Systematics of Otarrha, a new Neotropical subgenus of Chimarra (Trichoptera: Philopotamidae)

Systematic Entomology (2002) 27, 65±130

Systematics of Otarrha, a new Neotropical subgenus of Chimarra (Trichoptera: Philopotamidae) ROGER J. BLAHNIK Department of Entomology, University of Minnesota, St Paul, Minnesota, U.S.A.

Abstract. Subgenus Otarrha is established in genus Chimarra to include eighteen described species formerly placed either in subgenus Chimarra or unplaced to subgenus, and thirteen new species. All species are Neotropical, with collective distributions primarily in the Antilles (Greater and Lesser) and northern South America. One species occurs in southeastern Brazil and another species in Costa Rica and Panama. New species are described, recognized species redescribed (except for C. diannae and C. koki), and a key to the identi®cation of males in the subgenus is provided. Additionally, characters supporting monophyly of the subgenus and a phylogeny of its species are proposed. Described species transferred to this subgenus include Chimarra cubanorum Botosaneanu, C. diakis Flint, C. diannae Flint & Sykora, C. dominicana Flint, C. garciai Botosaneanu, C. guapa Botosaneanu, C. jamaicensis Flint, C. koki Botosaneanu, C. machaerophora Flint, C. patosa Ross, C. puertoricensis Flint, C. quadrifurcata Botosaneanu, C. retrorsa Flint, C. rossi Bueno-Soria, C. sensillata Flint, C. septemlobata Flint, C. septifera Flint and C. spinulifera Flint. Chimarra patosa is designated the type species of the subgenus. New species described in Otarrha include Chimarra amazonia (Peru), C. barinas (Venezuela), C. darlingtoni (Cuba), C. diaphora (Venezuela), C. incipiens (Venezuela), C. odonta (Brazil), C. parene (Peru), C. parilis (Peru), C. particeps (Peru), C. peruana (Peru), C. phthanorossi (Colombia), C. redonda (Dominican Republic) and C. tachuela (Venezuela). Two additional species are described and left incertae sedis to subgenus, Chimarra usitatissima Flint and C. angularis, sp.n. (Venezuela, Guyana). Introduction Chimarra Stephens is one of the largest genera of Trichoptera, and is nearly cosmopolitan in distribution. New World species of Chimarra have traditionally been placed in two subgenera, the nominotypical subgenus or subgenus Curgia. Neotropical members of subgenus Chimarra were recently revised (Blahnik, 1998), with the addition of many new species. These species were formally placed in a number of species groups, and relationships among species within these groups were proposed. Similarly, Curgia was recently revised by Flint (1998), who also recognized a number of distinct species groups for that subgenus and more than doubled the number of described species. Another subgenus, Chimarrita (Blahnik, 1997), was recently established for a distinctive group of Neotropical species formerly placed in subgenus Chimarra. The three Correspondence: Roger J. Blahnik, Department of Entomology, University of Minnesota, 219 Hodson Hall, 1980 Folwell Ave, St Paul, MN 55108, U.S.A. E-mail: [email protected] For this paper, pages in excess of ®fty were paid for by the author.

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papers cited above should be consulted for background information on the taxonomy of genus Chimarra. The species treated in this paper include the remainder of the known species of Neotropical Chimarra. Like members of subgenus Chimarrita, these species were formerly either placed in subgenus Chimarra, or their subgeneric placement was left undetermined. They are here placed in a new subgenus. The species included in this paper were previously informally referred to as the patosa group. Chimarra patosa was described by Ross (1956), who referred to that species and a number of similar South American forms (then undescribed), as being evolutionarily intermediate between Chimarra (Curgia) and Asiatic forms of Chimarra, typi®ed by the species Chimarra tsudai. The character used by Ross to indicate intermediacy was the retention in C. patosa of a narrow mesal lobe of tergum X separating two divided lateral lobes. However, Ross's argument that the mesal strap of Chimarra patosa represents a mesal lobe of tergum X may be in error because a similar structure in related species described in this paper could as easily be interpreted as an extension of tergum IX. Moreover, not all species in the patosa group 65

66 Roger J. Blahnik

Figs 1±3. 1, Chimarra machaerophora, head and thorax; 2, C. machaerophora, maxillary palp; 3, C. tachuela, wings (A) forewing (B) hindwing.

ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

Systematics of Otarrha possess a median lobe. Flint (1964) recognized the relationship of certain Antillean species to C. patosa, and listed some character similarities. These Antillean species lack a mesal process on tergum X, making Ross's evolutionary argument for the intermediacy of the patosa group between Chimarra (Chimarra) and Chimarra (Curgia) questionable: the character evidence is inconclusive at best. Additionally, the de®nition of the patosa group has been rendered somewhat vague. A formal description of characters diagnosing the patosa group, and designation of all species properly belonging to it, has never been made. Nevertheless, the group itself is well de®ned, distinctly different from subgenera Chimarra, Curgia and Chimarrita, and warrants its own subgeneric name. The relative relationships among these subgenera remain unresolved. However, recent molecular work (Blahnik, unpublished) suggests the likelihood that the subgenus recognized here, along with subgenera Curgia and Chimarrita, form a monophyletic, New World endemic group. Although it is probable that the species remaining in subgenus Chimarra, most of which occur in the Old World, form a monophyletic group, the group remains a diverse assemblage. It is not unlikely that additional subgenera will be recognized within this group as relationships of Old World Chimarra are better understood. Materials and methods Species determinations were made by examination of male genitalia. Male genitalia were cleared in cold 10±12% KOH solution overnight, and a syringe was used to empty the macerated contents from the abdomen. Female genitalia were prepared in the same way, but were additionally stained with Chlorazole Black dye before examination. The stain was prepared by dissolving several grains of the dye in glycerin. Genitalia remained in this solution for several minutes, and were then transferred to 80% ethanol to rinse off excess dye and stop the dyeing process. Genitalia were examined in small dishes of glycerin, and drawings made with use of an ocular grid. Terminology for male genitalia largely follows that of Schmid (1980, 1982), except that the term lateral lobe of tergum X is used instead of intermediate appendage. The term phallotremal sclerite complex is used to refer to internalized, sclerotized structures at the tip of the phallic apparatus, and consists of what has been previously referred to as a rod and ring structure and any additional associated sclerites. Terminology for structures of the head and thorax is given in Fig. 1. Venational terminology is given in Fig. 3A,B. Terminology for structures of the male genitalia is given in Fig. 4; female genitalic morphology is shown in Fig. 34. Forewing measurements are give to the nearest 0.1 mm. However, considerable variation in size occurs within species and the measurements only re¯ect the variability in the specimens examined (often a very limited sample). Female associations are presumptive, based on their co-occurrence with males. Lettering for the views of genitalia was standardized to aid in comparing the different species. Not all views are included ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

67

for each species, and this accounts for the non-consecutive lettering in some ®gures. Letters used for ®gures are, for male genitalia: A, lateral view; B, dorsal view of segments IX and X; C, inferior appendage, ventral view; D, inferior appendage, dorsal view; E, inferior appendage, caudal view; F, inferior appendages, oblique lateral view; G, phallic apparatus, lateral view; H, phallic apparatus, dorsal view; I, tergum VIII, dorsal view. Letters used for female genitalia are: A, lateral view; B, terga IX and X, dorsal view; C, segment IX and vaginal apparatus, ventral view. The oblique lateral view, when included, is provided to give an overall dimensional perspective of the pair of inferior appendages, but its orientation is not constant for all of the species. In general, the view is orientated caudally and somewhat ventrolaterally. Additional features illustrated in some ®gures are discussed in the corresponding ®gure legend. Holotype specimens deposited at the Museo de Historia Natural `Javier Prado' are on extended loan to the Smithsonian Institution. Unless otherwise stated, all material for newly described species is represented by pinned specimens. Method of curation is not indicated for described species. The following acronyms are used for depositories: CU, Cornell University, Ithaca, New York, U.S.A.; IBUNAM, Instituto BiologõÂa de la Universidad Nacional AutoÂnoma de MeÂxico, Mexico City, Mexico; INHS, Illinois Natural History Survey, Champaign, Illinois, U.S.A.; IZAM, Instituto de Zoologia Agricola, Maracay, Venezuela; MCZ, Museum of Comparative Zoology, Harvard, Massachusetts, U.S.A.; MJP, Museo de Historia Natural `Javier Prado', Universidad Nacional Mayor de San Marcos, Lima, Peru; MZUSP, Museu de Zoologia, Universidade de SaÄo Paulo, SaÄo Paulo, Brazil; NMNH, United States National Museum of Natural History, Smithsonian Institution, Washington, D.C., U.S.A.; RNHL, Rijksmuseum van Natuurlijke Histoire, Leiden, The Netherlands; UMSP, University of Minnesota Insect Collection, St Paul, Minnesota, U.S.A.; ZMUA, ZooÈlogisch Museum, Universiteit van Amsterdam, Amsterdam, The Netherlands; ZMUH, ZooÈlogische Institut und ZooÈlogisches Museum, UniversitaÈt von Hamburg, Hamburg, Germany. Cladistic analysis Phylogenetic analysis of ®fty-six characters was performed using PAUP 3.1.1 (Swofford, 1993). Chimarra koki and C. diannae were omitted from the analysis because these species were not examined during the study; however, their probable relationships are indicated in the species descriptions. Characters were polarized using seven outgroup species, including two species of Chimarrhodella, two members of subgenus Chimarrita, the incertae sedis species C. usitatissima and single representatives of subgenera Curgia and Chimarra. Characters used are listed in Appendix 1 and the data matrix is presented in Appendix 2. Both species of Chimarrhodella were designated as outgroup. All characters were treated as unordered and no differential weights were applied. The analysis was run using the heuristic option in PAUP, including 100 replicative searches.

68 Roger J. Blahnik Chimarra (Otarrha), subgen.n. Type species: Chimarra patosa Ross, 1956, present designation. Chimarra patosa group: Ross, 1956: 71; Flint, 1964: 23; Flint, 1971: 23; Flint, 1991: 28. Regardless of the evolutionary position of subgenus Otarrha within genus Chimarra, the group itself represents a well de®ned lineage, without obvious relationships to subgenera Curgia, Chimarrita or Chimarra. Character evidence supporting the monophyly of the latter subgenera was presented by Blahnik (1997), and evidence for the monophyly of Otarrha is presented in this paper. A diagnostic key to the subgenera, including species of Otarrha, can be found in the publication by Blahnik (1998). Otarrha is exclusively Neotropical in distribution, and particularly well represented in the Greater Antilles and the northern part of South America. It is also represented in the Lesser Antilles, but by species more similar to those on mainland South America than to those in the Greater Antilles. The subgenus is practically absent from Central America. Only one species, Chimarra rossi, has a distribution in lower Central America, including Panama and Costa Rica, and it has a close sister species in Colombia. Described species transferred to subgenus Otarrha include C. cubanorum Botosaneanu, C. diakis Flint, C. diannae Flint & Sykora, C. dominicana Flint, C. garciai Botosaneanu, C. guapa Botosaneanu, C. jamaicensis Flint, C. koki Botosaneanu, C. machaerophora Flint, C. patosa Ross, C. puertoricensis Flint, C. quadrifurcata Botosaneanu, C. retrorsa Flint, C. rossi Bueno, C. sensillata Flint, C. septemlobata Flint, C. septifera Flint and C. spinulifera Flint. New species described in this paper include C. amazonia, C. barinas, C. darlingtoni, C. diaphora, C. incipiens, C. odonta, C. parene, C. parilis, C. particeps, C. peruana, C. phthanorossi, C. redonda and C. tachuela. It is dif®cult to de®ne characters that universally apply to all species of Otarrha, but certain characters in combination are generally diagnostic. All species, except C. odonta, have the anterior setal warts divided and reduced venation in the hindwing, with R1 fused to the subcosta (or apparently so), Rs three-branched (®rst fork missing) and M two-branched. Males of most species also have enlarged, earlike preanal appendages and a ventral process that is relatively elongate, wider preapically than basally. Adult. Forewing length 3.3±6.9 mm; female usually somewhat larger than male. Colour generally brown or brownish to fuscous-black. Maxillary palp 5-segmented (Fig. 2): typically with ®rst segment very short, nearly globular; second segment about twice length of segment 1, apically with stiff bristlelike setae on mesal margin; third segment relatively elongate, about twice length of segment 2, inserted on outer apical margin of segment 2, narrow basally, with distinct dorsal bulge in basal half; segment four subequal to segment 4; segment 5 usually slightly

longer than segment 3. Labial palp 3-segmented. Head (Fig. 1) usually somewhat ¯attened, sometimes distinctly so, relatively elongate (with postocular parietal sclerite prominent, sometimes conspicuously so). Ocelli 3, large. Frontal setal wart divided into 3 setal areas or warts, an ovate ventral setal area and 2 oblique, dorsolateral setal areas; setal areas distinct and separated in some species, indistinct and merging in others. Dorsum of head with anterior, anteromesal, posterior and posterolateral setal warts; posterolateral setal warts large; anterior setal warts each divided into 2 small, circular setal warts (elongate and undivided in C. odonta). Prothorax with paired rounded mesal and lateral setal warts, mesal setal warts larger, sometimes somewhat irregular in shape (Fig. 1). Mesoscutellar setal wart rounded, not produced anteriorly, without anterior suture line. Spurs 1 : 4 : 4, foretibial spur prominent, meso- and metatibial spurs elongate. Male foretarsal claws not enlarged, symmetrical or nearly so. Forewing venation (Fig. 3) typical of Chimarra, with M4 absent; stem of Rs vein straight; s, r-m and m crossveins unpigmented and nearly linearly arranged; 2A and 3A anal veins both looped to 1A in proximal half of vein. Hindwing with venation reduced: R1 fused to subcosta, or nearly so, and Rs 3-branched (®rst fork absent) (in all species except C. odonta), M always 2-branched; anal loop very small. Male genitalia. Eighth segment narrow, frequently with paired projections (or mesal projection, or both) from posterodorsal margin (Figs 15B, 22I). Ninth segment synsclerous; anterior margin (in lateral view) broadly rounded, linear or with sinuous ventral expansion; segment almost always with sinuous dorsomesal and ventromesal invagination; in some species with (usually elongate, narrow) dorsomesal projection; posteroventrally with elongate ventral process, usually rounded apically and slightly constricted basally (tapering and acute in C. sensillata complex, Fig. 28A). Tergum X completely divided mesally, lobes typically without basal separation (Fig. 14B) or separated only slightly (Fig. 27B), infrequently widely separated (Fig. 24B); apically and ventrolaterally with numerous sensilla; lateral lobes either entire (Fig. 25A) or apically cleft (Fig. 23A), sometimes to base. Preanal appendages almost always large and ¯attened, earlike, with constricted base and rounded lateral margins (Fig. 27B). Inferior appendages 1-segmented, usually with distinct basal in¯ection (Fig. 20A), variously shaped, short or elongate. Phallotheca generally rather simple, tubular, with basodorsal expansion (typical of Chimarra, Fig. 24G). Endotheca membranous, short, eversible, sometimes with elongate dorsal expansion (Fig. 4G); phallic spines absent (typically) or 2±4 present (Fig. 20G) (sometimes apparently more nearly associated with phallotremal sclerite complex than endotheca). Phallotremal sclerite complex often elongate, half or more length of phallotheca (Fig. 26G), typically with rod and ring structure (Fig. 24G,H), rod often with ¯attened apicolateral projections (Fig. 23G), sometimes also with surrounding apical sclerite (Fig. 25G). Female genitalia. Sternum VII with short ventral process from posteromesal margin (Fig. 43A). Segment VIII short, synsclerous, with short apodemes from anterior margin (Fig. 43A) or apodemes absent (Fig. 49A); mid-laterally with ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

Systematics of Otarrha

69

Fig. 4. Chimarra amazonia, male genitalia.

row of elongate setae from posterior margin (Fig. 41A, setal bases illustrated, setae omitted for clarity); ventrally with diffuse and poorly de®ned paired setal areas (Fig. 45C) (setal areas well de®ned and separate in C. tachuela of sensillata species complex); posterolateral margin distinctly cleft below lateral setal area (Fig. 48A), often nearly to anterior margin of segment (Fig. 43A). Tergum IX short, wide, with apodemes from anterior margin (Fig. 46A,B); posteromesally with projecting setose area (Fig. 46B); sternum IX membranous. Tergum X composed of paired setose structures, each with bulbous and often distinctly sclerous basal region and less sclerous apical region (Fig. 43A); apically with elongate cercus. Vaginal apparatus typically short and membranous, with prominent anterior cuplike sclerite (Fig. 50C); usually also distinctly sclerotized apically (Fig. 49C), often with projecting elongate paired dorsal sclerites (these sometimes continuous with sclerous apicoventral region of rectal opening and dorsum of vaginal apparatus, Fig. 48D). Rectum usually opening apically (Fig. 48D), sometimes preapically (Fig. 43C). ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

Etymology. The subgeneric name is derived from the Greek otaros, meaning large-eared, in reference to the ¯attened, earlike preanal appendages of most species in this subgenus. Key to males of Otarrha species 1. ± 2(1). ±

Inferior appendages elongate, narrow (Figs 20A,C,D; 22A,C,D) .................................................. 2 Inferior appendages short, or relatively so (Figs 6A,C,D,F; 23A,C,D,F) (longest in members of sensillata complex, Fig. 28A,C,D) .................. 17 Inferior appendage with apex elongate, acuminate and very angled, retrorsely in¯ected (Fig. 24C,D) .............................................................. 3 Inferior appendage with apex not retrorsely in¯ected, sometimes bent at angle; apex sometimes acute, but not distinctly acuminate (Fig. 22C,D) ................ 5

70 Roger J. Blahnik 3(2).

Inferior appendage with very prominent rounded preapical projection, appearing apical because apex is retrorsely in¯ected (Fig. 24)............ quadrifurcata ± Inferior appendage without prominent rounded projection, sometimes widened preapically, apex obviously retrorsely directed (Fig. 10) .................. 4 4(3). Inferior appendage with apex elongate, acuminate (Fig. 10)....................................... dominicana ± Inferior appendage with apex short, acuminate (not illustrated) ........................................ diannae 5(2). Inferior appendage with distinct preapical, tinelike projection or branch on mesal surface (Fig. 20C,D) .............................................................. 6 ± Inferior appendage without preapical tine on mesal surface.................................................... 12 6(5). Tergum VIII with 2 very elongate projections from posterior margin (Fig. 20I) ............................. 7 ± Tergum VIII with short projections (Fig. 19B), or projections absent ........................................ 8 7(6). Projections from tergum VIII converging mesally, more or less uniformly covered with short spinelike setae (Fig. 20)...................................... patosa ± Projections from tergum VIII not noticeably converging, with elongate spinelike setae at apex (Fig. 21)........................................... peruana 8(6). Lateral lobes of tergum X very elongate and arched; segment IX much smaller in diameter than segment VIII; preanal appendages very large (Cuba) (Fig. 12).............................................. guapa ± Lateral lobes of tergum X neither greatly elongate nor arched; segment IX not much smaller in diameter than segment VIII; preanal appendages large but only relatively so (South America) .......................... 9 9(8). Inferior appendage with basal projection from mesal surface in addition to preapical projection, apex of ventral process of segment IX not ¯attened, truncate (Fig. 4) .......................................... amazonia ± Inferior appendages without basal projection; apex of ventral process of segment IX distinctly ¯attened, truncate (Fig. 19A)..................................... 10 10(9). Inferior appendages with mesal margin serrate proximally to tinelike projection, basally (Fig. 18C,D) or for entire length (Fig. 19C,D) . 11 ± Inferior appendage with mesal margin not serrate; tergum VIII with stout setae from posterior margin but not spinelike projections, tergum not narrowed dorsomesally (Fig. 17)............................ parene 11(10). Inferior appendages with mesal margin serrate only at base, not continuous to tine (Fig. 18).......... parilis ± Inferior appendage with serrations of mesal margin continuous to mesal tine (Fig. 19) .......... particeps 12(5). Tergum VIII with 2 elongate projections from posterior margin (Fig. 8) .......................... diakis ± Tergum VIII with short projections from posterior margin.................................................... 13 13(12). Mesal lobe of tergum X having sclerous projecting shelf; tergum X with short dorsolateral lobes, only slightly projecting; inferior appendage with mesal

±

14(13).

± 15(14). ± 16(15).

±

17(1). ± 18(17).

±

19(18). ± 20 (19). ± 21 (20). ±

curvature, not angularly bent (Fig. 9).................. .................................................... diaphora Mesal lobe of tergum X (if present) without sclerous projecting shelf; tergum X with elongate rodlike dorsolateral lobes (Fig. 22A,B); inferior appendages often with angular mesal in¯ection (Fig. 22C,D).... ............................................................ 14 Phallic apparatus with enormously elongate paired spines; lateral lobes of tergum X very strongly sclerotized; apex of inferior appendages unequally bi®d (Fig. 27)......................................... rossi Phallic spines often elongate but not enormously so; apex of inferior appendage not bi®d ................ 15 Ventrolateral lobes of tergum X short, rounded apically, not converging mesally (Fig. 22) ........... ............................................... phthanorossi Ventrolateral lobes of tergum X elongate, narrow apically, converging mesally (Fig. 30B) ........... 16 Apex of mesal lobe of tergum X with paired whiplike retrorsely directed projections; phallotremal sclerite complex modi®ed to form paired elongate needlelike dorsal spines and paired ventral sclerites (Fig. 26)........................................... retrorsa Apex of mesal lobe of tergum with paired acute projections not retrorsely directed; phallotremal sclerite complex with short dorsal spines and elongate ventromesal projection (Fig. 30)............. septifera Terga VII and VIII with paired dorsal projections; inferior appendage widened apically, basomesally with acute spine (Fig. 29) ............... septemlobata Tergum VII without dorsal projections; tergum VIII with projections present or absent; inferior appendage not as above............................................. 18 Lateral lobes of tergum X each deeply cleft apically; inferior appendage relatively elongate, apex truncate, much wider than base (Puerto Rico) (Fig. 23).................................... puertoricensis Lateral lobes of tergum X not apically cleft, sometimes with sclerous basal spine; inferior appendage not widened apically or with distinct apicodorsal projection (Fig. 6A).................................... 19 Tergum VIII with elongate dorsomesal projection (Jamaica) (Fig. 15) ..................... machaerophora Tergum VIII without elongate dorsomesal projection (but sometimes with paired processes from posterior margin of tergum VIII, Fig. 28B) ................... 20 Lateral lobes of tergum X very elongate, tapering and acute apically, projecting beyond inferior appendages (Fig. 11A) ............................................... 21 Lateral lobes of tergum X not greatly elongate, not or scarcely extending beyond inferior appendages (Fig. 16A) ............................................... 22 Lateral lobes of tergum X strongly curved, basodorsally with distinct short bi®d spine (Cuba) (Fig. 11)............................................ garciai Lateral lobes of tergum X without strong curvature, basodorsally without distinct spine (Jamaica) (Fig. 14)....................................... jamaicensis

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Systematics of Otarrha 22(20). Inferior appendage with prominent truncate incurved protuberance from apicoventral margin (Fig. 16) ..... ................................................................ odonta ± Inferior appendage without prominent protuberance or protuberance present on apicodorsal margin (Fig. 6A,F) .............................................. 23 23(22). Inferior appendage curved, tapering, apically dentate (Fig. 28C,D); tergum X short, simple, without prominent spines (sensillata species complex) .... 24 ± Inferior appendage otherwise; tergum X with basodorsal (Fig. 6A,B) or basolateral spines, or both, or spinules (Fig. 7A) ...................................... 27 24(23). Rod and ring of phallotremal sclerite complex with intricate surrounding sclerite, sclerite distinctly visible; phallotheca with prominent sclerotized apicoventral extension and elongate de¯exed dorsal extension (Fig. 13G) ................................... 25 ± Rod and ring of phallotremal sclerite complex without intricate surrounding sclerite, or not apparent; phallotheca without prominent sclerotized apicoventral extension, apicodorsal extension short, not distinctly de¯exed ...................................... 26 25(24). Ventral apex of phallotheca very elongate; inferior appendage with dentations extending to base as ridge on mesal surface (Fig. 28) ................... sensillata ± Ventral apex of phallotheca projecting but not greatly elongate; inferior appendage with dentations terminal only, not extending to base (Fig. 13) .................. .................................................... incipiens 26(24). Tergum VIII without projections from posterior margin; phallic apparatus without 2 small tacklike spines (Fig. 5) ................. .................. barinas ± Tergum VIII with short blunt projections from posterior margin; phallic apparatus with 2 small tacklike spines (Fig. 33) ....................... tachuela 27(23). Ventral process of segment IX (lateral view) very distinctly truncate apically (Dominican Republic) (not illustrated) ....................................... koki ± Ventral process not truncate apically ............... 28 28(27). Apex of inferior appendage sclerous, incurved, with several preapical teeth on ventral margin and incurved spine on proximal surface (Cuba) (Fig. 7) ........................................ darlingtoni ± Inferior appendage with rounded apicodorsal protuberance but without teeth or spines on ventral margin (Fig. 6A,F) .............................................. 29 29(28). Tergum X (dorsal view) narrow and tapering apically (Cuba) (Fig. 6)............................... cubanorum ± Tergum X (dorsal view) relatively broad, forming distinct rounded shoulders preapically (Hispaniola) (Fig. 31).................................................. 30 30(29). Tergum X with elongate basal spine and distinct small spine on mesal surface of inferior appendage (Haiti) (Fig. 31)................. spinulifera spinulifera ± Tergum X with short spines basally (on bulbous rounded structure), without spine on mesal surface of inferior appendage (Dominican Republic) (Fig. 32)............................ spinulifera galalcha ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

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Chimarra (Otarrha) amazonia, sp.n. (Figs 4A±D,G; 34A±C; 56) This species is similar to C. parene, C. parilis and C. particeps, but differs from them in several details of the male genitalia. The most notable differences are in the shape of the lateral lobes of tergum X, which lack whiplike apical and lateral projections, and the possession in C. amazonia of an acute basomesal projection on the inferior appendage, in addition to a preapical tine. A distinctive character of the phallic apparatus of this species is an elongate, lightly sclerous, dorsal sacklike structure, which terminates brushlike, with a number of small setae on the ventral surface. This structure is suggestive of those found in C. usitatissima and species of genus Chimarrhodella. A similar structure exists in the closely related C. parilis, although the apical setal brush is apparently absent in that species. The function of this structure is unknown. Adult. Forewing length (male) 4.9±5.4 mm (female) 5.5 mm. Colour nearly uniformly brown except head slightly darker, fuscous or blackish, with contrasting pale setal warts. Head only moderately elongate, postocular parietal sclerite not elongate. Male genitalia (Fig. 4). Tergum VIII with 2 short rounded projections from posterior margin. Abdominal segment IX (lateral view) with sinuous anterior margin, reaching its greatest width below mid-lateral point; in dorsal view with dorsomesal and ventromesal margins strongly invaginated; posteroventral process relatively elongate, projecting, slightly ventrally curved, rounded apically. Posterior margin of tergum IX (possibly mesal lobe of tergum X) with short mesal projection. Tergum X divided mesally, forming 2 sclerotized lateral lobes; lateral lobes each with acute sclerotized apical projection and rounded dorsal projection bearing numerous sensilla on dorsal margin, basally with several short setae or setaform sensilla. Preanal appendages of moderate size, ¯attened, earlike. Inferior appendage (in lateral view) angularly ¯exed at base, with elongate dorsal extension, appearing to narrow apically due to mesal curvature; with distinct mesal curvature in dorsal or ventral view, preapically with short tinelike projection, basally with acute mesal projection, apex of inferior appendage acute. Phallic apparatus slightly ¯exed basally and weakly developed dorsally, with elongate membranous sacklike extension ending in brushlike structure with numerous small setae on ventral surface, apex of phallotheca narrow, tubular, becoming membranous apically; endotheca with 2 elongate curved dorsal spines. Phallotremal sclerite complex composed of indistinct rod and ring structure, rod apically with short paired lateral extensions. Female genitalia (Fig. 34). Segment VIII short, with angular ventrolateral projections from posterior margin; anterior margin more or less linear in lateral view, with distinct short apodemes on lateral margin; posterior margin with elongate setae midlaterally, bordered ventrally by deep cleft, extending halfway or more to anterior margin; posterior margin in ventral view nearly linear, projecting at angle at lateral margins; posteroventral margin on either side of midline with several prominent setae. Tergum IX short, with moderately elongate

72 Roger J. Blahnik

Fig. 5. Chimarra barinas, male genitalia (G insets = dorsal view of apex of phallic apparatus and ventral view of phallotremal sclerite complex).

apodemes from anteroventral margin; posteromesally with projecting setose area, wide at base and abruptly sinuous and narrowed apically. Sternum IX membranous. Tergum X bilobed, each lobe divided into prominent rounded sclerous basal region and less sclerous short apical region with cercus at apex. Vaginal apparatus largely membranous, with prominent paired elongate conical sclerites on apicodorsal margin; sclerous projections continuously sclerous basally with dorsum of vaginal apparatus and venter of rectum; rectum opening apically; vaginal apparatus widened anteroventrally, with de¯exed cuplike sclerite. Material examined. Holotype, Y, PERU: Madre de Dios, Amazonia Lodge, Toma del Agua (stream), 12°52.22¢S 71°22.56¢W, 415 m, 29.vi.1993 (R. Blahnik and M.

Pescador) (MJP). Paratypes, PERU: Madre de Dios, 1Y, 1X, same data as holotype (NMNH, MJP). Etymology. This species takes its name from Amazonia Lodge on the Madre de Dios River in Peru, near which the type specimens were collected. Chimarra (Otarrha) barinas, sp.n. (Figs 5A±D,G; 56) This is the ®rst of four very similar species, which I will refer to here as the C. sensillata species complex. In addition to C. barinas, the complex includes C. incipiens, C. sensillata and C. tachuela. The male of C. barinas is most similar to C. tachuela, and shares with that species the absence of a distinct apical sclerite surrounding the rod and ring structure of the ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

Systematics of Otarrha phallotremal sclerite complex and a phallotheca without an acute ventral apex. It differs in having tergum VIII with short dorsal projections and in lacking 2 tacklike spines associated with the apex of the phallotremal sclerite complex. Adult. Forewing length (male) 4.9±5.1 mm. Colour nearly uniformly brown, except head and thorax slightly darker, fuscous or blackish. Head relatively elongate, ¯attened; postocular parietal sclerite large, distinctly elongate and in¯ated (slightly projecting below eye). Male genitalia (Fig. 5). Tergum VIII with short paired projections from posterior margin, rounded or subtruncate at apex, narrowly separated basally. Abdominal segment IX with anterolateral margin sinuously expanded ventrally; ventral expansion not greatly elongate, without mesal invagination; posteroventral process narrow, elongate, projecting, nearly uniform in width, rounded apically. Tergum X completely divided mesally into 2 simple and relatively short lobes, rounded apically, contiguous basally, each with numerous apical and ventrolateral sensilla; lobes distinctly angular and incurved basally, forming ridgelike extension on mesal surface. Preanal appendages rounded, constricted basally, of moderate size, only slightly ¯attened. Inferior appendage simple in pro®le; slightly upcurved and tapering in lateral view; with mesal curvature in dorsal view, apically with abruptly incurved sclerotized teeth. Phallic apparatus with tubular phallotheca, slightly constricted basally, apicodorsally with short, lightly sclerous, keelshaped projection; phallic spines absent. Phallotremal sclerite complex with very prominent rod and ring structure, extending more than half length of phallotheca, rod with gradually widened lateral platelike projections. Female. Unknown. Material examined. Holotype, Y, VENEZUELA: Barinas, 22 km NW Barinitas, 19.ii.1976 (C. M. and O. S. Flint, Jr) (NMNH). Paratype, VENEZUELA: Barinas, 1Y, 30 km NW Barinitas, 24.ii.1976 (C. M. and O. S. Flint, Jr) (NMNH). Etymology. This species is named barinas for the state in Venezuela where the type specimens were collected. Chimarra (Otarrha) cubanorum Botosaneanu (Figs 6A±D,F,G; 57) Chimarra cubanorum Botosaneanu, 1980: 69, 98±99, Fig. 3a±c, Y, Cuba (ZMUA). This species is nearly identical to C. spinulifera Flint, differing primarily in its provenance, size and minor details of tergum X. In C. cubanorum, the lateral lobes of tergum X in dorsal view are narrower and less abruptly incurved apically (less shouldered). Also, the structure of the spine at the base of tergum X is different in this species and in each of the two subspecies of C. spinulifera. As the differences are probably diagnostic, and there is no way to assess the signi®cance of the differences in allopatry, I retain this as a valid species. However, the differences between this species and C. spinulifera are hardly more distinctive than those between ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

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the subspecies of C. spinulifera. For additional character discussion, see the diagnosis under C. spinulifera. Adult. Forewing length (male) 3.6 mm. Colour (in alcohol) brown. Head ¯attened, postocular parietal sclerite only slightly elongate. Male genitalia (Fig. 6). Tergum VIII with short wide truncate mesal projection from posterior margin. Abdominal segment IX with anterolateral margin widest ventrad of midlateral, abruptly narrowing ventrally, gradually narrowing dorsally; dorsomesal and ventromesal margins strongly invaginated; posteroventral process very elongate, projecting, narrow basally, acute apically. Tergum X completely divided mesally into 2 sclerotized lobes, narrowly separated basally, tapering apically, with distinct mesal curvature in apical third; basomesally with rounded projection, bearing sensilla and prominent, sclerotized, irregular, spinelike projection; basolaterally with elongate curved spine; laterally with short spines and numerous sensilla. Preanal appendages large, ¯attened, earlike. Inferior appendage, in lateral view, short, rounded basally, with thumblike apicodorsal projection; with slight mesal curvature in dorsal or ventral views. Phallic apparatus with narrow tubular phallotheca, slightly constricted basally, expanded basodorsally; endotheca short, membranous. Phallotremal sclerite complex large, about half length of phallotheca, composed of rod and ring structure, ring with elongate dorsal extension. Female. Unknown. Material examined. Holotype, Y, CUBA: Oriente, Cupeyal, vi.1971 (I. Garcia) (ZMUA). Chimarra (Otarrha) darlingtoni, sp.n. (Figs 7A±D,F,G; 56) This species bears a general similarity to C. cubanorum and C. spinulifera, but is easily distinguished by the structure of its inferior appendages, which are less narrowed and possess several sclerous teeth. Additionally, unlike C. cubanorum and C. spinulifera, the inferior appendage possesses a distinct spine on the mesal surface. Adult. Forewing length (male) 4.4 mm. Colour more or less uniformly brown; setal warts of posterior part of head and thorax pale brown. Head only moderately elongate, postocular parietal sclerite not prominent or elongate. Male genitalia (Fig. 7). Tergum VIII unmodi®ed. Abdominal segment IX with anterolateral margin extended and broadly rounded; dorsomesal and ventromesal margins distinctly invaginated; posteroventral process elongate, projecting, rounded apically. Tergum X relatively simple, completely divided mesally into 2 elongate sclerotized lobes, each rounded apically with numerous sensilla along ventral margin; basodorsally with prominent posteriorly directed spine; basolaterally with several additional small spines. Preanal appendages large, ¯attened, earlike. Inferior appendage, in lateral view, short, rounded basally, apically with broadly sclerotized region; apicoventral margin with distinct teeth; mesal surface with prominent, hooked spine. Phallic apparatus with simple tubular phallotheca, expanded basodor-

74 Roger J. Blahnik

Figs 6, 7. 6, Chimarra cubanorum, male genitalia; 7, C. darlingtoni, male genitalia.

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Systematics of Otarrha sally; endotheca short, membranous. Phallotremal sclerite complex composed of rod and ring structure, ring narrow with elongate dorsal extension; rod elongate, nearly two-thirds length of phallotheca, with preapical ventral process. Female. Unknown. Material examined. Holotype, Y, CUBA: coast below Pico Turquino, 26±30.vi.1936 (Darlington) (MCZ). Etymology. I take great pleasure in naming this species after P. J. Darlington, Jr, eminent entomologist and biogeographer, who collected the type specimen. Chimarra (Otarrha) diakis Flint (Figs 8A±D,G,I; 56) Chimarra (Chimarra) diakis Flint, 1971: 23, Figs 35±37; Y, Brazil (NMNH). This is a distinctive species, similar only to C. diaphora, but unlikely to be confused with that species because of the more elongate processes on tergum VIII of the male. It differs from C. diaphora also in possessing lateral lobes of tergum X which are wider in dorsal view, and a mesal lobe with a shorter sclerous projection. The mesal lobe is one of the most distinctive characteristics of C. diakis and C. diaphora, and has no clear homology in other species with similarly shaped inferior appendages. Both of these species are also similar in possessing a short phallotheca with a distinctive pair of prominent, ¯attened spines (undoubtedly derived as a modi®cation of the phallotremal sclerite complex). Adult. Forewing length (male) 4.7 mm. Colour (in alcohol) pale brown. Head distinctly ¯attened, elongate; postocular parietal sclerite elongate. Male genitalia (Fig. 8). Tergum VIII with paired narrow elongate projections, narrowly separated mesally, each with numerous small spinelike projections. Abdominal segment IX widest mid-laterally, narrowed anterodorsally and ventrally; anterodorsal margin strongly invaginated mesally, anteroventral margin nearly linear mesally, slightly invaginated; posteroventral process elongate, projecting, rounded apically; (apparently) with elongate dorsomesal lobe from posterior margin, rounded apically and with sclerotized short upturned process before apex. Tergum X forming 2 short sclerotized lateral lobes, projecting dorsally and with numerous sensilla; lobes wide in dorsal view. Preanal appendages large, ¯attened, earlike. Inferior appendage, in lateral view, elongate, appearing tapered apically due to ¯attening of apex; in dorsal and ventral views, with pronounced mesal curvature; apically rounded and dorsoventrally ¯attened, with extreme apex forming short acute mesal projection. Phallic apparatus with short tubular phallotheca, distinctly expanded basodorsally; endotheca short, membranous. Phallotremal sclerite complex prominent, nearly length of phallotheca, composed of rod and ring structure and complex apical sclerite; ring distinct, rod elongate, apical sclerite with dorsal ring and elongate ventral troughlike sclerite, divided apically into paired ¯attened acute sclerites. Female. Unknown. ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

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Material examined. Holotype, Y, type no. 74118, BRAZIL: Gebiet Endstation, Rio Marauria, Bergbach II, etwa 350 m uÈber dem Meeresspiegel, schattig, starkes GefaÈlle uÈber GranitbloÈcke, 26.i.1963 (E. J. Fittkau) (NMNH). Chimarra (Otarrha) diannae Flint & Sykora Chimarra diannae Flint & Sykora, 1993: 48, 50±51, Figs 1±3; Y, St Lucia (NMNH). This species is probably most similar to C. dominicana, but also shares character similarities with the more recently described C. quadrifurcata. These three species together constitute a small group of closely related species, all described from islands in the Lesser Antilles. From C. dominicana it can be distinguished by having the apex of the inferior appendages much less elongate, toothlike rather than spinelike; from C. quadrifurcata it is distinguished by having the inferior appendage less developed preapically, merely enlarged and rounded rather than forming a distinctly protruding projection. Chimarra (Otarrha) diaphora, sp.n. (Figs 9A±D,G,I; 35A±C; 56) This species is similar to C. diakis, and I initially considered it a variant of that species. It is recognized as a species here based on diagnostic characters. The most distinctive differences lie in the structure of terga VIII, IX and X. The posterior processes of tergum VIII are short, as opposed to very elongate in C. diakis. Additionally, the lateral lobes of tergum X are narrow rather than wide in dorsal view, and the mesal lobe has a sclerotized process which is elongate and projecting, as opposed to short and upturned. Only additional collecting will reveal how widespread the respective forms are and whether they intergrade. Adult. Forewing length (male) 4.7±5.4 mm (female) 5.1± 5.5 mm. Colour more or less uniformly medium brown, except head blackish-brown, and proximal part of antennae with segments annulate, brown basally, pale stramineous apically. Head slightly ¯attened dorsally, postocular parietal sclerite moderately elongate (about as long as wide) Male genitalia (Fig. 9). Tergum VIII with short paired projections from posterior margin, each with numerous small spinelike projections. Abdominal segment IX widest mid-laterally, narrowed anterodorsally and ventrally; anterodorsal margin strongly invaginated mesally, anteroventral margin with less pronounced rounded mesal invagination; posteroventral process elongate, projecting, rounded apically; (apparently) with elongate dorsomesal lobe from posterior margin, rounded apically and with distinctly projecting sclerotized process before apex. Tergum X forming 2 short sclerotized lateral lobes, projecting dorsally, with numerous sensilla; lobes narrow in dorsal view. Preanal appendages large, ¯attened, earlike. Inferior appendage, in lateral view, elongate, appearing tapered apically

76 Roger J. Blahnik

Figs 8, 9. 8, Chimarra diakis, male genitalia (G inset = dorsal view and enlargement of phallotremal sclerite complex); 9, C. diaphora, male genitalia.

ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

Systematics of Otarrha due to ¯attening of apex; in dorsal and ventral views with pronounced mesal curvature; apically rounded and dorsoventrally ¯attened, with extreme apex forming short acute mesal projection. Phallic apparatus with short tubular phallotheca, distinctly expanded basodorsally; endotheca short, membranous. Phallotremal sclerite complex prominent, nearly length of phallotheca, composed of rod and ring structure and complex apical sclerite; ring distinct, rod elongate, apical sclerite with dorsal ring and elongate ventral troughlike sclerite, divided apically into paired ¯attened acute sclerites. Female genitalia (Fig. 35). Segment VIII moderately extended anteroventrally; anterolateral margin with short mesally directed apodemes; posterior margin with rounded projection mid-laterally, bearing elongate marginal setae, bordered ventrally by cleft from posterior margin, cleft relatively deep, extending halfway or more to anterior margin; segment ventrolaterally with indistinct sclerous suture line extending from posterior cleft to anterior margin; posterior margin in ventral view, linear with several prominent setae from posterior margin on either side of midline and numerous smaller setae on venter of segment. Tergum IX short with moderately elongate apodemes from anteroventral margin, posteromesally with subquadrate projecting setose area. Sternum IX membranous. Tergum X bilobed, each lobe divided into prominent rounded sclerous basal region and less sclerous apical region with cercus at apex. Vaginal apparatus with lightly sclerous collar at apex, ventrally with rounded indentation; at mid-length with prominent cupped sclerite, tubelike anteriorly and with dorsally emerging tube open posteriorly; anteroventrally with indistinct ¯attened sclerite. Material examined. Holotype, Y, VENEZUELA: T. F. Amazonas, Agua Blanca, 0°49¢N 66°08¢W, Cerro de la Neblina, 160 m, 20±21.iii.1984 (O. Flint and J. Louton) (NMNH). Paratypes, 5Y, 3X, VENEZUELA: T. F. Amazonas, same data as holotype (IZAM, NMNH, UMSP). Etymology. This species is named from the Greek word diaphoros, meaning different, and referring to the fact that, while being similar to C. diakis, it is also diagnostically different. Chimarra (Otarrha) dominicana Flint (Figs 10A±D,G,H; 36A±C; 56) Chimarra dominicana Flint, 1968b: 10, 12, Figs 12±17; Y, Dominica (NMNH). Malicky, 1983: 264; Botosaneanu, 1988: 221; Botosaneanu, 1990: 43; Flint & Sykora, 1993: 49. This species is most easily distinguished by the shape of the inferior appendage, which has its apex elongate, acuminate and very strongly retrorsely ¯exed. It is similar in the last respect to C. diannae and C. quadrifurcata, both of which also have inferior appendages with retrorsely ¯exed apices, but in C. diannae the apex of the inferior appendage is much shorter, ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

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toothlike rather than spinelike, and in C. quadrifurcata the acuminate apex is accompanied by an additional rounded preapical process. Chimarra quadrifurcata also has a pair of elongate phallic spines, which are absent in both C. dominicana and C. diannae. Adult. Forewing length (male) 4.5±4.6 mm (female) 5.2 mm. Colour more or less uniformly brown. Head distinctly ¯attened, elongate; postocular parietal sclerite elongate. Male genitalia (Fig. 10). Tergum VIII with short acute paired projections from posterior margin. Abdominal segment IX widest mid-laterally, much narrowed anterodorsally and ventrally; anterodorsal and anteroventral margins strongly invaginated mesally; posteroventral process relatively elongate, projecting, rounded apically; dorsomesally with short truncate lobe from posterior margin, only lightly sclerotized. Tergum X forming 2 sclerotized lateral lobes rather widely separated by mesal lobe of segment IX; each lateral lobe cleft longitudinally into dorsal and ventral portions, dorsal portion simple, short, rounded apically, with numerous sensilla; ventral portion more elongate, apically with 2 sclerotized protuberances, one distally on dorsal margin and other preapical on ventrolateral margin. Preanal appendages large, ¯attened, earlike. Inferior appendage elongate with pronounced mesal curvature; apically narrowed to elongate acuminate re¯exed tip; in dorsal and ventral views, with apical third of inferior appendage very strongly curved, acuminate apex re¯exed and mesally directed. Phallic apparatus with slightly curved tubular phallotheca expanded basodorsally; endotheca short, membranous. Phallotremal sclerite complex prominent, very elongate, nearly length of phallotheca, composed of rod and ring structure and complex apical sclerite; ring indistinct, rod very elongate, apical sclerite indistinct except apex strongly sclerotized, ¯attened, curved and with 2 upturned spinelike structures apically; preapically with lightly sclerotized, winglike extensions. Female genitalia (Fig. 36). Segment VIII elongate ventrally, anterior and posterior margins both extended; anterior margin undulate in lateral view, without apodemes; posterior margin with rounded projection mid-laterally, bearing elongate setae on margin, bordered ventrally by cleft from posterior margin, cleft only shallowly indentate; ventrolaterally with indistinct suture line extending from posterior cleft to anterior margin; posteroventral margin linear, posterolaterally with several prominent setae on either side of midline and numerous smaller setae on venter of segment. Intersegmentally with prominent receptacle for inferior appendage between segments VII and VIII. Tergum IX short with short apodemes from anteroventral margin, posteromesally with projecting setose area, truncate apically. Sternum IX membranous. Tergum X bilobed, each lobe divided into prominent rounded sclerous basal region and less sclerous apical region with cercus at apex. Vaginal apparatus relatively short and more or less sclerous in apical half; apex with sclerous collar bearing distinct notch on ventral margin; laterally with prominent rounded,

78 Roger J. Blahnik

Fig. 10. Chimarra dominicana, male genitalia (H = dorsal view and enlargement of apex of phallic apparatus and phallotremal sclerite complex).

sclerous projections; anteroventrally with cuplike receptacle located before apex of anterior margin. Material examined. DOMINICA: 1X allotype, 2.2 mi. E of Pont Casse, 11.v.1964 (O. S. Flint, Jr) (NMNH); 1Y paratype, 2.2 mi. E Pont Casse, 1.v.1964 (O. S. Flint, Jr) (NMNH); 1Y, 4X paratypes, 0.5 mi. W Pont Casse, 22±24.vii.1963 (O. S. Flint, Jr) (NMNH); 1X paratype, Pont Casse, 27±30.xi.1964 (P. J. Spangler) (NMNH); 4Y, 10X paratypes, 0.4 mi. E Pont Casse, (various dates) 21.iv.-23.vi.1964 (O. S. Flint, Jr) NMNH; 1Y, 5X paratypes, 0.5 mi. S Pont Casse, (various dates) 8.iv.-17.vi.1996 (O. S. Flint, Jr) (NMNH); 4X paratypes, 1.3 mi. E Pont Casse, (various dates) 29.iv.-11.vi.1964 (O. S. Flint, Jr) (NMNH); 1Y paratype, Fond Figues R., 400 m, 29.iv.1966 (R. J. Gagne) (NMNH); 1X paratype, Sylvania, 1800 m, 9.ii.1964 (Dale F. Bray) (NMNH).

Chimarra (Otarrha) garciai Botosaneanu (Figs 11A± D,F,G; 57) Chimarra garciai Botosaneanu, 1980: 99±101, Fig. 4a±c; Y, Cuba (ZMUA) . This species is most similar to C. jamaicensis, differing primarily in having tergum X more arched and with sensilla located laterally along the whole length of the tergum rather than concentrated basolaterally. It also differs in having an inferior appendage with a shorter apicomesal process. Adult. Forewing length (male) 4.7±4.8 mm. Colour (in alcohol) brownish. Postocular sclerite only moderately elongate. ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

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Fig. 11. Chimarra garciai, male genitalia.

Male genitalia (Fig. 11). Tergum VIII with short paired acute mesally directed processes from posterior margin. Abdominal segment IX widest mid-laterally, anterolateral margin broadly rounded, narrowed dorsally and ventrally, ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

posterolateral margin angularly projecting at level of inferior appendages; anterodorsal and anteroventral margins strongly invaginated mesally; posteroventral process elongate, projecting, narrowed basally, subacute apically. Tergum X divided

80 Roger J. Blahnik

Fig. 12. Chimarra guapa, male genitalia.

mesally, forming 2 elongate sclerotized lateral lobes; each strongly tapered and arched, with numerous sensilla along entire length of ventrolateral margin, apically with several small spinelike projections. Preanal appendages very large, ¯attened, earlike. Inferior appendage short, rounded basally, tapering apicodorsally, terminating in short projecting process from apicodorsal surface; ventral margin of process extending

onto mesal surface of appendage. Phallic apparatus with phallotheca narrow, tubular, expanded basodorsally; endotheca elongate, membranous, terminating in small patch of setae, dorsally bulging and membranous. Phallotremal sclerite complex composed of rod and ring structure, about one-fourth length of phallotheca; ring distinct, with dorsal projection, rod only relatively elongate. ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

Systematics of Otarrha Female. Unknown. Material examined. Holotype, Y, CUBA: Oriente, Cupeyal, vi.1971 (I. Garcia) (ZMUA). Chimarra (Otarrha) guapa Botosaneanu (Figs 12A± D,G; 37A±C; 56) Chimarra guapa Botosaneanu, 1977: 252±253, Figs 30±34, 34 bis; Y, Cuba (ZMUA); Botosaneanu, 1980: 92; Botosaneanu, 1994a: 459±460, Fig. 20. This is a distinctive species whose most diagnostic characteristics are found in the structure of the inferior appendage of the male, which is elongate and possesses a blunt, tinelike mesal process, and by the elongate arched lateral lobes of tergum X. In general form, the inferior appendages are super®cially similar to C. patosa and several other newly described Peruvian species, but quite unlike any other Antillean species. However, in the shape of segment IX, with very rounded anterior margins, and in the structure of the lateral lobes of tergum X and preanal appendages, the species is very much like other Antillean species, and this is where the true af®nities of C. guapa no doubt lie. Adult. Forewing length (male) 4.3±4.7 mm (female) 4.5 mm. Colour (in alcohol) pale brown, setal warts contrastingly paler in colour, pale stramineous. Head only moderately elongate, rounded dorsally (not ¯attened); postocular parietal sclerite not prominent, only moderately elongate. Male genitalia (Fig. 12). Tergum VIII with elongate narrow posteromesal projection, projection divided apically. Abdominal segment IX with anterolateral margin expanded, rounded, narrowed dorsally and ventrally; posterolateral margin angularly projecting at position of inferior appendages; anterodorsal and anteroventral margins with strong mesal invagination; posteroventral process elongate, projecting, prominent, slightly curved, narrowed basally, subacute apically. Tergum X divided mesally into 2 narrow sclerotized lateral lobes, each very elongate and arched, with numerous sensilla along ventral and apical margins, preapically with small spinelike projection; basoventral margin of lateral lobe with sclerotized bi®d spinelike projection. Preanal appendages very large, ¯attened, earlike. Inferior appendage elongate, narrowing to acute apex; in dorsal and ventral views, with mesal curvature, acute apex and branch or tine at mid-length on mesal surface. Phallic apparatus with phallotheca narrow tubular, slightly curved and relatively elongate, distinctly expanded basodorsally; endotheca short, membranous. Phallotremal sclerite complex short, composed of indistinct rod and ring structure and apical sclerite; ring with elongate dorsal projection, rod relatively short, apical sclerite detached, short, composed of ring and short decurved ventral sclerite. Female genitalia (Fig. 37). Segment VIII short; anterolateral margin nearly linear, slightly projecting at position of apodemes but apodemes undeveloped; posterior margin with rounded projection mid-laterally, bearing elongate setae on margin, bordered ventrally by deep cleft from posterior ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

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margin, extending nearly to anterior margin; posteroventral margin, in lateral view projecting and cupped; in ventral view rounded, with mesally separated setal areas bearing small group of elongate setae. Tergum IX short with short apodemes from anteroventral margin, posteromesally with subquadrate projecting setose area. Sternum IX membranous. Tergum X bilobed, each lobe divided into prominent rounded lightly sclerous basal region and less sclerous apical region with cercus at apex. Vaginal apparatus with yokelike sclerous region apically, sclerous ventrally and open dorsally; apicodorsally with prominent elongate projecting conical sclerites; rectum opening apically above sclerites; vaginal apparatus, largely membranous anteriorly, with de¯exed cuplike sclerite. Material examined. CUBA: Oriente, 3Y paratypes, RõÂo Jojo, 24.ii.1973 (L. Botosaneanu) (NMNH); 4Y, 1X paratypes, Mayari, Arroyo Seboruguito, 28.ii.1973 (L. Botosaneanu) (NMNH). Chimarra (Otarrha) incipiens, sp.n. (Figs 13A±D,G; 56) This is the second of four species recognized in the C. sensillata species complex. The species is most similar to C. sensillata, differing only in minor details. It shares with that species the presence of widely separated, short, acute projections from the posterodorsal margin of segment VIII, and a phallotheca with an acute ventral apex, but differs in lacking a toothed, basomesal ridge on the inferior appendage. However, the latter character in C. sensillata is probably homologous with the row of apical teeth found in C. incipiens, and simply represents an extension of this structure onto the mesal surface. Adult. Forewing length (male) 5.5±5.7 mm. Colour nearly uniformly brown, head and thorax slightly darker, fuscous or blackish. Head relatively elongate, ¯attened; postocular parietal sclerite large, distinctly elongate and in¯ated (slightly projecting below eye). Male genitalia (Fig. 13). Tergum VIII with short paired projections from posterior margin, acute or subacute apically, widely separated basally. Abdominal segment IX with anteroventral margin distinctly expanded, invaginated mesally; posteroventral process narrow, elongate, projecting, acute apically, gradually tapering from base. Tergum X completely divided mesally into 2 simple short apically rounded lobes, contiguous basally, each with numerous apical and ventrolateral sensilla; ventrolateral sensilla enlarged. Preanal appendages short, tapering, scarcely ¯attened. Inferior appendage simple in pro®le; in lateral view distinctly upcurved, tapering; in dorsal view noticeably short, with pronounced, almost angular, mesal curvature, apically with strongly incurved sclerotized teeth. Phallic apparatus with tubular phallotheca, slightly constricted basally; apicodorsally with lightly sclerous elongate de¯exed projection; apicoventrally with sclerous acutely projecting extension, slightly upcurved, not greatly elongate; phallic spines absent. Phallotremal sclerite complex with very prominent rod and ring structure, extending more than half length of phallotheca; rod with abruptly widened

82 Roger J. Blahnik

Fig. 13. Chimarra incipiens, male genitalia (G inset = ventral view of phallotremal sclerite complex).

lateral platelike projections; rod and ring structure subtended at mid-length by complex sclerite. Female. Unknown. Material examined. Holotype, Y, VENEZUELA: Distrito Federal, Bajo Seco, Est. Exp. Bajo Seco, c. 15 km NE Colonia Tovar, 2000 m, 22±25.i.1983 (O. S. Flint, Jr) (NMNH). Paratypes, VENEZUELA: Distrito Federal, 8Y, same data as holotype (IZAM, NMNH, UMSP). Etymology. This species is named incipiens because of the minimal and incipient differences that differentiate it from C. sensillata. Chimarra (Otarrha) jamaicensis Flint (Figs 14A±D,G; 38A±C; 56) Chimarra jamaicensis Flint, 1968a: 18, 21, 22, 25, 27, 29; Y, Jamaica (NMNH).

20,

Figs

This species is most similar to C. garciai, especially in the shape of the inferior appendages and the elongate lobes of tergum X. It differs in that the apical lobe of the inferior appendage is narrower and more mesal in position, and also in having a discrete sensillate region on the lateral margin of tergum X. Also, unlike C. garciai, C. jamaicensis lacks spinelike projections at the base of the lateral lobes of tergum X. Chimarra jamaicensis also is quite similar to C. machaerophora, but notably lacks the elongate dorsal projection found on tergum VIII of the males of that species. Adult. Forewing length (male) 5.2±5.9 mm (female) 5.1± 5.5 mm. Colour more or less uniformly brown. Head relatively elongate, ¯attened; postocular parietal sclerite prominent, elongate. Male genitalia (Fig. 14). Tergum VIII unmodi®ed dorsally. Abdominal segment IX broadly rounded anterolaterally, narrowed and mesally invaginated dorsally and ventrally, posterolateral margin with angular extension at ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

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Fig. 14. Chimarra jamaicensis, male genitalia (G inset = dorsal view of phallotremal sclerite complex).

level of inferior appendages; posteroventral process elongate, projecting, narrowed basally, tapering and rounded apically. Tergum X divided mesally, forming 2 elongate sclerotized upwardly directed lateral lobes; each strongly tapered apically, basolaterally with numerous sensilla, apically and basomesally with several small spinelike ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

projections. Preanal appendages very large, ¯attened, earlike. Inferior appendage short, rounded basally and apically, with distinct preapical projection from dorsomesal surface. Phallic apparatus with phallotheca narrow, tubular, expanded basodorsally; endotheca membranous. Phallotremal sclerite complex about one-third length of

84 Roger J. Blahnik

Fig. 15. Chimarra machaerophora, male genitalia.

phallotheca, composed of rod and ring structure; ring distinct, with elongate paired dorsal projections, rod only relatively elongate. Female genitalia (Fig. 38). Segment VIII short; anterior margin slightly projecting at position of apodemes, apodemes broadly rounded, undeveloped; posterior margin with rounded mid-lateral projection bearing elongate setae, projection bordered ventrally by deep cleft from posterior margin, extending nearly to anterior margin; posteroventral margin, in lateral view, projecting and distinctly cupped; in ventral view nearly linear, with rounded lateral margins; ventrally with row of elongate setae on posterior margin.

Tergum IX very short, with moderately elongate apodemes from anteroventral margin; posteromesally with projecting short broadly rounded setose area. Sternum IX membranous. Tergum X bilobed, each lobe divided into prominent rounded lightly sclerous basal region and less sclerous apical region with cercus at apex. Vaginal apparatus short, ventrally with lightly sclerous rounded crescentic sclerite; dorsolaterally with elongate paired sclerites, each ¯attened anteriorly, narrowed to short conical projection posteriorly; anteriorly with lightly sclerous de¯exed cupped sclerite; anterodorsally with projecting membranous region. ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

Systematics of Otarrha Material examined. JAMAICA: 29Y, 3X paratypes, St Andrew Parish, Hardwar Gap, Dicks Pond Trail, 25.vii.1962 (O. Farr and R. Flint) (NMNH); 22Y, 5X paratypes, same location, 16.vii.1963 (Flint and Farr) (NMNH); 2Y paratypes, St Andrew Parish, small stream, 1 1/8 mi. SW crossing Dicks Pond Trail, Hardwar Gap, 26.ix.1963 (W. L. Peters and T. H. Farr) (NMNH); 9Y, 4X paratypes, Portland Parish, Hardwar Gap, waterfall, 17.vii.1963 (Flint and Farr) (NMNH). Chimarra (Otarrha) koki Botosaneanu Chimarra (Chimarra) koki Botosaneanu, 1996: 11, Figs 7±12; Y, Dominican Republic (ZMUA). This species was only recently described and is not redescribed or reillustrated here. It is perhaps most similar to C. redonda, and like that and several other Antillean species it has an inferior appendage with an apicomesal toothlike projection. The truncate ventral process of segment IX of the male in this species is a character unique among the Antillean members of the subgenus and would serve to differentiate it from any of these species. A similarly developed ventral process is found in C. septemlobata, which, however, has distinctly different inferior appendages, as well as a mainland distribution. Chimarra (Otarrha) machaerophora Flint (Figs 15A±D, F,G; 39A±C; 56) Chimarra machaerophora Flint, 1968a: Figs 18, 19, 20, 26; Y, Jamaica (NMNH).

20±21,

This species is most similar to C. jamaicensis and C. garciai, although males can be easily distinguished by the possession of an elongate mesal process on tergum VIII, and by having tergum X shorter, wider and less tapered apically than these species. Additionally, the inferior appendage has an apical process which is very short and broad, as compared to narrow in the other two species. Adult. Forewing length (male) 6.0±6.5 mm (female) 5.8± 6.7 mm. Colour more or less uniformly brown. Head relatively elongate, ¯attened; postocular parietal sclerite prominent, elongate. Male genitalia (Fig. 15). Tergum VIII with elongate narrow mesal projection bearing scattered setae. Abdominal segment IX with anterolateral margin nearly linear, narrowed anterodorsally and ventrally, posterolateral margin slightly angularly projecting at level just below inferior appendages; anterodorsal and anteroventral margins invaginated mesally; posteroventral process elongate, projecting, narrowed basally, nearly acute apically. Tergum X divided mesally, forming 2 elongate sclerotized lateral lobes; each rather broadly rounded apically with numerous scattered sensilla concentrated basolaterally and apically. Preanal appendages large, ¯attened, earlike. Inferior appendage short, with short broad apicodorsal ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

85

projection extending onto mesal surface. Phallic apparatus with phallotheca short, tubular, expanded basodorsally; endotheca short, membranous. Phallotremal sclerite complex about half length of phallotheca, composed of rod and ring structure; ring large with dorsal projection, rod moderately elongate. Female genitalia (Fig. 39). Segment VIII short; anterior margin slightly projecting at position of apodemes, apodemes broadly rounded, undeveloped; posterior margin with rounded midlateral projection bearing elongate setae, projection bordered ventrally by cleft from posterior margin, extending halfway to anterior margin; posteroventral margin, in lateral view, projecting and somewhat cupped; in ventral view nearly linear, with lateral margins rounded and strongly re¯exed onto opposite surface; apicoventral margin with group of elongate setae on either side of midline and numerous smaller setae. Tergum IX very short, with moderately elongate apodemes from anterodorsal margin; posteromesally with short projecting broadly rounded setose area. Sternum IX membranous. Tergum X bilobed, each lobe divided into prominent rounded lightly sclerous basal region and less sclerous apical region with cercus at apex. Vaginal apparatus very short; ventrally with lightly sclerous inverted Y-shaped sclerite; dorsolaterally with pair of longitudinal sclerites, each ¯attened anteriorly, narrowed and slightly curled at apex; anteroventrally with lightly sclerous curled sclerite and mesal cuplike sclerite. Material examined. JAMAICA: 3Y, 1X paratypes, Portland Parish, Hardwar Gap, waterfall, 17.vii.1963 (Flint and Farr) (NMNH); 4Y, 9X paratypes, St Andrew Parish, Hardwar Gap, Dicks Pond Trail, 16.vii.1963 (Flint and Farr) (NMNH); 12Y, 6X paratypes, same location, 25.vii.1963 (O. Farr and R. Flint) (NMNH); 3X paratypes, same location, 26.ix.1963 (W. L. Peters and Farr) (NMNH). Chimarra (Otarrha) odonta, sp.n. (Figs 16A±G; 40A±C; 57) This species is unusual for the subgenus in having several characters that would seem to be primitive for the group, including a hindwing with Rs that is four-branched, R1 not fused to the subcosta and the possession of undivided anterior setal warts. The species is also primitive in having a very simple tergum X that possesses numerous sensilla and is completely divided mesally. Chimarra odonta can be readily identi®ed by the structure of the inferior appendages, which are short and possess a strongly mesally curved, blunt, toothlike process near the ventral apex. This is the only species in subgenus currently known from southeastern Brazil. Adult. Forewing length (male) 4.7±6.9 mm. Colour more or less uniformly brownish-fuscous, except head somewhat darker. Head ¯attened, postocular parietal sclerite prominent, elongate, in¯ated (extending below eyes). Palps only moderately elongate, third segment much longer than second and subequal to ®fth. Tarsal claws of male not enlarged, small, symmetrical. Setae of metatibiae prominent, bristly. Venation typical for Chimarra except M of hindwing 2-branched; R1 of

86 Roger J. Blahnik

Fig. 16. Chimarra odonta, male genitalia (G inset = dorsal view of phallotremal sclerite complex).

hindwing narrowly separated from subcosta; 3A of forewing looped to 1A; anal loop of hindwing small. Male genitalia (Fig. 16). Tergum VIII with broadly rounded, moderately projecting mesal extension. Abdominal segment IX with anteroventral margin somewhat extended, with distinct dorsomesal and ventromesal invagination; poster-

oventral process elongate, projecting, ventrally de¯exed, rounded apically. Tergum X simple in structure, completely divided mesally into 2 sclerotized lobes, each with numerous apical and apicoventral sensilla. Preanal appendages ¯attened, earlike. Inferior appendage, in lateral view, short, rounded basally, angular apicodorsally, with short truncate sclerotized ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

Systematics of Otarrha toothlike projection from ventromesal surface. Phallic apparatus with tubular phallotheca, expanded basodorsally; endotheca short, membranous. Phallotremal sclerite complex prominent, extending nearly length of phallotheca, composed of rod and ring structure and complex apical sclerite; ring broad and rod elongate, with 2 short detached spinelike structures preapically; apical sclerite composed of sclerous ring subtending apex of rod and 2 elongate ventral sclerites, each widened preapically and with abruptly upturned, acutely narrowed apex; mesally with detached ¯attened semicircular sclerite between ventral branches of apical sclerite. Female genitalia (Fig. 40). Segment VIII relatively short; anterior margin with obtusely angular apodemes, only weakly developed; posterior margin opposite apodemes also obtusely angular, with row of elongate setae extending ventrally along margin; ventrolaterally with deep cleft from posterior margin, extending more than halfway to anterior margin; ventromesally with pair of proximate sclerous raised setal warts, separated by pronounced mesal indentation. Sternum IX membranous, projecting and cupped apically, appearing as ventral extension from segment VIII. Tergum IX with moderately elongate apodemes from anteroventral margin; posteriorly setose with short truncate mesal projection. Tergum X bilobed, each lobe with sclerous basal collar and rounded setose cercus-bearing apical region. Vaginal apparatus relatively short, membranous apically and ventromesally, lightly sclerous ventrolaterally but with distinctly sclerous paired granular areas extending from ventral to apicolateral margin; anteroventrally with pair of short divergent foldlike projections. Material examined. Holotype, Y, BRAZIL: SaÄo Paulo, Est. Biol. Boraceia, 850 m, 3.ix.1977 (C. M. and O. S. Flint, Jr) (MZUSP). Paratypes, BRAZIL: Rio de Janeiro, 14Y, Parque National Tijuca, Represa dos Ciganos, 7.iv.1977 (C. M. and O. S. Flint, Jr) (MZUSP, NMNH); 2Y, Nova Friburgo, 950 m, 20.iv.1977 (C. M. and O. S. Flint, Jr) (NMNH); 9Y, km 54, 26 km E Nova Friburgo, 410 m, 19.iv.1977 (C. M. and O. S. Flint, Jr) (NMNH); 7Y, same location, 25.iv.1977 (C. M. and O. S. Flint, Jr) (NMNH); 1Y, km 17, 18 km S Teresopolis, 1180 m, 18±19.iv.1977 (C. M. and O. S. Flint, Jr) (NMNH); SaÄo Paulo, 3Y, same data as holotype (NMNH); 3Y, 1X, Rio Macac´ (®rst order), on RJ 116, km 65, 22°22.786¢S 42°33.945¢W, 933 m, 17.iii.1996 (Holzenthal, Rochetti, Oliveira) (UMSP); 1Y, Rio Macac´ (second order), on RJ 116, km 62, 22°23.201¢S 42°33.945¢W, 840 m, 17.iii.1996 (Holzenthal, Rochetti, Oliveira) (UMSP). Etymology. The name odonta, from the Greek for tooth, refers to the peculiar, blunt, incurved, toothlike process at the apex of the inferior appendage. Chimarra (Otarrha) parene, sp.n. (Figs 17A±D,G; 56) This species is similar to C. parilis and C. particeps. It differs in lacking serrations on the mesal surface of the inferior appendages. This species is most similar to C. parilis and shares with it the peculiar spine at the base of the ventral process of segment IX; it differs in the structure of the phallic apparatus, which in C. parene is less in¯ected, or bent, basally, ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

87

and less elongate and sclerous apically. This species is also similar to C. amazonia, but that species lacks the whiplike lateral and apical processes found on tergum X in C. parene, C. parilis and C. particeps. Adult. Forewing length (male) 4.6 mm. Holotype specimen incomplete, but general colour pale brown. Head somewhat ¯attened, postocular parietal sclerite moderately elongate (about as long as wide). Male genitalia (Fig. 17). Tergum VIII narrow mesally, without pronounced projections from posterior margin. Abdominal segment IX with anteroventral margin somewhat sinuously extended, invaginated dorsomesally and ventromesally; posteroventral process elongate, projecting, truncate apically, with rounded apicodorsal angle and pronounced spinelike projection basally; dorsomesally with very narrow elongate acuminate projection, only lightly sclerotized. Tergum X forming 2 sclerotized lateral lobes, each with acute dorsal projection possessing sensilla and short setae or setaform sensilla, and elongate lateral projection with 2 strongly sclerotized narrow acute whiplike projections, one apically and one preapically. Preanal appendages large, ¯attened, earlike. Inferior appendage, in lateral view, rounded basally, with strongly ¯exed elongate dorsal extension, narrowing apically; in dorsal or ventral views with distinct mesal curvature and bearing preapical tinelike projection and very small spine at base of tinelike projection. Phallic apparatus with short tubular phallotheca, expanded basodorsally and with acute apicoventral extension; endotheca short, membranous, with narrow elongate saclike projection basodorsally, and bearing 5 curved spines: 2 small, paired, apical; 2 stout, paired, preapical; and one short, stout, basal. Phallotremal sclerite complex composed of rod and ring structure, ring incomplete dorsally, rod short apically with small indistinct subtending sclerite. Female. Unknown. Material examined. Holotype, Y, PERU: Parene, El Campamiento, 16.vi.1920, Cornell University Expedition, lot Y 569 (CU). Etymology. This species is named parene for the type locality, the name penned on a label by Dr J. M. Kingsolver, who under Dr H. H. Ross, initiated but failed to complete, a study of genus Chimarra. The locality name, however, is very likely a misspelling of Perene. Chimarra (Otarrha) parilis, sp.n. (Figs 18A±D,G; 41A±C; 56) This species is most similar to C. parene, differing in several details, including the possession of minute serrations on the mesal surface of the inferior appendages, inferior appendages which are shorter and `blockier' (more squared basally) and a phallotheca that is more in¯ected basally and elongate apically. It differs from the following species in having serrations on the mesal margin of the inferior appendage which do not continue to the mesal tine and also in details of the shape of the inferior appendages and tergum X.

88 Roger J. Blahnik

Figs 17, 18. 17, Chimarra parene, male genitalia; 18, C. parilis, male genitalia.

ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

Systematics of Otarrha Adult. Forewing length (male) 5.2±5.8 mm (female) 5.2± 5.4 mm. Colour more or less uniformly dark brown; antennae indistinctly annulate intersegmentally with light brown; forewings with evident white marking at s crossvein. Head slightly ¯attened; postocular parietal sclerite moderately elongate. Male genitalia (Fig. 18). Tergum VIII narrow mesally, without pronounced projections from posterior margin. Abdominal segment IX with somewhat sinuous ventral extension of anterior margin; dorsomesal and ventromesal margins strongly invaginated; posteroventral process elongate, projecting, truncate apically, with acute apicodorsal angle and pronounced spinelike projection basally; dorsomesally with very narrow elongate acuminate projection, only lightly sclerotized. Tergum X forming 2 sclerotized lateral lobes, each with acute dorsal projection possessing sensilla and short setae or setaform sensilla, and elongate lateral projection with 2 strongly sclerotized narrow acute whiplike projections, one apically and one preapically. Preanal appendages large, ¯attened, earlike. Inferior appendage, in lateral view, rounded basally, with strongly ¯exed elongate dorsal extension, narrowing apically; in dorsal or ventral view with distinct mesal curvature and bearing preapical tinelike projection and second very small spine at base of tinelike projection; basomesally with very small serrations. Phallic apparatus with short tubular phallotheca, expanded basodorsally, rather strongly in¯ected basally, apically elongate and lightly sclerous; endotheca membranous, with narrow elongate saclike projection basodorsally, and bearing 5 curved spines: 2 very small, paired and apical; 2 stout, paired, preapical; and one short, stout and basal. Phallotremal sclerite complex composed of rod and ring structure, rod short with small indistinct subtending sclerite apically. Female genitalia (Fig. 41). Segment VIII short; anterolateral margin more or less linear, with angular lateral projection at point of apodeme but apodeme undeveloped; posterior margin with elongate setae mid-laterally, bordered ventrally by marginal cleft; posterolateral cleft moderately deep, extending about halfway to anterior margin; posterior margin, in ventral view with deep mesal emargination, laterally with about 4 elongate setae on either side of emargination. Tergum IX short, with relatively short, bent apodemes from anteroventral margin; posteromesally with short narrow rounded projection. Sternum IX membranous. Tergum X bilobed, each lobe divided into prominent rounded sclerous basal region and less sclerous apical region with cercus at apex. Vaginal apparatus with elongate conical apicodorsal sclerites emerging from sclerous region formed by dorsum of vaginal apparatus and ventral margin of rectum; rectum opening apically; vaginal apparatus sclerous apicoventrally, widening abruptly to rounded sclerous region extending ventrally and laterally, narrowed and widening again anteriorly to form pair of enlarged lobes; anteromesally with cuplike sclerite emerging between enlarged lateral lobes. Material examined. Holotype, Y, PERU: Madre de Dios, Amazonia Lodge, Toma del Agua (stream), 12°52.22¢S 71°22.56¢W, 414 m, 29.vi.1993 (R. Blahnik and M. Pescador) (MJP). Paratypes, PERU: Madre de Dios, 8Y, 3X, same data as holotype (MJP, NMNH, UMSP). ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

89

Etymology. This species is named parilis from a Latin word meaning equal or like, in reference to its similarity to C. parene. Chimarra (Otarrha) particeps, sp.n. (Figs 19A±D,G; 42A±C; 56) This species is similar to C. parene and C. parilis, but differs in a number of details, including the possession of inferior appendages with mesal serrations extending from the base to the mesal tine, and the possession of a tergum X which, in lateral view, appears more bulbous apically. Unlike C. parene and C. parilis, this species lacks a spine at the base of the ventral process of segment IX. Additionally, the inferior appendages are slightly more elongate, more rounded basally and have apices which are wider than C. parilis. The female vaginal apparatus is also different in structure from that of C. parilis, additionally supporting speci®c status for this taxon. Adult. Forewing length (male) 4.4±4.7 mm (female) 4.8± 5.2 mm. Colour (in alcohol) uniformly brown. Head ¯attened; postocular parietal sclerite moderately elongate. Male genitalia (Fig. 19). Tergum VIII very narrow mesally, with 2 short acute dorsolateral projections from posterior margin. Abdominal segment IX with anteroventral margin sinuously extended, strongly invaginated dorsomesally and ventromesally; posteroventral process elongate, projecting, blunt apically, widest basally and apically; dorsomesally with lightly sclerotized and very elongate narrow acuminate projection. Tergum X forming 2 sclerotized lateral lobes, each with acute dorsal projection possessing sensilla and short setae or setaform sensilla, and elongate lateral projection with 2 strongly sclerotized narrow acute whiplike projections, one apically and one preapically; apex of lateral lobe, in lateral view, more bulbous than C. parene. Preanal appendages large, ¯attened, earlike. Inferior appendage, in lateral view, narrow basally and dorsally ¯exed, with elongate apically narrowed dorsal extension; in dorsal or ventral view with distinct mesal curvature and bearing short preapical tinelike projection; mesal surface proximal to tine distinctly serrate. Phallic apparatus with short tubular phallotheca, expanded basodorsally and with acute apicoventral extension; endotheca short, membranous, with 2 pairs of curved spines: 2 short and narrow and 2 larger and stouter. Phallotremal sclerite complex indistinct, apparently composed of paired ventral rods and indistinct or incomplete ringlike structure. Female genitalia (Fig. 42). Segment VIII short; anterolateral margin more or less linear, with angular lateral projection at point of apodeme but apodeme undeveloped; posterior margin with elongate setae mid-laterally, bordered ventrally by marginal cleft; posterolateral cleft moderately deep, extending about halfway to anterior margin; posteroventral margin with shallow mesal emargination, laterally with several elongate setae on either side of emargination. Tergum IX short, with relatively short bent apodemes from anteroventral margin; posteromesally with short narrow rounded projection. Sternum IX membranous. Tergum X bilobed, each lobe divided into prominent rounded sclerous

90 Roger J. Blahnik

Fig. 19. Chimarra particeps, male genitalia.

basal region and less sclerous apical region with cercus at apex. Vaginal apparatus with stout conical apicodorsal sclerites, emerging from sclerous region formed by dorsum of vaginal apparatus and ventral margin of rectum; rectum opening apically; vaginal apparatus sclerous apicoventrally, widening abruptly to rounded sclerous region extending ventrally and laterally, narrowed slightly and widening again to form pair of short ventrolateral lobes; vaginal apparatus anteromesally with elongate narrowed membranous extension and de¯exed cuplike sclerite. Material examined. Holotype, Y (in alcohol), PERU: Huanuco, Tingo Maria, 672 m, 1±6.ii.1980 (J. B. Heppner) (NMNH). Paratypes, PERU: Huanuco, 24Y, 5X (in alcohol), same data as holotype (MSP, NMNH, UMSP).

Etymology. This species is named particeps from a Latin word meaning comrade, partner or sharer, in reference to the close relationship or partnership of this species with C. parene and C. parilis. Chimarra (Otarrha) patosa Ross (Figs 20A±D,G,I; 43A±C; 57) Chimarra patosa Ross, 1956: 71, Figs 109A,B, 147±149; Y, Peru (INHS); Fischer, 1971: 215. This species is most similar to C. peruana, sharing with it the very elongate processes from the posterior margin of ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

Systematics of Otarrha tergum VIII. It differs, however, in having the processes converging mesally, and by lacking a distinct clublike brush of setae at the apices of the processes. The inferior appendages are also somewhat different in structure. Adult. Forewing length (male) 4.8±5.2 mm (female) 5.2± 5.8 mm. Colour more or less uniformly dark brown; antennae indistinctly annulate with light brown. Head slightly ¯attened; postocular parietal sclerite moderately elongate. Male genitalia (Fig. 20). Tergum VIII very narrow mesally, with 2 elongate narrow strongly converging projections from posterior margin, each with numerous short spines. Abdominal segment IX with anterolateral margin nearly linear for twothirds of length but abruptly angularly narrowed near ventral margin, dorsomesal and ventromesal margins strongly invaginated; posteroventral process relatively elongate projecting, rounded apically. Tergum X (or posterior margin of tergum IX?) with elongate acuminate upturned mesal projection; laterally forming 2 sclerotized lobes; lateral lobes deeply cleft, each with elongate acute ventral projection and short dorsal projection with numerous apical sensilla or short setae or both. Preanal appendages relatively small, ¯attened, earlike. Inferior appendage, in lateral view, ¯exed at base, with elongate dorsal extension, appearing narrowed apically due to mesal curvature; in dorsal or ventral view with distinct mesal curvature, preapically with short tinelike projection, apex of inferior appendage blunt. Phallic apparatus with short phallotheca, strongly ¯exed basally and only weakly developed dorsally; endotheca membranous, with 2 elongate curved dorsal spines. Phallotremal sclerite complex composed of rod and ring structure; ring with dorsal extension, rod relatively short; apically with indistinct subtending sclerite. Female genitalia (Fig. 43). Segment VIII short, with projecting ventral margin; anterolateral margin more or less linear, with short lateral apodemes; posterior margin with elongate setae mid-laterally, bordered ventrally by deep marginal cleft, extending nearly to anterior margin; in ventral view with nearly quadrate mesal extension, posterior margin linear, with about 4 elongate setae on either side of midline and numerous smaller setae. Tergum IX short with short bent apodemes from anteroventral margin; in¯ated laterally; posteromesally with short broadly rounded projection. Sternum IX membranous. Tergum X bilobed, each lobe divided into prominent rounded sclerous basal region and less sclerous apical region with cercus at apex. Vaginal apparatus largely membranous, with lightly sclerous Y-shaped sclerite extending from dorsum of vaginal apparatus and branching apically; apex of branches with ®ngerlike ventral projection; rectum opening preapically, with dorsal pouch at opening; vaginal apparatus anteriorly with de¯exed cuplike sclerite. Material examined. PERU: Cuzco, 1Y, 1X, Paucartambo, Puente San Pedro, c. 50 km NW Pilcopata, km 152, 13°09¢S 71°26¢W, 1430 m, 30±31.viii.1989 (N. Adams et al.) (NMNH); 1Y, 1X, same location, 24.vi.1983 (R. Blahnik and M. Pescador) (UMSP); 2Y, Paucartambo to Pilcopata, Quebrada QuitacalzoÂn at Puente QuitacalzoÂn, 13°01.57¢S 71°29.97¢W, 1050 m, 25±27.vi.1983 (R. Blahnik and M Pescador) (MJP, NMNH); Madre de Dios, 1X, Amazonia Lodge, Toma del ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

91

Agua (stream), 12°52.22¢S 71°22.56¢W, 415 m, 29.vi.1983 (R. Blahnik and M. Pescador) (MJP). Chimarra (Otarrha) peruana, sp.n. (Figs 21A±D,G,I; 57) This species is similar to C. patosa, but differs in several details of the male genitalia. The most notable differences include the shape of the inferior appendages and the possession of dorsal processes of tergum VIII which are less strongly converging than those of C. patosa and terminate in a club of elongate setae. Adult. Forewing length (male) 4.9±5.1 mm. Colour brown. Head ¯attened, postocular parietal sclerite moderately elongate. Male genitalia (Fig. 21). Tergum VIII very narrow mesally, with 2 elongate narrow projections from posterior margin, each with scattered short spines and terminating in club of elongate setae. Abdominal segment IX with anterolateral margin nearly linear, rounded ventrally; dorsomesal and ventromesal margins strongly invaginated; posteroventral process relatively elongate, projecting, rounded apically. Tergum X (or posterior margin of tergum IX?) with elongate acuminate upturned mesal projection; laterally forming 2 sclerotized lobes; lateral lobes deeply cleft, each with elongate acute ventral projection and short dorsal projection with numerous sensilla or short setae. Preanal appendages relatively small, ¯attened, earlike. Inferior appendage with distinct mesal curvature, rounded apex and preapical tinelike projection on mesal surface; in lateral view with less angular basal in¯ection than C. patosa and with elongate curved dorsal extension, apparently narrowing apically (due to mesal curvature). Phallic apparatus with short phallotheca, strongly ¯exed basally and only weakly developed dorsally; endotheca membranous, with 2 elongate curved dorsal spines. Phallotremal sclerite complex composed of rod and ring structure; ring with dorsal extension, rod relatively short; apically with indistinct subtending sclerite. Female Unknown. Material examined. Holotype, Y, PERU: Aina, 1400 m, am back gestreift, 6.vi.1936, Heidelberg-SuÈd Peru Sammelreise 1936, Eing. Nr. 1937 (ZMUH). Paratypes, PERU: 3Y, same data as holotype (ZMUH). Etymology. This species is named peruana from the Spanish word for Peruvian, in reference to the country of origin of the holotype. Chimarra (Otarrha) phthanorossi, sp.n. (Figs 22A±D, G,H,I; 44A±C; 57) This species is most similar to C. rossi, but differs signi®cantly in having phallic spines which are much less elongate. It also differs in having inferior appendages with apices which are not bi®d. In a general sense, C. phthanorossi is also similar to C. retrorsa and C. septifera, but differs from these species in

92 Roger J. Blahnik

Figs 20, 21. 20, Chimarra patosa, male genitalia; 21, C. peruana, male genitalia.

ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

Systematics of Otarrha

93

Fig. 22. Chimarra phthanorossi, male genitalia.

lacking paired acuminate apices on the mesal lobe of tergum X. Adult. Forewing length (male) 4.8±5.0 mm (female) 4.8± 5.0 mm. Colour more or less uniformly dark brown; antennae ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

indistinctly annulate with light brown. Head distinctly ¯attened; postocular parietal sclerite moderately elongate. Male genitalia (Fig. 22). Tergum VIII with acute paired mesally directed projections from posterior margin. Abdominal

94 Roger J. Blahnik

Fig. 23. Chimarra puertoricensis, male genitalia (G inset = dorsal view of phallotremal sclerite complex).

segment IX with anterolateral margin nearly linear, abruptly narrowed dorsally and ventrally; dorsomesal and ventromesal margins strongly invaginated; posteroventral process elongate, projecting, rounded apically. Posterior margin of tergum IX with very short rounded mesal process. Tergum X forming paired lateral lobes, each divided into short lightly sclerotized apically rounded ventrolateral lobe and an elongate sclerotized rodlike dorsolateral lobe; rodlike lobe with minute spines basally on mesal margin and scattered sensilla laterally. Preanal appendages large, ¯attened, earlike. Inferior appendage elongate, curved, enlarged basally, strongly mesally ¯exed in apical third; in dorsal or ventral view with apex

¯attened, slightly enlarged preapically, abruptly narrowing to acute mesally directed apex. Phallic apparatus with phallotheca short, expanded basodorsally; endotheca short, membranous. Phallotremal sclerite complex composed of rod and ring structure and associated sclerites; dorsally with elongate symmetrically paired spines, approximately equaling phallotheca in length; apically with elongate structure subtending rod, lightly sclerous basally, apically forming pair of curved irregular sclerotized rods. Female genitalia (Fig. 44). Segment VIII relatively elongate, with protruding posteroventral margin; anterolateral margin more or less linear but with ventral margin ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

Systematics of Otarrha abruptly recessed; posterior margin with elongate setae midlaterally, bordered ventrally by shallow cleft; ventrolaterally with indistinct suture line extending from lateral cleft to posterior margin; tapering posteriorly in ventral view, posterior margin linear with about 4 prominent setae on either side of midline and numerous smaller setae. Tergum IX moderately elongate, with short apodemes from anteroventral margin; posteromesally with elongate tapering projecting setose area, wide at base and narrow at apex. Sternum IX membranous. Tergum X bilobed, each lobe divided into small rounded sclerous basal region and less sclerous elongate apical region with cercus at apex. Vaginal apparatus largely membranous, apicodorsally with sclerite, extending from ventral wall of rectum and widening apically; rectum opening apically; vaginal apparatus midventrally with elongate cupped sclerite; anteriorly widened to form projecting membranous pouch. Material examined. Holotype, Y, COLOMBIA: ChocoÂ, km 130, 86 km E Quibdo, 17.ii.1983 (O. S. Flint, Jr) (NMNH). Paratypes, 16Y, 2X, same location as holotype (NMNH); 22Y, 2X (pinned), 22Y, 5X (in alcohol), 52 km E Quibdo, 20.ii.1983 (O. S. Flint, Jr) (NMNH, UMSP). Etymology. This species is named phthanorossi from the Greek word phthano, meaning to do ®rst or anticipate, in reference to the relationship of this species to C. rossi Bueno-Soria, which it resembles in all character states but in a much less exaggerated and apomorphic form. Chimarra (Otarrha) puertoricensis Flint (Figs 23A± D,F,G; 45A±C; 56) Chimarra puertoricensis Flint, 1964: 23±24, Fig. 4B±C,F,I,K, L; Y, Puerto Rico (NMNH). The distinctive shape of the inferior appendages, which are relatively elongate, narrow at the base and wide at the apex, will distinguish this species from any other species in the subgenus. The deeply incised lateral lobes of tergum X are also unique among the Antillean species. Adult. Forewing length (male) 4.5 mm (female) 4.9 mm. Colour more or less uniformly dark brown; antennae indistinctly annulate with light brown. Head ¯attened; postocular parietal sclerite moderately elongate. Male genitalia (Fig. 23). Tergum VIII with short mesal and 2 dorsolateral projections from posterior margin. Abdominal segment IX with anterolateral margin broadly rounded; dorsomesal and ventromesal margins strongly invaginated; posteroventral process elongate, projecting, slightly curved, subacute apically; tergum IX with short rounded posteromesal process. Tergum X completely divided mesally into 2 divided sclerotized lobes, each with elongate narrow lateral cleft; apex of dorsal portion of lateral lobe slightly expanded, upturned, acute, ventral portion acute apically, both portions with several sensilla apically; lateral lobes basally with 2 stout spines. Preanal appendages large, ¯attened, earlike. Inferior appendage with distinct mesal curvature; in lateral view, moderately ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

95

elongate, narrow basally, gradually expanded to wide truncate apex; ventral angle of apex acute and sclerotized. Phallic apparatus with tubular phallotheca, expanded basodorsally; endotheca short, membranous. Phallotremal sclerite complex prominent, more than half length of phallotheca, composed of distinct rod and ring structure, ring with dorsal extension, rod with preapical lateral extensions. Female genitalia (Fig. 45). Segment VIII short; anterolateral margin more or less linear, with well developed lateral apodemes; posterior margin with elongate setae midlaterally, subtended ventrally by marginal cleft; posterolateral cleft moderately deep, extending about halfway to anterior margin; posterior margin, in ventral view, rounded mesally, with short membranous margin and group of elongate setae on either side of midline, laterally with sclerous projections. Tergum IX short, with moderately elongate apodemes from anteroventral margin; posterodorsal margin with mesal emargination, forming short rounded projections on either side of midline. Sternum IX membranous. Tergum X bilobed, each lobe divided into lightly sclerous basal region and less sclerous apical region with cercus at apex. Vaginal apparatus short, lightly sclerous, with short conical projections apicodorsally, emerging from union of ventral surface of rectum and dorsal surface of vaginal apparatus; conical projections continuously sclerous basally with dorsal surface of vaginal apparatus; rectum opening apically; vaginal apparatus posteroventrally becoming lightly sclerous and forming ventral collar preapically, anteriorly membranous with de¯exed cuplike sclerite. Material examined. PUERTO RICO: 1Y, 2X paratypes, Maricao, ®sh hatchery, 8±11.viii.1961 (Flint and Spangler) (NMNH); 1Y, same location, 23.xii.1962 (P. and P. Spangler) (NMNH). Chimarra (Otarrha) quadrifurcata Botosaneanu (Figs 24A±C,G,H,I; 46A±C; 56) Chimarra quadrifurcata Botosaneanu, 1994b: 43±47, Figs 29± 36; Y, Guadeloupe (ZMUA). This is a distinctive species, most similar to C. dominicana and C. diannae, both of which also possess inferior appendages that are strongly retrorsely curved and acuminate. It differs by having a distinctive, ¯attened preapical lobe on the inferior appendages, a pair of elongate phallic spines and a more elongate mesal lobe of tergum X (or apical extension of tergum IX). Adult. Forewing length (male) 5.7 mm (female) 5.1 mm. Colour (in alcohol) brown. Head distinctly ¯attened; postocular parietal sclerite elongate. Male genitalia (Fig. 24). Tergum VIII with short acute paired projections from posterior margin. Abdominal segment IX with anterolateral margin nearly linear or slightly sinuate, much narrowed dorsally and ventrally; anterodorsal and anteroventral margins strongly invaginated mesally; posterior

96 Roger J. Blahnik

Fig. 24. Chimarra quadrifurcata, male genitalia.

margin with pronounced expansion at level of inferior appendages; ventral process elongate, projecting, rounded apically, narrowed basally. Tergum IX dorsomesally with moderately elongate, lightly sclerotized lobe from posterior margin, wide basally and narrowed apically. Tergum X forming 2 sclerotized lateral lobes rather widely separated by

mesal lobe of segment IX; each lobe partially cleft longitudinally into dorsal and ventral projections; dorsal projection narrow and more elongate, with several sensilla at apex, ventral projection short, slightly curved, acute apically. Preanal appendages large, ¯attened, earlike. Inferior appendage, in lateral view elongate with pronounced mesal curvature; in ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

Systematics of Otarrha

97

Fig. 25. Chimarra redonda, male genitalia.

dorsal view narrowed apically to elongate acuminate retrorsely directed tip; preapically with rounded ¯attened lobe, appearing apical in lateral view. Phallic apparatus with relatively short phallotheca, expanded basodorsally; endotheca short, membranous. Phallotremal sclerite complex prominent, longer than phallotheca, composed of rod and ring structure and associated sclerites; dorsally with paired elongate symmetrical spines, ventrally with elongate lightly sclerous structure subtending rod, apically with 2 short sclerotized spines. Female genitalia (Fig. 46). Segment VIII elongate ventrally, anterior and posterior margins both extending; anterolateral margin sinuous, with angular projection at normal position of apodemes, apodemes undeveloped; posterior margin with ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

rounded projection mid-laterally, bearing elongate setae on margin, bordered ventrally by cleft from posterior margin; posterolateral cleft moderately indentate, less than halfway to anterior margin; ventrolaterally with distinct suture line extending from posterior cleft to anterior margin; ventral surface with distinct shallow broadly sinuous indentation; posterior margin in ventral view with acutely projecting apex, laterally with about 4 prominent setae on either side of midline and numerous smaller setae on venter of segment. Intersegmentally with prominent receptacle for inferior appendage between segments VII and VIII. Tergum IX short, with moderately elongate apodemes from anteroventral margin, posteromesally with elongate narrow projecting setose

98 Roger J. Blahnik area. Sternum IX membranous, intersegmental membrane between terga IX and X with shallow pocketlike invagination; Tergum X bilobed, each lobe divided into prominent rounded sclerous basal region and less sclerous apical region with cercus at apex. Vaginal apparatus largely membranous; apex with collar, sclerous dorsally, forming paired ¯attened ventral projections, open ventrally; rectum opening apically; vaginal apparatus anteroventrally with elongate cuplike receptacle located before apex of anterior margin. Material examined. GUADELOUPE: Basse Terre, 1Y, 1X paratypes, Riviere St Louis (Hauts de Macouba), 8.iv.1991 (L. Botosaneanu) (ZMUA). Chimarra (Otarrha) redonda, sp.n. (Figs 25A±D,F,G; 47A±C; 56) Botosaneanu, 1996: 11 (in reference to undescribed species). This species is readily identi®ed by the structure of the inferior appendages, which are ovate in lateral view and have a pronounced sclerous tooth preapically on the mesal surface. Adult. Forewing length (male) 4.3±4.6 mm (female) 4.4± 5.0 mm. Colour more or less uniformly brown, antennae indistinctly annulate with light brown. Head somewhat ¯attened, postocular parietal sclerite moderately elongate. Male genitalia (Fig. 25). Tergum VIII with small widely separated projections from posterior margin. Abdominal segment IX with anterolateral margin broadly rounded; dorsomesal and ventromesal margins strongly invaginated; posteroventral process elongate, projecting, narrow basally, rounded apically. Tergum X simple in structure, completely divided mesally into 2 sclerotized lobes, each with numerous lateral sensilla and small spinelike projections. Preanal appendages large, ¯attened, earlike. Inferior appendage, in lateral view, ovate; preapically with distinct sclerotized projection from dorsomesal surface. Phallic apparatus with tubular phallotheca, expanded basodorsally; endotheca short, membranous. Phallotremal sclerite complex prominent, about half length of phallotheca, composed of rod and ring structure and curved apical sclerite. Female genitalia (Fig. 47). Segment VIII short; anterolateral margin more or less linear with rounded projection at normal position of apodemes, apodemes undeveloped; posterior margin with elongate setae mid-laterally, subtended ventrally by deep marginal cleft; posterolateral cleft extending more than halfway to anterior margin; posteroventral margin rounded mesally, with prominent lightly sclerous margin with lateral projections; apex of rounded mesal projection with group of setae, not separated mesally. Tergum IX short, with moderately elongate apodemes from anteroventral margin; posterodorsal margin with short broad truncate setose projection. Sternum IX membranous. Tergum X bilobed, each lobe divided into lightly sclerous basal region with mesal projection and less sclerous bulbous apical region with cercus at apex. Vaginal apparatus short and lightly sclerous but with short conical sclerous projections apicodorsally, emerging from

dorsal surface of vaginal apparatus; conical projections continuously sclerous basally with dorsal surface of vaginal apparatus; rectum opening apically; vaginal apparatus anteroventrally becoming lightly sclerous and forming ventral collar preapically, anterior apex membranous, with de¯exed cuplike sclerite. Material examined. Holotype, Y, DOMINICAN REPUBLIC: La Palma, 12 km E of El RõÂo, 2±13.vi.1969 (Flint and Gomez) (NMNH). Paratypes, DOMINICAN REPUBLIC: 16Y, 8X, same data as holotype (NMNH, UMSP); 10Y, 2X, Cordillera Central, Parque Nacional Armando Bermudez, Arroyo Manuel Estrella, 25.iv.1995 (L. Botosaneanu) (ZMUA). Etymology. This species is named redonda from the Spanish word for round, in reference to its the rounded inferior appendages. Chimarra (Otarrha) retrorsa Flint (Figs 26A±D,G; 57) Chimarra (Chimarra) retrorsa Flint, 1974: 26, Figs 30±32; Y, Suriname (RNHL). This species is similar in general form to several other species in having a tergum X with rodlike dorsal lobes, and inferior appendages which are elongate, apically ¯attened and acute, and have an angular, mesal in¯ection. It is probably most similar to C. septifera, which also has the mesal lobe of tergum X membranous mesally (apparently) and possesses narrowly separated, sclerous lateral structures, each terminating in an acuminate point. Chimarra retrorsa is distinguished from C. septifera by having the apices of these structures whiplike and retrorsely directed. Adult. Forewing length (male) 4.0 mm. Colour (in alcohol) yellowish brown. Head somewhat ¯attened, postocular parietal sclerite moderately elongate. Male genitalia (Fig. 26). Tergum VIII with 2 short acute projections from posterior margin on each side of midline. Abdominal segment IX with anterolateral margin nearly linear, rounded dorsally and ventrally; dorsomesal and ventromesal margins strongly invaginated; posteroventral process elongate, projecting, rounded apically. Tergum X (or posterior margin of tergum IX?) with elongate mesal process de¯exed basally and with 2 dorsally recurrent whiplike processes at apex; tergum X dorsolaterally forming 2 sclerotized lobes, separated basally, each elongate, narrow and with small spinelike projections. Preanal appendages large, ¯attened, earlike. Inferior appendage elongate, angular basally and strongly mesally ¯exed in apical third; in dorsal or ventral view with mesal curvature, strongly mesally ¯exed in apical third, apex acute. Phallic apparatus with short tubular phallotheca, distinctly expanded basally; endotheca short, membranous. Phallotremal sclerite complex large, elaborate, composed of rod and ring structure; ring dorsally with 2 elongate narrow spinelike extensions, equaling length of phallic apparatus; rod distinct with elongate lightly sclerotized apicoventral sclerite forming 2 acute apical spines. Female. Unknown. ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

Systematics of Otarrha

99

Fig. 26. Chimarra retrorsa, male genitalia.

Material examined. Holotype, Y, SURINAME: Brownsberg, 400 m, 15.ix.1938 (Geijskes) (RNHL); 1Y paratype, Brownsberg, 10.viii.1958 (Geijskes) (NMNH). Chimarra (Otarrha) rossi Bueno-Soria (Figs 27A±D, G,H; 48A±D; 57) Chimarra (Chimarra) rossi Bueno-Soria, 1985: 22, Figs 12,13; Y, Costa Rica (IBUNAM); Holzenthal, 1988: 56; Blahnik & Holzenthal, 1992: 410. This species is unique and easily recognized by the extreme development of the male genitalia. In general features, the male genitalia resemble those of C. phthanorossi. However, the apicodorsal spines of the phallic apparatus are much more elongate and the apicoventral sclerite is more strongly ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

sclerotized and highly developed. In addition, the apices of the inferior appendages are bi®d rather than simple and acute, and tergum X forms lateral lobes which are very heavily sclerotized. Adult. Forewing length (male) 4.4±5.2 mm (female) 4.6± 6.2 mm. Colour more or less uniformly dark brown; antennae indistinctly annulate with light brown. Head ¯attened, postocular parietal sclerite moderately elongate. Male genitalia (Fig. 27). Tergum VIII with 2 short, acute projections from posterior margin. Abdominal segment IX with anterolateral margin nearly linear, angularly narrowed dorsally and ventrally; dorsomesal and ventromesal margins strongly invaginated; posteroventral process elongate, projecting, rounded apically. Tergum X (or posterior margin of tergum IX?) with short mesal process, continuous with 2 simple moderately elongate lightly sclerotized lateral lobes; tergum X dorsally forming 2 curved elongate narrow highly

100 Roger J. Blahnik

Fig. 27. Chimarra rossi, male genitalia. ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

Systematics of Otarrha sclerotized lobes, lobes separated basally, each with small spines on mesal and apical margins, basoventrally with small granular projections. Preanal appendages large, ¯attened, earlike. Inferior appendage elongate, curved, enlarged basally, rather strongly mesally ¯exed in apical third; in dorsal or ventral views with mesal curvature, mesally ¯exed in apical third, apex bi®d due to angular mesal notch, dorsal fork of bi®d apex slightly more elongate. Phallic apparatus very complex and conspicuous due to extreme modi®cation of phallotremal sclerite complex; phallotheca relatively short, basally expanded; endotheca membranous, saclike apically. Phallotremal sclerite complex large, elaborate, composed of rod and ring structure and highly modi®ed apical armature; dorsally with 2 very elongate narrow spinelike extensions; rod distinct, rather elongate, subtended by complex apical sclerite; apical sclerite modi®ed apically into pair of curved sclerotized rods, enlarged preapically, with extreme apex mesally curved. Female genitalia (Fig. 48). Segment VIII relatively elongate, with protruding posteroventral margin; anterolateral margin undulate; posterior margin with elongate setae mid-laterally on rounded projection, bordered ventrally by shallow cleft, extending less than halfway to anterior margin; ventrolaterally with distinct suture line extending from lateral cleft to posterior margin; posteroventral margin nearly linear, with about 4 prominent setae on either side of midline and numerous smaller setae. Tergum IX of moderate length, with moderately elongate apodemes from anteroventral margin; posteromesally with elongate tapering setose projection. Sternum IX membranous. Tergum X bilobed, each lobe divided into prominent sclerous basal region and less sclerous elongate apical region with cercus at apex. Vaginal apparatus largely membranous, apicodorsally with distinct sclerite extending from ventral surface of rectum; apex of apical sclerite mesally emarginate and with acute ventral projections, which nearly meet ventrally; rectum opening apically or near apically; vaginal apparatus ventrally with cuplike, posterior sclerite; anteriorly with dorsal hoodlike projection and elongate sacklike structure. Material examined. COSTA RICA: 3Y, 1X, Tuis, 18.vi.1967 (Flint and Ortiz) (NMNH); 1X, 3 mi. W Turrialba, 18±21.vi.1967 (Flint and Ortiz) (NMNH); Alajuela, 6Y, Reserva Forestal San RamoÂn, RõÂo San Lorencito and tribs, 10.216°N 84.607°W, 980 m, 13± 16.vi.1988 (C. M. and O. S. Flint, Holzenthal) (NMNH); 4Y, 27X, same location, 1±4.v.1990 (Holzenthal and Blahnik) (UMSP); 24Y, 13X, same location, 28± 30.vii.1990 (Holzenthal, Blahnik, MunÄoz) (UMSP); 3Y, 2X, same location, 2±4.vii.1986 (Holzenthal, Heyn, Armitage) (UMSP); 39Y, 23X, same location, 6± 10.iii.1991 (Holzenthal, MunÄoz, Huisman) (UMSP); 7Y, same location, 30.iii.-1.iv.1987 (Holzenthal, Hamilton, Heyn) (UMSP); 4Y, 3X, Quebrada Latas, 8.9 km NE Bajos del Toro, 10.269°N 84.260°W, 1030 m, 6.ix.1990 (Holzenthal, Blahnik, Huisman) (UMSP); 1Y, Cerro Campana, c. 6 km NW Dos RõÂos, 10.9°N 85.4°W, 640 m, 15±16.iii.1986 (Holzenthal and Fasth) (UMSP); 41Y, Cerro ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

101

Campana, RõÂo Bochinche trib., 6 km NW Dos RõÂos, 10.945°N 85.413°W, 600 m, 22±23.vii.1987 (Holzenthal, Morse, Clausen) (UMSP); 1Y, Quebrada Honda, 5.4 km S Crucero, 10.311°N 84.245°W, 650 m, 12.ii.1992 (Holzenthal, MunÄoz, Kjer) (UMSP); 2Y, P. N. [Parque Nacional] RincoÂn de la Vieja, Quebrada ProvisioÂn, 10.769°N 85.281°W, 810 m, 4.iii.1986 (Holzenthal and Fasth) (UMSP); Cartago, 1Y, 1X, La Suiza, 17.vi.1967 (Flint and Ortiz) (NMNH); 4Y, 1X, RõÂo Chitaria, route 10, 10 km NW RõÂo ReventazoÂn, 9.920°N 83.604°W, 740 m, 26.iii.1991 (Holzenthal, MunÄoz, Huisman) (UMSP); Guanacaste. 2Y, P. N. Guanacaste, EstacioÂn Pitilla, RõÂo OrosõÂ, 10.991°N 85.428°W, 700 m, 19±20.vi.1988 (C. M. and O. S. Flint, Holzenthal) (NMNH); 7Y, 5X, same location, 22±25.v.1990 (Holzenthal and Blahnik) (UMSP); 2Y, 1X, P. N. Guanacaste, EstacioÂn Maritza, RõÂo Tempisquito, 10.958°N 85.497°W, 550 m, 30±31.viii.1990 (Huisman, Blahnik, Quesada) (UMSP); 1Y, P. N. Guanacaste, c. 0.7 km N EstacioÂn Maritza, 10.96°N 85.50°W, 550 m, 31.viii.1990 (Huisman and Quesada) (NMNH); 5Y, Quebrada Garcia, 10.6 km ENE Quebrada Grande, 10.862°N 85.428°W, 470 m, 8.iii.1986 (Holzenthal & Fasth) (UMSP); 6Y, 1X, RõÂo Los Ahogados, 11.3 km ENE Quebrada Grande, 10.865°N 85.423°W, 470 m, 7.iii.1986 (Holzenthal and Fasth) (UMSP); 1Y, same location, 26.vi.1986 (Holzenthal, Heyn, Armitage) (UMSP); 8Y, RõÂo Gongora (sulphur mine), 4 km NE Quebrada Grande, 10.887°N 85.470°W, 590 m, 21.vii.1987 (Holzenthal, Morse, Clausen) (UMSP); 1Y, P. N. RincoÂn de la Vieja, Quebrada Zopilote, 10.765°N 85.309°W, el 785 m, 3.iii.1986 (Holzenthal) (UMSP); 9Y, RõÂo Mena, 4.2 km W Santa Cecilia, 11.059°N 85.448°W, 260 m, 11.iii.1986 (Holzenthal and Fasth) (UMSP); 2Y, RõÂo Aguacate, 0.5 km E Aguacate, 10.565°N 84.939°W, 590 m, 16.ii.1992 (Holzenthal, MunÄoz, Kjer) (UMSP); Heredia, 12Y, 9X, P. N. Braulio Carrillo, EstacioÂn El Ceibo, RõÂo Peje, 10.327°N 84.078°W, 480 m, 29±31.v.1990 (Holzenthal, Blahnik, MunÄoz) (UMSP); 48Y, 28X, P. N. Braulio Carrillo, EstacioÂn Magsasay, RõÂo Peje, 10.402°N 84.050°W, 130 m, 25±26.viii.1990 (Holzenthal, Blahnik, Huisman) (UMSP); LimoÂn, 4Y, 3X, P. N. Braulio Carrillo, Quebrada Gonzalez, 10.160°N 83.939°W, 12± 14.v.1990 (Holzenthal and Blahnik) (UMSP); 4Y, Reserva Biol. Hitoy Cerere, RõÂo Cerere (UMSP); 9Y, 2X, Reserva Biol. Barbilla, RõÂo Dantas, 15 km S Pacuarito, 9.994°N 83.443°W, 300 m, 27±30.i.1992 (Holzenthal, MunÄoz, Kjer) (UMSP); 1Y, same location, 1.ii.1992 (Holzenthal, MunÄoz, Kjer) (UMSP); Puntarenas, 3Y, 4X, Reserva Biol. Carara, Quebrada Bonita, 9.775°N 84.605°W, el 35 m, 18± 20.v.1990 (Holzenthal and Blahnik) (UMSP); 12Y, 3X, same location, 11.iii.1991 (Holzenthal, MunÄoz, Huisman) (UMSP); 6Y, 1X, RõÂo Jaba at rock quarry, 1.4 km W Las Cruces, 8.79°N 82.97°W, 1150 m, 9.viii.1990 (Holzenthal, Blahnik, MunÄoz) (UMSP); 2Y, same location, 14.vi.1986 (Holzenthal, Heyn, Armitage) (UMSP); 1Y, RõÂo Bellavista, c. 1.5 km NW Las Alturas, 8.951°N 82.846°W, 1400 m, 16±17.iii.1991 (Holzenthal MunÄoz, Huisman) (UMSP); 2Y, RõÂo Guineal, c. 1 km E Finca Helechales, 9.076°N

102 Roger J. Blahnik

Fig. 28. Chimarra sensillata, male genitalia.

83.092°W, 840 m, 22.ii.1986 (Holzenthal, Morse, Fasth) (UMSP); 3Y, Jardin Botanico R & C Wilson, trib. along Sendero del Agua, 8.80°N 82.96°W, 1180 m, 8.viii.1990 (Holzenthal, Blahnik, MunÄoz) (UMSP); 1Y, 1X, Quebrada Aguabuena, RincoÂn de Osa, 17.ix.1991 (F. MunÄoz) (UMSP); San JoseÂ, 1X, P. N. Braulio Carrillo, Carrillo, Quebrada Sanguijuela, 10.160°N 83.963°W, 800 m, 11± 12.vi.1988 (C. M. and O. S. Flint, A. ChacoÂn) (NMNH); 1Y, same location, 27.iii.1987 (Holzenthal, Hamilton, Heyn) (UMSP); 2Y, 1X, Reserva Biol. Carara, RõÂo Carara, in Carara, 9.778°N 84.531°W, 200 m, 14.iii.1991 (Holzenthal, MunÄoz, Huisman) (UMSP); 4Y, Reserva Biol. Carara, RõÂo del Sur, 1.5 km S of Carara, 9.769°N 84.531°W, 160 m, 13.iii.1991 (Holzenthal, MunÄoz, Huisman) (UMSP); PANAMA: Canal Zone, 5Y, RõÂo Frijoles, Pipeline Road, 26.ii.1985 (Flint and Louton) (NMNH); San Blas, 4Y, 2X, RõÂo Carti Grande, 2 km W Nusagandi, 5.iii.1985 (Flint and Louton) (NMNH).

Chimarra (Otarrha) sensillata Flint (Figs 28A±D,G; 56) Chimarra (Chimarra) sensillata, Flint, 1981: 12, Figs 29±32; Y, Venezuela (NMNH). This is the nominate species in the C. sensillata species complex, and the only species heretofore described. For a detailed discussion of the differences between the species in this complex, see `Chimarra sensillata species complex', which follows the species descriptions. Chimarra sensillata is most similar to C. tachuela, but differs in several minor characters, the most distinctive of which are the slightly longer dorsal processes of segment VIII, the more elongate ventral process of segment IX and the possession of a ridge on the mesal surface of the inferior appendage, which continues to the base of the appendage as an extension of the apical teeth. Adult. Forewing length (male) 4.9 mm. Colour more or less uniformly dark brown; antennae indistinctly annulate with ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

Systematics of Otarrha light brown. Head somewhat ¯attened, postocular parietal sclerite moderately elongate. Male genitalia. Tergum VIII with short paired projections from posterior margin, acute apically, widely separated basally. Abdominal segment IX with anterolateral margin distinctly expanded; anterodorsal and anteroventral margins with pronounced mesal invagination; posteroventral process narrow, very elongate, projecting, acute apically, tapering from base. Tergum X completely divided mesally into 2 simple short lobes, rounded apically, contiguous basally, each with numerous apical and ventrolateral sensilla, ventrolateral sensilla enlarged. Preanal appendages short, rounded, not (or scarcely) ¯attened. Inferior appendage simple in pro®le; in lateral view distinctly upcurved, tapering; in dorsal view noticeably short, with pronounced mesal curvature, apically with row of incurved sclerotized teeth extending to base as ridge on mesal surface of appendage; basomesally with prominent seta. Phallic apparatus with tubular phallotheca, slightly constricted basally; apicodorsally with lightly sclerous elongate de¯exed projection; apicoventrally with elongate sclerous acutely projecting extension; phallic spines absent. Phallotremal sclerite complex with very prominent rod and ring structure, extending more than half length of phallotheca; rod with abruptly widened lateral platelike projections; rod and ring structure subtended at mid-length by complex sclerite. Female. Unknown. Material examined. VENEZUELA: Aragua, 1Y paratype, Dos Riitos, 6 km N Rancho Grande, 4.ii.1976 (C. M. and O. S. Flint, Jr) (NMNH). Chimarra (Otarrha) septemlobata Flint (Figs 29A±D, G,I; 49A±C; 56) Chimarra (Chimarra) septemlobata Flint, 1991: 28, Figs 26± 28; Y, Colombia (NMNH). Chimarra septemlobata is unique, unlike any other described species, with several diagnostic characters. Diagnostic characters include the possession of a tergum VII with posterior projections, in addition to projections on tergum VIII; short, mesally curved inferior appendages, which have a broad apex and a basomesal spine; and a very prominent, apically truncate ventral process on segment IX. Adult. Forewing length (male) 5.5±6.5 mm (female) 5.9± 6.5 mm. Colour brownish black, head more distinctly black, antennae indistinctly annulate with pale brown. Head ¯attened, postocular parietal sclerite elongate, eyes relatively small. Male genitalia (Fig. 29). Terga VII and VIII both with paired projections from posterior margin, possessing minute spines; projections from tergum VII short, narrow, diverging basally; projections from tergum VIII more elongate, fused basally, diverging apically, apex decurved and indistinctly sclerotized, lacking spines. Abdominal segment IX widest mid-laterally, narrowed dorsally and ventrally; dorsomesal margin strongly invaginated, ventromesal margin only slightly ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

103

so; posteroventral process prominent, projecting, elongate, truncate apically. Tergum X (apparently) forming 2 pairs of lateral lobes, subequal in length; outer pair, in lateral view, narrow, elongate, with slight dorsal in¯ection near middle; inner pair of lobes much wider, uniform in width throughout length; both pairs of lobes with scattered sensilla. Preanal appendages large, ¯attened, earlike. Inferior appendage, in lateral view, moderately elongate, with angular basal in¯ection; appendage narrow basally, widening apically, apex truncate; in dorsal or ventral views with distinct mesal curvature, basally with distinct mesal spine, apically strongly sclerotized and weakly dentate. Phallic apparatus with tubular phallotheca, expanded basally; endotheca short, membranous. Phallotremal sclerite complex composed of rod and ring structure, extending half or more of length of phallotheca, without distinct apical sclerite. Female genitalia (Fig. 49). Segment VIII short; anterolateral margin more or less linear, without apodemes; posterior margin with elongate setae on short broadly rounded projection, subtended ventrally by marginal cleft; posterolateral cleft relatively deep, extending halfway or more to anterior margin; posterior margin nearly linear with acute sclerous projections laterally, in ventral view with truncate mesal projection. Tergum IX short, with short bent apodemes from anteroventral margin; posterodorsal margin with broadly rounded setose mesal projection. Sternum IX membranous. Tergum X bilobed, each lobe divided into lightly sclerous basal region with short mesal protuberance, and less sclerous apical region with cercus at apex. Vaginal apparatus very short and lightly sclerous; apicodorsally with prominent ¯attened sclerous projections, emerging from dorsal surface of vaginal apparatus; rectum opening apically; vaginal apparatus anteroventrally with lightly sclerous lateral margins, lateral sclerites converging posteriorly; anteromesally with de¯exed cuplike sclerite. Material examined. COLOMBIA: Antioquia, 1Y paratype, Quebrada La Iguana, 17 km NW Medellin, 14±15.ii.1984 (C. M. and O. S. Flint, Jr) (NMNH); 24Y, 2X paratypes, same location, 22.ii.1984 (C. M. and O. S. Flint, Jr) (NMNH). Chimarra (Otarrha) septifera Flint (Figs 30A±D,G,H; 57) Chimarra (Chimarra) septifera Flint, 1974: 25±26, Figs 27± 29; Y, Suriname (RNHL). This species is similar in general form to several other species in possessing a tergum X with rodlike dorsal lobes, and inferior appendages which are elongate, mesally in¯ected, and have apices that are ¯attened and narrow to an acute point. It is probably most similar to C. retrorsa, and shares with that species the possession of paired dorsomesal lobes of tergum X, which are narrow and apically acute. Chimarra septifera can be diagnostically distinguished from C. retrorsa by having the apices of these structures acute and apically in¯ected, but not whiplike or retrorse. Other diagnostic characters are found in the structure of the phallic apparatus, and include a phallotheca

104 Roger J. Blahnik

Fig. 29. Chimarra septemlobata, male genitalia (I = dorsal view of terga VII and VIII).

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Systematics of Otarrha

Fig. 30. Chimarra septifera, male genitalia (H = dorsal view of apex of phallic apparatus and phallotremal sclerite complex). ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

105

106 Roger J. Blahnik

Fig. 31. Chimarra spinulifera spinulifera, male genitalia.

with its ventral apex projecting, sclerous and bearing paired ®ngerlike dorsal projections; a phallotremal sclerite with a projecting apicomesal sclerite, and without elongate dorsal phallic spines. Adult. Forewing length (male) 4.7 mm. Colour (in alcohol) yellowish brown. Head somewhat ¯attened, postocular parietal sclerite moderately elongate. Male genitalia: Tergum VIII concavely rounded from median of posterior margin, forming 2 angular dorsolateral prominences. Abdominal segment IX with anterolateral margin

nearly linear, abruptly narrowed dorsally and ventrally; dorsomesal and ventromesal margins strongly invaginated; posteroventral process elongate, projecting, rounded apically. Tergum X (or posterior margin of tergum IX?) with elongate mesal process, membranous mesally and laterally forming 2 narrow elongate sclerites, each with acute apex; tergum X dorsolaterally forming 2 basally separated sclerotized lobes, each elongate, narrow, rounded apically and with several apical sensilla. Preanal appendages large, ¯attened, earlike. Inferior appendage, in lateral view, elongate, angular basally, ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

Systematics of Otarrha strongly mesally ¯exed apically; in dorsal or ventral view with pronounced mesal curvature, enlarged basally, apex angular, acute. Phallic apparatus with narrow tubular phallotheca, expanded basally; ventral apex of phallotheca elongate, sclerotized, with paired narrow ®ngerlike processes from dorsal margin; endotheca short, membranous. Phallotremal sclerite complex large, elaborate, composed of rod and ring structure and apical sclerite; ring with short dorsal projection, rod only moderately elongate, apical sclerite forming ring structure surrounding rod and ring structure, dorsally with short extension and ventrally with elongate narrow mesal extension, nearly length of phallotheca. Female. Unknown. Material examined. SURINAME, 1Y paratype, Brownsberg bergkreek bj Gonddelvers Kamp, 10.viii.1958 (Geijskes) (NMNH). Chimarra (Otarrha) spinulifera spinulifera Flint (Figs 31A±D,F,G; 57) Chimarra (Chimarra) spinulifera spinulifera Flint, 1968c: 151, 153, Figs 4±7; Y, Haiti (MCZ); Botosaneanu, 1996: 12. This subspecies is very similar to C. cubanorum, and also to the newly described subspecies, C. spinulifera galalcha, differing primarily in provenance and in possessing a distinct, small spine on the mesal surface of the inferior appendage. It also differs somewhat from those forms in having an elongate narrow spine emerging from the rounded protuberance at the base of the lateral lobes of tergum X. The apices of tergum X in both subspecies are slightly less narrowed than in C. cubanorum. These differences are all rather minor and I am unsure of their taxonomic signi®cance. Chimarra spinulifera and C. cubanorum are easily distinguished from other species of subgenus by the shape of the inferior appendages, which are short, rounded basally, each possessing a short, thumblike apicodorsal projection. Adult. Forewing length (male) 4.5 mm. Colour yellowish brown. Head distinctly ¯attened, postocular parietal sclerite elongate. Male genitalia (Fig. 32). Tergum VIII with posterior margin slightly projecting. Abdominal segment IX with anterolateral margins broadly rounded, widest ventrad of mid-lateral; dorsomesal and ventromesal margins strongly invaginated; posteroventral process very elongate, projecting, narrow basally, subacute apically. Tergum X completely divided mesally into 2 sclerotized lobes, narrowly separated basally, tapering apically, with distinct mesal curvature in apical third; basomesally with rounded projection, bearing several sclerotized spines, one narrow, elongate and projecting; basolaterally with stout, curved spine; laterally with short spines and numerous sensilla. Preanal appendages large, ¯attened, earlike. Inferior appendage with slight mesal curvature; in lateral view, short, rounded basally, with thumblike apicodorsal projection and small, sclerous spine on mesal surface. Phallic apparatus with narrow tubular phallotheca, slightly constricted basally, expanded basodorsally; endotheca membranous. Phallotremal ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

107

sclerite complex prominent, more than half length of phallotheca, composed of rod and ring structure, ring with dorsal extension, rod strongly upturned apically; rod subtended apically by indistinct, cupped sclerite. Female. Unknown. Material examined. Holotype, Y HAITI: Roche Croix, Mt La Hotte, 4000 ft, 14.x.1934 (Darlington) (MCZ). Chimarra (Otarrha) spinulifera galalcha Botosaneanu (Figs 32A±D,F,G; 50A±C; 57) Chimarra (Chimarra) spinulifera galalcha Botosaneanu, 1996: 11±12, Figs 13±18; Y, Dominican Republic (ZMUA). This subspecies was only recently described. I originally illustrated this form to represent C. spinulifera, without recognizing that it has some distinctive differences from the type. Subsequently, Botosaneanu described this new subspecies. Hopefully, the illustration here, and the comparative illustration of the nominotypical subspecies, will help to distinguish the two forms and lay a foundation for their subsequent taxonomic assessment. This subspecies differs from the nominotypical one in lacking a spine on the mesal surface of the inferior appendages and the spines on the rounded protuberances at the base of the lateral lobes of tergum X are all short. The latter character would also distinguish it from C. cubanorum. Additionally, the apicodorsal lobes of the inferior appendages appear to be less broadly rounded than in the nominotypical subspecies. Adult. Forewing length (male) 4.2±4.4 mm (female) 4.2± 4.8 mm. Colour (in alcohol) yellowish brown. Head distinctly ¯attened, postocular parietal sclerite elongate. Male genitalia (Fig. 32). Tergum VIII with short wide truncate mesal projection from posterior margin. Abdominal segment IX with anterolateral margins broadly rounded, widest ventrad of mid-lateral; dorsomesal and ventromesal margins strongly invaginated; posteroventral process very elongate, projecting, narrow basally, subacute apically. Tergum X completely divided mesally into 2 sclerotized lobes, narrowly separated basally, tapering apically, with distinct mesal curvature in apical third; basomesally with rounded projection, bearing pair of short sclerotized spines; basolaterally with stout curved spine; laterally with short spines and numerous sensilla. Preanal appendages large, ¯attened, earlike. Inferior appendage with slight mesal curvature; in lateral view, short, rounded basally, with thumblike apicodorsal projection. Phallic apparatus with narrow tubular phallotheca, slightly constricted basally, expanded basodorsally; endotheca membranous. Phallotremal sclerite complex prominent, less than half length of phallotheca, composed of rod and ring structure, ring with dorsal extension; rod apically with indistinct sclerite. Female genitalia (Fig. 50). Segment VIII short; anterolateral margin slightly projecting at position of apodemes, apodemes broadly rounded, undeveloped; posterior margin with elongate setae on broadly rounded, mid-lateral projection; posterolateral setae bordered ventrally by deep cleft from posterior margin, extending nearly to anterior margin; poster-

108 Roger J. Blahnik

Fig. 32. Chimarra spinulifera galalcha, male genitalia.

oventral margin slightly projecting, ventral margin nearly linear and bearing linear row of 4 elongate setae on either side of midline. Tergum IX short, with moderately elongate apodemes from anteroventral margin; posteromesally with setose projection, wide basally and truncate apically. Sternum IX membranous. Tergum X bilobed, each lobe divided into lightly sclerous basal region with sclerous mesal protuberance and less sclerous apical region with cercus at apex. Vaginal apparatus short and largely sclerous dorsally, apically diverging to forming paired, ¯attened sclerites; ventrally largely membranous; anteriorly with strongly de¯exed cupped sclerite.

Material examined. DOMINICAN REPUBLIC: La Estrelleta, 5Y, 8X, 4 km SE RõÂo Limpio, c. 760 m, 24± 25.v.1973 (D. and M. Davis) (NMNH). Chimarra (Otarrha) tachuela, sp.n. (Figs 33A±D,G; 51A±C; 56) This is the last of the four species in the C. sensillata species complex. For a detailed discussion of the differences between these species see C. sensillata species complex immediately ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

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109

Fig. 33. Chimarra tachuela, male genitalia (G inset = ventral view of phallotremal sclerite complex and apical spines).

following the species descriptions. This species is similar to C. barinas, but can be distinguished from that species, and the other species of the complex, by lacking projections from the posterodorsal margin of segment VIII. Another unique character is the presence of 2 small tacklike spines in the phallic apparatus, located apical to the phallotremal sclerite complex. Adult. Forewing length (male) 5.5±6.1 mm (female) 6.1 mm. Colour black or brownish black. Head distinctly ¯attened, postocular parietal sclerite elongate. Male genitalia (Fig. 33). Tergum VIII widest mesally, without posterodorsal projections. Abdominal segment IX with anterolateral margin sinuously expanded ventrally, ventral margin with slight mesal invagination; posteroventral process narrow, elongate, projecting, acute. tapering in apical half. Tergum X completely divided mesally into 2 relatively short simple lobes, subacute apically, contiguous basally, each with numerous apical and ventrolateral sensilla; lobes rather distinctly incurved basally, forming ridgelike extension on mesal surface. Preanal appendages rounded, constricted basally, of moderate size, only slightly ¯attened. Inferior ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

appendage simple in pro®le; in lateral view slightly upcurved, tapering, with apicomesal teeth visible (not strongly, mesally incurved); in dorsal view, with sinuous mesal curvature, narrowing, acute apically, apical teeth gradually incurved. Phallic apparatus with tubular phallotheca, slightly constricted basally, apicodorsally with short lightly sclerous projection, gradually tapering, not keeled dorsally; apicoventrally with short paired lateral lobes, not strongly sclerotized; phallic spines 2, short, tacklike, apical. Phallotremal sclerite complex with very prominent rod and ring structure, extending more than half length of phallotheca; rod with sinuously enlarged lateral platelike projections. Female genitalia (Fig. 51). Segment VIII very short; anterolateral margin nearly linear but with pair of short projecting lateral apodemes; posterior margin with several elongate setae mid-laterally, bordered ventrally by deep cleft from posterior margin, extending nearly to anterior margin; in ventral view with posterior margin not projecting and with pair of well de®ned setal warts on either side of midline, each with several prominent setae, surrounded by shorter setae. Tergum IX short with moderately elongate

110 Roger J. Blahnik Table 1. Character differences in C. sensillata group.

Posterior margin of tergum VIII separation of lobes shape of lobes Anteroventral margin of segment IX mesal invagination? Ventral process of segment IX (taper) shape of apex length Tergum X (length) shape of apex, dorsal view basal sensilla enlarged? tergum rolled under basoventrally with ridge on mesal surface? Inferior appendages (apical teeth extending to base as mesal ridge?) Dorsal apex of phallotheca apex keeled? Ventral apex of phallotheca (shape) apex projecting? apex sclerous? Phallotremal sclerite (sclerous apical structure) 2 tacklike apical spines

barinas

incipiens

sensillata

tachuela

bilobed narrow rounded or truncate weakly extended no none or slight + ± rounded moderate moderate rounded no

bilobed wide acute, rel. short strongly extended yes from base acute moderate short rounded yes

bilobed wide acute, mod. elongate strongly extended yes from base acute elongate short rounded yes

unlobed ± ± strongly extended yes from apical half acute moderate moderate relatively acute no

yes

no

no

yes

no short, upturned yes w/o ventral lobes no no

no elongate, de¯exed no tapering, acute yes yes

yes elongate, de¯exed no tapering, acute yes, strongly yes

no short no with paired ventral lobes no no or weakly

apparently absent absent

present absent

present absent

apparently absent present

apodemes from anteroventral margin; posteromesally with broadly rounded projection bearing numerous setae. Sternum IX membranous. Tergum X bilobed, each lobe bulbous, setose, with cercus at apex, and with small sclerous setose basomesal projection. Vaginal apparatus relatively short and almost wholly membranous; anteriorly with lightly sclerous de¯exed cupped sclerite. Material examined. Holotype, Y, VENEZUELA: Merida, La Campana, 12 km SE Santo Domingo, 24.ii.1976 (C. M. and O. S. Flint, Jr) (NMNH). Paratypes, VENEZUELA: Merida, 18Y, 3X, same data as holotype (NMNH); 1Y, P. N. (Parque Nacional) Sierra Nevada, Quebrada La Mucuy, 7 km E Tabay, 8.637°N 71.034°W, 2200 m, 18.i.1994 (Holzenthal, Cressa, RincoÂn) (UMSP); Tachira, 1Y, Quebrada Mesa del Palmar, 5 km S El Cobre, 7°59.851¢N 72°03.795¢W, 2370 m, 18± 20.iv.1995 (Holzenthal, Cressa, Gutic) (UMSP). Etymology. This species is named tachuela, from the Spanish word for tack, in reference to the pair of small tacklike spines at the apex of the phallotremal sclerite complex.

particular to the shape of tergum X and of the inferior appendages. The similarity is also, undoubtedly, due to the species being closely related. I initially considered the species to represent a single variable species, primarily because so many similar forms existed in a relatively restricted geographical region, and because each form was represented from either a single collection or collections in close geographical proximity. However, the variation is rather greater than ordinarily encountered in a single species and appears to occur primarily between, and not within, populations. I felt that I was being subjectively in¯uenced by a human perceptual bias, based on the structural simplicity of the species, and an unwillingness to create names for so many similar forms. Ultimately, I decided to give each of the `species' a name, because each is diagnosable and, upon careful examination, can be found to differ in a number of minor characters from the other `species'. The differences, although often subtle, are numerous. They are listed in Table 1 so that the differences can be more directly compared.

The Chimarra sensillata species complex

Species incertae sedis

The species of this complex include C. barinas, C. incipiens, C. sensillata Flint and C. tachuela. These four species are all similar, differing primarily in features of the development of tergum VIII, and subtle features of the shape of tergum X, the shape of the inferior appendages and characters of the phallic apparatus. The differences appear minor, in part, because the structural groundplan for the species is simple, referring in

The following two species are being treated as insertae sedis. They possess a similar hindwing venation to members of subgenus Otarrha and would key out here in the subgeneric key of Blahnik (1998). However, in other respects the species are very different from Otarrha and I do not believe that they belong within subgenus Otarrha proper. Except for the venational reduction, none of the other characters used to ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

Systematics of Otarrha

111

Figs 34±36. 34, Chimarra amazonia, female genitalia; 35, C. diaphora, female genitalia; 36, C. dominicana, female genitalia.

de®ne subgenus Otarrha are present. As noted in the phylogenetic analysis (below), venational reduction in the hindwing has arisen several times in genus Chimarra, and its presence here is not convincing evidence of a close relationship to Otarrha. In general, the character states possessed by ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

the following two species are simple and probably plesiomorphic, or peculiar and derived. Although these species also do not possess the characters used to de®ne subgenus Curgia (Blahnik, 1997, 1998; Flint, 1998), several characters suggest that they may be primitive

112 Roger J. Blahnik

Figs 37±39. 37, Chimarra guapa, female genitalia; 38, C. jamaicensis, female genitalia; 39, C. machaerophora, female genitalia.

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Systematics of Otarrha and basal members of that lineage. The problem with most of these characters is that they are either dif®cult to polarize or illde®ned and subject to homoplasy. Character similarities noted are as follow. (1) The presence in the females of a ventral process on segment VIII. This character is universally found in Chimarra (Curgia), and never in subgenera Chimarra or Otarrha. In the latter two subgenera, a ventral process is present on segment VII. In Chimarra (Chimarrita) two lineages are found, one with a ventral process on segment VII and one with a ventral process on segment VIII. (2) The structure of the anal veins of the forewing. Like members of subgenera Curgia and Otarrha, the 3A anal vein is looped onto the 1A anal vein distal to intersection of the 2A vein, with no connecting crossvein between 2A and 3A, as commonly occurs in subgenera Chimarrita and Chimarra. In members of subgenus Otarrha, the point of intersection is in the proximal half of the 1A vein, and in subgenus Curgia it is at half length or more. Additionally, the 3A vein has a rather distinct, angular basal in¯ection in Curgia. In both of these characters, the following two species resemble members of subgenus Curgia. (3) The size of the anal loop of the hindwing. In subgenera Chimarrita and Otarrha, the anal loop is typically very small. In subgenera Curgia and Chimarra, it is typically fairly large, and this is the case in the following two species. Chimarra angularis, sp.n. (Figs 52A±D,F,G; 54A,B; 58) This new species is similar to C. usitatissima, and the two could be easily confused. Diagnostic character differences of the male genitalia include structures of tergum X and the inferior appendages. Tergum X of C. angularis is shorter and, in lateral view, appears more bent than that of C. usitatissima. Also, the apex of the inferior appendage of C. angularis is shorter, wider and more rounded than C. usitatissima, and the ridge on the mesal surface of the inferior appendage is less

Fig. 40. Chimarra odonta, female genitalia.

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113

distinct and does not continue to the apex. Because both species are widespread and were found together at two widely separate locations, and because all material examined could be separated into one of the two forms, I am considering this a distinct species. Adult. Forewing length (male) 4.3±5.3 mm (female) 4.7± 5.4 mm. Colour uniformly medium brown, except coxae and femora yellowish brown and antennae annulate with light brown. Head short and rounded, postocular parietal sclerite short. Eyes very large. Maxillary palp 5-segmented, very short; second segment stouter and slightly longer than third, apicomesally with stout bristles. Labial palp 3-segmented. Setae of head elongate and dense. Forewing venation typical for Chimarra; s, r-m and m cross veins linearly arranged and unpigmented, forming indistinctly evident bar; anal veins with 3A looped to 1A. Hindwing with reduced venation: subcosta narrowly parallel to R (apparently fused), Rs 3-branched, M 2branched. Male genitalia (Fig. 52). Segment IX with anterior margin concave, narrow dorsally and with ventral margin distinctly produced; dorsally with paired projections from posterior margin, each with short scalelike or spinelike setae; posterior margin of segment IX very angularly produced mid-laterally; ventral process elongate, narrow, projecting, apically acute. Tergum X simple, with numerous sensilla; apically cleft for more than half of length; in lateral view bent, with 2 points of in¯ection about equally spaced. Preanal appendages short, rounded. Inferior appendage short, mesally curved, rounded basally, with short apicodorsal projection; apicodorsal projection short, apically rounded; mesal surface of appendage with distinct ridge, not extending to apex. Phallic apparatus with short tubular phallotheca, slightly expanded basodorsally; endotheca with elongate membranous apicodorsal projection bearing numerous short lightly sclerotized setae on ventral surface. Phallotremal sclerite complex composed of rod and ring structure extending more than half length of phallotheca, apically with subtending sclerite, wide at base and narrowing to elongate tapering decurved apex.

114 Roger J. Blahnik

Figs 41±43. 41, Chimarra parilis, female genitalia; 42, C. particeps, female genitalia; 43, C. patosa, female genitalia.

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Systematics of Otarrha

Figs 44±46. 44, Chimarra phthanorossi, female genitalia; 45, C. puertoricensis, female genitalia; 46, C. quadrifurcata, female genitalia.

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115

116 Roger J. Blahnik

Figs 47±49. 47, Chimarra redonda, female genitalia; 48, C. rossi, female genitalia, (D) lateral view of vaginal apparatus and terga IX and X; 49, C. septemlobata, female genitalia.

ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

Systematics of Otarrha Female genitalia (Fig. 54). Segment VIII narrow, synsclerous; anterior margin nearly linear, with short projecting lateral apodemes; posterolateral margin with 3 widely separated rounded setal warts on either side; posteroventral margin slightly produced; anteroventrally with short ventral process rounded or subacute at apex. Tergum IX short, with moderately elongate anteroventral apodemes; posterodorsally with short paired mesal projections, each with several short setae; preapically with several elongate spaced setae on either side of midline. Sternum IX membranous. Tergum X composed of paired bulbous lobes, each with numerous short setae, apically with paired cerci. Vaginal apparatus membranous and indistinct, anteriorly with cuplike sclerite. Material examined. Holotype, Y, VENEZUELA: T. F. Amazonas, Cerro de la Neblina, basecamp (rainforest clearing near RõÂo Baria), 0°50¢N 66°10¢W, 140 m, 26± 31.i.1985 (P. J. and P. M. Spangler, R. Faitoute, W. Steiner) (NMNH). Paratypes, VENEZUELA: Bolivar, 1Y, 1X, RõÂo Cuyuni, El Dorado, 10.ii.1976 (C. M. and O. S.

Flint, Jr) (NMNH); T. F. Amazonas, 1Y, RõÂo Orinoco, Puerto Ayacucho, 8.iii.1984 (O. S. Flint, Jr) (NMNH); 5Y, 10X, RõÂo Cataniapo, 10 km S Puerto Ayacucho, 9.iii.1984 (O. S. Flint, Jr) (NMNH); 2Y, 6X, San Carlos de RõÂo Negro, 23.i.1985 (P. and P. Spangler, R. Faitoute, W. Steiner) (UMSP); 1Y, San Carlos de RõÂo Negro, 1°56¢N 67°03¢W, 6±12.xii.1984 (R. L. Brown) (NMNH); 1Y, Cerro de la Neblina, basecamp (malaise trap in rainforest), 0°50¢N 66±10¢W, 140 m, 19.ii.1985 (P. J. and P. M. Spangler, R. A. Faitoute, W. E. Steiner) (NMNH). GUYANA: Mazaruni-Potaro District, 1Y (in alcohol), Takutu Mtns (forest clearing near streams), 6°15¢N 59°5¢W, 18.xii.1983 (P. J. Spangler and W. E. Steiner) (NMNH); 1Y, 1X (in alcohol) same location, 3±10.xii.1983 (W. E. Steiner and P. J. Spangler) (UMSP); 1Y, same location, 11.xii.1983 (P. J. Spangler, R. A. Faitoute, W. E. Steiner) (NMNH). Etymology. This species is named angularis for the short bent lateral lobes of tergum X, which alerted me to the possibility that this might represent a distinct species.

Figs 50, 51. 50, Chimarra spinulifera galalcha, female genitalia; 51, C. tachuela, female genitalia. ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

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118 Roger J. Blahnik

Figs 52, 53. 52, Chimarra angularis, male genitalia; 53, C. usitatissima, male genitalia (G inset = dorsal view of phallotremal sclerite complex).

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Systematics of Otarrha Chimarra usitatissima Flint (Figs 53A±D,G; 55; 58) Chimarra (Chimarra) usitatissima Flint, 1971: 24±25, Figs 43±45, Y, Brazil (NMNH); Flint, 1974: 29, Figs 40± 42. As discussed above, distinguishing differences between this species and C. angularis include a longer tergum X and an inferior appendage with a more acute apex, which possesses a mesal ridge extending to the apex. Adult. Forewing length (male) 4.9±5.2 mm (female) 5.3 mm. Colour uniformly medium brown, antennae annulate with light brown. Head short and rounded, postocular parietal sclerite short. Eyes very large. Maxillary palp 5segmented, very short; second segment stouter and slightly longer than third, apicomesally with stout bristles. Labial palp 3-segmented. Setae of head elongate and dense. Forewing venation typical for Chimarra; s, r-m and m cross veins linearly arranged and unpigmented, forming indistinctly evident bar; anal veins with 3A looped to 1A. Hindwing with reduced venation: subcosta narrowly parallel to R (apparently fused), Rs 3-branched, M 2-branched. Male genitalia (Fig. 53). Segment IX with anterior margin concave, narrow dorsally and with ventral margin distinctly produced; dorsally with paired projections from posterior margin, each with short scalelike or spinelike setae; posterior margin of segment IX angular, mid-laterally produced; ventral process elongate, narrow, projecting, apically acute. Tergum X simple, with numerous sensilla; apically cleft for more than half of length; in lateral view moderately elongate and bent, with 2 points of in¯ection, one basodorsally and one apically; intervening dorsal region more elongate than basal or apical regions. Preanal appendages short, rounded. Inferior appendage short,

119

mesally curved, rounded basally, with distinct apicodorsal projection; apicodorsal projection relatively elongate, tapering, apically acute; mesal surface of appendage with distinct ridge, extending to apex. Phallic apparatus with short tubular phallotheca, slightly expanded basodorsally; endotheca with elongate, membranous apicodorsal projection, bearing numerous short lightly sclerotized setae on ventral surface. Phallotremal sclerite complex composed of rod and ring structure, extending more than half length of phallotheca, apically with subtending sclerite, wide at base and narrowing to elongate tapering decurved apex. Female genitalia. Segment VIII narrow, synsclerous; anterior margin nearly linear, with short projecting lateral apodemes; posterolateral margin with 3 widely separated rounded setal warts on either side; posteroventral margin slightly produced; anteroventrally with short ventral process, distinctly acute at apex (Fig. 55). Tergum IX short, with moderately elongate anteroventral apodemes; posterodorsally with short paired mesal projections, each with several short setae; preapically with several elongate spaced setae on either side of midline. Sternum IX membranous. Tergum X composed of paired bulbous lobes, each with numerous short setae, apically with paired cerci. Vaginal apparatus membranous, anteriorly with cuplike sclerite. Material examined. VENEZUELA: Bolivar, 1Y, RõÂo Grande Res. Sta., 26 km E of El Palmar, 700 ft, 24± 25.iii.1978 (J. B. Heppner) (NMNH); T. F. Amazonas, 1Y, 1X, Cerro de la Neblina, basecamp, °51¢N 66°10¢W, 140 m, 20±24.iii.1984 (O. Flint and J. Louton) (NMNH). GUYANA: Mazaruni-Potaro District, 1Y, 1X, Takutu Mtns (forest clearing near streams), 6°15¢N 59°5¢W, 18.xii.1983 (P. J. Spangler and W. E. Steiner) (UMSP). SURINAME: 1Y, Tapanahoni RivierKamp, 11.v.1952 (D. C. Geijskes) (NMNH). ECUADOR: Napo, 2Y, 2X, Puerto Montufar,

Figs 54, 55. 54, Chimarra angularis, female genitalia; 55, C. usitatissima, female genitalia (A = lateral view of ventral process of segment IX).

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120 Roger J. Blahnik

Fig. 56. Distribution of Chimarra (Otarrha) species.

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Systematics of Otarrha

Fig. 57. Distribution of Chimarra (Otarrha) species.

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122 Roger J. Blahnik

Fig. 58. Distribution of incertae sedis species.

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Systematics of Otarrha 29.iv.1976 (J. Cohen) (NMNH); 2Y, 3X, Limoncocha, 15.vi.1977 (P. J. Spangler and D. R. Givens) (NMNH). PERU: Loreto, 1Y, Callicebus Res. Station, Mishana, RõÂo Nanay, 25 km SW Iquitos, 120 m, 17.i.1980 (J. B. Heppner) (NMNH); Madre de Dios, 2Y, 4X, RõÂo Tambopata Res., 30 km SW Puerto Maldonado, 290 m, 6±10.xi.1979 (J. B. Heppner) (NMNH). BRAZIL: Holotype Y (type # 7412), Rio Branquinho, bei Cachoeira, 21.vii.1961 (E. J. Fittkau) (NMNH); Minas Gerais, 3Y, Serra do Cipo, km 110, 8.x.1975 (Froehlich and Vanin) (NMNH); ParaÂ, 1Y (in alcohol), Rio GurupõÂ, vi.1963 (B. Malkin) (NMNH); Rondonia, 3Y, 8 km S Cacaulandia, 21.xi.1991 (D. Petr) (NMNH).

Phylogenetic considerations Monophyly of subgenus Otarrha Evidence for monophyly of subgenus Otarrha is based on the absence of apomorphic characters used to de®ne other subgenera of Chimarra, and the presence of a convincing number of unique synapomorphic characters. In addition, species of Otarrha share a number of character states that are probably plesiomorphic, but the retention of which serves as a re¯ection of their relatedness, because the characters, individually, are modi®ed in one or another of the other subgenera of Chimarra. The characters listed below are believed to be

Fig. 59. Cladogram for Chimarra (Otarrha) species. Nodes with decay index greater than one indicated. ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

123

124 Roger J. Blahnik synapomorphic and are used to infer the monophyly of Otarrha. Characters were polarized by the use of Chimarrhodella as an outgroup, and by comparison to character states in the other subgenera of Chimarra. (1) Anterior setal warts divided into 2 rounded setal areas (apomorphic); anterior setal warts elongate, diagonal, undivided (plesiomorphic). The apomorphic character state is found in all species of Otarrha except C. odonta, in which the plesiomorphic condition may be primitively retained. The plesiomorphic character state is found in Chimarrhodella and all other subgenera of Chimarra. (2) M of hindwing 2-branched (®rst fork lost) (apomorphic); M of hindwing 3-branched (plesiomorphic). The apomorphic character state is found in all species of subgenus Otarrha. Loss of the ®rst fork of M vein (M 2branched) is a character also found in some species of Chimarrita and in several lineages of subgenus Chimarra, interpreted as independent losses in each. Venational reduction in Philopotamidae seems to follow a similar course whenever it occurs. Only in the primitive Paulianodes is M of the hindwing 4-branched. In other genera, M is typically 3-branched (M4 absent). A corresponding reduction of M of the forewing (M4 absent) is a character used to de®ne Chimarrinae. Additional venational loss in Chimarrinae occurs in Chimarra, and inevitably involves only the hindwing. Venational loss occurs in a rather predictable sequence. The ®rst venational reduction involves a fusion of the subcosta with R1. Often the veins run narrowly parallel (apparently fused) rather than being completely fused, and may be divergent (apparently branched) near the apex. The ®rst vein to be lost is M4 (M becoming 2-branched); this character may accompany the ®rst reduction. The last reduction to occur is a loss of the ®rst fork of Rs (Rs 3-branched). All three of these venational reductions occur together in various lineages of Chimarra. (3) Subcosta of hindwing fused to R1 (apomorphic); subcosta of hindwing not fused to R1 (plesiomorphic). The apomorphic character state is found in all species of Otarrha except C. odonta, in which the plesiomorphic condition may be primitively retained. The plesiomorphic character state typi®es genera Chimarrhodella and Chimarra. However, a similar venational fusion occurs in some species of subgenera Chimarrita and Chimarra (see 2). (4) First fork of Rs of hindwing lost (Rs 3-branched) (apomorphic); Rs of hindwing 4-branched (plesiomorphic). The apomorphic character state is found in all species of Otarrha except C. odonta, in which the plesiomorphic condition may be primitively retained. The apomorphic character state typi®es genera Chimarrhodella and Chimarra. However, a similar venational loss occurs in some species of subgenera Chimarrita and Chimarra (see 2). (5) Third segment of maxillary palp constricted basally and with dorsal bulge in basal half (apomorphic); third segment of maxillary palp uniform in width or gradually expanded apically (plesiomorphic). The apomorphic character state is found in all members of subgenus Otarrha, although somewhat variably developed. A similar condition to that found in

Otarrha has not been found in other species of Chimarra, or Chimarrhodella, but only a limited number of species in the subgenera were surveyed for the character. (6) Ventral process of segment IX elongate, narrow and projecting, constricted basally and rounded apically (apomorphic); (plesiomorphic state?). The former character state typi®es members of subgenus Otarrha. In members of the sensillata species complex, the process is elongate, tapering and acute; the process is truncate apically in several other species, but the groundplan is similar in all cases. The similarity in the development of the structure among the species of Otarrha is used to infer their close relationship. However, the character cannot really be polarized because a ventral process is absent in Chimarrhodella and the process is very differently developed in other subgenera of Chimarra. When a ventral process occurs in other genera of Philopotamidae, it is often on one or more segments preceding segment IX but sometimes also variously developed on segment IX. In most cases, these processes are elongate, projecting and acute, and the inference that this is the plesiomorphic condition for Chimarra is reasonable. This character state typically occurs in subgenus Chimarrita, which is plesiomorphic for a number of other character states and may be a basal lineage for Chimarra as a whole. An elongate ventral process (whether rounded apically or acute) is exceptional among members of subgenus Curgia, and limited to only some lineages of subgenus Chimarra, most notably among species of the obscura group of eastern North America, and in certain African species. (7) Preanal appendages large, ¯attened and earlike in shape (apomorphic); preanal appendages elongate, linear (plesiomorphic). The apomorphic character state is absent (or dubious) only in a few species of Otarrha, including C. odonta and members of the sensillata species complex. The plesiomorphic condition occurs in Chimarrhodella and may represent the condition found in the common ancestor of Chimarra and Chimarrhodella. In genus Chimarra, preanal appendages are typically short, rounded and knoblike and this is probably the plesiomorphic condition for the genus (present in the common ancestor of surviving lineages). (8) Ventral setal areas of segment VIII diffuse, ill-de®ned, sometimes merging mesally (apomorphic); ventral setal areas paired, well de®ned and separate (plesiomorphic). The apomorphic character state occurs in all species of subgenus Otarrha for which female genitalia were available, except C. sensillata, which has paired, well de®ned ventral setal areas, possibly as a retention of the plesiomorphic character state. (9) Posterolateral margin of segment VIII with pronounced cleft (apomorphic); posterolateral margin of segment VIII uncleft (plesiomorphic). The apomorphic condition is found in all species of Otarrha for which female specimens were available and has never been found in species of other subgenera of Chimarra. It should be noted, however, that the cleft possibly re¯ects a natural constriction at the point of fusion of the tergum and sternum. (10) Posteromesal margin of tergum IX with projecting setose area (apomorphic); posterior margin of tergum IX unprojecting (plesiomorphic). The apomorphic character state ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

Systematics of Otarrha is found in all species of Otarrha for which female specimens were available. The number and arrangement of setae varies in other subgenera of Chimarra and in Chimarrhodella, but are not typically on a mesal projection. (11) Tergum X divided into (more or less) sclerous basal lobe and nonsclerous, apical cercus-bearing lobe (apomorphic); tergum X simple, rounded, and nonsclerous (plesiomorphic). The apomorphic character state is peculiar to Otarrha and found in all species examined, although the size, shape and sclerosity of the basal portion of the segment is variable. This basal, sclerous structure is considered a part of tergum X only by inference, as a comparable structure is absent in both Chimarrhodella and other subgenera of Chimarra. Phylogenetic analysis of subgenus Otarrha The cladistic analysis produced 320 equally most parsimonious cladograms of length = 111, CI = 0.685 and RI = 0.855. Despite the large number of cladograms obtained, the strict consensus (Fig. 56) is quite well resolved. An analysis was also performed using only the Chimarrhodella species as outgroup and deleting the other outgroup taxa, although retaining Chimarra usitatissima, as it potentially fell within the ingroup in the initial analysis. This resulted in eighty equally most parsimonious cladograms, the strict consensus of which had the same resolution and relative relationships among the ingroup taxa as the previous. The large number of cladograms obtained in the ®rst analysis resulted in part from a lack of resolution of relationships among the outgroup taxa. If Chimarra usitatissima was deleted from the analysis, the relative positions of the two most basal taxa, C. odonta and the sensillata species complex (as a whole), were reversed. In the analyses in which C. usitatissima was retained, the sensillata group was the most basal taxon, and when it was omitted C. odonta was the most basal taxon. This difference can be attributed to venational characters used in the analysis. Reasons for suspecting that these characters might be subject to homoplasy were discussed under character (2) in the section dealing with monophyly of subgenus Otarrha. Because of uncertainty about the phylogenetic position of C. usitatissima, and the relative placement of C. odonta and the sensillata species complex, the node for the latter two taxa was arbitrarily collapsed in the cladogram presented. Deleting other combinations of outgroup taxa produced various numbers of cladograms, but never changed the relationships or resolution of ingroup taxa. Decay indices (Bremer, 1988; Donoghue et al., 1992) for the remaining nodes are indicated on the cladogram (Fig. 56). Bootstrap values (Felsenstein, 1985) for the nodes were also attempted, but the large number of cladograms generated in some of the bootstrap runs rapidly exceeded the memory capacity of the computer used, even when nodes with a decay index greater than two were constrained. For reasons discussed above and under the descriptions of C. usitatissima and its close sister taxon C. angularis, these species are being retained as incertae sedis. Although it is possible that these species are basal taxa in subgenus Otarrha, ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

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as indicated in the phylogenetic analysis, this hypothesis is only weakly supported. This compares to the relatively strong support for monophyly of subgenus Otarrha otherwise de®ned. Chimarra usitatissima has a generalized morphology, including a simple, divided tenth tergum, which resembles basal species in subgenus Otarrha, but also various species in subgenus Curgia. Incidental homoplasy is most likely to affect inclusion of taxa at the base of a clade, especially when the morphology of the taxa is otherwise plesiomorphic, and there is some reason to believe that this is what is happening with C. usitatissima and C. angularis because the only characters indicating their possible inclusion in subgenus Otarrha are venational loss characters. Chimarra odonta is an interesting species with several characters that clearly suggest its placement within subgenus Otarrha, although in a basal or nearly basal position. It is interesting that, in addition to being a basal species, it is also the only species known from southeastern Brazil. Other species in subgenus Otarrha fall into four major clades. I have refrained from dividing the subgenus into formal species groups because of my uncertainty about the placement and relationships of several species, particularly the most basal taxa, C. odonta and the C. sensillata species complex, but also C. septemlobata and C. puertoricensis. One of these informal species groups is represented by the sensillata complex. As discussed above, there is some ambiguity about whether this species complex or C. odonta constitutes the most basal lineage in subgenus Otarrha. Like C. odonta, members of this species complex have a relatively simple, short, rounded tenth tergum with numerous apical sensilla. The species from the Greater Antilles probably all belong to a single monophyletic clade, although character evidence uniting the entire group is limited. The species with the most distinctive differences are C. guapa and C. puertoricensis. The development of elongate inferior appendages in C. guapa, giving the species a super®cial resemblance to C. patosa and related species, likely constitutes an independent apomorphic development. In most other respects, it is a typical member of the Antillean group of species. Chimarra puertoricensis, by contrast, is different in a number of respects from other species in this group, including the shape of the inferior appendages, the possession of deeply cleft lateral lobes of tergum X and lateral lobes somewhat separated mesally. I consider its close relationship to C. redonda doubtful, and would (subjectively) collapse the node of the cladogram indicating this relationship, although probably retaining it within the Antillean lineage. The two other major clades of Otarrha are both South American and very likely related to one another. Species in the subgenus from the Lesser Antilles fall here. Both groups have elongate, curved inferior appendages and lateral lobes of tergum X, which are partially or wholly cleft laterally and separated mesally by a sclerous process (reduced in some cases but still leaving the lateral lobes of tergum X separated mesally). Many of the species have the posterior margin of tergum VIII modi®ed. Most of the species also either have phallic spines or a phallotremal sclerite complex that is elongate and sclerous. The patosa group of species, with known members con®ned to Peru, all have a mesal tine on the

126 Roger J. Blahnik inferior appendage. The species belonging within this clade include, in addition to C. patosa itself, C. amazonia, C. parene, C. parilis, C. particeps and C. peruana. The other major clade of Otarrha in South America can be referred to as the retrorsa group. Except perhaps for C. dominicana, members of the retrorsa group of species are characterized by having the apex of the inferior appendage ¯attened, but with a projecting, acute apex. Many of these species have the lobes of tergum X rodlike, sometimes elongate, with apical sensilla. These species also possess separate ventrolateral processes, quite variably developed in the various species, which typically lack sensilla. These may either represent a ventral part of the lateral lobe of tergum X, resulting from the lateral lobes being wholly cleft laterally, or (perhaps more likely) some de novo structure derived from the periphallic membrane. Species rather clearly belonging within this clade include C. diakis, C. diannae, C. diaphora, C. dominicana, C. quadrifurcata, C. retrorsa, C. septifera, C. rossi and C. phthanorossi. The relationship of C. septemlobata, which is also an interesting and distinct species, is not clear, although it is probably related to, or af®liated with, one of the two major clades of South American species discussed above. Biogeography The geographical distribution of subgenus Otarrha, in general aspects, is similar to that of subgenus Chimarrita (Blahnik, 1997). Otarrha is widespread in the northern part of South America, from the Guyanas in the east to Peru in the west, with a single, basal or nearly basal species also found in southeastern Brazil. It is additionally found on all of the major islands of the Greater Antilles, and also represented in the Lesser Antilles, but by species with a closer relationship to species from South America than to those of the Greater Antilles. This is consistent with the more recent geological origin of the Lesser Antilles (Donnelly, 1988). Only one species, C. rossi, is known from Central America. Its known distribution is limited to Costa Rica and Panama, and it has a similar, but less apomorphic, sister species in Colombia. The absence of the subgenus in most of Central America is striking. The general distribution for the subgenus conforms to a South American Caribbean distribution track, discussed by Rosen (1976), and constitutes one of several area relationships between the biota of the Greater Antilles and that of continental areas. However, it differs from many taxa with this distribution in lacking a signi®cant Central American distribution. The recently described subgenus Chimarrita (Blahnik, 1997) has a similar distributional pattern. The origin and biogeographical relationships of the fauna of the Greater Antilles has been a source of interest and speculation for many years. There is considerable disagreement about the geological history of the area, in part due to the tectonic complexity of the region, both currently and historically because it involves several minor tectonic plates. The Greater Antilles themselves are located on the Caribbean plate, which owes its origin to a ¯ood basalt resulting from the separation of the North and South American tectonic plates

during the early Jurassic (Donnelly, 1988). Rosen (1976) provided a review of the many hypotheses for the origin of the Antilles, and discussed biogeographical implications of the theories. The considerable differences between the theories makes the whole ®eld of geological reconstruction of plate tectonic history seem speculative. However, there has been an improved understanding of plate tectonics gained in recent years, and this has made some of the early theories less likely. Donnelly (1988) summarized what is currently known about the origin of the Antilles from a geological perspective, and also discussed the restrictions this poses for biogeographical reconstructions. In general, the Antilles may have served as a proto-isthmus connecting South and Central America, as Rosen (1976) suggested, but its location at the site of the current isthmus is unlikely. Additionally, it was more likely a ®lter bridge, with intervening water gaps, and never a continuous landmass. A signi®cant connection of the Greater Antilles with South America has probably not occurred since the end of the Cretaceous or early Cenozoic, and this is the most probable date for the origin of the Antillean lineage of subgenus Otarrha. The relationships of the islands themselves have a complex history, resulting, in part, from the sinistral movement of the islands, commencing with the opening of the Cayman trough. This movement has stretched the island chain and brought different land areas into juxtaposition at various periods in the history of the islands. It has also, undoubtedly, been a factor in producing the close relationship that currently exists between the faunal elements of the respective islands. For instance, the large islands of Cuba and Hispaniola, as they exist today, are actually composed of island elements that have only recently been brought into juxtaposition. Current understanding of Caribbean geology supports a complex history for the Antillean fauna, involving inter-island exchange and augmentation of the original fauna from various mainland sources, depending on historic opportunities for dispersal and also on the dispersal capabilities of the various faunistic elements. In the light of the above discussion, it is interesting that Chimarra is represented on the Greater Antilles by three different subgenera, each of which is also widespread in South America. Of these, only subgenus Curgia is also widespread in Central America, and then only represented by a few lineages. In contrast, subgenus Chimarra is entirely absent from the Greater Antilles, although currently widespread and abundant in both Central America and the northern parts of South America. As suggested by Ross (1956), this subgenus is likely to have been an Old World immigrant to the New World. Its absence from the Greater Antilles makes it unlikely that it could have reached the New World before the early Cenozoic. Five species of Chimarra have been described from Oligocene amber deposits from the Dominican Republic (Wichard, 1983a, 1983b). Unfortunately, I have not been able to place these species to subgenus, in part because of the incomplete character information available in the species descriptions, but it is clear that they are not members of subgenus Chimarra. ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

Systematics of Otarrha As noted in the phylogenetic analysis, species of subgenus Otarrha from the Greater Antilles probably form a monophyletic group. The group has most of the apomorphic characters of the subgenus as a whole, and the species bear a close general resemblance to the major South American groups of the subgenus, the patosa and retrorsa groups. All of these species are characterized by large, ¯attened preanal appendages. Within the Antilles, the species from Cuba (all from the eastern part of the island) seem to have their closest af®nities either with each other, or with species from Hispaniola and Jamaica. The sole representative of the subgenus from Puerto Rico is, in many ways, the most distinctly different of the group. Among the South American representatives of the subgenus, most belong to either the retrorsa or patosa groups, which are probably sister taxa, as indicated by the phylogenetic analysis. The patosa group is only known from Peru, and the retrorsa group is found from the Guyanas, Venezuela, Colombia and lower Central America. The most primitive members of the subgenus include C. odonta from southeastern Brazil, and members of the Chimarra sensillata species complex from Venezuela. These areas also coincide with areas of diversi®cation within subgenus Curgia (Flint, 1998) and also subgenus Chimarrita (Blahnik, 1997). What is known of the distribution and phylogeny of subgenus Otarrha suggests an origin that includes the collective area of the Brazilian subregion of South America and also the Greater Antilles, but not Central America. The absence of the genus from Central America may be due to any of several causes: the subgenus may be historically absent from Central America; its absence may be only apparent due to inadequate collecting; or the subgenus may have become extinct there, possibly as a result of competition with members of subgenus Chimarra, which are currently abundant and widespread in the region and could have displaced subgenus Otarrha. The similar distribution of subgenus Chimarrita (Blahnik, 1997) might be used as evidence to support the ®rst explanation. However, it will undoubtedly be necessary to clarify the historical components that have produced the distributional patterns seen among the various lineages of organisms inhabiting the Greater Antilles before any consensus hypothesis can be proposed. Acknowledgements A number of people contributed in one way or another to this paper and I would like to thank them all. In particular, I would like to acknowledge the contributions of Professor Ralph Holzenthal, University of Minnesota, St Paul, who was my academic advisor and funded the research through two National Science grants awarded to him, BSR-8917684 and DEB-9400632, and Dr Oliver S. Flint, Jr, currently emeritus Curator of the Division of Neuropteroids, Smithsonian Institution, who generously made available the bulk of specimens on which the research was done, and also gave freely of his time and advice. I would also like to acknowledge my gratitude to the University of ã 2002 Blackwell Science Ltd, Systematic Entomology, 27, 65±130

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Minnesota Graduate School for funding provided through a Doctoral Dissertation Fellowship, and Dr Karl Kjer who assisted in the phylogenetic analysis and also provided funding under NSF grant DEB 96-32879. Finally, I thank those other people who contributed to the research, Dr Abelardo Sandoval who organized and sponsored the Smithsonian BioLat expedition to Peru that I accompanied, Dr L. Botosaneanu, Institute for Taxonomic Zoology, University of Amsterdam, who made specimens available and also shared information and insights, and Cleone T. Graham, Museum of Comparative Zoology, Harvard, also for making specimens available.

References Blahnik, R.J. (1996) The Systematics and Biogeography of the Genus Chimarra Subgenus Chimarra Stephens for the Neotropics (Trichoptera: Philopotamidae). PhD Dissertation, University of Minnesota, St Paul. Blahnik, R.J. (1997) Systematics of Chimarrita, a new subgenus of Chimarra (Trichoptera: Philopotamidae). Systematic Entomology, 22, 199±243. Blahnik, R.J. (1998) A revision of the Neotropical species of the genus Chimarra, subgenus Chimarra (Trichoptera: Philopotamidae). Memoirs of the American Entomological Institute, 59, 1±317. Blahnik, R.J. & Holzenthal, R.W. (1992) New species of Chimarra subgenus Chimarra Stephens from Costa Rica (Trichoptera: Philopotamidae). Proceedings of the Washington Entomological Society, 94, 409±438. Botosaneanu, L. (1977) TrichopteÁres (imagos) de Cuba, captures par moi-meme en 1973 (Insecta, Trichoptera). Fragmenta Entomologica, 13, 231±284. Botosaneanu, L. (1980) TrichopteÁres adultes de Cuba collectes par les zoologistes cubain (Trichoptera). Mitteilungen MuÈnschener Entomologische Gesellschaft, 69, 91±116. Botosaneanu, L. (1988) TrichopteÁres de la Martinique. Annales de la SocieÂte entomologique de France (N.S.), 24, 215±228. Botosaneanu, L. (1990) Results of a trichopterological (Insecta: Trichoptera) travel to the Lesser Antilles in 1989. Bulletin de l'Institute Royal des Sciences Naturelles de Belgique, Entomologie, 60, 39±40. Botosaneanu, L. (1994a) A study of the larvae of caddis¯ies (Trichoptera) from Cuba. Tropical Zoology, 7, 451±475. Botosaneanu, L. (1994b) Les TrichopteÁres de la Guadeloupe. Annales de la SocieÂte entomologique. Francaise (N.S.), 24, 215±228. Botosaneanu, L. (1996) Caddis ¯ies (Trichoptera) from the Dominican Republic (West Indies). II. All families except Hydroptilidae; with general observations for Hispaniola. Bulletin de l'Institute Royal des Sciences Naturelles de Belgique, Entomologie, 66, 5±26. Bremer, K. (1988) The limits of amino acid sequence data in angiosperm phylogenetic construction. Evolution, 42, 795±803. Bueno-Soria, J. (1985) Estudios en insectos acuaticos III. cinco nuevas especies de Chimarra Stephens (1829) de MeÂxico y Centro America. (Trichoptera: Philopotamidae). Folia EntomoloÂgica Mexicana, 63, 13±23. Donnelly, T.W. (1988) Geological constraints on Caribbean biogeography. Zoogeography of Caribbean Insects (ed. by J. K. Liebherr), pp. 15±37. Comstock Publishing Associates, Division of Cornell University Press, Ithaca, New York.

128 Roger J. Blahnik Donoghue, M.J., Olmstead, R.G., Smith, J.F. & Palmer, J.D. (1992) Phylogenetic relationships of Dipsacales based on rbcl sequences. Annals of the Missouri Botanical Garden, 79, 333±345. Felsenstein, J. (1985) Con®dence limits on phylogenies: an approach using the bootstrap. Evolution, 39, 783±791. Fischer, F.C.J. (1971) Trichopterorum Catalogus, Vol. XII (Supplement to Vol. I and II). Nederlandse Entomologische Vereniging, Amsterdam. Flint, O.S. Jr (1964) The caddis¯ies (Trichoptera) of Puerto Rico. University of Puerto Rico Agricultural Experiment Station. Technical Paper, 40, 1±80. Flint, O.S. Jr (1968a) The caddis¯ies of Jamaica (Trichoptera). Bulletin of the Institute of Jamaica, Science Series, 19, 1±68. Flint, O.S. Jr (1968b) Bredin-Archbold-Smithsonian Biological Survey of Dominica, 9. The Trichoptera (caddis¯ies) of the Lesser Antilles. Proceedings of the United States National Museum, 125(3665), 1± 86. Flint, O.S. Jr (1968c) New species of Trichoptera from the Antilles. Florida Entomologist, 51, 151±153. Flint, O.S. Jr (1971) Studies of Neotropical caddis¯ies, XII: Rhyacophilidae, Glossosomatidae, Philopotamidae, and Psychomyiidae from the Amazon Basin (Trichoptera). Amazonia, 3, 1±67. Flint, O.S. Jr (1974) The Trichoptera of Surinam, studies of Neotropical caddis¯ies, XV. Studies of the Fauna of Suriname and Other Guyanas, 55, 1±151. Flint, O.S. Jr (1981) Studies of Neotropical caddis¯ies, XXVIII: the Trichoptera of the RõÂo LimoÂn Basin, Venezuela. Smithsonian Contributions to Zoology, 330, 1±61. Flint, O.S. Jr (1991) Studies of Neotropical caddis¯ies, XLV: the taxonomy, phenology, and faunistics of the Trichoptera of Antioquia, Colombia. Smithsonian Contributions to Zoology, 520, 1±113. Flint, O.S. Jr (1998) Studies of Neotropical caddis¯ies, LIII: a taxonomic revision of the subgenus Curgia of the genus Chimarra (Trichoptera: Philopotamidae). Smithsonian Contributions to Zoology, 594, 1±131.

Flint, O.S. Jr & Sykora, J.L. (1993) New species and records of caddis¯ies (Insecta: Trichoptera) from the Lesser Antilles, with special reference to Grenada. Annals of Carnegie Museum, 62, 47± 62. Holzenthal, R.W. (1988) CataÂlogo sistemaÂtico de los TricoÂpteros de Costa Rica (Insecta: Trichoptera). Brenesia, 29, 51±82. Malicky, H. (1983) TrichopteÁres des Petites Antilles (Trichoptera). Resultats de la mission hydrobiologique Austrichienne-Francaise de 1979 aux de la Guadeloupe, de la Dominique et de la Martinique (Petites Antilles), partie II. Annalen des Naturhistorischen Museums in Wien, 85B, 263±271. Rosen, D.E. (1976) A vicariance model of Caribbean biogeography. Systematic Zoology, 24, 431±464. Rosen, D.E. (1985) Geological hierarchies and biogeographic congruence in the Caribbean. Annals of the Missouri Botanical Garden, 72, 636±659. Ross, H.H. (1956) Evolution and Classi®cation of the Mountain Caddis¯ies. University of Illinois Press, Urbana, Illinois. Schmid, F. (1980) Les Insectes et Arachnides du Canada, Partie 7: Genera des TrichopteÁres du Canada et des EÁtats Adjacents. Publication 1692. Agriculture Canada, Ottawa. Schmid, F. (1982) Revision des trichopteÁres Canadiens. II. Les Glossosomatidae et Philopotamidae (Annulipalpia). Memoires de la Societe Entomologique du Canada, 122, 1±76. Swofford, D.L. (1993) PAUP: Phylogenetic Analysis Using Parsimony, Version 3.1.1. Computer program distributed by the Illinois Natural History Survey, Champaign, Illinois. Wichard, W. (1983a) KoÈcher¯iegen des Dominikanischen Bernsteins ± II. Fossile Arten der Gattung Chimarra (Trichoptera, Philopotamidae). Mitteilungen MuÈnschener Entomologisch Gesellschaft, 72, 137±145. Wichard, W. (1983b) KoÈcher¯iegen des Dominikanischen Bernsteins ± III. Chimarra succini n. sp. (Stuttgarter Bernsteinsammlung: Trichoptera, Philopotamidae). Stuttgarter BeitraÈge zur Naturkunde, Series B, 95, 1±8.

Appendix 1. Characters used in the cladistic analysis. 0 = presumed plesiomorphic state; 1, 2, 3, 4, 5 = apomorphic states; ? = missing data.

Male genitalia

Adults 1. 2. 3. 4. 5. 6. 7.

Anterior setal warts: (0) elongate, undivided; (1) divided into 2 rounded warts. Foretibial spurs (number): (0) 2; (1) one. Anal vein of hindwing (loop): (0) without loop; (1) 2A looped to 1A. Anal veins of forewing: (0) 3A looped to 2A; (1) 3A looped to 1A; (2) 2A forked to 1A and 3A. R1 vein of hindwing: (0) = not fused to subcosta; (1) fused to subcosta. Rs vein of hindwing: (0) 4-branched; (1) 3-branched (®rst fork absent). M of hindwing: (0) 3-branched; (1) 2-branched.

Accepted 26 January 2001

8. 9. 10. 11. 12. 13.

Tergum VIII (narrowing of posteromesal margin): (0) not, or only slightly narrowed; (1) distinctly narrowed; (2) extremely narrowed. Tergum VIII (posterior processes): (0) absent; (1) with scabrous posterior processes, narrowly separated; (2) with scabrous posterior processes, widely separated. Posterior processes of tergum VIII (length): (0) absent or not extremely elongate; (1) extremely elongate. Ventral process of segment IX (shape): (0) elongate, acute; (1) elongate, apex rounded or truncate; (2) short; (3) absent. Apex of ventral process of segment IX: (0) apex not truncate; (1) apex truncate or subtruncate. Tergum X (sclerous mesal process): (0) lateral lobes of tergum X not separated by sclerous mesal process; (1) lateral lobes of tergum X separated by sclerous mesal process.

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Systematics of Otarrha 14 15. 16. 17. 18. 19. 20.

21. 22. 23. 24. 25.

26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36.

Lateral lobes of tergum X (development): (0) without rodlike dorsolateral lobe; (1) with very short, rodlike dorsolateral lobe; (2) with elongate rodlike dorsolateral lobe. Ventral process of segment IX (basodorsal spine): (0) absent; (1) present. Ventromesal lobes of tergum X: (0) absent; (1) short, rounded; (2) elongate, acute and converging. Tergum X (basomesal spines): (0) without spines; (1) with row of minute spines. Tergum X (ventrolateral spines): (0) without ventrolateral spines; (1) with ventrolateral spines. Mesal process of tergum X: (0) absent or narrow; (1) very wide. Lateral lobes (basal spinous process): (0) basal spinous process absent; (1) with mesally convergent spinelike process; (2) with basal spine or spinous process separated mesally. Lateral lobes of tergum X (length): (0) short or moderate in length; (1) very distinctly elongate (not rodlike). Lateral lobes of tergum X (shape): (0) not elongate, arched; (1) elongate, arched. Lateral lobe of tergum X (apex): (0) without retrorse whiplike process; (1) with retrorse whiplike process. Lateral lobe of tergum X (lateral margin): (0) without retrorse whiplike process; (1) with retrorse whiplike process. Posteromesal process of segment IX (sclerotized process): (0) without sclerotized spinelike process at midlength; (1) with sclerotized spinelike process at midlength. Posteromesal process of segment IX (shape): (0) absent or not acuminate; (1) short acuminate; (2) elongate acuminate. Lateral lobe of tergum X (shape of dorsal margin): (0) without acute dorsomesal projection; (1) with acute dorsomesal projection. Inferior appendage (length): (0) short; (1) elongate, narrow (with mesal curvature). Inferior appendage (apical teeth): (0) without dentate apex; (1) with dentate apex. Inferior appendage (shape of preapical tine): (0) without preapical tinelike process; (1) with preapical mesal tinelike process. Inferior appendage (accessory mesal tine): (0) without small spine at base of mesal tinelike process; (1) with small spine at base of mesal tinelike process. Inferior appendage (basomesal serrations): (0) serrations absent; (1) serrations present. Inferior appendage (shape of apex): (0) apex not widened and ¯attened dorsoventrally; (1) apex widened and ¯attened dorsoventrally. Inferior appendage (mesal in¯ection): (0) if curved, without strong angular in¯ection; (1) with very angular mesal in¯ection. Inferior appendage (apicodorsal process): (0) process absent; (1) with short rounded projecting process. Apex of apicodorsal process of inferior appendage: (0) absent; (1) dorsoventrally ¯attened and extending onto mesal surface; (2) protruding, not dorsoventrally ¯attened

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37. 38. 39. 40.

41. 42. 43.

129

and not extending onto mesal surface; (3) wide, dentate on ventral half; (4) toothlike and preapical, not protruding. Spines of phallic apparatus: (0) absent; (1) short; (2) elongate, paired, symmetric; (3) elongate, asymmetric. Phallotheca (basal in¯ection): (0) not or only slightly bent; (1) strongly bent. Phallotremal sclerite complex (length and sclerosity of apical sclerite): (0) without elongate sclerous apical sclerite; (1) with elongate sclerous apical sclerite. Apical sclerite of phallotremal sclerite complex: (0) membranous or short; (1) forming elongate dorsoventrally ¯attened plates; (2) elongate, upturned and spinelike; (3) forming elongate irregular sclerites; (4) forming elongate sclerous mesal process; (5) forming elongate membranous process with paired acute apical sclerites. Phallotheca (ventral apex): (0) not sclerous, elongate and acute; (1) sclerous, elongate and acute. Phallotheca (dorsal apex): (0) not forming projecting process; (1) forming short process; (2) forming elongate de¯exed process. Preanal appendages (shape): (0) elongate, narrow; (1) short, knoblike; (2) ¯attened, earlike.

Female genitalia 44. Posteroventral setal warts of segment VIII: (0) rounded, well de®ned; (1) not de®ned (diffuse). 45. Ventral process: (0) present on segment VIII; (1) present on segment VII. 46. Segment VIII (posterolateral margin): (0) without prominent cleft; (1) with prominent cleft. 47. Tergum IX (posterior margin): (0) without setose shelflike projection; (1) with setose shel¯ike projection. 48. Tergum X: (0) without sclerous basal lobe; (1) with sclerous basal lobe. 49. Segment VIII (length): (0) segment short, not elongate ventrally; (1) segment elongate ventrally.0 50. Segment VIII (posteroventral margin): (0) without membranous extension; (1) with membranous extension. 51. Segment VIII (sclerotized lateral suture line): (0) suture line absent; (1) suture line present. 52. Cuplike sclerite of vaginal apparatus: (0) located at anterior apex of vaginal apparatus; (1) located ventrally, before apex of vaginal apparatus. 53. Clasper receptacles: (0) without prominent intersegmental clasper receptacles; (1) with prominent intersegmental clasper receptacles between segments VII and VIII. 54. Vaginal apparatus (apical sclerites): (0) without paired apicodorsal sclerites; (1) with paired apicodorsal sclerites. 55. Segment VIII (notch): (0) without ventromesal notch on posterior margin; (1) with ventromesal notch on posterior margin. 56. Vaginal apparatus (ventral sclerotization): (0) without abruptly terminating ventral collar before cuplike sclerite; (1) with abruptly terminating, lightly sclerous ventral collar before cuplike sclerite.

130 Roger J. Blahnik Appendix 2. Character matrix for Chimarra (Otarrha) species.

Taxa

11111111112222222222333333333344444444445555555 12345678901234567890123456789012345678901234567890123456

C. amazonia 11111111201010000000000001010100000010000021111100000100 C. barinas 1111111000000000000000000000100000000000012????????????? C. cubanorum 1111111000100000010100000000000000120000002????????????? C. darlingtoni 1111111000100000010100000000000000130000002????????????? C. diakis 1111111010101100000000001001000010000011002????????????? C. diaphora 111111101010110000000000100100001000001100211111000?0100 C. dominicana 11111110001011010010000000010000110000120021111110111100 C. garciai 1111111000100000000211000000000000110000002????????????? C. guapa 11111110001000000002110000010000000000000021111100000101 C. incipiens 1111111000000000000000000000100000000000122????????????? C. jamaicensis 11111110001000000000100000000000001100000021111100000101 C. machaerophora 11111110001000000000100000000000001100000021111100000101 C. odonta 01110010001000000000000000000000000000100020111101000100 C. parene 1111111100111010000000110111011000001000002????????????? C. parilis 11111111001110100000001101110111000010000021111100000110 C. particeps 11111111001110000000001101110101000010000021111100000110 C. patosa 11111112211010000000000002010100000021000021111100000100 C. peruana 1111111221101000000000000201010000002100002????????????? C. phthanorossi 11111110001012011000000000010000110020130021111110110100 C. puertoricensis 11111110001010000100000000000000000000000021111101000101 C. quadrifurcata 11111110001011000010000000010000110020120021111110111100 C. redonda 11111110001000000100000000000000001400000021111101000101 C. retrorsa 111111110010?202000000100001000011002015002????????????? C. rossi 11111110001012011000000000010000110020130021111110110100 C. sensillata 1111111000000000000000000000100000000000122????????????? C. septemlobata 11111110101111010000000000000000000000000021111100000100 C. septifera 111111110010?202000000000001000011000014002??111???????? C. spinulifera 11111110001000000101000000000000001200000021111100000100 C. tachuela 11111110000000000000000000001000000000000120111100000000 Chimarrhodella costaricensis 0 0 0 0 0 0 0 0 0 0 3 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Chimarrhodella ulmeri 00000000003000000000000000010000000010001000000010000000 Chimarra (Chimarrita) neblina 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 1 1 0 0 ? 0 0 0 0 C. (Chimarrita) maldonadoi 0 1 1 2 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 0 0 0 0 0 0 0 ? 0 0 0 0 C. (Curgia) banksi 01110000002000000000000000000000000000000010000000000010 C. usitatissima 01111110000000000000000000000000001100000010000000000000 C. (Chimarra) emima 01120000002000000000000000000000000030001010100000000000

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