. nov., and the synonymy of Ba-

songa Möschler with Cliniodes is reinstated. Missing data for female genitalia and larval feeding habit are predicted with a simple method.

Additional key words: Stridulation, seed feeding, frontal structure, diagnosis, missing data

INTRODUCTION

The purpose of this paper is to describe and diagnose

a new, phylogenetically unique species of snout moth

from the West Indies. In the course of my studies on

the Neotropical Eurrhypini Leraut & Luquet

(Crambidae: Odontiinae), T.L. McCabe brought to my

attention a series of specimens that he collected in 1986

on Great Exuma, Bahama Islands. I subsequently

found in the Museum für Tierkunde (Dresden) one

specimen collected in 2000 by J.-P. Rudloff in eastern

Cuba. The male specimens signiﬁcantly resemble other

tropical Eurrhypini in having almost entirely white

forewings with the antemedial line nearly absent, a

black discocellar spot, and a simple, narrow, reddish

terminal band (Fig. 1). This pattern occurs in

Neotropical eurrhypines such as Argyrarcha Munroe,

Sobanga Munroe, and Cliniodes paradisalis (Möschler)

(Fig. 4), as well as in Palaeotropical genera such as

Autocharis Swinhoe and Pseudonoorda Munroe (Fig.

3).Dissection of males conﬁrmed its placement in the

Eurrhypini and more precisely with a group of genera

that share an unusual, hitherto unrecognized

modiﬁcation of the genitalic structures that characterize

the tribe (Figs. 8, 9) (Leraut & Luquet 1983). Its closest

relatives are restricted to the Palaeotropics, ranging

from Africa to New Guinea; some of these genera share

the wing pattern (e.g. Pseudonoorda), and others do not

(Clupeosoma Snellen). However, the new species also

shares symplesiomorphies with members outside the

group. Most strikingly, the frontoclypeal suture, just

above the base of the haustellum, is exaggerated in a

porcine snout of nearly circular shape (Fig. 5). This

appears to be homologous to the distinctly upturned

frontoclypeus of Autocharis and Dicepolia Snellen (Fig.

6). The new species also has plesiomorphic forewing

venation and genitalic androconia. A new genus

seemed necessary, but hesitating to add another name

to an already nomenclaturally atomized subfamily, I

tested its relationships by cladistic analysis.

The genitalic structures relate to the diagnosis of the

Eurrhypini. The tribe includes nearly two hundred

species in about four dozen genera, with greatest

diversity in Palaeotropical forests. The tribe has been

characterized by three apomorphies of the male

genitalia: (1) a pair of “lamelliform structures” on the

8th abdominal sternite (Leraut & Luquet 1983), (2) a

pair of large, square to oblong “squamiform structures”

attached to the vinculum (Minet 1980; Figs. 8, 9: Sq),

and (3) a plume of long scales in medial position on the

vinculum (Nuss & Kallies 2001). The second and third

structures have been confused in previous studies (see

Discussion), but in this paper, I identify the squamiform

structures with the two square membranes. The

lamelliform structures are bundles of a few robust

chaetiform setae cemented together and directed

posteriad. The chaetae apparently rub against the

numerous transverse ridges of the squamiform

structures. The structures been observed to have a

stridulatory function for male courtship in one species,

Syntonarcha iriastis Meyrick of Australia and Wallacea

(Gwynne & Edwards 1986), but none of the many other

species with the apparatus have yet been studied. The

squamiform and lamelliform structures in S. iriastis are

strongly modiﬁed, so the species was omitted from this

analysis.

In the new species and its Palaeotropical relatives, the

lamelliform structures are absent from the eighth

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sternite (S8) and appear to be functionally replaced by a

pair of lozenge- or ribbon-like ﬁelds on the

intersegmental membrane between S8 and the

vinculum, just anterior of the squamiform structures

(Fig. 8: Sep). These sclerotized ﬁelds bear transverse

ridges so that the outline and texture are reminiscent of

cuttlebones. Reﬂecting the terminology of Minet

(1980) and Leraut & Luquet (1983), I refer to these as

structurae sepiformes (Gk. sepion, cuttlebone).

The Odontiinae has not been subject to previous

phylogenetic analysis, except as a terminal taxon (Solis

& Maes 2002). The results demonstrate that the new

species is not closely related to several other

Neotropical odontiines, despite shared wing patterns

and head structures. In addition, explicit predictions

are derived from the phylogeny about unobserved

characters, including unknown feeding habits. Generic

transfers are made where evidence is sufﬁcient, but

broad nomenclatural changes are not made pending a

larger sample of species and characters (Hayden in

prep.).

MATERIALS AND METHODS

Taxa. The thirty-nine species selected for

phylogenetic analysis include Eurrhypini that share

similar wing pattern and/or sepiform structures. All

known Neotropical Eurrhypini that resemble the new

species in maculation are included. The type species of

genera were included wherever possible. Two

Odontiini serve as outgroups: Cynaeda dentalis (Denis

& Schiffermüller, 1775), the type species and genus of

the Odontiini, and Tegostoma comparale (Hübner,

1796). The ingroup taxa are listed in Table 1 with

information about slide preparations. For the external

characters of many species, additional specimens were

examined from the same collections. Species used in

this analysis were identiﬁed by comparison with original

descriptions, digital photographs of type material, and

revisions and faunal treatments.

The Neotropical eurrhypine genera were transferred

in Munroe (1995). Leraut & Luquet (1983: 528)

indicated that Hyalinarcha Munroe, Metrea Grote,

FIGS. 1–6. Dorsal habitus of wings and frontoclypeus. 1, Suinoorda maccabei holotype m, right half. 2, S. maccabei paratype f,

left half (reﬂected) (McCabe Coll.). 3, Pseudonoorda distigmalis f(Congo, Likouala Region: CMNH). 4, Cliniodes paradisalis (Ja-

maica, Moneague: AMNH). 5, S. maccabei m, frontal aspect of head. 6, Dicepolia ruﬁtinctalis, frontal aspect of head (USNM). Fc,

frontoclypeus; CH, prothoracic coxal hairs.

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Pseudonoorda, and Viettessa Minet belong in the

Eurrhypini based on possession of the apomorphic

structures. The following Old-World genera used in

this study are hereby placed in the Eurrhypini because

their type species possess the structures: Aeglotis Amsel,

Autocharis, Clupeosoma, Deanolis Snellen, Ephelis

Lederer, Hemiscopis Warren, Heortia Lederer,

Hydrorybina Hampson, Noordodes Hampson, and

Pitama Moore. The following misplaced species are

transferred to the Eurrhypini for the same reason:

Epipagis ocellata (Hampson 1916) and Mecyna catalalis

Viette 1953.

The terminal “Pseudonoorda brunneiﬂava” is a

composite of two species: the female is identiﬁed as

Pseudonoorda brunneiﬂava Munroe, and the male is a

closely related undescribed species. The new species

has the forewing postmedial line closer to the distal

wing edge and the terminal area yellow like the median

area, rather than violet. Consequently, the wing pattern

(char. 4) is coded as both states because P. brunneiﬂava

and P. metalloma have a violet terminal area.

The following sources provided specimens used in

this analysis: American Museum of Natural History,

New York (AMNH); Academy of Natural Sciences,

Philadelphia (ANSP); Carnegie Museum of Natural

History, Pittsburgh (CMNH); Canadian National

Collection of Insects, Arachnids and Nematodes,

Ottawa (CNC); Cornell University Insect Collection,

Ithaca (CUIC); Instituto Nacional de Biodiversidad,

Costa Rica (INBio); Muséum d'Histoire Naturelle,

Genève (MHNG); Museo del Instituto de Zoología

Agrícola, Maracay (MIZA); Muséum National

d'Histoire Naturelle, Paris (MNHN); The Natural

History Museum, London (NHM); New York State

Museum, Albany (NYSM); Musée Royal de l'Afrique

Centrale, Tervuren (RMCA); Naturalis, Leiden

(RMNH); Senckenberg Naturhistorische Sammlungen

Dresden (Museum für Tierkunde: MTD); United States

National Museum, Washington, D.C. (USNM);

Museum für Naturkunde, Humboldt Universität, Berlin

(ZMHB); Staatliche Naturwissenschaftliche Sammlung

Bayerns, München (ZSM).

Preparation. Specimens were examined under

tungsten-ﬁlament light with a Wild Heerbrugg

stereomicroscope. Measurements were taken with an

ocular micrometer at 1000X, and coloration was

inspected under incandescent light and compared with

the Methuen Handbook of Colour, 2nd ed. (Kornerup

& Wanscher 1967). Photomicrographs were taken with

a digital Nikon DX1 camera and Microptics

photomicrographer.

Specimens were dissected according to Robinson

(1976). Abdomens were macerated for 20 mins. in 10%

aqueous KOH solution in a simmering water bath and

dissected in water. After inspection of deciduous hairs

and scales, genitalia were cleaned with size 3/0 sable

hair brushes and a snipe pinfeather, stained with

Chlorazol black E, dehydrated 24hr in 95% ethanol, and

mounted in Euparal. Some dissections were stored in

glycerin for several months during the process of

character coding to observe 3-dimensional structures.

Wings were prepared by wetting in 95% ethanol,

immersion in acetone for 30s., and denudation of scales

in water. Wings were soaked in weak bleach solution

(20%) for 60s., brushed again, stained in Eosin-Y (1g /

70% ethanol) for ca. 1 hour, and dehydrated and

mounted as for the genitalia above.

Additional slides by E.G. Munroe and preparators in

the CNC were examined (Table 1). These were often

less informative, as the androconia, scales, and even the

squamiform structures were often removed.

Characters. Forty-eight characters were coded in

the context of a larger survey of Odontiinae (Appendix

A). Characters for phylogenetic analysis were drawn

from the external morphology of the head (4), wings

(11), tympanal organs (1), male genitalia (22), female

genitalia (9), and larval feeding habit (1). Terminology

follows Klots (1956), Kristensen (2004), and the

LepGlossary (Leptree Team 2008), and terms for

tympanic organs follow Maes (1995). Sources of

hostplant information are listed in Table 2. The states

were delimited with consideration of plausible

physiological and behavioral constraints while

maximizing grouping information for the taxon sample.

Characters are numbered from 0; see Appendix B for

descriptions of character states. In the Results and

Discussion, parenthesized numbers refer to

“(character:state).” Some characters were coded as

inapplicable if they depend on the presence or absence

of another character. Unobserved characters include

antennal ciliation for Dicepolia munroealis (Viette)

(char. 0), the vincular androconia for Autocharis

barbieri (Legrand), D. munroealis and Sobanga rutilalis

(Walker) (char. 30), female genitalia for Aeglotis

argentalis (Christoph) and Clupeosoma atristriatum

Hampson (chars. 37–45), and feeding habit for most

terminals (char. 46). Characters were coded for more

than one state where a structure either did not clearly

belong to one state or where the terminal was

polymorphic (exhibiting more than one discrete state

among specimens).

Phylogenetic analysis. The data matrix was

entered with WinClada v. 1.00.08 (Nixon 2002)

(Appendix B). Electronic ﬁles are available from the

author. The matrix was analyzed with parsimony with

TNT v. 1.1 for Windows (Goloboff et al. 2003, 2008) on

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TABLE 1. Species and specimen preparations included in the analysis. Slide numbers refer to preparations by the author (JEH) unless

indicated by preparator’s initials.

Taxon Sex Locality Slide no. Collection

Aeglotis argentalis (Christoph 1887) 1m Pakistan: Kohistan 270 MTD

Argyrarcha margarita (Warren 1892) 2m, 1f Brasil: Rio Gr. do Sul 126, 127, 427 CUIC

1f Br: Sta. Catharina 2760 M. d’A. CNC

Autocharis albiplaga (Hampson 1913) 1m, 1f South Africa: Cape Prov. 93, 119 USNM

Autocharis barbieri (Legrand 1965) 1m, 1f Seychelle Is. 94, 120 USNM

Autocharis fessalis (Swinhoe 1887) 1m, 1f India: Orissa 49, 50 USNM

Autocharis mimetica (Lower 1903) 1m, 1f Australia: NT 189, 190 NYSM

Cliniodes costimacula (Hampson 1913) 1m Venezuela: Aragua: Rancho Grande 48 USNM

1f Costa Rica: Prov. Limón 210 INBio

Cliniodes opalalis Guenée 1854 1m Peru: Pasco Dept. 59 CUIC

1m Peru: Huanuco 61 AMNH

1m Venezuela: Tachira 64 MIZA

1m Peru: Divisoria 66 USNM

1m Bolivia: Cochabamba 162 CNC

1m Costa Rica: Puntarenas: Monte Verde 195 AMNH

1f Costa Rica: Puntarenas: Monte Verde 172 CUIC

1f Jamaica 1107 M. d’A. CNC

Cliniodes ostreonalis (Grote 1882) 1m, 1f Canada: Que.: Norway Bay 137, 138 CNC

1m Canada: Ont.: Merivale 1319 EGM CNC

1f Canada: Ont.: Merivale 2911 M. d’A. CNC

Cliniodes paradisalis (Möschler 1886) 1m Jamaica: Port Antonio 17 AMNH

1m Jamaica: St. Andrew Parish 269 ANSP

1m Jamaica 2553 EGM CNC

1f Jamaica: Trelawny 154 CNC

Clupeosoma atristriatum Hampson 1917 2m PNG: New Britain: near Keravat 121, 122 CNC

Clupeosoma sericiale (Hampson 1896) 1m, 1f Philippine Is: Luzon 53, 54 USNM

Clupeosoma orientalale (Viette 1954

[1953]) 1m Madagascar Est 258 MNHN

1f Madagascar Est 259 RMCA

Clupeosoma vohilavale (Marion & Viette

1956) 1m, 1f Madagascar: Maroantsetra 256, 257 MNHN

Cynaeda dentalis (Denis & Schiffer- 1m, 1f Germany: Rheingau: Loreley 38, 39 USNM

Deanolis sublimbalis Snellen 1899 1m Malaysia: Sabah: Kinabalu NP 130 USNM

1f Philippine Is: Samar 254 ZMHB

Dicepolia munroealis (Viette 1960) 1m Madagascar: Lakato route Holotype, prep. unkn. MNHN

1f Madagascar: Anosibe 288 MHNG

Dicepolia roseobrunnea (Warren 1889) 1m Rio Iça 175 CUIC

1m Fr. Guiana: St. Jean de Maroni BM 22137 NHM

1m Brasil: Upper Amazon, Codajas BM 22138 NHM

1m Brasil: Estado Paraíba 111,915 Heinrich USNM

1f Brasil: Estado Paraíba 224 USNM

Dicepolia ruﬁtinctalis (Hampson 1899) 1m Guyana 79 CUIC

1m Venezuela: Amazonas 81 MIZA

1f Bolivia: Cochabamba 167 CNC

1m, 1f Mexico: Ver.: Jalapa 232, 233 USNM

1m, 1f Peru: Avispas 234, 235 CNC

1m, 1f Panama: Canal Zone 236, 237 USNM

1m Bolivia: Cochabamba 238 USNM

1f Brasil: Distr. Fed. 239 USNM

1m Brasil: Rondonia 246 LACM

Ephelis cruentalis (Geyer 1832) 1m Hamfelt Coll. 44 USNM

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TABLE 1. (continued)

Taxon Sex Locality Slide no. Collection

Ephelis cruentalis (Geyer 1832) 1m Hamfelt Coll. 44 USNM

1m [unknown] 330 CMNH

1f Italy: Basilicata: Monticchio 45 USNM

1f Turkey: Dorah Rober 271 RMNH

Epipagis ocellata (Hampson 1916) 1m Congo: Lulua 307 RMCA

1f Congo: Bokuma 308 RMCA

Eurrhypis pollinalis (Denis & Schiffermüller 1775) 1m, 1f [illeg.] Hamfelt Coll. 132, 133 USNM

Hemiscopis suffusalis (Walker 1866 [1865]) 1m, 1f Philippines: Los Baños 196, 320 USNM

1m Philipplines: Mindanao 417 CNC

1m Sri Lanka: Kandy 1652 EGM CNC

1f China: Hainan Is. 78 CUIC

1f Sri Lanka: Colombo 1653 EGM CNC

Heortia dominalis (Lederer 1863) 1m, 1f Philipplines: Mindanao 324, 325 RMNH

Heortia vitessoides (Moore 1885) 1m Cambodia 75 CUIC

1m Sri Lanka: Galle Distr. 326 USNM

1f Sri Lanka: E. Distr. 152 USNM

1m Indonesia: Borneo 2555 EGM CNC

Hyalinarcha hyalinalis (Hampson 1896) 1m PNG: Dagua Rd., Wewak 115 CNC

1f PNG: Morobe 116 CNC

1m PNG: Wewak 3028 DK CNC

2m India: Assam: Margherita 3024 DK, 3032 DK CNC

Hydrorybina polusalis (Walker 1859) 1m Laos 74 CUIC

1f Sri Lanka: Kandy 4819 DK CNC

1f Philippines: Luzon 321 ZMHB

Mecyna catalalis Viette 1953 1m Madagascar: Analamazaotra 279 MNHN

1f Madagascar: Anosibe route 280 MNHN

Mecynarcha apicalis (Hampson 1898) 1m, 1f Guyana: Rupununi 51, 52 USNM

1f ? Ellsworth Collection 155 CUIC

1m Brasil: Amazonas 161 CMNH

Mimoschinia rufofascialis (Stephens 1834) 1m, 1f USA: Virgin Is.: Kingshill 57, 58 CUIC

1m USA: AZ: Pima Co. 87 CUIC

1f USA: TX: Uvalde 88 CUIC

1f Mexico: B.C. Sur 272 CMNH

Noordodes magniﬁcalis (Rothschild 1916) 1m, 1f New Guinea: Irian Barat 249, 250 ZSM

Pitama hermesalis (Walker 1859) 1m, 1f Malaysia: Sabah 197, 204 USNM

Pseudonoorda brunneiﬂava Munroe 1974 1f Indonesia: N. Sulawesi 261 RMNH

Pseudonoorda sp. near brunneiﬂava Munroe 1974 1m Philippine Is: Luzon 260 MTD

Pseudonoorda distigmalis (Hampson 1913) 1m Cameroon: Efulen 20 CMNH

1m, 1f Cameroon: Efulen 117, 118 CNC

Pseudoschinia elautalis (Grote 1881) 1m, 1f USA: Arizona 139, 140 CUIC

Sobanga rutilalis (Walker 1862) 1m Venezuela: Amazonas 141 CUIC

1m, 1f Brasil: Amazonas 142, 143 CUIC

1m Brasil: Téffé 1117 M. d’A. CNC

Suinoorda maccabei sp. nov. 3m, 1f Bahamas: Great Exuma 186, 187, 188, 274 CUIC

1m Cuba: Holguín 273 MTD

Tegostoma comparale (Hübner) 1m, 1f S. Russia 27, 28 USNM

Viettessa bethalis (Viette 1958) 1m Cameroon: Bonenza 264 RMCA

1f Congo: Paulis [Isiro] 319 RMCA

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a Dell Latitude D610 PC. The commands were “rseed

1; hold 1000; collapse auto; mult= hold 10 replic 20;”

The same commands can be implemented by selecting

Settings / Memory / Max. trees = 1000, then entering

search parameters under Analyze / Traditional Search.

“Collapse auto” ensures that branches with no

apomorphies are collapsed.

During preliminary character analysis, additivity was

explored for some characters, and results that implied

homoplasy in complex characters were reanalyzed. In

ﬁnal analysis, characters were treated as non-additive

and equally weighted, and polarity was determined

from the results of unconstrained analysis with C.

dentalis as the primary outgroup (Nixon & Carpenter

1993). Inapplicable data are represented by “-” and

missing data by “?,” but both are analyzed allowing all

possible states to be considered. The matrix was

analyzed as-is, but to assess tree length correctly, a

second matrix was analyzed where the polymorphic

terminals were divided into multiple terminals (Nixon &

Davis 1991).

The apomorphies of terminals and clades are

interpreted to be their diagnoses (Farris 1979).

Character evolution was traced with the Character

Diagnoser function of Winclada and TNT. Fast and

slow optimization herein respectively refer to

accelerated and delayed transformation (Swofford &

Maddison 1987) and were implemented with WinClada.

Bremer support values for clades were calculated by

expanding memory for trees, generating many

suboptimal trees up to Nextra steps, and searching

among those for the shortest tree that lacks a given node

(“hold 80000; subopt N; bbreak=ﬁllonly; bsupport”).

The values of missing data were predicted to be those

that best agree with the optimality criterion in

simultaneous analysis. For parsimony, this is all possible

permutations for unscored cells that add no extra steps.

See Wilkinson (1995: Table 1) for an antecedent.

Possible states were restricted to those actually

observed in scored taxa. Predictions were made by

inspecting mapped characters with WinClada, then

running a simple procedure ﬁle in TNT to check the

ambiguous cases. The ﬁle, available from the author,

uses the “xread =” command to replace “?’s” with

alternative states.

RESULTS

Phylogenetic analysis. Searches found 1 tree of 187

steps (consistency index = 0.35, retention index = 0.67)

(Fig. 7). Most of the cells coded for more than one state

represent uncertainty of state assignment. There are

four real polymorphisms: the wing pattern (char. 4) of

the new species and the composite Pseudonoorda

brunneiﬂava, and the feeding habit (char. 47) of two

other species (Table 2, footnotes). Splitting these four

into eight terminals, each pair differing only in the state

of the polymorphism, resulted in the same topology

with length 191. Branches without support are

collapsed as polyotomies. Character state changes are

reported in Appendix C.

TABLE 2. Sources of hostplant data.

Species References

Autocharis barbieri Gerlach and Matyot 2006

Autocharis fessalis Beeson 1961, Browne 1968

Cliniodes opalalis Janzen & Hallwachs 2005

Cliniodes ostreonalis* McDunnough 1931, Munroe 1961, Hayden 2008

Cynaeda dentalis Slamka 2006, Huertas Dionisio 2007

Deanolis sublimbalis Waterhouse 1998

Dicepolia roseobrunnea de Oliveira 1942 [1941]

Eurrhypis pollinalis Slamka 2006, Huertas Dionisio 2007 (for E. guttulalis (Herrich-Schäffer))

Hemiscopis suffusalis** Beeson 1961, Tominaga 1999, Turner 1908

Heortia vitessoides Munroe 1977, Singh et al. 2000

Hydrorybina polusalis Meyrick, E. 1938 MS, in Robinson et al. 2001

Mimoschinia rufofascialis Heinrich 1921, Leech 1949

Pseudoschinia elautalis Mann 1969

Tegostoma comparale Huertas Dionisio 2007

* Folivore on Thymelaeaceae and Rhamnaceae. Coded as polymorphic.

** H. suffusalis on Dipterocarpaceae and Phyllanthaceae, but H. purpureum (Inoue) and H. violacea (Lucas) on Thymelaeaceae.

Phyllanthaceae produce alkaloids like those in Thymelaeaceae. Coded as polymorphic.

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The sepiform clade (node 64, S) is strongly supported

as monophyletic, with Bremer support of 5 (i.e. the next

shortest trees from which the clade is absent are 5 steps

longer) (Bremer 1988). The clade has ﬁve

unambiguous apomorphies: lamelliform structures

absent (21:0), sepiform structres present (22:1, uniquely

derived), distal edge of squamiform structures rounded

(26:1, uniquely derived, reversed to squarish in the

Pseudonoorda brunneiﬂava-group), squamiform

structures with central enations (27:1), cervix bursae

large and sclerotized (40:2).

The new species is the ﬁrst-diverging terminal in the

sepiform clade. The Palaeotropical species that

constitute the remainder of the clade (node 63) share

three unambiguous synapomorphies: Rs1stalked with

Rs2+3 (11:1), loss of S8 piluli (23:0), and elongate

ovipositor (38:1). The results indicate that

Pseudonoorda is paraphyletic.

The sister group of the sepiform clade (node 42, L)

includes medium- to large-bodied species of global

FIG. 7. Cladogram (187 steps, CI = 0.35, RI = 0.67). Numbers above branches refer to nodes (Appendix C), and numbers below

are Bremer support values (suboptimality of trees lacking the node). NA, Nearctic; NT, Neotropical; PA, Palaearctic; PT,

Palaeotropical; E, Eurrhypini; L, leaf-feeding clade; S, sepiform clade. *A. margarita is also known from Madagascar (Munroe

1974). **C. ostreonalis is Eastern Nearctic.

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distribution. The clade’s two apomorphies are an

extension of the forewing costal streak through the

discal spot that approximates or reaches the postmedial

line (6:1) and the shallow depth of the sacci

tympanorum extending underneath S2 (15:1). The

group includes several members that feed on

Thymelaeaceae, a plant family that produces unusual

toxic terpenoid compounds (Evans 1986).

Deactivation of char. 47 (feeding habit) results in

almost no change. Three trees of 183 steps are

recovered that differ in minor aspects that do not affect

the sepiform clade, thymelaeacean clade, or any

nomenclatural conclusions drawn here. Combining

states 2 and 3 into one state (general folivory) returned

the same topology with one fewer step.

Systematic section. Clupeosoma orientalale (Viette)

comb. nov. and Clupeosoma vohilavale (Marion &

Viette) comb. nov. are transferred from Autocharis, as

they share synapomorphies with Clupeosoma sensu

Munroe (1974b) (see Discussion). Hyalinarcha

hyalinalis (Hampson) comb. nov. is transferred from

Boeotarcha Meyrick, as it is closely related to the type

species H. hyalina (Hampson), females of which were

not available for study. Noordodes magniﬁcalis

(Rothschild) comb. nov. is transferred from Noorda

Walker.

The synonymy of Basonga Möschler, 1886 (type

species B. paradisalis Möschler) with Cliniodes

Guenée, 1854, last recognized in Klima (1939), is

revived, as is the combination Cliniodes paradisalis

(Möschler). Metrea Grote, 1882 is synonymized with

Cliniodes, syn. nov., resulting in the combination

Cliniodes ostreonalis comb. nov.

Suinoorda Hayden gen. nov.

Diagnosis. Unambiguous autapomorphies: female

with entire forewing solid orange, concolorous with

terminal area (4:1, 2; Fig. 2); lateral arms of gnathos

approximated to lower corners of uncus (36:1).

Symplesiomorphies not shared with rest of sepiform

clade: forewing Rs1not stalked with Rs2+3 (char. 11:0;

Fig. 10); piluli present on S8 (char. 23:1; Fig. 12: Pi);

androconium of long, straight setae present near base of

valval costa, dorsal side (28:1; Fig. 18: BA); ovipositor

short (38:0; Fig. 16). Otherwise sharing the

apomorphies of the sepiform clade of Eurrhypini (see

above).

Two other diagnostic characters have ambiguous

optimization: forewing antemedial line absent (8:0,

autapomorphic with slow optimization or

synapomorphic with fast optimization); basicostal

androconium present as ﬁeld of deciduous hairs from

base of costa (28:1, Fig. 18: BA; autapomorphic with

slow optimization or symplesiomorphic with fast

optimization).

Etymology: Latin sus, swine, referring to the shape

of the frontoclypeus, and also the reﬂexive pronoun,

referring to taxonomic rank; plus Noorda Walker.

Gender, feminine.

Type species: S. maccabei Hayden sp. nov.

S. maccabei Hayden sp. nov.

Description. Male habitus (ﬁgs 1, 5, 13, 14, 15): Head. Frons

ﬂat and oblique. Frontoclypeus expanded in a low cylindrical

protuberance, lateral edges of ridge curling down and inward almost

to secondary contact above pilifers; lower edges of frontoclypeus

crenulate (ﬁg. 5: Fc). Frons with smooth, yellow-beige scales, dark

brown laterally on protuberance. Vertex rough, yellow-beige. Scales

between antennae and eyes white. Ocelli present. Chaetosemata

absent. Antennae round and smooth, about 3/5 length of forewings,

cilia dense and longer than in female. Labial palpi dark brown above,

white below; not greatly exceeding length of head, length 0.90±0.03

mm from base under head to apex; porrect or slightly drooping with

third meron downturned ~30° from porrect axis. Maxillary palpi short

(0.36±0.03 mm), brown, terminal scales not strongly dilated. Pilifers

small and separate from lower corners of frontoclypeus. Haustellum

well-developed, basal vestiture white.

Thorax. Collar, distal tegulae and dorsal thorax yellow-beige.

Tegulae proximally brownish red. Ventral thorax and coxae white.

Forelegs: coxa white mixed with light gray, with patch of long, yellow-

gray, backward-sweeping hairs extending the length of anterior margin

of coxa (ﬁg. 5: CH); femur and tibia gray, the latter with epiphysis

hidden in long scales, without androconium; tarsomeres white, all but

the basal one with small dorsal gray patch. Midlegs: femur and inner

surface of tibia white; outer tibial surface orange fading to yellow

before white, with gray patch at joint with femur; tibia bearing

androconium of straight, white hairs as long as tibia (ﬁg. 14: An); inner

tibial spur 3 times length of outer; tarsi as for foreleg. Hindlegs:

uniformly white, no androconia; spurs as fore midleg; tarsi as for

foreleg.

Abdomen. Praecinctorium white-scaled, monolobate, as broad as

deep. Abdomen dorsum yellow (hindmost scale row white), venter

white. Male abdomen 4.8±0.1 mm in length excluding genitalia.

Genitalic androconia pale yellow. Eighth sternite without scales (ﬁg.

15: S8).

Forewing. Length, 7.2 – 7.7 ±0.1mm, width, 3.6 – 4.0 ±0.1mm (n

= 6). Upper side pale yellow with scales tipped in yellow, salmon or

fuscous, giving irrorate appearance in fresher specimens. Costa

thickly scaled and colored, with longer, setiform scales on edge;

proximally gray along leading edge; distally and inner proximally dark

salmon-orange. Antemedial line represented by minuscule brown

spot behind costa at 1/5 length and faint spot or line along dorsal edge

at 3/8 length. Discal spot large, oval, dark brown, beyond 1/2 length

of wing, 0.42 ±0.06mm wide by 0.66 ±0.06mm long. Postmedial line

narrow, dark brown, smoothly curving convexly from costa (at Rs4) to

CuA1and concavely from CuA2to tornus. Postmedial area 1.0±0.1

mm wide, orange or dark salmon. Terminal fringe dark gray or

fuscous. Dorsal edge of forewing without scale tufts. Fovea absent.

Forewing underside pale yellow, with costa and terminal area orange.

Male-type retinacular hook and female-type retinaculum both

present.

Hindwing. Length, 6.2 ±0.1mm by 3.6±0.1 mm; lustrous white,

with long, sparse hairs on anal area. Anal edge smoothly curved.

Terminal area pale orange with some scales brown-tipped in darker

specimens. Postmedial line faint, brown, from M2to anal fold, where

it meets the terminal edge; white behind anal fold (a few brown scales

extend proximally along anal fold in fresher specimens). Terminal

fringe yellowish brown. Hindwing underneath translucent white, with

yellow costa and terminal fringe. Male frenulum a single bristle.

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FIGS. 8–12. Genitalia and wings. 8, Suinoorda maccabei mgenitalia, stridulatory apparatus (JEH 187). 9, Cliniodes ostreonalis m

genitalia, stridulatory apparatus, squamiform structures slightly displaced (JEH 137, CNC). 10, S. maccabei mwing venation (JEH

274). 11, S. maccabei mgenitalia, entire (JEH 187). 12, same, eighth abdominal segment. En, enation of squamiform structure;

F, ﬁeld of scales in fringe on tergum 8 posterior edge; Fr, frenulum; Lm, lamelliform structure; Lo, lobe of S8; Pi, location of piluli;

Sep, sepiform structures; Sq, squamiform structures; S8, eighth sternite; T8, eighth tergite.

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Dense patch of small, erect scales between costa and Sc+R, from base

of wing to the divergence of Sc+R1and Rs; patch extended discretely

but more sparsely onto anterior half of discal cell (ﬁg. 13: P).

Female habitus (ﬁg. 2): Structure and coloration as in male,

except dorsum of head, thorax and forewings almost uniformly

salmon-orange. Tegulae and lateral collar dark brown; midleg tibia

upper surface more strongly orange, without androconia. Antennal

cilia shorter than in male, but antenna itself of same thickness.

Forewing medial and terminal areas concolorous, with the latter

slightly more densely scaled. Forewing antemedial line absent; discal

spot and postmedial line indistinctly indicated by light-brown scales.

Retinaculum a patch of scales as in male. Two frenular bristles.

Hindwing underside without subcostal patch of scales. Hindwing

terminal area darker yellow than rest of hindwing, but lines absent.

Abdomen 4.4±0.2 mm in length excluding ovipositor; exceeding

hindwings by about one third length.

Venation (ﬁg. 10): Forewing: Sc meeting margin 2/3 from wing

apex. R1basally straight and moderately broadened. Rs1basally

straight, not stalked with Rs2+3. Rs2+3 with stalk curved to approximate

Rs1closely. Sc and radial veins except Rs4distally curved toward

anterior margin, the ﬁrst three veins suddenly and sharply so. Rs4

unstalked, basally curved toward Rs2+3, distally meeting margin below

wing apex. Cell complete, 7/12 length of wing. M1nearly straight,

from upper corner of discal cell. M2and M3straight, from lower

corner of cell. CuA1and 2straight, arising from cell basad from tip.

1A straight, complete; 2A looped and joining 1A at half its length.

Hindwing: Sc+R1stalked to about 1/3 length from point of

departure from cell, partly joined to Rs to just beyond discal cell. Base

of Sc+R1+Rs strongly broadened. Discal cell short: anterior portion

1/3 length of wing, distally demarcated by short, straight spur of M1

parallel to body axis; posterior portion demarcated by straight vein

running diagonally to nearly 1/2 length of wing. M1straight. M2and

M3basally approximate, not stalked, from lower corner of cell. CuA1

from corner of cell; CuA2from cell at just more than 2/3. CuP and

1A+2A straight; 3A basally straight and broad, distally slightly bowed

anteriad before meeting margin.

Tympanal organs (ﬁg. 17): Bullae tympani oval, length twice

width. Edges of fornix tympani straight and meeting in blunt, elbow-

like right angle. Sacci tympani round, not hypertrophied (ﬁg. 17: ST);

rami tympani (transverse edge) on same level as elbow of fornix

tympani. Processus tympani a broad, nearly circular lobe. Puteoli

tympani shallow. Venulae not evident. S2 a rounded rhombus, wider

than long, bluntly protruding over S3; median third of S2 also

emergent and slightly more sclerotized than lateral areas.

Male genitalia (ﬁgs 8, 11, 12, 18): Eighth segment. T8

membranous, longer than wide; lateral sclerotized edges of T8 with

anterior ends ending in shallow, posteriad-facing invaginations,

posterior ends fading before fringe; transverse bar absent. Transverse

fringe of long, broad, keeled scales present along posterior edge of T8

(ﬁgs 12, 15: F). Lateral ends of fringe separate as distinct lobes in

pleuron 8, slanted dorso-anterior to ventro-posterior and closely

adjacent to posterior corners of S8. S8 roughly rectangular, parallel-

FIGS. 13–19. Habitus and genitalia. 13, Suinoorda maccabei underside of mwings. 14, S. maccabei mmesothoracic tibia, lateral

aspect. 15, S. maccabei mabdomen, lateral aspect. 16, S. maccabei fgenitalia (JEH 188). 17, same, tympanal organs. 18, S. mac-

cabei mgenitalia, dorsal aspect (JEH 187). 19, Autocharis barbieri mgenitalia, ventral aspect (JEH 94, USNM). An, androconium

of long hairs; BA, basicostal androconium; C, colliculum; CvB, cervix bursae; DS, ductus seminalis; F, tergum 8 posterior fringe; P,

patch of scales on hindwing subcostal area; ScS, scoop-shaped scales along costa; ST, saccus tympani; S8, sternite 8 (naturally de-

nuded); VA, vincular androconium (pleural A8-9 intersegmental).

sided; anterior quarter underneath S7 and bilobate, with broadly

triangular emargination and medially sclerotized to halfway point.

Edges sclerotized, most strongly along anterior third. Posterior edge

with squared corners and central third broadly, roundly emarginate,

without projections. Central area of S8 trapezoidal, slightly raised,

with corners at sternal margin at anterior third to posterior

emargination. Posterolateral corners of S8 with area of ﬁne

nondeciduous setae (piluli, char. 22; ﬁg. 12: Pi). Lamelliform

structures absent. Androconia of pleuron 8 on large, lance-ovate

patches: dorso-anterior end rounded and adjacent to anterior corners

of T8, ventro-posterior end tapered and connected directly to the

sides of S8 at the point where S8 emerges from underneath S7.

Genitalia. Uncus an isosceles triangle about twice as high as width

at base, with expanded, ﬂat-topped, bilobate apex bearing conspicuous

tufts of nondeciduous setae at tip of each lobe; rows of fainter hairs on

lateral edges of uncus, but elsewhere bare. Arms of gnathos

transversely straight, with broad bases fused to tegumen and medial

area narrow; bases of gnathos arms close to base of uncus. Gnathos

central element parallel-sided, produced 210±15µm, with minute

granules at bluntly acute tip; ventrally deeply indented beyond

(posterior of) the level of the arms. Tegumen almost as long as

vinculum (680±15µm), without ornamentation; with long, straight-

sided lateral arms descending to valvae. Membrane between arms

with diffuse deciduous hairs, not grouped into discrete patches. Juxta

smooth, occupying most of the area inside vinculum and sides

parallelling edges of vinculum; base nearly circular, as high as

squamiform structures, with concave emargination where base of

vinculum bows upward; apex of juxta abruptly cuspidate and acute,

unornamented and not projecting. Valvae lyriform, 1,770±15µm

along costa. Costa sinuate, proximally convex and distally concave,

edge basad of apex membranous and bearing short, smooth, unkeeled

scales that curve over edge of costa (ﬁg. 18: ScS). Costa sclerotized to

apex, with small tuft of short setiform hairs on a slightly out-turned

apex. Distal edge of valva straight. Saccular edge of valva roughly

mirroring costa: subapically concave, medially convex, and basally

indented to accommodate squamiform structures. Convex area of

sacculus with dense, broad scales. Medial area of valva basally

sclerotized and distally membranous with striae and sparse, ﬁne setae.

Area below costa longitudinally depressed as narrow, shallow pocket.

Reverse side of valva with androconium 380±15µm from base of valva,

in submarginal position (behind shallow pocket); base of ﬁeld a

transverse row, 150±15µm long (ﬁg. 18: BA). Vinculum 1,050±20µm

high, roughly square. Transtilla absent. Saccus not developed,

medially concave. Androconia present lateral of vinculum, of straight

hairs as long as valvae (ﬁg. 18: VA). Basal area of androconia extended

to 500±15µm in length, running up behind (dorsal of) vinculum

toward tegumen. Phallus 1,680±15µm long, 170±15µm at narrowest

point, gently curved. Vesica with numerous small cornuti along most

of length.

Squamiform and associated structures. Posteroventral vinculum

(facing juxta) with medial, sclerotized, bicapitate boss. S8–9

intersegmental membrane extended over vinculum, bearing

squamiform structures lateral of boss. Medial pluma of unkeeled

scales not obvious. Membrane laterally extended as semi-sclerotized

arms bearing short tufts of scales parallel to lateral edges of

squamiform structures. Membrane anteriad of vinculum with

thickened pair of longitudinal, parallel ﬁelds bearing transverse ridges

at intervals of about 15µm (ﬁg. 8: Sep). Ridged ﬁelds narrow, each

45±15µm wide and 450±15µm long, with further sclerotization

extending 250±15µm anteriad. Squamiform structures round,

675±15µm long by 400±15µm wide, symmetrical; stiff membranes

attached on lower edge to vinculum (ﬁg. 8: Sq). Medial edge of

squamiform structures strongly sclerotized and smoothly arcuate in

hemiellipse extending over distal end of squamiform structures.

Lateral edges clavate and longitudinally striate, with narrow base and

broad termination before membranous laterodistal area that does not

not meet the arc of the medial edge. Central area of squamiform

structures ﬁnely striate. Basicentral area of each squamiform

structure with digitate process or enation directed medially, adjacent

to boss of accessory sclerite (ﬁg. 8: En). Enations curving around

ridged areas when the latter are folded against them.

Female genitalia (ﬁg. 16): Ovipositor short, 750±15µm long by

450±15µm deep, with A9 not longer than deep. Ovipositor lobes soft

and truncate. Anterior apophyses extending to colliculum; posterior

apophyses not extending beyond anterior edge of A9. Colliculum

short, about twice as long as wide (ﬁg. 16: C). Ductus bursae between

colliculum and corpus bursae proper expanded in an irregularly

chamber (cervix bursae: CvB), demarcated from corpus bursae by

slight constriction and end of sclerotization. Cervix bursae irregularly

but mostly sclerotized. A few granular hooklets irregularly distributed

on the signum. Ductus seminalis narrow and arising from

unsclerotized area of anterior ductus bursae (DS). Corpus bursae

proper (anterior of sclerotized cervix) 1900±30µm long, longitudinally

pleated and bearing two convex, granular signa on opposite sides of

the bursa, the larger and posterior of the two being transversely ovate,

and the anterior one nearly circular.

Type material. Holotype. m: BAHAMAS—Great Exuma—

Simons Pt, 23.31.50—75.47.30, 12 April 1986, Tim L. McCabe / [red

label] HOLOTYPE Suinoorda maccabei Hayden. T.L. McCabe

Collection. Paratypes. BAHAMAS: 1m, same data as holotype

except: 10 April 1986 / J.E. Hayden Slide No. 187 m(JEH Coll.). 1m,

same data as holotype except: 14 April 1986 / J.E. Hayden Slide Nos.

186 m, 274 wings (CUIC). 2mm, same data as holotype except: 15 April

1986 (CUIC and T.L. McCabe Collection). 1f, same data as holotype

except: 15 April 1986 / J.E. Hayden Slide No. 188 f(CUIC). CUBA:

1m: [green label] Cuba, Holguin, Rafael Freyre, Piedra Picar, dry

forest, 9.vi.2000, leg. J.-P. Rudloff / J.E. Hayden Slide No. 273 m

(MTD). CUIC type no. 7356.

Diagnosis: As for genus.

Biology: Unknown. See Missing Data.

Distribution: Commonwealth of the Bahamas:

Great Exuma Island; Republic of Cuba: Holguín

Province (ﬁg. 20).

Etymology: I take pleasure in naming this species

after Dr. Tim McCabe, Curator of Entomology at the

New York State Museum, who collected and ﬁnely set

the Bahamian series.

Variation: The Cuban specimen differs from the

Bahamian ones in having a slightly narrower

frontoclypeal arch and the lateral arms of the gnathos at

a slightly more obtuse angle to the median element.

These features are not of speciﬁc distinction.

Similar species: Suinoorda differs from similar

species in the possession of a highly, almost circularly

arched frontoclypeal margin. Autocharis and Dicepolia

have an obtusely to acutely angulate frontoclypeus, and

Noordodes and P. brunneiﬂava have a low, rounded

arch. S. maccabei also differs in the sexually dimorphic

forewing coloration, and the sharp apical curvature of

the forewing radial veins is unique. Similar species

outside the sepiform clade lack the sepiform structures.

Among similar Neotropical Eurrhypini, Dicepolia

species have long labial palpi, are uniformly colored

brownish orange or rosy brown, and have a dark, narrow

postmedial line farther from the terminal margin.

Female S. maccabei differ in having no trace of ante-

and postmedial lines. Cliniodes paradisalis (ﬁg. 4) has

short, upturned labial palpi typical of Cliniodes. Both

have a smooth postmedial line, but it extends much

farther basad along the posterior wing margin in C.

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FIG. 20. Known distribution of S. maccabei. Starred localities: Simon’s Point, near Georgetown, Great Exuma Island, Bahama

Is., and near Rafael Freyre, Holguín Province, Cuba.

paradisalis, approximating the obvious antemedial line.

The forewing has a discal spot continuous with the

costal streak, and the hindwing is suffused more

strongly. Argyrarcha and Sobanga differ strongly from

S. maccabei in maculation, including a well-developed

antemedial line and the costal streak extending through

the discal spot toward the postmedial line. These also

have an expanded T8 posterior fringe, which in

Argyrarcha and Mecynarcha Munroe is cape-like and

larger than any single tergite.

Among Palaeotropical genera, the red-and-white

Autocharis species have long labial palpi, some (A.

fessalis Swinhoe) have hindwing M2+3 stalked, and some

have a more strongly developed spot of color on the

hindwing anal vein. In Autocharis species, the shape of

the forewing postmedial line tends to be more angulate

on the veins. Pitama hermesalis and Viettessa have a

straight frontoclypeus, a much broader postmedial

band, and most species are much larger in size. Like

some of the Neotropical taxa, Viettessa species have a

costal streak extending through the discal spot toward

the postmedial line. Pseudonoorda brunneiﬂava and

Noordodes have Rs1stalked with Rs2+3. Noordodes has a

postmedial line that steps sharply basad along the

cubital veins. The ground color is yellow and the

markings, dark violet.

Character evolution. The four unambiguous

symplesiomorphies of Suinoorda argue for its early

divergence. The frontoclypeal arch (char. 3:2, ﬁgs 5, 6)

evolved once or twice from a transversely straight

margin (3:0), depending on fast or slow optimization: in

the last common ancestor of Hyalinarcha and

Suinoorda (ﬁg. 7: node 44) or independently in nodes

52 and 64. The length of antennal sensilla (char. 0) is

informative: except for four autapomorphic reversals, it

characterizes nodes 43 and 56, and has ci = 0.42, ri =

0.78.

Forewing maculation with a white or light-colored

median and distinctly darker terminal area (4:1) evolved

one to three times from states 0 or 2 and was lost several

times. It appeared above T. comparale (node 46) and

was retained as the fundamentally plesiomorphic

condition along most of the phylogeny. Under slow

optimization, it evolved once at node 46, but under fast

optimization, it evolves independently in C. paradisalis

and Autocharis (node 50). The concolorous maculation

in both female S. maccabei and Pseudonoorda sp. near

brunneiﬂava, regarded as split terminals, is recovered as

independent reversals from the white/red condition.

The stalking of Rs1with Rs2+3 (char. 8:1) is mainly a

feature of the sepiform clade above Suinoorda (node

63), but it evolved independently in Hydrorybina.

Some species lack the stridulatory apparatus.

Lamelliform structures (21:1; ﬁg. 9: Lm) were gained

with the Eurrhypini (node 46) and lost in S. rutilalis and

the sepiform clade. Likewise, the squamiform

structures (square membranes: char. 24; ﬁgs 8, 9: Sq)

were gained at node 46 and lost in S. rutilalis, rendering

characters 25, 26, and 27 inapplicable. The outgroup

Cynaeda dentalis is classiﬁed in Odontiini, which is

deﬁned by the absence of the lamelliform and

squamiform structures (Leraut & Luquet 1983).

The S8 piluli (char. 23) were gained at node 46

(Eurrhypini) and lost at nodes 41 (the Aeglotis-

Argyrarcha clade), 63 (the sepiform clade above

Suinoorda), and in H. hyalinalis. Some but not all of

these clades have the stridulatory apparatus absent or

modiﬁed (e.g. asymmetrical squamiform structures at

node 47, evolved from symmetrical structures). The

basicostal androconium on the dorsal side of the costa of

the male valve (28:1; ﬁgs 18, 19: BA) evolved from the

absent condition (28:0) at node 44, the last common

ancestor of Hyalinarcha, Suinoorda, and Aeglotis. It

was secondarily lost at nodes 41, 63, 53 (Cliniodes spp.),

and in M. catalalis. An androconium situated halfway

along the costa, at the end of a sclerotized extension

(28:2), is a synapomorphy of derived Cliniodes species

(node 57), and it evolved from the absent condition. A

gnathos with arms basally approximate to the lower

uncus (36:1) evolved three times from the not-

approximated condition (36:0) at nodes 47, 53, and in

Suinoorda.

The ovipositor length (char. 38) is homoplastic (ci =

0.12), but it contains some grouping information (ri =

0.46). It is long in the Palaeotropical sepiform species,

but its short length in Suinoorda contributes to

grouping that clade with the large-bodied species of

node 42.

Stem boring and leaf mining (47:0) is the primitive

eurrhypine feeding habit, and it is restricted to the

outgroups (Odontiini) and Eurrhypis Hübner and is

predicted for Ephelis. Seed- or fruit-feeding (47:1)

evolved from it and is the most general larval feeding

habit. It is known for the three distantly related groups

Mimoschinia Munroe + Pseudoschinia Munroe,

Dicepolia roseobrunnea (Warren), and Deanolis, and

predicted for the intervening taxa. External folivory on

Thymelaeaceae (47:2) is restricted to the clade of node

42 and evolved once or twice from seed-feeding, in

Hemiscopis and in node 54, or earlier depending on

optimization. Folivory on other plant families (47:3)

evolved independently from seed-feeding in Autocharis

and from either seed- or Thymelaeaceae-feeding in

Hemiscopis + Hydrorybina. Predictions for unscored

data are reported in Table 3.

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TABLE 3. Predicted missing data. The following states add no extra steps to the cladograms when analyzed together.

Species Character Predicted states

A. argentalis 38. Ovipositor length short

39. Colliculum short

40. Duct. burs. sclerotization absent

41. Appendix of duct. burs. absent

42. Corp. burs. signum present (one or two)

43. Corp. burs. signum, shape round

44. Ductus seminalis origin from ductus bursae

45. Ductus seminalis width narrow

46. Pleats on cervix bursae absent, or present

47. Feeding habit seeds/fruits, or folivore on Thymel., or folivore on other family

A. margarita 47. Feeding habit seeds/fruits, or folivore on Thymel., or folivore on other family

A. albiplaga 47. Feeding habit folivore on other family

A. barbieri 31. Vincular androconia present

A. mimetica 47. Feeding habit seeds/fruits

C. costimacula 47. Feeding habit folivore on Thymel.

C. paradisalis 47. Feeding habit folivore on Thymel.

C. atristriatum 38. Ovipositor length long

39. Colliculum short

40. Duct. burs. sclerotization absent

41. Appendix of ductus bursae absent

42. Corpus bursae signum present (one or two)

43. Corp. burs. signum, shape round

44. Ductus seminalis origin from ductus bursae

45. Ductus seminalis width enlarged

46. Pleats on cervix bursae present

47. Feeding habit seeds/fruits

C. orientalale 47. Feeding habit seeds/fruits

C. sericiale 47. Feeding habit seeds/fruits

C. vohilavale 47. Feeding habit seeds/fruits

D. munroealis 0. Antennal sensilla longer in male

31. Vincular androconia absent

47. Feeding habit seeds/fruits

D. ruﬁtinctalis 47. Feeding habit seeds/fruits

E. cruentalis 47. Feeding habit stem borer or leaf miner

E. ocellata 47. Feeding habit seeds/fruits, or folivore on Thymel., or folivore on other family

H. dominalis 47. Feeding habit folivore on Thymel.

H. hyalinalis 47. Feeding habit seeds/fruits

M. catalalis 47. Feeding habit seeds/fruits, or folivore on Thymel., or folivore on other family

M. apicalis 47. Feeding habit seeds/fruits, or folivore on Thymel., or folivore on other family

N. magniﬁcalis 47. Feeding habit seeds/fruits

P. hermesalis 47. Feeding habit seeds/fruits, or folivore on Thymel., or folivore on other family

P. brunneiﬂava 47. Feeding habit seeds/fruits

P. distigmalis 47. Feeding habit seeds/fruits

S. rutilalis 31. Vincular androconia present

47. Feeding habit seeds/fruits, or folivore on Thymel., or folivore on other family

S. maccabei 47. Feeding habit seeds/fruits

V. bethalis 47. Feeding habit seeds/fruits, or folivore on Thymel., or folivore on other family

DISCUSSION

The recent collection of this new species recalls

Hampson’s comment on the Bahamian moth fauna:

“[…] I should expect a thorough exploration of the

other islands [than Nassau and Andros] to add

considerably to the list of species” (1901). Hampson

studied only specimens from those islands and Abaco

(1901, 1904); had he seen it, he probably would have

assigned it to Noctuelia Guenée or Noorda Walker

following his classiﬁcation of Pyraustinae (Hampson

1899a). I ﬁnd no reference to a species ﬁtting this

description in other studies of Bahamian and Cuban

Lepidoptera (Smith et al. 1994 and citations therein).

The distribution of this species in the Bahamas and

Cuba (Fig. 20) can be explained by the subaerial

exposure of the Bahama Platform during the last glacial

maximum, when the islands were separated by only the

narrow Old Bahamas Passage (Miller & Miller 2001).

The Cuban locality is about 270km due south of the

type locality, but this distribution is congruent with that

of other Lepidoptera (Miller & Simon 1998). The

species may be expected to be distributed more widely

in eastern Cuba and the islands that are part of the

Great Bahama Bank. Another species, Cautethia exuma

McCabe (Sphingidae), was described from the same

locality and is apparently endemic to Great Exuma

(McCabe 1984).

The ﬂora of the type locality is a mix of native and

naturalized species (T. McCabe pers. comm. 2007):

Caesalpinia L., Casuarina L., coconut, Coccoloba L.,

Erithalis G. Forst., Ficus L., frangipani (Plumeria rubra

L.), Key Lime, Malpighia L., Mimosa L., orange, and

“old woman's tongue,” a legume. The Cuban locality is

mixed forest: mostly secondary growth with native dry-

forest undergrowth and relictual vegetation in the

vicinity characteristic of mogotes; dominant trees

include Acacia Mill., Caesalpina, Coccoloba, coconut,

Ficus, Gymnanthes lucida Sw., Hibiscus elatus Sw.,

Mimosa, Plumeria L., and Trichilia L. (J.-P. Rudloff

pers. comm. 2008). If the larva is frugivorous on a host

common to these areas, Coccoloba, Ficus, or the

legumes are candidates.

The vicariant biogeographic relationship between

Suinoorda and the rest of the sepiform clade, which is

distributed from tropical West Africa to Fiji, parallels

other relationships in the cladogram (Fig. 7). Cliniodes

(node 53) is derived with respect to Asian and African

genera. Dicepolia (node 67), with both Neotropical and

Malagasy species, is related to genera and species

distributed around the Indian Ocean. The placement of

Malagasy D. munroealis is a sampling artifact; additional

evidence (Hayden 2009) indicates that the Malagasy

and Neotropical members constitute two monophyletic

clades. Argyrarcha margarita, related to the strictly

Neotropical Mecynarcha and Sobanga, is recorded from

Brazil and Madagascar (Munroe 1974a), and Aeglotis is

Central Asian. Vicariant relationships between

Neotropical and Palaeotropical taxa occur in many

Lepidoptera (Holloway & Nielsen 1999), although such

relationships involving Antillean endemics are better

known for other insect orders (Liebherr 1988).

The discovery of congeneric species in the Greater

Antilles is probable. Evidence from paleogeography

and lepidopteran biogeography suggests that Hispaniola

and Puerto Rico are most closely related to the eastern

Cuban landmass (Iturralde-Vinent & MacPhee 1999;

Liebherr 1988; Fontenla 2003), and the continued

discovery of even conspicuous moth taxa in Hispaniola

indicates that the fauna remains poorly known (Rawlins

& Miller 2008).

The phylogeny includes about half of the genera in

the Eurrhypini (Nuss et al. 2008), and it will be tested

by an analysis with more taxa and characters (Hayden in

prep.). It would be premature to confer a formal name

and rank on the sepiform clade, as such an act would

relegate the rest of the Eurrhypini to a paraphyletic

nominotypical taxon.

There is terminological uncertainty about the

squamiform structures. Nuss & Kallies (2001)

distinguished two different structures: a plume of long

scales in medial position on the vinculum (not shown

here) and a pair of “rifﬂed membranes” lateral of the

plume. They interpreted the scale plume to be the

“structures squamiformes” (Minet 1980; Leraut &

Luquet 1983), and they considered the membranes to

be a third eurrhypine synapomorphy. The “squamiform

structures” of Minet (1980: Figs. 5, 9) probably instead

refer to the membranes (M. Nuss pers. comm. 2009).

The illustrations of Leraut & Luquet could refer to

either structure (1983: Figs. 13, 19), and Munroe

neither illustrated nor elaborated on his reference to

“specialized scalelike sclerotizations associated with the

juxta and vinculum midventrally” (1972: 137).

Regardless of terminology, the long plume and the large

membranes frequently occur together, although the

plume was not observed in dissections of Suinoorda.

The plume is deciduous, whereas the membranes are

less easily removed. One hypothesis is that the

membranes and plume are developmental homologs:

the vinculum fundamentally bears scales in a transverse

row, of which two scales are hypertrophic. The rifﬂes or

striations are the longitudinal ridges, which bend

laterally so that the lamelliform structures rub across the

ridges.

Autocharis was reinstated by Amsel (1970) and

OLUME

63, N

UMBER

199

received numerous species removed from Noorda

Walker (Viette 1990; Shaffer & Munroe 2007). The

results indicate that the small, white-and-violet species

and the gray species form one monophyletic group. In

addition to the genitalia and apomorphies of node 50,

Autocharis species can be distinguished from

Pseudonoorda by having long male antennal sensilla and

plesiomorphic forewing venation. D. munroealis and

related species will have been transferred in another

publication (Hayden 2009).

Suinoorda maccabei requires a new genus because it

cannot be accommodated by any available genus

without substantial recircumscription. The next best

alternative would be to transfer the whole sepiform

clade to the oldest available name, Clupeosoma Snellen,

1880. Such an act would stabilize nomenclature by

mitigating the proliferation of genera with one or few

species, which is a problem in the Odontiinae (Nuss et

al. 2008). However, synonymy would disrupt several

established combinations, including the pest species in

Deanolis Snellen (Waterhouse 1998).

An important consideration for generic

circumscription is the generic diagnosis or deﬁnition,

which affects the management of specimens identiﬁed

only to that rank. Historically, most of the genera

included in this study were created to accommodate

species that did not ﬁt any pre-existing generic

diagnosis. The problem lies not with the general

practice of ﬁtting species into generic deﬁnitions but

rather with two speciﬁc issues. First, most of the

deﬁnitions predate Munroe’s circumscription of the

Odontiinae (1961), so genera were differentiated from

distantly related taxa by means of inappropriate

characters. Second, the diagnoses are typological,

therefore untestable and arbitrarily exact—that is, new

monotypic genera were deﬁned with any number of

specious characters (e.g. Munroe 1974a). Cladistic

parsimony equates diagnoses with synapomorphies,

making diagnoses both testable and efﬁcient (Farris

1979). Past nomenclatural debates (e.g. Ehrlich &

Murphy 1982 et seqq.) could have beneﬁted from

consideration of diagnoses. In the case of Clupeosoma,

none of the synapomorphies of node 64 are found in

previous deﬁnitions (Snellen 1880; Hampson 1897).

The characters that ﬁgure in Snellen’s original diagnosis,

revived by Munroe (1974b), are either invariant in the

Eurrhypini or dispersed across the cladogram (chars. 2,

7, 10, 11, 12). Of these, the pearly, metallic scales on

the body and wings (7:2) are shared among the

Malagasy and New Guinean species. With the transfer

of the Malagasy species, Clupeosoma is delimited at

node 60 with the following diagnosis: the forewing

medial area is concolorous with postmedial/terminal

area (4:2, except C. orientalale, which has an irregularly

darker medial area), the underside of the body and the

wings have pearly, metallic scales (7:2), and the costa of

the genitalic valva is apically produced in a straight point

(32:1).

One might criticize the proposal of a genus for a

single species. As suggested above, congeners probably

exist in related areas of endemism. More saliently,

Suinoorda is the ﬁrst odontiine genus proposed on

explicit phylogenetic grounds, and other monotypic

genera (Basonga,Metrea) are synonymized on the same

grounds. Suinoorda should accommodate any species

that share its derived characters: the sexually dimorphic

wing pattern (4:1,2), the absence of the forewing

antemedial line (8:0), and the gnathos arms near the

base of the uncus (36:1).

Pseudonoorda is recovered as paraphyletic, but

nomenclatural changes would require study of more

species, including the type species P. minor Munroe.

Munroe (1974b) recognized these two groups of

Pseudonoorda: one centering on P. distigmalis

(Hampson) and P. nigropunctalis (Hampson), and

another on P. brunneiﬂava. The latter group includes at

least P. brunneiﬂava, P. metalloma (Lower), P. photina

(Tams) and an undescribed species mentioned by

Munroe (1974b). The male of the coded terminal may

be Munroe’s undescribed species. Deanolis sublimbalis

Snellen, the red-banded mango borer of Southeast Asia

and New Guinea (confused in Munroe 1974b with the

spilomeline Decelia terrosalis Snellen), is closely related

to Pseudonoorda species of the ﬁrst group.

Cliniodes is restricted to South America and the West

Indies, except the eastern Nearctic C. ostreonalis (see

Hayden 2008). Cliniodes is diagnosed here by three

apomorphies: loss of the androconium from the base of

the valval costa (28:0), proximity of the gnathos arms to

the uncus (36:1), and a cervix bursae with pleated

sclerotization (46:1). Furthermore, C. ostreonalis (=

Metrea) and C. paradisalis (= Basonga) share with

derived Cliniodes species the loss of yellow forewing

scales and a juxta with a recurved, apical hook. Male

genitalia (not ﬁgured) indicate that C. ostreonalis is

closely related to the Andean C. glaucescens (Hampson)

and two other species, whereas C. paradisalis is related

to C. opalalis Guenée and C. euphrosinalis Möschler.

Epipagis ocellata and Mecyna catalalis are misplaced

in their respective genera, which both belong in the

Spilomelinae. Described in Sameodes Snellen, E.

ocellata is related to the other African species Epipagis

ﬂavispila (Hampson), Hapalia iospora (Meyrick), both

of which are here transferred to Eurrhypini, and

possibly some of the species placed in Epascestria by

Maes (2002). Viette (1990: 90) published the

200

200 J

OURNAL OF THE

EPIDOPTERISTS

’ S

OCIETY

combination “A[utocharis] catalalis (Viette), 1953,” but

it is not explicit whether the transfer was for Mecyna

catalalis Viette, 1953a: 136 or for Dichocrocis catalalis

Viette, 1953b: 208. Regardless, the species would be

misplaced in Autocharis, so it will be transferred in a

future work.

The speciﬁc epithets combined with Tegostoma and

Clupeosoma terminate with -ale (ICZN 1999: Artt. 31.2,

34.2) because these generic epithets are unambiguously

neuter. The original spellings are with -alis, as was

Zeller’s combination T. comparalis (1847: 581). The

Greek neuter ending -ma, -mat- is a common sufﬁx, as

in somatic, stomatic. Indeed, the meanings of

“Tegostoma” and “Clupeosoma” evidently refer to

character states 3:1 and 7:2, although that fact does not

rationalize the present choice of diagnosis. The

problems associated with gender agreement (Sommerer

2002) are acknowledged but beyond the scope of this

paper.

Predicting missing data is a strong empirical rationale

for phylogenetic classiﬁcation. Given some characters,

one can infer the states of other characters to an

arbitrary degree of precision. Subsequent observations

conﬁrm predictions, thereby expediting ﬁeld- and

labwork. For uunscored cells in characters with n

states, there are nupossible permutations, requiring

zero to many extra steps. These predictions follow

explicit rules (Fitch 1971) and are replicable.

The results indicate that Suinoorda larvae feed on

seeds or fruits. One extra step is required if the larvae

feed on something else, but without more information

about ecology and biology, there is no indication what

else that would be. It is possible that the feeding habit

is a ﬁfth unobserved state, such as specialization on

another tissue or family, though allowing for an

unknown state requires process assumptions about

character evolution or evolutionary rates.

Most of the predictions herein are precise (Table 3).

The least certain area involves the eight species,

unscored for feeding habit, above node 42 and below

node 54. Those above node 54 are all known or

predicted to be external folivores on Thymelaeaceae,

which their large body size and aposematism

corroborate. Taken individually, the other eight species

could assume any of the three habits other than stem-

boring/leaf-mining. However, only 17 of the 38possible

permutations add no extra steps. For example, Mecyna

catalalis would feed on Thymelaeaceae only if (1) all the

others do the same or if (2) P. hermesalis, V. bethalis

(Viette), and E. ocellata do the same and all the others

are seed/fruit feeders. The same situation obtains for E.

ocellata with regard to generalist folivory (state 3).

Although there is one tree with substantial support,

ambiguous character mapping clearly complicates

phylogenetic predictivity.

Fieldwork will be necessary to observe the feeding

habits of Suinoorda and any broader distribution or

undiscovered congeners; the conclusions above are

intended to facilitate that research. The phylogenetic

results and generic diagnoses will likewise require

testing with more evidence. This analysis demonstrates

that the new species is more closely related to

Palaeotropical genera than to the known Neotropical

ones. The Eurrhypini includes many other genera, and

the present sample focuses on the sepiform clade and

the superﬁcially similar Neotropical species.

Elucidating the relationships of the other major groups

will depend on description of novel characters and

reexamination of traditional ones.

ACKNOWLEDGEMENTS

I thank James K. Liebherr for his support, discussions about

methodology and West Indian biogeography, and critical com-

ments. I give thanks to Bernard Landry, Patrice Leraut, Joel

Minet, Michael Shaffer, and especially Alma Solis for advice

about pyraloid systematics and Odontiinae in particular. Dis-

cussions with M. Nuss about odontiine systematics, methodol-

ogy, the exigencies of cataloging, and the proliferation of generic

names in the subfamily led me to pursue the full cladistic analy-

sis; his comments on an earlier draft greatly improved this paper.

I kindly thank Tim McCabe and Jan-Peter Rudloff collecting

and lending the specimens and for accounts of the habitats. I

am also grateful to many people for making additional material

available: J. Barbut, U. Buchsbaum, P. Clavijo, A. Herrera, D.

Lafontaine, W. Mey, James Miller, E. van Nieukerken, J. and W.

de Prins, J. Rawlins, K. Tuck, and J. Weintraub. I thank K.

Nixon, M. Luckow, and the Bailey Hortorium systematics group

for comments on the drafts and stimulating discussions. The

Willi Hennig Society generously distributed the software gratis.

This work was supported in part by NSF Doctoral Dissertation

Improvement Grant DEB-0808415 to J. Liebherr and by the

Arthur Rawlins Endowment.

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Please see Appendices on next 4 pages

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Appendix A: Characters.

Head

0. Antennal sensilla: sensilla short in both sexes = 0; male antenna thicker = 1; male sensilla longer = 2; sensilla long

in both sexes = 3.

1. Labial palpi, aspect: porrect = 0; upturned = 1. Difﬁcult to discern for species with very short palpi, such as C.

ostreonalis.

2. Labial palpi, length: shorter than or equal to head = 0; longer than length of head = 1.

3. Frontoclypeal margin: straight = 0; prong = 1; ^-shaped arch = 2 (ﬁgs 5, 6: Fc). Despite its exaggeration, the

nearly circular structure in Suinoorda is homologized with the sharply angled shape in other taxa. At the other

extreme, some species have an slightly bowed arch. The double prong, two horizontal projections above the

haustellum shared by Eurrhypis and Tegostoma, is common in many Odontiini.

Wings

4. Forewing medial area, color: terminal and medial areas translucent or paler than postmedial line = 0; terminal

area dark, medial area pearly white or yellow = 1 (ﬁgs 1, 3, 4); terminal and medial areas dark = 2 (ﬁg. 2); terminal

area lighter than medial area = 3. In practice, states 0 and 2 may be hard to discern, e.g. comparing Dicepolia to

Hemiscopis and Hydrorybina. In state 0, the red or violet terminal forewing band is fundamentally absent, and any

coloration affects the whole forewing de novo. In state 2, the terminal band is fundamentally present, and the medial

area is suffused with the same coloration. State 1 broadly characterizes the sharply different coloration between

medial and postmedial areas, seen in the white/red pattern of Suinoorda, Autocharis, etc., the Schinia-like pattern of

Mimoschinia, and the modiﬁed aposematic pattern of Heortia.

5. Forewing medial area suffusion: absent = 0; with yellow or orange scales = 1. The yellow scales may be diffusely

dispersed among whiter scales (E. ocellata), or may occupy the entire wing. In some cases (Heortia), the pattern is

implicated in aposematic mimicry.

6. Extension of forewing costa: absent = 0; costa approximating or fused to postmedial line through discal spot = 1.

Applicable only for char. 4 state 1.

7. Sheen of wings and body scales: matte = 0; opalescent, silky, or hyaline = 1; metallic = 2. The difference between

states 0 and 1 can best be distinguished on the hindwings. State 2 is restricted to the legs, lower thorax and underside

of the wings in C. orientalale and C. vohilavale, whereas in other species C. atristriatum and C. sericiale, it also occurs

on the dorsal side of the wings.

8. Antemedial line: absent or reduced = 0; present = 1.

9. Proximity of forewing postmedial line to distal wing margin: narrow (PM line near distal margin) = 0; broad (PM

line ca. 3/5 from base of wing) = 1.

10. Forewing fovea: absent = 0; present = 1.

11. Forewing Rs1: not stalked with Rs2+3 = 0 (ﬁg. 10); stalked with Rs2+3 = 1.

12. Hindwing M2, M3: arising separately from cell = 0; stalked = 1.

13. Color in terminal hindwing anal area: not suffused = 0 (ﬁgs 1–4); suffused = 1. Inapplicable where the

hindwing lacks any trace of color. The absence or strong reduction of coloration on the anal area, at least indicated

by the postmedial line’s abrupt cessation or convergence with the margin at A1, is also common in Glaphyriinae,

Evergestinae, Pyraustinae s. str. and a few Spilomelinae.

14. Hindwing anal margin, emargination: absent = 0; present = 1. The round notch in the basal half of the

hindwing anal area appears to articulate with the metathoracic legs.

Tympanal organs

15. Saccus tympani, medial rounding: rounded and deep: = 0 (ﬁg. 17: ST); medially oblong: = 1. The pair of sacci

extend anteriad under S2. The outline of state 0 is like a D, and state 1 is more like a “|)” shape.

Male genitalia

16. A8 tergite posterior fringe: not expanded = 0; expanded, square, like whole tergite = 1. All Odontiinae have a

distinct fringe of hairs on the posterior edge of T8. The fringe is normally a narrow strip (state 0; ﬁg. 12: F); state 1

describes cases where the scale-bearing ﬁeld is expanded, up to a length:width ratio of 1:1. Syntonarcha also has an

expanded fringe (Gwynne & Edwards 1986, ﬁg. 2: “cuticular ﬂap”).

17. S8 bilobate: straight or monolobate = 0; weakly bilobate (lobe depth less than width) = 1 (ﬁg. 12); strongly

bilobate (lobe depth exceeds width) = 2. The anterior, internally extended lobes of sternite 8 evidently serve as

muscle insertion sites for the retraction of the genitalia. In state 2, the medial concavity is deeper than the width of

either lobe.

18. S8 posterior median projection: absent = 0 (ﬁg. 12); present = 1. This is in the same plane as the rest of the

sclerite (cf. characters 19, 20).

19. S8 posterior median projection, shape: simple, triangular = 0; bilobate or trapezoidal = 1. Inapplicable if char.

18 absent.

20. S8 posterior edge: straight = 0; excavate or concave = 1 (ﬁg. 12); boss or saddle-horn = 2 (ﬁg. 9). In state 2, the

medial posterior margin emerges out of the plane of the rest of S8.

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APPENDIX A (continued)

21. Lamelliform structures: absent = 0 (ﬁg. 12); present = 1 (ﬁg. 9: Lm).

22. Sepiform structures: absent = 0 (ﬁg. 9); present = 1 (ﬁg. 8: Sep).

23. Piluli of S8 (posterolateral of lamelliform structures): absent = 0; present = 1 (ﬁg. 12: Pi). These sensillae occur

posterolaterally of lamelliform structures, closer to the posterior corners of S8. See also Nuss & Kallies 2001: ﬁg. 2.

24. Squamiform structures: absent = 0; present = 1 (ﬁgs 8–10). These are the paired “rifﬂed membranes” of Nuss

& Kallies (2001: ﬁg. 1).

25. Squamal symmetry: symmetrical = 0 (ﬁgs 8–10); asymmetrical = 1. See Gwynne & Edwards 1986 for an

asymmetrical example not included in this study.

26. Squamiform structures, distal edge: squarish = 0 (ﬁg. 9); rounded = 1 (ﬁgs 8, 10).

27. Squamal enations: absent = 0; present = 1 (ﬁg. 8: En). These fold around the sepiform structures where the

latter are present, but the presence of a similar mid-squamal protuberance in Hemiscopis warrants coding them as a

separate character.

28. Basicostal androconia: absent = 0; medium-length to long scales from ﬁeld at base of costa = 1 (ﬁgs 18 & 19:

BA); long scales from discrete ﬁeld at end of sclerotized costa = 2.

29. Main patch of nondeciduous setae: not differentiated = 0; massive, robust setae = 1. Restricted to Heortia and

Cliniodes. In C. paradisalis, this is represented by a single, hook-like, basally directed spine near the apex of the

valve. Like other sensilla chaetica, it does not absorb Chlorazol stain.

30. Scoop-shaped scales: absent = 0; present = 1 (ﬁg. 18: ScS). These broad scales have ﬁne, closely spaced ridges,

pearly lustre, and are shaped like potato chips. They commonly arise from the dorsal side of the valve and curve over

the edge. They are often apparent by their lustre and dense packing on the valval edges of partly exposed genitalia.

31. Vincular androconia: absent = 0; present = 1 (ﬁg. 18: VA). Common to most Pyraloidea, this is the pair of

androconia on the pleural portion of the A8-A9 intersegmental membrane. Often called “coremata” (e.g. Sutrisno

2002, char. 30; Landry 1995, “coremata on intersegmental membrane VIII-IX”).

32. Apex of costa: not projecting = 0; straight hook = 1; blunt sigmoid (apex out-turned) = 2 (ﬁg. 19); downcurved

hook fused to pleated ﬂap = 3.

33. Juxta apex: not differentiated = 0; recurved hook = 1.

34. Gnathos apex: blunt and linguiform = 0; sharp, acute = 1; hatchet-shaped = 2.

35. Gnathos sides: inverse V = 0; inverse Y, with medial process = 1 (ﬁg. 11); compact base with ventral notch = 2;

transverse bar = 3. Some gnathoi coded as state 0, including S. maccabei, have bent lateral arms and approximate

state 1, but the bends lie well below the medial junction, more like a capital upsilon.

36. Gnathos base-uncus base: gnathos base not articulating with uncus lower corners = 0; gnathos base near or

articulating with uncus lower corners = 1 (ﬁg. 11).

37. Phallus: not spiral = 0; spiral = 1.

Female genitalia

38. Ovipositor length: short = 0 (ﬁg. 16); long = 1.

39. Colliculum: short = 1 (ﬁg. 16: C); long (most of length of ductus) = 2.

40. Ductus bursae sclerotization: absent = 0; smooth, extending along the long, narrow ductus bursae = 1; rough,

on limited area of cervix bursae = 2. In many Eurrhypini, including Suinoorda, the ductus bursae is expanded as a

cervix bursae (ﬁg. 16: CvB). In this sample of taxa, no clear distinction could be made between the typically long,

narrow ductus and a short, inﬂated cervix bursae. Species with state 1 have the ductus bursae mostly but weakly

sclerotized, whereas state 2 represents a limited area that is often granular. Suinoorda and Noordodes share a large

cervix bursae occupied by massive, irregular sclerotization.

41. Appendix of ductus bursae: absent = 0; present = 1.

42. Corpus bursae signum: absent = 0; present (1 or 2) = 1 (ﬁg. 16).

43. Corpus bursae signum shape: round = 0 (ﬁg. 16); linear = 1. Inapplicable if char. 42 absent.

44. Ductus seminalis origin: from ductus bursae = 0 (ﬁg. 16: DS); from corpus bursae, with signum extension = 1.

45. Ductus seminalis basal width: narrow = 0; enlarged = 1.

46. Pleats or wrinkles on cervix bursae: absent = 0; present = 1; present over both cervix and corpus bursae = 2.

State 2 represents cases where there is no clear distinction between the plications of the cervix and corpus bursae. In

Suinoorda, pleats occur on the corpus, but they are clearly not continuous from the irregularly shaped sclerotization

of the cervix.

Larva

47. Feeding habit: tube-dwelling stem borer or leaf miner = 0; seed and fruit pulp feeder = 1; folivore,

Thymelaeaceae = 2; folivore, other = 3. The states reﬂect plausible physiological and metabolic constraints. The

external silken tube of Eurrhypis and Tegostoma is probably a modiﬁcation of an ancestrally internal gallery, retained

for feeding on thin-branched hostplants. Thymelaeaceae produce many unique feeding deterrents (Maistrello et al.

2005), and folivory on this family is rare in Lepidoptera (Robinson et al. 2008). How correctly the states are deﬁned

will emerge as new records are obtained (see the predicted values).

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PPENDIX

B:Data matrix. ?: unobserved, -: inapplicable. More than one state: A: [01], B: [02], C: [12], D: [23]

Terminal Character Number

0 5 10 15 20 25 30 35 40 45

| | | | | | | | | |

Cynaeda dentalis----------------

200000-0-0100000000-00000---00010010000100110000

Tegostoma comparale-------------

200120-010000100000-00000---000100100011000-0000

Aeglotis argentalis-------------

D010101110000001010-110010000010001300??????????

Argyrarcha margarita------------

0000101110000001100-210011000010001210010010001?

Autocharis albiplaga------------

2012100110000000020-010110001011202000012011000?

Autocharis barbieri-------------

2012A00110000000021111011000101?2011001200110003

Autocharis fessalis-------------

2012100100001000020-1101100010113010001220110003

Autocharis mimetica-------------

2012100100000000020-01011000101120200012200-000?

Cliniodes costimacula-----------

000021-110000101010-210110000011000010012010101?

Cliniodes opalalis--------------

310020-110000100010-2101100021110112100220101012

Cliniodes ostreonalis-----------

1A0000-110000-01010-210110000011010310012010101D

Cliniodes paradisalis-----------

1100101110000100010-C1011000211001101001200-101?

Clupeosoma atristriatum---------

001020-2001110001110101010110011101000??????????

Clupeosoma orientalale----------

000030-200010000011010101011001110100011001001A?

Clupeosoma sericiale------------

000020-2001110001110101010110011101000110010011?

Clupeosoma vohilavale-----------

10A020-210010000011010101011001110000011001001A?

Deanolis sublimbalis------------

001010010001001001111010101100110020001100100101

Dicepolia munroealis------------

?01200-110000001011011011000101?30110101010-00A?

Dicepolia roseobrunne-----------

201200-11000000002100101100010113011010101100001

Dicepolia rufitinctalis---------

001200-1100010000210110110001011301101010010000?

Ephelis cruentalis--------------

2000100010000000010-01011000000100100001000-000?

Epipagis ocellata---------------

1000111110000001010-210110001011001000110110100?

Eurrhypis pollinalis------------

2001111010000100010-01011000000100100001000-0010

Hemiscopis suffusalis-----------

001020-1110001010110210110011011000000010010000D

Heortia dominalis---------------

1010111110000101100-110110001111000000012010000?

Heortia vitessoides-------------

1100111111000101110-1101100011110000000120100022

Hyalinarcha hyalinalis----------

200200-110000000010-210010001010301100010011001?

Hydrorybina polusalis-----------

001020-111010001021011011000101100000011000-0013

Mecyna catalalis----------------

000021-1100000010110110110000011001A0001000-000?

Mecynarcha apicalis-------------

000020-110000101100-110011000000001210010010001?

Mimoschinia rufofascialis-------

2000100010000000010-11011000000100100011100-0021

Noordodes magnificalis----------

0002100110010000010-101010110011001000112010001?

Pitama hermesalis---------------

1000100101000001020-010110001011001100010010000?

Pseudonoorda brunneiflava-------

0002A00100010000010-001010010011001000A12000011?

Pseudonoorda distigmalis--------

00101001000100100111101010110011002000110010010?

Pseudoschinia elautalis---------

2000100010000000010-11011000000100100001100-0001

Sobanga rutilalis---------------

0000101110000001100-00000---000?001210010010001?

Suinoorda maccabei--------------

0002C00100000000010-10111011101100101001201000A?

Viettessa bethalis-------------

1000101101000001020-B1011000101100100011000-000?

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Cynaeda dentalis :

No autapomorphies

Tegostoma comparale :

3: 0 --> 1

13: 0 --> 1

38: 0 --> 1

Aeglotis argentalis :

0: 0 --> 23

2: 0 --> 1

Argyrarcha margarita :

20: 1 --> 2

Autocharis albiplaga :

38: 1 --> 0

39: 2 --> 1

Autocharis barbieri :

No autapomorphies

Autocharis fessalis :

12: 0 --> 1

Autocharis mimetica :

42: 1 --> 0

Cliniodes costimacula :

0: 1 --> 0

Cliniodes opalalis :

0: 1 --> 3

35: 0 --> 2

39: 1 --> 2

Cliniodes ostreonalis :

4: 12 --> 0

35: 0 --> 3

Cliniodes paradisalis :

31: 1 --> 0

42: 1 --> 0

Clupeosoma atristriatum :

2: 0 --> 1

Clupeosoma orientalale :

4: 2 --> 3

Clupeosoma sericiale :

No autapomorphies

Clupeosoma vohilavale :

0: 0 --> 1

8: 0 --> 1

34: 1 --> 0

Deanolis sublimbalis :

No autapomorphies

Dicepolia munroealis :

15: 0 --> 1

17: 2 --> 1

42: 1 --> 0

Dicepolia roseobrunnea :

20: 1 --> 0

Dicepolia ruﬁtinctalis :

0: 2 --> 0

12: 0 --> 1

Ephelis cruentalis :

No autapomorphies

Epipagis ocellata :

38: 0 --> 1

41: 0 --> 1

Eurrhypis pollinalis :

3: 0 --> 1

5: 0 --> 1

6: 0 --> 1

13: 0 --> 1

46: 0 --> 1

Hemiscopis suffusalis :

13: 0 --> 1

20: 1 --> 2

27: 0 --> 1

Heortia dominalis :

2: 0 --> 1

17: 1 --> 0

Heortia vitessoides :

1: 0 --> 1

9: 0 --> 1

46: 0 --> 2

Hyalinarcha hyalinalis :

20: 1 --> 2

23: 1 --> 0

31: 1 --> 0

46: 0 --> 1

Hydrorybina polusalis :

11: 0 --> 1

17: 1 --> 2

38: 0 --> 1

46: 0 --> 1

Mecyna catalalis :

5: 0 --> 1

Mecynarcha apicalis :

4: 1 --> 2

13: 0 --> 1

Mimoschinia

rufofascialis :

38: 0 --> 1

46: 0 --> 2

Noordodes magniﬁcalis :

No autapomorphies

Pitama hermesalis :

6: 1 --> 0

35: 0 --> 1

Pseudonoorda

brunneiﬂava :

20: 1 --> 0

26: 1 --> 0

42: 1 --> 0

Pseudonoorda

distigmalis :

No autapomorphies

Pseudoschinia elautalis :

No autapomorphies

Sobanga rutilalis :

20: 1 --> 0

21: 1 --> 0

24: 1 --> 0

Suinoorda maccabei :

36: 0 --> 1

Viettessa bethalis :

38: 0 --> 1

42: 1 --> 0

Node 41 :

23: 1 --> 0

31: 1 --> 0

Node 42 :

6: 0 --> 1

15: 0 --> 1

Node 43 :

0: 2 --> 0

43: 1 --> 0

Node 44 :

7: 0 --> 1

30: 0 --> 1

42: 0 --> 1

Node 45 :

20: 0 --> 1

47: 0 --> 1

Node 46 :

17: 0 --> 1

21: 0 --> 1

23: 0 --> 1

24: 0 --> 1

Node 47 :

16: 0 --> 1

17: 1 --> 0

25: 0 --> 1

36: 0 --> 1

Node 48 :

20: 1 --> 0

34: 1 --> 2

Node 49 :

35: 1 --> 0

40: 0 --> 2

Node 50 :

38: 0 --> 1

39: 1 --> 2

47: 1 --> 3

Node 51 :

2: 0 --> 1

17: 1 --> 2

Node 52 :

32: 0 --> 3

35: 0 --> 1

Node 53 :

28: 1 --> 0

36: 0 --> 1

46: 0 --> 1

Node 54 :

13: 0 --> 1

34: 1 --> 0

40: 0 --> 2

Node 55 :

5: 0 --> 1

Node 56 :

0: 0 --> 1

Node 57 :

15: 1 --> 0

28: 0 --> 2

29: 0 --> 1

34: 0 --> 1

Node 58 :

5: 1 --> 0

33: 0 --> 1

Node 59 :

10: 0 --> 1

12: 0 --> 1

16: 0 --> 1

Node 60 :

4: 1 --> 2

7: 1 --> 2

32: 0 --> 1

Node 61 :

3: 2 --> 0

18: 0 --> 1

40: 2 --> 0

Node 62 :

45: 0 --> 1

Node 63 :

11: 0 --> 1

23: 1 --> 0

38: 0 --> 1

Node 64 :

21: 1 --> 0

22: 0 --> 1

26: 0 --> 1

27: 0 --> 1

40: 0 --> 2

Node 65 :

2: 0 --> 1

14: 0 --> 1

19: 0 --> 1

34: 1 --> 2

46: 1 --> 0

Node 66 :

41: 0 --> 1

Node 67 :

37: 0 --> 1

43: 1 --> 0

Node 68 :

2: 0 --> 1

9: 0 --> 1

34: 1 --> 0

Node 69 :

4: 1 --> 2

18: 0 --> 1

Node 70 :

16: 0 --> 1

29: 0 --> 1

Node 71 :

30: 1 --> 0

Node 72 :

40: 0 --> 1

Node 73 :

8: 1 --> 0

9: 0 --> 1

17: 1 --> 2

APPENDIX C: Unambiguous character state changes of nodes for cladogram (ﬁg. 7). Format: “character: primitive state --> de-

rived state.” Obtained with TNT commands “Optimize / Synapomorphies / List synapomorphies.”

Update on the status of Citheronia sepulcralis in the Bahamas (Saturniidae: Ceratocampinae)

Article

Full-text available

Jan 2018

A molecular phylogeny for the pyraloid moths (Lepidoptera: Pyraloidea)

Conference Paper

Full-text available

Nov 2012

We present the first detailed molecular estimate of relationships across the subfamilies of Pyraloidea, and assess its concordance with previous morphology-based hypotheses. Maximum likelihood analyses yield trees that differ little among data sets and character treatments and are strongly supported at all levels of divergence. Subfamily relationships within Pyralidae, all very strongly supported, differ only slightly from a previous morphological analysis, and can be summarized as Galleriinae + Chrysauginae (Phycitinae (Pyralinae + Epipaschiinae)). In Crambidae the molecular phylogeny is also strongly supported, but conflicts with most previous hypotheses. Among the newly-proposed groupings is a wet-habitat clade comprising Acentropinae + Schoenobiinae + Midilinae, and a provisional mustard oil clade containing Glaphyriinae, Evergestinae and Noordinae.

A molecular phylogeny for the pyraloid moths (Lepidoptera: Pyraloidea) and its implications for higher‐level classification

Article

Oct 2012

Pyraloidea, one of the largest superfamilies of Lepidoptera, comprise more than 15 684 described species worldwide, including important pests, biological control agents and experimental models. Understanding of pyraloid phylogeny, the basis for a predictive classification, is currently provisional. We present the most detailed molecular estimate of relationships to date across the subfamilies of Pyraloidea, and assess its concordance with previous morphology‐based hypotheses. We sequenced up to five nuclear genes, totalling 6633 bp, in each of 42 pyraloids spanning both families and 18 of the 21 subfamilies, plus up to 14 additional genes, for a total of 14 826 bp, in 21 of those pyraloids plus all 24 outgroups. Maximum likelihood analyses yield trees that, within Pyraloidea, differ little among datasets and character treatments and are strongly supported at all levels of divergence (83% of nodes with bootstrap ≥80%). Subfamily relationships within Pyralidae, all very strongly supported (>90% bootstrap), differ only slightly from a previous morphological analysis, and can be summarized as Galleriinae + Chrysauginae (Phycitinae (Pyralinae + Epipaschiinae)). The main remaining uncertainty involves Chrysauginae, of which the poorly studied Australian genera may constitute the basal elements of Galleriinae + Chrysauginae or even of Pyralidae. In Crambidae the molecular phylogeny is also strongly supported, but conflicts with most previous hypotheses. Among the newly proposed groupings are a ‘wet‐habitat clade’ comprising Acentropinae + Schoenobiinae + Midilinae, and a provisional ‘mustard oil clade’ containing Glaphyriinae, Evergestinae and Noordinae, in which the majority of described larvae feed on Brassicales. Within this clade a previous synonymy of Dichogaminae with the Glaphyriinae is supported. Evergestinae syn. n. and Noordinae syn. n. are here newly synonymized with Glaphyriinae, which appear to be paraphyletic with respect to both. Pyraustinae and Spilomelinae as sampled here are each monophyletic but form a sister group pair. Wurthiinae n. syn., comprising the single genus Niphopyralis Hampson, which lives in ant nests, are closely related to, apparently subordinate within, and here newly synonymized with, Spilomelinae syn. n.

TAXONOMIC STUDY OF FAMILY PYRALIDAE(LEPIDOPTERA) FROM FAISALABAD PUNJAB PAKISTAN

Thesis

Full-text available

Oct 2020

Family Pyralidae includes Pyralid moths, snout moths or grass moths.These belong to order Lepidoptera. The members of this family are polyphagous in nature. The labial palpi of snout moth projected forward. Pyralid consider as one of the diverse family of insect pest which cause huge losses to important Agricultural crops worldwide. The snout moths are important and establish worldwide in tropical and subtropical ecosystems. It is adiverse group with 5000 species belong to five major subfamilies namely Pyralinae, Phycitinae, Epipaschiinae, Galleriinaeand Chrysauginae worldwide. The 30 species of 25 genera of family Pyralidae are collected from Pakistan. In the present study snout moths were collected from different locations of Faisalabad from infected weeds, trees, plants and shrubs etc. Collection of snout moth will be done with the help of Ariel net and other suitable equipment’s. The specimenswerecollected and preserved in wooden boxes by using suitable pins. Later, these were shifted in insectarium, Department of Entomology University of Agriculture Faisalabad. Identification of specimens were done by using different taxonomic keys and available relevant literature.

First report of the genus Neadeloides Klima, 1939 (Lepidoptera, Crambidae, Pyraustinae) in Laos, with description of a new species

Article

Full-text available

Aug 2020

The genus Neadeloides Klima, 1939 is a small genus of the subfamily Pyraustinae (Lepidoptera, Crambidae). It was established for the type species Hoterodes cinerealis Moore, 1867 as the objective replacement name for Adeloides Warren, 1892, a homonym of Adeloides Blanchard, 1845 of Curculionidae in Coleoptera (Singh et al. 2019; Nuss et al. 2003–2020). Recently, this genus was studied by Singh et al. (2019), with the inclusion of three species: N. cinerealis (Moore, 1867) and N. nubilus Singh, Ranjan & Singh, 2019 from India, and N. glaucoptera (Hampson, 1896) from Bhutan. Species of Neadeloides can be distinguished by the following characters. In the male, the antenna are very long, about twice the length of the forewing; forewing narrow, with a produced apex; termen extremely oblique; in the female, antenna almost same length of the forewing; forewing wider than male; in the male genitalia, uncus broad, inverted U–shaped, ventrally concave, with a small, sub–apical lateral projection on both sides; valva elongated flap–like, membranous at base and sclerotized from middle to apex; costal margin rather concave, nearly straight; dorsum of valva outwardly angled at middle or roughly straight; sacculus process hook–shaped or broad rectangular, with setose apical flap (Singh et al. 2019).

Borboletas e mariposas de Santa Catarina: uma introdução

Book

Full-text available

Jun 2016

O estado de Santa Catarina possui um conhecimento significativo acerca de sua biodiversidade, possuindo informações sobre suas aves, mamíferos, peixes, anfíbios, formigas, e flora em geral, por meio dos trabalhos da Flora Ilustrada Catarinense iniciados por Raulino Reitz. Com o objetivo de contribuir para este conhecimento, o presente trabalho teve como base a lista das espécies de Lepidoptera (borboletas e mariposas) registradas em Santa Catarina e compiladas por Piovesan e colaboradores, com acréscimo de novos registros pessoais e informações de diversos livros e artigos adicionais, resultando no registro de 1637 espécies de borboletas e mariposas. Neste trabalho são apresentadas informações sobre origem, evolução e biologia em geral da ordem Lepidoptera (borboletas e mariposas) e informações sobre 37 famílias desta ordem que ocorrem em Santa Catarina e mais de 300 fotos.

Bioecology of Heortia vitessoides Moore (Lepidoptera: Pyralidae: Odentiinae), a major defoliator of Aquilaria malaccensis Lam. (Indian eagle wood)

Article

Full-text available

Jan 2000

Surveys conducted in Jorhat, and Golaghat Districts of Assam revealed that Heortia vitessoides Moore was a major insect pest of agar-oil yielding tree, Aquilaria malaccensis. Although the pest is active for eight months from early March to October but trees in the region were defoliated twice in a season. Forty to hundred per cent defoliation was observed during May/June in 53.0-77.6 % trees while during Aug.lSept 21.66-39.33% trees were affected. The intensity of attack was more in the trees grown in open than under shade. There were five to six generations in a year. First generation started with the emergence of adults from the hibernating pupae of previous year. The female laid 400-600 eggs in two to three masses after two to four days of mating. Life cycle from egg to adult was completed within 35.8 days. A polyhedrosis virus was found to infect this pest in the field.

Création d'une sous-famille particulière pour Noorda Walker, 1859 et définition d'un nouveau genre parmi les Odontiinae [Lep. Pyralidae]

Article

Jan 1980

Joël Minet

The Biogeography of the West Indian Butterflies (Lepidoptera): An Application of a Vicariance/Dispersalist Model

Chapter

Jun 2001

A phylogenetic analysis of the Australian Glyphodes Guen{u00E9}e and allied genera (Lepidoptera: Pyraloidea: Crambidae: Pyraustinae)

Thesis

Jan 1999

Hari Sutrisno

Nouvelles Pyrales de Madagascar [Lepidoptera]

Article

Jan 1978

Pierre Viette

New species of geometers and pyrales from the Khasia Hills

Article

C. Swinhoe

To agree or not to agree - The question of gender agreement in the International Code of Zoological Nomenclature

Article

Jan 2002

Manfred Sommerer

The new (4th) edition of the International Code of Zoological Nomenclature still requires in its Articles 31.2 and 34.2 that an adjectival species-group name be in agreement with the gender of the name of the genus it is at any time associated with. Prominent and influential publications on the taxonomy of Lepidoptera expressly chose to ignore the gender agreement requirements of the (corresponding previous) Code, and to use the specific name as given in the original description. For most lepidopterists of our time it is, by lack of knowledge in Latin and Greek, impossible to ascertain unambiguously the gender of the generic names in Lepidoptera. Since strict application of the gender agreement provisions of the Code in the nomenclature of Lepidoptera would, in the course of progress in systematics, require continuous changes of epithets of specific names, the task of updating the names in electronic databases of large lepidopteran groups is beyond the manpower and financial resources of museums and scientific institutions. It is therefore practically not possible to apply those rules of gender agreement. Regrettably, the International Commission on Zoological Nomenclature did not accept those arguments for the latest version of the Code. The author explains that in lepidopterology there has never been a tradition of 'classic purity' as advocated by the Code. Given the priority of the principles of stability and permanence of zoological names the author proposes that all lepidopterists follow the example of leading authors in taxonomy and disregard the gender agreement requirements of the Code. The leading lepidopterists' societies should encourage their members in this respect. The Societas Europaea Lepidopterologica (SEL) with about 600 members has, on 4 June, 2002, passed an appropriate resolution (which is reproduced in the Appendix).

Dioptine moths of the Caribbean region: Description of two new genera with notes on biology and biogeography (Lepidoptera : Notodontidae : Dioptinae)

Article

Feb 2008

Two new species of dioptine moths (Lepidoptera: Noctuoidea: Notodontidae: Dioptinae) are described from the Caribbean island of Hispaniola, and placed in two new monobasic genera. These are the first records of subfamily Dioptinae in the Caribbean Region. Eremonidia mirifica new genus, new species, is a plesiotypic member of the tribe Dioptini, and Caribojosia youngi new genus, new species, is placed in the tribe Josiini. Both species are described from cloud forest habitats in the Sierra de Neiba (Dominican Republic and adjacent Haiti); E. mirifica is also known from similar habitats in the Cordillera Central (Dominican Republic). The larvae of C. youngi are described feeding on mature leaves of Passiflora sexflora Jussieu (Violates: Passifloraceae). Habitats, relationships, ecological associations, behavior, and conservation issues for these species are discussed.

Titanio caradjae (Rebel, 1902) comb. n., transferred from Brachodidae (Sesioidea) to Crambidae (Pyraloidea)

Article

Jan 2001

Brachodes caradjae (Rebel, 1902) is transferred from Brachodidae (Sesioidea) to Titanio Hübner, [1825] (Pyraloidea: Crambidae: Odontiinae). The holotype as well as male and female genitalia are figured. In this context, we give short insights into the systematics of the Odontiinae and provide the hypothesis, that the diagnostic characters 'structurae squamiformes' and 'structurae lamelliformes' given by Leraut & Luquet (1982) for the Eurrhypini Leraut & Luquet, 1982 can be regarded as synapomorphies of this tribe. Further, we present a possible third synapomorphy for the Eurrhypini, a paired riffled membrane situated ambilateral to the 'structurae squamiformes' of the vinculum.

Biogeography of Antillean Butterflies (Lepidoptera: Rhopalocera): Patterns of Association among Areas of Endemism

Article

Dec 2003

Jorge Fontenla

An association analysis among areas of butterfly endemism in the West Indies was developed. The following analytical tools were used: Parsimony analysis of endemicity (PAE), Principal Components Analysis (PCA) and Panbiogeography. PAE and PCA's results yielded three main associations : Cuba-Bahamas (BAH)-Isle of Pines (PIN)-Cayman Islands (CAY), Hispaniola (HIS), Puerto Rico (PRI)-Jamaica (JAM)- Virgin Islands (VIR), and Leeward and Winward islands (LEE-WIN). Panbiogeography produced the following generalized tracks: Cuba-BAH, Cuba-PIN, Cuba-HIS, Cuba-CAY, Cuba-HIS-BAH, Cuba-HIS-PRI, HIS-PRI, HIS-JAM, Cuba-HIS-JAM-PRI and LEE-WIN. Cuba is the "mass center" of shared species and HIS the "mass center" of endemism. It was considered that both islands represent "biogeographic attractors" in the complex Antillean scenario. Panbiogeographic analysis produced results more in agreement with the real complexity and reticulation of spatial relationships among areas.

A new genus of Caribbean Odontiinae with Palaeotropical affinities (Lepidoptera: Crambidae)

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